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© 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.

Open Geosci. 2015; 7:342–361

Research Article Open Access

Cajus G. Diedrich* and Udo ScheerMarine vertebrates from the Santonian coastal carbonates ofnorthwestern Germany – a tool for the reconstruction of a Proto-North Sea Basin intertidal dinosaur-exchange bridgeDOI 10.1515/geo-2015-0020

Received December 10, 2013; accepted November 03, 2014

Abstract: A diverse vertebrate fauna, dominated by sharkteeth, is recorded from conglomerates within the lime-stones of the Upper Cretaceous (Santonian) Burgstein-furt Formation of northwestern Germany. The conglomer-ate beds comprise carbonatic, glauconitic and phosphatenodules, as well as Triassic, Jurassic and Cretaceous extr-aclasts. The Burgsteinfurt Formation conglomerates con-tain �ning-upwards parasequences 2–20 cm in thickness,interpreted as tempestite layers within a unit formed bylarger-scale Milankovitch Cycles. The presence of the in-oceramid Sphenoceramus patootensis and belemnite Go-nioteuthis granulata indicate a late Santonian age for theunit. The studied vertebrate fauna from the Weiner Eschlocality consists of 20 selachian species (14 macroselachi-ans and 6 microselachians), a few teleosts, rare marinemosasaur remains, and one tooth from a theropod di-nosaur. 95% of the vertebrates in the assemblage are de-positionally autochthonous, with the remaining materialreworked from older underlying Cenomanian–Coniacian(lower Upper Cretaceous) limestones. On the basis of ob-served sedimentary structures, the scarcity of deep-seaselachians, and the dominance of the Mitsukurinidae(59% of the preserved shark fauna) in the fossil assem-blage, the unit is interpreted as a shallow (0–3 metresdeep), subtidal, nearshore environment, or even subaerialcarbonate-sand islands, located on the southern marginof a submarine swell. The presence of a Santonian thero-pod in this deposit, and other dinosaur records in north-ern Germany, together support the interpretation of ashort-lived uplift event with strong upwelling in�uence forthe Northwestphalian-Lippe submarine swell north of theRhenish Massif in the southern Proto- North Sea Basin. Anewmigration model for dinosaurs moving along carbon-ate coasts or intertidal zones of shallow carbonate-sand is-lands in Central Europe is presented, which may explainthe scattered distribution of dinosaur remains across Eu-rope in the Upper Cretaceous.

Keywords: Selachian fauna; teleosts; mosasaurs; thero-pod dinosaur; late Santonian (Late Cretaceous); nearshore

coastal and intertidal environments; upwelling; subma-rine swell; northwestern Germany; southern Proto- NorthSea Basin; Europe

1 IntroductionIn northwestern Germany, Upper Cretaceous shark teethhave been used in the past to interpret the bathymetry,salinity and temperature of the southern Proto- NorthSea Basin of central Europe [1–5]. In addition, sharksfrom middle Eocene transgressive gravels (shark-teethbonebeds) in the same region of northern Germany werealso analysed to further understand the uplift of a sub-marine swell, called the Northwestphalian-Lippe Swell,which began in the early Cenomanian [6]. This studypresents the �rst tectonic signs of the uplift of this swell,based on localities near Ochtrup in the northwestern-most part of the Münsterland Cretaceous Basin, where,like much of this basin, Coniacian–Santonian sedimentscrop out extensively ([7, 8]; Figure 1). A new map of lateSantonian palaeogeography, presented here, di�ers fromthe previous ‘fully marine, subtidal Santonian’ palaeo-geographic models of the Cretaceous Münsterland Basin([7, 9–12]; Figure 1).

The Late Cretaceous marine vertebrate locality dis-cussed here (Figure 1) is a small, long-abandoned quarrylocated on a hill called Weiner Esch, 5 km south ofOchtrup in northwestern-most Westphalia, northwesternGermany (topographical map Metelen, coordinates lati-tude 25° 80’50” S and longitude 57° 84’82” E). The quarryis protected by a preservation order, as well as being a nat-ural and palaeontological monument.

Previously, only rare, badly preserved, isolatedselachian teeth have been recorded from Weiner Esch,

*Corresponding Author: Cajus G. Diedrich: Private ResearchInstitute PaleoLogic, Petra Bezruce 96, CZ-26751 Zdice, Czech Repub-lic, E-mail: [email protected] Scheer: Ruhr Museum,Fritz-Schupp-Allee 15, D-45141 Essen,Germany, E-mail: [email protected]

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Marine vertebrates from the Santonian coastal carbonates of northwestern Germany | 343

Figure 1: A: Map showing the study area within central Europe. B–C: Geographic position of the Santonian vertebrate and dinosaur bonelocalities of northwestern Germany discussed in this paper (Map of Münster Cretaceous Basin after [7, 11]; location of vertebrate fossil sitesafter: 1. Aachen: [1]; Halle- Ascheloh: [37]; Höver: [4, 55]).


