Meded. Tert. Kwart. Geol. 171-180 2 1987 Werkgr. 24(1-2) pp. figs Leiden, June

Evolution and stratigraphical distribution of Oligocene and Miocene Scaphopoda in the North Sea Basin — a preliminary report

by

Ronald Janssen

Forschungsinstitut Senckenberg,

Frankfurt am Main

F.R.G.

Janssen, Ronald. Evolution and stratigraphical distribution of

Oligocene and Miocene Scaphopoda in the North Sea Basin — a

preliminary report. — Meded. Werkgr. Tert. Kwart. Geol., 24(1-2):

171-180, 2 figs. Leiden, June 1987.

As a result of a taxonomical revision of Oligocene and Miocene scapho-

pods ofthe North Sea Basin observations on stratigraphical distribution

well of as as on evolutionary lineages species are presented. There are

from the the two important lineages reaching Latdorfian, resp. Rupelian, to the Vierlandian (Fissidentalium novaki-lineage and Antalis

Miocene geminata-lineage) and one (Reinbekian to Syltian) lineage

(based on Fissidentalium “badense” auct.). Assemblages of scaphopod

species characterize different facies in the Rupelian, but stratigraphical

units in the Hemmoorian. Most previous assumptions as to identity of

species and their development in the North Sea Basin appeared to be

incorrect.

Dr R. Janssen, Forschungsinstitut Senckenberg, Senckenberganlage

25, D-6000 Frankfurt am Main, Federal Republic of Germany.

Contents: 172 Samenvatting, p.

Zusammenfassung, p. 172

172 Introduction, p.

of in the North Sea Basin, 174 Evolutionary development scaphopods p.

Stratigraphical distribution of Oligocene species, p. 175

Stratigraphical distribution of Miocene species, p. 177

Some 179 general considerations, p.

179 Notes, p.

180 Acknowledgements, p.

180 References, p. 172

Samenvatting

miocene in Noordzee- Evolutie en stratigrafische verspreiding van oligocene en Scaphopoda het

bekken—een voorlopig rapport.

Door middel van een kritische taxonomische herziening van de oligocene en miocene Scapho-

het Noordzeebekken werd de toe te poda van mogelijkheid geschapen Scaphopoda passen voor een

stratigrafische ouderdomsbepaling. Zowel evolutiereeksen als kenmerkende soortenassociaties

kunnen nu voor stratigrafische doeleinden worden gebruikt. Van belang zijn vooral drie ontwikke-

nl. Fissidentalium novaki - novaki - lingsreeksen, (Von Koenen) n. subsp. polypleurum Seifert (Latdorf tot

Antalis - Goldfuss — - Vierland), n. sp. geminata geminata n. subsp. n. sp. (Rupel tot Vierland) en

Fissidentalium - Zimmermann “badense” - n. sp. floratum (= auct.) floratum n. subsp. (Reinbek tot

Sylt).

vooral verschillende niveau’s Karakteristieke soortengezelschappen zijn kenmerkend voor van

Hemmoor-ouderdom. Ook in het komen soortcombinaties die hier echter Rupelien typische voor,

andere meer tot bepaalde facies beperkt zijn. Soortenuitwisselingen met Tertiairbekkens hebben alleen tijdens het Hemmoor bestaan met het Burdigalien van de Aquitaine. Directe betrekkingen tot het Mioceen in niet van het Paratethys-gebied zijn, tegenstelling tot bestaande opvattingen, aantoon-

Het is dat alle door in 1959 aanzien de baar. gebleken Seifert gemaakte veronderstellingen ten van evolutie der in het soortdeterminaties soorten Noordzeebekken gebaseerd zijn op onjuiste en derhalve niet houdbaar.

Zusammenfassung

Durch eine kritische taxonomische Revision der Scaphopoden des Oligozäns und Miozäns im Nord-

der seebecken wurde die Grundlage geschaffen für die Verwendung von Scaphopoden bei stratigra- phischen Altersbestimmung. Es lassen sich nunmehr sowohl evolutive Reihen als auch charakteristi- sche für die Von Artenvergesellschaftungen Stratigraphie nutzen. Bedeutung sind vor allem drei Ent-

Fissidentalium - - wicklungslinien: novaki (von Koenen) novaki n. subsp. polypleurum Seifert (Latdorf-

ium bis Antalis - - - Vierlandium), n. sp. geminata Goldfuss geminata n. subsp. n. sp. (Rupelium bis

und Fissidentalium - Zimmermann “badense” - Vierlandium) n. sp. floratum (‘ auct.) floratum n. subsp.

