Upper Cretaceous Belemnites from the Vomb Trough in Scania, Sweden

Home , Actinocamax, Belemnella, Belemnellocamax, Belemnitella, Kristianstad Basin, Pachydiscus

WALTER KEGEL CHRISTENSEN

UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH IN SCANIA, SWEDEN

UPPSALA 1986

SVERIGES GEOLOGISKA UNDERSÖKNING

SER. Ca AVHANDLINGAR OCH UPPSA TSER I A4 NR 57

WALTER KEGEL CHRISTENSEN

UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH IN SCANIA, SWEDEN

UPPSALA 1986 ISBN 91-7158-353-X ISSN 0348-1352

In memory of the outstanding Swedish palaeontologist and biostratigrapher Johan Christian Moberg (1854-1915) who monographed the Upper Cretaceous belemnites of Sweden a century ago in 1885. Moberg used the belemnites biostrati graphically and propose d a partly new subdivision of the Upper Cretaceous. The publication of the monograph was the culmi nation of Moberg's first field of activity: the Cretaceous of Sweden. Later, he mainly focussed on the Cambro-Silurian of Scania. Seen in a European context, Moberg's study was only the third monograph dealing with Upper Cretaceous belem nites, following monographs on the English belemnites by Sharpe (1853) and German belemnites by Schliiter (1876).

A contribution to IU S PROJECf TORNQUIST UNES O (IGCP Accession Number 86)

Textkartorna är från sekretessynpunkt godkända för spridning. Lantmäteriverket 1986-03-26

Address:

Walter Kegel Christensen GeologicaJ Museum 0ster Voldgade 5-7 DK-1350 Capenhagen

Fotosats: ORD & FORM AB Tryck: Offsetcenter AB, Uppsala 1986 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TRO UGH

CONTENTS

Abstract ...... 3 ActinocamaxMiller ...... 23 Introduction ...... 4 A. verusMiller ...... 23 Material ...... 5 Gonioteuthis Bayle ...... 24 General geologic framework ...... 5 G. westfalica (Schliiter) ...... 25 The Vom b Trough ...... 6 G. westfalicagranulata (Stolley) ...... 25 Description of Iocalities ...... 7 G. granulataquadrata (Stolley) ...... 26 Eriksdal ...... 7 G. quadratascaniensis?Christense n ...... 27 Gonioteuthisspp...... 27 ~~~~~~:ga . ::::::::::::::::: ::::: ::: : ::::::::: :::::::::::::::::::::::: :: :::: ~ Belemnellocamax Naidin ...... 27 Kullemölla-Lyckås ...... 10 B. exgr.grossouvrei(Janet) ...... 28 Kullemölla l boring ...... 11 B. m. mammillatus(Ni!sson) ...... 29 Rödmölla- Tosterup ...... 11 B. balsvikensis (Brotzen) ...... 29 Summarizing remarks ...... 13 Belemnitella d'Orbigny...... 30 Stratigraphy ...... 13 B.propinquagroup ...... 30 Generaloutline ...... 13 'Actinocamax' cf.lundgreni Stolley...... 30 The subdivision of the Upper Cretaceous ofSweden ...... 14 B. p. propinqua(Moberg) ...... 30 Stratigraphic conclusions ...... 15 B. praecursor group...... 31 Depositional history of the Vomb Trough during the Upper B. alphaNaidin ...... 32 Cretaceous ...... 18 B. ex gr. alp ha NaidinlpraecursorStolley ...... 34 Systematic part ...... 19 Belemnitella taxa from the highest Lower Campanian- Morphologyofthe guard and terminology ...... 19 Lower Maastrichtian ...... 34 Measurements and abbreviations ...... 19 B. mucronata (Schlotheim) ...... 35 Biometric methods ...... 20 B. aff.langeiJe!etzky ...... 36 Classification ofBelemnitellidae ...... 20 Summary ...... 38 The species concept in belemnites ...... 20 Acknowledgments ...... 39 Family Belemnitellidae ...... 22 References ...... 39

ABSTRACT

Christensen, W.K., 1986: Upper Cretaceous belemnites from the taxa from the uppermost Lower Campanian-Lower Maastrich Vomb Trough in Scania, Sweden. Sveriges geologiska undersökning. tian are discussed and the concept of the majority of these taxa Ser. Ca, No. 57, pp. 1-57. Pis. 1-7. Uppsala 1986. is shown to be in a state of chaos. Manuscript delivered in January 1985 . The localities in the Vomb Trough are described and placed Upper Cretaceous belerunites from classic outcrops and one in an international stratigraphic fraruework on the basis of the boring in the Vomb Trough in Scania , Sweden, are described, belerunites faunas, which are compared to faunas from western including univariate and bivariate biometric analyses. Europe and the Russian Platform. The Vomb Trough is a narrow, elongated, asymmetrical The 'Westfalicus conglomerate' at Rödmölla-Tosterup graben initiated in early Mesozoic times. The maximum thick yielded Belemnitella propinqua and Gonioteuthis westfalica and ness of the Upper Cretaceous strata is a little less than 800 m, is from the Lower-Middle Santonian. The mar! at Eriksdal is and the dominant sediments are glauconitic, calcareous, clayey referred to the Upper Middle Santonian on the basis of Belem siltstones deposited in a fairly shallow-water marine environ nitella propinqua and Gonioteuthis westfalicagranulata. Lower ment. Several conglomerates with extra-basinal clasts and non Middle Santonian strata with Gonioteuthis westfalica are possi sequences occur in the Santonian and Campania n, at !east close bly also present at Eriksdal. The outcrops at Kullemölla-Lyckås to the margin of the trough. contain Belemnitella alpha, Gonioteuthis granulataquadrata, Thirteen belemnite taxa from the Santonian-Campanian are Actinocamax verus, and Belemnellocamax ex gr. grossouvrei, recognized representing the following genera: Actinocamax and are assigned to the basal Lower Campanian. The loose Miller, Gonioteuthis Bayle, Belemnellocamax Naidin, and blocks at Kåseberga contai ning Belemnitella ex gr. alphal Belemnitella d'Orbigny. The nominate subspecies and two praecursor and Gonioteuthis granulataquadrata or Gonioteuthis other subspecies of Actinocamax verus are discussed; the sub quadrata are from the Lower Campanian, possibly the basal species dnestrensis is placed in synonymy of the nomina te sub Lower Campanian. The 'Verus conglomerate' at Rödmölla• species, and i t is suggested that the subspecies fragilis ma y be a Tosterup is sandwiehed between beds from the basal Santonian geographic subspecies. Actinocamax (Paractinocamax) is dis and the uppermost Lower Campanian, and ma y therefore be of cussed and placed in synonymy of Belemnellocamax. Two taxa middle to late Santonian and/or early Early Campanian age. of Belemnitella, B. alphaand B. praecursor, from the lower and The calcareous sandstones and conglomerates (Tosterup con middle Lower Campanian are tentatively recognized and fully gloruerate) at Rödmölla-Tosterup span the Lower-Upper Cam discussed. Measurements of critical interna! characters of the panian boundary. The beds with Belemnellocamax holotype of Belemnitella praecursor are reported. Belemnitella mammillatus, Belemnitella mucronata, and Gonioteuthis quad-

3 WALTER KEGEL CHRISTENSEN rata scaniensis? are from the uppermost Lower Campanian, propinqua, and Belemnitella ex gr. alpha/praecursor. The previ whilst the beds with Belemnellocamax balsvikensis are from the ous biostratigraphical dating of the upper part of the boring lower Upper Campanian. The lower part of the Köpinge using foraminifera is supportedon belemnite evidence. Maas sandstone is referred to the uppermost Lower Campanian on trichtian belerunites have not been met with, although Maas the basis of Belemnellocamax mammillatus and Belemnitella trichtian ammonites and foraminifera are known from the mucronata, whereas the upper part is assigned to the middle trough. Earlier records of stratigraphically important fossils, Upper Campanian on the basis of Belemnitella mucronata and including ammonites and echinoderms, are reviewed. Belemnitella aff. langei. The following belemnite species from The present belemnite study has thus demonstrated the pre the Santonian-basal Lower Campanian are recorded from the sence of Upper Cretaceous strata from the Lower-Middle San Kullemölla boring: Actinocamax verus, 'Actinocamax' lund tonian, basal Lower Campanian, uppermost Lower Campa greni, Gonioteuthis westfalica, Gonioteuthis spp., Belemnitella nian-lower Upper Campanian, and middle Upper Campanian.

Introduction

Upper Cretaceous belerunites from Sweden were studied (1910), Lundegren (1930 to 1936), Brotzen (1936 to already in the first half of the 19th century by Wahlenberg 1966), and Christensen (1970 to 1982). Other strati (1818), Nilsson (1826, 1827), Blainville (1825-1827, graphically important fossil groups such as ammonites 1827), and d'Orbigny (1840); these authors, however, and inoceramid bivalves have been recorded from the only recorded Belemnitella mucronata (Schlotheim) and Vomb Trough but, unfortunately, none of these groups Belemnellocamax mammillatus (Nilsson). have been revised recently. However, recent comments The first detailed study on the Swedish belerunites was by various authors on ammonites and inoceramid presentedin the classical monograph on the Upper Cre bivalves described from the area by earlier workers are taceous cephalopods of Sweden by Moberg (1885). In summarized here (see seetian on description of this work he described nine species, forms, and varieties localities). Lastly, it should be ad de d that a significant of belemnites, which were assigned to the genera description of the foraminiferal fauna from Eriksdal in Actinocamax Miller and Belemnitella d'Orbigny. In 1894 the Vomb Trough was presented by Brotzen (1936). Moberg reviewed the fauna and commented briefly upon The histories of subdivision of Upper Cretaceous the taxa described earlier. Swedish Upper Cretaceous strata in Sweden and Germany are closely connected. belerunites have also been studied by Schliiter (1876), de The first reliable scheme was that of Schliiter (1870). Morgan (1882), Stolley (1897, 1930), Jeletzky (1949b) Stolley (1897) subdivided the Senanian of Germany and and, more recently, by the present author (Christensen Sweden, mainly on the basis of belemnites, into (top to 1970 to 1982). bottom): Mucronatenkreide, Quadratenkreide ( = Christensen (1975a) monographed the Upper Cretace Mammillatenkreide), Granulatenkreide, and Westfalicus ous belerunites from the Kristianstad Basin in Scania on kreide. This classification was used subsequently in Swe the basis of new and old collections. In the present paper den by most authors, even as late as Bergström & Sund the belerunites from the Vom b Trough are monographed quist (1978), but translated into: mucronatakrita, mainly on the basis of collections made in the last part of mammillatuskrita, granulatuskrita, and westfalicuskrita. the 19th century and the beginning of the 20th century. Christensen (1985) reviewed the Albian to Maastrichtian The faunas are campared to and correlated with belem of southern Sweden and Bornholm. nite faunas from other parts of Europe, and the outcrops During the Upper Cretaceous, two belemnite subpro are placed in an international stratigraphic framework. vinces have been recognized within the North European In the Swedish Upper Cretaceous, belemnites, above Province (Christ~nsen 1975a, 1976, 1982; see Fig. 11.) all other macrofossils, are of fundamental importance in The Balto-Seandian belemnite faunas can be considered biostratigraphy and correlation. This is also the case for as mixtures of the faunas found in the Central Russian the Upper Cretaceous of the Vom b Trough. The present and Central European Subprovinces, respectively. They belemnite stratigraphy is the result of progressive refine are, therefore, very important from a stratigraphic point ments by a series of workers, notably Schliiter (1870 to of view, because the y provide a basis for earrelation be 1897), De Geer (1881 to 1889), Lundgren (1876 to 1895), tween the two subprovinces (cf. Fig. 8). Moberg (1881 to 1910), Stolley (1897 to 1932), Hennig

4 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

Material

The majority of the belerunites studied we re placedat my century. The collections of J.C. Moberg and A. Lunde disposal from the following institutions: Geological Sur gren constitute a substantial part of the studied material, vey of Sweden, Uppsala (SGU); Swedish Museum of and the belerunites figured by Moberg (1885) are also Natural History, Section of Palaeozoology, Stockholm included in the stud y. (RM); Geological Institute, University of Lund (LM); The outcrops in the Vom b Trough are not accessible or Geological Institute, University of Stockholm (GI); and very badly exposed today. I have, therefore, only been Geological Museum, University of Capenhagen able to make new belemnite collections from one local (MGUH). These collections generally were made in the ity, Rödmölla at Tosterup. last part of the 19th century and the beginning of the 20th

General geologic framework

During the Mesozoic the block-faulted NW-SE trending ziemmanouil1982), and the thickness of the Upper Cre Fennoscandian Border Zone acted as a transition zone taceous was found to be about 785 m in the Köpingsberg between the s table Fennoscandian Shield and the subsid boring (Anderegg, Norting & Skoglund 1968; Chat ing Permo-Mesozoic Danish Subbasin, which is part of ziemmanouil 1982). Limestones, including chalk and the Danish-Potish Trough. The border zone in Scania is marty chalk, predominate in the Malmö area, and the characterized by grabens and horsts trending NW-SE and thickness of the Upper Cretaceous is about 1640 m (Nor is erossed by faults with a main direction NNE-SSW, ting & Skoglund 1977). The thickness of the Upper Cre creating a mosaic of tilted blocks (Figs 1-2). taceous stages in various areas in southern Sweden is Late Lower to Upper Cretaceous deposits are found shown in Fig. 3. northeast of the border zone in the Kristianstad and Båstad Basins and at Särdal, further north along the Swedish west coast. Within the border zone, sediments are recorded from the Vomb Trough and the istand of Bornholm. Southwest of the border zone sediments are known from the Malmö area which, from a tectonic and sedimentologic point of vie w, betongs to the Dan is h Sub basin. The tithology and thickness of the late Lower to Upper Cretaceous strata are highly variable within the different areas. Biocalcarenites and glauconitic sandstones prevail in the basins northeast of the border zone, and the thick ness of the strata is lessthan200m (Christensen 1975a). In two borings in the southern part of the Hanö Bay the thickness of the Cenomanian-Campanian is about 540 m, and sandstonesand calcarenites predominate (Norling &

Skoglund 1977; Norting 1981; Norting in Kumpas 1980). o 50 km .l In the Vomb Trough the Upper Cretaceous sedimentary deposits are characterized by detritat clastic sediments Fig. l. Main structural trends in southern Sweden and adj acent areas. such as glauconitic calcareous, clayey siltstones (Chat- Modified from Gravesen, Rolle & Surlyk (1982).

5 WALTER KEGEL CHRISTENSEN

FEN NOSCAN DIAN \ \ SHIELD \ 1 \ \ \. \ \. \. \. \. \. \. ~~ o~ ', HANÖ BAY <>~ ' BALTIC ~ '$> ' ' SEA ' " " ' " " ' " • Simrishamn ' " " ' ' " ' ' " ' ' Upper Cretaceous ' ' !:5:9 Danian o 25km e Town l l

Fig. 2. Map showing the distribution of the Upper Cretaceous and Danian at the base of the Quaternary, and the approximate location of the Fennoscandian Border Zone. Modified from Gry (1960) and Bergström et al. (1982).

cally active during the Mesozoic and the thickness of the The Vomb Trough Upper Cretaceous strata varies due to differential subsi Norling (1982) and Chatziemmanouil (1982) described dence. The greatest thickness is found in the southeast the trough as a narrow, elongated, asymmetrical graben ern part of the trough (.A.nderegg et al. 1968; Chat which was initiated in the early Mesozoic (Fig. 4) . The ziemmanouil 1982). length of the trough is about 80 km; its width is about 7 Chatziemmanouil (1982) analysed the Upper Cretace km towards the northwest and 11 km towards the south ous of the trough on the basis of five bo rings placed al o ng east. The trough is bounded to the northeast by the large the axis of the trough. The borings were dated by means scale Fyledalen Fault and to the southwest by the of foraminifera, and all Upper Cretaceous stages were Romeleåsen Horst, a Precambrian granitic fault block. recognized, although Lower Maastrichtian deposits were The Herrestad Uplift, an east-west trending horst in the only found in the Köpingsberg borings immediately southern part of the trough, divides the trough into two south of the Herrestad Uplift. parts. According to Chatziemmanouil (1982) the tectonic Several informallithostratigraphic units, such as Eriks elements bordering the trough: the Romeleåsen Horst, dal mar!, Kullemölla mar! , Lyckås mar!, Kåseberga mar! , Herrestad Uplift, and the Lower Palaeozoic Plateau Köpinge sandstone, Valleberga sandstone, Rödmölla northeast of the trough, were important source areas conglomerate, and Tosterup conglomerate, have been during the Upper Cretaceous. The trough was tectoni- established for various sedimentary rocks in the trough in

6 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

grafiska kartan över Sverige (Topographical maps of c Sweden), scale 1:50.000. Thickness c Kristianstad Basin 18 - - 55 105 6 1885; Hägg 1930, 1935 , 1954; and Lundegren 1935b). ------Hanö Bay 75 25 40 400 50 - The Swedish materialofthese groups needs revision, bu t ------same recent published comments are included here. ? - Bås ta d Bas i n 12 - - ? The localities are Iisted in alphabetical order. ------50- Vom b Trough 60 90 85 390 160 50 ------Eriksdal Malmö area 65 50- 75- 300- 350- 500- 75 100 350 400 700 ------Moberg (1882b, 1884, 1910a) indicated two marl-pits at 85 20 180 Bornholm (?) Eriksdal (Fig. 5) , the geographical coordinates of which are: Fig. 3. Thickness of Albian-Maastrichtian stages in various areas in southern Sweden and Bornholm, Denmark. Sources: Christensen Eriksdal A- 10 Y stad NV/20 Tornelilla SV; VB 251594 (1975a), Norling & Skoglund (1977) , Norling (1981) , and Chat Eriksdal B -10 Ystad NV/20 Tornelilla SV; VB 256593 ziemmanouil (1982). According to Moberg (1884) about 10 m of greyish yellow sandy mar! with a conglomeratic bed about 35 cm thick in the middle part of the seetian were exposed at the last part of the 19th and the beginning of the 20th E riksdal B; the sedimentary strata dipped a bo ut 20° centuries. Som e of the se informal u nits have been u sed as towards southwest. Moberg reported that fossils occur formal lithostratigraphic units by later authors. How red very commonly in the sediments. Lundegren (1934, ever, these u nits are very bad! y de fin ed or not defined at 1935a, in Brotzen 1936) mentioned that he had cleaned all, and they are, therefore, used informally in the pre the seetian at Eriksdal B in 1931 and collected fossils bed sent paper. by-bed. Brotzen (1960: 11) claimed that the pit was filled up with Quaternary sand and grave!. This statement is, Description of localities however, not correct. When I visit ed the o uterops in 1979, they were covered by scree and overgrown but can Belerunites from five localities and one boring were easily be cleaned. I succeeded in collecting same few studied (Fig. 4). All outcrops are situated in the north belerunites from the topmost part of the sequence at eastern part of the trough close to the Fyledalen Fault. Eriksdal B. Moberg (1884) did not comment upon Eriks Most of the belemnite material placed at my disposal is dal A . labelled on! y with the name of the outcrop, and the exact The belerunites exaroined in the course of the present harizon within the seetian is not indicated. This fact study originate by and !arge from the collections made by el e ar! y reduces the stratigraphic potential of the material. J.C. Moberg in the last century and by A. Lundegren in However, most outcrops on ly expose som e fe w me tres of 1931. The following taxa are recognized: Gonioteuthis sediments, which can be referred to a specific belemnite westfalicagranulata (Stolley), Belemnitella propinqua zone. The outcrops are not accessible or are very badly propinqua (Moberg), and Actinocamax ve rus Miller. The exposed today, and consequently i t was necessary to re! y locality can thus be referred to the Upper Middle Santo on earlier worksin order to describe the localities. I have nian on belemnite evidence. lt should, however, be especially relied upon the studies by Lundgren (1876, added that in addition to G. westfalicagranulata, same 1880), Moberg (1881, 1882a, b, 1884, 1910a, b) , De Mor tew westfalica-like specimens with a relatively shallow gan (1882), Hadding (1927), Hägg (1930, 1935, 1954), pseudoalveolus occur at Eriksdal (see p. 26). These Lundegren (1933a,b, 1935a,b) and Brotzen (1938, 1945, specimens ma y indicate that strata from the Lower Midd 1958) . Recent! y, the o uterops at Rödmölla-Tosterup le Santonian also are present at Eriksdal. were discussed in great detail by Gravesen (1977) . Hägg (1930:69-70) and Stolley (1930:168) recorded The outcrops are shown in Figs 4-6 and, where possi the crinoid Uintacrinus from Eriksdal and, according to ble, their coordinates are given according to Topo- both authors, this should indicate the presence of the

7 WALTER KEGEL CHRISTENSEN

N D Danian

l1 l l Upper Cretaceous 1 • Locality • Town o Bore-hole

Fig. 4. Map of the Upper Cretaceous of the Vom b Trough at the base of the Quaternary showing outcrops and bore-holes. Modified from Bergström et al. (1982).

