Palaeobiogeography and migration in the Late Cretaceous belemnite family Belemnitellidae

WALTER KEGEL CHRISTENSEN

Christensen, W.K. 1991. Palaeobiogeography and migration in the Late Cretaceous belemnite family Belemnitellidae. -Acta Palaeontologica Polonica 42, 4,451495.

The Late Cretaceous belemnite family Belemnitellidae Pavlow, I9I4, which includes nine genera and two subgenera, occurs only in the Northern Hemisphere, that is in the North European and North American palaeobiogeographical Provinces of the North Temperate Realm, in addition to the northern margin of the Tethyan Realm in Europe. The North European Province is subdivided into the Central European, Central Russian and Baltoscandian Subprovinces. The centre of origin and dispersal of the belemnitellids lay in the North European Province and all known genera and subgeneraoccur there. The belemnitellids immigrated intermittently into the Tethyan Realm (at least nine times) and the North Ameńcan Province (at least six times). The majority of the species occurring in the Tethyan Realm are conspecific with those from the North European Province, whereas the species occurring in the North American Province are endemic, with a few exceptions. The endemic species probably evolved by allopatric speciation from initial migrants. More than a score migrations have been recognized within the subprovinces of the North European Province. The palaeogeographical distńbution and migration pat- terns of the belemnitellids were to a certain extent controlled by eustatic sea-level changes, cool or warm climatic phases and competition, although the cause of several migration events cannot be satisfactorily explained at present.

Key words: Belemnitellidae, Late Cretaceous, palaeobiogeography,migration, Northern Hemisphere.

Walt e r Ke g eI C hri s te ns en [wkc @ sav lk. g eomu s. ku. dk ], Ge ol o g i cal M us e um, U niv e r s ity of C op enhagen, Q ster Voldgade 5-7,1DK- I 3 5 0 C openhagen, D enmark.

Introduction

During the Cenomanian, the earliest stage of the Late Cretaceous, belemnites had a tripartite distribution. The family Belemnitellidae Pavlow, 1914 inhabited the North Temperate Realm (Boreal Realm of authors), the family Dimitobelidae Whitehouse, 1924 inhabited the South Temperate Realm (Austral Realm) and the family Belemnop- seidae Naef, 1922 inhabited the Tethvan Realm. The latter familv became extinct in the 458 Late Cretaceous belemnitelllds; CHRISTENSEN

(s a IJJ Numberof genera ui (5 Genera and subgeneraof Belemnitellidae and subgenera o * 123456 U) tttttl I 65.4 .e I ss \ = .s E o ,i ) 71.3- IIE

L ( o- o =€-j=l t-rJ

JEĘ

80.6- \ q;rL -.Ę-Tfi TF LU 83.5- IET- E IF \ z.t T ^i: T s f.ę a t A= 86.3- ttt :> z. ()=OM { _l-i E -Yl- , 88.7- i- x $ ( CE: Ę PM !s ł E o c 93.3- Ę * z.: IIJ M a (J I 98_5 ; f

Fig. 1. Statigraphical range and inferred phylogeny of belemnitellid generaand subgenera(modified from Christensen 1997).Stage abbreviations after Harland et al. (1989):CEN - Cenomanian,TUR - Turonian, CoN-Coniacian,sAN_Santonian,CPM-Campanian,MAA_Maastńchtian,E-early,M-middle, L - late. Ages in Ma after Obradovich (1994). middle Cenomanian, and, consequently,the belemnites had a bipolar distribution during the remaining part of the Late Cretaceous.Thus, a Tethyan Realm cannot be defined on the basis of belemnites alone after the Cenomanian, although it can be rccognizedon the basis of otherfossil groups,including rudists,ammonites, echinoids, actaeonellidgastropods and larger foraminifera. The belemnitellids appearedin the early Cenomanian,some way abovethe base of the substage,and became extinct at theMaastrichtian-Danian boundary. The dimitobe- lids appearedin the Aptian and disappearedin the early Maastrichtian (Doyle & Zinsmeister1988; Doyle & Howlett 1989;Crame et al. 1996).Itseems, however, that the dimitobelids became almost extinct in the middle early Campanian (Crame et al. 1996). The last belemnopseid genera Parahibolites Stolley, 1919 and Neohibolites

Figf . Late Cretaceousbelemnite stratigraphy of northwestEurope, Baltoscandiaand the Russian Platform (modified from Christensen 1997). Abbreviations: A. - Actinocamax, BIc. - Belemnellocatnax,BIn. - Belemnella, Blt. - Belemnitella, Gc. - Goniocamcuc,Gt. - Gonioteuthis,N. - Neohibolites,P. - Praeacti- nocamax,E-early,M-middle,L-late,Lo-lower,U-upper.Verticalaxisnottoscale. ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 459

ZONAL ZONAL BELEMNITES, J IA. BELEMNITEZONES. BELEMNITES, GES RUSSIAN PLATFORM NW EUROPE BALTO.SCANDIA

LlBIn. kazimiroviensis |\--B In. ka zi m irov ie n si s < (E) <-l t_ junior j> El Blt. lunior lBlt. (Vtt\

F Bln. fastigata I OL BIn. cimbrica cc :! B. sumensis - Bln. sumensis q) a- BIr-btr* q) B. lanceolata q) Bln. oseudobtusa m LU B. licharewi Bln. lanceolata Bln. lanceolata

z Blt. minorll B.l. najdini qL B.l. langei f Blt. minor I B.l. minor o- q UJ L Blt. woodi Blt. mucronata

ri I Gt. praewestfalica Gc. lundgreni zM Gc. lundgreni

L zTi rvl o ff

F P. plenus triangulus

P nlanttą P. plenus P. plenus z z { rr > lvl P. primus (NE) '/- z .,. ._ P. primus P. primus /ż LIJ .,/ E N. ultimus N. u/lrmus(SW) 460 Late Cretaceous belemnitelllds: CHRISTENSEN

Stolley, 1911 appearedin the Aptian and became extinct in the early and middle Cenomanian,respectively (Combómorel et al, 1981). The belemnitellidswere stenothermalshelf dwellers,but it seemsthat the breeding, spawning,hatching, and, possibly for the females at least, dying grounds,were inner neritic, shallow water environments(Christensen 1976,199'7). The dimitobelids were also stenothermalshelf dwellers(Doyle & Howlett 1989).Since both the belemnitel- lids and dimitobelids were stenothermalshallow-water dwellers, deep and warm water in the Tethyan Realm may have acted as a physical barrier and precluded spread of thesefamilies. Species of Neohibolites andParahibolites migratedaway from the TethyanRealm into both the North Temperateand SouthTemperate Realms during Aptian to Cenoma- nian time (Stevens1973; Doyle 1988, 1992;Doyle & Howlett 1989).In the Cenoma- nian they are recorded as far north as northwestGermany and southernand eastern England, where they may be locally abundant.The two generahave been recorded from deep marine and continentalslope deposits,as well as inner neritic sediments, and the habitat of these genera may thereforehave been surface oceanic, but they probablyspent their breeding season inshore (Doyle & Howlett 1989).They werethus eurythermal,adapted to life in warm temperateas well as tropical waters. Christensen(1997) discussed the taxonomyand evolutionaryhistory of the Belem- nitellidae and recognized nine genera and two subgenera:Prąeąctinocamax Naidin, 1964,Actinocamax Miller, 1823,Belemnocamax Crick, 1910, GoruioteuthisBayle, 1878,Belemnellocamax Naidin, 1964,Goniocamar Naidin,l9G, Belemnitellad'Or- bigny, 1840,Belemnella (Belemnella)Nowak, 1913,Belemnella (Pachybelemnella) Schulz, 1979,Belemnella (Neobelemnella)Naidin, I975, and Fusiteuthis Kongiel, 1962 (Fig. 1). He provided diagnoses for the genera and subgenera,figured repre- sentative species of the genera, and reviewed previous classifications by Naidin (I964b) and JeletzĘ (unpublishedmanuscript t972 for the Treatiseon Invertebrate Pąlaeontology).Belemnella (Neobelemnella),Belemnocamax, and Fusiteuthis are monotypic,but the latteris most likely anomen dubium(Christensen 1997). The belemnitellids are of fundamentalimportance in biostratigraphyand correlation in Europe, especially during the Coniacian to Maasfrichtianstages, because they were common,widely distributedand their fossilizationpotential is high (Christensen1990b). Twenty six belemnite zones have been establishedin norttrwestEurope and a little less on the Russian Platform (Frg. 2). Some of the northwestEuropean zones have been subdividedrecenfly (Christensen1995, 1996, 1997;Christensen & Schulz 1997).The first occurrence of a zonal index belemnite in a specific area is due either to an evolutionaryevent or a migration event.It has been assumedthat the Late Cretaceous belemnitezones have isochronous boundaries, but Christensen(1996) showed that some of the Maastrichtianzones have slightly or highly diachronousboundaries, reflecting migrations.Nielsen (1995:p. 56),referring to trilobites,noted that: 'It is well-knownthat althoughbiostratigraphy ideally correlatesthe appearanceand extinction of taxa, it very ofteninstead mirrors faunalmigrations relating to palaeoenvironmentalchanges...'. It is, consequently,of greatimportance for biostratigraphyand correlationto unravel the cause of the first occurrencesof index belemnitesin various areas. The aim of this paper is to discuss the migration of the belemnitellids within the palaeobiogeographicalframework of Christensen(1975,1976,1988, 1993b,1997). ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 461

Palaeobiogeographical framework

Kauffman (1973) established a quantitative scheme for defining Cretaceous biogeo- graphical units on the basis of bivalve molluscan genera and subgenera. He recognized the following units, in decreasing order: realms (>75Vo endemic genera), regions (50-:75Voendemic genera), provinces (25-50Vo endemic genera), subprovinces (10- 25Vo endemic genera) and endemic cenffes (S-lOVo endemic genera). Itis obvious that this quantitative approach cannot be applied to animal groups high in the food-chain, as for example belemnites. Howevęr, the major biogeographical units of Kauffman are applicable to the Late Cretaceous belemnites, and Christensen (1975, 1976, 1988, 1990b, I993b, 1997) recognized realms, provinces and subprovinces. The North Temperate Realm comprises the North American and North European Provinces, and the latter includes the Central European, Central Russian and Balto- scandian Subprovinces. Christensen (1997) showed that the centre of evolution and dispersal of the belemnitellids lay in the North European Province for the following reasons: (1) they are common and all known genera and subgenera occur there; (2) the earliest belemniteI|id, Prąeactinocamax primus (Arkhangelsky, IgIf), appeared in the early Cenomanian, some way above the base of the substage, in this province; (3) the earliest species of the younger genera occurred either only there or appeared earlier there than elsewhere (Table 1). He also showed that the genera Praeactinocamax and Belemnitellą are more widely distributed than the remaining genera, because they occur in both the North European and North American Provinces, in addition to the northern margin of the Tethyan Realm in Europe. He ranked the genera in the following way with respect to their palaeogeographical distribution, from the largest to the smallest areal extent (Table 1): (1) Praeactinocamax and Belemnitella: (2) Gonioteu- this, Belemnellocamax andBelemnella; (3) Actinocamax) (4) Goniocamax; and (5) Be- lemnocamax. The Central European and Central Russian Subprovinces of Christensen (1975) are well-detined from the middle Coniacian to the boundary between the early and late Campanian, that is a period around 6-:7 Ma (Mega annuum = 106 y"*r), and are charactęrizedby independently evolving belemnite lineages. The Gonioteuthis stock inhabited the Central European Subprovince and the Goniocamax-Belemnitella stock inhabited the Central Russian Subprovince (Christensen 19'15, 1976, 1988, 1990b, 1993c,1997; Christensen & Schulz 1997). The belemnite faunas of Baltoscandia show affinity to those of the Central Russian Subprovince at certain times, and to those of the Central European Subprovince at other times, as will be shown below. However, in the early Coniacian and latest early Campanian-early late Campanian the belemnite faunas of Baltoscandia are so distinct that it seems appropriate to regard them as belonging to a specific subprovince, which was called the Baltoscandian Subprovince by Christensen (1997). This subprovince is charactęrizedby the species Goniocamax lundgreni (Sto|ley' 1897) in the early Conia- cian and the genus Belemnellocąmąx in the latest early Campanian and early late Campanian. At other times during the Late Cretaceous the subprovinces are less distinct and no subprovinces are recognized in the middle and late late Campanian and Maastrichtian. 462 I.ate Cretaceous belemnitellŻs; CHRISTENSEN

Table 1. The occunence of belemnitellid generaand subgenerain the North Europeanand North American Provinces of the North TemperateRealm and the Tethyan Realm (after Christensen 1997).Fusiteuthis is mostlikely anomendubium.T)tegeneraandsubgeneraarerankedwithrespecttotheirpalaeobiogeographi- cal distribution. Nine genera and two subgeneraoccur in the North European Province, five genera and two subgenerain the Tethyan Realm and essentially two genera in the North American Province. The numbers l-lf referto the frst appeamncesof the generaand subgenerain the various palaeobiogeographicalunits. Detailed data are not available for the subgeneraof Belemnella. Abbreviations: 1 - early Cenomanian,2 - lateCenomanian,3-middleTuronian,4-earlySantonian,5-latestSantonian,6-latestSantonian--earliest Campanian,T-middleConiacian,S-earlyCampanian,9-earlySantonian,lO-lateSantonian,ll-early Turonian, 12 - latestSantonian+arliest Campanian.

