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 species, 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 (Mutterlose, JV. repentinus. The similarity in various characters in Schmid & Spaeth 1983), Hibolithes became extinct P. primus and A7, repentinus may be a case of synchro- in the basal Aptian in the Boreal Realm, in the middle nous homeomorphy. Barremian in the Mediterranean Province and at the It can thus be concluded that the origin of the Belem- Hauterivian-Barremian boundary in the Indo-Pacific nitelidae is poorly known. Province (Mutterlose 1988). The time gap between Jeletzky (1946, unpublished MS 1972) further sug- the latest belemnopseids and the earliest belemnitellid gested that all belemnitellid genera are derived from is thus at least about 20 million years (Ma). the Cenomanian Praeactinocamax rootstock, imply- Naidin & Alekseev (1975) suggested that/1, primus ing that the family Belemnitellidae is a monophyletic was derived from the early Middle CenomanianNeohi- taxon. This suggestion is, however, open to discus- bolites repentinus Naidin & Alekseev, 1975, which sion. Praeactinocamax and Belemnocamax are most occurs in the Crimea. This suggestion was based on likely not derived from the same ancestral species

CO LU Ü 2 Genera and subgenera oi Belemnitellidae Number of genera and subgenera CO 1 2 3 4 5 6 65.4 •S? I 71.3 JJLî g S JS "B S ^5 ^3 -Q CQ p. rï O CL O OC LU CL •S CL S X I ! 80.6- i CL Ü 83.5" < M 86.3- 05 T iËf 88.7- DC — D M

93.3- U 2 — LU M Ü ï 98.5

Fig. 1. Stratigraphical range and inferred phylogeny of belemnitellid genera and subgenera. Stage abbreviations after Harland et al. (1989). Ages in Ma after Obradovich (1994).

Christensen: Late Cretaceous Belemnitellidae 61 Belemnopseidae Belemnitellidae

Para- Neohibolites Praeactinocamax Actinocamax Gonioteuthis Blc. hib. (O LU

65.4

71.3

Q. Q." 3U

80.6 a IÜL 83.5

86.3 CO 8? 88.7

93.3

B. = Belemnocamax Parahib. = Parahibolites 98.5 Blc. =» Belemnellocamax

Fig. 2. Stratigraphical range diagram of Late Cretaceous belemnopseids and belemnitellids from the North European Province. 1, northwest Europe; 2, Russian Platform; 3, Bornholm, Denmark. Stage abbreviations after Harland et al. (1989). Ages in Ma after Obradovich (1994).

(Christensen 1993c). Moreover, it has been suggested pre-Turonian species of Praeactinocamax, that is P. by Jeletzky (unpublished MS, 1972) and Naidin (1964 primus or P. plenus. Oddly enough, Naidin (1964b: b, Fig. 35) that Actinocamax, which appears in the 171 ) also noted thatActinocamax may be derived from Early Turonian (Naidin 1964b), is derived fromPrae- a small Albian-Cenomanian species of Neohibolites. actinocamax. This suggestion is, however, very ques- In conclusion, the Belemnitellidae is probably a poly- tionable, because the two genera differ in several criti- phyletic taxon. cal characters, including the size of the guard, the Goniocamax enters at the base of the Coniacian (Fig. length of the cone-shaped alveolar fracture, the size 1) (Christensen & Schulz, in press). Naidin (1964b) and shape of the juvenile guard and the growth rela- derived this genus from the Late Turonian species tionship (see diagnoses). Christensen (1993b) and Gonioteuthis (Goniocamax) medwedicicus, which is Christensen & Schulz (in press) showed that growth here placed in Praeactinocamax (see below). Chri- is isometric in Actinocamax, and, consequently, both stensen (1988) suggested that the Belemnitella stock, juvenile and adult specimens are short and stout. including Goniocamax lundgreni, was derived from a Growth is allometric inPraeactinocamax (see below); Turonian-Coniacian species ofPraeactinocamaxfrom juvenile specimens are long and slender (needle- the Central Russian Subprovince. At that time, it was shaped) and adult specimens are stouter. Thus, it seems erroneously believed that G. lundgreni, the earliest not very likely that Actinocamax is derived from a species of the genus, was Late Coniacian in age. G.

62 Bulletin of the Geological Society of Denmark Belemnitellidae

Belemnella Belemnella Goniocamax Belemnitella (Belemnella) (Pachybelemnella) Î2 uî CD < .§ å 65.4

