Open Geosci. 2015; 7:342–361

Research Article Open Access

Cajus G. Diedrich* and Udo Scheer Marine vertebrates from the Santonian coastal carbonates of northwestern Germany – a tool for the reconstruction of a Proto- North Sea Basin intertidal dinosaur-exchange bridge

DOI 10.1515/geo-2015-0020 coastal and intertidal environments; upwelling; subma- Received December 10, 2013; accepted November 03, 2014 rine swell; northwestern Germany; southern Proto- North Sea Basin; Europe Abstract: A diverse vertebrate fauna, dominated by shark teeth, is recorded from conglomerates within the lime- stones of the Upper Cretaceous (Santonian) Burgstein- furt Formation of northwestern Germany. The conglomer- 1 Introduction ate beds comprise carbonatic, glauconitic and phosphate nodules, as well as Triassic, Jurassic and Cretaceous extr- In northwestern Germany, Upper Cretaceous shark teeth aclasts. The Burgsteinfurt Formation conglomerates con- have been used in the past to interpret the bathymetry, tain ning-upwards parasequences 2–20 cm in thickness, salinity and temperature of the southern Proto- North interpreted as tempestite layers within a unit formed by Sea Basin of central Europe [1–5]. In addition, sharks larger-scale Milankovitch Cycles. The presence of the in- from middle Eocene transgressive gravels (shark-teeth oceramid Sphenoceramus patootensis and belemnite Go- bonebeds) in the same region of northern Germany were nioteuthis granulata indicate a late Santonian age for the also analysed to further understand the uplift of a sub- unit. The studied vertebrate fauna from the Weiner Esch marine swell, called the Northwestphalian-Lippe Swell, locality consists of 20 selachian species (14 macroselachi- which began in the early Cenomanian [6]. This study ans and 6 microselachians), a few teleosts, rare marine presents the rst tectonic signs of the uplift of this swell, mosasaur remains, and one tooth from a theropod di- based on localities near Ochtrup in the northwestern- nosaur. 95% of the vertebrates in the assemblage are de- most part of the Münsterland Cretaceous Basin, where, positionally autochthonous, with the remaining material like much of this basin, Coniacian–Santonian sediments reworked from older underlying Cenomanian–Coniacian crop out extensively ([7, 8]; Figure 1). A new map of late (lower Upper Cretaceous) limestones. On the basis of ob- Santonian palaeogeography, presented here, diers from served sedimentary structures, the scarcity of deep-sea the previous ‘fully marine, subtidal Santonian’ palaeo- selachians, and the dominance of the Mitsukurinidae geographic models of the Cretaceous Münsterland Basin (59% of the preserved shark fauna) in the fossil assem- ([7, 9–12]; Figure 1). blage, the unit is interpreted as a shallow (0–3 metres The Late Cretaceous marine vertebrate locality dis- deep), subtidal, nearshore environment, or even subaerial cussed here (Figure 1) is a small, long-abandoned quarry carbonate-sand islands, located on the southern margin located on a hill called Weiner Esch, 5 km south of of a submarine swell. The presence of a Santonian thero- Ochtrup in northwestern-most Westphalia, northwestern pod in this deposit, and other dinosaur records in north- Germany (topographical map Metelen, coordinates lati- ern Germany, together support the interpretation of a tude 25° 80’50” S and longitude 57° 84’82” E). The quarry short-lived uplift event with strong upwelling inuence for is protected by a preservation order, as well as being a nat- the Northwestphalian-Lippe submarine swell north of the ural and palaeontological monument. Rhenish Massif in the southern Proto- North Sea Basin. A Previously, only rare, badly preserved, isolated new migration model for dinosaurs moving along carbon- selachian teeth have been recorded from Weiner Esch, ate coasts or intertidal zones of shallow carbonate-sand is- lands in Central Europe is presented, which may explain

the scattered distribution of dinosaur remains across Eu- *Corresponding Author: Cajus G. Diedrich: Private Research rope in the Upper Cretaceous. Institute PaleoLogic, Petra Bezruce 96, CZ-26751 Zdice, Czech Repub- lic, E-mail: [email protected] Keywords: Selachian fauna; teleosts; mosasaurs; thero- Udo Scheer: Ruhr Museum,Fritz-Schupp-Allee 15, D-45141 Essen, pod dinosaur; late Santonian (Late Cretaceous); nearshore Germany, E-mail: [email protected] © 2015 C.G. Diedrich and U. Scheer, licensee De Gruyter Open. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 License. The article is published with open access at www.degruyter.com. Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 343