344 | C.G. Diedrich and U. Scheer

and little of this material was described in detail [7–9, 13].This publication records a well-preserved assemblage in-cluding selachians, some teleosteans, and a few reptileteeth with the aim of better understanding the shark, �sh,and reptile biodiversity of the upwelling-in�uenced up-per Santonian shallow-marine carbonate sediments ofthe northern Münsterland Cretaceous Basin, within thesouthern Proto- North Sea Basin. In this contribution, the�rst Santonian dinosaur remain fromnorthern Germany ispresented. Together with yet unpublished �nds from twoother southern Proto- North Sea Basin localities – WeinerEsch, Lingen (Lower Saxony), and a new locality (con-taining dinosaur remains and a comparable shark faunato Weiner Esch) north of the Harz Mountains (Lower Sax-ony), these dinosaur �nds are important to the improvingof the palaeogeographic map of Ziegler 1990 [12]. It allowsthe interpretation of coastline and island positions, andtherefore the reconstruction of the swell’s uplift history.

2 Material and methodsA small collection of material from Weiner Esch wasmade by K. Niemeyer between 1966 and 1975, and his fewselachian teeth are now deposited in the Geomuseum ofthe Westphalian Wilhelms-University Münster, Germany.A further small, but important collection, including thesingle fragment of a dinosaur tooth as well as the onlyrelict of a chimaera, was later made by E. Wijnker (Nether-lands), which was donated to the Ruhr Museum (Essen,Germany) and is housed together with the huge collec-tion of K.-H. Hilpert. In addition to these three privatecollections, the current authors etched three kilograms ofsediment collected from Conglomerate Bed II (Figure 2)with formic acid to obtain additionalmicroselachians, pla-coid denticles and teleostean teeth. Of thismaterial, small-sized fossils were photographed with a REM, and largerteeth photographedusing a binocularmicroscope,with alldrawings produced using a binocularmirror (Figures 3–7).As the terrestrial reptile material (Figure 8) has proved im-portant for interpreting palaeogeography (Figure 9), twomore sites from northern Germany are mentioned herein,although thedinosaurmaterial from these localities awaitsdetailed investigation and is as yet unpublished. This un-publishedmaterial includes fossils from a site north of theHarz Mountains, stored in a private collection; and a col-lection from Lingen, deposited in the Museum of NaturalHistory Oldenburg (Staatliches Museum für NaturkundeOldenburg).

Institutional abbreviations: Geomuseum of the West-phalian Wilhelms-University Münster, Germany (GPI),Ruhr Museum Essen, Germany (RE).

3 Geology

3.1 Stratigraphy and age

In [7] described the 8.5 m section of limestone at WeinerEsch as ‘Weiner Schichten’, a unit now includedwithin theBurgsteinfurt Formation (upper Santonian– lowerCampa-nian) withWeiner Esch as its stratotype [11]. Due to quarryback�lling prior to the site’s protection, only �ve metresof these sediments are still exposed (Figure 2). This sec-tion is informally subdivided herein into ‘Lower Conglom-erate Beds’ and ‘Upper Sandy Limestone Beds’. At thislocality, the Burgsteinfurt Formation discordantly over-lies the older lower Santonian Emscher Formation [8] dueto a tectonic uplift that created a submarine swell (theOchtrup Swell) as the northeastern prolongation of theDutch High during the Santonian [8, 10]. During this time,Upper Triassic, Jurassic and Lower Cretaceous sedimentsbecame inverted and eroded during an alpinotype subher-cynic tectonic period (Wernigeröde Phase sensu [7]), form-ing the Northwestphalian-Lippe Swell. With its parallelmountain ranges – Wiehengebirge and Teutoburger Wald– this former swell separates the Münsterland Basin fromthe rest of the North German lowland. This tectonic eventexplains the presence of conglomerates, extraclasts, andeven the reworked shark teeth seen within our material(see Taphonomy section). Most vertebrate remains (95%)seem to be autochthonous within the sediment, an inter-pretation followed by other authors [7, 8], although someof the selachian species from Weiner Esch noted in thispaper obviously originate from reworked Cenomanian–Coniacian, or even middle Santonian layers (Table 1). Acomparable taphonomic situation is found in the Eoceneshark-gravels of the Fürstenau Formation (Lower Sax-ony, about 80 km northeast of Weiner Esch) of north-ernGermany,which also contains reworked Cenomanian–Turonian species [14].

The sediments in Weiner Esch are considered lateSantonian in age (= krsa4; [7, 8]) based on the presenceof the inoceramid Sphenoceramus patootensis and cal-citic rostrae of the belemnites Gonioteuthis granulata andActinocamax verus. These index taxa were collected bythe present authors from a number of di�erent conglomer-ate beds (especially from Conglomerate Bed II; Figure 2),again supporting the traditional interpretation of this unit



Figure 2: Stratigraphy of the Burgsteinfurt Formation (upper Santonian) exposed at the Weiner Esch vertebrate locality, located nearOchtrup in northwestern Germany (Chronostratigraphic data after [11]). Shark teeth and other macrofaunal remains are preserved at thebases (= conglomerates) of several parasequence sets (storm deposits I–XVIII).



as late Santonian in age. Unfortunately, other more bios-tratigraphically diagnostic taxa, such as ammonites, aremissing from this unit.