Uintacrinus Zone from the Lower Upper Santonian. It Therefore, it is not necessary to infer the presence of has been shown, however, by Ulbrich (1971) and Ernst & Upper Santonian rocks at Eriksdal. Schulz (1974) that Uintacrinus occurs bothin the Uinta Lastly, i t should be mentioned that the monograph of crinus/G. granulara Zone from the Lower Upper Santo Brotzen (1936) on the foraminiferal fauna from Eriksdal nian and the subjacent G. westfalicagranulata Zone from was based on material collected by A. Lundegren in 1931 the Upper Middle Santonian. from Eriksdal B. According to Birkelund & Bromley (1979) one specimen of Hauericeras cf. pseudogardeni (Schluter) has been f o und at Eriksdal; the specimen was figured as Ammonites n.sp.? by Moberg (1885 , PI. 2:7). H. Kåseberga pseudogardeni occurs in the Munster Basin in the so called 'Santon 4' and 'Campan l' of Arnold (1964), and Loose blocks consisting of light, porous, yellow to grey these units were correlated by Ernst (1968 , Table l) with ish-yellow marlstones from the Upper Cretaceous are the Marsupites/G . granulara Zone of the Upper Upper known from immediately west of the small village of Santonian and the G. granulataquadrata and/. lingua/G. Kåseberga at the sea-shore on the south coast (Lundgren quadrata Zones of the lowermost Campanian (Fig. 9). 1880; Moberg 1882a; Lundegren 1935b; and Hägg 1935 , The presence of H. cf. pseudogardeni may thus indicate 1939). the presence of Upper Santonian rocks at Eriksdal. How Hägg (1939) mentioned that he had collected two ever, Ulbrich (1971) recorded H. cf. pseudogardendram belemnites from a loose block, and they were referred to the G. westfalicagranulata Zone and the lower G. as Gonioteuthis quadrata (Blainville) and Belemnitella granulara Zone of the German Demoeratic Republic. mucronata (Schlotheim). I have examined the two speci-

8 UPPER CRET ACEOUS BELEMNJTES FROM THE VOMB TROUGH

century and the first third of the 20th century. Stolley ~~ (1897) subdivided the 'Mucronatenkreide', mainly on the l~ERIKSDAL basis of ammonites, in to three zones (bottom to top): a \\ A• "B zone with Hoplitoplacenticeras coesfeldiense, a zone with Bostrychoceras polyplocum, and a zone with Scaphites constrictus. Hägg (1930) and Lundegren (1933a) sug *Mill gested, on the basis of the ammonites recorded from the sandstone, that the sandstone apparently embraces all + Church D KULLEMOLLA BENE· C~*-E l STAD three zones. [ Building Lundegren (1933a) studied the Köpinge sandstone along the logelstorp brook between Valleberga and Ingelstorp. Towards the east the sandstone was exposed O 23kmBAL~t~\l from c. 500 m northeast of the Valleberga Church to c. 500 m northwest of the church, and Lundegren recorded ~~--~~~ HOGEST~ Belemnellocamax mammillatus from this area. He conse quently referred the sandstone in this area to the Fig. 5. Map showing outcrops at Eriksdal, Kullemölla, and Lyckås. After Moberg (1884). mammillatus Zone and showed that it was older than assumed earlier. Hägg (1930: 13) had collected B. mammillatus in situ at Valleberga Church already in mens and another specimen of Gonioteuthis in the collec 1913, but apparently he did not realisethe importance of tion of the GeologicaJ Institute, Stockholm University. the discovery. Hägg (1954) restricted the name Köpinge The specimen referred to as B. mucronata by Hägg is sandstone to sandstone of mucronata Zon e a ge as u sed by here assigned to B. ex gr. alpha/praecursor (see discus Stolley (1897 , 1930), while similar sandstones of sion below). A precise determination of the two speci mammillatus Zone age sensu Stolley were referred to as mens of Gonioteuthis is not feasible, but they are tenta Valleberga sandstone. tively ascribed to G. granulataquadrata (Stolley) or G. According to Lundegren (1933a) no outcrops were quadrata (see discussion below). found from c. 500 m northwest of Valleberga Church to c. The Ioose blocks yielding the belemnites mentioned 600 m towards northwest. From this point and c. 1400 m above are assigned to the Iower part of the Lower Cam towards west-southwest of logelstorp Church, Lunde panian on belemnite evidence. gren recorded only Belemnitella mucronata from the Ammonites and inoceramid bivalves were recorded sandstone; it was therefore assigned to the mucronata from the Kåseberga blocks by earlier authors, but this Zone sensu Stolley. Lundegren mentioned that pebbles material needs revision. of Silurian shales were found throughout the sequence. Brotzen (1938) , who visited the outcrops between Val leberga and logelstorp in 1937, contirmed the observa tions of Lundegren, and mentioned that he had seen Ioose blocks consisting of typical Tosterup conglomera te Köpinge (see below) in this area. In the southeastern part of the Vom b Trough south ofthe The majority of the fossil material (including belem Herrestad Up lift , a yellow, high! y calcareous sandstone nites) kept in Swedish geological institutions is on ly la bel rich in glauconite, the so-called Köpinge sandstone, has led 'Köpinge', and this fact obviously reduces the strati been recorded frommanysmall outcrops, the majority of graphic val u e of the material. However, in addition to which are not accessible today. The sandstone was found, belemnites only labelled 'Köpinge', I have studied notably, northeast of Y stad at Köpinge (Fig. 4). The belemnites collected by A. Lundegren in 1932, F. Brot Köpinge sandstone and its fauna were dealt with particu zen in 1937, and T. Birkelund and F. Brotzen in 1960 lady by Schliiter (1870, 1876), Lundgren (1876) , Moberg from the sandstone between Valleberga and Ingelstorp. (1884, 1885), Stolley (1896, 1897), Grönwall (1912) , In the material generally la belled 'Köpinge' two taxa of Lundegren (1933a) , Brotzen (1938, 1945, 1966) , and Belemnitella are recognized: B. aff. langei Jeletzky and Hägg (1954), who also Iisted the outcrops. B. mucronata (Schlotheim). B. aff.langei is the dominant The Köpinge sandstone was referred to the 'Mu form and wasfiguredas B. mucronata by Moberg (1885, cronatenkreide' by authors in the last part of the 19th PI. 6:14-16, 19). The Köpinge sandstone yielding these

9 WALTER KEGEL CHRISTENSEN taxa probably belongs to the middle Upper Campanian, Ancyloceras? elegans Moberg was figured by Moberg viz. the B. mucronata Zone and possibly the basal B. (1885 , PI. 3:10). According to Blaskiewicz (1980:28-29) 'minor' Zone (see below). this specimen may be referred to Neancyloceras The Köpinge sandstone in the eastern part of the Val phaleratum (Griepenkerl), a species which Blaskiewicz leberga-Ingelstorp area yielded Belemnellocamax used as an index fossil for the lowermost zone of the mammillatus mammillatus (Nilsson) and Belemnitella Upper Campanian in Poland. This zone also yielded T. mucronata (Schlotheim) and belongs to the uppermost spiniger. Lower Campanian. Moberg (1885, PI. 3:1) figured Ammonites oldhami The Köpinge sandstone in the western part of the Sharpe. This specimen should be assigned to Pachydiscus Valleberga-Ingelstorp area yielded B. mucronata and B. koeneni Grossouvre which occurs in the N. phaleratum aff. langei. Both taxa occur throughout the sedimentary and B. polyplocum Zones in Poland (Blaskiewicz sequence; B. mucronata is the dominant species in the 1980:42). lower part of the sequence, w h ile B. aff. langei prevails in Moberg (1885) also recorded Hoploscaphites eonstric the upper part of the sequence. The Köpinge sandstone tus (Sowerby) and Anagaudryceras cf. lueneburgense in this area can thus be referred to the middle Upper (Schhiter) from the Köpinge sandstone. The presence of Campanian. these two species ma y indicate that part of the sandstone Belemnellocamax balsvikensis (Brotzen), w hi ch occurs is from the Maastrichtian. The specimens, however, may in the basal Upper Campanian, has not been recorded have been collected from loose blocks in the Köpinge from the Köpinge sandstone. Moreover, Maastrichtian area. In this connection it is of interest that Chat belerunites are also unknown from the sandstone. It can ziemmanouil (1982) recorded lower Maastrichtian strata thus be concluded that the Köpinge sandstone on belem from the Köpingsberg boring, which is situated in the ni te evidence can be assigned partly to the uppermost former area of outcrop of the Köpinge sandstone. Lower Campanian and partly to the middle Upper Cam Stokes (1975) reported Micraster glyphus (Schliiter) panian. from Köpinge. M. ex gr. schroederilglyphus occurs in Species of ammonites and echinoids of biostratigraphic Germany from the B. mucronata/G. quadrata gracilis value, recorded from the Köpinge sandstone by earlier Zone to the P. stobaei!G. papillosa basiplana Zone workers, have been the subject of published comments (Ernst & Schulz 1980). Iisted bel o w. Stokes (1979) recorded Diplodetus idae (Cotteau) Pachydiscus stobaei (Nilsson) was figured by Moberg from Köpinge and mentioned that specimens of this (1885 , PI. 2: 1-2, 5) . This species appears in Germany in species were variously labelled as Micraster coran the uppermost Lower Campanian B. mucronata/G. guinum, M. corbovis, M. idae, and Brissopsis? cretacea. quadrata gracilis Zone, continues inta the G. vulgaris Zone, and is used as an index fossil for the P. stobaei!G. papillosa basiplana Zone in the lower part of the Upper Campanian (Khosrovschahian 1972). Trachyscaphites spiniger (Schliiter) was figured by Moberg (1885, PI. 3:6). This species occurs in Germany Kullemölla-Lyckås in the lower part of the Upper Campanian from the P. stobaei!G. papillosa basiplana Zone to the G. vulgaris Moberg (1882b, 1884) indicated seven outcrops at Zone (Schmid & Ernst 1975; Ernst, Schmid & Klischies Kullemölla and on e o uterop at Lyck ås (Fig. 5) . Accord 1979). T. spiniger seems to have the same vertical range ing to Moberg (1884) the mar! pit at Lyckås (locality I) as the important guide fossil Hoplitoplacenticeras vari showed the same sequence as the !arge marl pit referred (cf. Schmid & Ernst 1975). to as Kullemölla D (Stora Grafven). About 8m of grey Bostrychoceras (Mobergoceras) junior (Moberg) was ish-yellow sandy marl which dipped about 30° towards figured by Moberg (1885 , PI. 3:14) . This species occurs in northwest we re exposed at the latter locality. Locality Germany in the B. mucronata/G. quadrata gracilis Zone Kullemölla E (Qvarnrännan) exposed a slightly deeper (Schmid & Ernst 1975). According to Moberg the speci part of the sequence than Kullemölla D. The other pits at men came from the B. mucronata Zone as used earlier. l t Kullemölla only showed seetians of small thicknesses, should be stressed, however, that the Köpinge sandstone and they were not treated in detail by Moberg. Today the was considered to belong only to the B. mucronata Zone outcrops are not accessible or are covered by scree and in the last part of the 19th century (see a bo ve). overgrown.

10 UPPER CRET ACEOUS BELEMNITES FROM THE VOMB TRO UGH

The outcrops at Kullemölla-Lyckås have yielded the westfalica by Lundegren (1935b) , came from the 440 m following belemnites: Gonioteuthis granulataquadrata, leve!. Belemnitella alpha, Actinocamax verus, and Belem Representatives of the genus Gonioteuthis we re found nellocamax ex gr. grossouvrei, and can thus be referred to in the 60-390 m in tervaL Lundegren (1935b) also re the basal Campanian, Zone of G. granulataquadrata. ported Gonioteuthis from the 390-440 m interval, but the A . Hennig (pers. comm. in Schliiter 1897:46) reported belemnite specimens found in this interval are not deter Marsupites from Lyckås. This record, however, was not minable at the genus leve! with one exception (see described in detail and has not been verified by subse a bo ve). The specimens of Gonioteuthis are weil-pre quent workers. The record of Marsupites at Lyckås is served and in some cases complete. They cannot, how therefore considered dubious for the time being. ever, be assigned with certainty to species, because it is necessary to analyse a certain number of specimens, sa y at !east 8-10, in order to make a reliable specific determi nation. Only the earliest member of the Gonioteuthis lineage , G. westfalica, is easily recognizable owing to the structure of its anterior and, and several specimens of this Kullemölla l boring taxon were identified from the 330-390 m interval. The boring was made in 1918-1919 and described by A single specimen of B. propinqua came from the 248 Gavelin (1919) and Lundegren (1933b , 1935b). It was m leve!; this specimen was referred to as B. mucronata by commented upon later by Brotzen (1945, 1958) . Lundegren (1935b:8). Three specimens of B. ex gr. Recently, Chatziemmanouil (1982) reexamined the alpha/praecursor came from the uppermost 20 m of the Upper Cretaceous of the boring, and four other borings boring; these specimens also were referred to as B. muc in the trough, from a sedimentologic and biostratigraphic ronata by Lundegren (1935b:8). point of vie w. Guy-Ohlson (1982) studied the palynologi The stratigraphic range of the belerunites found in the ca! content of the basal part of the bo ring and showed that Kullemölla l boring are discussed in the systematic part a major unconformity is present between the Lower of the paper. The biostratigraphic subdivision of the Jurassic and Lower Cretaceous. upper part of the boring by Chatziemmanouil (1982) can The drill-hole was placed c. 850m westsouthwest ofthe be supported on belemnite evidence. The uppermost Kullemölla Mill rather close to the outcrops at part of the boring yielded B. ex gr. alpha/praecursor and Kullemölla mentioned above (Figs. 4--5). A little more is thus from the lower Lower Campanian, as are the than 600 m of Upper Cretaceous sediments, rnaini y con sediments outcropping n ear by. sisting of glauconitic, calcareous siltstone, were found in the boring. Lundegren (1933b, 1935b) dated the Upper Cretace ous strata on the basis of belemnites, ammonites, and inoceramid bivalves. The belerunites are revised in the Rödmölla-Tosterup present paper; the two other groups still need revision. Chatziemmanouil (1982) subdivided the Upper Cretace The outcrops at Rödmölla-Tosterup (Fig. 6) were dealt ous of the boring on the basis of foraminifera and, with by Lundegren (1876) , Moberg (1881, 1884, 1888, according to this author, the 28-128 m in terval belongs to 1910b), de Morgan (1882) , Hadding (1927) , and more the Campanian, the 128-515 minterval to the Santonian, recently by Gravesen (1977). Gravesen reviewed in great the 515-575 m interval to the Coniacian, the 575-614 m detailliterature concerning Rödmölla-Tosterup and also interval to the Turonian, and the 614--639 m interval to provided new information about some of the outcrops. the Aptian-Cenomanian. The following outline relies heavily on his review. The following belerunites are recognized from the bor Moberg (1910b) distinguished five outcrops at Röd• ing: Actinocamax ve rus Miller, 'A '. cf. lundgreni Stolley, mölla-Tosterup, and they were referred to as CV:l-5. Gonioteuthis westfalica (Schliiter), G. spp., Belemnitella Three outcrops (CV: 1-3) were found in the southeastern propinqua (Moberg), and B. ex gr. alpha!praecursor. slope of the valley of the Rödmölla brook; locality CV:4 A . verus was found in the 40-360 m interval, and the was the !arge Tosterup mar! pit; and locality CV:5, situ species is most common in the 40-190 m interval. ated a little southwest of the mar! pit, was found in the Altogether, 42 specimens have been examin ed. A single western slope of the valley. At present, CV:l is very specimen of 'A.' cf. lundgreni, earlier referred to as G. badly exposed; CV:2 was reexcavated by P. Gravesen

11 WALTER KEGEL CHRISTENSEN and others (see Gravesen 1977); and at CV:3loose blocks of Tosterup conglomera te we re found recent! y in a culti N vated field (Gravesen 1977). CV:4 and CV:5 are not accessible today. The geographical coordinates are as follows: 1 Rödmölla CV:l- 10 Ystad N0/20 Tornelilla SO; VB 37054805, Rödmölla CV:2- 10 Ystad N0/20 Tornelilla SO; VB 36954800, *Mill Rödmölla CV:3- 10 Ystad N0/20 Tornelilla SO; VB 36854790. O 250 500m Locality CV: l showed former! y the contact between the Lower Jurassic and the Upper Cretaceous sequence. Fig. 6. Map showing outcrops at Rödmölla-Tosterup. After Moberg According to Moberg (1910b) the almost vertical Upper (1910b). Cretaceous sediments rested upon Lower Jurassic sandstone. The basal Cretaceous bed, the so-called 'Westfalicus conglomerate' of Hadding (1927), is a - coarse-grained, greyish-yellow sandstone rich in shell The 'Verus conglomerate' is followed upwards by cal- fragments with numerous small dark pebbles of Lower careous sandstones in which several conglomerates or Jurassic sedimentary rocks. Two hard beds each 50 cm conglomeratic beds are present. These conglomerates thick and approximately 2 m apart were present in the are characterized by their content of pebbles of Silurian lower part of the 'Westfalicus conglomera te'. Moberg shales (Colonus Shale) and Lower Jurassic sandstones (1910b) reported Gonioteuthis westfalica and (Hadding 1927), and were referred to as Tosterup con Actinocamax verus from this part of the sequence. I have gloruerate by Moberg (1884, 1910b) and 'Mammillatus examined specimens of G. westfalica labelled Tosterup 7 conglomerate' by Hadding (1927). In the former Tos by J.C. Moberg in 1887. On the basis of the adherent terup mar! pit (sketched by the Morgan 1882, Fig. 7), the matrix, it can be inferred with great probability that the calcareous sandstone was overlain by two conglomerates specimens came from the 'Westfalicus conglomerate'. approximately 40 cm apart, which in turn was succeeded Moreover, I have seen one specimen of Belemnitella upwards by calcareous sandstone; the sedimentary sequ propinqua from the conglomerate. On the basis of these ence dipped about 10° towards the southeast (de Morgan taxa the conglomerate can be assigned to the Lower 1882; Moberg 1910b). Moberg (1910b) mentioned that Middle Santonian. Moberg (1910b) mentioned that two Upper Cretaceous strata could be followed from south of hard beds, more or less similar to those mentioned the pit to the Rödmölla brook, and conglomerates and above, were present about 50 m to the south. These beds sandstones alternated along this stretch. Moberg (1910b) did not yield any fossils. recorded Pachydiscus stobaei, Trachyscaphites spiniger, The 'Westfalicus conglomerate' is succeeded upwards Scaphites roemeri?, in addition to Belemnellocamax by loose arenaceous beds which in turn are overlain by mammillatus and Belemnitella mucronata from the cal another conglomerate, the so-called 'Verus conglome careous sandstone underlying the conglomerates. He rate' of Hadding (1927) and Rödmölla conglomerate of also mentioned that the same fossils, by and !arge, occur Moberg (1910b). The 'Verus conglomerate' is rather red in the conglomerates. I have studied some few speci similar to the 'Westfalicus conglomerate', only differing mens of B. mammillatus from this pit, and the sediments by containing more and larger pebbles, which consist of can therefore be referred to the uppermost Lower Cam Lower Jurassic rocks, notably ferrugenious, dark brown, panian. fine-grained sandstones and day iron-stones (Hadding Moberg (1910b) also recorded the Tosterup conglom 1927). This conglomerate yielded A. verus (see Moberg erate from locality CV:2, and from this outcrop he 1910b ). The 'Verus conglomera te' and the subjacent and reported B. mammillatus and one specimen of superjacent arenaceous beds are less inclined than the 'Actinocamax quadratus Blainville var.'. This specimen is beds of the 'Westfalicus conglomerate' (cf. sketch of this most Iikely the same specimen as collected by J. C. part of the section by de Morgan, 1882, Fig. 8). Moberg in 1887 at Tosterup 5 and figured by Christensen

12 UPPER CRET ACEOUS BELEMNITES FROM THE YOMB TROUGH

(1975a:20, Fig. 19). Locality CV:2 was reexcavated by P. Upper Campanian at that locality. Gravesen and others. When I visited the locality, it was about 6 m high and 8 m wide, and showed calcareous sandstone with pebbles of Silurian shales. In some hori Summarizing remarks zons the pebble content was so high that the sediment could be classified as a conglomerate. This outcrap The oldest part of the Upper Cretaceous sequence at yielded the following belemnites: B. mammillatus (very Rödmölla-Tosterup can be ascribed to the Lower/Middle common), B. mucronata (rare), and G. quadratascanien Santonian. The so-called 'Verus conglomerate' is sand sis? Christensen (very rare). The calcareous sandstone wiehed between beds from the Lower/Middle Santonian and Tosterup conglamerate at this outcrop can thus be and uppermost Lower Campanian, and it may therefore assigned to the uppermost Lower Campanian on belem be referred to the upper Santonian and/or lower Lower nite evidence. Campanian. The alternating calcareous sandstone and Typical Tosterup conglamerate was also found at conglomerates (Tosterup conglamerate) w h ich follows CV:3, and here the conglamerate yielded B. balsviken upon the 'Verus conglomerate' are from the uppermost sis. Consequently, the conglomera te is from the basal Lower Campanian and the basal Upper Campanian.