71.3

80.6

IÜL 83.5 < M 03 86.3

88.7 3 M

93.3

F. = Fusiteuthis 98.5 B. (N.) = Belemnella (Neobelemnella) Fig. 2. Continued.

lundgreni appears, however, at the base of the Conia- (1964b) noted that the origin of the genus is unknown. cian. Christensen & Schulz (in press) noted that the Christensen (1988) suggested that Belemnellocamax origin of G. lundgreni is unknown. may be derived from one of the large Cenomanian- Belemnitella appears at the base of the Santonian Turonian species of Praeactinocamax from Europe (Christensen & Schulz, in press). B. schmidi Chri- due to the size of the guard, overall morphology and stensen & Schulz (in press), the earliest species of the ontogeny. It is noteworthy, however, that the time gap genus, may have evolved from Goniocamax lundgreni between the two genera is at least 2.5 Ma. by allopatric speciation. Belemnella enters at the base of the Maastrichtian Gonioteuthis enters in the Middle Coniacian (Ernst (Schulz 1979). It has been suggested that Belemnella & Schulz 1974). Christensen & Schulz (in press) sug- may be derived from Belemnellocamax, because both gested that the earliest species, G. praewestfalica Ernst genera have large guards, long and slender juvenile & Schulz, 1974, may have been derived from Gonioca- guards and a small Schatzky Distance. Furthermore, max lundgreni. the vascular imprints branch off the dorso-lateral dou- Belemnellocamax appears probably at the base of ble furrows posteriorly at an angle exceeding 30 de- the Santonian (Christensen 1986). Jeletzky (unpub- grees in both genera (Nowak 1913; Jeletzky 1949a, lished MS 1972) derived this genus from a Turonian 1951; Christensen 1975a). However, the latest spe- species of Actinocamax (Praeactinocamax). Naidin cies of Belemnellocamax, B. balsvikensis (Brotzen,

Christensen: Late Cretaceous Belemnitellidae 63 The number increases to three to four in the Coniacian Praeactinocamax Belemnitella and reaches a maximum of six in the Early Santonian. Afterwards the number decreases to four from the Ul Middle Santonian to basal Early Campanian and three LU in the late Early Campanian. Only one genus, Belem- nitella, occurs in the Late Campanian, except its basal part. The early Late Campanian Belemnellocamax balsvikensis occurs commonly only in Scania and is virtually absent elsewhere (see discussion below). 65.4 Three genera and subgenera occur in the Early Maa- strichtian and the number decreases to two or three in the Late Maastrichtian, depending the systematic sta- 1 tus of Fusiteuthis. Belemnitella is a very long-ranging genus, occur- 71.3, ring from the Santonian to the Maastrichtian, c. 21 Ma longevity. The generaActinocamax and Praeacti- nocamax are long-ranging, c. 11-12 Ma longevity. The s5 longevity of the generaBelemnellocamax and Gonio- O.Û. teuthis is around 6-7 Ma. Goniocamax and the subgenera of Belemnella are short-ranging, around 2-4 Ma longevity. 80.6, The stratigraphical ranges of the belemnitellids of SU the North European and North American Provinces, 83.5, -Jo_- in addition to the Tethyan Realm are shown in Fig- OL ures 2-4. 86.3,

O M 88.7, Systematic Palaeontology Class Cephalopoda Cuvier, 1795 93.3, Subclass Coleoidea Bather, 1881 Superorder Belemnoidea Hyatt, 1884 LU M C _. Order Belemnitida Zittel, 1895 98.5 Suborder Belemnopseina Jeletzky, 1965 Family Belemnitellidae Pavlow, 1914 Fig. 3. Stratigraphical range diagram of Late Cretaceous [ICZN 1985, Opinion 1328, name no. 572] belemnitellids of the North American Province. A. = Actinocamax. Stage abbreviations after Harland et al. Diagnosis. - Belemnopseina with a conical depres- (1989). Ages in Ma after Obradovich (1994). sion, an alveolus, in anterior part of guard, which is connected through ventral fissure with surface; surface markings consist of dorso-lateral longitudinal depressions and double furrows, single lateral furrows, 1960), is from the early Late Campanian, and the ear- more or less distinct vascular imprints, which cover liest species of Belemnella, B. lanceolata (Schlotheim, all or parts of guard, longitudinal striae, and granules, 1813), is from the earliest Maastrichtian. The two which may form corrugated transverse lines; pro- genera are thus separated by a considerable time gap, ostracum narrow and tongue-shaped, and phragmo- estimated to be about 6-7 Ma. Schulz (1979) suggested cone with a dorsal keel, which fits in a dorsal furrow that Belemnella may have evolved from an unknown in alveolus; guard prolonged ventrally around ventral species of Belemnellocamax or an unknown, closely fissure in a tongue-like extension; a shorter dorsal allied, genus in the late Late Campanian. extension may also be present. Belemnocamax appears at the base of the Middle Cenomanian, has a very short age range and its origin Remarks. - The guard is usually the only part of the is unknown (Christensen 1993c), Fusiteuthis enters skeleton preserved in belemnitellids, and its external in the latest Maastrichtian (Kongiel 1962), but is most and internal characters are used for taxonomic classi- likely a nomen dubium (discussed later). fication. These include 1) size of the guard, 2) shape The number of genera and subgenera of the Belem- of the guard, 3) structure of the adorai end, 4) surface nitellidae is shown in Figure 1. One to two genera markings, 5) internal characters, including the alveo- and subgenera occur in the Cenomanian and Turonian. lar angle, fissure angle, Schatzky distance, and the

64 Bulletin of the Geological Society of Denmark shape of the bottom of the ventral fissure, and 6) ontogeny. These were discussed in detail by Christensen P G. B/c Belemnitella Belemnella (1986). The Riedel-Quotient of Ernst (1964) is the CO ratio of the length of the guard divided by the depth •S GE S 1 ilskii •a

of the pseudoalveolus. The Riedel-Index of Ernst & josjn ST A

Schulz (1974) is the depth of the pseudoalveolus as a AG E percentage of the length of the guard." The Birkelund P ci 1 i iroviens Index of Christensen (1995) is the length from the •9 »s •g apex to the protoconch divided by the dorso-ventral ci. s p S- Q. 8- •S c3i diameter at the protoconch. 65.4 The guard is completely calcified only in the gen-