3.2 Sedimentology

The sedimentary section at Weiner Esch is dominated byarenitic limestones containing 85-95% calcium carbon-ate [7, 8]. Calcite shell producers, such as the lamelli-branchs Sphenoceramus and Pycnodonte, plus crinoids,belemnites, bryozoans and balanids, are present in themajority of bioclasts. Of this group, crinoid remainsdominate (50%), followed by bryozoans (30%; see also[7]), belemnites, echinoids, oysters, balanids, and in-oceramids. These fossils are often rounded, sorted ac-cording to grain size, and are seldom greater thanfour centimetres in length, although belemnites and in-oceramids are occasionally larger and sometimes com-pletely preserved. The microfauna is dominated by sand-agglutinating foraminifers [8]. As a result of the late dia-genetic dissolution of aragonite, no gastropods and only afew other lamellibranchs are preserved in the assemblage.

The middle part of the section of the BurgsteinfurtFormation (Figure 2) visible at Weiner Esch contains atleast 18 thin �ning-upward sets (Figure 2), with vertebrateremains and invertebrate marine fossils remarkably en-riched at the base (2-10 cm) of each of these sets. The Con-glomerate Beds II and III with their high density of extr-aclasts and fossil remains possibly represent ‘large-scale’Milankovitch sequence boundaries (Figure 2). Fossilifer-ous beds are typical for such boundaries in carbonate sys-tems (e.g. [6]), where each parasequence might representcoastal storm events, compared to other shallow-marinecarbonate storm deposits with similar �ning-upward sets(e.g. [15]). The nearshore shallow-coastal subtidal facies ofthe submarine Ochtrup Swell, thewestern adjacent part orprolongationof theNorthwestphalian-LippeSwell,was in-�uenced by storm sediments transported across the swellto the levee (i.e. the southeastern slope) from the north-west and north, as indicated by the orientation of shal-low subtidal channels and cross bedding within this fa-cies [8]. Gutter-casts and cross-bedded carbonate sands,or channels, �lled with fossiliferous conglomerates, char-acterize the shallowmarine coastal conditions around thisswell [7, 8].

The conglomerates are predominantly composed ofslightly to well-rounded extraclasts of Triassic, Jurassic,and Lower Cretaceous marls and dolomites, dark brownclaystones, asphaltites, red sandstones and claystones,but also includes smaller, rounded Upper Cretaceous (Tur-

onian) limestone pebbles. Jurassic ammonite fragmentshave been found within these pebbles (body chamberspreserved 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 theconglomerates, mainly formed of carbonate particles andmarine invertebrate remains, additionally contains quartzand glauconite grains. Especially together with the nu-merous phosphorite nodules seen in these conglomerates,they indicate the in�uence of upwelling on the deposi-tional environment [7, 8].

4 Paleontology

4.1 Faunal taphonomy

At Weiner Esch, the vertebrate fauna is particularly richin Conglomerate Beds I and II but fossils are found in allconglomeratic layers (Figure 2). Only teeth and small bonefragments are present in this assemblage due to the sort-ing e�ects of palaeocurrents, with selachians mainly rep-resented by teeth and few dermal scales. Based on thepreservation of sharp cutting edges (especially nearly allOdontaspidae) on about 95% of the selachian teeth, itseems clear that these teethwere not transported over longdistances or extensively reworked. The roots of these au-tochthonous teeth are very soft, due to late diagenesis,and therefore aremostly removed in Scapanorhynchus andother selachian taxa. In contrast, there is a small num-ber of allochthonous teeth in the assemblage, identi�edbased on their rounded cusps and roots (Table 1). Havingpassed through diagenesis prior to the deposition of thesediments at Weiner Esch, the roots of the allochthonousteeth are ‘prefossilized’ and often incompletely preserved.

4.2 Vertebrate biodiversity and assemblagecomposition

Listings of the diverse marine-invertebrate and vertebratemacrofauna from Weiner Esch have been previously pub-lished [3, 7, 8], although these lists typically used older‘morphotype systematics’, not easy to correlate with sev-eral recent faunal revisions. The extremely confusingnomenclature of shark teeth is a particular problem thatneeded to be revised in the future. For example, the toothmorphotaxon Cretoxyrhina is now known to belong to thegenus Isurus [16], which was described using skeletal re-mainsmore than just teeth. Similar revisions have recently



Figure 3: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Paraorthacodus recurvus, anteriortooth, a–b. labial, RE No. 3786/10. 2. Paraorthacodus recurvus, posterior lateral tooth, a, c. lateral, b. occlusal, RE No. A 3782. 3. Ptychoduscf. polygyrus, tooth, a. lateral, b–c. occlusal, GPI No. A1C-2-1 (from reworked Cenomanian layers). 4. Hexanchus microdon, lateral toothof 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, RENo. 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, RENo. A4815. 12. Scyliorhinus (or Crassescyliorhinus) a–b. labial, RE No. A4816. 13. Pteroscyllium sp., tooth crown, a. lingual, b–c. labial, RENo. A 4812/1.