Figure 1: A: Map showing the study area within central Europe. B–C: Geographic position of the Santonian vertebrate and dinosaur bone localities of northwestern Germany discussed in this paper (Map of Münster Cretaceous Basin after [7, 11]; location of vertebrate fossil sites after: 1. Aachen: [1]; Halle- Ascheloh: [37]; Höver: [4, 55]). 344 Ë C.G. Diedrich and U. Scheer and little of this material was described in detail [7–9, 13]. Institutional abbreviations: Geomuseum of the West- This publication records a well-preserved assemblage in- phalian Wilhelms-University Münster, Germany (GPI), cluding selachians, some teleosteans, and a few reptile Ruhr Museum Essen, Germany (RE). teeth with the aim of better understanding the shark, sh, and reptile biodiversity of the upwelling-inuenced up- per Santonian shallow-marine carbonate sediments of 3 Geology the northern Münsterland Cretaceous Basin, within the southern Proto- North Sea Basin. In this contribution, the 3.1 Stratigraphy and age rst Santonian dinosaur remain from northern Germany is presented. Together with yet unpublished nds from two other southern Proto- North Sea Basin localities – Weiner In [7] described the 8.5 m section of limestone at Weiner Esch, Lingen (Lower Saxony), and a new locality (con- Esch as ‘Weiner Schichten’, a unit now included within the taining dinosaur remains and a comparable shark fauna Burgsteinfurt Formation (upper Santonian – lower Campa- to Weiner Esch) north of the Harz Mountains (Lower Sax- nian) with Weiner Esch as its stratotype [11]. Due to quarry ony), these dinosaur nds are important to the improving backlling prior to the site’s protection, only ve metres of the palaeogeographic map of Ziegler 1990 [12]. It allows of these sediments are still exposed (Figure 2). This sec- the interpretation of coastline and island positions, and tion is informally subdivided herein into ‘Lower Conglom- therefore the reconstruction of the swell’s uplift history. erate Beds’ and ‘Upper Sandy Limestone Beds’. At this locality, the Burgsteinfurt Formation discordantly over- lies the older lower Santonian Emscher Formation [8] due to a tectonic uplift that created a submarine swell (the Ochtrup Swell) as the northeastern prolongation of the 2 Material and methods Dutch High during the Santonian [8, 10]. During this time, Upper Triassic, Jurassic and Lower Cretaceous sediments A small collection of material from Weiner Esch was became inverted and eroded during an alpinotype subher- made by K. Niemeyer between 1966 and 1975, and his few cynic tectonic period (Wernigeröde Phase sensu [7]), form- selachian teeth are now deposited in the Geomuseum of ing the Northwestphalian-Lippe Swell. With its parallel the Westphalian Wilhelms-University Münster, Germany. mountain ranges – Wiehengebirge and Teutoburger Wald A further small, but important collection, including the – this former swell separates the Münsterland Basin from single fragment of a dinosaur tooth as well as the only the rest of the North German lowland. This tectonic event relict of a chimaera, was later made by E. Wijnker (Nether- explains the presence of conglomerates, extraclasts, and lands), which was donated to the Ruhr Museum (Essen, even the reworked shark teeth seen within our material Germany) and is housed together with the huge collec- (see Taphonomy section). Most vertebrate remains (95%) tion of K.-H. Hilpert. In addition to these three private seem to be autochthonous within the sediment, an inter- collections, the current authors etched three kilograms of pretation followed by other authors [7, 8], although some sediment collected from Conglomerate Bed II (Figure 2) of the selachian species from Weiner Esch noted in this with formic acid to obtain additional microselachians, pla- paper obviously originate from reworked Cenomanian– coid denticles and teleostean teeth. Of this material, small- Coniacian, or even middle Santonian layers (Table 1). A sized fossils were photographed with a REM, and larger comparable taphonomic situation is found in the Eocene teeth photographed using a binocular microscope, with all shark-gravels of the Fürstenau Formation (Lower Sax- drawings produced using a binocular mirror (Figures 3–7). ony, about 80 km northeast of Weiner Esch) of north- As the terrestrial reptile material (Figure 8) has proved im- ern Germany, which also contains reworked Cenomanian– portant for interpreting palaeogeography (Figure 9), two Turonian species [14]. more sites from northern Germany are mentioned herein, The sediments in Weiner Esch are considered late although the dinosaur material from these localities awaits Santonian in age (= krsa4; [7, 8]) based on the presence detailed investigation and is as yet unpublished. This un- of the inoceramid Sphenoceramus patootensis and cal- published material includes fossils from a site north of the citic rostrae of the belemnites Gonioteuthis granulata and Harz Mountains, stored in a private collection; and a col- Actinocamax verus. These index taxa were collected by lection from Lingen, deposited in the Museum of Natural the present authors from a number of dierent conglomer- History Oldenburg (Staatliches Museum für Naturkunde ate beds (especially from Conglomerate Bed II; Figure 2), Oldenburg). again supporting the traditional interpretation of this unit Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 345