North European NońhAmerican Genera and subgenera Tethyan Realm Province Province

Praeactinocamnx +l +2 +'

Belemnitella +4 +5 +6

B elemne lla (B eI e rnn eII a) + +

B elemne lla (P ac hybelemne lla ) + +

B el emnelLa (N eob elemnella ) + +

Gonioteuthis +7 +8

Belemnellocamax +9 +10

Actinocamnx +11 (+)12

Goniocamnx +

Belemnocamax +

Fusiteuthis +

The middle Coniacian-middle early Campanianbelemnite faunas of Baltoscandia are of greatimportance, because they include species of both the Gonioteuthis and Goniocamax-Belemnitella stocks,and thusprovide a basis for correlationbetween the two subprovinces(Frg. 2). England belongs to the Central European Subprovince as defined on belemnites, becausethe genus Gonioteuthis predominatesin the Santonianand early Campanian. However, two loosely defined faunal provinces have been recognized in England (Wright & Wright l95I; Peake & Hancock 1961;Reid 1971, 1976;Wood & Smith 1978; Mortimore 1983; Mortimore & Wood 1986; Whitham 1993). The northern province includes Yorkshire, Lincolnshire and the northern part of Norfolk, and it shows faunal affinity to northernGermany and Poland. The southernprovince com- prises Kent, Surrey,Sussex, Hampshire, Wiltshfue and Dorset in southernEngland and belongs to the Anglo-Paris Basin. The two provinces are separatedby the 'transitional province', which comprises the Chiltern Hills and East Anglia as far north as central Norfolk. The middle CenomanianBelemnocamąx boweri Crick, 1910is recordedonly from the northernprovince (Christensen1993c), and Santonianand early Campanian speciesof Gonioteuthisare more common in the northernprovince thanin the southern province (Mitchell 1994,1995).Gonioteuthis is very common in the lower Campanian ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 463 near Ipswich in the 'transitionalprovince'and also in northernNorfolk in the southern part of the northernprovince. During the Late Cretaceousthe belemnitellids immigrated intermittentlyinto the North American Province and the northern European part of the Tethyan Realm. They migratedalso within the subprovincesof the North EuropeanProvince.

North European Province

This province extendsfrom heland in the west to the Ural Mountains and beyond in the east (Fig. 3). In addition to the belemnitellids,Cenomanian speciesof the belem- nopseid generaNeohibolites andParahibolites arealso known to occur there.

[-l Land [nTm Central Russian : Central European _- Boundarybetween North Furopean L-J areas ll l ,il,'I Subprovince -l Subprovince Provinceand TethvanRealm

Fig. 3. Map showing the Central European and Central Russian Subprovinces of the North European palaeobiogeographicalProvince, as defined by the Gonioteuthis and Goniocamax-Belemnitella stocks, respectively(modified from Christensen 1976).The BaltoscandianSubprovince (indicatedby crosshatch- ing) is distinguishedby Goniocamax lundgreni inthe early Coniacian and the genusBelemrnnellocamax in the latest early and early late Campanian. Late Cretaceous land and sea areas represent maximum inundationfor all stages.The boundariesare not reliable in detail and the biogeographicunits are typically gradationalin character.Abbreviations: A - Azerbaijarrin the Caucasus,B _ the island of Bornholm in thę Baltic Sea, Ba - Bavaria in southern Germany, C - Chartreuse in the Subalpine Chain in France, M-MangyshlakinKazakhstan,N-NorthernCalcareousAlpsinAustria,R-Romania,S-Scaniain southem Sweden. The.Vocontian Basin in southeastFrance and the Corbidres in the easternpart of the French Pyrónóes are situated south of the map.

Cenomanian The Cenomanian belemnite fauna of northwest Europe is diverse and includes four genera and five species: the belemnitellids Praeactinocafturx primus, P. plenus (Blain- ville, 1825_|827) and Belemnocąmax boweri, in addition to the belemnopseids Neo- 'cf. hibolites ultimus (d'Orbigny, 1845) and Parahibolites tourtiae' (Weigner, 1909) (Christensen 1990a, 1993c; Christensęn et al. 1992; Christensen, Juignet et al. 1993) 464 Late Cretaceous belemnitellids: CHRISTENSEN

Belemnopseidae Belemnitellidae Nerlhibolites Praeactinocamax Actino- Gonioteuthis BIc. hib. B camax

(5 a Ą ,t !g o IJJ o s F J t) Ą O) o :ts g e o ?Ą cĄ ui !) ,, (/) c) (/t B 'Ą q o ,t 3 lr) s Ą 1) 5 l) (')! s .h (/, C' fl u, (Ą c = ,,) B $ g Ę @ q) rt - 3 c B Ę e l) I l) I q) Ę q) o (t) q) E tr 3 o /) o h (/)= 3 o $ R Ą I s o o :< s h $ ś h IB stt s - e F .Ą q) D q) t 5 s q) r) o (/) ś rJ = s - q) E Ę D o o V, V) e s h .Ą q) q) $ 3 o h o 0) B ś q) g a) lJ P Ę q) s q) B :IJ a F (Ą o ) XJ s F r t o g x $ Ś f ó E a, I o I o o. qJ o- $ 9 I e o ś 3) ści ś1) (D a t5.4

21.3-

o- o- UJ Ę| 1 J

Fig. 4. Stratigraphical range diagram of Late Cretaceous belemnopseids and belemnitellids from the North European Province (modified from Christensen 1997). Abbreviations: Parahib. - Parahibolites, B. - Belemnocamax, Blc. - Belemnellocamnx, B. (N.) - Belemnella (Neobelemnella), F. - Fusiteuthis,I - northwest Europe, 2 - Russian Platform, 3 - Bornholm, Denmark. Stage abbreviations as in Fig. 1. Ages in Ma after Obradovich (1994).

(Fig. 4). During the early and middle Cenomanianthe belemnopseidswere distributed in the Central European Province, whereasP primus occuned mainly in the Central Russian Subprovinceand on Bornholm, Denmark.It theręforeseems that Baltoscandia was part of the Central Russian Subprovince in the middle Cenomanian. It can be mentionedin passingthat Packer et ąl. (1989)recorded east European foraminifera from the early middle CenomanianArnager GreensandFormation of Bornholm. The genusPąrahibolites occurred mainly in the Balkan{rimęan_Caucasian area (Ali-Zade 1972),and Stevens(1973) noted that it was not recordednorth of the line delimiting the southernextent of Praeactinocąmąx.However, Stolley (1920)recorded the extręmelyrare spęcies ,cf. Pąrahibolites tourtiae,from the earliestCenomanian of the Mtinsterland and Subhercynian CretaceousBasins in the Central European Sub- province.Neohibolites ultimus is mainly of early early Cenomanianage but rangedinto the middle Cenomanian (Christensen et al. 1992). The highest record is from the Acanthoceras rhotomagenseZone, Turrilites acutus Subzone of northern France (Christensen,Juignet et ąl. L993). Christensen (1990a) showed that Praeactinocamaxprimus appearedin the early Cenomanian,some wav above the base.and continuedinto the early middle Cenoma- ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 465

Belemnitellidae Belemnella Belennella B, fuimnat Belemnitella (Belemnella)(PadryMunndla)N.F. c o Ę (s o u, N o @ c c f o I,JJ N o B o o o 5 ah o o \ Y c N u) E ui F !u D |Ą N o (J E !s tr t, o s rs 't) .c r ro fl o r E N !! i/) a Ę o o B a = h (Ą x ii F ) I t 5 o N 0) F o o 0 t\ q) E E E q) n r = F B B Ę o o \ (! o N N o 0) o o o o c o o B Ś q) F o N o o F 5t t9 o E o b B o o ts F o F (Ą - F ą ś |Ę Ę Ę Ę r (/) 0) o 5 EN * F x $ 5 N N N s (Ąa o F $ ) o a E Ą a a. N s t) l=.s c a o s s E () o N a s o o t, B a 65-4 I t. I I I ! 71.9 I I I I >L I (L I o I I T I UJ Ę, I ona

Fig.4. (continueĄ. nian in the Central Russian Subprovince.It has beenrecorded as Belemnitesląnceola- tus fromBed A of thecondensed Cenomanian limestone succession of Devon, southern England, whichis of early Cenomanianage (see Christensen 1990a). According to C.J. Wood (personal communication August 1997) this belemnite (BGS GSM 101028, housed in the Geological Survey Museum, British Geological Survey, Keyworth) is misidentified and belongs to Neohibolites ultimus. It has been shown recently that although P primus occurs rarely in the Central European Subprovincęit has an acme in this subprovincein the middle Cenomanian, at the base of the Turrilites costatusSubzone of theAcanthoceras rhotomngenseZone. P. primus occurs at the base of the marly bed of couplet C1 (see below) or its correlativesin the Anglo-Paris Basin in southernEngland, where it is relatively rare, and in the Cleveland Basin in easternEngland (Paul et al. 1994;Gale 1995;Mitchell et ąl. 1996)' in addition to the Lower Saxony and Mtinsterland Basins in northwest Germany (Christensen1990a; Christensen et al. 1992).Moreover, it has beenrecorded also from the early middle Cenomanian glauconitic sands in Northern Ireland (see Christensen1990a). Gale (1990, 1995) provided a composite cyclostratigraphyfor the Cenomanian Stageof Europe, which is graduatedinto ZLZprecessioncycles (modeat 21,000years). He numberedthe couplets, consisting of a marl bed below and a chalk bed above, 466 Inte Cretaceous belemnitelllds; CHRISTENSEN