Stratigraphy

General outline were introduced by Coquand (1856, 1857) , who charac The Upper Cretaceous Series includes the following six terized them by oysters, rudists, brachiopods, echinoids, stages: Cenomanian, Turanian, Coniacian, Santonian, and ammonites. Campanian, and Maastrichtian. All stages except the The Senanian has its type locality in the Paris Basin Maastrichtian have their type localities in France. The southeast of Paris at the town of Sens. According to Cenomanian type locality is situated in the Paris Basin, d'Orbigny (1842) the Senanian included all beds younger southwest of Paris, in the environs of the Le Mans. The than the Turanian and olderthan the Danian. Maastrich Turanian is based on seetians in Touraine. The Conia tian strata are not recorded from the Paris Basin and cian, Santonian, and Campanian type Iocalities are in the therefore some authors have excluded the Maastrichtian Aquitaine Basin, while the Maastrichtian type locality is from the Senonian. Haneock & Kennedy (1981) the ENCI-quarry at Maastricht in southeastern Holland. regarded the Senonian as a superstage, while Harland, When d'Orbigny (1842) began to subdivide the Upper Cox, Llewellyn, Pickton, Smith and Walters (1982) con Cretaceous in to stages, he first recognized two: the Turo sidered it as a subseries. Since the Senanian means diffe nian below and Senanian above. In 1847 d'Orbigny rent things to different authors and represent 23-24 Ma realized t hat the Turonian contained two different faunas (Obradovich & Cobban 1975; van Hinte 1976; Lanphere and the lower part was separated as the Cenomanian. & Jones 1978; Kennedy & Odin 1982; and Harland et al. d'Orbigny (1852) discussed at length the Cenomanian 1982), it is suggested here that the Senanian should no and Turonian stages, and the two units were charac longer be employed. The Subcommission on Cretaceous terized by various fossil groups, such as gastrapods, Stratigraphy of the IUGS does not recommend the for bivalve molluscs , foraminifera , and ammonites. The malization of the Senanian (Birkelund, Hancock, Hart, Cenomanian was referred to as the Second Zone of Ru Rawson, Remane, Robaszynski, Schmid & Surlyk 1984). dists and the Turanian as the Third Zone of Rudists. The Upper Cretaceous stages were loosely defined and The Coniacian, Santonian, and Campanian stages the characterizing fossils were in some cases of little

13 WALTER KEGEL CHRISTENSEN

biostratigraphic value. Moreover, i t has later been shown that the bases of the majority of the Cretaceous stages are The subdivision of the Upper characterized by hiati in the type areas (Birkelund et al. Cretaceous of Sweden 1984). Consequently, many diverging points of view regarding the delimitation of the stages have been put Christensen (1975a:20-23) surveyed in great detail the forward in the literature. Moreover, attempted earrela zonation and earrelation of the Upper Cretaceous strata tion of the sedimentary successions in France and Hol of Sweden, and consequently on! y a review is given here. land with successions in other areas has introduced inum In the Upper Cretaceous of Sweden, belemnites have erable complications. In an attempt to clarify the situa been shown to be of fundamental importance in bio tion, the Subcommission on Cretaceous Stratigraphy stratigraphy and correlation. The present belemnite heJd a symposium on Cretaceous Stage Boundaries in stratigraphy is the result of progressive refinements by a Capenhagen in the autum of 1983. At this meeting a set series of workers, and the studies of Schliiter (1870, of stage boundary proposals and recommendations were 1876), Moberg (1885, 1894), Stolley (1897, 1930), formulated (Birkelund et al. 1984). Ammonites have Jeletzky (1948, 1949b, 1951, 1958), and Christensen been considered by many authors to be the most signifi (1970a,b, 1971 , 1973, 1975a) are particularly relevant. cant group in the subdivision of the Upper Cretaceous The early subdivision of the Upper Cretaceous strata stages. As mentioned above, both d'Orbigny and of Sweden and Germany is intimately connected, reflect Coquand used various fossils for defining the stages, and ing the activities of German paleontologists in Sweden ammonites can hardly be considered more important during the last part of the 19th century. Moberg and other than a number of other groups. In northern Europe, Swedish palaeontologists apparently also visited German ammonites have been shown to be important for the localities. The first reliable subdivision of the Swedish zonatio n of the type Cenomanian and type Turonian and Upper Cretaceous was made by Schliiter (1870) , who its correlatives. Nevertheless, general agreement on the distinguished (top to bottom): greensand with Belem Albian-Cenomanian and Cenomanian-Turonian bound nitella mucronata and biocalcarenites with Belem aries and the subdivision of the Turonian has not been nellocamax mammillatus. Later on more belemnite zones achieved (cf. Wright & Kennedy 1981; Robaszynski were added by Moberg (1881 , 1882a, 1885 , 1894). Stolley 1982; Alcayde 1983; Kennedy 1984; Birkelund et al. (1897, 1930) subdivided the Senonian of German y and 1984). Sweden into (top to bottom): Mucronatenkreide, Quad The 'standard ammonite zonation' of the Coniacian ratenkreide = Mammillatenkreide, Granulatenkreide, Maastrichtian was reviewed by Kennedy (1984) and , in and Westfalicuskreide = Emscher. This zonation was the opinion of the present author, Kennedy el ear! y used subsequently in Sweden by most authors, translated demonstrated that i t is not workable. On the other hand, into mucronatakrita, mammillatuskrita, granulatuskrita, belemnites, inoceramid bivalves, and echinoids are of and westfalicuskrita. Stolley (1897 , 1930) correlated the importance of local zonations of the Coniacian-Maas Swedish strata with Belemnellocamax mammillatus with trichtian in northern Europe, and these local zonations German strata containing Gonioteuthis quadrata. This can be correlated with little difficulties with in that region, earrelation was nol generally accepted in Sweden, and except to the type Coniacian-Campanian. other proposals were put forward by Lundegren (1932a , The Coniacian-Maastrichtian of northern Germany 1934, 1936). has been carefull y studied during the last twenty years by When discussing the zonation of the Swedish Upper a number of workers (see review by Ernst, Schmid & Cretaceous it should be born in mind that up to 1930 the Klischies 1979; Schulz, Ernst, Ernst & Schmid 1984), and zonation was only concerned with the Santonian-Maas work is in progress on the Coniacian-Maastrichtian of trichtian. Pre-Santonian Upper Cretaceous strata were Great Britain (see Bailey, Gal e , Mortimore, Swiecicki & first recorded from the Båstad Basin by Lundegren Wood 1983, 1984). (1932b) and Stolley (1932), in the Malmö area by Brotzen The definition of the Coniacian-Campanian stages (1945) and from the Kristianstad Basin by Christensen ehosen in the present paper is shown in Fig. 7, and the (1970a). belemnite zonations of the Upper Cretaceous in NW Christensen (1975a) discussed at length the age of the Europe, Balto-Scandian, and on the Russian Platform zon e with B. mammillatus and concluded that this zone is are shown in Fig. 8. from the uppermost Lower Campanian and is equivalent

14 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

YO RKSHI RE and PHOSPHATIC W GERMANY S. ENGLAND NORFOLK N. IRELAND LINCOLNSHIRE CHALK , FRANCE

grimmensis/ granulasus Zone Ballymagarry Chalk ~n. Paramoudra Ch alk langel Zon e z Q; ~ n. ------Portrush C halk z n. polyplocum Zone seeston Ch atk c :;J g! "o Garron Chalk w / B.'minor ' o N ~ N -::_(a !.t_ ö!'-_ Sp2~~ ::_E[ej ~ ~ G lena rm Chalk =r: ~ ------;;"' Q 4: vulgaris Zone B. mu cronat a Zone "' Weybourne Chalk c u ~ o B ;;; e ------u a: n. ~ ~ Bal li ntoy Chalk -;;:; stobaeijbasiplana Zone " Eaton Chalk E UJ Q; E CL ai ------ai CL ;t Al" ::J E. c oni ca/ mucronata Zone Basal mu cronata Ch alk La rr y Bane Chalk ~ - ..Q.g_ gracili s/ mucronata Zone G. quadra ta Applinocrinus c1 Zon e cretaceus - Subzon e g! z conicajpapitlosa Zone o ~ N B z ;;; 4: n. papillosa Zone <;;"' Boheeshane Chalk CL Q; Gonioteuthis Zone -o --- ~ n. se nenensis Zone ------Hagenowia------(= 'zone of granul aled "'~ upper unit with 4: n. blackmorei Ac tinocamax' A :;J G. quadrata and u pilulaj senonensis Zone Harizon sensu Brydon e) " a: ------" B. praecu rsor UJ pil ula Zone Abundant S. binodesus c ;;; O. pi lula "o lingua Zone N s: n. Ottasler Subzone l. IIJonnmnmmon o l inguaj quadrata Zone pilula Zone _J Q; ------l~I~mW~fWIIJfll middle unit with Zon e 2"' ;t E. depressula ö. Creggan Chalk G . quadrata and E. gra nulataquadrata Zone Subzone ö O. pilula II I Il Il Il l III N Cloghastucan Chalk Marsupites/granulata Zone Marsupites testudinarius Zone Marsupites Zone M arsupites Zone ::1 z ::J l ~ Uintacrinus/granulata Zone Uintacrinus socialis Zone Uintacrinus Zone Uintacrinus Zone N Galboly Chalk z :i Cloghfin Sponge Beds J~ltJJl~Ilm[lower u nit with JitJJJI.A .verus o westfalicagranulata Zone 1- flintless c -~.:..c_o_r~n.9.:.~9_G..:.2r:§'~:.. z ~ c ha l k "o 4: cordiformis;Westfalica Zone M . coranguinum Zone M. coranguinum Zone Hagenowia ------N (/) rostrata Zone K i lcoan Sands E _J undulatoplicatus Zone i! flinty '3 z ::J praewestfalica Zone chalk "c < "' u involutus Zone o ~ ~ " z koeneni Zone ~ . Q>" M . deci piens Zone u c g...~ de formis Zone M .cort estudinarium Zone M .cortestudinarium Zone M . normanniae Zone ~rS

Fig. 7. Strati graphic earrelation di agram of the Coni acian-Campanian. Sources: Schulz (1978) , Ernst et al. (1979), and Ernst & Schulz ( 1980) for western German y; Bailey et al. (1983) for southern England; Peake & Haneock (1970) and C .J. Wood (MS, and pers. comm. 1983) for Norfolk; Wood (1981) for Yorkshire and Lincolnshire ; Fletcher (1977) , Fletcher & Wood (1978 , 1982) fo r Northern lreland; and Jarvis (1980) for France.

to the German G. quadrata gracilisl B. mucronata Zone correlatable with the lowermost part of the Mucronaten (cf. Figs 8-9). Belemnellocamax balsvikensis was recog kreide in Germany. nized by Brotzen (1960) and described by Christensen (1975a). Before 1960, specimens of B. balsvikensis were assigned to B. mammillatus. Brotzen regarded B. bals vikensis to be from the uppermost Lo w er Campanian, Stratigraphic conclusions but Christensen (1975a) suggested that the species is from the basal Upper Campanian. This suggestion was The belemnite faunas of the Vomb Trough have been contirmed by Christensen & Schulz (1976), who showed correlated with belemnite faunas from western Europe that the zone with B. balsvikensis and B. mucronata is and the Russian Platform (Fig. 8) . Formal belemnite equivalent to the German E. conica/ B. mucronata Zone zones are not established because extensive, continuous and a part of the superjacent P. stobaei/G. papillosa seetians are not present in the Vomb Trough or other basiplana Zone. The mammillatuskrita of Sweden there areas in southern Sweden. fore spans the uppermost Lower Campanian-basal Stratigraphic ages of the belemnite localities are Upper Campanian, and the upper part of the summarized in Fig. 9. The oldest part of the Upper mammillatuskrita (the zone with B. balsvikensis) is Cretaceous sequence at Rödmölla-Tosterup ('West-

15 W ALTE R KEGEL CHRISTENSEN

Belemnite zones, Zonal belemnites, Zonal belemnites, NW Europe Balto-Seandia Russian Plattorm

....: ....: ....: ti::> B. casimiravensis ti::> t;::> B. casimiravensis ro ro ro ro ro - ro :E _J B. junior ~_J ~_J B. junior ::> ::> ::> c B. fastigata c c ro . ~ . ~ ~::> B. ci mbrica .E=> .E !l:! B. sumensis (.) (.) Q) ·c B. sumans1s ·c .g c: c;; c;;- c;; E B. Janceolata ro B. obtusa ro ro ro ro .l!? B. licharawi :E ..J B. psaudobtusa ~_J

t ro B. 'tangei .iii B. l. najdini c a. c c O) ro ~ ------ro . ~ ::> c: B. l. Jangai ·c- ::> ::> ~ batsvikensisj B. mucronata .Q B .

B. mammillatusj G. q. scaniansis B. mucronatajG.q. gracilisj G. q. gracilisj B. mucronata ::> B. mucronata B. mammillatus -~ c G. q. gracilis c ro a. c ro . ~ ·c: Q; c c ro a. ro ro a. a. a. a. :J E ::> E E~ ro G. q. quadrata ro ro B. alphajB. praecursorj () () () ~ G. q. quadrata

::> B. praacurso( G. granulsta ::> G. granulata - ::> G. granulata ::> _J c c c ro ro ro c c westfalicagranulata G. wastfalicagranu/ata/ c .3 G. o B. propinqua o c B. propinqua/ @:E ~:E ro (/) (/) A'. lundgrani ui/icus G. w. westfalica ::> ~ : w. wastfalicajB. propinquaj (/)_J lundgrani - --_J A. _J ------_J ------c=> G. westfalica praewestfalica c=> c ro _ A'. lundgreni ro- . ~ ::> A'. lundgrani (.) ~:E ------: ~:E ro c_ c_ ------c o o o ()_J O ..J ()_J

c::> c::> c ro - ro::> ·c: :E . c_~:E ·c: e- o o- :J ..J ~_J :J ..J F F! F A. ptanus triangulus

A. p/enus A. p/enus c @::> @::> ro c c ·c: ::> A. plenus ro - ro - ro E ~:E ------~:E ------o c c c

16 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

STANDARD LOCA ZONATION, BELEMNITE ZONAL NW-GERMANY ZONES, BELEMNITES, BELEMNITES LITIES NW-EUROPE BALTO-SCANDIA grimmensis/ ~ l--"9.:..:'a:.cnc::u:..:loc::s::::us'----'=Z'-"o.:..:n:::.e---l B. '!angei, Z a. langei Zone "'

gracilis/mucronata zone G. quadrata gracilisj B. mammillatus,IG.q.scaniensis B. mucronata B. mucronata ll~ Z - conica jpapi Il osa Zone G. quadrata gracil is "' ~ ~~------1------~ ~ Z a. papillosa Zone .o c: ., ä: Cii 1------1 o g "' ~ ? senenensis Zone u "'Q; ~ a. ~ ~ ii .o :: ~ O pilulajsenonensis Zone G.quadrata quadrata '§ ? N «J""' ., ~ >:: ffi t pilula Zone e ~ ct1 ct1 ~ ? ~ ~·------1 ~ o -?- ~ ~ ~ o ..... linguajquadrata Zone L :J m ~ ~ <,; ~ ~~------1------4~~~~~~~~--1-~+-~ o e- ~ ~ ,~~.·t"lrt-tllt-~~~~lt-t-~-+-+.~1+-~-1·- ..Q granulataquadrata Zone G. granulataquadrata g: ~~~~~lataquadrataj ~ c: >:: ., ~d l ~ ., Il ~--I--M-a_ffi_u-pi-te-sfi~g-rn-n-ul-at_a_Z_on_e_~------~~~~------~~~~ ~~=·41~.-~~~~ri~~+-~• 4-+~ . ~~~~~4-+-~l~.,

z => G. granulsta _u_ G. granulats (!) ~ ~ ~ ~ ~ ~ ? ~ <( Uintacrinu5j9ranulata Zone L E åi M ci : cti 3 ~ ~ ~ Z -11------~---:------t""?!"-::-::===~=::-;--+·-=E-1 "' § ;:: : 11 O westtalicagranulata Zone G.westfalicagrnnulata G.westfa/icagranu/ataj s ~ ,t ; (!jhlt+"Hr+++:+-~·o++~H-++11 ++1·~-+-1 ~:?:~---:---:-~--~-I------:~~B.~p~ro~p~i~nq~u~a~~--~--l_:-~~.~~t+r+.rtJ·~-+~4-~-f-HI+-i-~-+~-~l~l+~l+,lr---

Fig. 9. Diagram showing the stratigraphic distribution of the belemnites occurring in the Vomb Trough, and the age of the localities according to the present study. In addttwn, the verttcal dtstnbutwn of belemmtes used as mdex fosstls is also shown.

falicus conglomerate') is referred to the Lower-Middle The outcrops at Kullemölla-Lyckås, which yielded Santonian on the basis of Gonioteuthis westfalica and Gonioteuthis granulataquadrata, Belemnitella alpha, Belemnitella propinqua. The outcrop at Eriksdal B is Actinocamax verus, and Belemnellocamax ex gr. assigned to the Upper Middle Santonian on the basis of grossouvrei, are ascribed to the basal Campanian, Zone G. westfalicagranulata and B. propinqua. Possibly, also of G. granulataquadrata. The loose blocks at Kåseberga Lower Middle Santonian strata we re exposed at Eriksdal from which B. ex gr. alphalpraecursor and G. granulata (see above). quadrata or G. quadrata have been collected, ma y also be Outcrops with Upper Santonian sediments are not from the basal Campanian, but i t cannot be excluded that recorded with the possible exception that the 'Verus they are slightly younger. comglomerate' at Rödmölla-Tosterup may be of that The alternating calcareous sandstones and conglome a ge. rates (Tosterup conglomera te) at Rödmölla-Tosterup

Fig. 8. Upper Cretaceous belemnite zones in western Europe, Balto-Scandia, and the Russian Platform. Sources: Christensen (1975a), Naidin & Kopaevich (1977) , Naidin (1979a, 1979b , 1981, 1983) , Schulz (1978) , Ernst et al. (1979) , and Ernst & Schulz (1980).