Table1:

Verteb

rate

teethreco

veredfro

mco

nglomerates

with

intheup

perS

antonian

(Upp

erCr

etac

eous

)San

dySa

ndylim

estone

sof

Weine

rEsc

h,ne

arOc

htru

p,no

rthw

estern

Germ

any(all

materialisstored

attheRu

hrMus

eum

Esse

n,Ge

rman

y).

Spec

ies

Material

Agerang

eSe

lach

iiParaorthacodus

recurvus

(Trautsc

hold,

1877

)2(1

anterio

rand

1lateraltoo

th)

Coniac

ian–

Campa

nian

Ptycho

duscf.polygurus

Agas

siz,

1835

2(1

rootless

and1toothfra

gmen

t)(Rew

orke

dfro

mup

per

Ceno

man

ian

strata)

Hexanchu

smicrodon(Aga

ssiz,1

843)

15(3

lateral,

4an

terolateral,

8lateral

teeth

and

fragm

ents

and

some

tooth

cusp

s)

Ceno

man

ian–

Maa

stric

htian

Protosqu

alus

cf.sigei

Capp

etta,1

977

3inco

mpleteteeth

Ceno

man

ian–

Campa

nian

Heterodontus

rugosus(Aga

ssiz,1

843)

7(2

anterio

rand

5lateraltee

th)

Coniac

ian–

Campa

nian

Chiloscyllium

cf.greenei

(Cap

petta

,197

3)1an

terio

rtoo

thlower

Ceno

man

ian–

Campa

nian

Carcha

riassubu

lata

(Aga

ssiz,1

844)

37(17an

terio

r,4an

terolateral,10

lateral

teeth,

6cu

sps)

Ceno

man

ian–

Campa

nian

Scapanorhynchu

sraph

iodon

(Aga

ssiz,

1844

)20

9(15

anterio

r,12

lateralteeth,

182

cusp

s)Ce

noman

ian–

Campa

nian

Megarhizodonmacrorhiza(Cop

e,18

75)

2(1

anterio

rand

1lateraltoo

th)

(Rew

orke

dfro

mAlbian

–Cen

oman

ian)

Isurus

mantelli

(Aga

ssiz,1

843)

30(1

anterio

r,2lateraltee

th,2

7cu

sps)

Coniac

ian–

Campa

nian

Lamna

append

iculata

pachyrhiza

(Her-

man

,197

5)44

(3an

terio

r,on

esy

mph

y-se

al/a

nteriolateral,

20lateral

teeth,

and20

crow

ns)

Coniac

ian–

Maa

stric

htian

Lamna

arcuata(W

oodw

ard,

1894

)5(1

anterio

laterala

nd4lateraltee

th)

Ceno

man

ian–

Campa

nian

Cretodus

semiplicatus

(Mün

ster

inAg

as-

siz,

1843

)1lateraltoo

thCe

noman

ian–

Campa

nian

Protolam

nasokolovi

(Cap

petta

,198

0)3

(1an

terolateral,

one

lateral,

and

one

toothcrow

n)(Rew

orke

dfro

mAp

tian–

Ceno

man

ian)



Table1:

Table1c

nd.

Spec

ies

Material

Agerang

eProtolam

naacutaMüller&

Died

rich,

1999

2an

terio

rtee

thCe

noman

ian–

Santon

ian

Paranomotodon

angu

stidens

(Reu

ss,

1845

)8(1

anterio

rand

7lateraltee

th)

Ceno

man

ian–

Maa

stric

htian

Palaeohypotodu

sbronni

(Aga

ssiz,1

838)

2lateraltee

thCa

mpa

nian

–Maa

stric

htian

Squalicorax

kaup

i(Ag

assiz,

1835

)42

(2an

terio

r,on

elateraltee

th,3

9cu

sps)

Coniac

ian–

Campa

nian

Scyliorhinus

elonga

tus(D

avis,1

887)

1lateraltoo

thof

theup

perjaw

Santon

ian

Scyliorhinus

cf.blotiCa

ppetta,1

980

1tooth

(Rew

orke

dfro

mCe

noman

ian)

Scyliorhinus

sp.o

rCrassescyliorhinus

sp.

1tooth

(Rew

orke

dfro

mCe

noman

ian)

Pteroscyllium

sp.

3teethcrow

nsCe

noman

ian–

Santon

ian

Chim

aerif

ormes

Ischyodu

scf.bifu

rcatus

Case

,197

81jaw

fragm

ent

Santon

ian–

Maa

stric

htian

Teleos

tei

Pycn

odon

tidae

inde

t.1tooth

Cretac

eous

Coelodus

sp.

