New marine reptile fossils from the Late Jurassic of Poland with implications for vertebrate faunas palaeobiogeography

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New marine reptile fossils from the Late Jurassic of Poland with implications for vertebrate faunas palaeobiogeography b _ Błazejowski _ Daniel Tyborowskia,* , Błazej a b

Museum of The Earth, Polish Academy of Sciences, Aleja Na Skarpie 20/26, 27, 00-488 Warszawa, Poland Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warszawa, Poland

A R T I C L E I N F O

A B S T R A C T

Article history: Received 10 May 2019 Received in revised form 19 September 2019 Accepted 19 September 2019 Available online xxx

_ A new marine vertebrate assemblage from the Late Jurassic (late Kimmeridgian) at Krzyzanowice near _ in the NE margin of the Holy Cross Mountains in Poland is described. This new locality is rich in Iłza fossils of coastal and pelagic reptiles. The most frequent fossils are plesiochelyid turtle shell fragments _ and pliosaurid skull bones and teeth. The Krzyzanowice vertebrate assemblage is similar to the Late Jurassic Boreal/Sub-Boreal localities of the Kimmeridge Clay in Great Britain and Svalbard Archipelago in the Arctic, in the presence of pliosaurids and long-necked plesiosaurids. However, plesiochelyid turtles and crocodylomorphs are similar to those from the Mediterranean/Sub-Mediterranean sites of the northern border of the Tethys Ocean, as, for example, in the Swiss Jura Mountains and Southern _ vertebrate fauna demonstrates that, during the Germany. This unique composition of the Krzyzanowice Late Jurassic this new locality was located in the transitional palaeobiogeographic line referred to in this paper as the “Matyja-Wierzbowski Line”. The new palaeobiogeographical reconstructions of Late _ locality and other sites with Jurassic of Europe are based on the composition of the Krzyzanowice similar turtle-pliosaurid faunas which formed a long-term, stable ecological sympatry in marine ecosystems of the European Archipelago. © 2019 Published by Elsevier Ltd on behalf of The Geologists' Association.

Keywords: Pliosauroidea Testudinata Crocodylomorpha Late Jurassic Palaeobiogeography Poland

1. Introduction Since 2012 the Upper Jurassic strata of the Holy Cross Mountains margin in Poland have produced a wealth of vertebrate remains (Kin _ _ and Błazejowski, 2012; Kin et al., 2013; Błazejowski et al., 2016; Tyborowski, 2016, 2017; Tyborowski et al., 2016). The vertebrate assemblages typically have a European Late Jurassic composition that includes actinopterygian fish, ichthyosaurs, crocodyliforms, turtles and rare pterosaurs (Kin et al., 2013; Tyborowski et al., 2016). The best known of these localities, the Owadów-Brzezinki Quarry in the Łód z Voivodeship (Fig. 1), yields the Caturus giganteus, “Furo microlepidotes” and Orthocormus teyleri, and the marine reptiles Cryopterygius kielanae and Owadowia borsukbialynickae _ (Błazejowski et al., 2015; Tyborowski, 2016, 2017; Szczygielski et al., 2018). The occurrence of characteristic species of ammonites (Matyja and Wierzbowski, 2016) indicate an early/late Tithonian _ boundary (middle Volgian) age (Błazejowski et al., 2016). In 2010, a new Upper Jurassic (Oxfordian) locality (the Morawica quarry) with marine reptile remains was described in the Holy Cross Mountains region (Fig. 1) which yielded a partially articulated cranial skeleton

and isolated bones and teeth of a large ichthyosaur Ophthalmosauridae indet. (Tyborowski et al., 2018). In an another quarry at Zalas near Kraków, plesiosaurian teeth from the Upper Jurassic (lower Oxfordian) strata have been described (Lomax, 2015). This paper describes the discovery of a new and very rich assemblage of marine vertebrates from the late Kimmeridgian _ carbonate sequence at the Krzyzanowice site in the Holy Cross Mountains of Poland and preliminary descriptions of the marine reptiles (turtles, pliosaurs, plesiosaurs and crocodiles). This paper also discusses the first results of a taxonomic analysis of the _ vertebrate assemblage from the Krzyzanowice site, which, it is hoped, will be helpful for other palaeobiogeographical studies of the Kimmeridgian and Tithonian turtle and sauropterygian communities across Europe. In addition, mechanisms responsible for concentrating the vertebrate fossils are considered in terms of palaeoenvironment and taphonomy. Institutional abbreviations. – MZ, Museum of The Earth, Polish Academy of Sciences, Warsaw, Poland; ZPAL, Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland. 2. Geological and palaeontological context

* Corresponding author. E-mail address: [email protected] (D. Tyborowski).

The strata described here represent the youngest part of the Jurassic succession preserved below the Lower Cretaceous deposits

https://doi.org/10.1016/j.pgeola.2019.09.004 0016-7878/© 2019 Published by Elsevier Ltd on behalf of The Geologists' Association.