A 1-5I,BI-45, Cl-46,D1-49 andBl-L7 andprovided an integratedstratigraphy for the Cenomanian,comprising ammonite,inoceramid and planktic foraminiferanbiostrati- graphy,sequence stratigraphy and cyclostratigraphy.Couplet Cl was depositedduring the beginning of a transgression(TST = transgressivesystems tract) following an eustatic sea-level lowstand (LST), some distance above a sequenceboundary (Gale I990;Pal| et aI. 1994;Mitchell et ąl. 1996).Praeactinocamax primus is accompanied by apulse faunaof smallbivalves, which consistsof speciesthat are identical or closely similar to speciesof Jefferies' (196f) North Boreal faunal group from beds 4{ of the late Cenomanian Plenus Marls of the Anglo-Paris Basin. He interpretedthese as migrantsfrom the north and east,indicative of a cool climatic phase,which was called the Plenus Cold Event by Gale & Christensen(1996). The cool climatic phase in the early middle Cenomanianis here called the Primus Cold Evęnt. The duration of this eventwas very short and less thanfl ky. Thus, P. primus abruptlyinvaded the Central European Subprovince coincidently with a rapid rise of sea-leveland a cool climatic phasein the early middle Cenomanian. Christensen(1990a) described a little less than 100specimens of Praeactinocamnx primus from theprimus bed of the Wunstorf pit nearHannover in northwestGermany, which equateswith couplet Cl of the Anglo-Paris Basin (Gale 1995;Mitchell et al. 1996). This large number of specimens may falsely indicate that P primus occurs commonly there comparedwith its occunence in C1 couplets elsewhere.This is not the case, however,but rdlatesto the facts that many private collectors have collected therefor sęveralyeals and the comparativelygood exposureof the bed in question.The sampleof P primlzs from Wunstorf containsall growthstages. Christens en et ąl. (L992) recorded a juvenile specimen from the Mi.insterlandBasin and I have seen another juvenile specimen from couplet Cl at Abbot's Cliff betweenDover and Folkestone, southernEngland. It thereforeseems that P primus bred and spawnedeverywhere in the Central European Subprovince. The stratigraphicalranges of Neohibolites ultimus and Praeactinocam.axprimus overlap,but, accordingto my knowledge,they have neverbeen recorded from the same bed in the Central European Subprovince.The two species occur in the early middle Cenomanianof southernEngland, but P primus occurs at a slightly higher stratigraphi- cal level, i.e. coupletC1, thanN. ultimus,which is from known only one specimenfrom the earliestmiddle Cenomanian(Christensen et al. 1992). P raeac tinocamax primus is followed upwards by P raeactino c amax p lenus and the two species form an evolutionary lineage (Christensen 1974, 1990a). P. plenus is recorded from the middle Cenomanian to the earliest early Turonian on the Russian Platform (Naidin 1981). It occurs mainly in Beds 4-6 of the Plenus Marls of the Anglo-Paris Basin (Jefferies 1962,1963)but a single specimenhas also beenfound in Bed 8 (Gale & Christensen 1996).The Plenus Marls are of middle late Cenomanian age,Metoicocerąs geslinianumZone. P plenusoccurs also in thegeslirtianumZone of northwestGermany (Christensen 1990a). Wood & Mortimore(1995) and Gale (1995) noted that it occurs in the top unit of the trĘartite Plenus Bed in the Lower Saxony Basin, where it is an extreme rarity. This unit is infened to correlatewith Jefferies' Bed 4 of the Plenus Marls. P. plenus also occurs in the Plenus Bed in the Mtinsterland Basin. Marcinowski (1972)recorded P. primus andP.plenus from thęlate Cenomanian conglomeratesat Glanów in thę Polish Jura Chain. Christensen (1990a) suggested, ACTA PALAEONTOLOGTCA POLOMCA (42) (4) 467 however, that the specimen referred to as P. primus by Marcinowski was either reworked or, more likely, a slenderspecimen of P. plenus. Prąeactinocamax plenus has thus a very limited stratigraphical distribution in northwestEurope, becausethe duration of theM. geslinianumZone was estimatedto be about 110 ky (Gale 1990).Beds 4-6 of the Plenus Marls or its correlativeswere deposited during a significant fall of sea temperature,the Plenus Cold Event (see above),coincidently with the beginningof a transgression(TST), following an eustatic sea-level lowstand, slightly above a major sequence boundary (Gale 1990, 1995; Mitchell et al. 1996). Thus, P. plenus invaded northwestEurope in the middle late Cenomanianduring a cool climatic phaseand arapidrise of sea-level.In addition,lack of competition of the Tethyan belemnopseids,which became extinct in the middle Cenomanian may also have been of significance for this southwardsmigration. P plenus even locally invaded the northern margin of the TeĘan Rea]m (see later discussion). The very small and anomalous-lookingBelemnocamax boweri has a very limited stratigraphicaland geographicaldistribution. It occurs in the early middle Cenoma- nian, basal part of the Turrilites costarzs Subzone,of the Lower Saxony and Miinster- land Basins in northwestGermany, in additionto the Cleveland Basin and at Hunstan- ton, Norfolk, in eastern England (Christensen 1993c). B. boweri co-occurs with Prąeactinocamaxprimus in the early middle Cenomanian.The highest record of B. boweri is from the base of the Nettleton Stone at South Ferriby in east England (Christensen et aI. 1992), which is of late middle Cenomanian age,Acanthocerąs jukesbrowneiZone (Mitchell et al. 1996).

Discussion. - Belemnocamaxboweri andthelast speciesof Neohibolites became extinct in the middle Cenomanian,and Praeactinocąnnx plenus of theprimus_plenus lineage disappearedin the late Cenomanian in northwestEurope and in the earliest early Turonian on the Russian Platform (Fig. 4). This belemnite extinction, which lasted aboutf .5 Ma, may be part of the well-known, global middle Cenomanian+arly Turonian mass extinction,which was discussedat length by Kauffman & Hart (1996) and summarizedby Barnes et al. (1996).It is associatedwith a near-peakMesozoic eustatic sea-level highstand, a global warming peak, greenhouseclimate reflecting elevatedCOz and Oceanic Anoxic Event II. It is, however,difficult to imagine exactly which of thesefactors were responsiblefor the world-wide extinction of the euryther- mal, surface oceanic belemnopseidsand the extinction of some stenothermal,shelf dwelling belemnitellids in Europe.

Turonian-early Coniacian Central European Subprovince. - After the belemniteextinction around the Cenomanian-Turonian boundary,belemnitellids are very rare or absentin the Turo- nian and Coniacian of the Central European Subprovince. They occur very rarely in England, Germany, the Czech Republic and southern Sweden and are not recorded from Northern keland, France, Poland and the Crimea. Neither are belemnites reported from the early Santonian of the Crimea (Naidin 1973) and Poland (Cieś- liński 1963). 468 Late Cretaceous belemnitellids : CHRISTENSEN

The small faunafrom the Central EuropeanSubprovince and Baltoscandia,consist- ing of a little more than a dozen specimens,was revised by Christensen(1982), who recognized the following species: Praeactinocamax bohemicus (Stolley, 1916), P paderbornensis (Schliiter, 1894), P. strehlelzsls(Fritsch & SchlÓnbach, 187f), and Goniocąmaxesseniełzsis (Christensen, 1982) (Fig. a). The lectotypeand paralectotype of the late Turonian P strehlensi,scame from the Pliiner limestone of Strehlen near Dresden.Christensen recorded five specimensof P bohemicus:(1) theholotype which camefrom the Plżinerlimestone near Kostic in the CzechRepublic; it is of late Turonian age;(2) two specimensfrom the Plżinerlimestone at Strehlen;these are of late Turonian age;(3) one specimenfrom the middle Coniacian of Yorkshire in England; and (4) one specimen of supposedly late Trnonian age from Sfirdal in southern Sweden. Three granulatedbelemnite apical fragmentsof supposedlylate Turonian age, from SŻirdal, were tentatively assigned to P bohemicus by Christensen (1982). T\e holotype, by monotypy,of P.paderbornensis,from the MtinsterlandBasin,is of late early Coniacian age (Kaplan & Kennedy 1994).The holotype, by monotypy,of G. esseniensis,from the Mtinsterland Basin, is of early, but not earliest, early Coniacian age (Kaplan & Kennedy 1994). Kośtak(1996) described six specimens,including theholotype, of the late Turonian Praeactinocamaxbohemicu,s, one specimenof the late TuronianP. atr.bohemicus and two specimensof Goniocamax lundgreni of late early Coniacian age from the Bohe- mian Basin in the CzechRepublic. The three species of Praeactinocąmax,P, bohemicus,P paderbornensis and P strehlensis, differ from contemporaneousspecies of this genus from the Central Russian Subprovince and the North American Province (seebelow) and their origin is unknown (Christensen1982).In contrast, the early Coniacian Goniocamax esseniensis is very closely allied to and may be conspecific with G. lundgrerel(Christensen & Schulz 1997). The holotype of G. esseniensisis adult and may be regarded as an occasionalmigrant from the BaltoscandianSubprovince, as may the two specimensof G. lundgreni from the Bohemian Basin. Central Russian Subprovince. - During Turonian and early Coniacian time belemnitesare more common in the Central Russian Subprovincethan in the Central European Subprovince, but they have a very limited area of distribution, occurring almost ęxclusively in the Volga district (Naidin 1981). The following species are recorded:Praeactinocaftu)x intermedir,rs (Arkhangelsky, Igtf), P coronarrzs(Makhlin, |965)' P planus (Makhlin, 1965),P. medwedicicus(Naidin, 1964) and P matesovąe (Naidin, L9G), in addition to various subspeciesof Actinocamoc verus Miller, 1823 (Naidin 1964b;Makhlin 1965) (Fig. a). However, only P intermediusoccurs com- monly there.P. plenus triangulus Naidin, 1964is used as an index speciesfor the early early Turonian of the Central Russian Subprovince (Naidin 1981) (Fig. 2), but Chris- tensen(1974) placed this subspeciesin synonymy with the nominoĘpical subspecies. Baltoscandia. - As shown above,Turonian belemnites are very rarein Baltoscan- dia, but it should be borne in mind that Turonian rocks have been recordedonly from one outcrop in this area,the temporat/ exposureat Sżirdalon the Swedish west coast (Bergstróm et al. 1973).Since Praeąctinocamax bohemicu.toccurs at this locality it seemsthat Baltoscandia was partof the CentralEuropean Subprovince in the Turonian. ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 469

Goniocamnx lundgreni appearedat the base of the Coniacian of Bornholm and persistedinto the early Santonian(Christensen & Schulz 1997)(Figs 4,5). The early Coniacian of Baltoscandiais characterizedby G. lundgreni,although it is rare. Chris- tensen& Schulz (1997)recorded only about60 specimensfrom the early Coniacian of Bornholm and thesewere collected over a period of about 100 years. Actinocąmąx verus antefragills Naidin, t964 is early Turonian in age on the Russian Platform. Christensen & Schulz (1997) recorded two specimensof A. verus cf . antefragilis fromthe late early and middle Coniacian of Bomholm (Fig. a). These were considered as stray specimens from an unknown area, where the subspecies survived from the early Turonian to the early early Coniacian. One may get the impression that the belemnite species diversity is high in the Turonianand early Coniacian (Fig. a). Christensen(1982: p.77) noted,however, that some of the species of Praeactinocamałcare distinguished on the basis of minor differences and may eventually prove to be in part conspecific. More material is necessaryin order to solve this problem.