Figure 2: Stratigraphy of the Burgsteinfurt Formation (upper Santonian) exposed at the Weiner Esch vertebrate locality, located near Ochtrup in northwestern Germany (Chronostratigraphic data after [11]). Shark teeth and other macrofaunal remains are preserved at the bases (= conglomerates) of several parasequence sets (storm deposits I–XVIII). 346 Ë C.G. Diedrich and U. Scheer as late Santonian in age. Unfortunately, other more bios- onian) limestone pebbles. Jurassic ammonite fragments tratigraphically diagnostic taxa, such as ammonites, are have been found within these pebbles (body chambers missing from this unit. preserved in sideritic nodules; [8]) or as limonitized frag- ments. The size of these extraclasts generally does not ex- ceed one centimetre in diameter. The sand fraction of the 3.2 Sedimentology conglomerates, mainly formed of carbonate particles and marine invertebrate remains, additionally contains quartz The sedimentary section at Weiner Esch is dominated by and glauconite grains. Especially together with the nu- arenitic limestones containing 85-95% calcium carbon- merous phosphorite nodules seen in these conglomerates, ate [7, 8]. Calcite shell producers, such as the lamelli- they indicate the inuence of upwelling on the deposi- branchs Sphenoceramus and Pycnodonte, plus crinoids, tional environment [7, 8]. belemnites, bryozoans and balanids, are present in the majority of bioclasts. Of this group, crinoid remains dominate (50%), followed by bryozoans (30%; see also 4 Paleontology [7]), belemnites, echinoids, oysters, balanids, and in- oceramids. These fossils are often rounded, sorted ac- cording to grain size, and are seldom greater than 4.1 Faunal taphonomy four centimetres in length, although belemnites and in- oceramids are occasionally larger and sometimes com- At Weiner Esch, the vertebrate fauna is particularly rich pletely preserved. The microfauna is dominated by sand- in Conglomerate Beds I and II but fossils are found in all agglutinating foraminifers [8]. As a result of the late dia- conglomeratic layers (Figure 2). Only teeth and small bone genetic dissolution of aragonite, no gastropods and only a fragments are present in this assemblage due to the sort- few other lamellibranchs are preserved in the assemblage. ing eects of palaeocurrents, with selachians mainly rep- The middle part of the section of the Burgsteinfurt resented by teeth and few dermal scales. Based on the Formation (Figure 2) visible at Weiner Esch contains at preservation of sharp cutting edges (especially nearly all least 18 thin ning-upward sets (Figure 2), with vertebrate Odontaspidae) on about 95% of the selachian teeth, it remains and invertebrate marine fossils remarkably en- seems clear that these teeth were not transported over long riched at the base (2-10 cm) of each of these sets. The Con- distances or extensively reworked. The roots of these au- glomerate Beds II and III with their high density of extr- tochthonous teeth are very soft, due to late diagenesis, aclasts and fossil remains possibly represent ‘large-scale’ and therefore are mostly removed in Scapanorhynchus and Milankovitch sequence boundaries (Figure 2). Fossilifer- other selachian taxa. In contrast, there is a small num- ous beds are typical for such boundaries in carbonate sys- ber of allochthonous teeth in the assemblage, identied tems (e.g. [6]), where each parasequence might represent based on their rounded cusps and roots (Table 1). Having coastal storm events, compared to other shallow-marine passed through diagenesis prior to the deposition of the carbonate storm deposits with similar ning-upward sets sediments at Weiner Esch, the roots of the allochthonous (e.g. [15]). The nearshore shallow-coastal subtidal facies of teeth are ‘prefossilized’ and often incompletely preserved. the submarine Ochtrup Swell, the western adjacent part or prolongation of the Northwestphalian-Lippe Swell, was in- uenced by storm sediments transported across the swell 4.2 Vertebrate biodiversity and assemblage to the levee (i.e. the southeastern slope) from the north- composition west and north, as indicated by the orientation of shal- low subtidal channels and cross bedding within this fa- Listings of the diverse marine-invertebrate and vertebrate cies [8]. Gutter-casts and cross-bedded carbonate sands, macrofauna from Weiner Esch have been previously pub- or channels, lled with fossiliferous conglomerates, char- lished [3, 7, 8], although these lists typically used older acterize the shallowmarine coastal conditions around this ‘morphotype systematics’, not easy to correlate with sev- swell [7, 8]. eral recent faunal revisions. The extremely confusing The conglomerates are predominantly composed of nomenclature of shark teeth is a particular problem that slightly to well-rounded extraclasts of Triassic, Jurassic, needed to be revised in the future. For example, the tooth and Lower Cretaceous marls and dolomites, dark brown morphotaxon Cretoxyrhina is now known to belong to the claystones, asphaltites, red sandstones and claystones, genus Isurus [16], which was described using skeletal re- but also includes smaller, rounded Upper Cretaceous (Tur- mains more than just teeth. Similar revisions have recently Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 347

