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The Late Cretaceous Belemnite Family Belemnitellidae: Taxonomy and Evolutionaryhistory

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The Late Cretaceous Belemnite Family Belemnitellidae: Taxonomy and Evolutionaryhistory The Late Cretaceous belemnite family Belemnitellidae: Taxonomy and evolutionaryhistory WALTER KEGEL CHRISTENSEN Christensen,W. K.: The Late Cretaceous belemnite family Belemnitellidae: Tax­ onomy and evolutionary history. Bulletin of the Geological Society of Denmark, Vol.44,pp.59-88.Copenhagen, 1997-03-15. https://doi.org/10.37570/bgsd-1998-44-04 The Late Cretaceous belemnite familyBelemnitellidae Pavlow, 1 914 occurs only in the Northern Hemisphere and includes nine genera and two subgenera: Praeactinocamax Naidin, 1964b, Actinocamax Miller, 1823, Belemnocamax Crick, 1910,Gonioteuthis Bayle, 1878,Belemnellocamax Naidin, 1964b,Gonio­ camax Naidin, 1964b, Belemnitella d'Orbigny, 1840, Belemnella (Belemnella) Nowak, 1913, Belemnella (Pachybelemnella) Schulz, 1979, Belemnella (Neobelemnella) Naidin,1 975 and Fusiteuthis Kongiel,1962. The latter is most likely a nomen dubium. Diagnoses of the genera and subgenera are provided. The origin of the family is poorly known. The number of genera and subgenera fluctuated during the Late Cretaceous. It was one to two in the Cenomanian, increased gradually to a maximum of six in the Early Santonian,decreased gradu­ ally to one during most of the Late Campanian and increased to two or possibly three in the Maastrichtian. The belemnitellids occur in the North European and North American palaeobiogeographical Provinces of the North Temperate Realm, in additon to the northern European margin of the Tethyan Realm. The centre of evolution lay in the North European Province and all known genera and subgenera occur there. Species of five genera and two subgenera occur on the northern European margin of the Tethyan Realm and the majority of these are conspecific with species from the North European Province. Species of essentially two gen­era occur in the North American Province and these are endemic, with a few exceptions. Key words: Belemnitellidae, taxonomy, evolutionary history, Northern Hemi­ sphere,L ate Cretaceous. W. K. Christensen, Geological Museum, University of Copenhagen, (t)ster Vold­ gade 5-7, DK-1350 Copenhagen, Denmark; E-mail: [email protected]; 3rd October 1996. During the Late Cretaceous, except in the Cenoma­ 1981).Th erefore,a Tethyan Realm cannot be defined nian,belemnites had a bipolar distribution. The Late on the basis of belemnites alone after the Cenomanian, Cretaceous family Belemnitellidae Pavlow, 1914 in­ but it can be recognized on the basis of other fossil habited the North Temperate Realm(= Boreal Realm groups,such as rudists, ammonites, echinoids, actae­ of authors) and the late Early and Late Cretaceous onellid gastropods and larger foraminifera. (Aptian to Maastrichtian) family Dimitobelidae White­ The Cretaceous dimitobelids were monographed by house,1924 inhabited the South Temperate Realm( = Stevens (1965) and dealt with later by Doyle (1985, Austral Realm). The last genera of the familyBelem­ 1987a,1987b, 1988, 1990, 1992),Doyle & Zinsmei­ nopseidae Naef,1922, the mid-Cretaceous(Aptian to ster (1988) and Doyle & Howlett (1989). Cenomanian)Neohibolites Stolley,1911 andParahi­ The belemnitellids were distributed in the North bolites Stolley, 1919 inhabited the Tethyan Realm,but American and North European palaeobiogeographical occurred also in the North Temperate and South Tem­ Provinces of the North Temperate Realm,in addition perate Realms, where they may be locally abundant. to the northern part of the Tethyan Realm in Europe They became extinct in the Earlyand Middle Ceno­ (Christensen 1975a, 1976, 1988, 1993b). The centre manian(Combemorel, Christensen, Naidin & Spaeth of evolution of the belemnitellids lay in the North Christensen : Late Cretaceous Belemnitellidae 59 European Province, and they invaded the North Ame- comprise A. manitobensis (Whiteaves, 1889), A. aff. rican Province and Tethyan Realm at various times plenus and A. walked Jeletzky, 1961. Three granu- during the Late Cretaceous. The belemnitellids are lated taxa from this province: A. groenlandicus Birke- common in the North European Province and all lund, 1956, A. aff. groenlandicus and A. sternbergi known genera and subgenera are recorded. They are Jeletzky, 1961 were placed in Gonioteuthis (Gonioca- generally rare in the North American Province, and max) with a query. essentially two genera are recorded. The belemnitellids The classification of Naidin was discussed by J. A. occur rarely in the Tethyan Realm in Europe and five Jeletzky in his unpublished manuscript for the Trea- genera and two subgenera are reported. tise on Invertebrate Palaeontology (copy obtained by The belemnitellids were stenothermal shelf dwell- me in 1972; it is referred to below as