Middle Coniacian-early Campanian During this peńod, which lastedabout 6_-7 Ma, the Central EuropeanSubprovince was characterizedbythe Gonioteuthis lineage, whilstthe Central Russian Subprovincewas characterized by the Goniocamax-Belemnitella lineage. The middle Coniacian- -middle early Campanianbelemnitefaunas of Baltoscandiainclude speciesof boththe Gonioteuthis and Goniocamnx-Belemnitella stocks. Belemnellocamax ex gr. gros- souvrei (Janet, 1891) and Actinocamax verus verus are widely distributed in the Santonianand early Campanianof the North EuropeanProvince. The first is very rare, and the latteris common. Central Buropean Subprovince. - The Gonioteuthis lineage includes seven speciesand subspecies,in ascendingorder: G. praewestfalicaErnst & Schulz, !974, G. westfalica (ScŁiliter, 1874), G. westfalicagranulata(Stolley, 1897), G. granulata (Blainville, 1827), G. granulataquadrara (Stolley, 1897), G. quadrata quadrata (Blainville,1827), and G. quadrata gracilis (Stolley, 1892)(Fig. a). The ęarliestmember of this lineageis very rare.Gonioteuthis praewestfalica occlxs very rarely in the late middle and late Coniacian of Liigerdorf nearHamburg, northwest Germany @rnst & Schulz 1974). A single specimen was recorded from the late Coniacian of Kent, southernEngland by Christensen(1991) and anotherspecimen of supposedly late Coniacian age was reported from Helgoland by Wood & Schmid (1991).Kaplan & Kennedy G99Ą recordedtwo specimensof lateConiacian age from Augustdorf and Paderborn-Elsen,respectively, in the Mtinsterland Basin. A single specimen has been collected from the late Coniacian chalk of northern France (C.J. Wood personalcommunication August 1997).The specimensof G. praewestfa- lica fuomKent and northernFrance may be regardedas a stray specimens. The early and middle Santonianbelemnite fauna of southernEngland is diverseand includes speciesfrom both the Central EuropeanSubprovince, Gonioteuthis westfalica andG. westfalicagranulata,ffid the Central Russian Subprovince,Goniocamax,Iund- greni and Belemnitella propinqua (Moberg, 1885), in addition to the widespread Belemnellocamąxexgr.7rossouvreiandActinocąmaxverusverus (Christensen 1991). 410 Late Cretaceous belemnitellids: CHRISTENSEN

G. lundgreni, B. propinqua, and B. ex gr. grossouvrei are extremely rare, as is G. Iundgreniin the early Santonianof the MtinsterlandBasin (Ernst t964a; Christensen 1973, 1991).The specimensof G. lundgreniand B. propinqua from northwestGer- many and southernEngland may be regardedas occasional migrantsfrom the north. The late Santonianbelemnite faunas of the Central European Subprovinceinclude Gonioteuthis granulata, in addition to the widespreadBelemnellocamax ex gr. gros- souvrei and Actinocamax verus.In contrast,the eady and middle early Campanian faunas comprise species of Gonioteuthis with subordinatespecies of Belemnitella, A. verusand B. ex gr.grossouvrei. Mixed Gonioteuthis/Belemnitellaassemblages occur at three distinct horizons in the early Campanianof the Central EuropeanSubprovince and Baltoscandia,that is in earliest, middle and latest early Campanian time. In Scania, northern Germany and northeast Belgium, the two genera co-occur in the earliest early Campanian: G. granulataquadrata and B. alpha Naidin, L964 in Scania (Christensen 1975, 1986, I99L); G. granulataquadrataand B. praecursor Stolley, 1897at Braunschweig (Ernst I964b, 1968; Christensen 199 1 ); G. q. quadrata,B. alpha,,B. aff. mucronata"/prąecur- sor' or'8. aff. senior/praecursor'inthe Bottrop Marl in thesouthwestern corner of the Miinsterland Basin (Ernst I964b; Christensen 1986, t99l); G. granulataquadrata, G. q. quadrata and B. praecursor in the Stromberg beds in the easternpart of the Miinsterland Basin (Kaplan et aI. 1996); and G. q. quadrata and B. praecursor at Hallembayein northeastBelgium (Christensen& Schmid 1987;Christensen 1991). The next mixed assemblagesabove consist of Gonioteuthis q. quadrata and Belemnitella praecursor and occur in Northern Ireland (Fletcher & Wood t978), Norfolk and Suffolk in easternEngland, as węll as in southernEngland (Christensen 199I) and northernFrance (Jarvis 1980).These are middle early Campaniapin age, that is the traditional Hagenowią blackmorei Horizon of England, which equates with the German offaster pilula/Galeolą senoruensisZone and the lower part of the G. senonensisZone (Christensen1991). A single specimen of B. praecursor was recorded from the middle early Campanian G. senonensisZone of Liigerdorf by Christensen& Schmid (1987). The highestmixed assemblages,consisting of Gonioteuthisquadrata/Belemnitella mucronata are latest early Campanian in age. At Lźigerdorfnear Hamburg and Mis- burg-HÓver near Hannover in northwestGermany G. quadrata gracilis and B. mucro- nata (Schlotheim,1813) occur in the so-calledOverlap Zone of Schmid (1953),which equateswith the gracilis-mucronata Zone of Ernst (1963a, 1963b).The assemblage comprises G. quadrata scaniensls Christensen, 1975 and B. mucronata in Scania (Christensen1975, 1986). Mixed assemblagesof this age occur also in England, Poland and the SubhercynianCretaceous Basin. Thus, it can be concluded that Belemnitella alpha invaded Baltoscandia and the Central European Subprovincein the earliestearly Campanian.B. praecursor entered the Central European Subprovince twice in the early Campanian,in the earliest and middle early Campanian.B. mucronataimmtgratedinto Baltoscandianand the Central European Subprovincesin the latest early Campanian. The populations of Belemnitellą alpha from Scania (Christensen1986) and of B. praecursor from the early early CampanianInocerąmus lingua/Gonioteuthis quadrata Zone of northeast Belgium (Christensen & Schmid 1987) and the middle early ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 471

Campanian of East Harnham in southern England (Christensen 1991) include all growth stages,indicating that thesespecies bred there.On the contrary,populations of B. praecursor from Stiffkey in Norfolk and from Porchesterand Shawford in southern England (Christensen1991), as well as from localities in northernFrance (Christensen unpublished),consist only of adults,which may be consideredas stray specimens.The populationsof B. mucronatafrom the latestearly Campanianof Scania and northwest Germany comprise all growth stages(Christensen 1975, 1986, 1995),implying that it bred there. Belemnitella praecursor is more common at Hallembaye than elsewhere in the Central European Subprovince. Christensen & Schmid (1987) estimatedthat it ac- countsfor about4O%o of thebelemnitefauna there. In contrast,B. praecursor comprises only 34%o of the belemnitefauna from the middle early Campanianphosphatic chalk of Hardivillers and Ribemont in northernFrance (Jarvis 1980).Jarvis (1980)noted that Actinocamax verus does not occur in the phosphatic chalk and suggestedthat it was oustedby Gonioteuthis.However, this suggestionis incorrect, becauseActinocamax becameextinct slightly earlier. Central Russian Subprovince. - This subprovince is charactertzedby the Go- niocamnx-Belemnitellalineage, which includes, in ascendingorder: G. Iundgreni,B. schmidi Christensen & Schulz, 1997,B. propinqua, B. prcłecursor,B. alpha, and B. mucronata(Fig. a). As mentioned above, G. Iundgrent appearedat the base of the Coniacian in the BaltoscandianSubprovince. It enteredthe Central Russian Subprovincein the middle Coniacian (Naidin 1964b). Gonioteuthis has not been recorded in this subprovince east of the Ukrainian Syneclise, with the exception of the doubtful reports of G. praewestfalicą and G. ?praewestfalicafrom the Coniacian of Mangyshlak, Kazakhstan (Marcinowski et al. 1996). I suspect, however, that the specimens of these taxa should be placed in Praeąctinocamax arąlensis (Arkhangelsky, 19L2) and/or P, mujnakensis (Naidin, 1964),twospecies occurring in the middle and late Coniacian of the Muinak Peninsula on the Aral Sea (Naidin I964b). These species were considered as subspeciesof Gonioteuthis(Goniocamax) westfalica by Naidin (1964b)and Makhlin (1965),but theywere placed inPrąeactinocamaxby Christensen (1997), who noted,however, that they may be transitionalforms betweenPraeactinocamcu and Goniocąnnułx. Naidin (I964a) recordedBelemnitella praecursor from the late Santonian of the Central Russian Subprovince and it thus appearedearlier there than in the Central European Subprovince. As discussed later, it enteredthe Tethyan Realm in the late Santonian. Baltoscandia. - The middle and late Coniacian belemnite fauna of Bornholm includes Goniocamnx lundgreni with subordinate G. birkelundae Christensen & Schulz, t997 (only known from Bornholm) andActinocaftKtxveruscf . antefragllis (see above) (Fig. 5). In the late Coniacian G. lundgreni accountsfor about 957o of the belemnitefauna. G. birkelundaeis closely allied to G. lundgreni and probably evolved from that speciesby allopatric speciationelsewhere; it immigratedlater to Bornholm. The early Santonianbelemnite fauna of Bornholm is extraordinarily diverse and Christensen & Schulz (1997) recorded four genera and nine species:Actinocamax 472 Late C retaceous b elemnitellzds: CHRISTENSEN

Ernst& Schulz(1974) Schulzet al. (1984) Christensen & Schulz a Schulz(1996) (1997) (ĄLIJ F. Lógerdorf, NW Germany Bornholm, Denmark U)

Faunal zones Gonioteuthis Zones Belemnitezones Belemnite assemblage zones z rogatae/ westfalicagranulata westfalic (Rl:11.0-12.5) z westfalica rogalae/westfalica n )t >ńE (Rl: 8.5-11.0) exposed

z Gt. westfalica/A. v. verus/Blt. westfalica coranguinum/ Gonioteuthis oropinoua/Gc.striatus/Gt. ernsti Z1 (Rt< westfalica 8.5) westfalica F z (Rr < 8.s) Btt. striatus a- Gt. westta Iica/G c. Iu ndgren i/ Blt. propinqua cc pachti/ Gonioteuthis Gc. lundgreni/Gt.praewestfalica/ qF undulatoplicatus Gonioteuthis Blt. oropinąua uJ- exlremelvrare praewestfalica Gc. lundgreni/Gt.praewestfz Gc. birkelundae/Blt.schm idi

bucailli/ Goniocamax lundgreni/ praewestfalica praewestfalica Gon iocam ax bi rkel undae z involutus/bucailli Goniocamax Ó tvt no belemniles lundgreni Goniocamax lundgreni/ z koeneni Actinocamax verus ct. o- antefragilis

tr not e poseo Goniocamax lundgreni

Fig 5. Stratigraphicaldiagram, showing Coniaciarr--earlySantonian belemńte zones and assemblagezones of Bornholm, in addition to middle Coniacian-middle Santonianfaunal zones and GonioteuthisZones of Lźigerdorf(modified from Christensen& Schulz 1997)' Abbreviations:A. _ Actinocatlnx, Blt, - Belemnitel- la,Gc.-Goniocamnx,Gt-Gonioteuthis,E-early,M-middle,L-late.RlistheRiedellndexofErnst & Schulz (l97 4) indicating the depthof the pseudoalveolusas a percęntageof the length of the guard.Full namesof the non belermite index-speciesare as follows: Micraster rogalae,M. coranguinum,M. bucailli, Sphenoceramuspachti, Cladoceramus undulatoplicatus,Volviceramusinvolutus,V lcoeneni.Vertical axis not to scale. verus verus, Gontoteuthispraewestfalica, G. westfalicą, G. ernsti Christensen & Schulz, 1997,Goniocamnx lundgreni lundgreni, G. birkelundae,G. striatus Christen- sen & Schulz, 1997, Belemnitella schmidi and B. propinqua (Fig. 5). The genus Belemnellocamax,which occurs elsewherein the early Santonian,is notrecordedfrom Bornholm. It is in fact the only genus occuffing in the early Santonianthat does not occur on Bornholm. Gonioteuthispraewestfalica is of early early Santonianage on Bornholm (Fig. 5), whereasit is of late middle and late Coniacian age in northwestGermany and of late Coniacian age in England and France (seeabove). It thereforespread northwards in the earliest Santonian. G. westfalica is, by and large, of late early Santonian age on Bornholm (Fig. 5). In the early early Santonian the genus Gonioteuthis is fairly common on Bornholm and extremęlyrare in northwestGermany. All growth stagesof G. praewestfalicaand G. westfalicaoccur on Bornholm, indicating thatthese taxa bred there.In contrast,the specimensof G. praewestfąIicąand G. westfalica fromthe chalk of Lżigerdorfare adult. ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 473