Christensen: Late Cretaceous Belemnitellidae 65 (guard up to 55 mm long) with isometric growth; gen- age; it possibly also occurs in beds of the same age in erally long cone-shaped alveolar fracture; ventral fis- western Europe (Naidin 1964b). A. quasiverus occurs sure, ventral furrow, and ventral notch absent; single in the Late Santonian of Crimea. lateral furrows weakly developed or absent; granulation present in some species. Genus Praeactinocamax Naidin, 1964b Discussion. — Naidin (1964b) recognized three subgenera of Actinocamax: A. (Actinocamax), type species Type species. - Belemnites plenus Blainville, 1825- A. verus; A. (Praeactinocamax), type species A. 1827, p. 376, PI. 11 bis : 3, by original designation by plenus; and A. (Paractinocamax), type species A. Naidin (1964b: 34). grossouvrei. Christensen (1986) placed A. (Paractinocamax) in Emended diagnosis. — Medium-sized to large belem- synonymy wiihBelemnellocamax, type species Belem- nitellids (guard up to 115 mm long) with allometric nites mammillatus (see later discussion). The legiti- growth; adult specimens stouter and more lanceolate macy of the subgenera Actinocamax and Praeactino- in ventral view than juvenile specimens; usually with camax was initially questioned by Christensen (1982, short cone-shaped alveolar fracture, but some species 1986, 1991, 1993b, 1994), but they were recognized with a very shallow pseudoalveolus; ventral fissure by Gale & Christensen (1996) and Christensen & absent; ventral furrow and ventral notch sometimes Schulz (in press), because they differ in several criti- present; juvenile guard long and slender (needle- cal characters (see discussion above). Subgenus Prae- shaped). actinocamax is here elevated to full generic rank due to these differences. Discussion. - The genus is generally not granulated, Actinocamax has been the subject of excessive sub- but granulation may occur very rarely in some spe- division by Russian palaeontologists. Four species and cies and one species, P. groenlandicus, is granulated. nine subspecies have been established: A. verus (in- The earliest belemnitellid, P. primus, enters some cluding six subspecies), A. laevigatus (including three way above the base of the Early Cenomanian in the subspecies), A. minutus Glazunova, 1972 andA. quasi- North European Province and is followed upwards by verus Naidin, 1953. These were discussed by Chri- the Late Cenomanian P. plenus. The two species are stensen & Schulz (in press), who recognized A. v. ve- closely allied and form an evolutionary lineage rus,A. verus subfragilis Naidin, 1964b, A. verus ante- (Christensen 1974, 1990a). These species are not fragilis Naidin, 1964b, A, laevigatus and A. quasi- granulated. verus. In addition to the two large Cenomanian species, I also include in this genus medium-sized to large spe- Distribution. - Actinocamax occurs mainly in the cies from the Turonian-Early Santonian of the North North European Provinces and is recorded from the European and North American Provinces (Figs 2-3). Early Turonian to the boundary between the lower and Species from the Central European Subprovince com- upper part of the Early Campanian (Fig. 1). It occurs prise: bohemicus, strehlensis and paderbornensis. extremely rarely in the North American Province. The They were revised by Christensen (1982), who showed stratigraphical ranges of the species are shown in Fig- that bohemicus may be granulated, whereas the oth- ures 2 and 4. ers are not granulated. A. v. verus is widely distributed and occurs com- Species from the Central Russian Subprovince in- monly in the North European Province. In northwest clude: planus (Makhlin, 1965), coronatus (Makhlin, Europe it is recorded from the Santonian and early 1965), intermedius, matesovae, medwedicicus, Early Campanian. On Bornholm it enters in the latest mujnakensis (Naidin, 1964b) and aralensis (Arkhan- Early Santonian, slightly above the appearence of gelsky, 1912). These species are not granulated. Gonioteuthis westfalica westfalica (Christensen & Species from the North American Province com- Schulz, in press). In offshore chalks it is most comprise: manitobensis, cobbani, groenlandicus, walkeri, mon in the late Late Santonian; it has an acme occur- sternbergi, aff. primus and aff. groenlandicus (Fig. rence in the upper part of the early Late Santonian 3). P. groenlandicus is granulated (Birkelund 1956) Uintacrinus Zone of southern England. On the Rus- and P. cobbani is very rarely granulated (Christensen sian Platform it enters in the Coniacian. 1993b). P. sternbergi was established by leletzky A. verus antefragilis is recorded from the Early (1961) on the basis of a single specimen, which dif- Turonian of the Russian Platform, and A. verus cf. fers from P. manitobensis only by being granulated. antefragilis was reported recently from the late Early Jeletzky (1961) suggested that P. aff. groenlandicus and Middle Coniacian of Bornholm (Christensen & from central West Greenland, which was similarly Schulz, in press). A. verus subfragilis occurs in the erected on the basis of a single, granulated specimen, late Coniacian of the Russian Platform. may be either a geographical subspecies or a morpho- A. laevigatus occurs on the Russian Platform in the logical variety of P. sternbergi. Christensen & Hoch so-called 'Pteria '-beds of earliest Early Campanian (1983) suggested that P. sternbergi should be placed

