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Palaeobiogeography and Migration in the Late Cretaceous Belemnite Family Belemnitellidae

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Palaeobiogeography and Migration in the Late Cretaceous Belemnite Family Belemnitellidae 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 <E Blt. mucronata Blt. mucronata Blc. balsvikensis/ Blt. mucronata Gt.q. gracilis/ Blc, mammillatus/ Blt. mucronata/ z Blt. mucronata/ Gt. o. qracilis/ Blt. mucronata q. Zl Gt. scaniensis Btc.'mźmmiltatus CL Gt. q. gracilis U B lt. a lpha/Blt. praec urso r/ Gt.q. quadrata Gt. q. quadrata -- G (f Blt. praecu rsor/A. Iaevigatus/ ŚE Gt. gran ulataq u adrata/ LU Gt. granulataquadrata Gt. granulataquadrata Blt. alpha ( Pten'a-beds') U Blt. praecursor/ L Gt. granulata Gt. granulata 1; Gt. granulata z- Gt. westfalicag ran u lata/ z Gt. westfa licag ran u lata pM Blt. propinqua r propinqua z U Blt. Gt. westfalica/ a- Gt. westfalica I propinqua G Blt. Lc E Gonioteuthis Gc. lundgreni/ Gc. lundgreni uilicus extremelv rare Gt. praewestfalica 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)
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