Gonioteuthisernsti occurs very rarely in the latestearly Santonianof Bornholm and the Miinsterland Basin in northwestGermany. It is a close ally of G. westfalica and may have evolved by allopatric speciationfrom this specieselsewhere (Christensen & Schulz 1997).It later immigrated to Bornholm and the Mi.insterlandBasin. The late early Santonian Goniocamax striatus is recorded only from Bomholm. It is closely allied to G. birkelundae and is considered as the lineal descendentof this species (Christensen& Schulz 1997).Theearlięst early SantonianBelemnitellą schmidi occurs very rarely on Bornholm and in the Central Russian Subprovince.This species may have evolved from Goniocamaxlundgreniby allopatricspeciation in an unknown arca; it later enteredBornholm and the Central Russian Subprovince(Christensen & Schulz 1997).B. propinqua is most likely the lineal descendentof B. schmidi. Actinocamax verus verus appea$ in the early Coniacian of the Central Russian Subprovince (Naidin 1964b)(Fig. a). on Bornholm it ęnterstogether wtth Gonioca- max striatus in the latest early Santonian,above the extinction level of Goniocamax lundgreni and slightly later than Gonioteuthis westfalica (Fig. 5). A. verus verus appearsalso in the early Santonian of the Central European Subprovince. Thus, A. verus verus immigratedfrom the Central Russian Subprovinceinto the Central Euro- pean Subprovince and Baltoscandiain early,but not earliest,Santonian time. There is a major faunal turnoverin the belemniteassemblages of Bornholm in the middle early Santonian(Christensen & Schulz 1997)(Fig. 5). Belemnite assemblages of middle Coniacian to early early Santonian agę are characterizedby Goniocamąx Iundgreni,which accountsfor about85-95%o of thebelemnite faunas, with subordinate Actinocamax verus cf. antefragilis, Gonioteuthispraewestfalica, Goniocąmax birke- lundae, Belemnitella schmidi and B. propinqua. Assemblagesof late early Santonian age arc characterizedby Gonioteuthis westfalica, which comprises 90-95Vo of the belemnite faunas, with subordinateActinocamax verlłs verus, Gonioteuthis ernsti, Goniocamax lundgreni, G. striatus and Belemnitella propinqua. Bornholm was thus part of the Central Russian Subprovince from middle Coniacian to early early Santo- nian time and becamepart of the Central EuropeanSubprovince in late early Santonian time. Baltoscandia remained part of the Central European Subprovince until latest early Campanian time, becauseGonioteuthis predominates(Christensen 1973, 1975, 1986,1993a). The latestearly Campanianand early late Campanianbelemnite fauna of Scania are characterizedby the genusBelemnellocam.ax: B. mammillatus (Nilsson, 1826)in the latest early Campanian and B. balsvikerzsis(Brotzen, 1960) in the early late Campa- nian. Both species are extremely common in Scania and all growth stages occur, implying that the species bred there. B. mammillatus co-occurs with Belemnitella mucronata, Gonioteuthis quadrata scaniensis and Belemnellocamax ex gr. gros- souvrei, and it comprises about 90-957o of the belemnite faunas (Christensen 1975: table 3). Belemnellocam,axbalsvikensrs co-occurs with Belemnitellamucronata and the former comprisesabout 95Voof the belemnitefaunas (Christensen1976). Belemnellocąmąxmammillątus and B. balsvikensis are extremely rare outside Scania. A little more than 100 specimens of B. mnmmillatus are recorded from the Central Russian Subprovince,in addition to northwestGermany and northernPoland in the Central EuropeanSubprovince (Christensen 1975, 1986;Olszewska 1990).B. bąIsvikensishas not been recorded outside Scania, except for two specimens from 474 Late Cretac eous belemnitelhds; CHRISTENSEN

Misburg near Hannover in northwestGermany (Christensen& Schulz 1976).To my knowledge, the specimensof B. mammillalas occurring outside Scania are adult and they may be regardedas stray individuals buried outsidetheir normal habitat,indicat- ing belemnitemigrations to the south and east.The two specimensof B. balsvikensis from northwestGermany include a juvenile and an adolescentindividual. The presence of the juvenile specimen at Misburg is an enigma, since it is very unlikely that the distancefrom Scania to Misburg (about500 km) couldbe coveredby active swimming. Therefore, Christensen & Schulz (1976) suggestedthat it may have drifted to the Misburg area during heavy storms,either as a gas-filled carcass or clinging to drift- wood or sea-weed.

Late Campanian This substageis characterizedbyBelemnitella,because three genera became extinct at or near the boundarybetween the early and late Campanian.Gonioteuthis disappeared at the boundary,Actinocamax slightly below, and Belemnellocamax slightly above (Fig. 1). The duration of this major belemnite extinction is estimatedto be around 2.5 Ma. Kauffman & Hart (1996) did not record a regional or global faunal extinction event or turnover aroundthe early and late Campanianboundary. However, there are significant microfaunal changesin the European chalks during this interval (Bń|ey et aI. 1983).For the time being, this belemniteextinction is enigmatic. As mentioned above the early late Campanian Belemnellocamaxbalsvikensis is virtually restrictedto the BaltoscandianSubprovince. Thus, only one species,Belem- nitella mucronata,occurred in the earliestlate Campanianof the remaining part of the North EuropeanProvince (Fig. a). The speciesdiversity increasedupwards and several large and small speciesof Belemnitella occurredin the middle and late late Campanian €ig. a). The belemnite fauna of the late late Campanian Beeston Chalk of Norfolk is particularly diverse and includes five taxa of Belemnitella: the very large speciesB. minor I JeletzĘ I95|, the large speciesB. pauli Christensen,t995, in addition to the small speciesB. langei Jeletzky,1948,,B. najdini Kongiel, 1962,and B. sp. 1 (Chris- tensen1995) (Fig. 6).B. minor I predominates,while the othersare rare. For instance, B. minor I constitutesabout 60Vo of thefauna in the Caistor St. Edmundspit (Christen- sen 1995:p. 15).B. najdini appearedsligthly laterthan B. langei in theBeeston Chalk. In the latest Campanian ParamoudrazChalk the very large speciesB. minor II Chris- tensen, 1995 predominatesand the small taxon B. ex gr. langeilnajdini is rare. The ParamoudrarChalk, which is sandwichedbetween the Beeston Chalk and Paramou- drazChalk hasyielded B. minorll andrare B. pauli. Thus,small speciesof Belemnitella enteredNorfolk three times during the late late Campanian. After the major belemnite extinction event around the early and late Campanian boundary,the genusBelemnitella expandedits area of distributionin late Campanian time and invaded the entire North European Province, where it occurs commonly. It even enteredthe TethyanRealm (see later discussion).This extinction event differs, therefore,from that at the Cenomanian-Turonianboundary, when the belemnitellids retreatedinto refugia (see above). ACTA PALAEONTOLOGTCA POLOMCA (4f) (4) 415

Nońo|k ńE Belemitezones, óń NW Europe Belemnitella Belemnitella zones subzones fastigata -F not exposed 9L cimbica cr s F o sumensis CD s obtusa minorll E pseudobtusa (= minorlll ol Christensen 1995) ui lanceolata minorll / exgr.langeil najdini langei minorll z minorll z s L .€ minorl / langei / najdini / ś pauli minor minorl minorl I langei eo o c0 minorl LU F woodi J E mucronata mucronata not exposed

Fig. 6. Stratigraphical conelation diagram, showing conventional late Campanian Belemnitella zones (JeletzĘ 1951)' early MaastrichtianBelemnella zones (Schulz 1979)and inforrnal Belełnnitellazones and subzonesof Norfolk (Christensen1995) (modified from Christensen 1995,1996). The Beeston Chalk is placed in the zone of B. minor I, the ParamoudrarChalk in the subzoneof B. minorll, and the Paramoudraz Chalk in the subzoneof B. minor II and B. ex gr.langei/najdinl. Abbreviations: E - early, L- late.Vertical axis not to scale.

Maastrichtian The genus Belemnella appearedsuddenly at the base of this stage and continued to its end. It is widely distributedin the North European Province and has also been recorded from the northernpart of the Tethyan Realm (see later discussion). Belemnell.a almost oustedBelemnitella and,consequenfly, the early Maastrichtianbelemnite faunas of the North EuropeanProvince consist of speciesof Belemnella with subordinatespecies of Belemnitella.Therarity of Belemnitellnin the early Maastrichtianmay be measuredby the following observations.Belemruellą lnnceolnta (Schlotheim' 1813) co-occurswith Belemnitelląminor sensuChristensen (I97 5) andB. mucrotufiain theearliest Maastrich- tian of the abandonedBalsvik pit in Scania,southern Sweden andB. lanceolataaccounts for about90Vo of theassemblage (Christensen 1975). Belemnitella pulchra Schulz, 1982 occurs togetherwith species of Belemnell.ain the middle early Maastrichtian of Krons- moor near Hamburg in northwest Germany, and it accounts for about 0.37o of the assemblage(Schulz 1982).The early Maasrichtian belemnite fauna of the Maastricht, Lidge and Aachen areaconsists of speciesof Belemnitell.a(about 207o) and Belemnella (about807o) (compiled on thebasis of datain Keutgen& van der Tuuk 1990). Belemnellą ląnceolata appeurredin the middle part of the B. IąnceolątąZone in Denmark (Schulz L979),probably reflecting a belemnite immigration into Denmark (Christensen1996). The earliest Maastrichtian of Norfolk is not exposed and the early early Mapqtrichtianbelemnite sequenceof Norfolk is totally anomalousand not repeatedelslwhere in Europe (Christensen 1996). It is in ascending order: (1) a mixed BóIómnellalBelemnitellaminor II assemblage;(2) only Belemnitella 476 Late Cretaceous belemnitelhds: CHRISTENSEN minor II; (3) a mixed BelemnellalBelemnitellaminor II assemblage;and (4) Belem- nellą. This belemniteSequence may be explainedby two successiveimmigrations of Belemnella into the Norfolk area. Christensen(1995) established Belemnitella minor III from the earĘ early Maas- trichtian of Norfolk and noted thatthis subspeciesdiffers only from the latestCampa- nian B. minor II by its smaller guard.I have subsequentlystudied abundant material of B, minor from the early early Maastrichtian Craie Phosphatóede Ciply of the Mons Basin in southernBelgium, and it is now known that this supposeddissimilarity is incorrect.Therefore, B. minor trI is placedin synonymywith B. minorll. It has bęen suggestedthat the causeof thę nearly total replacementof Belemnitella by Belemnella at the base of the early Maastrichtian was due to a climatic cooling (JeletzĘ 1951)' andBelemnellą has beenenvisaged as a genusadapted to life in cold water on the basis of its geographicaldistribution (Jeletzky 1951) and morphology (Kongiel 1962).However, palaeotemperaturesof northwestGermany, registeredby oxygen isotope data (6'ó0), contradict this suggestion.The palaeotemperaturewas about 15'C in the early late Campanian, increased to about 17"C in the earliest Maastrichtian and decreasedto about 15.C in the middle early Maastrichtian (SchÓn- feld & Schulz L996).Moreover, If Belemnellawas a cold watergenus, one may assume that it was able to migrate northwards,but it has not been recordednorth of Denmark and NorthernIreland, which lay aboutpalaeolatitude 45"N in the Maastrichtian(Smith et al. 1994).As a matterof fact,Belemnella immigrated southwardsinto the northern Tethyanmargin in Europe (seelater discussion). Belemnella (Belemnella?)praearkhangelskii Naidin, 1964 is an east European species,which is recorded from Ukraine and the Saratow district in Russia (Naidin t975).In northwestEurope it occurs in a very nźfTowhorizon, namely in the middle part of the middle early MaastrichttanBelemnella sumensisZone at Hemmoor and Kronsmoor in northwest Germany (Schulz L979) and at Altembroeck in northeast Belgium (Keutgen in Jagt et al. 1995;Keutgen 1997).Unhońzoned specimensfrom Mpns Klint, Denmark probably came from the same stratigraphicallevel (Schulz 1979). Thus, its appearancein northwest Europe is probably due to a westwards migration(Schulz 1979:p. 53). Belemnella dtsappearedslightly below the boundary between the early and late Maastrichtian at Kronsmoor in northwest Germany, whilst two specimens of B. (Pachybelemnella)cf . cimbrica Birkelun d, 1957have been recorded from the early late Maastrichtian Belemnitella junior Zone of Denmark (see Christensen 1996).Belem- nella was replaced by Belemnitella junior Nowak, t913, which ranged through the entirelate Maastrichtianand occurs in northwestEurope, Poland, westernUkraine and Crimea in the North European Province, in addition to Azerbaijan in the Caucasus in the northern part of the Tethyar,Realm. Christensen (1996: fig. 3) showed that it appeared distinctly later in Denmark (base of the late late Maastrichtian) than in northwestGermany and the Maastricht area(at or immediately above the base of the late Maastrichtian). He suggested,therefore, that the appearanceof B. junior in Denmark was due to either a northwards migration of this species in the latę late Maastrichtian or lack of suitable shallow-waterhabitats during the early late Maas- trichtian. Belemnites are extremely rare in the early late Maastrichtian chalk of Denmark (Christensen1996, 1997),which is known only from outcropssituated in the ACTA PALAEONTOLOGTCA POLOMC A (4f) (4) 477 basinal part of the Dańsh Subbasin. Belemnites are more common in the late late Maastńchtian cha]k of Denmark, which is known mainly from outcropslocated in the marginal part of the basin. Belemnella (Neobelemnella)kazimiroviensis (Skołozdrówna, 1932) is an eastEu- ropeanspecies, which rangedthrough the entire late Maastńchtian on the easternpart of the Russian Platform Naidin 1973, L975).Christensen (1996: ftg.4) showedthat it appearedlater in Denmark (late late Maastrichtian) and much later in the Maastricht area(latest late Maastrichtian),implying a westwardsspread of this species.Machalski (1996) demonstratedthat B. (N.) kazimiroviensis appearedearlier in Poland than in Denmark on ammoniteevidence, that is in theupper part of the early lateMaastrichtian. He thus provided supportingevidence for the westwardsmigration of this species. The belemnitellidsbecame extinct at the Cretaceous-Palaeogeneboundary, as did many other fossil groups, including the ammonites.This major extinction event was discussedby Kauffman & Hart (1996) and summarizedby Barne-sat aI. (1996).It is associatedwith a worldwide iridium anomaly,major 613Oand6180 excursionsand a global sea-levelfall. The cause of the iridium anomaly has been discussedsince 1980 and attributedto the impact of a major bolide or volcanism. It is beyond the scope of this paperto discussthe end Cretaceousmass extinctionsin detail (seeKeller 1996 and Macleod et aI. 1997for two recentreviews). Kennedy (1989) noted that the disappearanceof the ammonitesand bęlemnitesat or close to the Maastrichtian-Danian boundary is a particular event in a general phenome- non. Kennedy (1993)suggested that the ammoniteand belemniteextinction at the end of the Cretaceous may have been the result of the collapse of marine plankton and sub- sequentcut-off of food supply for the tiny ammoniteand belemnitehatchlings. He noted, however,that equally small hatchlings of other coleoid groups survived the crisis.