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Fig. 1. Geological sketch map of the Holy Cross Mountains in Poland, showing locations of the Upper Jurassic bone-bearing palaeontological sites (Morawica = Oxfordian; _ Krzyzonowice = Kimmeridgian; Owadów-Brzezinki = Tithonian).

in the north-eastern margin of the Holy Cross Mountains which _ crop out mostly between the villages of Krzyzanowice and Malenie _ (Fig. 2A). They consist of several rock units described north of Iłza _ in the older geological studies of Pozaryski (1948); Da˛browska (1953, 1957): (1) grey marly clays with intercalations of Nanogyra lumachelles, several tens of meters in thickness; (2) bioclastic packstones and grainstones (“conglomerates”) composed mostly of oyster shell hash, oyster-brachiopod lumachelles, and sandstones of shell detritus with glauconite; all these deposits contain larger or smaller amounts of siliciclastic grains – quartz and fragments of dark flints, their total thickness ranges up to 4–8 m; (3) yellow-coloured limestones (“canary-yellow limestones”) with a rich fauna of gastropods, composed mostly of large nerineids (Fig. 2D) including a diversified assemblage of Nerinea, Cryptoplocus, Ptygmatis, Pseudonerinaea and Nerinella (Karczewski, 1960), bivalves and brachiopods – which is the main level of bone accumulations (“turtle limestones”) (Borsuk-Białynicka and Młynarski, 1968). This rock unit may reach up to 6 m in thickness, but it may be thinner in some places; (4) yellowish marls without macrofauna but with ostracods; up to 2 m in thickness. The directly overlying deposits are clays and marls which yielded the ammonites of Valanginian age. According to Da˛browska (1957) there exists a small angular unconformity within the Jurassic deposits (between units 2 and 3) resulting from local tectonic movements; this, however, has been indirectly questioned by Kutek (1962) who recognized that the deposits of unit 3 are crossbedded. These youngest Jurassic deposits of the north-eastern

margin of the Holy Cross Mountains (Da˛browska, 1953, 1957; and earlier papers cited therein) were deposited in a shallow marine environment, substantially of a very shallow-water depth during sedimentation of the biodetrital limestones of unit 2. The large facies contrast between the grey marly clays with the intercalation of Nanogyra lumachelles of unit 1 of an open-marine environment, and the biodetrital limestones of unit 2 of a very shallow-water environment could have resulted from sea-level oscillation marked by shallowing (?regression) and the subsequent appearance of new transgressive deposits (unit 3) (Kutek, 1962). Biostratigraphical documentation of the deposits in question is, however, poor. The deposits of unit 1 could possibly be of Upper Kimmeridgian age but have so far yielded only “indistinguishable fragments of ammonites bearing perisphinctid-like ribbing” (Gutowski, 1998), whereas the yellow limestones with nerineid gastropods have so far yielded a single specimen referred to as “Cardioceras” Amoeboceras (Nannocardioceras) cf. anglicum (Fig. 2B) by Da˛browska (1957) which indicates a higher part of the Upper Kimmeridgian (possibly the Autissiodorensis Zone – Kutek and Zeiss, 1997) (Fig. 3). The new vertebrate assemblage is dominated by the bones of turtles (mainly shell fragments) and a pliosaurid sauropterygian jaw fragment and isolated teeth (Fig. 5). Bones of long-necked plesiosaurs and teeth of crocodylomorphs are very rare at this locality. Other vertebrate remains, including actinopterygian fish teeth, are also rare. Among the new fossils, the most interesting are the remains of turtles and large pliosaurs. This is the easternmost

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_ Fig. 2. Palaeontological excavations and invertebrate fossils from the Krzyzonowice site. (A) Geological profile of the uppermost Kimmeridgian sediments. (B) Ammonite. _ (C) Field works in the Krzyzanowice site. (D) Large gastropod Nerinea.

locality with turtle remains from the Upper Jurassic of Europe and the third find of large pliosaurids in this part of the world, after finds of the superpredatory pliosaurs from Svalbard and England (Hurum et al., 2012; Benson et al., 2013). 3. Material and methods _ site was The first time that the fossiliferous Krzyzanowice  ska, Gwidon Jakubowski investigated was in 1962 by Teresa Maryan (Museum of Earth, Polish Academy of Sciences), Zofia KielanJaworowska and Magdalena Borsuk-Białynicka (Institute of Paleobiology, Polish Academy of Sciences). At that time, the scientists found turtle remains and fragments of a pliosaur jaw. Turtle remains were described by Borsuk-Białynicka and Młynarski (1968). In the summer and autumn of 2018, the team led by Daniel Tyborowski performed the first paleontological excavations _ in Krzyzanowice (Fig. 2), which produced new, very rich bone material. The collected fossils are housed in the Museum of The Earth, Polish Academy of Sciences, Warsaw and in the Institute of Paleobiology, Polish Academy of Sciences, Warsaw. The studied material is preserved as original bone fragments and also as silicon/latex casts. All vertebrate silicon/latex casts have been prepared manually in the Institute of Paleobiology, Polish