(Reinbekium bis Syltium).

Charakteristische Artenassoziationen kennzeichnen vor allem verschiedene Niveaus innerhalb

der Hemmoor-Stufe. Auch in der Rupelstufe gibt es spezielle Artengesellschaften, die hier allerdings mehr anderen auf bestimmte Faziesbereiche beschränkt sind. Beziehungen zu Tertiärbecken bestehen während des Hemmooriums der nur zum Burdigal Aquitaine. Direkte Beziehungen zum

Miozän der Paratethys lassen sich entgegen bisherigen Annahmen mit Scaphopoden nicht nachweisen. Es hat sich gezeigt, daß alle von Seifert 1959 vorgestellten Hypothesen über die evolutive

Entwicklung der Arten im Nordseebecken auf falschen Artbestimmungen beruhen und somit nicht haltbar sind.

Introduction

Molluscs have always played an important role in Tertiary research. Especially gastropods and bivalves have attention for of attracted purposes stratigraphy. Contrary to that, however, the 173

have less The for that be scaphopods been more or neglected. reason seems to that scaphopod species

usually look quite uniform in shell-form and sculpture, so that species discriminationoften is very dif-

tubes which ficult. Moreover, scaphopod are frequently broken, renders species identification even

more difficult. It is true that scaphopods of the North Sea Basin have been dealt with in several faunal

Koenen Anderson monographs, e.g. Nyst (1845), von (1882, 1892), Speyer (1870), Kautsky (1925),

R. A.W. but Seifert deals (1964), Janssen (1978), Janssen (1984), only one paper, (1959), exclusively

with scaphopods of the North Sea Basin. In this paper only the species of the Late Oligocene and

Miocene were taken into consideration and the taxonomical and nomenclatural results were mainly

ideas older Koenen and the based on the of authors such as von (1882) Kautsky (1925). Nevertheless,

revision of Seifert still is the only systematic base which has been followed by all later authors.

For the first time, Seifert tried to elaborate specific differences by means of statistical methods.

the distribution of the known him and their Resting upon stratigraphical species as to according to

he scheme of the of that in the morphological similarity proposed a evolutionary development group

North Sea Basin. These hypotheses on the evolution of species have been referred to by other authors

with from other Goddeeris dealing scaphopods basins, e.g. Caprotti (1979). Later on, (1976, 1978)

Miocene but this work continued. So there is started a critical re-evaluation of some species, was not

Basin no modern critical and comprehensive revision of Tertiary scaphopods of the North Sea and ad-

jacent areas.

When well fossil in the ofthe curating Recent as as scaphopod material collections Senckenberg-

Museum I realized that number of (Frankfurt am Main) a hitherto generally accepted species iden-

tifications (e.g. of Miocene species from Northern Germany with Recent species like D. entalis Linne

or with those originally described from the Miocene of the Vienna Basin, like D. badense Homes or

mutabile Homes) can no longer be maintained. This led me to start a comprehensive revision of all

Oligocene and Miocene scaphopods of the North Sea Basin and adjacent basins (Mayence Basin,

Paris the examination of Basin). Especially microsculptural patterns with a Scanning Electron

facilitated refined discrimination and of of Microscope (SEM) a species recognition complexes very

but is all the and similar, different species. One important result that previous hypotheses on identity evolutionary relationships of species were found to be totally untenable.

and The work is based mainly on material from classical outcrops on an evaluation of samples from the borehole Broekhuizenvorst (see van Rooijen et al., 1984). It was possible to establish some well-founded evolutionary lineages which can be used in stratigraphy, as well as some well-defined

in in of the species assemblages. However, some details, particularly transitional zones lineages, observations need closer control materialfrom may some stratigraphical by densely sampled borings.

Insofar it be that amendmentswill be with may possible some turn out to necessary, especially regard

to the exact stratigraphical range of the species. But nevertheless the results presented here allow the

of in be it members of in use scaphopods stratigraphy, as evolutionary lineages or form of defined

in much better than it has been species assemblages, a way possible up to now.