1tooth

Cretac

eous

Enchodus

sp.

7teeth

Cretac

eous

Reptilia

Mos

asau

ridae

inde

t.(Leiodon/Clid

astes)

3toothfra

gmen

tsSa

nton

ian–

Maa

stric

htian

Tyrann

osau

ridae

inde

t.1toothfra

gmen

tSa

nton

ian–

Maa

stric

htian



Figure 4: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Carcharias subulata, anteriortooth 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. A3786/01. 3. Carcharias subulata, lateral tooth of the upper jaw, a–b. labial, RE No. A 3786/06. 4. Scapanorhynchus raphiodon, anteriortooth, a–b. labial, RE No. A 3784/09. 5. Scapanorhynchus raphiodon, anterior tooth, a–b. labial, RE No. A 3784/08. 6. Scapanorhynchusraphiodon, 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.



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. Isurusmantelli, lateral tooth, labial, RE No. A 6530.



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 appendiculatapachyrhiza, lateral tooth, a–b. labial, c. lateral, d. lingual, RE No. A 3778/1.



been published for the white (Carcharodon) and mega-tooth sharks (Otodus: see [5]). As these shark species arealready well described in the literature, none of the �ndsfromWeiner Esch are described herein again (see [2, 5, 16–44]). Shark scales remain the most di�cult fossils fromWeiner Esch to identify, even to genus level [37], due to thechangingmorphology of scales over di�erent body regions[34]. Teeth of teleosts, mosasaurs theropods are also di�-cult to identify to genus level as a result of poor preserva-tion, fragmentation, or a lack of characteristic features.

According to the references mentioned above, 20di�erent selachian taxa (14 macroselachians and 6 mi-croselachians) have been identi�ed from the Weiner Eschlocality of the Burgsteinfurt Formation (upper Santo-nian). Especially Scapanorhynchus teeth could be placedwithin the jaw according to associated tooth sets fromAmerica [45], whereas for other species modern sharkdentitions were compared [46]. The investigated assem-blage consisted of 599 determinable (Table 1, Figure 9)and 135 non-determinable selachian teeth. The assem-blage comprises 59% Scapanorhynchus raphiodon (Agas-siz, 1843) (Figures 4.4–8), 8% Carcharias subulata (Agas-siz, 1844) (Figures 4.1–3), 7% Squalicorax kaupi (Agas-siz, 1835) (Figure 7.10), 6% Isurus mantelli (Agassiz, 1843)(Figures 5.1–3), 5% Lamna appendiculata pachyrhiza (Her-man, 1975) (Figures 6.1–4), 4%Hexanchusmicrodon (Agas-siz, 1835) (Figures 3.4–5), 2% Paranomotodon angusti-dens (Reuss, 1845) (Figure 7.7), 1% Lamna arcuata (Wood-ward, 1894) (Figure 7.1), and 1% Heterodontus cf. rugosus(Agassiz, 1839) (Figure 3.7–8). All other recorded specieseach represent less than 1% of the assemblage: Proto-squalus cf. sigei Cappetta, 1977 (Figure 3.6), Chiloscyl-lium cf. greenei (Cappetta, 1973) (Figure 3.9), Paraorthaco-dus recurvus (Trautschold, 1877) (Figures 3.1–2), Megarhi-zodon macrorhiza (Cope, 1875) (Figures 7.3–4), Ptychoduscf. polygyrus Agassiz, 1835 (Figure 3.3), Cretodus semi-plicatus (Münster in Agassiz, 1843) (Figure 7.2), Proto-lamna sokolovi (Cappetta, 1980) (Figures 7.5–6), Proto-lamna acuta [37], Palaeohypotodus bronni (Agassiz, 1843)(Figures 7.8–9), Scyliorhinus elongatus (Davis, 1887) (Fig-ure 3.10), Scyliorhinus cf. bloti Cappetta, 1980 (Figure 3.11),Scyliorhinus sp. or Crassescyliorhinus sp. (Figure 3-12).Pteroscyllium sp. (Figure 3.13). Chimerae are representedusing illustrations of [47] by a single jaw identi�ed as Is-chyodus cf. bifurcatus Case, 1978 (Figure 8.1). Bony �shesare only represented by one tooth of Pycnodontidae indet.(Figures 8.3), and oneCoelodus sp. (Figure 8-2) and few En-chodus sp. (Figure 8.4).

Marine reptiles from this locality are repre-sented by teeth fragments of Mosasauridae indet.(Leiodon/Clidastes) (Figures 8.5–6) (as per the genus de-

scriptions in [48–51]), whereas dinosaurs are recorded bya single tooth fragment belonging to a theropod (Tyran-nosauridae) [52] (Figure 8.7).