Tethyan Realm

Belemmnitellids invaded the northern European margin of the Tethyan Realm inter- mittently during the Late Cretaceous,and species of five genera and two subgenera have been recorded (Fig. 7): Praeactinocamax, Gonioteuthis,Belemnellocamnx, Be- lemnitella, Belemnella (Belemnella),B. (Pachybelemnella)and B. (Neobelemnella) (Christensen 1997).The majority of the species occurring in the Tethyan Realm are conspecific with thosefrom the North EuropeanProvince and thus provide a basis for correlation.Endemic speciesof Belemnitella of latę Campanian age occur in Austria, Azerbaijan, Bulgaria and Romania. Praeactinocamaxplenus was recordedrecently from the middle late Cenomanian of les Lattes in the Vocontian Basin, Alpes-Maritime of southeastFrance (Gale & Christensen 1996),in a bed equivalent to Jefferies' Bed 4 of the Plenus Marls of the Anglo-Paris Basin. Thus, it was the first belemnnitellid that invaded the TeĘan Realm. P plenus occurs theretogether with elementsof the North Boreal faunal group of Jefferies (1962) (see above). The small sample of P plenus, consisting of nine specimens, includes all growth stages, indicating that the species bred there. The occurrence of P plenus at les Lattes extends its distribution southwardsby nearly 600 km from the southernmostlocality in the Anglo-Paris Basin. This southwards 478 Late Cretac eous b elemnitelhZs; CHRISTENSEN

P. G. Blt Belemnitella Belemnella U) := z (s a s s $ śe .a o IU Q) (') o Ę tri o o I Ę (Ą Ę 6 P Ę o F E N s E a ł: e o o € g g O) Ę Ę .s a ś a 3 ś o J \ s Ę so) 0) (') E E o E B o o- o E *s s o- x P x CL o il N N $ o. U' Bs o a. o o Ę o a s .s a. B) 6s.4 L ? I +$

E I II *8 71.3 ! *7 T I ? I I I -6 >L ? L I I I I o_ y5 tJJ I I ? Ę ! I \4 JE I I I 80.6 T I

83.5 *3 +2 86.3

88.7

93.3 L I -1 IIJ M

98.5 E

Fig. 7. Stratlgraphical range diagram of Late Cretaceous belemnitellids occurring in the TetĘan Realm (modified from Christensen 1997). Abbreviations: P - Praeactinacamax, G. - Gonioteuthis, BIc. - Belemnellocamax.Stage abbreviationsas in Fig. 1. Ages in Ma afterObradovich (1994). belemnite migration into the TetĘan Realm was due to a significant fall of sea temperature,the Plenus Cold Event, working in concert with the development of suitable shallow water habitats (Gale & Christensen 1996). In addition, lack of competitionof the TeĘan belemnopseidsmay also have been of significance for this migration. Belemnitellids are not recorded from the Turonian to middle Santonian of the TeĘan Realm. In contrast,they occur in the North Ameńcan Province during this time (seebelow). Belemnitellapraecursor,Gonioteuthis sp. andBelemnellocamax ex gr.grossouvrei were recordedfrom the late Santonianand earliest early Campanian of the Corbiires in the easternpart of the French Pyrónóes by Christensen et aI. (1990: fig. 1) and Christensen,Bilotte, & Hansotte(1993). Atthę classicsectionbetween Sougraigne and Croutets,B. praecursor occars in the latest late Santonianand B. ex gr. grossouvrei occurs in the earliestand latestlate Santonian.Gonioteuthis sp. and Belemnellocamax ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 479 ex gr. grossouvreloccur in the earliest early Campanian atLaBastide. B. praecursor is recorded also from the late Santonianand earliest early Campanian of Azerbaijan (Ali-Zade 1972). The early Campanian Gonioteuthis quadrata and late Campanian Belemnitellą mucronatawere recordedfrom the Aquitaine Basin by previous authors(summarized by Sóronie-Vivien 1972: pp. 135_136),but these records need to be confirmed. Previous records of B. mucronatąfrom the northernmargin of the TethyanRealm, for instance the Balkans and Turkey, also need to be confirmed, becausemost previous authorslumped aI|Belemnitella mucronąta-likeforms, that is small and large species of Belemnitella,as wellas speciesof Belemnella,in B. mucronatą. Late Campanian species and subspecies of Belemnitella from Romania were describedbyNeagu & Georgescu(1991), Bulgariaby Stoyanova-Vergilova& Jolkicev (1993)and Azerbaijan by AIi-Zade (1972).They recordedB. mucronata,B. minor, B. langei and B. najdini, as well as subspeciesof these taxa, all known from the North European Province, in addition to some local species and subspecies.However, the descriptions of the species from Romania and Bulgaria leave much to be desired, becausethe internal charactersare rarely studied or not studied at all, and the quality of the platesin the paperby Neagu & Georgescu(1991) is extremelypoor. Combómorel (1996)described a small sampleof Belemnitelląsp., consisting of five nearly-completespecimens and about 30 fragments,from the Sub-Alpine Chain at Chartreusenear Grenoble in the Savoie, southeastFrance (Fig. 3). This speciesis late late,but not latest,Campanian in age and may be conspecific with B. hoeferl (Schloen- bach, 1867)(Christensen in press).However, a specific determinationof B. sp. is not possible, because the internal characters are unknown. Combómorel figured only nearly-complete specimens, which are fully-grown, but all growth stages occur (R. Combómorel personal communicationFebruary 1997).B. sp. formed, therefore, a breedingpopulation at Chartreuse. The little known, but valid latelate, but not latest,Campanian Belemnitella hoeferi was describedby Christensen (in press) on the basis of material from the Gschlief- grabentectonic window of the northernUltrahelvetic series and the Gosau Group of the Northern Calcareous Alps, Austria (Fig. 3). It belongs to the B. mucronata group of Christensen( 1995),and can be distinguishedfrom most speciesof this group on the basis of the slenderguard. The sampleof B. hoeferi from the GschliĘraben includes all growth stages,indicating that the species bred there. Christensęnsuggested that B. hoeferi evolved by allopatric speciation from an initial migrant, which may have invaded the northernEuropean margin of the TetĘan Realm coincidently with the late late Campanian transgressionof Hancock (1990, 1993). Christensen also described B. cf . minor II from the latest late Campanian of the Gosau Group. Thus, this species invaded the TetĘan Realm slightly later thanB. hoeferi. These recordsof speciesof Belemnitella indicatethat this genusspread southwards into the TethyanRealm severaltimęs during the late Campanian. The genusBelemnitellahas not been recordedfrom the early Maastrichtian of the Tethyan Realm, with one exception. Schulz & Schmid (1983) recorded a single, fully-grown specimenof B. pulchra from the middle early Maastrichtian,lower part of the Belemnella sumensis Zone, of Bavaria in southernGermany (Fig. 3), implying a southwardsmigration at that time. 480 Inte Cretaceous belemnitelhds: CHRISTENSEN

Ali-Zade (I97f) recorded Belemnella lanceolata lanceolata and B. sumensisoc- cidentalis Birkelund, 1957from the early Maastrichtianof Azerbujan (Fig. 3). Schulz (1979) placed the first in synonymy with the early early Maastrichtian B. (Pachybe- lemnella) inflata (ArkhangelsĘ 1912) and the latter in synonymy with the middle early Maastrichtian B. (8.?) praearkhangelskii. Moreover, Naidin (1975) and Schulz (L979) reportedB. (8.?) praearkhangelskii from westernKazak*rstan and Azerbaijan. B. (B.) gracilis (ArkhangelsĘ, I9I2) occurs abundantlyin the middle early Maastrich- tian, lower part of the Belemnella sumensis Zone, of Bavaria in southernGermany (Schmid & Schulz 1979; Schulz & Schmid 1983). All growth stages are present, implying that the speciesbred there. Therefore,the genus Belemnella immtgratedinto the TethyanRealm twice in the early Maastrichtian, viz. in the earliest Maastrichtian and in the middle early Maas- trichtian.The lattermigration is coeval with the B. pulchra migration. Belemnitellajunior andBelemnella (Neobelemnella)Icazimiroviensis are recorded from theentire late Maastrichtianof Azerbaijan in Caucasus(Ali-Zade lg7f),implying a southwardsimmigration of these species into the TethyanRealm at the base of the late Maastrichtian. Alt-Zade (1972)recorded B.(N.) l

North American Province

This province includes Greenland, the Western Interior of North America, in addi- tion to the Atlantic and Gulf coasts of the USA (Figs 8, 9). Belemnitellids are ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 48r

Fig. 8. Approximate distribution of land and sea in North America and Greenland during Turonian and Coniacian time. 1 - Kangerlussuaq area, central East GreenLand;f - Geographical Society and Traill islands, central East Greenland;3 - Svartenhukand Nuussuaq peninsulas,central West Greenland. Dots show occurrencesof Turońan to SantonianPra eactinocarnax.Arrows show probablymigration routes. The distributionof land and sea is based on JeletzĘ (I97I) and Williams & Stelck (1975) for North America and Christensen(1993b) for Greenland.Williams & Stelck (1975) postulatedan extensionof the Western Interior Sea to central West Greenland across the area of the Hudson Bay and Baffin Island. In contrast, JeletzĘ (197I) suggesteda marine link betweenthe WesternInterior and centralWest Greenlandthrough the CanadianArctic Archipelago.

generallyvery rare and belemnopseidshave not beenrecorded. Christensen (1993b, 1997)showed that the belemnitefaunas consist essentially of speciesof Praeactino- camax (Turonian to early Santonian)and Belemnitella (IatestSantonian to Maas- trichtian)(Fig. 10).The genusActinocąmax is representedonly by one Specimenof A. verus? from centralEast Greenland(Jeletzky in Donovan 1954)(see below) and two specimens of A. aff. laevtgatus Arh,hangelsky,I9I2 from Kansas (Jeletzky t96I). However,Jeletzky (1961)noted that the two specimensfrom Kansas may be juveniles of one of the Specięsof Praeactinocamax.The middle Tuionian belemnite fauna seemsto be rather diverse,but this is probably due to excessive subdivision (Christensen1997)