17 WALTER KEGEL CHRISTENSEN span the uppermost Lower Campanian-lower Upper Chatziemmanouil also estimated water depths and Campanian (the zone with Belemnellocamax mam concluded that the depth of water in the Coniacian millatus and Belemnitella mucronata and the superjacent ranged from 100-600 m at Kullemölla and 150-200 m at zone with Belemnellocamax balsvikensis and B. mucro Köpingsberg. W ater depth decreased in the early Santo nata) . The oldest part of the Köpinge sandstone is also nian to 100-200 m in both areas. In the middle and upper from the uppermost Lower Campanian and was found in Santonian a further decrease took place, and water the eastern part of the Valleberga-Ingelstorp area. The depths were estimated to be 30-100 m at Kullemölla and Köpinge sandstone from the western part of the Val less than 50 m at Köpingsberg. These sea-levet fluctua leberga-Ingelstorp area and the former outcrops around tions were considered to be of eustatic and local tectonic Köpinge can be assigned to the middle Upper Campa origin. Chatziemmanouil did not include information nian on the basis of B. mucronata and B. aff. langei. The from surface outcrops in the trough , although these pro zone with B. balsvikensis was not recognized in the vide important additional data. Several conglomerates, Köpinge area. some of which are of considerable thickness, and non Maastrichtian ammonites have been recorded from the sequences occur in the Santonian and Campanian but Köpinge area, and Lower Maastrichtian strata were none of these were identified in the borings. found in the Köpingsberg boring. Maastrichtian belem At Rödmölla-Tosterup, Lo w er Jurassic rocks are over nites, however, have not been recorded. lain by Upper Cretaceous strata with a conglamerate ('Westfalicus conglomerate') from the Lower-Middle Santonian at the base. The Tosterup congtornerate is thick at Rödmölla-Tosterup and betongs to both the Depositional history of the Vomb uppermost Lower Campanian and basal Upper Campa nian. The congtornerate is also known from the Val Trough during the Upper Cretaceous leberga-Ingelstorp area. Parts of the upper Santonian Formation of the trough was probably initiated in the and the Lower Campanian, with the exception of the Triassic since the oldest sediments recorded are of that uppermost Lower Campanian, are not present at Röd• age (Norling 1982; Chatziemmanouil 1982). The max mölla-Tosterup. On the basis on information from out imum thickness of strata preserved in the trough is about crops, it can therefore be inferred that transgressive 1500 m and occurs close to Köpingsberg in the southern pulses took place in the early Santonian and late Lower part of the trough (Norling 1982:76). Campanian, at !east in the Rödmölla-Tosterup area. Chatziemmanouil (1982) analysed the depositional The zone with Belemnellocamax balsvikensis and history of the Vomb Trough during the Upper Cretace Belemnitella mucronata is not recognized in the Val ous on the basis of five borings placed along the trough leberga-Ingelstorp area, bu t the lack of this zone ma y be axis. Three borings were situated in the northwestern due to bad exposures in that area. part and two in the southeastern part. The maximum The youngest Cretaceous sediments are from the thickness of the Upper Cretaceous is a little lessthan 800 Maastrichtian, probably the Lower Maastrichtian. In the m, found in the Köpingsberg boring (Anderegg et al. Köpingsberg boring the Lower Maastrichtian is termi 1968; Chatziemmanouil1982). The analysis was basedan nated by a conglamerate with coal and plantremains and seismic methods, sedimentologic analyses, and palaeon is overlain by Quaternary strata (Chatziemmanouil tologic/ecologic studies of foraminifera. The dominant 1982). Thismay be attributed to erosion of still younger sediments are glauconitic, calcareous, clayey siltstones Cretaceous strata during the Tertiary and Quaternary deposited in a fairly shallow-water environment. Periods. A large-scale world-wide regression, however, sedimentation during the Upper Cretaceous therefore took place towards end of the Cretaceous, and this may probably kept pace with subsidence. The targest rate of also explain the absence of sediments from the upper part sedimentation was in the Coniacian and Santonian and, of the Maastrichtian. In the Kristianstad Basin sediments consequently, the major subsidence occurred in this of late Maastrichtian age are also missing (Christensen period. 1975a).

18 UPPER CRETACEOUS.BELEMNITES FROM THE VOMB TROUGH

Systematic part

Morphology of the guard and terminology Moreover, the Schatzky distance is measured along the The guard is usually the only part of the skeleton pre wall of the alveolus by Naidin (1952 , 1959 , 1974) and served, and externa! and interna! characters are used for Ernst (1964a) . I have used the original definition as pro taxonornie classification. The characters are shown in posed by Jeletzky (1949a). Fig. 10. The term conellae (Hölder 1955) is used descriptively The following characters generally are considered to for conical tuberdes which may cover the wall of the be of taxonornie value in describing Upper Cretaceous alveolus or pseudoalveolus. In some species the conellae belemnites belonging to the Belemnitellidae Pavlov: (1) are covered by the 'white-layer' sensu Christensen size of guard, (2) shape of guard, (3) structure of the (1975a:29). anterior end, ( 4) surface markings, (5) interna! charac The 'Riedel-Quotient' (Ernst 1964a) is the ratio of ters, and (6) ontogeny. length of guard divided by depth of pseudoalveolus, and Belemnites which are completely calcified anteriorly the 'Schlankheits-Quotient' (Ernst 1964a) is the ratio of are provided with an alveolus which houses the phrag length of guard divided by dorso-ventral diameter at the mocone. In some species, the anterior end was not com anterior end. pletely calcified, and depending upon the degree of cal cification, the anterior end may be conical with an alveo lar fracture , flat with a pit in its centre, or developed as a concave pseudoalveolus. Measurements and abbreviations Surface markings include darso-lateral longitudinal depressions, darso-lateral double furrows , lateral A list of measured characters and abbreviations is given furrows , vaseular imprints, longitudinal striation, and below (cf. Fig. 10). granulation. Total length of guard (L) The interna! characters are alveolar angle , fissure Length from apex to protoconch (LAP) angle, Schatzky distance, and the shape of the bottom of Dorso-ventral diameter at protoconch (DVDP) the ventrat fissure. The alveolar angle is the angle of the Lateral diameter at protoconch (LDP) alveolus measured in the median plane. It is measured in Dorso-ventral diameter at alveolar end (DVDAE) the anterior part of the alveolus for reasons given by Lateral diameter at alveolar end (LDAE) Christensen (1975a). The fissure angle is the angle be Maximum lateral diameter (MLD) tween the wall of the alveolus and the straight Iine which Schatzky distance (SD) connects the intersection points of the bottom of the Alveolar angle (AA) ventrat fissure on the wall of the alveolus and the outer Fissure an g le (FA) margin of the guard. The Schatzky distance is defined as 'Riedel-Quotient' (RO) the distance from the anterior part of the protoconch to 'Schlankheits-Quotient' (SQ) the beginning of the bottom of the ventrat fissure mea sured along the longitudinal axis of the guard (Jeletzky Linear measurements were made with a vernier caliper 1949a). These definitions were used by Birkelund (1957) , to an accuracy of 0.1 mm, and an gles were measured with Kongiel (1962) and Christensen (1975a). In this context i t a goniometer ocular fitted on a WILD stereomicroscope should be pointed out that these characters are not mea to an accuracy of O.SO . sured in the same way by all authors. For instance, Naidin In earher studies of Belemnitella and Belemnella, (1952, 1959, 1974) and Schulz (1979) defined the fissure numerous other characters have been measured ( e.g. angle as the angle between the longitudinal axis of the length of reconstructed guard, depth of alveolus of recon guard and the straight Iine connecting the intersection structed guard, etc.), but these measurements are func points ofthe bottom ofthe ventrat fissure. The difference tions offive basic measurements: LAP, DVDP, SD, FA, between the two definitions isthat the fissure angle in the and AA, and are therefore of little taxonornie value, as latter case includes the half of the alveolar angle. are also the ratios obtained from these measurements.

19 WALTER KEGEL CHRISTENSEN

Biometric methods tinocamax), and A. (Paractinocamax), and two subgen era of the genus Gonioteuthis: G. (Gonioteuthis) and G. The variability of the belemnite species and subspecies is (Goniocamax); these five subgenera were treatedlater as analysed using univariate and bivariate statistical genera by Naidin & Kopaevich (1977) and Naidin (1981) methods and is summarized by descriptive statistics, his among others. This classification is not followed here for tograms, and scatter diagrams. The statistics were calcu reasons given by Christensen (1982) . The subgenus Par lated according to standard formulae presented by Simp actinocamax is furthe r discussed below. son, Roe & Lewontin (1960) and Sokal & Rohlf (1969). The statistical methods and tests used in the present paper were discussed at length by Christensen (1975a:31-33) . The species concept in belerunites I have earlier discussed the disadvantages of using The modern species concept in palaeontology is based on ratios in palaeontologic studies (Christensen 1973 , 1974, populations, rather than a few specimens (the so-called 1975a), especially in cases where growth is allometric. In typological-morphological species concept). Species are the present paper, I have on! y calculated ra tios in those studied by analysing the variation of homogenous popu instances where the small number of available specimens lations from restricted stratigraphic intervals by means of presents bivariate analysis. various biometric methods. This concept has only In the univariate analysis the estimates of the following recently been applied to Upper Cretaceous belemnites statistics were calculated: arithmetical mean value (X) ; (Birkelund 1957; Kongiel 1962; Ernst 1964a, 1968; standard deviation (SD); and the coefficient of variation Christensen 1971 , 1973 , 1974, 1975a, 1975b, herein; (CV). In addition , the observed range (OR) is reported, Schulz 1979; and J arvis 1980) , and biometric studies have and N is the number of specimens. shown that many belemnite species and subspecies estab The regression Iine is written: y = a + bx , and the lished by earlier authors are nothing but morphological original measurements were used in the calculations, variants. because of the linear trend on ordinary graph paper and Joysey (1956:84--88) , Bettenstaedt (1968:356), Schulz the homoseedastic variance around the regression Iine. (1979:37) , and Callomon & Birkelund (1982:353) have The estimates of the following statistical parameters we re discussed 'horizontal classification' and 'vertical classifi calculated: the slope (b) and standard deviation of the cation'. l t should be mentioned that Joysey used the slope (SDb) ; the intercept on the y-axis (a) and the stan terms chronologic for 'horizontal classification' and var dard deviation of the intercept (SDa) ; the variance ietal for 'vertical classification'. 'Horizontal classifica (SD2yx) and the standard deviation (SDyx ) of the regres tion' is b ase d upon the analysis of the variation of popula sion Iine; and the correlation coefficient (r) . N is the tions from successive horizons, and can be considered as number of specimens. The correlation coefficients were an approximation to the biological species concept. In testedfor significance by using Table Y in Rohlf & Sokal 'vertical classification' , similar specimens from succes (1969) and t-tests on the y-intercepts were performed in sive horizons are given the same name. 'Horizontal order to see if the intercept differed significantly from classification' was favoured by the authors mentioned zero. The regression Iines of two sam p les we re compared above, and has also been used by the present author in the way described by Hald (1957:571- 579). (Christensen 1971 , 1973 , 1975a, and herein). Ernst (1964a, 1968) defined species of Gonioteuthis on Classification of the Belemnitellidae the basis of analysis of populations from successive hori zons ('horizontal . classification'). Nevertheless, the The following genera within the Belemnitellidae have stratigraphic ra n ges of species of Gonioteuthis were illus received general recognition: Actinocamax Miller, trated in diagrams using 'vertical classification'. In that Gonioteuthis Bayle, Belemnellocamax Naidin, Belem way, two or three species of Gonioteuthis occur in the nitella d'Orbigny, Belemnella Nowak, Belemnocamax same horizon. For example, G. westfalica, G. west Crick, and Fusiteuthis Kongiel. The two last-mentioned falicagranulata, and G. granulara were shown to co-occur genera are monotypic. in the upper part of the G. westfalicagranulata Zone (see Naidin (1964b) recognized three subgenera of the Ernst 1966:137 ; Fig. 6). This Iine of action should be genus Actinocamax: A. (Actinocamax), A. (Praeac- discouraged.

20 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

® ®

ntral fissure

alveolus dorsal field darso-lateral longitudinal depression protoconch so

vaseular imprints

darso-lateral double furrow LAP apical- Iine

L growth Iines

Fig. l O. Figures showing the morphologic elements of the guard. A and B: Belemnitella mucronata from the Gerrnania IV quarry at Misburg/ Hannover. Neotype proposed by Chri stensen et al. (1975) . A: Ventro lateral view. B: View of split guard. C: View of ground guard of Belemnellocarnax mammillatus; schematized drawing. L = length of guard; LAP = length from apex to protoconch; DVDAE = dorso ventral diameter at alveolar end ; DVDP = dorso-ventral diameter at protoconch; D = depth of pseudoalveolus; SD = Schatzky distance; FA = fissure angle; and AA = alveolar angle. After Christensen (1975a).

21 WALTER KEGEL CHRISTENSEN

and characterized by independently evolving belemnite lineages; the Gonioteuthis stock inhabited the Central Family Belemnitellidae Pavlov, 1914 European Subprovince and the Belemnitella stock (including 'A'. lundgreni) inhabited the Central Russian Class Cephalopoda Cuvier, 1794. Subdass Coleoidea Subprovince. In other periods of the Upper Cretaceous Bather, 1888. Order Belemnitida Zittel, 1895. Suborder the subprovinces are less distinct and ma y disappear com Belemnopseina J eletzky, 1965. pletely. The Balto-Seandian belemnite faunas can be consi Distribution. - Belemnitellidae are restricted to the dered as mixtures of the belemnite faunas from the two Upper Cretaceous and are reported from the Lower subprovinces in the upper Coniacian-Lower Campanian, Cenomanian to the Upper Maastrichtian inclusive. and provide therefore a base for correlation. Near the Christensen (1975a, 1976, 1982) reviewed the Lower-Upper Campanian boundary, the belemnite palaeobiogeography of the Upper Cretaceous belem faunas of Balto-Seandia are characterized by the genus nites of Europe (see also Combemorel, Christensen, Belemnellocamax: B. mammillatus in the uppermost Naidin & Spaeth 1981), and consequently only a brief Lower Campanian and B. balsvikensis in the lower outline is given here. Belemnitellidae characterize the Upper Campanian. North Temperate Realm, but occur also in the northern After the extinction of the genera A ctinocamax, part of the Tethyan Realm. Belemnites belonging to Gonioteuthis, and Belemnellocamax near the Lower Belemnitellidae are unknown from the southern part of Upper Campanian boundary, the genus Belemnitella the Tethyan Realm and the South Temperate Realm. spread all over the North European Province in the The North European Province was subdivided in to two Upper Campanian, and has even been recorded from the subprovinces: the Central European and the Central northern part of the Tethyan Real m. Howeve r, no well Russian Subprovinces (Fig. 11) . These subprovinces are defined subprovinces can be recognized in the Upper well-defined in the upper Coniacian-Lower Campanian Campanian and Maastrichtian.

Central Russian E=3 Central European Boundary between Nort h European IIIIIIIIIIlJ Subprovince ~ Subprovince - - -- Province and Tethyan Real m

Fig. 11. Distribution of Upper Cretaceous biogeographic units in Europe based on belemnites. Upper Cretaceous land and sea areas represent maximum inundation for all s tages. The boundaries of land areas are not reliable in detail and the biogeographic units are typically gradational in character. After Christensen (1976).

22 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

Genus Actinocamax Miller, 1823 sharply demarcated from the surface of the guard; A. v. fragilis was differentiated by having a high cone-shaped Type species. - Actinocamax ve rus Miller, 1823 alveolar fracture which is up to one quarter of the entire length of the guard, and the fracture generally is Distribution. - Actinocamax ranges from the Lower asymmetric; and A. v. dnestrensis was distinguished by Cenomanian to the middle Lower Campanian inclusive. having a shallow pseudoalveolus. The variation of the lt is distributed in the North European and North Ameri structure of the anterior end is due to differential calci can Provinces. fication. Arkhangelsky (1912) and Naidin (1964b) considered A. v. verus and A. v. fragilis as geographic subspecies; A. v. verus has its main distribution in western Europe and A. v. fragilis on the Russian Platform. Birkelund Actinocamax ve rus Miller, 1823 (1957) and Christensen (1973) did not differentiate the PI. l, fig. l three 'subspecies', because they occur tagether at out Material.- Eriksdal: 116 complete specimens and alveo crops in Sweden and Denmark. lar fragments; Kullemölla: 154 complete specimens and Two samples of A. verus, from Eriksdal (upper Middle alveolar fragments; Lyckås: 3 complete specimens. In Santonian) and Kullemölla (basal Campanian), were addition, many fragments of the posterior part of the analysed with respect to the structure of the alveolar end. guard of a small belemnite most Iikely belonging to A. With respect to the structure of the anterior end both verus are present from Eriksdal and Kullemölla. sam p les showed a series of forms ranging from specimens Kullemölla boring: 42 specimens. with a high cone-shaped alveolar fracture to the speci mens with a shallow pseudoalveolus with all intermediate Short description. -An Actinocamax with a small, stout forms. Nevertheless, the specimens of the two samples guard which is lanceolate in lateral and ventraJ views. were subdivided into three groups, although it was The guard is flattened ventrally and the anterior end is difficult in some cases to decide to which group the speci compressed. The anterior end may have a low or high mens should be placed. Group l consists of specimens cone-shaped alveolar fracture, be flat , or have a shallow with a lo w cone-shaped alveolar fracture (ve rus-like pseudoalveolus. There is a small pit in the anterior end specimens), group 2 consists of specimens with a high for housing the posterior part of the phragmocone. The cone-shaped alveolar fracture (fragilis-like specimens), alveolar fracture is symmetric or asymmetric, and in the and group 3 of specimens with a shallow pseudoalveolus latter case the dorsal side is more incised than the ventraJ (dnestrensis-like specimens). The result is shown in Table side. The alveolar fracture may be sharply demarcated l, and it can be seen from this table that most specimens from the surface of the guard or may continue gradually in the two samples belong to group l. Some few speci into the surface of the guard. The anterior end exhibits mens in both samples belong to groups l and 3. In the concentric growth layers of the guard andradiating ribs. opinion of the author the sam p les ex hi bit a normal varia Darso-lateral longitudinal depressions are faintly tion with respect to the structure of the anterior end. developed, and vaseular imprints are weakly developed Similar variations are also seen in other species of or not present. The ventraJ fissure and ventraJ furrow are not preserved. The surface of the guard generally is Actinocamax, i.a. A. plenus (see Christensen 1974) and in G. westfalica (see Ernst 1964a; Christensen 1975a). lt covered by granules which may form corrugated trans verse Iines. Affinity. - A. ve rus kan be distinguished easily from TABLE l. Estimates of relative abundance of three groups of other species of Actinocamax by its small size and stout Actinocamax verus from E riksdal (upper Middle Santonian) and guard. Specimens of A. ve rus ha ving a flat anterior end or Kullemölla (basal Campanian). Group l contains specimens with a iow cone-shaped alveolar fracture (verus-like specimens); group 2 includes a shallow pseudoalveolus differ from juvenile specimens specimens with a high cone-shaped alveolar fracture (fragilis-like speci of G. westfalica in being stouter and more lanceolate in mens); and group 3 contains specimens with a shallow pseudoalveoius (dnestrensis-iike specimens). ventraJ view. Naidin (1964b) distinguished the following 'subspecies' of A. verus: A . v. verus, A. v. fragilis Ark Locality Group 1 Group 2 Group 3 hangelsky, 1912, and A. v. dnestrensis Naidin, 1952. A. v. E riksdal 100 (86%) 9 ( 8%) 7 (6%) 116 verus was characterized by having a low cone-shaped Kullemölla 122 (79%) 26 (17%) 6 (4%) alveolar fracture which generally is asymmetric and 154

23 WALTER KEGEL CHRISTENSEN is therefore suggested that A. verus dnestrensis is placed of the anterior part of the guard. In G. westfalica the in synonymy of A . v. verus. A. v. verus and A. v. fragilis anterior end may be convexly conical, flat , or developed may be considered as geographic subspecies if it can be as a shallow pseudoalveolus, while in stratigraphically demonstrated that specimens with a high cone-shaped younger representatives the depth of the pseudoalveolus alveolar fracture (fragilis-like specimens) prevail on the increases and ma y be up to 113 of the entire length of the Russian Platform. guard. Simultaneausly with the development of deeper pseudoalveolus the guard becomes increasingly stout and Remarks. - A. verus was recorded by Moberg (1884, large, reaching a maximum in G. quadrata. Another 1888, 1910b) from the 'Westfalicus conglomerate' and characteristic feature is the gradual development of 'Verus conglomerate' at Rödmölla-Tosterup. I have not granulation. The oldest member, G. westfalica praewest seen A. verus from these conglamerate and the speci falica, does not possess granulation (Ernst & Schulz, mens appear to be lost. 1974) . The succeeding taxon, G. w. westfalica, shows a Distribution. - A. verus is wide-spread in the North widevariation with respect to that character; some speci European Province. mens carry scattered granules on the dorsal and/or vent A . verus appears in northwestern Europe in the upper ra( side of the guard, and sometimes the gran u les appears most part of the Lower Santonian /. undulatoplicatus to be arranged in longitudinal rows. The same pattern is Zone and continues into the middle Lower Campanian also valid for G. westfalicagranulata. In stratigraphically O. pilula Zone sensu germanico (Ernst 1963a; Christen younger species, however, the granulation becomes a sen 1975a). In off-shore chalks A. verus is most common very prominent character. In samples of G. westfalica the in the Upper Santonian (see Ernst & Schulz 1974; Bailey shape of the guard is highly variable, and a great propor et al. 1983), w h e reas in near-shore sedimentary facies, for tion of the guards are lanceolate in ventral view. In instance in Sweden and the Subhercynian Cretaceous, it younger population the variation in the shape of the occurs commonly also in the basal Campanian (Uibrich guard is smaller and guards which are lanceolate in vent 1971; Christensen herein). ra( view are relatively rare. The cross-section of the According to N aidin ( 1864b) A. v. fragil is occurs on the anterior end is oval to pointed oval in G. westfalica and Russian Plattorm from the Upper Turonian to the lower subrectangular to subquadrate in stratigraphically part of the Lower Campanian ('Pteria beds'), A. v. verus younger representatives with all intermediate stages. In from the Coniacian to the Lower Campanian, and A. v. the uppermost Lower Campanian, the Gonioteuthis dnestrensis in the Santonian. stock is characterized by a return of earlier features: e.g. reducing length of the guard, diminishing depth of the pseudoalveolus, and increasing slenderness of the guard. The Gonioteuthis lineage provides a good tool for stratigraphic purposes. It is , however, necessary to ana Genus Gonioteuthis Bayle, 1879 lyse homogeneous samples of a certain size in order to make a reliable specific determination, and limited mate Type species. - Belemnites quadratus Blainville, 1827 rial has only little stratigraphic value (Ernst 1964a; Remarks. -The evolutionary lineage of Gonioteuthis, in Christensen 1975a). Only the earliest member of the ascending order: G. westfalica praewestfalica Ernst & lineage, G. westfalica, is easily recognizable owing to its Schulz, G. w. westfalica (Schliiter), G. westfalica structure of the anterior end. granulata (Stolley), G. granulara (Biainville), G. Ernst (1963a , 1963b, 1964a, 1964b, 1966, 1968) and granulataquadrata (Stolley), G. quadrata (Blainville), Ernst & Schulz (1974) characterized populations of and G. quadrata gracilis (Stolley) , has been studied espe Gonioteuthis on the basis of mean values of various cially by Stolley (1897, 1916, 1930) , Ernst (1963a , 1963b, ratios. Christensen (1975a) discussed the use of these 1964a, 1964b, 1966, 1968) Ernst & Schulz (1974), Chri ratios and suggested that the characters in question stensen (1975a, 1975b), and Jarvis (1980). should be analysed by bivariate statistical methods. This The lineage is an outstanding example of phyletic proeecture was followed by Jarvis (1980). gradualism, vi z. slow gradual transformation of a suite of In studies on samples of Gonioteuthis, the following characters within populations of Gonioteuthis through characters are important: (1) length of guard, (2) length time. The Gonioteuthis stock existed for about 10 Ma and of guard vs. depth of pseudoalveolus, and (3) length of the general trend in evolution is the gradual calcification guard vs. dorso-ventral diameter at alveolar end.