5 Discussion

5.1 Comparison of Mesozoic sharkassemblages in northwestern Germany

The relatively diverse (20 species) shark assemblage pre-served at Weiner Esch demonstrates some similarities tothe fauna of 38 early Cenomanian shark species recordedat Ascheloh near Halle/Westphalia, in the TeutoburgerWald Mountains ([37]; Figure 9). At both sites, the numberof macroselachian species is comparable, with 12 speciesrecorded at Halle-Ascheloh and 14 at Weiner Esch (Fig-ure 9). At Halle-Ascheloh, the greater abundance of mi-croselachians allowed greater amounts of research on thisgroup, accounting for the increased diversity (26 species)in comparison to themicroselachians ofWeiner Esch. Fau-nal mixing seen at both sites seems to be related to tapho-nomic or sedimentary processes, speci�cally slumping atHalle-Ascheloh, and sedimentary condensation at WeinerEsch. Reworked teeth are present in theWeiner Eschmate-rial originating mainly from Cenomanian–Turonian strataof northwestern Germany (Figure 9; Tables 1-2).

The absence of Hemipristidae or Rhinobatidae, anddominance of Mitsukurinidae (goblin sharks), especiallyScapanorhynchus raphiodon (59% of the shark fauna), andlamnids in the Weiner Esch fauna generally indicates thein�uence of boreal and shallow-marine upwelling condi-tions (Table 1, Figure 9). Therefore, the assemblage showspalaeoecological similarities to the early Cenomaniancarbonate-sand shark fauna of HalleAscheloh, where theMitsukurinidae also dominate the macroselachians (Fig-ures 1, 9; [37]). In contrast to the living deep-water sharkMitsukurina [53], it seems that Cretaceous Mitsukurinidaewere associated with cold-water and upwelling conditionsin shelfal regions e.g. the Western Interior Seaway [42]and the Proto- North Sea Basin; both areas under north-ern boreal upwelling in�uence. Like other shark groupsthat have changed life habits during their evolution, thesame is interpreted here for the Mitsukurinidae. Repre-sentatives of this family with lateral cusp developmentreduction, serration, or changes in tooth formula withinthe jaws - are already evident in the early Late Cretaceous[5, 16, 29, 44]. The late Santonian Weiner Esch selachianfauna is most similar to the early Campanian greensandfauna of the Vaals Formation in the Aachen area ([1], Ta-



Figure 7: Sharks from the upper Santonian conglomerates of Weiner Esch (northwestern Germany). 1. Lamna arcuata (Woodward), lateraltooth, 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.



ble 1) near the German–Belgian–Netherlands border. TheAachen selachian fauna contains a number of species sim-ilar to Weiner Esch (Table 2), and was also deposited in ashallownearshoremarine environment, as indicatedby itslithology (glauconitic carbonate sands; [1]). However, mi-croselachians in theAachen assemblagewere only studiedusing small samples, and percentages of individual sharkspecies were not quoted, thereby hampering comparisonsto other northwest German shark assemblages.

Conversely, a number of open marine early and lateCampanian vertebrate and selachian faunas from Ger-many can be seen to be considerably di�erent from theassemblage at Weiner Esch [3]. The only similarities arethe presence of deep-sea or cool-water species at WeinerEsch, compared with the Squalidae (Protosqualus) thatare particularly well represented in the Late Cretaceous ofnorthern and northwestern Europe (e.g. [2, 37]; Table 2).The microselachian genera Chiloscyllium and Scyliorhinusrecorded at Weiner Esch are also present in these Campa-nian deposits, although represented by di�erent species,all of which indicate the in�uence of subtropical, warmsurface-water from Tethys in the environment [27, 28]. Anearly Campanian selachian fauna (12 species) collectedfrom marly limestones at Höver, near Hannover (LowerSaxony, northern Germany), di�ers from the Weiner Eschassemblage particularly in the microselachians [4]. Thisdi�erence seems to be a result of the palaeoenvironment,which at Höver was an open marine facies with more orless continuous sedimentation, lacking sedimentary con-densation or vertebrate bone-bed formation, and thereforereducing the likelihood of �nding teeth. Among the low-abundance sharkmaterial recovered fromHöver, the dom-inant squalomorph sharks (six neritic and pelagic species,including Cretascymnus westphalicus and Centroscymnuspraecursor) are considered deep-water selachians [3] liv-ing in a deep basin, agreeing with the interpreted palaeo-geography of the ProtoNorth Sea Basin at this time (Fig-ure 9, and [12]).

Comparison of the Weiner Esch assemblage to thesharks collected from the Middle Eocene Fürstenau For-mation from Bippen and Dalum (Lower Saxony) wasalso considered important, as a mixing of reworkedCenomanian–Turonian shark teeth within a predomi-nantly autochthonous fauna was also observed in thislatter formation (Figures 1, 9; [14, 44]). The Weiner Esch(Santonian) and Bippen (Eocene) assemblages have com-parably high percentages of sand sharks (Odontaspidae),and similarities in the upwelling in�uence within thefauna [14]. Therefore, it is suggested that an abundance ofsand sharks is a good marker for identifying very shallow,coastal marine palaeoenvironments, as also seen in the

lower Cenomanian Halle-Ascheloh site ([37]; Figure 9). Inthe Eocene localities, sand sharks are even higher in pro-portion than in the Santonian fauna (Figure 9), which �tsthe interpreted palaeogeography of very shallow marineconditions along the southern margin of the Proto- NorthSea Basin during this time [14].