Figure 3: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Paraorthacodus recurvus, anterior tooth, a–b. labial, RE No. 3786/10. 2. Paraorthacodus recurvus, posterior lateral tooth, a, c. lateral, b. occlusal, RE No. A 3782. 3. Ptychodus cf. polygyrus, tooth, a. lateral, b–c. occlusal, GPI No. A1C-2-1 (from reworked Cenomanian layers). 4. Hexanchus microdon, lateral tooth of the lower jaw, a–b. lingual, RE No. A 3789/2. 5. Hexanchus microdon, lateral tooth of the upper jaw, a–b. lingual, RE No.A 3789/2. 6. Protosqualus cf. sigei, tooth crown, labial, RE No. A 4813/1. 7. Heterodontus cf. rugosus, anterior tooth, a–b. labial, RE No. A 4810/1; 8. Heterodontus rugosus, anterolateral tooth, a–b. lateral, c. occlusal, RE No. A 3780/2. 9. Chiloscyllium cf. greeni, anterior tooth, labial, RE No. A 4814. 10. Scyliorhinus elongatus, lateral tooth, a–b. lingual, c. lateral, d. labial, RE No. A 4811. 11. Scyliorhinus cf. bloti, a–b. labial, RE No. A4815. 12. Scyliorhinus (or Crassescyliorhinus) a–b. labial, RE No. A4816. 13. Pteroscyllium sp., tooth crown, a. lingual, b–c. labial, RE No. A 4812/1. 348 Ë C.G. Diedrich and U. Scheer Cenomanian–Campanian Coniacian–Maastrichtian (Reworked from Aptian–Cenomanian) Cenomanian–Campanian strata) Cenomanian–Maastrichtian 209 (15 anterior,cusps) 12 lateral teeth, 182 44seal/anteriolateral, (3 20and 20 crowns) lateral anterior, teeth, one1 lateral tooth symphy- tooth crown) Cenomanian–Campanian teeth, 6 cusps) 2 (1 anterior and 1 lateral tooth) Coniacian–Campanian teeth and fragmentscusps) and some tooth (Her- (Agassiz, (Trautschold, (Cope, 1875) 2 (1 anterior and 1 lateral tooth) (Reworked from Albian–Cenomanian) (Cappetta, 1973) 1 anterior tooth lower Cenomanian–Campanian Agassiz, 1835 2 (1 rootless and 1 tooth fragment) (Reworked from upper Cenomanian (Münster in Agas- (Agassiz, 1843) 7 (2 anterior and 5 lateral teeth) Coniacian–Campanian Cappetta, 1977 3 incomplete teeth Cenomanian–Campanian (Agassiz, 1843) 15 (3 lateral, 4 anterolateral, 8 lateral (Cappetta, 1980) 3 (1 anterolateral, one lateral, and one (Agassiz, 1844) 37 (17 anterior, 4 anterolateral, 10 lateral sigei greenei (Woodward, 1894) 5 (1 anteriolateral and 4 lateral teeth) Cenomanian–Campanian (Agassiz, 1843) 30 (1 anterior, 2 lateral teeth, 27 cusps) Coniacian–Campanian polygurus cf. cf. cf. Megarhizodon macrorhiza Isurus mantelli Lamna appendiculata pachyrhiza man, 1975) Lamna arcuata Cretodus semiplicatus siz, 1843) Protolamna sokolovi Scapanorhynchus raphiodon 1844) Hexanchus microdon Protosqualus Heterodontus rugosus Chiloscyllium Carcharias subulata Ptychodus SpeciesSelachii Paraorthacodus recurvus 1877) Material Age range Vertebrate teeth recovered from conglomerates within the upper Santonian (Upper Cretaceous) Sandy Sandy limestones of Weiner Esch, near Ochtrup, northwestern Germany (all Table 1: material is stored at the Ruhr Museum Essen, Germany). Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 349 8 (1 anterior and 7 lateral teeth) Cenomanian–Maastrichtian ) 3 tooth fragments Santonian–Maastrichtian sp. 1 tooth (Reworked from Cenomanian) (Reuss, (Agassiz, 1838) 2 lateral teeth Campanian–Maastrichtian Case, 1978 1 jaw fragment Santonian–Maastrichtian (Davis, 1887) 1 lateral tooth of the upper jaw Santonian Leiodon/Clidastes Cappetta, 1980 1 tooth (Reworked from Cenomanian) Crassescyliorhinus (Agassiz, 1835) 42 (2 anterior, one lateral teeth, 39 cusps) Coniacian–Campanian Müller & Diedrich, 1999 2 anterior teeth Cenomanian–Santonian bloti sp. 3 teeth crowns Cenomanian–Santonian bifurcatus cf. sp. or cf. sp. 7 teeth Cretaceous sp. 1 tooth Cretaceous Table 1 cnd. Pteroscyllium Chimaeriformes Ischyodus Teleostei Pycnodontidae indet.Coelodus Enchodus Reptilia Mosasauridae indet. ( Tyrannosauridae indet. 1 tooth 1 tooth fragment Cretaceous Santonian–Maastrichtian Table 1: SpeciesProtolamna acuta Paranomotodon angustidens 1845) Palaeohypotodus bronni Squalicorax kaupi Scyliorhinus elongatus Scyliorhinus Scyliorhinus Material Age range 350 Ë C.G. Diedrich and U. Scheer

Figure 4: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Carcharias subulata, anterior tooth of the upper jaw, a–b. labial, RE No. A 3790/1. 2. Carcharias subulata, anterolateral tooth of the upper jaw, a–b. labial, RE No. A 3786/01. 3. Carcharias subulata, lateral tooth of the upper jaw, a–b. labial, RE No. A 3786/06. 4. Scapanorhynchus raphiodon, anterior tooth, a–b. labial, RE No. A 3784/09. 5. Scapanorhynchus raphiodon, anterior tooth, a–b. labial, RE No. A 3784/08. 6. Scapanorhynchus raphiodon, anterolateral tooth, a–b. labial, RE No. A 3784/01. 7. Scapanorhynchus raphiodon, lateral tooth, a–b. labial, RE No. A 3779/05. 8. Scapanorhynchus raphiodon, tooth, lingual with typical striation, RE No. A 3784/12. Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 351