The taxonomical-nomenclatural revision of the scaphopods of the North Sea Basin will be

in the future in the "Archiv fur Molluskenkunde". the published near In present paper only the results and concerning stratigraphical, geographical facies-connected distribution, as well as the evolution of the shall be If more important species presented. new species are involved they will be mentioned in avoid the introduction of nomina nuda. As far open nomenclature, just to as necessary for understanding, remarks on some species are given as annotations at the end of this paper. 174

EVOLUTIONARY DEVELOPMENTOF SCAPHOPODS IN THE NORTH SEA BASIN

Seifert (1959) listed five species for the Middle and Late Oligocene and ten species for the Miocene.

The form which from ribbed species according to Seifert a single evolutionary lineage leads the

Rupelian Dentalium kickxii Nyst, via the Late Oligocene D. geminatum Goldfuss, to the Miocene

D. badense Homes. From this main stem the line of D. polypleurum Seifert - D. bouei Deshayes splits off

the offshoots should Koenen and during Late Oligocene. Miocene be the species D. dollfusi von

D. novemcostatum mutabile Homes. Among the unsculptured species the Miocene D. vitreum Gmelin is

from the D. Lamarck. This supposed to descend Oligocene fissura comparatively simple picture can,

be maintained turned that neither the however, no longer since it out taxonomical assumptions (cor-

the rect species identifications), nor supposed relationships between species are right. Especially the

and in but "central" species kickxii badense the sense of Seifert are not independant species, merely complexes of several quite distinct taxa.

— the kickxii complex

It Antalis kickxii (Nyst) is a species restricted to the facies of the Rupel Clay. apparently disappeared

at the end of the Rupelian. Populations from the basal Rupelian sandy facies (Meeressand of the

Mayence Basin, Berg Sands in Belgium), which have hitherto been attributed to kickxii, belong in

reality to two different species of two separate evolutionary lineages: the first can be regarded as a

MiddleOligocene subspecies of the Early Oligocene Fissidentalium novaki (von Koenen) (note 1) and

the Late The second is leads to Oligocene species polypleurum Seifert. an as yet undescribed species

(note 2), which seems to be the precursor of the Chattian Antalis geminata (Goldfuss). From geminata

the youngest Chattian populations can be separated as an independant subspecies, which gave rise to

further in the Vierlandian. So there different from a new species are two lineages, ranging Early,

Middle the Miocene Vierlandian. Both have with resp. Oligocene, to Early nothing to do true Antalis kickxii (Nyst).

— the badense complex

and is According to Seifert subsequent authors Dentalium badense Homes present during the entire

from the Vierlandian to the with the of Gramian. studies have Miocene, up Syltian, exception My

demonstrated that also in this case not a single species is involved, but a complex of several differing

forms. Moreover, none of the species present in the North Sea Basin can be identified with genuine

badense from the Vienna Basin Miocene (note 3).

The Vierlandian form is a descendant from the Late Oligocene geminata, which became extinct

during the Vierlandian. The occurrence cited for the Hemmoorian is based—apart from misiden-

tifications the record of for the fauna of Hemmoor. —mainly upon Kautsky Kautsky's material,

however, was derived from glacial melting water deposits. It contained, besides true Hemmoorian

from mica All elements, also Reinbekian species and even specimens the Late Miocene clays. the

“badense” from Hemmoor that I have been able from Late material to examine, was evidently derived

" " Miocene Also the citation of badense the Reinbekian is incor- sediments. from by Seifert based on

identified in of Anderson rectly material, this case “dollfusi” auct. non von Koenen. (1964: 194) men-

remarkable of “badense” in the Baumann collection tioned an especially lot from the locality Woltrup,

in Lower This does from that near Osnabriick Saxony. sample, however, certainly not originate

of Reinbekian but from Late Miocene this and locality unquestioned age, a locality (note 4). So, large characteristic species, thought to be badense, appears for the first time in the Langenfeldian, for which 175

it is an important index fossil. Gramian representatives seem to be known only from wells (Hinsch,

From the the Morsum Cliff of 1977; 1979). Syltian at (Island Sylt) a rich population is known, which

merits distinction from A Reinbekian in- “badense” as a separate subspecies. common species, always

correctly identified as D. dollfusi von Koenen by Seifert, Anderson and subsequent authors (note 5)

may be considered as the forerunner of the Late Miocene “badense”. An intermediateform occurs in

the lowermost in well uppermost Reinbekian, resp. Langenfeldian deposits (Giihlitz Mecklenburg;

Broekhuizenvorst, The Netherlands).