5.2 Uplift reconstructions of the submarineNorthwestphalian-Lippe swell

The Cenomanian Halle-Ascheloh site, concentration ofvertebrate remains resulted from marine slumping alongthe margin of a submarine swell and elongated carbon-ate platform, the Northwestphalian-Lippe Swell, locatedat the northern margin of the Münsterland CretaceousBasin (Figure 9, [6]). The uplift of this feature began dur-ing the early Cenomanian [54], �rst causing large-scaleslumping on the swell’s southern slopes (Figures 1, 9). Inthe middle late Cenomanian, the carbonate platform alsochanged tectonically along its southern margin, with con-tinuous high subsidence, reaching the deep-water con-ditions required for the deposition of shark and �sh-rich black shales during the second Cretaceous OceanicAnoxic Event (= OAE II) in the Osning Depression (Fig-ure 9; [54, 55]). The hercynic-controlled tectonic uplift isbest exposed at Halle and Bielefeld (Ostwestfalendammsite), both of which lie along the southern slope of thesubmarine swell, where upper Turonian – lower Conia-cian units contain large carbonate ‘phacoids’ of slumpedCenomanian–Turonian consolidated carbonates up to sev-eral meters in size [56, 57]. It is demonstrated here thatthe swell regions (Ochtrup Swell, and others, Figure 9)must have already been uplifted to subaerial or shallowintertidal conditions (i.e. carbonate-sand islands) duringthe late Santonian, thereby separating the MünsterlandCretaceous Basin (Figure 9) from the Proto- North SeaBasin. Similar in age to the hercynian uplift event are thenorthern German Harz Mountains, of which its earliest ‘is-land’ stage has beenpreviously dated asmiddle Santonian[58, 59].

5.3 Dinosaur migration along intertidalzones or islands in northern Germany

[9] interpreted the sediments at Weiner Esch and thenearby Seller Esch site (no longer exposed) as ‘coastal sed-iments’ and was the �rst to propose the existence of ‘is-lands’ in northwestern Germany during the Late Creta-ceous. In the [12] palaeogeographic maps of Europe, inter-



Figure 8: Bony �shes chimaeras, and marine and terrestrial reptiles from the upper Santonian conglomerates of Weiner Esch (northwesternGermany). 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.



Table2:

Compa

rison

oftheLa

teCr

etac

eous

selach

ianfaun

asof

Germ

any,

centralE

urop

e,with

theglob

alCr

etac

eous

selach

ianreco

rd,a

ndthepa

laeo

ecolog

ical

preferen

cesof

know

ntaxa

.

Taxo

nPa

laeo

geog

raph

yAg

eBa

thym

etry

Clim

ate

Selach

ii

Aachen

Coesfeld

WeinerEsch

Hannover-Höver

NortheEurope

NorthAmerica

NorthAfrica

Cenomanian

Turonian

Coniacanian

Santonian

Campanian

Maastrichtian

benthic

nectic

neritic

pelagic

tropical/subtroppical

temperate

boreal

eurytherm

Paraorthacodus

recurvus

(Trautsc

hold)

xx

xx

––

xx

xx

Ptycho

duspolygu

rusAg

assiz

xx

xx

––

xx

xx

Hexanchu

smicrodon(Aga

ssiz)

xx

xx

x–

––

–x

xx

Protosqu

alus

cf.sigei

Cape

ttax

xx

––

––

xx

xx

Heterodontus

rugosusAg

assiz

xx

x–

–x

xx

Chiloscyllium

cf.greenei

Cape

ttax

xx

xx

xx

––

––

xx

xCarcha

riassubu

lata

(Aga

ssiz)

xx

x–

––

xx

xx

Scapanorhynchu

sraph

iodonAg

assiz

xx

xx

x–

––

xx

xx

Isurus

mantelli

(Aga

ssiz)

xx

xx

x–

––

xx

xLamna

append

iculatapachyrhiza

(Herman

)x

xx

xx

xx

––

––

xx

xPlicatolam

naarcuata(W

oodw

ard)

xx

x–

––

–x

xx

Cretodus

semiplicatus

(Mün

ster)

xx

xx

x–

––

–x

xx

Protolam

nasokolovi

Cape

ttax

x–

––

xx

xProtolam

naacutaMüller&

Died

rich

xx

x–

––

xx

xx

Paranomotodon

angu

stidens(Reu

ss)

xx

xx

x–

––

–x

xx

Palaeohypotodu

sbronni

(Aga

ssiz)

xx

––

–x

xx

Squalicorax

kaup

i(Ag

assiz)

xx

xx

x–

––

xx

xScyliorhinus

cf.elong

atus

Cape

ttax

xx

x–

––

xx

xScyliorhinus

cf.blotiCa

petta

xx

xx

––

–x

xx

Pteroscyllium

sp.