66 Bulletin of the Geological Society of Denmark Table 1. Univariate analysis of Praeactinocamax pienus cies may be transitional forms between the genera from the Plenus Marls of the Betchworth Limeworks, Eng- Praeactinocamax and Goniocamax. land. RQ = Riedel-Quotient; RI = Riedel-Index; N = number of specimens; X = mean value; SD = standard deviation; Distribution. - Praeactinocamax occurs in the North CV = coefficient of variation; OR = observed range. American and North European Provinces, in addition to the northern part of the Tethyan Realm in Europe. Character N X SD CV OR It appears in the Early, but not earliest, Cenomanian RQ 10 39.6 15.8 39.8 23.3-87.6 and continues into Early Santonian. The stratigraphical RI 10 2.8 0.9 31.6 1.3- 4.3 ranges of the species are shown in Figures 2-4.

Genus Belemnocamax Crick, 1910

in synonymy with P. manitobensis, because granula- Type species. - By monotypy Belemnocamax boweri tion may occur very rarely in this species. The spe- Crick, 1910, p. 364, PI. 28: 1-2. cies from the North American Province were discussed by Christensen (1993b). Diagnosis. - Very small belemnitellid (guard up to 25 Birkelund (1956) referred one specimen of unknown mm long), conical in lateral and ventral views, and age from central West Greenland to P. aff. primus. This with acute apex; anterior end with pseudoalveolus; determination is open to discussion, because the speci- Riedel-Quotient from about 4 to 7; walls of pseudo- men is enclosed in matrix and only the side is exposed. alveolus sometimes with conellae; guard with long» It is so poorly preserved that it cannot be orientated broad, and deep ventral groove extending from the with certainty. Moreover, P. primus occurs in the anterior end; length of groove 1/3 to 1/4 of length of Lower and Middle Cenomanian in the North Euro- guard; dorso-lateral longitudinal depressions distinct, pean Province, but Cenomanian belemnitellids are not delimiting the dorsal field; dorso-lateral depressions recorded elsewhere in the North American Province. from anterior end and almost to apex; single lateral P. primus usually has a short cone-shaped alveolar furrows present; guard with longitudinal striae, but fracture. Specimens with a flat anterior end are rare otherwise smooth. arid specimens with a shallow pseudoalveolus are extremely rare (Christensen 1990a). A specimen from Discussion. -Christensen (1993c) described/?, boweri Bornholm, MGUH 7834, figured by Birkelund (1957, on the basis of a small sample, consisting of 18 speci- PI; 1: 2) and Christensen (1990a, Fig. 3C), has a mens, from the limestone pit at Wunstorf, west of Riedel-Quotient of c. 46 and a Riedel-Index of c. 2. Hannover, Lower Saxony Basin. In addition, three Christensen (1974) analyzed a large sample of P. specimens from Dörenthe, Münster Basin, and eight plenus from the Plenus Marls of the Betchworth specimens from eastern England, including the holo- Limeworks in southern England. He showed that most type, were also studied. specimens have a low cone-shaped alveolar fracture, B. boweri differs from all other belemnitellid gen- and that about 7% of the specimens have a shallow era by the size and shape of the guard, and by having pseudoalveolus, with all intermediate forms. I have a long, broad, and deep ventral groove. subsequently analyzed the variation of the Riedel- The ancestry of B. boweri is unknown. J. A. Jeletzky Quotient and Riedel-Index of the specimens with a (unpublished MS 1972) suggested that it probably shallow pseudoalveolus (Table 1). represents an early independent offshoot of a still un- P. manitobensis include morphological variants with known pre-Cenomanian form between Neohibolites either a low cone-shaped alveolar fracture, a flat an- and Actinocamax, and that this aberrant genus died terior end or a shallow pseudoalveolus (Jeletzky 1950, out without issue. 1961). According to Jeletzky (1961, PI. 1: 2), specimen no. 7936-2 has a Riedel-Quotient between 17.5 Distribution. - B. boweri is very rare and has a very and 19.4. restricted distribution. It occurs in the early Middle Five species from the Russian Platform have a shal- Cenomanian of the Lower Saxony and Münsterland low pseudoalveolus or a flat anterior end with a cen- Basins in northwest Germany, the Cleveland Basin in tral pit. The Riedel-Quotient is 11 to 13 in the Late eastern England and at Hunstanton, Norfolk. Coniacian P. aralensis and Late Turonian P. matesovae, 16 to 18 in the Late Turonian P. coronatus, 13 to 25 in the Late Coniacian P. mujnakensis and about 15 in the Late Turonian P. medwedicicus. In contrast Genus Gonioteuthis Bayle, 1878 to Naidin (1964b) and Makhlin (1965), who placed these species in Gonioteuthis (Goniocamax), they are Type species. -Belemnites quadratus Blainville, 1827, here assigned to Praeactinocamax, because the pseu- p. 62, PI. 1:9, by original designation of Bayle (1878, doalveolus is relatively shallow. However, these spe- caption to PI. 22: 6-8).