Academy of Sciences. Selected specimens were drawn with a camera lucida. All skeletal elements consist of detached and disarticulated bones of various sizes. The described fossils are thus difficult to distinguish at the species level because it was not possible to compare their size and proportions, so they may represent one or several taxa. The analyzed vertebrate material consists of 6 described specimens representing isolated or disarticulated parts of the axial skeleton (plesiosaur) jaw and tooth (pliosaur) and shell (turtles). Specimen descriptions are based on standard characters that are commonly used in the literature, and can be found in most recent datasets. The described material is only a small part of the enormous collection of marine _ reptile bones from the Krzyzanowice site. In this paper we describe only 6 of the most informative specimens: MZ VIII Vr-71-72 and ZPAL V-KRZ/32-33. Quantitative analysis of vertebrate material was made on one piece of turtle bone-bearing breccia and other specimens seen on the limestone were counted. Each isolated bone fragment was treated as a separate object. Due to the high degree of fragmentation of almost all marine reptile bones (plates, teeth, flat skull bones), we decided to count each specimen individually despite the mode of preservation. For example, small and broken parts of turtle costals had the same value as a complete large

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_ Fig. 3. Reconstructed food web for the marine components of the Krzyzanowice community. Lines with arrows indicate the simplified assumed feeding pathways: (a) large pliosaurid sauropterygian reptile (Pliosaurus sp.); (b) large elasmosaurid plesiosaur (Elasmosauridae indet.); (c) small teleosaurid crocodylomorph (Machimosaurus sp.); (d) ammonite; (e) pachycormiform actinopterygian fish; (f) pycnodontiform actinopterygian fish; (g) plesiochelyid turtle (Plesiochelyidae indet.); (h) oysters (Nanogyra) and other bivalves (Pleuromya); (i) brachiopods; (j) large gastropods (Nerinea, Cryptoplocus, Ptygmatis, Pseudonerinaea and Nerinella).

pliosaur jawbone during the count. In total, 395 bone elements were counted. These comprise 248 fragments of turtle shell, 115 unidentified bone fragments, 12 fragments of pliosaurid skull and teeth, 10 bones from long-necked plesiosaurs (axial skeleton), 7 fragments of crocodiles (teeth and tooth crowns) and 3 actinopterygian teeth. The turtle remains belong to many individuals. The term “turtle limestone”, as used in this paper means any Upper Jurassic limestone from the border of the Holy Cross Mountains rich in vertebrate remains, among which turtle bones dominate.

show well-preserved surfaces presenting important microanatomical features such as Haversian canals and ornamentation (depending on the marine reptile group). All flat bones such as the plates of the turtle shell are oriented parallel to bedding. There are zones of contrasting bone density and this probably suggests periodical accumulation of skeletal fragments on the slopes of ripples during current activity.

4. Palaeoenvironment and taphonomy of vertebrate fossils

TESTUDINES Batsch (1788) EUCRYPTODIRA Gaffney (1975) PLESIOCHELYIDAE Baur (1888) Plesiochelyidae gen. et sp. indet. _ _ (Holy Cross Locality and horizon: Krzyzanowice near Iłza Mountains, Central Poland), Upper Kimmeridgian (Autissiodorensis Zone). Material: Fragmentary carapace and a single left costal; all elements collectively labelled MZ VIII Vr-71 (Fig. 4). All carapace fragments were collected together, and they are regarded here as belonging to two individuals. Description and comparison: Isolated left first costal (Fig. 4A-B) of a medium-sized individual (length, 46 mm; medial width, 27 mm) is incomplete and eroded on its proximal edge. The bone was probably slightly larger during the life of the animal. This costal is a very thick element (17 mm). The posterior edge of the bone is very convex distally. There are no costoperipheral fontanelles. These characteristic features are consistent with the Late Jurassic Plesiochelyidae turtles (Anquetin et al., 2014), especially with the genus Tropidemys known from the Late Jurassic Solothurn Turtle Limestone of Switzerland (Meyer, 1994; Püntener et al., 2014) and the Kimmeridge Clay Formation of southern England (Anquetin and Chapman, 2016). There are no crest-like

_ The vertebrate remains from the Krzyzanowice site contains mainly fragments of turtle shells, jaw bones and teeth of large pliosaurs, axial elements of long-necked plesiosaurs, as well as undeterminable tooth crowns of marine crocodiles and small actinopterygian fish teeth. Turtle plates, crocodile and pliosaur jaws, teeth and other dermal elements, as well as possibly some elements of visceral skeleton derived from the bone breccias are preserved as original bone tissue in fine-grained limestones, forming bone-beds. The quartz grains from these turtle-rich limestones are well-rounded and sorted. The turtle shell fragments, sauropterygian teeth, jaws and axial elements, fish teeth and other skeletal parts have suffered only slight distortion, and show such anatomical details as microsculpture of plate surfaces and blood vascular canals of dentary bones and teeth and can be examined from the silicon or latex casts. This method is easy to apply and allows the observation of every detail of the bone morphology including microstructure. Occasionally, small parts of the bones and teeth have beenmodified by diagenetic processes. Some bone accumulations contain rare, highly abraded and isolated bone fragments, although the surfaces of most of the bones are rarely eroded. In most cases, the vertebrate remains