24 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

Distribution. - Gonioteuthis is known from the Middle 25 % Coniacian I. involutus Zone to the boundary between the Lower-Upper Campanian, and the extinction leve) of the genus has been proposed by authors as the boundary for the Lower and Upper Campanian. 15 The genus had its evolutionary centre in northwestern Europe and is found almost exclusively in the Central European Subprovince. A few representatives of the 5 Lin genus are reported from the northernmost part of the mm Tethyan Realm. 30 40 50 60 Fig. 12. Histogram of length of guard (L) of Gonioteuthis westfalica granulata from Eriksdal. The numbers above the bars are the actual number of specimens. Gonioteuthis westfalica (Schlöter, 1874) PI. l, figs 2-3 Type. - The specimen figured by Schli.iter (1876, PI. 53: lO) was designated as lectotype by Ernst & Schulz (1974:50) . Material. - Eriksdal: 88 complete specimens and many fragments. These specimens were, by and !arge, col Material. - Rödmölla-Tosterup: 10 specimens. Kulle lected by J.C. Moberg in the last part of the nineteenth mölla boring, 330-390 m interval: c. 15 specimens. century and by A. Lundegren in 1931 (see above). The Remarks. - The earliest member of the Gonioteuthis Lundegren collection came from Eriksdal B (Lundegren lineage, G. westfalica, is closely comparable to known 1935a:577, footnote 3), and the specimens collected by species of Actinocamax, especially with regard to struc Moberg probably came from the same locality. ture of the anterior end, which may have a low cone Biometry. - 88 complete specimens were analysed. A shaped alveolar fracture, be flat, or developed as a very histogram of length of guard is shown in Fig. 12. The size shallow pseudoalveolus. The species is placed in the frequency distribution was testedfor normality using the genus Gonioteuthis on account of its undoubted relation Kolmogorov-Smirnov test for goodness of fit. The test ship to later members of the lineage. The taxon can be statistic D is 0.0420 with 88 degrees of freedom, which is recognized without difficulties owing to its structure of not significant (P>20% ), implying that the distribution the anterior end, even if only a small number of speci does not differ significantly from normality at the 5% mens is available. leve!. The mean length of the guard is 47.4 mm and the The specimens from Rödmölla-Tosterup we re la belled standard deviation is 8.4 mm. Tosterup 7 by J.C. Moberg in 1887, but on the basis of the The scatter plot of length of guard vs. depth of adherent matrix it can be inferred with great probability pseudoalveolus is shown in Fig. 13, and the estimates of that they came from the 'Westfalicus conglomerate' at the statistical parameters of the regression analysis are locality CV:1 (cf. Moberg 1910b and above). given below. Stratigraphy. - G. westfalica praewestfalica occurs in West Germany in the Middle-Upper Coniacian, and G. D = -0.0718 + 0.1081 L; N = 88; r = 0.6496; SDa = w. westfalica occurs in the upper Lower Santonian and 0.6498; SDb = 0.0151; SDyx = 1.0573. Lower Middle Santonian (Ernst & Schulz 1974); (see Figs 8-9). The value of the earrelation coefficient is very highly significant (P<0.1% with 86 degrees of freedom). A t test on the y-intercept showed that the intercept does not differ significantly from zero (ta = 0.1105 with 86 degrees Gonioteuthis westfalicagranulata (Stolley, 1897) of freedom; P>90%) w hi ch im p lies an isometric relation PI. 2, figs 4,6 ship of the two characters. Type. - The specimen figured by Stolley (1897, PI. 2:16; The Gonioteuthis sample was campared to a sample of PI. 3:6) was designated as lectotype and refigured by G. westfalicagranulata from the quarry 'Mathilde' in Christensen (1975b:128; PI. 10:1, Fig. 2A). northwest Germany (see Ernst 1968:278, Fig. 7). The

25 WALTER KEGEL CHRISTENSEN

D in 10 mm

8 • • • • ..• • .. 6 • ••• ...... ~ • • • • ·...... • •• 4 l • • • • • • • • • • 2 • Lin mm

10 20 30 40 50 60 Fig. 13. Scatter diagram and regression Iine for Gonioteuthis wes/falicagranulala from Eriksdal. L= length of guard; D = depth ofpseudoalveolus. • = J .C. Moberg Coli. ; • = specimens figured by Moberg (1885) ; .Å. = A. Lundegren Coli.; + = mean values.

estimates of the statistical parameters of this sample are falicagranulata from Eriksdal also contains some few as follows: specimens from the subjacent G. westfalica Zone. Stratigraphy. - G. westfalicagranulata occurs in the D = 0.1018 + 0.1106 L; N = 51; r = 0.4809; SDa Upper Middle Santonian (G. westfalicagranulata Zon e). 1.5940; SDb = 0.0288; SDyx = 1.1775.

The value of the correlation coefficient is very highly Gonioteuthis granulataquadrata (Stolley, 1897) significant (P90% ), implying an isometric relationship Christensen (1975b:130; PI. 10:2, Fig. 2B). of the two characters. The regression Iines of the two sam p les were compared Material. - Kullemölla: 11 specimens, 6 of which are and the test revealed that the variances, slopes and posi complete; Lyckås: 10 specimens, 3 of which are com tions of the two Iines do not differ at the 5% leve l. On the plete. basis of the bivariate analysis the sample of Gonioteuthis Remarks. - Only nine complete specimens are available from Eriksdal is referred to G. westfalicagranulata. It from Kullemölla and Lyckås. The mean length of the should be pointed out, however, that some very few guard of these specimens is 52.4 mm; the mean 'Riedel specimens with a relative! y shallow pseudoalveolus occur Quotient' is 5.6; and the mean 'Schlankheits-Quotient' is in the sample from Eriksdal. Five specimens have the 5.8. According to Ernst (1964a: 169) G. granulataquad following 'Riedel-Quotient': 15 .6, 16.8, 16.8, 19.0, and rata has a mean 'Riedel-Quotient' between 5.0-6.0. The 20.6, and they are rather westfalica-like. One of these specimens were also plotted on a scatter diagram of specimens is figured on PI. 1:5. In the population of G. length of guard vs. depth of pseudoalveolus for a sample westfalicagranulata from 'Mathilde', specimens with a of G. granulataquadrata from 'Weinberg' quarry (cf. 'Riedel-Quotient' above 15 do not occur. The presence Ernst 1964a:120; 1968:278; Christensen 1975a:38) . The of specimens of Gonioteuthis with a very shallow specimens from Kullemölla-Lyckås showed good agree pseudoalveolus at Eriksdal may be explained in one or ment to the sample from 'Weinberg'. The author, there more of the following ways: (l) the sample from Eriksdal fore, does not hesitate to refer the specimens from contains a larger number of specimens than the sample Kullemölla-Lyckås to G. granulataquadrata. from 'Mathilde' and therefore shows a greater variation, (2) the sample from Eriksdal is slightly older than the Stratigraphy.- G. granulataquadrata occurs in the lower sample from 'Mathilde', or (3) the sample of G. west- most Lower Campanian (G. granulataquadrata Zone).

26 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

Gonioteuthis quadrata scaniensis? Christensen, 1975 55 .5 mm, 'Riedel-Quotient' is 4. 7, and 'Schlankheits PI. l , fig . 9 Quotient' is 5.8. Belemnitella ex gr. alpha/praecursor is also known from the loose blocks, and the two specimens Type. - The specimen figured by Christensen (1975a, PI. are therefore tentatively assigned to G. granulataquad 2:4) is the holotype. rata or G. quadrata. Material.- Rödmölla-Tosterup, lo c. 2: on e alveolarfrag ment; Rödmölla: one complete specimen; and Tosterup 5: one alveolar fragment? Remarks.- Synonyms were given by Christensen (1975a) Genus Belemnellocamax Naidin, 1964 who also full y described the species (including biometric analysis) on the basis of material from the Kristianstad Type species. - Belemnites mammillatus Nilsson, 1826 Basin. Remarks. - The evolutionary lineage of Belemnello T wo of the specimens are small and s lender. The camax, in ascending order, B. ex gr. grossouvrei (Janet), important characters of the complete specimen are as B. mammillatus (Nilsson) , and B. balsvikensis (Brotzen), follows: length of guard is 46 .5 mm, 'Riedel-Quotient' is was studied recently by Christensen (1975a, 1976). The 6.0, and 'Schlankheits-Quotient' is 5.4. lt is not possible general trend in evolution is the gradual calcification of on the basis of on! y two specimens to refer them safely to the anterior end of the guard, which also becomes more species or subspecies. The specimen from loc. 2 was slender and less lanceolate in ventraJ view through time. collected tagether with B. mammillatus and B. muc Normall y, the genus is not granulated, bu t a few granu ronata. In the Kristianstad Basin , G. quadrata scaniensis lated specimens belonging to B. ex gr. grossouvrei and B. occurs tagether with B. mammillatus and B. mucronata, mammillatus have been recorded (Moberg 1885; Naidin and the specimen from loc. 2 is therefore tentatively 1964b; Christensen 1982). assigned to G. quadrata scaniensis. The complete speci Naidin (1964b) referred species of the grossouvrei men is on! y la belled Rödmölla and was ear!i er referred to group to A . (Paractinocamax), because they were as B. mammillatus by R. Hägg. The specimen might have claimed to have a short juvenile guard in contrast to come from loc. 2. The specimen labelled Tosterup 5 by species belonging to Belemnellocamax. The observation J .C. Moberg in 1887 was discussed by Christensen by N aidin, however, seems to be based on a single speci (1975a:21). men of B. grossouvrei pseudotoucasi (Naidin 1964b:72, G. quadrata scaniensis and G. q. gracilis are closely Fig. 15). I have studied the split guard of two specimens comparable. Both subspecies are small and slender but from Scania: (l) the holotype of B. grossouvrei var. scaniensis differs from gracilis by having a more shallow ornatus (Moberg, 1885) from Kullemölla (basal Campa pseudoalveolus. nian), and (2) a specimen from Flackarp in the Kristian Distribution. - G. quadrata scaniensis has up to now only stad Basin (highest Lower Campanian). Both specimens been recorded from the uppermost Lower Campanian of have a Iong and elongated juvenile guard. Moreover, Dr. Sweden. M.-G . Schulz, Kiel , has stud ie d three specimens of the grossouvrei group from the basal Campanian of Braunschweig, and these specimens also have Iong and elongated juvenile guards (M.-G. Schulz, pers. comm. 1984). In addition to these observations, I have seen Gonioteuthis spp. juvenile guards of the grossouvrei group from England PI. l, figs 10-11 and the Kristianstad Basin, and these guards are also very Remarks. -Representatives of Gonioteuthis occur in the elongated. In this context it should be noted that Prof. 60-390 m interval of the Kullemölla boring. Some of D .P . Naidin has not seen any juvenile or adoloscent these specimens are complete and weil-preserved (PI. specimens of the grossouvrei group from the Russian 1:11). lt is , however, not possible to assign them safely to Plattorm (D.P. Naidin in litt. 1984). species, except for G. westfalica, owing to the small On the basis of these observations, tagether with the number of specimens (see above). fact that species of the grossouvrei group show strong Two specimens of Gonioteuthis from loose blocks at affinity to B. mammillatus in several morphological Kåseberga have been studied. On e specimen (PI. 1: lO) characters, the grossouvrei group is referred to Belem has the following im portant characters: length of guard is nellocamax in the present p a per.

27 W ALTER KEGEL CHRISTENSEN

Distribution. - Belemnellocamax is recorded from the help solving these taxonornie problems. Lower Santonian (possibly highest Coniacian) to the According to de Grossouvre (1899: 133) B. ex. gr. lower Upper Campanian. grossouvrei should occur commonly in the Corbieres reg B. ex gr. grossouvrei is widely distributed, but rare in ion of the F ren ch Pyrenees. This record, however, has the North European Province. B. mammillatus is not been substantiated by later authors. extremelycommon in Scania, but rare outside that area: it has been recorded from northern Germany, Poland, Stratigraphy. - B. ex gr. grossouvrei is recorded in Ger and the eastern part of the Russian Platform. B . bals many from the G. granulataquadrata Zooe of the basal vikensis also occurs commonly in Scania, but outside this Campanian and the G. quadrata gracilisl B. mucronata area it is unknown except for a recent find of two speci Zone of the uppermost Lower Campanian. In Sweden, mens from oarthero Germany (Christensen & Schulz the group occurs in the basal Campanian at Kullemölla 1976). lt can thus be concluded that the area of distribu (herein) and in the uppermost Lower Campanian in the tion of the genus Belemnellocamax gradually diminished Kristianstad Basin (Christensen 1975a). The group is through its stratigraphic range. recorded from the Upper Santonian and possibly the lowermost Campanian in France and from the Upper Santonian and Lower Campanian on the Russian Plat form. Belemnellocamax ex gr. grossouvrei (Janet, 1891) Five specimens of the B. grossouvrei group have been Pl. 2, figs 1-2 recorded from England. (1) Crick (1906) described B. grossouvrei from flin ty ch al k at Firnber, Yorkshire (Fig. Material. - Kullemölla: five specimens. 7). According to C.J. Wood (pers. comm. 1983) the Remarks. - Specimens of the B. grossouvrei group are specimen came from high Coniacian or basal Santonian characterized by their !arge, ventrally flattened guards, Chalk. (2) Crick (1907) recorded B. blackmorei from which are lanceolate to strongly lanceolate in ventraJ West Harnham and according to C.J. Wood (pers. view. Moreover, they have a shallow pseudoalveolus, the comm. 1983) it came from the top part of the O. pilula cross-seetian of w h ich is triaugular. Zone or the base of the G. quadrata Zone sensu anglico. Specimens of the grossouvrei group occur rarely in (3) Brighton (1930) reported B . grossouvrei from Ruston western Europe and about 30 specimens are recorded Parva in Yorkshire. This specimen is from the O. pilula from Germany, England, France, and Sweden. These Zone sensu anglico (C.J. Wood, pers. comm. 1983). (4) specimens have been assigned to the following eight taxa: Bailey et al. (1983) recorded B . grossouvrei from Mottis B. grossouvrei (Janet, 1891) , B. toucasi (Janet, 1891) , B. fon t, Brydone loc. 1067. lt probably came from the alfridi (Janet, 1891), B. depressus (Andreae, 1895), B. Hagenowia blackmorei Harizoo (C.J. Wood, pers. depressus var. fusiformis (Andreae, 1895), B. marn comm. 1983). (5) Crick (1907) described Actinocamax mi/latus var. germanica (Stolley, 1930), B . mammillatus sp. from Fletcher & Co. 's pit at Gravesend; this specimen var. ornatus (Moberg, 1885), and B. blackmorei (Crick, is of late Coniacian or ear ly Santonian age (C. J. Wood, 1907). pers. comm. 1983) . The specimen was referred to the On the Russian Platform, the grossouvrei group is Mammillata group by Jeletzky (1949a) and the B. slightly more common than in Western Europe, and grossouvrei group by Christensen (1975a). It has a rather about 60--70 specimens have been recorded by Russian deep pseudoalveolus, the walls of which are covered by palaeontologists (e.g. Nikitin, 1958; Naidin 1964b; conellae and is strong! y flattened ventrally. The systema Glazunova 1972) . The specimens from Russia have been tic position of the specimen is uncertain, and it may assigned to B. grossouvrei pseudotoucasi (Naidin, 1964) belong to the 'A.' lundgreni group. In addition to the and B. g. pseudoalfridi (Naidin, 1964) by Naidin (1964b ), specimens mentioned above I have recently found two B . cf. toucasi and B. alfridi by Glazunova (1972), and B. specimens in the collections of British Museum (Natural toucasi var. seimensis (Nikitin, 1958) by Nikitin (1958). History): An adoloscent specimen from Micheldever, To sum up, eleven taxa have been established within Hants, (C 44382) was found among a sample of speci the grossouvrei group on the basis of relatively little mens la belled G. westfalica. lt is probably of basal Santo material. In the opinion of the author this group has been nian age (C.J. Wood, pers. comm. 1983). Anotherspeci the subject of excessive subdivision. Hopefully, the cur men came from East Harnham near Salisbury (C 44331). rent revision of this group on the basis of German and This specimen is probably of basal G. quadrata Zone age Swedish material by Prof. F. Schmid, Hannover, will sensu anglico.

28 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

To sum up, the grossouvrei group occurs from the 1975a:47). The estimates of the statistical parameters of Lower Santonian (possibly highest Coniacian) to the the sample from lgnaberga are given bel o w. boundary between the Lower and Upper Campanian. D = 0.1756 + 0.1098 L; N = 69; r = 0.6921; SDa 1.1031; SDb = 0.0140; SDyx = 1.4345.