Greenland Belemnitellids are extremely rare in Greenland and about 30 specimenshavę been recorded.Twenty Specimenshave been dęscribedfrom central East Greenland:two adult Specimensof the middle Turonian Praeactinocamcłxcf. manitobensis flrhi- teaves,1889) from the Kangerlussuaqarea (Fig. 8) (Christensen& Hoch 1983),in 48f Late Cretaceous belemnitelhZs: CHRISTENSEN additionto one specimenof Actinocamaxverus? and 13 specimensof Belemnitellaex gr.alpha/praecursor from the latest Santonian-earliestCampanian of the Geographi- cal Society island (Fig. 8) (Jeletzkyin Donovan 1954;Christensen 1993b, I997).The small sample of B. ex gr. alpha/praecursorincludes all growth stages,implying that the group bred there.The remaining materialis undeterminable. The small fauna of central West Greenland (Fig. 8), consisting of nine specimens, was describedby Birkelund (1956).She recordedthree specimensof the late early- middle Coniacian Praeactinocamax sp., four specimens of the early Santonian P groenlandicus (Birkelund, 1956),one specimenof P. cf . primus and one specimenof P. aff. groenlandicus(- ?P. manitobensis,see below). The age of the two last-men- tioned specimensis unknown (Birkelund 1956).P. primus occurs in the early and middle Cenomanian of the North European Province. The validity of Birkelund's determinationof P cf. primus was questionedby Christensen(1997), because the specimen is very poorly preserved and Cenomanian belemnites are not recorded elsewherein the North American Provincę. Moreover, Dr H. Nphr-Hansen,Copen- hagen has kindly studied dinoflagellate cysts obtained from the matrix of this spe- cimen, and theseare most likely of Coniacian-Santonianage (personalcommunication 20 February 1997).

North America The following speciesof Praeactinocamax have been recordedfrom North America: P manitobensisand P aff. plenus (describedby Jeletzky 1950, 196I); P. sternbergi (JeletzĘ 1961)and P walkeń (Jeletzky,1961) (described by JeletzĘ 196I);and P cobbąni (Christensen,1993). P. sternbergiis probablya junior Synonymof P mani- tobensis,as is P aff. groenlandicus(Christensen & Hoch 1982;Christensen 1997). Four species of Belemnitellą have been reported from the United States. Jeletzky (1955)described a single,adult specimen of theEuropean species B. praecursor from Kansas. Ieletzky (1960, I96f) recognizedthree late Campanianand Maastrichtian species:B. americana(Morton, 1830),B. bulbosaMeek, 1876,and '8. cf. bulbosa', the concept and stratigraphyof which are poorly known. They were discussed by Christensen(1997). Praeactinocamcłxmanitobensisis widely distributedin the middle Turonianof the Western lnterior, from Manitoba, Saskatchewan.Alberta and British Columbia in Canada southwardsto Kansas (Cobban I99I: fig. 1) (Fig. 8). P aff. plenu,vwas establishedon thebasis of a singlespecimen from Manitoba(discussed by Christensen 1993b)and P wąlkerion thebasis of two specimensfrom Kansas.Therefore, both taxa are poorly known. Seibertz & Spaeth(1995) described Prąeąctinocamax cf . mąnitobensisfrom the early Turonianof northernMexico (Fig. 8). Christensen(1997) regarded the validity of this determinationas opento questionfor thefollowing reasons:(1) the anteńorend with thecritical characters is missingin thespecimens; (2) P manitobensishas not been recordedpreviously south of Kansas (Cobban I99I): (3) P manitoberzslsis of middle Turonian age elsewhere;and (4) belemnitellidshave not been recordedso far south elsewhere;northern Mexico lay at palaeolatitude15-20'N. ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 483

Fig. 9. Approximate distribution of land and sea in North America and Greenland during early Maastrichtian time. Dots show the occunences of late Campanian and Maastrichtian Belemnitella. Arrows indicate migration routes. Modified after Williams & Stelck (I975).

Praeactinocamax cobbąni occurs in the late early and middle Coniacian of Monta- na and Wyoming, and the sample from Montana, consisting of about 150 specimens, includes all growth stages,implying that the species bred there (Christensen 1993b). Belemnitella ąmericanahas a flood occunence in the Mound Laurel and lowest paft of the Navesink F'ormations of the northern Atlantic Coastal Plain of New Jersey, Delaware and Maryland (Owens & Sohl 1973 ftg.4; Sohl 1987). This species is of late late Campanian and early Maastrichtian age (Christensen 1997). All growth stages are present, indicating that the species bred there. B. americana sensu Jeletzky (I96f) is an extremely variable species, and the range of variation of both external and internal characters is, in fact,larger in this species than in any other species of Belemnitella. Jeletzky (I96f) established seven varieties of B. americana, and var. americana is the most common variety; it constitutes 6U957o of the populations studied by him. He suggested thatB. americana is derived from a species of the European late Campanian Belemnitellalineage. 'Belemnitella Jeletzky (1960, I96f) recorded the younger late Maastrichttan cf. bulbosa' from the southern Atlantic Coastal Plain and Gulf Coastal Plain of the Carolinas, Alabama, Mississippi and Texas. He failed to describe this taxon but noted that it has a larger Schatzky Distance than B. americana.Ihavę studied two specimens of Belemnitellafrom the late Maastrichtian Prairie Bluff Formation of Alabama. These are closely similar to B. ąmericąna yar. americana with regard to shape, slenderness and surface markings, but are slightly smaller and have a larger Schatzky Distance, a smaller fissure angle and a smaller alveolar angle. 484 Late Cretac eous b elemnitellids; CHRISTENSEN

The late Maastrichttan Belemnitellą bulbosa is poorly known, because it was establishedon thebasis of only a few specimensfrom the Fox Hills Formation of South Dakota and the internal charactersare unknown. This species.is small and extremely slender as interpretedon the basis of its types. I have studied 10 specimęnsof Belemnitella sp. from the Pierre Shale of South Dakota,Baculites baculus andB. clinolobatusZones(Christensenunpublished). These zones are of early, but not earliest,Maastrichtiarr age.B. sp. is closely similar to B. bulbosa with regardto size and slendernessand may be conspecific with this species. I have also studieda single specimenof B. cf . bulbosafromthe Jeletzkytesnebrascen- sisZone of the Pierre Shale of SouthDakota.This zone is late late Maastrichtianin age and ranges into the highest Maastrichtian Mtcula prinsii Subzone CC26B of the nannofossil sequence.If B. sp. is conspecific with B. bulbosa,,then the latter ranges throughthe Maastrichtian,except its basal and uppermostpart. Thus, the stratigraphi- calrange of B. bulbosamay overlapwith thatof B. americąnąin the early Maastrich- tian and B. bulbosamay be partly coeval with '8. cf . bulbosa'. Jeletzky (1960)suggested that,B. cf . bulbosą,and B. bulbosa arederived from B. americana. Howęver, this suggestionis open to discussion,because the two taxa are poorly known. Discussion. - It can bę concluded that the majority of the belemnitellids of the North American Province are endemic. The endemic species include the Turonian- early Santonian species of Prąeąctinocąmaxand the late Campanian_Maastrichtian species of Belemnitella. Moreover, these have a punctuated,strongly discontinuous stratigraphicaldistribution and do not representan evolutionarylineage (Christensen 1993b,1997).They probably evolved from European speciesby allopatric speciation from initial migrants. It might be worth emphasizingthat, in contrastto Europe, the only Maastrichtian belemnitellids in North America areBelemnitella. Three Europeantaxa occur in the latestSantonian--earliest Campanian of the North American Province:Belemnitella praecursor in the WestemInterior, and Actinocamax verus?andBelemnitellą ex gr.alpha/praecursorin centralEast Greenland (see above). Thus, the belemnitellids immigratedinto the North American Province at least six times during the Late Cretaceous.These immigrations are in ascendingorder (Fig. 10): I) Praeactinocamaxmąnitobensisandother poorly known speciesof this genusin the middle Turonian. 2) Praeactinocęmaxcobbani and P sp. in the late early Coniacian. 3) Praeactinocamaxgroenlandicus inthe early Santonian. 4) Actinocanrcmverus?, Belemnitella praecursor andB. ex gr.alpha/praecursor in the latestSantonian-earliest Campanian. 5) Belemnitellaamericanainthe late late Campanian. 6) Belemnitella bulbosa probably in thę early, but not earliest,early Maastrichtian. The earliest belemnitellids from the North American Province are of middle Turonian age,an unusual situation in view of the very limited nea of distribution of belemnitellids in the Turonian of the North European Province (see above). The belemnitellids disappearedin the late Maastrichtian,some way below the Maastrich- tian-Danian boundary. ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 485

Praeactinocamax A. Belemnitella a (s O (,) z IJJ o o ui (5 .e € 3 o o (Ą t Ę f, s \ a_ $ tr a o o F q) s St) E o a os .ś Ę E s a I Q) s .ĘI € r $l) Ę t () _4. F ś Ot a. it (l) E * F o p g il Ss s E o a J Ę Ę (a s (U (E o O) I a. o) $ a 65.4

L

E I 6 ? 2 I 71.3

L I <-$ (L o- [! F IE

80.6 E' o-L o- uiE 83.5 <_Ą

86.3 <-3

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+'l 93.3 L z- UJM o_ 98.5 E

Fig. 10. Stratigraphicalrange diagram of Late Cretaceousbelemnitellids of the North American Province (modified from Christensen 1997). Stage abbreviationsas in Fig. I; A. - Actinocamax. Ages in Ma after Obradovich (1994\.

The endemic species of Praeactinocamnx occur in central East Greenland, central West Greenlandand the WesternInterior Basin of Canada and the USA (Fig. 8). They probably immigrated into the North American Province by a northern route, from northwestEurope via Greenland and Arctic Canada to the Western Interior Seaway (Jeletzky1950, t971; Birkelund 1965;Christensen & Hoch 1983)(Fig. 8). Belemnitellaąmericanatsrecorded from theAtlantic CoastalPlain, from Maryland and northwards,and '8. cf . bulbosa'from the Atlantic Coastal Plain southof Maryland and the Gulf Coast Plain. These may have immigratedvia a southernroute (Fig. 9). B. bulbosa ęnteredthe WesternInterior from the south. The majority of the belemnitellid species of the North American Province are endemic, and, consequently,intercontinental correlation based on belemnites is not possible. 486 Late Cretaceous belemnitellids; CHRISTENSEN