5. Systematic palaeontology

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Fig. 4. Plesiochelyid turtle (Plesiochelyidae indet.) shell remains, MZ VIII Vr-71 _ from the Krzyzanowice site, Holy Cross Mountains, Poland. (A) Isolated left first costal in ventral view. (B) Isolated left first costal in dorsal view. (C) Carapace fragment in ventral view. (D) Carapace fragment in dorsal view.

structures on the ventral surface of the bone a feature that is diagnostic for Tropidemys langii species (Püntener et al., 2014). Tropidemys (Pelobatochelys) blakii lacks this feature and the condition of this morphology in Tropidemys seebachi is unknown. However, the suture with the nuchal and the region where the crest-like articular site for the first thoracic rib are present in Tropidemys langii are not preserved in this specimen. Therefore, we are unable to say which species of Tropidemys this specimen belongs to. This specimen can be confidently assigned to the Plesiochelyidae indet. The carapace fragment (Fig. 4A-B) of a large individual consists of the right three costals, single left costal and two neurals. The first right costal (length, 102 mm; medial width, 35 mm) is incomplete and damaged distally. The distal part of the second right costal (length, 79 mm; medial width, 29 mm) is also damaged and broken off. The third right costal (length, 98 mm; medial width, 30 mm) is also incomplete distally. The ribs are present on the ventral surfaces of all three right costal plates. Small bone fragments from another costal are preserved at anterior margin of the first right costal. From the left side of the carapace, only a small fragment of the first costal is preserved. This costal plate (length, 51 mm; medial width, 29 mm) is broken distally. The left costal extends proximally. On the ventral surface of the left costal there is a welldeveloped rib. The right and left costals of this carapace fragment are slightly arched posteriorly. The neurals are approximately hexagonal, up to 32 mm in length and up to 27 mm in maximum width (anterior region of the bones). The right anterolateral margin

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of the first neural plate is broken. The anterolateral margins of these neurals are short (10 mm). The posterolateral margins are much longer (15 mm). Anteriorly, contact with the preceding neural is concave and gently rounded. Posteriorly, contact with the following neural is formed by two oblique facets. The presence of a gently sagittal keel on the dorsal surface of the neurals is diagnostic for the members of Tropidemys (Püntener et al., 2014). None of the other Late Jurassic European turtles have keeled neurals (Püntener _ et al., 2014). The new specimen from Krzyzanowice lacks keeled neurals. The elongated morphology of these plates, with the anterolateral margins significantly shorter than the posterolateral ones, is compatible with Tropidemys (Pelobatochelys) blakii. The concavity of the anterior margin of the neural plates is recognized as more developed in this species than that present in the other members of plesiochelyid turtles (Pérez-García, 2015). This specimen is interpreted as Plesiochelyidae indet. Given that all the fragments of turtle bones were collected from the same layer of limestone, exhibit the same mode of preservation and are similar anatomically, all of them are confidently assigned to a single family Plesiochelyidae in many different ontogenetic stages. SAUROPTERYGIA Owen (1860) PLESIOSAURIA de Blainville (1835) PLIOSAURIDAE Seeley (1874) THALASSOPHONEA Benson and Druckenmiller (2013) Pliosaurus Owen (1841) Pliosaurus sp. _ _ (Holy Cross Locality and horizon: Krzyzanowice near Iłza Mountains, Central Poland), Upper Kimmeridgian (Autissiodorensis Zone). Material: Large fragment of the upper jaw (premaxilla) and a large fragment of the lower jaw (dentary) with preserved teeth (Fig. 5) and a silicon cast of an isolated tooth (Fig. 6). All bone and teeth elements collectively labelled MZ VIII Vr-72. Description and comparison: A large fragment of the upper jaw (left premaxilla) within limestone block (Fig. 5B). The premaxilla forms the lateral surfaces of the snout. The body of the premaxilla is mediolaterally broad and dorsoventrally low. The premaxilla bears 10 alveoli. The first alveolus (mesialmost) is highly recurved, with a minimum diameter (17 mm) approximately half that of the third alveolus (42 mm). The reduction of the first premaxillary alveolus is interpreted as a Pliosaurus + Brachaucheninae synapomorphy (Benson and Druckenmiller, 2013; Benson et al., 2013). In Brachaucheninae however, the first alvelous is much more reduced than it is in the members of Pliosaurus (Benson et al., 2013). This condition in Pliosaurus is considered autapomorphic (Benson et al., 2013). The premaxillary alveoli from second to ten of MZ VIII Vr-72 are the largest, demonstrating the presence of a heterodont dentition. This specimen differs from some Kimmeridge Clay Formation pliosaurids, in which the distalmost alveolus is slightly reduced (Taylor and Cruickshank, 1993). The premaxilla has conspicuously convex lateral margins. The premaxillary dentition is separated from the maxilla by a region of compact bone forming a diastema with a characteristic “zig-zag” suture. This diastema and premaxilla-maxilla junction are preserved in the psterior-most region of the specimen. In dorsal and lateral views, the surfaces of premaxilla are fractured and covered by ‘wrinkles’ and foramina. The anteriormost surface of the specimen is eroded on its dorsal portion. The premaxillary teeth are preserved in positions 2–5, 7 and 9. All of the right premaxillary teeth are preserved. All of the premaxillary teeth are trihedral which is an autapomorphy of Pliosaurus (Benson et al., 2013). Specimen MZ VIII Vr-72 is interpreted as Pliosaurus sp. on following combination of characters: the reduction of first premaxillary alveolus; the presence of diastema between the premaxilla and maxilla forming a characteristic “zig-zag” suture; trihedral morphology of the premaxillary teeth.