Comparison of the regression Iines of the two samples Belemnellocamax mammillatus mammillatus (Nilsson, revealed that their variances, slopes, and positions do not 1826) differ significantly at the 5% leve!. 1t is therefore con PI. 2, figs 3-5 cluded that the sample from Rödmölla-Tosterup is of the Material.- Rödmölla-Tosterup, loc. 2: about 100 speci same stratigraphic age as the sample from lgnaberga new mens; Köpinge sandstone at Valleberga: 17 specimens. quarry, viz.latest Early Campanian (Christensen 1975a). Remarks. - Synonyms were given by Christensen Stratigraphy. - B.m. mammillatus occurs in the upper (1975a), who also described the species in detail (includ most Lower Campanian in beds correlatable with the ing biometric analysis) on the basis of material from the German G. quadrata gracilis/B. mucronata Zone. Kristianstad Basin in Scania. Biometry. - A sample of 25 complete specimens from Rödmölla-Tosterup, loc. 2 was analysed. The scatter p Iot Belemnellocamax balsvikensis (Brotzen, 1960) of Iength of guard vs. depth of pseudoalveolus is shown in PI. 2, figs 6--7 Fig. 14 , and the estimates of the statistical parameters are as follows: Material.- Rödmölla-Tosterup, Ioc. 3: c. 20 specimens, the majority of which are incomplete. D = - 1.7631 + 0.1422 L; N = 25; r= 0.7951; SD. = Remarks. - Synonyms were given by Christensen 1.6565; SDb = 0.0229; SDyx = 1.3264. (1975a), who also fully described the species (see also Christensen & Schulz 1976). Moreover, Christensen The value of the earrelation coefficient is very highly analysed biometrically populations of the species from significant (P<0.1% with 23 degrees of freedom) . A t localities in the Kristianstad Basin and discussed the test on the y-intercept showed that the intercept does not affinity of B.m. mammillatus and B. balsvikensis. differ significantly from zero (ta = 1.0643 with 23 degrees of freedom; 30%>P>20% ), which implies an isometric Stratigraphy. - B. balsvikensis occurs in the lower Upper relationship of the two characters. Campanian in beds corresponding to the German E. The sample of B. m. mammillatus from Rödmölla-Tos conica/ B. mucronata Zon e and the lower part of the P. terup was campared to a sample of the same species from stobaei/G. papillosa basiplana Zone (see Christensen & lgnaberga new quarry ( described by Christensen Schulz 1976: Fig. 3, and Figs 8-9, herein).

Din 12 mm • 8 •

Linmm

10 20 30 40 50 60 70 80 90 100 Fig. 14. Scatter diagram and regression Iine for Belemnellocamax mammillatus from Rödmöllafrosterup, loc. 2. L= length of guard; D= depth of pseudoalveolus; + = mean values.

29 W AL TER KEGEL CHRISTENSEN

Genus Belemnitella d'Orbigny, 1840 Material. - Kullemölla boring, 440 m leve!: one specimen. Type species. - Belemnites mucronatus Schlotheim, 1813 Remarks. - The specimen is incomplete: the posterior Remarks. - The International Commission on Zoologi part and the most anterior end of the guard are missing. ca! Nomenciature was recently petitioned (Christensen, The depth of the pseudoalveolus seems to have been Ernst, Schmid, Schulz & Wood 1982) to use its plenary relatively deep, and the guard is lanceolate in ventraJ powers to ratify the neotype for Belemnitella mucronata vie w. Moreover, the guard has longitudinal s tri a e and proposed by Christensen, Ernst, Schmid, Schulz & vaseular markings. The specimen is therefore referred to Wood (1975) and that the name should be attributed to as 'A'. cf. lundgreni. Schlotheim, 1813, and not to Link, 1807, its original author. This petition is still pending. Distribution. - 'A'. lundgreni occurs commonly in the Central Russian Subprovince and in Balto-Scandia. A Distribution. - Belemnitella is known from the Lower few representatives are known from the Central Euro Santonian to the uppermost Maastrichtian. The genus is pean Subprovince. The species occurs in the Middle Con widely distributed in the North European Province and iacian-Middle Santonian. has even been recorded from the northern part of the Tethyan Realm. It also occurs in the North American Province.

Belemnitella propinqua group Belemnitella propinqua propinqua (Moberg, 1885) The B. propinqua group includes 'Actinocamax' lund PI. 3, fig. 2 greni Stolley from the Coniacian-Middle Santonian and Type.- The specimen figured by Moberg (1885 , PI. 5:25) B. propinqua (Moberg) from the Lower-Middle Santo is the holotype. nian. B. propinqua is generally considered to be the earliest representative of the genus. It is a well-defined Material. - Eriksdal: three specimens; Rödmölla-Tos• species and was redescribed by Christensen (1971, 1973) . terup, loc. 1: one specimen; and Kullemölla boring, 248 I t evolved from 'A'. lundgreni. m leve!: one specimen. 'A'. lundgreni was placed in Gonioteuthis Remarks. - Christensen (1971, 1973) described the (Goniocamax) by Naidin (1964b) to gether with several species in detail on the basis of Swedish and Danish other species, including the earliest members of the material, including the holotype. Christensen (1971) Gonioteuthis lineage. This suggestion was criticized by gave synonyms of the species and showed that the con Ernst & Schulz (1974) and Christensen (1982). Ernst & cept of B.p. propinqua was misinterpreted by Russian Schulz suggested that subgenus Goniocamax with the palaeontologists. The following taxa were considered to type species 'A'. lundgreni should be elevated to a genus be synonyms of B. p. propinqua: Actinocamax pro or considered a subgenus of Belemnitella. They also sug pinquus? Moberg, 1885 , Belemnitella mucronata mut. gested that on ly 'A'. lundgreniandits ancestors should be anterior Stolley, 1897, B. ex gr. mirabilis Jeletzky, 1948, assigned to Goniocamax. The suggestion by Ernst & A. propinquus ra v ni Birkelund, 1957 , Gonioteuthis Schulz must await further studies and is outside the scope jeletzkyi Kongiel, 1962, and B. rylskiana Nikitin, 1958. ofthis p a per. 'A'. lundgreni, however, is placed here in in Naidin (1974) employed a different concept of B.p. the B. propinqua group, because it is closely allied to B. propinqua from that of Christensen (1971, 1973, and propinqua. herein). He distinguished between B.p. propinqua from the Lower Santonian and B.p. rylskiana from the upper Lower-Upper Santonian, in addition to the dubious B.p. mirabilis Ar khangelsky, 1912 from the Santonian of 'Actinocamax' cf. lundgreni Stolley, 1897 northern Kazakhstan (J eletzky 1949b). PI. 3, fig. l Distribution. - B.p. propinqua is distributed in the Cen Type.- The specimen figured by Stolley (1897, PI. 3:18) tral Russian Subprovince and in Balto-Scandia. It occurs wasdesignatedas lectotype by Birkelund (1957:14). in the uppermost Lower-Middle Santonian.

30 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

B. praecursor var. media is a morphologically inter Belemnitella praecursor group mediale form between var. praecursor and var. muc Two taxa are tentative! y placedin this group: B. praecur ronatiformis, and the varieties " . .. are on! y morphologi sor Stolley and B. alpha Naidin. cal varieties of the same specific type (parts of the same B. praecursor was established on the basis of a single populations; ... " (Jeletzky 1955:482). They are there specimen from the lowermost Campanian, G. granulata fore, following § 45 of the Code, to be regarded as of quadrata Zone, of the Broitzem quarry at Braunschweig infrasubspecific rank. On the other hand, var. praecursor (Stolley 1897; see also Ernst 1964b, 1968). lt was charac was on! y recorded from West German y, and the ear!i est terized by having a smooth guard, and the interna! populations of B. praecursor were said to consist mainly characters were unknown, because Stolley did not split of var. media and var. praecursor, w h ile stratigraphically the guard. The holotype was supposed to have been younger populations were said to be dominated by var. misplaced or lost during the Second World War mucronatiformis. If these suggestions are shown to be (Jeletzky, 1955), but the specimen was found in 1976 in correct by future studies, the varieties ma y be considered the Geologisch-Paläontologisches Institut und Museum to be geographic or chronologic subspecies. in Kiel (PI. 3:4). lt was subsequently split by Dr. M.-G. After Jeletzky's study of B. praecursor, specimens of Schulz, Kiel, who kindly supplied the following measure Belemnitella from the uppermost Santonian and lower ments of the critical characters: length from apex to and middle Lower Campanian were assigned to either B. protoconch is 65.0 mm; darso-ventraJ diameter at pro praecursor or B. mucronata by authors (e.g. see Nikitin toconch is 16.2 mm; lateral diameter at the protoconch is 1958; Kongiel1962). Ernst (1964b), however, described 16.9 mm; Schatzky distance is c. 10 mm; fissure angle is specimens of Belemnitella from the basal Campanian of 6-7°; alveolar angle is c. 20°; and the bottom of the the Munster Basin and showed that these specimens ventraJ fissure is almost straight. differed from both B. praecursor and B. mucronata. The Jeletzky (1955) discussed at length the evolution of specimens were referred to as B. aff. mucronata senior Santonian and Campanian species of Belemnitella and Nowak, B. aff. mucronata!praecursor, and B. aff. senior/ the concept of B. praecursor. His study was based mainly praecursor. on Russian material in addition to same few specimens Naidin (1964a) described and figured B. mucronata from England and Scania. Jeletzky's concept of B. alpha from the lower Lower Campanian of the Russian praecursor was broader than that of Stolley, and he also Flatform and included the English specimen, referred to included specimens of Belemnitella with more or less weil as B. ex gr. praecursor by Jeletzky (1955 , PI. 58:1) , in this developed vaseular markingsand longitudinal striae in B. subspecies. praecursor. According to Jeletzky, B. praecursor has a Christensen (1975a) described B. alpha from Scania Iong, more or less slender guard, which is slightly to and discussed the relationships of B. alpha, B. praecur moderately lanceolate in ventraJ view and slightly to sor, and B. mucronata. According to Christensen moderately conical in lateral view. The fissure angle is (1975a:52) B. alpha is closely comparable to B. praecur 5-10°, the alveolar angle a verages 20--22°, the Schatzky sor with respect to surface markingsand the y both lack a distance is 6-10 mm, and the bottom of the ventraJ fissure well-defined mucro. Christensen also claimed that the is straight or nearly straight. interna! characters were similar in the two species, and Jeletzky distinguished the following varieties of B. that the main characters separating B. alpha from B. praecursor: var. praecursor, var. media Jeletzky, 1955, praecursor isthestout guard. This supposed similarity is and var. mucronatiformis Jeletzky, 1955, but failed to now known to be incorrect. The comparison was based present an appropriate biometric analysis of the var on a biometric analysis of a sample of B. alpha from ieties. B. praecursor var. praecursor was said to be Scania and the available information of the variation of characterized by havinga completely smooth guard; var. B. praecursor. I have recent! y analysed the variation of a media by having numerous, faint, longitudinal striae in sample of B. praecursor from the basal Campanian of the addition to darso-lateral double furrows and single Hallembaye quarry in Belgium (see below) , and it is lateral furrows; and var. mucronatiformis by having a apparent that the important interna! characters of the fairly strongly sculptured guard with vaseular imprints two species differ. lt should be stressed here that and longitudinal striae and a more or less distinct mucro. although the two species differ in several critical charac Moreover, var. mucronatiformis is most! y s! en der and ters, it is not possible on the basis of on! y a few specimens samewhat lanceolate in ventraJ vie w. toassign them safely to either B. alpha or B. praecursor.

31 WALTER KEGEL eHRISTENSEN

Belemnitella alpha Naidin, 1964 Biometry. - The specimens from Kullemölla and Lyckås PI. 3, fig . 5; PI. 4, figs 1-3 were analysed biometrically as a single sample. Measure ments of a small sample of B. alphafrom Ringelestätt in 1894 Belemnitella mucronata Schlotheim the Kristianstad Basin, analysed by Christensen (1975a), Moberg, p. 77 are also reportedfor comparison. 1935 Belemnitella mucronata (Schlotheim) - Hägg, p. 65 ; PI. 10:8-9 1955 Belemnitella ex gr. B. praecursor Stolley - Jeletzky , PI. 58: l 1956 Belemnitella mucronata alpha Schatzky Univariate analysis (nom. nud.) - Naidin , p. 338 1959 Belemnitella mucronata alpha Shatskij B.alpha from Kullemölla-Lyckås: (nom. nud.) - Moskvine & Najdine , p. 505 e haracter N x SD ev OR 1960 Belemnitella mucronata alpha Schatzky LAP 12 55. 1 10.1 18.3 41.8-68.0 (nom. nud.) - Naidin, p. 51 DVDP 12 15.8 3.6 22.8 11.5-20.3 1964a Belemnitella mucronata alpha Naidin: pp. SD 7 11.0 1.3 11.8 9.6-12.6 89-90; PI. l :2 FA 3 13.3 0.6 4.4 13.0-14.0 1964b Belemnitella aff. muronata senior Nowak - AA Il 20.3 1.1 5.4 18.5-22.0 Ernst, pp. 181-182; PI. 1:2, 5 1974 Belemnitella mucronata alpha Schatzky B. alphafrom Ringelestätt (Christensen 1975a): (nom. nud)- Naidin, p. 218 , PI. 75 :3 e haracter N x SD ev OR 1975a Belemnitella alp ha Naidin- Christensen, pp. LAP 23 53.3 12 .1 22.7 33 .8-73.0 51-52; PI. 6:1-6 DVDP 23 15 .2 3.2 21.1 9.8-19.8 1980 Belemnitella mucronata alpha Schatzky so 19 10 .3 2.3 22.3 6.5- 13 .6 (nom. nud .) - Naidin , p. 91 ; PI. 10:8-9; PI. FA 21 12 .5 5.4 43.2 5.0-24.0 11:7; Textfigs 21:6, 22:3-5 AA 19 20.0 1.3 6.5 17.0-22.0

History and type. - B. mucronata alpha was used as a Bivariate analysis manuscript name by Schatzky in 1924 (see Naidin The scatter plot of length from apex to protoconch vs. 1964a:89, footnote 2). Naidin (1964a:89; PI. 1:2) dorso-ventral diameter at protoconch of the specimens described and figured the taxon under discussion and from Kullemölla-Lyckås and Ringeleslätt is shown in Fig. hence Naidin was regarded to be the author by Christen 15. The regression Iine of the Ringeleslätt sample is also sen (1975a). A lectotype should be designatedfrom the shown and the estimate of the statistical parameters of collection of D. P. Naidin. this sample are given bel o w. Material. - Kullemölla: 14 specimens; Lyckås: 7 speci mens. B. alpha from Ringeleslätt (Christensen 1975a): Short description.- A Belemnitella with a !arge, ventrally DVDP = 2.2011 + 0.2441 LAP; SDa = 1.2355; SDb = flattened guard, which is almost cylindrical or very 0.0226; SDyx = 1.2828; r = 0.9204; N = 23. slightly lanceolate in ventrat view , and high conical in The value of the earrelation coefficient is very highly lateral vie w. Common ly a mucro is on ly slightly delimited significant, while a t-test on the y-intercept gave a value or not present. which is not significant at the 5% leve) , implying an The relationship of length from apex to protoconch vs . isometric relationship of the two characters (Christensen dorso-ventral diameter at protoconch is isometric (see 1975a:52). below), and the ratio of these characters varies from about three to about four. Discussion. -The mean values of Iength from apex to The shape of the bottom of the ventrat fissure generally protoconch, Schatzky distance, fissure angle, and alveo is straight or may be straight with an outward bend just lar angle of the specimens from Kullemölla-Lyckås and below the ventrat surface. The walls of the alveolus may the sample from Ringeleslätt were campared by t-tests, be covered by conellae. The fissure angle is small, the which showed no significant differences at the 5% leve!. Schatzky distance is !arge, and the alveolar angle is about It is obvious from Fig. 15 that the specimens from 20° (see below). Kullemölla-Lyckås are scattered around the regression The surface of the guard normally has weakly Iine of B. alpha from Ringeleslätt. The specimens from developed vaseular imprints and longitudinal striae. Kullemölla-Lyckås are therefore referred to B. alpha.

32 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TRO UGH

DVDP in mm 20

8 • • 4 2

LAPin mm 10 20 30 40 50 60 70 80

Fig. 15. Scatter diagram and regression Iine of Belemnitellaalpha and Belemnitella praecursor. LAP = length from apex to protoconch; DVDP = dorso-ventral diameter at protoconch; + = mean values. o = B. alpha from Kullemölla/Lyckås; • = B. alphafrom Ringeleslätt, Kristianstad Basin; • = B. praecursor from the C.P.L. quarry at Hallembaye, Belgium ; o = B. praecursor from Russian Asia; * = holotype of B. praecursor. Regression Iine marked l is calculated on the basis of a sample of B. alphafrom Ringeleslätt, and regression Iine 2 on the basis of a sample of B. praecursor from Hallembaye.

Affinity.- B. alpha is closely comparable to B. praecur value of 3.1167 with 44 degrees of freedom, w hi ch is very sor and B . mucronata. The variation of a sample of B. high ly significant (P<0.1%), implying an allometric rela praecursor from the basal Campanian of the C.P.L. tionship of the two varia tes. This means that the ra tios of quarry at Hallembaye in Belgium was analysed recently length from apex to protoconch divided by dorso-ventral (Christensen & Schmid, in prep.), and the results are diameter at protoconch change during growth. Juvenile reported below. On e specimen from Hallembaye is and actaloscent specimens generally are slimmer than figured on PI . 5:1 . adult specimens. The holotype of B. praecursor (PI. 3:4) and some few specimens of B . praecursor from Russian Asia (Emba Univariate analysis and west Kazakhstan areas) are also plotted in Fig. 15 B. praecursor from Hallembaye quarry: (see also PI. 5:2). The holotype is situated very close to Charact er N x SD ev OR the regression Iine of B. praecursor from Hallembaye. LAP 46 60.9 11.8 19.4 20.1-79.2 The Russian specimens are scattered around the regres DVDP 60 15.3 3.6 23.3 4.9-19.0 sion Iine of B. praecursor. They are generally slimmer, SD 57 6.4 1.3 20.9 4.2-10.2 FA 52 17.6 5.0 28.2 10.0--28.0 however, than the Hallembaye specimens, and most of AA 59 20.1 0.9 4.3 18.0--22.0 them are situated below the regression line. The sample of B. praecursor from Hallembaye was campared to the sample of B. alphafrom Ringeleslätt. Bivariate analysis The t-tests on the mean values of length from apex to The scatter plot of length from apex to protoconch vs . protoconch, Schatzky distance, and fissure angle showed dorso-ventral diameter at protoconch of the sample of B. that the mean values differed significantly at the 5% praecursor is shown in Fig. 15 , as is the regression Iine. leve!. The regression Iines of the two samples were also The equation on the regression line is as follows. compared. No significant differences were found in the B. praecursor from Hallembaye: variances and slopes of the two lines. A t-test on the position of the lines, however, gave a value of 7.634 with DVDP = -2.0831 + 0.2854 LAP; N= 46; r= 0.9208; 65 degrees of freedom, which is very highly significant SDa = 0.6684; SDb = 0.0108; SDyx = 0.8534. (P<0.1%). This means that B. alphagenerally is more The value of the earrelation coefficient is very highly stout than B. praecursor. significant (P<0.1%). A t-test on the y-intercept gave a On the basis of the uni-and bi vari a te analyses i t can be