Figure 5: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Isurus mantelli, anterior tooth, a– b. labial, c. lateral, d. lingual, RE No. A 3868. 2. Isurus mantelli, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3772/1. 3. Isurus mantelli, lateral tooth, labial, RE No. A 6530. 352 Ë C.G. Diedrich and U. Scheer

Figure 6: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Lamna appendiculata pachyrhiza, anterior tooth, a–b. lingual, RE No. A 3779/02. 2. Lamna appendiculata pachyrhiza, symphyseal or anteriolateral tooth, a–b. labial, RE No. A 3774/01. 3. Lamna appendiculata pachyrhiza, anterior tooth, a. lateral, b. labial, c. lingual, RE No. A 3774/10. 4. Lamna appendiculata pachyrhiza, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3778/1. Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 353 been published for the white (Carcharodon) and mega- scriptions in [48–51]), whereas dinosaurs are recorded by tooth sharks (Otodus: see [5]). As these shark species are a single tooth fragment belonging to a theropod (Tyran- already well described in the literature, none of the nds nosauridae) [52] (Figure 8.7). from Weiner Esch are described herein again (see [2, 5, 16– 44]). Shark scales remain the most dicult fossils from Weiner Esch to identify, even to genus level [37], due to the 5 Discussion changing morphology of scales over dierent body regions [34]. Teeth of teleosts, mosasaurs theropods are also di- 5.1 Comparison of Mesozoic shark cult to identify to genus level as a result of poor preserva- tion, fragmentation, or a lack of characteristic features. assemblages in northwestern Germany According to the references mentioned above, 20 dierent selachian taxa (14 macroselachians and 6 mi- The relatively diverse (20 species) shark assemblage pre- croselachians) have been identied from the Weiner Esch served at Weiner Esch demonstrates some similarities to locality of the Burgsteinfurt Formation (upper Santo- the fauna of 38 early Cenomanian shark species recorded nian). Especially Scapanorhynchus teeth could be placed at Ascheloh near Halle/Westphalia, in the Teutoburger within the jaw according to associated tooth sets from Wald Mountains ([37]; Figure 9). At both sites, the number America [45], whereas for other species modern shark of macroselachian species is comparable, with 12 species dentitions were compared [46]. The investigated assem- recorded at Halle-Ascheloh and 14 at Weiner Esch (Fig- blage consisted of 599 determinable (Table 1, Figure 9) ure 9). At Halle-Ascheloh, the greater abundance of mi- and 135 non-determinable selachian teeth. The assem- croselachians allowed greater amounts of research on this blage comprises 59% Scapanorhynchus raphiodon (Agas- group, accounting for the increased diversity (26 species) siz, 1843) (Figures 4.4–8), 8% Carcharias subulata (Agas- in comparison to the microselachians of Weiner Esch. Fau- siz, 1844) (Figures 4.1–3), 7% Squalicorax kaupi (Agas- nal mixing seen at both sites seems to be related to tapho- siz, 1835) (Figure 7.10), 6% Isurus mantelli (Agassiz, 1843) nomic or sedimentary processes, specically slumping at (Figures 5.1–3), 5% Lamna appendiculata pachyrhiza (Her- Halle-Ascheloh, and sedimentary condensation at Weiner man, 1975) (Figures 6.1–4), 4% Hexanchus microdon (Agas- Esch. Reworked teeth are present in the Weiner Esch mate- siz, 1835) (Figures 3.4–5), 2% Paranomotodon angusti- rial originating mainly from Cenomanian–Turonian strata dens (Reuss, 1845) (Figure 7.7), 1% Lamna arcuata (Wood- of northwestern Germany (Figure 9; Tables 1-2). ward, 1894) (Figure 7.1), and 1% Heterodontus cf. rugosus The absence of Hemipristidae or Rhinobatidae, and (Agassiz, 1839) (Figure 3.7–8). All other recorded species dominance of Mitsukurinidae (goblin sharks), especially each represent less than 1% of the assemblage: Proto- Scapanorhynchus raphiodon (59% of the shark fauna), and squalus cf. sigei Cappetta, 1977 (Figure 3.6), Chiloscyl- lamnids in the Weiner Esch fauna generally indicates the lium cf. greenei (Cappetta, 1973) (Figure 3.9), Paraorthaco- inuence of boreal and shallow-marine upwelling condi- dus recurvus (Trautschold, 1877) (Figures 3.1–2), Megarhi- tions (Table 1, Figure 9). Therefore, the assemblage shows zodon macrorhiza (Cope, 1875) (Figures 7.3–4), Ptychodus palaeoecological similarities to the early Cenomanian cf. polygyrus Agassiz, 1835 (Figure 3.3), Cretodus semi- carbonate-sand shark fauna of HalleAscheloh, where the plicatus (Münster in Agassiz, 1843) (Figure 7.2), Proto- Mitsukurinidae also dominate the macroselachians (Fig- lamna sokolovi (Cappetta, 1980) (Figures 7.5–6), Proto- ures 1, 9; [37]). In contrast to the living deep-water shark lamna acuta [37], Palaeohypotodus bronni (Agassiz, 1843) Mitsukurina [53], it seems that Cretaceous Mitsukurinidae (Figures 7.8–9), Scyliorhinus elongatus (Davis, 1887) (Fig- were associated with cold-water and upwelling conditions ure 3.10), Scyliorhinus cf. bloti Cappetta, 1980 (Figure 3.11), in shelfal regions e.g. the Western Interior Seaway [42] Scyliorhinus sp. or Crassescyliorhinus sp. (Figure 3-12). and the Proto- North Sea Basin; both areas under north- Pteroscyllium sp. (Figure 3.13). Chimerae are represented ern boreal upwelling inuence. Like other shark groups using illustrations of [47] by a single jaw identied as Is- that have changed life habits during their evolution, the chyodus cf. bifurcatus Case, 1978 (Figure 8.1). Bony shes same is interpreted here for the Mitsukurinidae. Repre- are only represented by one tooth of Pycnodontidae indet. sentatives of this family with lateral cusp development (Figures 8.3), and one Coelodus sp. (Figure 8-2) and few En- reduction, serration, or changes in tooth formula within chodus sp. (Figure 8.4). the jaws - are already evident in the early Late Cretaceous Marine reptiles from this locality are repre- [5, 16, 29, 44]. The late Santonian Weiner Esch selachian sented by teeth fragments of Mosasauridae indet. fauna is most similar to the early Campanian greensand (Leiodon/Clidastes) (Figures 8.5–6) (as per the genus de- fauna of the Vaals Formation in the Aachen area ([1], Ta- 354 Ë C.G. Diedrich and U. Scheer