xx

xx

––

––

xx

xMegarhizodonmacrorhiza(Cop

e)x

xx

x



tidal zoneswere not delineated at all for the Santonian. Forthe new interpretation presented herein, the shark faunasand position of three dinosaur localities (Figure 9) wereused to interpret a preliminary newpalaeogeographicmapof the Münster Cretaceous Basin and its northern realm inthe late Santonian. It seems, based on Santonian dinosaurremains atWeiner Esch (tooth �gured herein), Lingen, andnorth of the Harz Mountains (represented by as yeat un-described postcranial longbones and fragments) in north-ern Germany (Figures 1, 9), that the uplifted submarineswell regions of northern Germany and the Netherlands(the Dutch High, Ochtrup High and Northwestphalian-Lippe High) must have been periodically intertidal or evenshallow carbonate-sand islands.

The idea of dinosaurs using beaches and intertidalareas as possible seasonal migration zones is becomingmore favoured, with many new discoveries of dinosaurtracks recorded in European Middle Triassic [60, 61] orUpper Jurassic [62] coastal or intertidal beach deposits,and examples such as recently described Cenomanian–Turonian trackways on the Adriatic Carbonate Platformand in central Europe [63]. Only such intertidal ‘bridges’can explain the distribution of Santonian dinosaur recordsalong European Cretaceous islands or peninsula coasts.

Within Europe, the Santonian dinosaur remains ge-ographically closest to Weiner Esch are described fromwest and central France (theropod teeth and a few bonefragments from Charente-Maritime, Maine-et-Loire, Indre-et-Loire, Vendee: [52]) and from Hungary (a few herbivo-rous dinosaur teeth (Rhabdodontidae indet., Nodosauri-dae indet.) and theropod remians of uncertain relation-ship from Iharkút, Bakony Mountains: [64]. These rare,scattered remains indicate awidespread distribution of di-nosaurs across Europe in the Santonian, supporting theidea that at this time the area contained several islandsand small landmasses ([12], Figure 9).

6 Conclusions

The late Santonian marine vertebrate fauna of WeinerEsch, exposed near Ochtrup in northern Germany, pro-vides important insights into the palaeogeography andhistory of the submarine Northwestphalian-Lippe Swell,thereby helping to �ll a gap in knowledge of this region’sevolution during the Late Cretaceous. Rare dinosaur re-mains recovered from three localities along the swell, sup-port the interpretation of an already uplifted environment

on the Ochtrup and Northwestphalian-Lippe Swells thatconnected the Dutch High with the uplifting Harz Moun-tains in the southern Proto- North Sea Basin of centralEurope. This swell started to uplift in the early Cenoma-nian, and later formed a back-arc-like depression (the Os-ning Depression), which was �lled with vertebrate-richblack shale layers in the late Cenomanian – early Turo-nian. Tectonic activities continued in this region in thelate Turonian – early Coniacian with slumping of largerphacoids along the swell slopes, �nally forming subaerialcarbonate-sand intertidal or island conditions in the lateSantonian. In the Eocene, the Northwestphalian-LippeSwell was uplifted to form a peninsula or islands, initi-ating the formation of the Wiehengebirge / TeutoburgerWald Mountains north of the Rhenish Massif. The mainvertebrate material obtained from Weiner Esch consistedof marine shark teeth (20 species), that together indicatethe in�uence of cold, upwelling (based on the presenceof glauconite and phosphorite nodules), boreal water dur-ing the late Santonian, with periodic, seasonally warmsurface-water in�uences derived from Tethys. The highpercentage of sand sharks (59%), especially S. raphiodon(Mitsukurinidae), seen in this assemblage is considereda result of this cold water and the upwelling in�uence,plus the concentration of shelfal shark teeth in a coastalcarbonate-sand facies. The bathymetry of the swell regionduring the late Santonian is estimated between intertidalto shallow subtidal (greater than 0 m water depth, and0–5 m along the swell margins). Additionally, rare thero-pod remains recovered from this site, along with other di-nosaur bones collected from two more sites on and alongthe submarine swell, indicate the seasonal migration orfaunal exchange of dinosaurs along coasts or intertidalzones, likely between islands, of Europe in the Late Cre-taceous.

Acknowledgement: The authors are very grateful to K.-H. Hilpert, Dr. K. Niemeyer and E. Wijnker, who collectedmost of the material described herein. Dr. M. Bertling isthanked for providing permission to study the materialstored in the Geomuseum of the Westphalian Wilhelms-University Münster, Germany. Sta� at PaleoLogic pro-duced all teeth drawings, illustrations and other graphicwork. Museological work on the material was sponsoredby the Ruhr Museum, and PaleoLogic sponsored the re-search. Finally, we thank M. Reinhardt for preparation ofthe binocular photos, and Dr. H.-M. Weber for supplyingus with REM photos.



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 TeutoburgerWald and Wiehengebirge mountain ranges developed in northern Germany from the southern Proto- North Sea Basin .



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