Some other, but only shortlived evolutionary lineages are represented in the Oligocene by the

species/subspecies Pseudantalis geinitzi (von Koenen) - sandbergeri (Bosquet) in the Latdorfian and

and — also in the and Rupelian, Episiphon sulcatulus (von Koenen) ottoi (Sharp & Pilsbry), Latdorfian

But these can be used for since discrimina- Rupelian. species hardly stratigraphical purposes, specific

tion in these lineages is only possible with abundantand well-preserved material at hand.

between smooth and The relationship the speciesfissura Lamarck vitreum Gmelin, as supposed by

has real basis: Pseudantalis is Eocene in the Basin Seifert, no fissura an species not occurring North Sea

(but it might be the predecessor of the Latdorfian geinitzi). Dentalium vitreum was originally described

from the Italian Pliocene, its true identity being somewhat doubtful to me (but see Caprotti, 1979:

who considers it of D. Da The Miocene of the North Sea 238, as a synonym vulgare Costa). species

identified is here indicated Antalis exlamarcki Basin, usually as vitreum, as n. subsp.

STRATIGRAPHICAL DISTRIBUTION OF OLIGOCENE SPECIES

Von Koenen described number of from the (1892) a species Early Oligocene (Latdorfian), among

and Fissiden- which the following more common wide-spread forms characterize the Latdorf stage:

" F. Koenen. talium acutum (Hebert), novaki (von Koenen), “Dentalium” dekagonum von

In the Middle there is also number of restricted Oligocene a large species, partially to special

facies. novaki Antalis and Pseudantalis confined Fissidentalium n. subsp., n. sp. sandbergeri (Bosquet) are

Antalis and P. to the basal sandy facies and the Nucula Clay (in Belgium). Especially n. sp. sandbergeri,

however, have a large geographical distribution, not only including the North Sea Basin, but also the

In the Mayence and Paris Basins. Antalis kickxii (Nyst) is an index species for the Rupelian clay facies.

well in also the is Nucula Clay as as the Rupel Clay species Rhabdus parallelus (Zinndorf) found,

be whereas Episiphon ottoi (Sharp & Pilsbry) and Siphonodentalium (?) microceras (Zinndorf) seem to

This confirmation material col- restricted to the upper part of the Rupel Clay section. needs by more

lected from short intervals of the Rupel Clay in Belgium and the Netherlands.

The fauna of the Late is in and scaphopod Oligocene quite poor species. Typical common

A. elements for the Chattian are geminata (Goldfuss), A. pseudofissura (R. Janssen) and Fissidentalium polypleurum (Seifert). The latter species reaches just beyond the Oligocene/Miocene boundary into the

Vierlandian.

The stratigraphical distribution of the more important Oligocene species is shown in fig. 1. 176

oceras

: S.micr llelus R.para n.ssp. Lnata sp. sp. . •

.a ■ gem. A.n A.n A.geminal A. aergeri nitzi

' gei ' P.sand P.

.toi atulus E.ot E.sulc n.ssp. aleurum vaki .no

-poly F

F F.novak]

jtum :kxii F.ac scaphopods. A.ki( agonum Oligocene

D.dek of

Basin Mayence Clay Meeressand distribution Neochatt Eochatt Ruisbroek Sand Rupel Nucula Clay Berg Sands Stratigraphical

CHATTIAN RUPELIAN LATDORFIAN 1. Fig. 177

STRATIGRAPHICAL DISTRIBUTION OF MIOCENE SPECIES

those which descendants During the Vierlandianessentially only species are present, are ofOligocene

forerunners. During the Hemmoorian, however, a large number of new forms suddenly appears,

without affinities to older North Sea Basin species. Several of them demonstrate close relationships

with species from the Burdigalian in the Aquitaine Basin (SW France): Pseudantalis aturensis

'' (Cossmann & Peyrot), Antalis exlamarcki n. subsp., “Dentalium” dumasi n. subsp., Gadila benoisti

Dischides These restricted houthalenensis (Goddeeris), subpolitus (Cossmann & Peyrot). species are to the