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_ Fig. 5. Large pliosaurid sauropterygian reptile (Pliosaurus sp.) jaws and teeth, MZ VIII Vr-72 from the Krzyzanowice site, Holy Cross Mountains, Poland. (A) Dentary in lateral view. (B) Praemaxilla in medial view.

A large fragment of the right dentary is preserved in the same limestone block (Fig. 5A). The preserved fragment of the dentary is 132 mm long. The surface of the bone is wrinkled. The dentary bears 5 alveoli. The dentition from the dentary is anisodont just like in the premaxillary dentition. All of the dentary teeth are preserved. The crowns of all teeth are robust, conical, gently recurved lingually and display gently longitudinal ridges throughout. The longitudinal ridges are regularly and evenly spaced throughout. These ridges disappear near the tip of the tooth crown. The tooth crown apex is lingually recurved. The total length of the largest dentary tooth measured from the base of the preserved part of the tooth root to the apical-most portion of the crown is 68 mm. The morphology of all preserved dentary teeth is most similar to the shapes and morphologies of several Middle and Late Jurassic pliosaurids (Massare, 1987; Benson et al., 2013). The broad-based conical shape and general morphology of the tooth are characteristic features of a posterior tooth of the Pliosauridae (Seeley, 1874; Benson et al., 2013). The best comparative example of the Late Jurassic pliosaurids is Pliosaurus. The genus Pliosaurus comprises many isolated bones, skeletons and teeth that were described from

several Upper Jurassic localities in many places of the world (Benson et al., 2013). Six currently valid species of Pliosaurus are known (Benson et al., 2013). The genus Pliosaurus is based on seven autapomorphies (Benson et al., 2013). One of these autapomorphies includes trihedral or subtrihedral teeth (Benson et al., 2013). The preserved dentary teeth share trihedral morphology. Hence the studied specimen most likely is Pliosaurus and it is interpreted as Pliosaurus sp. A silicon cast of a large isolated tooth (Fig. 6) measures 57 mm in apicobasal length. The apicalmost 30 mm represent the tooth crown and the basalmost 27 mm are interpreted as the root (Fig. 6A). This specimen is conical, curved and its cross-section has flattened labial and lingual surfaces (“subtrihedral”). Its distal surface bears prominent, apicobasally-oriented carinae. The labial surface of the tooth bears many apicobasal enamel ridges (Fig. 6B). The surface of the tooth is flattened. This feature is present in many Late Jurassic pliosaurid teeth (Taylor and Cruickshank, 1993; Knutsen et al., 2012; Benson et al., 2013) and also in pliosaurids from the Early and Middle Jurassic (Benson et al., 2011). This feature is also an autapomorphy of the genus Pliosaurus (Knutsen,

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TELEOSAURIDAE Geoffroy Saint-Hilaire (1831) Machimosaurus von Meyer (1837) Machimosaurus sp. _ _ (Holy Cross Locality and horizon: Krzyzanowice near Iłza Mountains, Central Poland), Upper Kimmeridgian (Autissiodorensis Zone). Material: a single tooth number ZPAL V-KRZ/33. Description and comparison: Small and fragile teeth and tooth _ crowns of crocodylomorphs are common in the Krzyzanowice site. ZPAL V-KRZ/33 is an isolated tooth crown. This specimen displays a robust conical morphology. The apex is rounded. The enamel surface is not smooth but presents numerous wrinkles. The crown has a circular cross-section. Several pronounced vertical striae ornate the enamel and this specimen is interpreted as Machimosaurus sp. 6. Palaeobiogeography and bioprovincialism

Fig. 6. Isolated tooth of the large pliosaurid sauropterygian reptile (Pliosaurus sp.) in _ axial view, MZ VIII Vr-72 from the Krzyzanowice site, Holy Cross Mountains, Poland. (A) Silicon cast. (B) Sketch drawing.