Christensen: Late Cretaceous Belemnitellidae 67 • ' 3

m i

f 1 '14

I i 12

'16 ¥17 21

Plate 1

68 Bulletin of the Geological Society of Denmark Diagnosis. — Medium-sized belemnitellids (up to 85 mm long), with a short cone-shaped alveolar fracture Plate 1 , :;,-,- or pseudoalveolus of varying depth (Riedel-Quotient Figs 1-3. Praeactinocamax pienus (Blainville, 1825-1827), from 3 to 20); guard usually only slightly flattened or Sedgwick Museum, B76709, Betchworth Limeworks, Sur- not flattened at all ventrally; generally subcylindrical rey, England, Plenus Marls, middle Late Cenomanian; 1, or cylindrical in ventral view and cylindrical in lat- dorsal view; 2, ventral view; 3, view of the anterior end showing concentric growth layers and radial ridges, x2. A eral view; juvenile guard short and stout; surface mark- large species of Praeactinocamax. Figured by Christensen ings, including dorso-lateral longitudinal depressions (1974, PI. 4: 4). and double furrows, vascular imprints, striae, and gran- Figs 4—5. Praeactinocamax cobbani Christensen, 1993, ules, well developed; Schatzky distance small, usu- holotype, U.S. Geological Survey, USGS 21411/16, Kevin ally 0.5 to 4.5 mm; bottom of ventral fissure com- Member, Marias River Shale Formation, Montana, Scaphi- monly sine-shaped forming a large angle with the wall tes preventricosa/Inoceramus deformis Zone, early Midd-of the pseudoalveolus; vascular imprints branch off le Coniacian; 4, dorsal view; 5, lateral view. A medium- dorso-lateral double furrows posteriorly at an angle sized species of Praeactinocamax. Figured by Christensen less than 30 degrees; relationship between length of (1993b, Figs 5.17-20). guard and dorso-ventral diameter at the alveolar end Figs 6-7. Actinocamax verus vents Miller, 1823, Geologi- cal Museum, Copenhagen, MGUH 23718, Bavnodde generally allometric. Greensand Formation, Bornholm, Denmark, west of Bavn- odde, between beds 3 and 5, Early Santonian; 6, dorsal Discussion. - Naidin (1964b) recognized two sub- view; 7, lateral view. Figured by Christensen & Schulz (in genera of Gonioteuthis: G. (Gonioteuthis), type spe- press, PI. 1: 2). cies Belemnites quadratus andG. (Goniocamax), type Figs 8-10. Gonioteuthis quadrata quadrata (Blainville, species Actinocamax lundgreni. Christensen & Schulz 1827), Geological Museum, Copenhagen, MGUH 17529, (in press) elevated subgenus Goniocamax to full ge- Vaals Formation, CPL quarry, Hallembaye, Belgium, lower neric rank, following an earlier suggestion by Ernst part of Inoceramus lingua/Gonioteuthis quadrata Zone, early, but not earliest, Early Campanian; 8, dorsal view, 9, & Schulz (1974) (see discussion below). lateral view; 10, view of the anterior end, xl.5. A species The Middle Coniacian to Early Campanian evolu- with a deep pseudoalveolus and well-developed granula- tionary lineage of Gonioteuthis includes, in ascend- tion. Figured by Christensen & Schmid (1987, PI. 3: 8- ing order, G. praewestfalica, G. westfalica, G. 11). westfalicagranulata (Stolley, 1897), G. granulata Figs 11-12. Gonioteuthis westfalica (Schlüter, 1874), Geo- (Blainville, 1827), G. granulataquadrata (Stolley, logical Institute, Lund, Sweden, LO 4859, Ringeleslätt, 1897), G. q. quadrata and G. quadrata gracilis Scania, Sweden, early Santonian; 11, dorsal view; 12, view (Stolley, 1892) (Fig. 2). It has been studied in great of the anterior end, x2. A specimen of G. westfalica which detail by German authors, including Stolley (1892, has a flat anterior end with a central pit. Figured by 1897,1916,1930), Ernst (1964,1966,1968), Ernst & Christensen (1975a, PL 2: 4). Figs 13-15. Belemnellocamax mammillatus (Nilsson, Schulz (1974) and Ulbrich (1971), in addition to 1826), Geological Museum, Copenhagen, MMH 13078, Christensen (1975a, 1975b, 1986,1991,1994), Chri- Ignaberga new quarry, Scania, Sweden, latest Early Cam- stensen & Schmid (1987), Christensen & Schulz (in panian; 13, dorsal view; 14, ventral view; 15, view of the press), Jarvis (1980) and Jagt, Kennedy, Burnett, anterior end, xl.5. Figured by Christensen (1975a, PI. 3: Christensen & Dhondt (1995). In addition to the mem- 6). bers of this evolutionary lineage the following spe- Figs 16-17. Goniocamax lundgreni (Stolley, 1897), Geo- cies and subspecies are placed in Gonioteuthis: the logical Museum, Copenhagen, Denmark, MMH 1690, Early Santonian G. ernsti Christensen & Schulz (in Arnager Limestone Formation, west of Amager, Bornholm, press), which occurs on Bornholm and in the Münster Early Coniacian; 16, dorsal view; 17, ventral view. Fig- Basin and G. quadrata scaniensis Christensen, 1975a ured by Christensen & Schulz (in press, PI. 2: 1). Figs 18—19. Belemnocamax boweri Crick, 1910, Geologi- from the latest Early Campanian of Scania (Chri- cal Museum, Copenhagen MGUH 22068 (cast), Wunstorf, stensen 1975a, 1986) (Fig. 2). According to Chri- Lower Saxony, Germany, early Middle Cenomanian.Acara- stensen (1975a, 1986) the latter taxon is a geographi- thoceras rhotomagense Zone, Turrilites costatus Subzoneca; l subspecies, which is closely allied to G. quadrata 18, ventral view, x3; 19, lateral view, x3. Figured by gracilis. Christensen (1993c, Fig. 3F). The Gonioteuthis lineage survived for around 7 Ma Figs 20-21. Belemnitella mucronata (Schlotheim, 1813), and has been considered as an example of phyletic neotype, Niedersächsiches Landesamt für Bodenforschung, gradualism, viz. gradual transformation of a suite of Hannover, Germany, kca 5/2, Germania IV pit at Misburg characters through time. The general trends in evolu- near Hannover, middle part of the basiplana/spiniger Zone, early, but not earliest, Late Campanian; 20, ventral view; tion are: (1) gradual calcification of the anterior end 21, lateral view. A large species of Belemnitella with a stout of the guard, (2) increasing size and stoutness of the guard. Figured by Christensen et al. (1975, PI. 1: 1). guard and (3) gradual development of granulation (Ernst 1964, Ernst & Schulz 1974). All specimens are coated with ammonium chloride, and In the oldest member of the lineage, the late Middle are natural size unless otherwise stated. Coniacian-early Early Santonian G. praewestfalica,