Hemmoorian. endemic in the An- Furthermore some apparently species are present North Sea Basin,

' talis dollfusi (von Koenen), and Lentigodentalium “costatum” mutabile' auct. A. dollfusi is confined to

the lowermost Hemmoorian (Behrendorfian) of Belgium (Edegem Sands). The so-called “dollfusi”

from the Reinbekian is not identical with the form of the Edegem Sands, but represents an as yet

has undescribed species. The Species previously identified as novemcostatum mutabile or costatum nothing

the mutabile Homes Vienna with from the to do with true from the Basin, or costatum Sowerby English

Pliocene. the undescribed which is Hem- It is, on contrary, yet another taxon (note 6), typical for the

moorian and the Reinbekian. Two different forms can be distinguished, characterized by the constant

is in Hemmoorian number of ribs (8 resp. 10-11). The form with 10-11 ribs present the Early and in

in Hemmoorian the Reinbekian, whereas the other with only 8 ribs occurs the later (Winterswijk-

Miste, resp. Houthalen).

in the refined distribu- Goddeeris (1978) (note 7) reported a preliminary paperon stratigraphical

of Miocene The observed succession tion of some species from the lower part the Belgian sequence.

results of based material from classical and the well Broek- agrees with the my study on outcrops on

huizenvorst in the is of well-defined Netherlands. It possible, especially by means species

various the classical localities into assemblages, to distinguish horizons, respectively to arrange a relative sequence according to their age. The Edegem Sands are characterized by the true A. dollfusi,

D. “costatum”/“mutabile” (10-11 ribs) and the first occurrence of Dischides subpolitus. The Miste Bed of

the Aalten Member in the Netherlands and the Houthalen Sands in Belgium yield a rich assemblage,

D. with A. exlamarcki n. subsp., D. dumasi n. subsp., “costatum”/“mutabile” (8 ribs), Gadila benoisti

houthalenensis, Dischides subpolitus and other species (cf. A.W. Janssen, 1984). In the Antwerp Sands

like A. exlamarcki and D. species n. subsp. Dischides subpolitus are absent, whereas “costatum”/“muta- bile” (10-11 ribs) and Gadila benoisti houthalenensis persist. In addition Gadila dingdenensis occurs for the

first time. In the Reinbekian the only remaining species are Gadila dingdenensis and D. “costatum”/

is for first time. Thus considerable “mutabile and one species, D. “dollfusi ” introduced the a ”, auct.,

impoverishment takes place, which continues during the Langenfeldian. The large and conspicuous

is the characteristic of this Two species “badenseÆ auct. (= floratum) only species stage. further

and Pulsellum introduced the species, Polyschides sp. sp. (both undescribed) are during Langenfeldian and the Gramian and the range through Syltian.

A common and conspicuous element in Miocene faunas is the species Pseudantalis aturensis

with smooth and tube. This misidentified (Cossmann & Peyrot), a polished species was previously as

D. entalis which aturensis from the Hemmoorian Linne, is a Recent species. P. occurs until the

and has therefore value for Langenfeldian no special stratigraphical purposes.

The results on the stratigraphical distribution of the Miocene species are summarized in fig. 2. 178

jmasi ssp.

n D.dt n.sp.

um )isti 'nensis Pulsel) G.benc houthale

enensis , G.dinge n.sp. marcki D. Ldes A.exla n.ss Polysch olitus D.subp

. ratum P ratum .sp. 3 F.flo n.s F.flo F.n

.sp. -11) -11) 3) (f (10- L.n (10 sleurum L !

! y ol -P

F 1 A.dollfusi 1 scaphopods. rensis

ta P.atu • Miocene Sp sp?

of A.n A.gerr n.s distribution Antwerpen Miste Houthalen Edegem SYLT GRAM LANGENFELDE REINBEK HEMMOOR VIERLAND NEOCHATT Stratigraphical 1

Aquitan. 2. Messin. 1 Torton. Serravallian Langhian Burdigalian

1 Fig. 179

SOME GENERAL CONSIDERATIONS

The scaphopod faunas of the Oligo-Miocene in the North Sea Basin developed from the comparative- ly rich assemblage of the Early Oligocene. The lineages of several species continue through Middle and Late Oligocene times. At the beginning of the Rupelian a conspicuous innovation of the fauna

of the fauna is observed the Chattian the occurs, whereas an impoverishment during and Vier-

in which elements introduced. It is in the Hemmoorian that landian, hardly any genuine new are only

lot of show These forms character the Hemmoorian again a new species up. new give a special to

connection the North Sea Basin and the Basin in SW fauna, demonstrating a between Aquitaine