2012; Benson et al., 2013). Benson et al. (2013) suggests that this feature may by an autapomorphy of Pliosaurus kevani. This specimen is also interpreted as Pliosaurus sp. PLESIOSAUROIDEA Gray (1825) ELASMOSAURIDAE Cope (1869) Elasmosauridae gen. et sp. indet. _ _ (Holy Cross Locality and horizon: Krzyzanowice near Iłza Mountains, Central Poland), Upper Kimmeridgian (Autissiodorensis Zone). Material: A single pectoral centrum (Fig. 7) number ZPAL V-KRZ/32. Description and comparison: Bones of the long-necked plesiosaurs are represented only by elements of the axial skeleton. ZPAL V-KRZ/32 is an isolated pectoral centrum. It is much wider than it is long and high. The articular surfaces of the bone are ovoid and amphicelous (Fig. 7A-B). These surfaces share well-defined margins without ventral notch. A large two articular facets for the neural arch are present on the dorsal surface of the bone (Fig. 7C). The ventral surface is gently convex and bears two very large foramina subcentralia. These foramina are separated from each other. The lateral surfaces of the bone as a whole are smooth and lack foramina. In view of its proportions and shape, in particular the ovoid morphology of the articular surface, which is wider than it is long and high, this centrum could belong, among Plesiosauroidea, to an indeterminate elasmosaurid. This centrum lacks the ventral notch that gives the articular surface a typical “binocular” morphology. The ventral notch is absent in elasmosaurids older than Late Cretaceous (Sachs and Kear, 2014) but is common in Late Cretaceous forms (Bardet et al., 1999; Druckenmiller and Russell, 2008). CROCODYLOMORPHA Hay (1930) THALATTOSUCHIA Fraas (1901)

Studies of Late Jurassic (Kimmeridgian/Tithonian) marine reptile diversity have demonstrated the presence of separate palaeobiogeographical zones in Europe during this time (Fig. 8), each containing a distinct fauna (Hurum et al., 2012; Anquetin and Chapman, 2016). As noted by these authors, the Late Jurassic is especially well-suited for palaeobiogeographical studies on plesiochelyid turtles, pliosaurids, plesiosaurids and ophthalmosaurid ichthyosaurs as it has yielded a high number of individuals and a wide diversity of well-preserved specimens. A simplified palaeobiogeographical system has recognized two main zones, a Boreal province and a Mediterranean province (Druckenmiller et al., 2012; Hurum et al., 2012; Anquetin and Chapman, 2016). Ophthalmosaurid ichthyosaurs are present in both zones whereas large pliosaurids and long neck plesiosaurids are unique to the Boreal/Sub-Boreal province (Knutsen et al., 2012; Benson et al., 2013), whereas plesiochelyid turtles are characteristic for the Mediterranean/Sub-Mediterranean province (Anquetin and Chapman, 2016). Localities with Late Jurassic strata across Europe and Russia have yielded abundant vertebrate remains, mainly of Kimmeridgian and Tithonian age (Druckenmiller and Knutsen, 2012; Druckenmiller et al., 2012; Knutsen et al., 2012; Benson et al., 2013; Anquetin et al., 2014; Anquetin and Chapman, 2016). Arguably, the best known are the Kimmeridge Clay Formation outcrops between Dorset and Yorkshire of southern and eastern England which have yielded abundantisolated, semiarticulated and articulated remains of marine reptiles (e.g. Benson et al., 2013; Anquetin and Chapman, 2016). These marine reptile faunas contain cryptodiran turtles (Plesiochelys, Tropidemys), crocodylomorphs (Machimosaurus, Dakosaurus, Metriorhynchus, Steneosaurus), plesiosaurs (Kimmerosaurus, Colymbosaurus), large pliosaurids (Pliosaurus, Liopleurodon) and ophthalmosaurid ichthyosaurs (Ophthalmosaurus, Brachypterygius, Nannopterygius). The members of the pliosaurid genus Pliosaurus were evidently the top predator in the local food chain (Benson et al., 2013). Turtle material from the Kimmeridge Clay contains three taxa unique to this formation: Tropidemys (Pelobatochelys) blakii, Enaliochelys chelonia and Tholemys passmorei. The rest of the tetrapod fauna from the Kimmeridge Clay represents members of plesiochelyids (Plesiochelys etalloni, Tropidemys langii), thalassemydids (Thalassemys hugii, Thalassemys bruntrutana) and eurysternids (Anquetin and Chapman, 2016). The Late Jurassic ecosystems of the Kimmeridge Clay belong to the Sub-Boreal Seaway and represent the British Sub-Boreal biogeographic province (Benson et al., 2010). Coeval assemblages have also been described from the Swiss Jura Mountains (Solothurn and Porrentruy) and the Hannover area of Lower Saxony in Germany and from Wattendorf in Bavaria, Germany (Portis, 1878; Püntener et al., 2014). These localities

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_ Fig. 7. Elasmosaurid plesiosaur (Elasmosauridae indet.) isolated pectoral centrum, ZPAL V-KRZ/32 from the Krzyzanowice site, Holy Cross Mountains, Poland. (A) Articular view. (B) Lateral view. (C) Dorsal view.