Christensen: Late Cretaceous Belemnitellidae 69 Plate 2

70 Bulletin of the Geological Society of Denmark less than 10% of the specimens are granulated, whereas niensis is 5.1 (Christensen 1975a), while it is ±4.5 in in the succeeding species, the late Early-early,Middle gracilis (Ernst 1964). Santonian G. westfalica, about the half of the speci- Mitchell (1994, 1995) showed that populations of mens are granulated (Christensen & Schulz, in press). Gonioteuthis from the Late Santonian Marsupites Granulation becomes a very promiment character in testudinarius and Uintacrinus socialis Zones of York- younger species. With respect to the depth of the shire are phylogenetically retrograde in comparison pseudoalveolus, Christensen & Schulz (in press) with coeval populations from northwest Germany. He showed that G. praewestfalica has a deeper pseudo- also demonstrated that populations of Gonioteuthis alveolus than G. westfalica from the early Early from the early Early Campanian of Yorkshire and Santonian, and the depth of the pseudoalveolus is vir- northwest Germany have reached the same evolution- tually identical in G. praewestfalica and G. westfalica ary development. from the late Early Santonian. Thus, the depth of the It can thus be concluded that the evolution of Gonio- pseudoalveolus first decreases from the late Middle teuthis lineage with respect to the depth of the pseudo- Coniacian to the early Early Santonian and this trend alveolus is not as straigthforward as supposed earlier. is reversed in the late Early Santonian. The depth of the pseudoalveolus first decreases and Ernst (1964) and Christensen (1991) noted that co- then increases in its earliest members, and the evolu- eval populations of G. q. quadrata from the middle tionary development of populations of the lineage may Early Campanian chalk of Lägerdorf and marl of be delayed in some areas. Misburg/Höver in northwest Germany differ with respect to the depth of the pseudoalveolus. The speci- Distribution. —Gonioteuthis had its evolutionary cen- mens from Misburg/Höver have a slightly deeper tre in northwest Europe and is recorded almost exclu- pseudoalveolus. The specimens from Lägerdorf are sively from the Central European Subprovince. It oc- thus phylogenetically retrograde compared to coeval curs very rarely in the northern part of the European specimens from Misburg/Höver. G. quadrata scanien- Tethyan Realm. The genus existed from the late Midd- sis from Scania is also phylogenetically retrograde in le Coniacian to the boundary between the Early and comparison with the coeval G. quadrata gracilis from Late Campanian (Fig. 1). The stratigraphical ranges northwest Germany. The mean Riedel-Quotient of sca- of the species are shown in Figure 2.