33 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH concluded that B. praecursor is moreslender, has a smal two alveolar and one apical fragment (see PI. 3:3). These ler Schatzky distance, and a larger fissure angle than B. have weakly developed vaseular imprints and well alp ha. Moreover, B. praecursor reaches a larger size of developed longitudinal striae. The two alveolar frag the guard than B. alpha. This is evident both from the ments see m to have small fissure an gles and the bottom of mean values of the length from apex to protoconch and the ventral fissure seems to be straight in both specimens. the observed range of this characters. Furthermore, the One of the alveolar fragment has a thin 'white-layer' and relationship of length from apex to protoconch vs. dorso part of the bottom of the alveolus is covered by small ventral diameter at protoconch is isometric in B. alpha conellae. and allometric in B. praecursor. The four specimens are referred to as B. ex gr. alphal B. alpha differs from B. mucronata in lacking a well praecursor, because they have weakly developed vaseu defined mucro and by having weaker developed vaseular lar imprints and well-developed longitudinal striae. In imprints. The relationship of length from apex to pro addition, the fissure angle seems to be small, the bottom toconch vs. dorso-ventral diameter at protoconch of the of the ventral fissure seems to be straight, and two speci two species is similar, but B. alphahas alarger guard than mens have conellae in the alveolus. As mentioned above B. mucronata. In addition, B. alphahas alarger Schatzky it is not possible on the basis of a few specimens of distance and a smaller fissure angle than B. mucronata Belemnitella toassign them safely to either B. alpha or B. (see Christensen, 1975a:52). praecursor. The specimen from Kåseberga was earlier referred to Distribution. - B. alpha has its main distribution in the B. mucronata by Hägg (1939) , as were the specimens Central Russian Subprovince and in Balto-Scandia. lt from the Kullemölla bo ring (Lundegren 1935b). has been recorded alsofrom northern German y and Eng land. Stratigraphy. - The range of B. alp/w is discussed above. The stratigraphic range of B. alpha in western Europe B. praecursor occurs in Western Germany in the G. was discussed by Christensen (1975a) who concluded that granulataquadrata Zone and the succeeding J. lingua/G. the species occurs in the lower Lower Campanian. At quadrata Zone (Ernst 1946b) . A single specimen was Kullemölla-Lyckås the taxon occurs tagether with G. collected from the G. senonensis Zone of Lägerdorf (M. granulataquadrata from the basal Campanian. In Russia, G. Schulz, pers. comm.). Jarvis (1980) recorded B. B. alpha occurs in the upper part of the lower Lower praecursor from the Lower Campanian phosphatic chalk Campanian and the middle Lower Campanian and is of France where it occurs tagether with G. q. quadra ta used as an index fossil for the middle Lower Campanian (Fig. 7) . Fletcher & Wood (1978:93 and Fig. 17) recorded (e.g. see Naidin & Kopaevich 1977; Naidin 1979, 1983). B. praecursor from the Boheeshane A Chalk from the Lower Campanian of Northern Ireland (Fig. 7). In Russia, B. praecursor appears in the Upper Santonian and continues into the lower and middle Lower Campa nian (Naidin & Kopaevich 1977; Naidin 1979, 1983). Belemnitella ex gr. alpha Naidinlpraecursor Stolley PI. 3, fig . 3; PI. 4, fig. 4 Material. - Kåseberga: one specimen; Kullemölla bor Belemnitella taxa from the highest Lower Campanian ing, 0--20 m interval: three specimens. Lower Maastrichtian Remarks.-The specimen from Kåseberga (PI. 4:4) is More than a score of species, subspecies, and varieties of relatively stout and the LAP:DVDP ratio is 3.4 Belemnitella from the highest Lower Campanian-Lower (52.2:15.4). The Schatzky distance is about 10 mm. The Maastrichtian have been erected, and the majority of fissure angle cannot be measured because the anterior these taxa were established by east european workers. part of the alveolus is missing. However, the fissure angle The systematics of many of these taxa are in a state of seems to have been small and the bottom of the ventral disorder and they are poorly understood. fissure seems to have been straight. The guard is slightly Some species and subspecies have received recognition lanceolate in ventral vie w and high conical in lateral vie w. in many publications after they were established (e.g. B. It has well-developed longitudinal striae. The 'white 'minor' Jeletzky and B. 'langei' Jeletzky). Other species layer' is thin and conellae cover part of the wall of the names have been used recently in publications by west alveolus. european authors ( e.g. B. najdini Kongiel and B. post The material from the Kullemölla boring consists of erior Kongiel; see Fletcher & Wood 1978; Schulz 1978,

34 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

1982; Schulz & Schmid 1983). Some of these taxa are 1964b Belemnitella mucronata mucronata discussed below and in the description of B. mucronata (Schlotheim) - Ernst, PI. l :6 1972 Belemnitella mucronata (Link) sensu lato - and aff. B. langei. Christensen, p. 322 , Fig. l B. 'minor' and B. 'langei' are used as index fossils in the 1974 Belemnitella mucronata mucronata Upper Upper Campanian, but the concept of the two (Schlotheim)- Naidin , p. 216; PI. 74:4-5 , PI. species was in a state of disorder until recent! y. Christen 76:4-5 sen et al. (1975) discussed the two taxa and pointed out 1975a Belemnitella mucronata mucronata (Link) Christensen, p. 32 ; PI. 7:1-3, PI. 8:1-4, PI. that they can only be interpreted with respect to their 9:1-6, PI. 10 :1- 2, PI. 11:1-3, Figs 22A-B holotypes, because statistically evaluated populations 1975 Belemnitella mucronata mucronata from the respective type localities are not available. The (Schlotheim) - Christensen et al. p. 40 ; PI. interna! characters of the holotypes were unknown 1:1-3, PI. 2:1-2, PI. 3:1-5, Fig. 3 before 1975. 1979a Belemnitella mucronata mucronata (Schlotheim) - Naidin, p. 85; PI. 1:7, PI. 3:9 The holotypes have now been split and it was revealed 1980 Belemnitella mucronata mucronata by Christensen et al. (1975) that the interna! characters of (Schlotheim) - Naidin p. 89; PI. 2:3, 8 the holotype of B. minor do not fit with those given in the original diagnosis (Jeletzky 1951) . Moreover, it was also shown that the holotype of B. minor falls within the History and type. - It is generally agreed that B. muc variation of the 'type population' of B. mucronata pro ronata should be interpreted with reference to the speci posed by Christensen et al. (1975). mens figured by Arkhangelsky (1912, PI. 9:3, 9, 23, 26, With respect to B. langei, Christensen et al. (1975) PI. 10:10). (See Birkelund & Rasmussen 1956; Birkelund showed that the interna! characters of the holotype agree 1957; Jeletzky 1964; Naidin 1971; and Christensen et al. weil with the original diagnosis for the species (Jeletzky 1973.) Christensen et al. (1975) therefore proposed a 1948). They concluded that it is a well-defined species, neotype for B. mucronata from the lower part of the and the main characters separating it from B. mucronata Upper Campanian of the quarry GERMANIA IV at are the !arge relative length, the !arge fissure angle, the Misburg/Hannover, NW Germany. In the present paper bottom of the ventraJ fissure which is complexly bent, B. mucronata is interpreted with respect to the proposed and its lanceolate shape in ventraJ view. neotype and type-series. The concept of B. 'minor' and B. 'langei' were misin Material.- Rödmölla-Tosterup, loc. 2: se ven specimens; terpreted by most earlier workers. According to Schulz Köpinge: c. 10 specimens; and Valleberga-Ingelstorp: 21 (1978:78, 81) B. langei sensu Jeletzky probably only specimens. occurs in western Europe at a specific horizon at Kroons moor (West Germany), and all other records of B. langei Description. - A Belemnitella with a !arge, ventrally from western Europe should probably be assigned to B. flattened guard, which is almost cylindrical or slightly najdini. lanceolate in ventraJ vie w and high conical in lateral vie w. A mucro is well-defined. Belemnitella mucronata (Schlotheim, 1813) The relationship between length from apex to pro PI. 5, figs 3-4; PI. 6, figs 1-2 toconch and dorsoventral diameter at protoconch is isometric, and the ratio of these characters varies from 1807 Belemnites mucronatus Link, p. 9 1885 Belemnitella mucronata (Schlotheim) - about 2.5 to about 4.5 , being 3.0--3.7 in most specimens. Moberg, p. 56; PI. 6:13, non PI. 6:14-16, 19 The shape of the bottom of the ventraJ fissure is vari 1912 Belemnitella mucronata (Schlotheim)- Ark able: it may be straight, straight with an outward bend hangelsky, p. 600; PI. 9:3, 9, 23, 26; Pl.l0:10 just below the ventraJ surface, undula ting, or s-shaped. It 1913 Belemnitella mucronata mut. senior Nowak, is , however, straight or straight with an outward bendin p. 395; PI. 42 :22 1915 Belemnitella mucronata var. panderosa Sin most specimens. The fissure angle varies from 5-47°, bu t zow, p. 147; PI. 8:11- 12 is commonly between 15-30°. The Schatzky distance var 1951 Belemnitella mucronata mut. senior Nowak ies from 4.5-15.0 mm, and is generally between 5.5-10.0 - Jeletzky, p. 81 ; PI. 2:1, ? PI. 1:4 mm with a mean value of about 8 mm. The alveolar angle 1955 Belemnitella mucronata (Schlotheim) un varies from 18.0--24.0° with a mean value of about 20°. named early variety- Jeletzky, p. 480 ; PI. 57:1 Adult specimens have distinct vaseular imprints, 1964 Belemnitella mucronata (Link) - Jeletzky, dorso-lateral double furrows and double depressions. Fig. l , PI. 1:1-4 Longitudinal striae usually are present.

35 WALTER KEGEL CHRISTENSEN

Remarks. -The variation of three samples of B. muc fissure angle, for example, is larger (c. 40--90°) in B. ronata from the highest Lower Campanian-Lower posterior than in B. mucronata. Specimens of Belem Upper Campanian of Scania and German y was analysed nitella very closely comparable to B. mucronata from the by Christensen (1975a) and Christensen et al. (1975) who basal Maastrichtian of Northern Ireland, Western Ger demonstrated significant differences in various charac many, and Belgium we re tentative! y assigned to B. post ters. For example, specimens from Scania attain alarger erior by Fletcher & Wood (1978), Schulz (1978) and size of the guard t han the specimens from German y. Schulz & Schmid (1983). No evidence was presented that Christensen (1975a) suggested that the differences might the interna! characters ofthese specimens were studied. I be due to ecological conditions or that the samples were have studied one specimen of Belemnitella from the basal of slightly different age. Additional work on the variation Maastrichtian of Sweden. The fissure angle of this speci of sam p les of Belemnitella of the same age from different men is 15 .0°, and it was therefore referred to B. muc areas and samples of Belemnitella collected bed-by-bed ronata (Christensen 1975a, PI. 11:1). through a !arge, continuous section are necessary in B. minor, as interpreted with respect to the holotype, order to solvethis problem. Work is in progress on saru falls within the variation of B. mucronata. According to ples of Belemnitella collected bed-by-bed from the high the original diagnosis by Jeletzky (1951) B. minor is est Lower Campanian-Lower Upper Campanian of distinguished from B. mucronata by its greater fissure Misburg/Höver near Hannover by Dr. M.-G. Schulz and angle (gene rally mo re than 30--40°), and the bottom of the author. the ventraJ fissure which is complexly bent. Distribution.- B. mucronata is wide-spread in the North Affinity. - B. mucronata is closely allied to B. praecursor European Province and is also recorded from the north and B. alpha (see discussion above) . ern part of the Tethyan Realm. Nowak (1913) distinguished B. mucronata mut. senior It has its main occurrence in the highest Lower Cam Nowak from the lower 'Mucronatenkreide' and B. muc panian-Lower Upper Campanian and is used as a guide ronata mut. junior Nowak from the upper 'Mucronaten fossil for this part of the stratigraphic column. The taxon kreide', m addition to Belemnella lanceolata is also recorded from the Upper Upper Campanian and (Schlotheim) from the middle 'Mucronatenkreide'. basal Maastrichtian. According to Nowak (1913 :402) m ut. senior is most Iikely identical to B. mucronata sensu Arkhangelsky. Bir kelund (1957) expressed the same point of view. Chri stensen et al. (1975) discussed the relationship of B. Belemnitella aff. langei Jeletzky, 1948 mucronata and B. senior and concluded that it remains PI. 6, figs 3-4; PI. 7, figs 1-4 questionable whether B. senior really is an independent 1885 Belemnitella mucronata Schlotheim- Moberg, p. species or whether it is an extreme variant of B. muc 56; PI. 6:14-16, 19; non PI. 6:13 ronata. On the basis of the descriptions and figures of the two taxa by Arkhangetsky (1912) and Nowak (1913), Material. - Köpinge sandstone at Köpinge and Val respectively, in addition to the statement by Nowak, B. leberga - Ingelstorp: c. 25 specimens. senior is here considered a junior synonym of B. muc Diagnosis. -A Belemnitella with a small, slender guard ronata. On the other hand, !arge, stout, strongly vas which is markedly lanceolate in ventraJ view and ven cularized specimens of Belemnitella do occur around the trally flattened. Vaseular imprints are distinct and adult Lower-Upper Campanian boundary in various areas, specimens are provided with a mucro. The fissure angle is e.g. England, Germany, and Sweden. These specimens small and the bottom of the ventraJ fissure is straight. earlier were referred to as B. mucronata senior or B. senior by authors. F u ture studies ma y show if these speci Description.- A Belemnitella with a small slender, ven mens should be referred to a new species or subspecies, trally flattened guard which is markedly lanceolate in or merely considered a morphological variant of B. ventraJ view and cylindrical or high conical in lateral mucronata. vie w. Adult specimens are provided with a distinct B. posterior was established by Kongiel (1962) from mucro. the uppermost Campanian and lower Maastrichtian of The relationship between length from apex to pro Poland. According to Kongiel, B. posterior shows strong toconch and dorso-ventral diameter at protoconch is resemblance to B. mucronata in its externa! characters isometric (see below), and the ratio of these characters but differs fromthat species in its interna! characters. The varies from 3.9 to 5.8, being 4.3-5.0 in most specimens.

36 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

pVDP 1nmm 12

LAP in mm

10 20 30 40 50 60

Fig. 16. Scatter diagram and regression Iine of Belemnitella aff.langei from the Köpinge sandstone. LAP = length from apex to protoconch; DVDP = dorso-ventral diameter at protoconch; + = mean values. o = specimens from 'Köpinge' ; • = specimens from Valleberga/Ingelstorp. The specimen marked l was figured by Moberg (1885 , PI. 6: 14) as B. mucronata. This specimen was subsequently referred to as B. n. s p. aff. mucronata (=B. pseudolanceo/ata Jeletzky) by Jeletzky (1951) and Naidin (1960) , B. mucronata var. mobergi by Nikitin (1958) , and B. minor by Kongiel (1962). The specimen is refigured herein on PI. 6:3. For comparison the holotypes of B. langei (.å.2) and B. minor (.å.3) are also plotted.

The ventraJ fissure generally is straight and the fissure The holotypes of B. langei and B. minor are also plot angle is small (see below). The Schatzky distance varies ted in Fig. 16 for comparative purpose. The holotype of from about 5-10 mm with a mean value of about 7 mm. B. langei (PI. 7:5) is situated very close to the regression The alveolar angle varies from 18-22° with a mean value Iine and is within the range of B. aff. langei. The holotype of about 20°. of B. minor is also situated close to the regression Iine bu t Adult specimens have distinct vaseular imprints, falls outside the range of B. aff. langei. darso-lateral double furrows and darso-lateral double Remarks. - B. aff. langei is closely comparable to B. depressions. Longitudinal striae usually are present. !ange i with re gard to overall shape and size of the guard, Bi ornetry. -The specimens from the Köpinge sandstone but differs from that species by having a smaller fissure at Köpinge and Valleberga-Ingelstorp were treatedasa angle, and the bottom of the ventraJ fissure is straight in single sample and analysed biometrically. contrast to B. langei. B. aff. langei may be considered as an ancestor of B. langei. The taxan is referred to as B. aff. Univariate analysis langei due to the limited amount of material the strati graphic contra! of which is poor. Character N x SD ev OR LAP 21 44.5 6.1 13.8 31.6--54.8 Affinity. -B. aff. langei is distinguished from B. muc DVDP 23 9.4 2.4 26.1 7.1-12.2 ronata by its small slender guard which is markedly lan SD 14 7.1 1.7 23.4 4.9-10.2 ceolate in ventraJ vie w. With re gard to surface markings FA lO 17.4 6.4 3.7 9.5-31.0 AA 14 19.8 l. O 4.9 18.0--22.0 and inner characters, B. aff. langei and B. mucronata are closely comparable. Bivariate analysis B. aff. langei differs from the holotype of B. langei and The scatter diagram of Iength from apex to protoconch specimens of B. langei sensu Naidin from Russia (see PI. vs. dorso-ventral diameter at protoconch is shown in Fig. 7, figs 5-7) by its small fissure angle and being more 16. The equation of the regression Iine is given below. flattened ventrally. Moreover, the bottom of the ventraJ fissure is straight and not complexly bent as in B. langei. DVDP = - 1.0310 + 0.2425 LAP; N= 21; r= 0.8636; B. aff. langei and B. pseudolanceolata Jeletzky are SDa = 1.4746; SDb = 0.0328; SDyx = 0.9018. closely comparable with respect to shape of guard, sur The value of the earrelation coefficient with 19 degrees face markings and size of fissure angle. B. aff. langei, of freedom is very highly significant (P<0.1%). The t however, differs from B. pseudolanceolata by havinga test on the y-intercept gave a value of 0.6992 with 19 smaller guard and a larger Schatzky distance. degrees of freedom, which IS not significant B. aff. langei shows resemblance with regard to shape (50%> P>40% ), implying an isometric relationship of of guard to B. mucronata var. mobergi, B. gorkiana, and the two characters. B. mucrolanceolata from the Upper Campanian of the

37 WALTER KEGEL CHRISTENSEN

Ukrainian Syneclise. These taxa were erected by Nikitin Distribution. - B. aff. langei has up to now only been (1958), but unfortunately were inadequately described recorded from the Vomb Trough in Scania. Here, it and figured; they are therefore poorly understood. occurs in the upper part of the Köpinge sandstone. At According to Naidin (1974) these three taxa are junior Valleberga-Ingelstorp, B. aff. langei and B. mucronata synonyms of B. l. langei, B. langei minor, B. pseudolan occur together; B. mucronata is the dominant form in the ceolata, and possibly B. langei najdini. lower part of the sequence and B. aff. langei in the upper Specimens very closely comparable to B. aff. langei part. The zone with B. mucronata and B. balsvikensis is occur in the uppermost Lower Campanian and Lower not identified in the Valleberga-Ingelstorp area (see Upper Campanian in Scania (see Christensen 1975a:56; above). It is therefore suggested t hat B. aff. lang ei occurs PI. 9:3, 6; PI. 11:2-3). These specimens were considered in the middle Upper Campanian, that is in beds w h ich can to be morphological variants of B. mucronata because be correlated with the upper part of the B. mucronata they only constitute an insignificant part of the Belem Zone and possibly the lower part of the B. 'minor' Zone nitella samples from these stratigraphic horizons, and (Fig. 9). they fall within the variation of B. mucronata in other important characters.

Sommary

- Belerunites from five localities and one boring in the subspecies are discussed. It is concluded that subspecies Vomb Trough have been studied. The localities, most of dnestrensis should be placed in synonym y of A . v. verus which lie in the periphery of the trough very close to the and that the subspecies fragilis is dubious. Samples of large-scale Fyledalen Fault, are described and placed in Gonioteuthis from outcrops in the trough are campared an international stratigraphic fra·mework on the basis of to Germansamples from well-known stratigraphic levels. the belemnites. The study is based mainly upon material The subgenus Actinocamax (Paractinocamax) of Naidin collected in the last half of the 19th century and the first is discussed and considered to be a synonym of Belem third of the 20th century. Earlier records of other strati nellocamax. The stratigraphic range of B. ex gr. graphic important fossils , such as ammonites and echino grossouvrei is fully discussed. derms, are reviewed. Two species of Belemnitella from the lower and middle The belemnite faunas are correlated with belemnite Lower Campanian are tentatively recognized: B. alpha faunas from western Europe and the Russian Platform, and B. praecursor. The measurements of critical charac but formal belemnite zones are not established. ters of the holotype of B. praecursor are reported. The The depositional history of the Vomb Trough during relationship of B. alphaand B. praecursor is full y discus the Upper Cretaceous is outlined. The dominant sedi sed on the basis of the available literature, in addition to ments are glauconitic, calcareous, clayey siltstones depo biometric analysis of samples of B. alphaand B. praecur sited in a fairly shallow-water environment, and the max sor, respectively. imum thickness of the Upper Cretaceous is a little less Belemnitella taxa from the highest Lo wer Campanian than 800m. Several conglomerates, some of which are of Lower Maastrichtian are discussed, and the concept of considerable thickness, and non-sequences occur in the many of these taxa is shown to be in a state of chaos. B. Santonian and Campanian. The youngest sediments are mucronata senior or B. senior is considered a synonym of from the Maastrichtian, probably the Lower Maastrich B. mucronata. The interna! characters of the holotype of tian. The sea appears to have withdrawn from the trough B. minor do not fit with those given in the original diag in the Upper Maastrichtian, as it also did in the Kristian nosis, and i t falls within the variation of B. mucronata. B. stad Basin farther northeast. langei and B. posterior are also discussed. Thirteen belemnite taxa, representing the genera B. aff. langei is closely comparable to B. langei in its Actinocamax Miller, Gonioteuthis Bayle, Belemnello externa! characters, by differs from that species in its camax Naidin, and Belemnitella d'Orbigny, are described inner characters. Other species or subspecies of Belem or discussed. nitella with guards markedly lanceolate in ventraJ view The nominate subspecies of A. verus and two other from the Upper Upper Campanian are also discussed.