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Fig. 8. Palaeobiogeography of the Late Jurassic (Kimmeridgian/Tithonian) marine reptile faunas of Eurasia. Boreal marine reptile faunas are located in the Arctic (A, Hurum et al., 2012) and Western Russia (R, Arkhangelsky and Zverkov, 2014). Mediterranean marine reptile assemblages are in Swiss Jura Mountains (S, Anquetin and Chapman, 2016) and Southern Germany (G, Anquetin and Chapman, 2016). The palaeontological sites with both Boreal and Mediterranean marine reptile faunas are located in Southern England (E, Benson et al., 2013) and Central Poland (P, Tyborowski et al., 2016). Palaeobiogeographical map courtesy of R. Blakey.

contain diverse plesiochelyid turtle fauna including the following taxa: Plesiochelys etalloni, Craspedochelys jaccardi, Tropidemys langii and Tropidemys seebachi (Portis, 1878; Anquetin et al., 2014; Püntener et al., 2014; Anquetin and Chapman, 2016). Thalassemydids were also present in Swiss Jura Mountains during the Late Jurassic. As in the Kimmeridge Clay Formation, turtles of the species Thalassemys hugii and Thalassemys bruntrutana are both present in the Solothurn turtle limestones of Switzerland (Püntener et al., 2015). The Late Jurassic localities of the Swiss Jura Mountains and southern Germany are dominated by turtles and clearly belong to the Sub-Mediterranean seas of the Tethys Ocean (Fig. 8). The Arctic region has also yielded some Late Jurassic (Tithonian) marine reptiles. The assemblage known from the Agardhfjellet Formation (Hurum et al., 2012) is characterized by the cooccurrence of large pliosaurids (Pliosaurus funkei), plesiosaurids (Spitrasaurus wensaasi, Spitrasaurus larseni, Djupedalia engeri) and ophthalmosaurid ichthyosaurs (Cryopterygius kristiansenae, Janusaurus lundi, Palvennia hoybergeti) (Druckenmiller and Knutsen, 2012; Druckenmiller et al., 2012; Knutsen et al., 2012; Roberts et al., 2014). These marine reptile faunas are clearly endemic and different from other marine reptile faunas in the Late Jurassic Europe (Fig. 8). They are considered to belong to a different

biogeographical province, termed the Boreal Realm (Hurum et al., 2012). A similar boreal fauna of marine reptiles (large pliosaurids, plesiosaurids and ophthalmosaurid ichthyosaurs) is present in western Russia (Arkhangelsky and Zverkov, 2014). Although most marine reptilian specimens have been found as _ isolated bones and teeth at Krzyzanowice, of note is the cooccurrence from the Late Jurassic (late Kimmeridgian) of both plesiochelyid turtles and large pliosaurids, as well as elasmosaurid plesiosaurs and small crocodylomorphs. Consequentlythe Krzy_ zanowice marine reptile assemblage appears to have been a palaeobiogeographically unique ecosystem in containing both Mediterranean/Sub-Mediterranean turtles and Boreal/Sub-Boreal pliosaurids. To summarize, these two groups of the Late Jurassic marine reptiles formed a stable ecological sympatry along the entire contact zone of the warm Tethys Ocean and colder SubBoreal seas during the Kimmeridgian – Tithonian. Thus, the Late Jurassic ecosystems in Europe, with plesiochelyid turtles and large pliosaurids are “palaeobiogeographical hubs” located at the border of both Sub-Mediterranean/Mediterranean and Boreal/Sub-Boreal provinces. In order to systematize this phenomenon, we propose the term Matyja-Wierzbowski Line (Fig. 8), which may be understood as: the hypothetical faunal/biogeographical boundary separating the