Figure 7: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Lamna arcuata (Woodward), lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3777/1. 2. Cretodus semiplicatus, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3779/13. 3. Megarhizodon macrorhiza, anterior tooth, a–b. labial, RE No. A 3802/07. 4. Megarhizodon macrorhiza, lateral tooth, a–b. labial, A 3847/10. 5. Protolamna sokolovi, anterolateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3778/02. 6. Protolamna sokolovi, anterior or lateral tooth, a–b. labial, RE No. A 3792/7. 7. Paranomotodon angustidens, lateral tooth, a–b. labial, RE No. A 3773/2. 8. Palaeo- hypotodus bronni, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3779/01. 9. Palaeohypotodus bronni, lateral tooth, a–b. labial, c. lingual, d. lateral, RE No. A 3804. 10. Squalicorax kaupi, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3774/05. Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 355 ble 1) near the German–Belgian–Netherlands border. The lower Cenomanian Halle-Ascheloh site ([37]; Figure 9). In Aachen selachian fauna contains a number of species sim- the Eocene localities, sand sharks are even higher in pro- ilar to Weiner Esch (Table 2), and was also deposited in a portion than in the Santonian fauna (Figure 9), which ts shallow nearshore marine environment, as indicated by its the interpreted palaeogeography of very shallow marine lithology (glauconitic carbonate sands; [1]). However, mi- conditions along the southern margin of the Proto- North croselachians in the Aachen assemblage were only studied Sea Basin during this time [14]. using small samples, and percentages of individual shark species were not quoted, thereby hampering comparisons to other northwest German shark assemblages. 5.2 Uplift reconstructions of the submarine Conversely, a number of open marine early and late Northwestphalian-Lippe swell Campanian vertebrate and selachian faunas from Ger- many can be seen to be considerably dierent from the The Cenomanian Halle-Ascheloh site, concentration of assemblage at Weiner Esch [3]. The only similarities are vertebrate remains resulted from marine slumping along the presence of deep-sea or cool-water species at Weiner the margin of a submarine swell and elongated carbon- Esch, compared with the Squalidae (Protosqualus) that ate platform, the Northwestphalian-Lippe Swell, located are particularly well represented in the Late Cretaceous of at the northern margin of the Münsterland Cretaceous northern and northwestern Europe (e.g. [2, 37]; Table 2). Basin (Figure 9, [6]). The uplift of this feature began dur- The microselachian genera Chiloscyllium and Scyliorhinus ing the early Cenomanian [54], rst causing large-scale recorded at Weiner Esch are also present in these Campa- slumping on the swell’s southern slopes (Figures 1, 9). In nian deposits, although represented by dierent species, the middle late Cenomanian, the carbonate platform also all of which indicate the inuence of subtropical, warm changed tectonically along its southern margin, with con- surface-water from Tethys in the environment [27, 28]. An tinuous high subsidence, reaching the deep-water con- early Campanian selachian fauna (12 species) collected ditions required for the deposition of shark and sh- from marly limestones at Höver, near Hannover (Lower rich black shales during the second Cretaceous Oceanic Saxony, northern Germany), diers from the Weiner Esch Anoxic Event (= OAE II) in the Osning Depression (Fig- assemblage particularly in the microselachians [4]. This ure 9; [54, 55]). The hercynic-controlled tectonic uplift is dierence seems to be a result of the palaeoenvironment, best exposed at Halle and Bielefeld (Ostwestfalendamm which at Höver was an open marine facies with more or site), both of which lie along the southern slope of the less continuous sedimentation, lacking sedimentary con- submarine swell, where upper Turonian – lower Conia- densation or vertebrate bone-bed formation, and therefore cian units contain large carbonate ‘phacoids’ of slumped reducing the likelihood of nding teeth. Among the low- Cenomanian–Turonian consolidated carbonates up to sev- abundance shark material recovered from Höver, the dom- eral meters in size [56, 57]. It is demonstrated here that inant squalomorph sharks (six neritic and pelagic species, the swell regions (Ochtrup Swell, and others, Figure 9) including Cretascymnus westphalicus and Centroscymnus must have already been uplifted to subaerial or shallow praecursor) are considered deep-water selachians [3] liv- intertidal conditions (i.e. carbonate-sand islands) during ing in a deep basin, agreeing with the interpreted palaeo- the late Santonian, thereby separating the Münsterland geography of the ProtoNorth Sea Basin at this time (Fig- Cretaceous Basin (Figure 9) from the Proto- North Sea ure 9, and [12]). Basin. Similar in age to the hercynian uplift event are the Comparison of the Weiner Esch assemblage to the northern German Harz Mountains, of which its earliest ‘is- sharks collected from the Middle Eocene Fürstenau For- land’ stage has been previously dated as middle Santonian mation from Bippen and Dalum (Lower Saxony) was [58, 59]. also considered important, as a mixing of reworked Cenomanian–Turonian shark teeth within a predomi- nantly autochthonous fauna was also observed in this 5.3 Dinosaur migration along intertidal latter formation (Figures 1, 9; [14, 44]). The Weiner Esch zones or islands in northern Germany (Santonian) and Bippen (Eocene) assemblages have com- parably high percentages of sand sharks (Odontaspidae), [9] interpreted the sediments at Weiner Esch and the and similarities in the upwelling inuence within the nearby Seller Esch site (no longer exposed) as ‘coastal sed- fauna [14]. Therefore, it is suggested that an abundance of iments’ and was the rst to propose the existence of ‘is- sand sharks is a good marker for identifying very shallow, lands’ in northwestern Germany during the Late Creta- coastal marine palaeoenvironments, as also seen in the ceous. In the [12] palaeogeographic maps of Europe, inter- 356 Ë C.G. Diedrich and U. Scheer