38 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

ACKNOWLEPGMENTS

I am grateful to several friendsand colleagues for allowing me Prof. Dr. T . Birkelund read the manuscript critically and to study belemnites in their care. offered helpful suggestions. Dr. J .S. Peel im proved the Eng I have benefited great ly from discussions with Dr. M.-G. lish. I wish to express my sincere thanks for this help. Schulz, Mr. C.J. Wood , Prof. Dr. F. Schmid, Prof. Dr. G. I gratefully acknowledge the technical assistance of Mr. C. Ernst, Prof. Dr. D.P. Naidin, and Dr. J .A. Jeletzky about Rasmussen, Mr. S.L. Jakobsen, Mr. J. Aagaard, and Mr. J. systematic and stratigraphic problems connected with Upper Sommer with preparation of illustrations. Cretaceous belemnites.

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39 WALTER KEGEL CHRISTENSEN

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BERGSTRÖM , B. systematisk framstäldt. - Sver. geol. unders. C63 , 45 pp. Stock HOLLAND , K. LARSSON , E. NORUNG and U. SIVHED: Guide to excur holm. (In Swedish.) sions in Scania. - Sver. geol. unders. Ca54, 75-81. Uppsala. - 1885: Cephalopoderna i Sveriges kritsystem. 2. Artsbeskrifning. - NORLING, E ., and SKOG LUND, R., 1977: Der Siidwestrand der Sver. geol. unders. C73 , 65 pp, Stockholm. (In Swedish.) Osteuropäischen Tafel im Hereich Schwedens. -Z. angew. Geol. - 1888: Om fördelningen af Sveriges vigtigare kritförekomster på två 23, 449-458. Berlin. skilda bäcken. - Geol. Fören. Stockh. Förh. 10 , 308-327. Stock NOWAK , J. , 1913: Untersuchungen iiber die Cephalopoden der oberen holm.. (In Swedish.) Kreide in Polen. 3. Teil. -Bull. Acad. Sci . Cracovie B, 335-412. -1894: Uber schwedische Kreidebelemniten. - Neues Jb. Miner. Geol. Cracovie. Paläont. 2, 69-78. Stuttgart. 0BRADOVICH, J .D., and COBBAN , W.A. , 1975: A time-seale for the Late 41 WALTER KEGEL CHRISTENSEN

Cretaceous of the Western Interior of North America.- Geol. Ass. SCHULZ , M.-G., and ScHMID , F. , 1983: Das Ober-Maastricht von Can. Spec. Pap. 13 , 31- 54. Montreal. Hemmoar (N-Deutschland): Faunenzonen-Gliederung und Korre 0RBIGNY , A. o ', 184(}-1842: Paleontologie fran~aise: Terrains cretaces. lation mit dem Ober-Maastricht von Dänemark und Limburg. - l. Cephalopodes. 1-120 (1840) ; 121-430 (1841); 431-662 (1842). News!. Stratigr. 13 , 21- 39. Berlin/Stuttgart. Paris. SHARPE, D., 1853: Description of the fossil remains of Mollusca found in - 1842-1843: Paleontologie fran~aise: Terrains cretaces. 2. Gas the Chalk of England. Cephalopoda.- Palaeontogr. Soc. Mem. 68 teropodes. 1-224 (1842) ; 225-456 (1843).- Paris. pp. London. 1848-1851: Paleontologie fran~aise: Terrains cretaces. 4. SIMPSON, G.G., RoE, A., and LEWONTIN , R.C. , 1960: Quantitative Brachiopodes. 1-32 (1847(1848)); 33-104 (1848); 105-390 (1851). Zoology. Revised Edition. 440 pp.- New York/Chicago/San Fran Paris. cisco/Atlanta: Harcourt, Brace & World Inc. - 1852: Cours Elementaire de Paleontologie et de Geologie Strati SINzow , I. , 1915: On the Upper Cretaceous sediments of the Saratov graphique 2, 383-847. -Paris. province. - Zapiski Imp. Miner. Ob-wa, Ser. 2, 50 , 133-162. St. PEAKE, N.B. , and HANCOCK , J .M. 1970: The Upper Cretaceous of Petersburg. (In Russian.) Norfolk. Revised edition. Trans. Norfolk Norwich nat. Soc. SoKAL, R.R. , and RoHLF, F.J., 1969: Biometry- The Principles and 293-339. Norfolk. practise of stalistics in biological research. 776 pp.- San Francisco: ROBASZYNSKI, F., (Coordinateur) 1982: Le Turanian de la region-type: W.H. Freeman and Co. Saumurois et Touraine. Stratigraphie, biozonations, sedimen STOKES, R.B. , 1975: Royaumes et provinces fauniques du crt~tace et a b tologie.- Bull. Cent. Rech. Expl. Prod. Elf-Aquitaine 6, 119-125. lissur la base d'une etude systematique de genre Micraster.- Me m. Pau. Mus. nat. Hist. Nat. C31, 94 pp. Paris. RoHLF, F.J. , and SoKAL , R.R., 1969: Statistical Tables. 253 pp.- San - 1979: The echinoid genus Diplodetus from the Santonian to Danian of Francisco: W.H. Freeman and Co. Northern Europe. - Neues Jb. Geol. Paläont. Mh . 10, 619-630. ScHLUTER , C. , 1870: Berich t ii ber eine geognostisch-paläontologische Stuttgart. Reise im sudlichen Schweden. - Neues Jb. Miner. Geol. Paläont. STOLLEY, E., 1896: Einige Bernerkungen ii ber die o be re Kreide Jg. 1870, 929-969. Stuttgart. insbesondere von Liineburg und Lägerdorf.- Arch. An trop. Geol. -1876: Cephalopoden der aberen deutschen Kreide. Teil2.- Palaeon Schlesw.-Holst. l , 139-176. Kiel/Leipzig. tographica 24 , 123-263. Stuttgart. -1897: Ueber die Gliederung des norddeutschen und baltischen Senon - 1897: U e ber einige exocyclische Echiniden der baltisch en Kreide und sowie die dasselbe characteriserenden Belemniten. - Arch. deren Bett.- Z . Dt. Geol. Ges. Jg. 1897, 18-50. Berlin. Antrop. Geol. Schlesw.-Holst. 2, 216--302. Kiel/Leipzig. SCHMID, F., and ERNST , G. , 1975: Ammoniten aus dem Campan der - 1916: Neue Beiträge zur Kenntnis der norddeutschen aberen Kreide. Lehrter Westmulde und ihre stratigraphische Bedeutung. l. Teil: I-IV.- Jber. Niedersächs. Geol. Ver. 9, 95-104. Hannover. Scaphites, Bostrychoceras und Hoplitoplacenticeras.- Ber. Naturh. - 1930: Einige Bernerkungen iiber die Kreide Siidskandinaviens. - Ges. 119, 315-359. Hannover. Geol. Fören. Stockh. Förh. 52, 157-190. Stockholm. ScHULZ, M.-G., 1978: ZurLitho-und Biostratigraphie des Obercam -1932: Die Belemniten De Geer's von Båstad.- Geol. Fören. Stockh. pan-Untermaastricht von Lägerdorf und Kroonsmoor (SW-Hol Förh. 54 , 498-499. Stockholm. stein). - News!. Stratigr. 7, 73-89. Berlin/Stuttgart. ULBRICH, H., 1971: Mitteilungen zur Biostratigraphie des Santon und - 1979: Morphometrisch-variationsstatistische Untersuchungen zur Campan des mittieren Teils der Subhercynen Kreidemulde. - Phylogenie der Belemniten-Gattung Belemnella im Untermaas Freiberger Forschungshefte C267, 47-71. Berlin. tricht NW-Europas.- Geol. Jb. A47, 3-157. Hannover. WAHLENBERG , G., 1818: Petrificata telluris Svecanae. 116 pp.- Upp - 1982: Erster Nachweis der Belemnitengattung Belemnitella (B. sala. pulchra n. sp.) im mittieren Untermaastricht NW-Deutschland. Wooo, C .J., 1981: Upper Cretaceous. In P. KENT (Ed.): E astern Eng Geol. Jb. A61, 279-293. Hannover. land from the Tees to the Wash, 92-115. - Inst. Geol. Sci. London. SCHULZ, M.-G., ERNST, G. , ERNST , H., and ScHMID , F., 1984: Conia WRIGHT , C. W. , and KEN NED Y, W.J. , 1981: The Ammonoidea of the cian to Maastrichtian stage boundaries in the standard seetian for Plenus Maris and the Middle Chalk. - Palaeont. Soc. Monogr. 148 the Upper Cretaceous white chalk of NW Germany (Lägerdorf• pp. London. Kroonsmoor-Hemmoor): Definitions and proposals. -Bull. geol. Soc. Denmark 33 , 203-215. Copenhagen.

The International Commission on Zoological Nomenciature has desig nated, by use of its plenary powers, specimen number kca 5/2 in the collections of the Niedersächsisches Landesamt fiir Bodenforschung, Hannover, BRD, as neotypefor Belemnites mucronatus Schlotheim, 1813 (see Opinion 1328; Bull. zoo!. Nomencl. 42, 222-225, September 30 , 1985). The neotype was described and figured by Christensen et al. (1975 , PI. 1: 1).

42 UPPER CRETACEOUS BELEMNITES FROM THE VOMB TROUGH

PLATES 1-7 Photographed specimens were coated with ammonium chloride. Figures are of natural size except where stated.

43 WALTER KEGEL CHRISTENSEN

PLATE l

Fig. l. Actinocamax verus Miller from Lyckås. Basal Campa Fig. 7-8. Gonioteuthis granulataquadrata (Stolley) from nian. Lyckås. Basal Campanian. A: Dorsal view. A : Dorsal view. B: Lateral view. B: Lateral view. LM LO 5724t C: VentraJ view. D : View of the anterior end. Figs. 2-3. Gonioteuthis westfalica (Schli.iter) from the so-called 7D is x 1.5 and 8D is x 2. 'Westfalicus conglomerate' at Rödmölla-Tosterup. Lower Fig. 7: The specimen is deposited in GeologicaJ Institute, Stock Middle Santonian. holm. 2A: Dorsal view. Fig. 8 is MGUH 16887. 2B: View of the anterior end, x 2. 3A: VentraJ view. Fig. 9. Gonioteuthis quadrata scaniensis? Christensen from 3B: View of the anterior end, x 2. Rödmölla-Tosterup. Uppermost Lower Campanian. Fig. 2 is SGU Type 5208 . A: Dorsal view. Fig. 3 is SGU Type 5209. B: Lateral view. C: VentraJ view. Fig. 4. Gonioteuthis westfalicagranulata (Stolley) from Eriks D: View of the anterior end, x 2. daL Upper Middle Santonian. LM LO 5725t. A : Dorsal view . B: Lateral view. Fig. 10. Gonioteuthis sp. [possibly G. granulataquadrata C: View of the anterior end, x 2. (Stolley) or G. quadrata (Blainville)] from Kåseberga ~ Lower RM Mo 150734. part of Lower Campanian. A: Dorsal view. Fig. 5. Gonioteuthis westfalicagranulata (Stolley) or G. west B: Lateral view. falica (Schli.iter) from Eriksdal. C: VentraJ view. A: Dorsal view. D: View of the anterior end , x 2. B: Lateral view. The specimen is deposited in Geological Institute, Stockholm. C: View of the anterior end, x 3. The 'Riedel-Quotient' of this specimen is 16.8. The specimen Fig. 11. Gonioteuthis sp. from the Kullemölla boring, level200 was figured by Moberg (1885, PI. 5:20) as Actinocamax m. granulatus Blainville forma westfalica. A: Dorsal view. SGU Type 3902. B: Lateral view. C: VentraJ view. Fig. 6. Gonioteuthis westfalicagranulata (Stolley) from Eriks D: View of the anterior end, x 2. daL Upper Middle Santonian. SGU Type 5210 . A: Dorsal view. B: Lateral view . C: View of the anterior end, x 2. RM Mo 150744.

44 28 4C

5A 58

1A 18 2A 38 3A 4A 48 80 5C

6A 68 6C ?A 78 7C BA BB BC

100

9A 98 9C 10A 108 10C 110 11A 11 B 11C WALTER KEGEL CHRISTENSEN

PLATE 2

Fig. l. Belemnellocamax ex gr. grossouvrei (Janet) from 3C: Ventral view. Kullemölla. Basal Campanian. Dorsal view. The specimen was 3D: View of the anterior end, x 3. figuredas Actinocamax depressus Andreae by Hägg (1935 , PI. 4A: Lateral view. 10:5-6). 4B : Ventral view. LM LO 3237t. 4C: View of the anterior end, x 1.5 . 5A: Dorsal view. Fig. 2. Belemnellocamax ex gr. grossouvrei (Janet) from 5B: Lateral vie w. Kullemölla. Basal Campanian. 5C: Ventral view. A: Dorsal view. 5D: View of the anterior end, x 1.5. B: Lateral vie w. Fig. 3 is SGU Type 5211. C: Ventral view. Fig. 4 is SGU Type 5212. D: View of the middle part of the dorsal field showing granula Fig. 5 is SGU Type 5213. tion, x 2. E: View of the anterior end, x 2. Figs. 6-7. Belemnellocamax balsvikensis (Brotzen) from Röd• Holotype of Actinocamax mammillatus var. ornatus Moberg mölla-Tosterup, loc. 3. Lower Upper Campanian. (1885 , PI. 5:26). 6A: Dorsal view. LM LO 774t. 6B: Lateral vie w. 6C: Ventral view. Figs. 3-5. Belemnellocamax mammillatus (Nilsson) from Val Fig. 7 shows the inner characters of the pseudoalveolus, x 2. leberga. Uppermost Lower Campanian. Fig. 6 is MGUH 16888 (ex P. Gravesen Coli.) 3A: Dorsal view . Fig. 7 is MGUH 16889. 3B: Lateral vie w.

46 2E

1 2A 28 2C 3A 38 3C

4C 30 50 7

4A 48 5A 58 5C 6A 68 6C WALTER KEGEL CHRISTENSEN

PLATE 3

Fig. l. 'Actinocamax' lundgreni Stolley from the Kullemölla B: Lateral view. boring, leve! 440 m. C: VentraJ view. A: Dorsal view. D: View of the split anterior end showing inner characters. B: View of the anterior end, x 3. The specimen was figured by Stolley (1897 , PI. 3:24). [t is SGU Type 5214. housed in the Geologisch-Paläontologisches Institut, Kiel.

Fig. 2. Belemnitella propinqua (Moberg) from the Kullemölla Fig. 5. Belemnitella alpha Naidin from Kullemölla. G. boring, leve! 248 m. View of the split anterior end, x 3. granulataquadrata Zone of the basal Campanian. SGU Type 5215. A: Dorsal view . B: Lateral view. Fig. 3. Belemnitella ex gr. alpha Naidinlpraecursor Stolley C: VentraJ view, light from the left emphasizing longitudinal from the Kullemölla boring, G-20 m interval. View of the split striae. anterior end, x 3. D: VentraJ vie w, light from upper left corner emphasizing SGU Type 5216. vaseular imprints. E : View of the split anterior end showing inner characters, x 2. Fig. 4. Belemnitella praecursor Stolley from the Broitzem The specimen was figured by Hägg (1935, PI. 10:8- 9) as B. quarry at Braunschweig. G. granulataquadrata Zone of the mucronata (Schlotheim). basal Campanian. Cast of holotype. LM LO 3239t. A: Dorsal view.

48 18

2 3 40 4A 48

5A 58 5C 50 WALTER KEGEL CHRISTENSEN

PLATE 4

Figs. 1-3. Belemnitella alpha Naidin from the G. granulata Fig. 3. Specimen from Lyckås. quadrata Zone of the basal Campanian. A: Dorsal view. Fig. l. Specimen from Kullemölla. B : VentraJ view. A: Lateral view. C: Lateral view. B: VentraJ view. D: View of the split anterior end showing inner characters, x 3. SGU Type 5217. LM LO 5726t. Fig. 2. Specimen from Lyckås. A: Dorsal view. Fig. 4. Belemnitella ex gr. alpha Naidinlpraecursor Stolley B: Lateral view. from Kåseberga. Lower part of Lower Campanian. C: VentraJ view. A: VentraJ view. D: View ofthe split anterior end showing inner characters, x 3. B: View of the split anterior showing inner characters. MGUH 16890. The specimen was referred to as B. mucronata (Schlotheim) by Hägg (1939). RM Mo 158065

50 2A 28 2C

1A 18 20

3A 38 3C 30 4A 48 WALTER KEGEL CHRISTENSEN

PLATE 5

Fig. l. Belemnitella praecursor Stolley from the C.P .L. quarry Fig. 3. Belemnitella mucronata (Schlotheim) from the Köpinge at Hallembaye, Belgium. The specimen was collected from the sandstone at Valleberga. Uppermost Lower Campanian. 'Smectite' 25 cm below its upper limit. G. quadrata Zone of the A : Lateral view. lower Lower Campanian. B: VentraJ view . A : Dorsal view. SGU Type 5218. B: Lateral view. C: VentraJ view. Fig. 4. Belemnitella mucronata (Schlotheim) from the Köpinge D: View of the split anterior end showing inner characters, x sandstone at Köpingebro. The specimen presurnably came 1.5 . from the lower part of the Upper Campanian. MGUH 16891. A: Dorsal view. B: Lateral view. Fig. 2. Belemnitella praecursor Stolley from the Aktjubinsk C: VentraJ view. area, western Kazakhstan, USSR. Basal Campanian. D : View of the split anterior end showing inner characters, x A : Dorsal view. 1.5. B: Lateral view. RM Mo 158066. C: VentraJ view. D: View of the split anterior end showing inner characters, x 1,5. MGUH 16892.

52 2A 1A 18 1C 28 2C

40 3A 38 4A 48 4C WALTER KEGEL CHRISTENSEN

PLATE 6

Fig. l. Belemnitella mucronata (Schlotheim) from the Köpinge Fig. 3. Belemnitella aff. langei Jeletzky from the Köpinge sandstone at Valleberga. Basal Upper Campanian. sandstone at Köpinge. Probably middle Upper Campanian. A : Dorsal view. A: Dorsal view. B: VentraJ view. B: Lateral view. C: View of the split anterior end showing inner characters. C: VentraJ view. MGUH 16893. D: View of the split anterior end showing inner characters. The specimen was figured as B. mucronata (Schlotheim) by Fig. 2. Belemnitella mucronata (Schlotheim) from the Köpinge Moberg (1885, PI. 6:14). sandstone at Valleberga. The specimen probably came from the LM LO 776. highest Lower Campanian or basal Upper Campanian. A: Dorsal view. Fig. 4. Belemnitella aff. langei Jeletzky from the Köpinge B: Lateral view. sandstone at Köpinge. Probably middle Upper Campanian. C: VentraJ view. A: VentraJ view. D: View of the split anterior end showing inner characters. B: Lateral view. RM Mo 158067. C: View of the split anterior end showing inner characters, x 2. SGU Type 5219.

54 20

1A 18 1C 2A 28 2C

3A 38 3C 30 4C 4A 48 WALTER KEGEL CHRISTENSEN

PLATE 7

Figs. 1-4. Belemnitella aff. langei Jeletzky from the Köpinge Fig. 5. Belemnitella langei Jeletzky from the quarry sandstone at Köpinge. Probably middle Upper Campanian. Schetschkowy-Gory at Seim , Ssumy Area, USSR. Cast of Fig. l. A: Dorsal view. holotype. Uppermost Campanian, 1-2 m below the 'Lan B: Lateral view. ceolatenschichten'. C: Ventrat view. A: Dorsal view. D: View of the split anterior end showing inner characters, x B: Lateral view. 1.5. C: Ventrat view. SGU Type 5220 Geol. Surv. Canada No. 47/96. Fig. 2. A: Dorsal view. B: Lateral view. Figs. 6-7. Belemnitella langei Schatzky sensu Naidin from the C: Ventrat view. Upper Upper Campanian. R . Don, USSR. Ex D.P. Naidin's D: View of the split anterior end showing inner characters, x 2. Coli. SGU Type 5221. A: Dorsal view. Fig. 3. A: Dorsal view. B: Lateral view. B: Lateral view. C: Ventrat view. SGU Type 5222. D : View of the split anterior end showing inner characters, x 2. Fig. 4. A: Dorsal view. Nate the !arge fissure angle. B: Lateral view. Fig. 6 is MGUH 16894. C: Ventrat view. Fig. 7 is MGUH 16895 . SG U Type 5223.

56 1A 18 1C 10 20 2A 28 2C 3A 38

4A 48 4C

60 70

5A 58 se 6A 68 6C 7A 78 7C

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