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ecozones/palaeobiogeographical realms of the warm Tethys Ocean and colder seas at the north, a transitional zone between Mediterranean/Sub-Mediterranean Province and Boreal/Sub-Boreal Province. It is recognised by the co-occurrence of plesiochelyid turtles (Mediterranean fauna) and large pliosaurid reptiles (Boreal fauna). The name of this new palaeobiogeographical belt commemorates two eminent researchers of Late Jurassic paleobiogeography – Bronisław Andrzej Matyja and Andrzej Wierzbowski. The MatyjaWierzbowski Line extends through Dorset and Yorkshire regions of England, between Fennoscandia and the Bohemian Massif and through the Teisseyre-Tornquist Zone. So, the paleogeography of the Matyja-Wierzbowski Line is similar and parallel to the TransEuropean Suture Zone. Paleontological localities and horizons such as the Kimmeridge Clay Formation, Owadów-Brzezinki Quarry and _ Krzyzanowice are representatives of the sites located at this important palaeobiogeographical line. Acknowledgements We wish to thank Andrzej Wierzbowski (Faculty of Geology, University of Warsaw, Poland) for useful advice on biostratigraphy and palaeontology of the Late Jurassic of Europe. We thank the mayor _ Mr. Przemysław Burek, for his help during the organisation of of Iłza, fieldwork. Special thanks to Gwidon Jakubowski (Museum of the Earth, Polish Academy of Sciences, Warsaw, Poland) and Magdalena Borsuk-Białynicka (Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland) for the invaluable support and enable  ski (Institute of _ research in the Krzyzanowice site, Marian Dziewin Paleobiology, Polish Academy of Sciences, Warsaw, Poland) and Dariusz Nast (Museum of the Earth, Polish Academy of Sciences, Warsaw, Poland) for help with photographs, Piotr Gieszcz (Faculty of Geography and Regional Studies, University of Warsaw, Poland) and all students of the Faculty of Geology of the University of Warsaw in Poland for the help during fieldwork in 2018 and 2019. We sincerely thank the two anonymous reviewers and the editor Malcolm Barrie Hart for many helpful suggestions and comments. References Anquetin, J., Chapman, S.D., 2016. First report of Plesiochelys etalloni and Tropidemys langii from the Late Jurassic of the UK and the palaeobiogeography of plesiochelyid turtles. Royal Society Open Science 3, 150470. Anquetin, J., Deschamps, S., Claude, J., 2014. The rediscovery and redescription of the holotype of the Late Jurassic turtle Plesiochelys etalloni. PeerJ 2, e258. Arkhangelsky, M.S., Zverkov, N.G., 2014. On a new ichthyosaur of the genus Undorosaurus. Proceedings of the Zoological Institute RAS 318, 187–196. Bardet, N., Godefroit, P., Sciau, J., 1999. A new elasmosaurid plesiosaur from the Lower Jurassic of southern France. Palaeontology 42, 927–952. Batsch, A.J.G.C., 1788. Versuch einer Anleitung, zur Kenntniß und Geschichte der Thiere und Mineralien. Akademische Buchhandlung, Jena, Germany. Baur, G., 1888. Osteologische Notizen über Reptilien (Fortsetzung II). Zoologischer Anzeiger 11, 417–424. Benson, R.B.J., Bates, K.T., Johnson, M.R., Withers, P.J., 2011. Cranial anatomy of Thalassiodracon hawkinsii (Reptilia, Plesiosauria) from the Early Jurassic of Somerset, United Kingdom. Journal of Vertebrate Paleontology 31, 562–574. Benson, R.B.J., Butler, R.J., Lindgren, J., Smith, A.S., 2010. Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates. Proceedings Royal Society B 277, 829–834. Benson, R.B.J., Druckenmiller, P.S., 2013. Faunal turnover of marine tetrapods during the Jurassic–Cretaceous transition. Biological Reviews 89, 1–23. Benson, R.B.J., Evans, M., Smith, A.S., Sassoon, J., Moore-Faye, S., Ketchum, H.F., Forrest, R., 2013. A Giant Pliosaurid Skull from the Late Jurassic of England. PLoS ONE 8, e65989 doi:http://dx.doi.org/10.1371/journal.pone.0065989. _ Błazejowski, B., Gieszcz, P., Tyborowski, D., 2016. New finds of well preserved Tithonian (Late Jurassic) fossils from the Owadów-Brzezinki Quarry, Central Poland: a review and perspectives. Volumina Jurassica 14, 123–132. _ Błazejowski, B., Lambers, P., Gieszcz, P., Tyborowski, D., Binkowski, M., 2015. Late Jurassic jaw bones of halecomorph fish (Actinopterygii: Halecomorphi) studied with X-ray microcomputed tomography. Palaeontologia Electronica 18.3.52A, 1–10. Borsuk-Białynicka, M., Młynarski, M., 1968. The first finding of the Mesozoic marine turtle Tretosternon aff. punctatum Owen, 1848 in Poland. Prace Muzeum Ziemi 12, 217–222. Cope, E.D., 1869. Synopsis of the extinct Batrachia and Reptilia of North America, Part I. Transactions American Philadelphia Society New Series 14, 1–235.

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Westbury, Wiltshire. Philosophical Transactions of the Royal Society B: Biological Sciences 341, 399–418. Tyborowski, D., 2016. A new ophthalmosaurid ichthyosaur from the Late Jurassic of Owadów-Brzezinki Quarry, Poland. Acta Palaeontologica Polonica 61, 791–803. Tyborowski, D., 2017. Large predatory actinopterygian fishes from the Late Jurassic of Poland studied with X-ray microtomography. Neues Jahrbuch für Geologie und Paläontologie 283, 161–172. _ Tyborowski, D., Błazejowski, B., Krystek, M., 2016. Szcza˛tki gadów z górnojurajskich wapieni w kamieniołomie Owadów-Brzezinki (Polska srodkowa). Przegla˛d Geologiczny 64, 564–569 in Polish with English summary. Tyborowski, D., Skrzycki, P., Dec, M., 2018. Internal structure of ichthyosaur rostrum from the Upper Jurassic of Poland with comments on ecomorphological adaptations of ophthalmosaurid skull. Historical Biology doi:http://dx.doi.org/ 10.1080/08912963.2018.1559308.

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