Synopsis and possible causes of belemnite migrations

The centreof belemnitellid evolution lay in the North EuropeanProvince, becauseall known genera and subgeneraoccur there and the first members of the genera and subgeneraappeared in this province. As shown above, speciesof five generaand two subgenerainvaded the northernEuropean margin of the Tethyan Realm at least nine times during the Late Cretaceous (Fig. 7). Species of essentially only two genera immigrated into the North American Province at least six times during the Late Cretaceous(Fig. 10).More than a score of migrationshave beenrecognized within the subprovincesof the North EuropeanProvince. Prąeactinocamax immigrated westwardsinto northwestEurope twice during the Cenomanian:P primus in the earliestmiddle Cęnomanianand P plenus in the middle late Cemnomanian.The durationof theseimmigration eventswas very shortlived and they took place coincidently with a rapid rises of sea-level and cool climatic phases, the so-called Primus and Plenus Cold Events. P. plenus is more widely distributedin the North European Province than P primus and it even enteredthe TethyanRealm, probably due to lack of competition from the belemnopseids,which became extinct worldwide in the middle Cenomanian. Since all growth stagesof Praeactinocamaxprimus occur in the Central European Subprovince there were breedingpopulations of this species in this subprovince.All growth stagesof P. plen r,soccur in the westernpart of the North European Province, as well as in the TethyanRealm, implying that it bred there. Belemrtocąmaxboweri became extinct in the middle Cenomanian and P plenus disappearedin the middle late Cenomanian in western Europe and in the earliest Turonian on the Russian Platform. This belemnite extinction, which included both belemnopseidsand belemnitellids,lasted about f.5 Ma and was probably part of the well-known, global middle Cenomanianto early Turonianmass extinction (see above). After the crisis around the Cenomanian-Turonianboundary belemnitellids were extremely rare in the Central European Subprovince and had a very limited area of distribution in the Central Russian Subprovince during the Turonian and early Conia- cian. Moreover, they did not invade the TethyanRealm again until the late Santonian. It appearsthat the Europeanbelemnitellids retreated into refugiain theTuronian, which may have been situatedin the Central Russian Subprovince. While the belemnitęllidsretręated northwards in Europe during the Turonian,they entered,via a northernroute, the North American Province for the first time. Praeac- tinocamaxappeared in the middle Turonian of the WesternInterior of Canada and the United States (P. manitobensis and other poorly known species of this genus) and central East Greenland(P cf. manitobensis). The northwardsretreat of belemnitellidsin Europe, the migration of belemnitellids into the North American Province via a northernroute and the lack of belemnitellids in the TethyanRealm in the Turonian may be causally related.Jenkyns et al. (1994) provided a palaeotemperaturecurve' derived from oxygen isotope data(60Ió), for the Cenomanian to Santonian of East Kent, which lay about palaeolatitude40"N. They showed that the palaeotemperaturereached a maximum of 27_28.C in thę earliest Turonian.Moreover, they notedthat this temperaturecompares well with a palaeotem- peratureof 32-33"C, obtainedfrom fish teethfrom Israel,which lay aboutpalaeolati- ACTA PALAEONTOLOGTCA POLONTCA (4f) (4) 481 tude 10'N. This Late Cretaceous warming peak may be the cause of the northwards retreat of the belemnitellids in the Turonian. The Baltoscandian Subprovince was repopulated by Goniocamax lundgreni inthe early Coniacian and this species spread to the Central Russian Subprovince in the middle Coniacian. The Central European Subprovince was repopulated by Gonioteu- this praewestfalica in the late middle Coniacian. Goniocamax esseniensis entered the Central European Subprovince in the early, but not earliest, Coniacian, G. lundgreni spread to this subprovince in the late early Coniacian and Actinocamax verus cf . antefragilis invaded the Baltoscandian Subprov- ince in the late early Coniacian. These may be considered as occasional immigrants. Prąeactinocamax immigrated, via a northern route, into the North American Province in the late early Coniacian and persistęd into the middle Coniacian: P. cobbani appeared in the Westęrn Interior of the United States and P sp. entered central West Greenland. The reasons for these colonization and migration events cannot be satisfactorily explained at present. Christensen (I97 6: p. I25) suggested that the parallel evolution of the Gonioteuthis and Goniocamax-Belemnitella stocks in the adjacent Central European and Central Russian Subprovinces during the middle Coniacian to early Campanian was due to mutual competition, because the two stocks are likely to have had the SźImegeneral ecological requirements. It seems that species of the Goniocamax-Belemnitella stock attempted to colonize the Central European Subprovince five times during the early Coniacian to early Campanian, namely in the early Coniacian (G. lundgreni and G. esseniensis), early Santonian (G. lundgreni and B. propinqua), earliest early Campanian (8. alpha and B. praecursor), middle early Campanian (B. praecursor) and latest early Campanian (8. mucronata). It was not until the late Campanian, however, that B. mucronata succesfully colonized the entire North European Province, probably due to thę cessa- tion of competition of the genera Actinocamax, Gonioteuthis and Belemnellocąmax. At the Samę time as Goniocamax lundgreni and Belemnitella propinqua invaded the Central European Subprovince in the early Santonian, Praeactinocamax groenlan- dicus entered central West Greenland, Actinocamax verus verus immigrated into thę Central European Subprovince and Baltoscandia and Gonioteuthis praewestfolica spread northwards to Bornholm. In addition, Gonioteuthis ernsti appeared in the Mtinsterland Basin and Baltoscandia, and Belemnitella schmidi entered Baltoscandia and the Russian Platform from an unknown area. Thus. several southwards and northwards migrations occurred during early Santonian time. Some of the species were occasional immigrants, which did not form breeding populations, for instance G. lund- greni and B. propinqua. Other species formed breeding populations, among others A. verus verus and G. praewestfalica. In Baltoscandia there was a major faunal turnover in the middle early Santonian. Belemnite assemblages below are predomi- nated by Goniocamax and above by Gonioteuthis. The cause of the migrations in the Santonian and the belemnite turnover in the early Santonian cannot be satisfactorily explained for the time being. Belemnitella praecursor and Belemnellocąmąx ex gr. 7rossouvrei invaded the Tethyan Realm in the late Santonian and Gonioteuthis entered this realm in the earliest early Campanian. B. praecursor, B. ex gr. alpha/praecursor and Actinocamax verus? 488 Late Cretaceous belemnitelllds: CHRISTENSEN

spreadto the North American Province in the latest Santonian-earliestearly Campa- nian. B. alpha and B. praecursor immigrated into Baltoscandia and the Central European Subprovince,respectively, in the earliest early Campanian.Thus, the genus Belemnitellamigratedboth northwardsand southwardsaroundthe Santonian-Campa- nian boundary.Again, some of these taxa were occasional migrants,for instanceB. praecursor in theUnited Statesand the Corbiires, B. ex gt.grossouvrei in theCorbiires andA. verus?in centralEast Greenland.In contrast,B. alpha andB. praecursor formed breeding populations in Baltoscandia and the Central European Subprovince. These migrationsoccurred during the time of relatively low sea-level(Schulz et al. 1984;Haq et al. 1987;Hancock1990,1993). Belemnitellapraecursor invadedthe Central European Subprovince in the middle early Campanian and formed breeding populations. This migration occurred coin- cidently with a rapid rise of sea-level (Hancock 1993). This is the Offaster pilula transgressionof German workers (Niebuhr et al. 1997,among others). BelemnitelląmucronatainvadedtheCentral European andBaltoscandian Subprov- inces in the latestearly Campanian,where it formed breedingpopulations. At the same time Belemnellocamaxmammillatus migrated southwardsinto the Central European Subprovince and eastwardsinto the Central Russian Subprovince,but it did not breed there. These migrations occur coincidently with a rapid rise of sea-level (Hancock 1993). This is the Belemnitella mucronata transgressionof German workers, e.g. Niebuhr et al. (1997). Three out of four generadisappeared at or near to the boundarybetween the early and late Campanian, namely Actinocamax, Gonioteuthis and Belemnellocamax,ffid only Belemnitella survived. This genus spreadsouthwards in the earliestlate Campa- nian and occurred in the entire North European Province, probably due to lack of competitionof the extantgenera. It even invaded the TeĘan Realm. It seemsthat small speciesof Belemnitellą,that is B. langei' B. najdini and B. ex gr. langei/najdini,immtgratedinto the Norfolk threetimes duringthe late late Campanian. Belemnitella americana enteredthe North American Province and B. hoeferi and B. sp. invaded the Tethyan Realm in the late late Campanian, where they formed breeding populations.These speciesmay have evolved by allopatric speciationfrom initial migrants,which immigrated northwardsand southwardsduring a rapid rise of sea-level (Hancock 1990, 1993).B. cf. minor II invaded Austria slightly later in the latest late Campanian. The genusBelemnella almostoustedthe genusBelemnitellatnthe early Maastńch- tian.Belemnellais widely distributedin the North Europe Province and even invaded the TeĘan Realm. Belemnitella was virtually absentin the TeĘan Realm during early Maastrichtiantime. The anomalousearly early Maastrichtianbelemnite succession of Norfolk may be explainedby two successiveimmigrations of Belemnella. Belemnella tmmigratedinto the TethyanRealm twice during the early Maastrich- tian, B. (Pachybelemnella) tnflata near the base of the Maastrichtian, and B. (8.) gracilis and B. (8.?) praearkhangelskii in the middle early Maastrichtian.B. gracilis formed breeding populations. B. praearkhangelskii migrated also westwards into northwestEurope at the sźLmetime. ACTA PALAEONTOLOGTCA POLONTCA (42) (4) 489

The faunalturnover at thebase of the Maastrichtian,southwards migrations of early Maastrichtian species BeIemneIIątnto the TethyanRealm and virtrral absenceof the genus Belemnitellą in the early Maastrichtian of this realm, as well as other early Maastrichtianmigrations events are apparentlyneither linked to palaeotemperaturenor to sea-levelchanges. These faunal turnoverand migration eventsremain to be solved. Belemnitellajunior and Belemnella (Neobelemnella)kazimiroviensis invaded the Tethyan Realm in the earliest late Maastrichtian.These migrations took place coin- cidently with a rapid rise of sea-level (Hancock 1990, 1993). B. junior migrated northwardsto Denmark in the late late Maastrichtian.B. (N.) knzimiroviensisspread westwardsfrom the the easternpart of the Russian Platform during the late Maastrich- tian and arrived in Poland in the early late Maastrichtian,in Denmark in the late late Maastrichtian and in the Maastricht areain the latestlate Maastrichtian. Thus, it appearsthat the distributionpattern and migration eventsof thebelemnitel- lids to a certain extentcan be explained by sea-levelchanges, cool or warm climatic phases,and mutual competition,but the causeof severalmigrations cannot be satisfac- torily explained at the present.

Acknowledgments

I am grateful to the journal referees, R. Marcinowski (Instifute of Geology, UniversĘ of Warsaw, Poland) and C.J. Wood (Croydon, Surrey, Great Britain), for critical and constuctive comments on the manuscript, and the staff of the Geological Museum, University of Copenhagen, for technical support.

References

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Paleobiogeografia i migracje późnokredowych belemnitów z rodziny Belemnitellidae

WALTER KEGEL CHRISTENSEN

Streszczenie

Belemnitellidae Pavlow, I9I4 to późnokredowarodzina belemnitów, obejmująca dziewtęćrodzajów i dwa podrodzaje.Przedstawiciele rodziny występowali wyłą- Czniena półkuli północnej'tj. w prowincjach paleobiogeograficznychpółnocnoeu- ropejskieji północnoamerykańskiejw obrębiepółnocnego królestwaklimatu umiar- kowanego (North TemperateRealm) oraz na północnych obrzeżlachkrólestwa te- tydzkiego (TethyanRealm) w Europie. W obrębieprowincji północnoeuropejskiej mozna wyróźnić trzy podprowincje: środkowoeuropejską,środkoworosyjską oraz bałtoskandzką.obszaręm, na którym belemnitellidy powstawały i z którego się rozprzestrzeniałyna inne obszary, była prowincja północnoeuropejska.Znane są stąd wszystkie rodzaje i podrodzaje wyróznione w obrębie rodziny. Z prowincji północnoeuropejskiejbelemnitellidy zaplszczaty się na obszary królestwa tetydz. kiego (co najmniej dziewięć razy) oraz prowincji północnoamerykańskiej(co naj- mniej sześĆrazy). Większośćgatunków występującychna obszarze królestwa te- tydzkiego znanych jest takze Z prowincji północnoeuropejskiej,natomiast wie- kszośćgatunków prowincji północnoamerykańskiejto endemity. Te ostatniepo- wstawałyZapewne na drodze specjacji allopatrycznej z imigrantów. W obrębie prowincji północnoeuropejskiejodnotowano ł'ącznieponad dwadzieściawydarzeń migracyjnych między jej poszczególnymi subprowincjami. Niektóre zmiany w roz- mieszczeniu belemnitellidów mozna wiry,aćz eustatycZnyfinoscylacjami poziomu oceanu światowego,wahaniami klimatycznymi, atakżLetłlmaczyćodzia|ywaniami konkurencyjnymi mtędzy poszczególnymi gatunkami. Przy czyny wielu w y darzefi migracyjnych pozostająjednak niejasne.