Genus Belemnellocamax Naidin, 1964b [= Actinocamax (Paractinocamax) Naidin, 1964b, p. Plate 2 62] Figs 1-2. Belemnitella najdini Kongiel, 1962, British Geo- logical Survey, Nottingham, BGS CJW 5478, Caistor, Nor- folk, Beeston Chalk, between Flint 9-10, late Late Campa- Type species. - Belemnites mammillatus Nilsson, nian; 1, ventral view; 2, view of the split anterior end, x 1.5. 1826, p. 340, by original designation of Naidin (1964b: A small, slender species of Belemnitella. Schatzky Dis- 153). tance small, 7.7 mm; fissure angle large, 120.5 degrees; Birkelund Index, 4.7. Figured by Christensen (1995, PI. 4: Diagnosis. - Large belemnitellids (up to 130 mm 1-4). long), with a shallow to relatively deep pseudoalve- Figs 3-5. Belemnitella minor Jeletzky, 1951 form II Chri- olus; guard markedly flattened ventrally, lanceolate stensen, 1995, holotype, Natural History Museum, Lon- or subcylindrical in ventral view, and subcylindrical don, BMNH C43553, Whitlingham, Norfolk, Paramoudra, to slightly lanceolate in lateral view; dorso-lateral lon- Chalk, late, but not latest, Late Campanian; 3, ventral view; 4, lateral view; 5, view of the split anterior end. A stout gitudinal depressions and double furrows usually well- and very large species of Belemnitella with well-developed defined, vascular imprints faint or absent; Schatzky vascular markings. Schatzky Distance large, 12.0 mm; fis- distance small, 0-2 mm; vascular imprints branch off sure angle small, 15.5 degrees; Birkelund Index, 3.4. Fig- dorso-lateral double furrows posteriorly at an angle ured by Christensen (1995, PI. 7: 3-6; Fig. 20D). of about 60 degrees; relationship between length of Figs 6-7. Fusiteuthis polonica Kongiel, 1962, holotype, guard and dorso-ventral diameter at protoconch Bochotnica, Poland, latest Maastrichtian; 6, ventral view; strongly allometric; juvenile guard very long and slen- 7, ventral view of the anterior part of the guard, approxi- der (needle-shaped). mately x6. Figured by Kongiel (1962, PI. 1: 1-3). Figs 8-10. Belemnella lanceolata (Schlotheim, 1813), Geo- logical Museum, Copenhagen, MMH 13115, Balsvik, Discussion. — The evolutionary lineage of Belemnello- Scania, Sweden, earliest Early Maastrichtian; 8, ventral camax includes, in ascending order, B. ex gr. gros- view; 9, lateral view; 10; view of the split anterior end, x2. souvrei, B. mammillatus andß. balsvikensis (see Chri- Figured by Christensen (1975a, PI. 12: 4). stensen 1986). The general trends in evolution are the gradual calcification of the adorai end and the shape All specimens are coated with ammonium chloride, except changes in which the guard becomes more slender and Figs 6-7, and are natural size unless otherwise stated. less lanceolate. The genus is usually not granulated,

Christensen: Late Cretaceous Belemnitellidae 71 but a few granulated specimens of B. ex gr. grossouvrei mens from northwest Germany. Thus, the geographi- and B. mammillatus have been recorded (Christensen cal distribution of the genus was gradually reduced 1986). during its stratigraphical range. Naidin (1964b) placed species of the grossouvrei group in his new subgenus Actinocamax (Parac- tinocamax), type species A. grossouvrei, and mammil- Genus Goniocamax Naidin, 1964b latus in his new genus Belemnellocamax. He distinguished A. (Paractinocamax) fromBelemnellocamax Type species. -Actinocamax lundgreni Stolley, 1897, on the basis of the size of the juvenile guard and p. 285, PL 3: 16-20, non 15, by original designation claimed that A. (Paractinocamax) has a short juve- of Naidin (1964b: 104). nile guard, while Belemnellocamax has a long and slender juvenile guard. Christensen (1986) noted, how- Emended diagnosis. - Small to medium-sized (up to ever, that specimens of the grossouvrei group from 80 mm long) non-granulated belemnitellids, with a west Europe have a very elongated juvenile guard. shallow to relatively deep pseudoalveolus; guard He therefore placed the grossouvrei group in Belem- markedly flattened ventrally, lanceolate in ventral view nellocamax, and A. (Paractinocamax) was considered and subcylindrical in lateral view; juvenile guard short a junior synonym of Belemnellocamax. Christensen and stout; guard with dorso-lateral longitudinal de- (1991) figured two juvenile specimens of B. ex gr. pressions and double furrows, vascular imprints, and grossouvrei from southern England, both of which are longitudinal striae; longitudinal striae usually more very elongated. distinct than vascular markings; Schatzky distance The grossouvrei group is rare; about 60 specimens small, 2 to 4 mm; bottom of ventral fissure commonly are known from west Europe, and about 60-70 speci- straight or slightly curved forming a medium-sized mens are recorded from the Russian Platform. The angle, about 30 to 50 degrees, with wall of pseudo- specimens from west Europe have been assigned to alveolus; vascular imprints branch off dorso-lateral eight species and varieties: grossouvrei, toucasi (Janet, double furrows posteriorly at an angle less than 30 1891), alfridi (Janet, 1891), depressus (Andreae, degrees; allometric relationship of length of guard and 1895), depressus var. fusiformis (Andreae, 1895), dorso-ventral diameter at protoconch; adults stouter mammillatus var. germanica (Stolley, 1930), mam- than juveniles. millatusvar. ornatus (Moberg, 1885) and blackmorei (Crick, 1907). Discussion. - Christensen & Schulz (in press) raised Naidin (1964b) distinguished five subspecies of B. the subgenus Gonioteuthis (Goniocamax) to full ge- grossouvrei: depressus, toucasi and alfridi from west neric rank and included the type species and its close Europe, and two new subspecies, pseudotoucasi and allies in Goniocamax, that is G. birkelundae Chri- pseudoalfridi, from the Russian Platform. In addition, stensen & Schulz (in press), G. striatus Christensen Nikitin (1958) established B. toucasi var. seimensis & Schulz (in press), G. esseniensis (Christensen, 1982) and Glazunova (1972) recorded B. cf. toucasi and B. and G. mirabilis (Arkhangelsky, 1912). alfridi from the Russian Platform. Goniocamax is closely similar to Gonioteuthis, but To sum up, eleven taxa have been established within differs from that genus by being non-granulated, flat- the grossouvrei group on the basis of relatively little tened ventrally, lanceolate in ventral view, and the material. Thus, the group has been the subject of ex- bottom of the ventral fissure is usually straight or cessive subdivision by previous authors. The group is slightly curved forming a medium-sized angle with currently being revised by W. K. Christensen and M.- the wall of the pseudoalveolus. G. Schulz, Kiel on the basis of material from west Europe. Distribution. - Goniocamax occurs mainly in the Cen- tral Russian Subprovince and Balto-Scandia. It oc- Distribution. -The genus appears probably at the base curs very rarely in the Central European Subprovince. of the Santonian and continues into the early Late It appears at the base of the Coniacian and continues Campanian (Figs 1-2). The Santonian-Early Cam- into the Early Santonian (Christensen & Schulz, in panian B. ex gr. grossouvrei is widely distributed but press) (Fig. 1). The stratigraphical ranges of most of rare in the North European Province; it occurs from the species are shown in Figure 2. Scania in southern Sweden in the north to the Corbi- ères in the French Pyrénées in the south. The latest Early Campanian B. mammillatus is extremely com- Genus Belemnitella d'Orbigny, 1840 mon in Scania, but rare outside this area; a little over [ICZN 1985, Opinion 1328; name no. 2269] 100 specimens are recorded from northern Germany, Poland and the eastern part of the Russian Platform Type species. - Belemnites mucronatus Schlotheim, (Christensen 1975a). The early Late Campanian B. 1813, p. Ill, by subsequent designation by Herr- balsvikensis is also extremely common in Scania, but mannsen (1846: 105); ICZN Opinion 1328 (1985); outside this area it is unknown except for two speci- name no. 2979.