France. During the Reinbekian there is again a process of species reduction, continuing until the Late

Miocene. The Late Miocene (Langenfeldian to Syltian) is characterized by a quite independant fauna rather in of these has its in the Reinbekian. There relation- poor species. Only one roots are no ships with the faunaof the northern Pliocene. The previously supposed connections with the Miocene of the Paratethys (Vienna Basin), as inferred by some species identifications, cannot be substantiated

the basis of on Scaphopoda.

NOTES

1. Fissidentalium novaki (von Koenen). Populations from the Middle Oligocene can be distinguished

subspecifically from typical Early Oligocene novaki by details of the sculpture. The form from the Berg Sands

has been identified up to now as acutum. The real Fissidentalium acutum (Hebert), however, is restricted to the

Latdorfianand clearly distinguishable from novaki by details of the sculpture.

2. Antalis This is the in the Meeressand in the identified as kickxii n. sp. species present Mayence Basin, by

be from kickxii and the of Sandberger and laterauthors. It can distinguished by a differing sculpture presence

an apical notch.

different of 3. The true Fissidentalium badense (Homes) from the Vienna Basin has a very type sculpture,

especially in the juvenile stage, and lacks the distinct secondary striation between the primary ribs. To the

North Sea Basin species the namefloratum Zimmermanncan be applied.

from the in this from characteristic of 4. Sediment extracted scaphopods sample yielded, apart a fragment

foraminiferal identified Dr C.H. to Carinastarte, a typical Langenfeldian assemblage, kindly by von Daniels,

whom I am grateful for his cooperation.

5. The Reinbekian species exhibits a striking similarity, especially in some details of the sculpture, with the

dimensions and the of Late Miocenefloratum. It can be distinguished by its much smaller presence an apical

slit. Its sculpture is completely different from that of the true dollfusi from the Edegem Sands.

6. This form is characterized by its sculpture, consisting of either 8 or 10-11 sharp-edged primary ribs and

regularly developing secondary riblets. It cannot be connected with the real mutabile from the Vienna Basin

is the in different Miocene or with costatum Sowerby from the British Pliocene. Very striking occurrence,

horizons of the Hemmoorian, oftwo forms with a constantly differing numberofribs (8 or 10-11). As further

differentiating characteristics are not present, and in the absence ofanobvious evolutionary trend, these two

forms are not considered to represent subspecies.

7. Goddeeris (1978) unfortunately used some nomina nuda, the identity of which can only indirectly be infer-

with the two forms octo- red from his notes on occurrence and frequency. So it seems that his D. antverpiense,

“costatum”/“mutabile” D. be resp. decemstriatum, equals my Lentigodentalium n. sp. auct.). decagonum might

is exlamarcki dumasi n. subsp. and D. houthalenensis perhaps n. subsp. 180

ACKNOWLEDGEMENTS

My thanks are due to all colleagues who supported the taxonomical revision by the loan of material.

I have thank A.W. Especially to my friend Janssen (Rijksmuseum van Geologie en Mineralogie,

for the loan of substantial material from Leiden), a very Belgium and for reading the English text of

this Dr S. Ritzkowski paper, (Geologisch-Palaontologisches Institut, Gottingen), for making

the available the scaphopod material of von Koenen collection (including all the types), and Mrs Dr

E. Kuster-Wendenburg (Ubersee-Museum, Bremen) for the loan of material from the Baumann col-

Messrs F. Weinbrecht and K. Giirs also lection. (Gliicksburg) (Frankfurt am Main) provided in-

and valuable from their for which I Further- teresting samples private collections, am very grateful.

more I have to thank colleagues B. Sliggers (Rijks Geologische Dienst, Haarlem), for providing the

scaphopod samples from the Broekhuizenvorst bore-hole, and Dr C.H. von Daniels

(Niedersachsisches Landesamt fur Bodenforschung, Hannover), for his identification of a

foraminiferal assemblage.

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Manuscript received 10 March 1987, revised version accepted 2 April 1987.