Figure 8: Bony shes chimaeras, and marine and terrestrial reptiles from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Ischyodus cf. bifurcatus, RE No. A 6527. 2. Coelodus sp., tooth, occlusal, RE No. A 3894/1. 3. Pycnodontidae indet., tooth, a–b. occlusal, RE No. A 3894/2. 4. Enchodus sp., tooth, a–b. lateral, RE No. A 3788. 5. Mosasaur (Leiodon/Clidastes) tooth fragment, a. lateral. b. dorsal, c. lingual, RE No. A 3776; 6. Mosasaur (Leiodon/Clidastes) tooth fragment, labial, RE No. A 3813. 7. Theropod (Tyrannosauridae) dinosaur tooth fragment, a. lateral, b. lingual/labial, RE No. A 6528.

Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 357

eurytherm

boreal

temperate

rpcl subtroppical tropical/

pelagic

neritic

nectic

benthic

Maastrichtian

Campanian

Santonian

Coniacanian

Turonian

Cenomanian

ot Africa North

ot America North

oteEurope Northe

anvr Höver Hannover-

enrEsch Weiner

Coesfeld Aachen (Herman) x x x x x x x – – – – x x x (Reuss) x x x x x – – – – x x x Agassiz x x x x x – – – x x x x (Cope) x x x x Capetta x x x x – – – x x x (Trautschold) x x x x – – x x x x (Agassiz) x x – – – x x x Capetta x x x x x x x – – – – x x x (Münster) x x x x x – – – – x x x Agassiz x x x – – x x x Capetta x x x – – – – x x x x (Woodward) x x x – – – – x x x Agassiz x x x x – – x x x x (Agassiz) x x x x x – – – – x x x Capetta x x x x – – – x x x Capetta x x – – – x x x (Agassiz) x x x – – – x x x x (Agassiz) x x x x x – – – x x x Müller & Diedrich x x x – – – x x x x greenei sigei elongatus bloti (Agassiz) x x x x x – – – x x x sp. x x x x – – – – x x x cf. cf. cf. cf. Comparison of the Late Cretaceous selachian faunas of Germany, central Europe, with the global Cretaceous selachian record, and the palaeoecological preferences of known taxa. Paraorthacodus recurvus Ptychodus polygurus Hexanchus microdon Protosqualus Heterodontus rugosus Protolamna acuta Paranomotodon angustidens Palaeohypotodus bronni Squalicorax kaupi Scyliorhinus Chiloscyllium Protolamna sokolovi Table 2: TaxonSelachii Carcharias Palaeogeography subulata Isurus mantelli Lamna appendiculata pachyrhiza Plicatolamna arcuata Cretodus semiplicatus Age BathymetryScyliorhinus Pteroscyllium Climate Megarhizodon macrorhiza Scapanorhynchus raphiodon 358 Ë C.G. Diedrich and U. Scheer tidal zones were not delineated at all for the Santonian. For on the Ochtrup and Northwestphalian-Lippe Swells that the new interpretation presented herein, the shark faunas connected the Dutch High with the uplifting Harz Moun- and position of three dinosaur localities (Figure 9) were tains in the southern Proto- North Sea Basin of central used to interpret a preliminary new palaeogeographic map Europe. This swell started to uplift in the early Cenoma- of the Münster Cretaceous Basin and its northern realm in nian, and later formed a back-arc-like depression (the Os- the late Santonian. It seems, based on Santonian dinosaur ning Depression), which was lled with vertebrate-rich remains at Weiner Esch (tooth gured herein), Lingen, and black shale layers in the late Cenomanian – early Turo- north of the Harz Mountains (represented by as yeat un- nian. Tectonic activities continued in this region in the described postcranial longbones and fragments) in north- late Turonian – early Coniacian with slumping of larger ern Germany (Figures 1, 9), that the uplifted submarine phacoids along the swell slopes, nally forming subaerial swell regions of northern Germany and the Netherlands carbonate-sand intertidal or island conditions in the late (the Dutch High, Ochtrup High and Northwestphalian- Santonian. In the Eocene, the Northwestphalian-Lippe Lippe High) must have been periodically intertidal or even Swell was uplifted to form a peninsula or islands, initi- shallow carbonate-sand islands. ating the formation of the Wiehengebirge / Teutoburger The idea of dinosaurs using beaches and intertidal Wald Mountains north of the Rhenish Massif. The main areas as possible seasonal