unpublished MS ers, but it appears that the breeding, spawning, hatch- 1972). He placed Goniocamax in synonymy withAcfi- ing, and, possibly for the females at least, dying nocamax (Praeactinocamax) and considered Actino- grounds, were inner neritic, shallow water environ- camax (Paractinocamax) as a subgenus of Belem- ments. nellocamax. Jeletzky regarded granulation to be of The belemnitellids are of fundamental importance great taxonomic significance (see below), and in con- in biostratigraphy and correlation in Europe, because trast to Naidin (1964b) he placed the granulated or they are common, widely distributed and their fos- supposedly granulated species paderbornensis, bohe- silization potential is high (Christensen 1990b). micus, groenlandicus and sternbergi in Actinocamax The aim of this contribution is to discuss the sys- (Actinocamax). He placed the species westfalicus, tematic palaeontology and evolutionary history of the lundgreni, manitobensis and intermedius in Actinoca- belemnitellids. Representative species of the genera max (Praeactinocamax). It has been shown subse- are figured on Plates 1-2. quently that paderbornensis is not granulated (Chri- stensen 1982) and manitobensis may be granulated (Christensen & Hoch 1983). Classification of the Belemnitellidae In this paper I recognize nine genera and two subgenera within the Belemnitellidae: Praeactinoca- Up to 1964, five genera within the Belemnitellidae max Naidin, 1964b, Actinocamax Miller, 1823, had received general recognition by most authors, in- Belemnocamax Crick, 1910, Gonioteuthis Bayle, cluding Wright & Wright ( 1951 ) : Actinocamax Miller, 1878, Belemnellocamax Naidin, 1964b, Goniocamax 1823, Gonioteuthis Bayle, 1878, Belemnocamax Naidin, 1964b, Belemitella d'Orbigny, 1840, Belem- Crick, 1910,£e/emmte//ad'Orbigny, 1840 and #e/em- nella (Belemnella) Nowak, 1913, Belemnella (Pachy- nella Nowak, 1913. In contrast, Kongiel (1962) did belemnella) Schulz, 1979, Belemnella (Neobelem- not recognize Belemnella. nella) Naidin, 1975 and Fusiteuthis Kongiel, 1962 Naidin (1964b) proposed a new classification of the (Fig. 1). Belemnocamax and Fusiteuthis are monö- belemnitellids without a true alveolus, that is species typic, but the latter is most likely a nomen dubium with a convexly conical alveolar fracture, a flat ante- (see below). The classification of Naidin and his ge- rior end or a pseudoalveolus (see below). He erected neric assignment of various species are discussed fur- two new subgenera of Actinocamax, A. (Praeactino- ther below. camax) and A. (Paractinocamax); one new genus, Belemnellocamax; and one new subgenus of Gonio- teuthis, G. (Goniocamax). He placed two small spe- cies, A. verus Miller, 1823 and A laevigatus Arkhan- Origin and evolutionary history of the gelsky, 1912, in Actinocamax (Actinocamax); two large species, A. primus Arkhangelsky, 1912 and A. Belemnitellidae plenus (Blainville, 1825-1827), in Actinocamax The earliest species of the Belemnitellidae, Praeac- (Praeactinocamax); one species, A. grossouvrei Janet, tinocamax primus, appears abruptly in the Early 1891, in Actinocamax (Paractinocamax); and one Cenomanian, some way above the base of the stage species, A. mammillatus (Nilsson, 1826), inBelemnel- (Christensen 1990a). Jeletzky (1946, unpublished MS locamax. 1972) suggested that the belemnitellids are derived The following species from the North European from Aptian belemnopseids, either Neohibolites ewal- Subprovince were placed in Gonioteuthis (Gonioca- di (Strombeck, 1861) or N. clava Stolley, 1911, be- max): A. lundgreni Stolley, 1897, the type species, A. cause the shape of the guard and the structure of the westfalicus (Schlüter, 1874) and A. intermedius alveolar end of these species and the earliest belem- Arkhangelsky, 1912, as well as two new species, nitellids are closely similar. Doyle (1987a, 1988a, matesovae Naidin, 1964b and medwedicicus Naidin, 1992) suggested that the belemnitellids may have 1964b. A. strehlensis (Fritsch and Schlönbach, 1872), evolved from a northern endemic stock and the A. bohemicus Stolley, 1916 and A. paderbornensis dimitobelids from a southern endemic stock of the Schlüter, 1894 were placed in the subgenus with a Tethyan belemnopseid Hibolithes de Montfort, 1808 query. Species from the North American Province during the later part of the Early Cretaceous. The sug- 60 Bulletin of the Geological Society of Denmark gestions by Jeletzky and Doyle suffer the disadvan- the similarity of the shape of the guard and the struc- tage, however, that the earliest belemnitellid is sepa- ture of the alveolar end, as well as other characters, in rated from Neohibolites and Hibolithes, respectively, the two species. The suggestion by Naidin & Alekseev by a considerable time gap. N. ewaldi is middle Aptian is not tenable, because P. primus appears earlier than and N. clava is early Late Aptian in age
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