migration zones is becoming vertebrate material obtained from Weiner Esch consisted more favoured, with many new discoveries of dinosaur of marine shark teeth (20 species), that together indicate tracks recorded in European Middle Triassic [60, 61] or the inuence of cold, upwelling (based on the presence Upper Jurassic [62] coastal or intertidal beach deposits, of glauconite and phosphorite nodules), boreal water dur- and examples such as recently described Cenomanian– ing the late Santonian, with periodic, seasonally warm Turonian trackways on the Adriatic Carbonate Platform surface-water inuences derived from Tethys. The high and in central Europe [63]. Only such intertidal ‘bridges’ percentage of sand sharks (59%), especially S. raphiodon can explain the distribution of Santonian dinosaur records (Mitsukurinidae), seen in this assemblage is considered along European Cretaceous islands or peninsula coasts. a result of this cold water and the upwelling inuence, Within Europe, the Santonian dinosaur remains ge- plus the concentration of shelfal shark teeth in a coastal ographically closest to Weiner Esch are described from carbonate-sand facies. The bathymetry of the swell region west and central France (theropod teeth and a few bone during the late Santonian is estimated between intertidal fragments from Charente-Maritime, Maine-et-Loire, Indre- to shallow subtidal (greater than 0 m water depth, and et-Loire, Vendee: [52]) and from Hungary (a few herbivo- 0–5 m along the swell margins). Additionally, rare thero- rous dinosaur teeth (Rhabdodontidae indet., Nodosauri- pod remains recovered from this site, along with other di- dae indet.) and theropod remians of uncertain relation- nosaur bones collected from two more sites on and along ship from Iharkút, Bakony Mountains: [64]. These rare, the submarine swell, indicate the seasonal migration or scattered remains indicate a widespread distribution of di- faunal exchange of dinosaurs along coasts or intertidal nosaurs across Europe in the Santonian, supporting the zones, likely between islands, of Europe in the Late Cre- idea that at this time the area contained several islands taceous. and small landmasses ([12], Figure 9).

Acknowledgement: The authors are very grateful to K.- H. Hilpert, Dr. K. Niemeyer and E. Wijnker, who collected 6 Conclusions most of the material described herein. Dr. M. Bertling is thanked for providing permission to study the material The late Santonian marine vertebrate fauna of Weiner stored in the Geomuseum of the Westphalian Wilhelms- Esch, exposed near Ochtrup in northern Germany, pro- University Münster, Germany. Sta at PaleoLogic pro- vides important insights into the palaeogeography and duced all teeth drawings, illustrations and other graphic history of the submarine Northwestphalian-Lippe Swell, work. Museological work on the material was sponsored thereby helping to ll a gap in knowledge of this region’s by the Ruhr Museum, and PaleoLogic sponsored the re- evolution during the Late Cretaceous. Rare dinosaur re- search. Finally, we thank M. Reinhardt for preparation of mains recovered from three localities along the swell, sup- the binocular photos, and Dr. H.-M. Weber for supplying port the interpretation of an already uplifted environment us with REM photos. Marine vertebrates from the Santonian coastal carbonates of northwestern Germany Ë 359

Figure 9: Palaeogeographic development (bathymetry and facies) of the Northwestphalian-Lippe Swell based on vertebrate faunas, espe- cially the frequency of shallow-marine sand and goblin sharks (Odontaspidae, Mitsukurinidae), for the A. Lower Cenomanian (based on [6, 37, 55]); B. Upper Santonian (data described herein); and C. Middle Eocene (based on [55]). From this swell, the present Teutoburger Wald and Wiehengebirge mountain ranges developed in northern Germany from the southern Proto- North Sea Basin . 360 Ë C.G. Diedrich and U. Scheer

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