72 Bulletin of the Geological Society of Denmark Species of Belemnitella, NW Europe

Fig. 5. Stratigraphical range and inferred phylogeny of Santonian-Maastrichtian species of Belemnitella. Modified from Christensen (1995). Stage abbreviations after Harland et al. (1989). Ages in Ma after Obradovich (1994).

Diagnosis. - Small to large belemnitellids (length from ceptable to calculate the mean value of the Birkelund apex to protoconch up to 80 mm) with a deep alveo- Index in species of Belemnitella, because the relation- lus; anterior end of guard completely calcified and ship of the length from the apex to the protoconch prolonged ventrally along ventral fissure in a tongue- versus the dorso-ventral diameter is generally isomet- like extension; well developed dorso-lateral longitu- ric (Christensen 1995). dinal depressions and straight double furrows, in ad- More than two dozen species, subspecies, and vari- dition to single lateral furrows; longitudinal striae may eties of Belemnitella from the Late Campanian and be present; vascular imprints branch off double fur- Early Maastrichtian have been established. Many of rows posteriorly at an angle less than 30 degrees; ju- these are poorly understood, because they were com- venile guard short and stout; alveolar angle 17 to 24 monly erected on the basis of relatively little mate- degrees, Schatzky distance long, copmmonly larger rial, and the variation of the critical characters was than 4 mm; relationship of length from apex to rarely studied (Christensen 1988,1993a). protoconch and dorso-ventral diameter at protoconch Christensen & Schulz (in press) established Belem- generally isometric. nitella schmidi, which appears at the base of the Santonian. They considered this species as the earli- Discussion. - Christensen (1995) introduced a classi- est member of Belemnitella. It is a transitional form fication of size ranges of species of Belemnitella based possessing characters in common with the genera on the length from the apex to the protoconch. This is Goniocamax and Belemnitella. It probably evolved as follows: (1) guard small; length from apex to from Goniocamax lundgreni by allopatric speciation. protoconch less than 55 mm; (2) guard large; length B. propinqua (Moberg, 1885) enters some way above from apex to protoconch between 55 and 65 mm; (3) the base of the Santonian and is probably the lineal guard very large; length from apex to protoconch descendent of B. schmidi (Christensen & Schulz, in larger than 65 mm. He also introduced a classifica- press). B. propinqua was considered previously to be tion of the relative length of species of Belemnitella the earliest member of Belemnitella by Jeletzky based on the Birkelund Index. This is as follows: (1) (1949b, 1955), Naidin (1964a, 1964b, 1974) and Chri- guard stout; mean value of Birkelund Index less than stensen (1971, 1973, 1991), although some authors, 4; (2) guard slender; mean value of Birkelund Index 4 including Birkelund (1957) and Glazunova (1972), to 5; (3) guard very slender; mean value of Birkelund placed it in Actinocamax. Index larger than 5. It should be stressed that it is ac- Christensen (1995) analyzed the evolutionary trends

Christensen: Late Cretaceous Belemnitellidae 73 of Belemnitella from the Early Santonian to the Late Distribution. - Belemnitella as interpreted here ap- Maastrichtian on the basis of biométrie analyses of pears at the base of the Santonian and continues to the 43 samples, representing 20 species and subspecies. top of the Maastrichtian (Fig. 1). It is recordedfrom He tentatively recognized two lineages of Belemnitella the North European and North American Provinces, (Fig. 5): in addition to the northern part of the Tethyan Realm. The stratigraphical ranges of the species are shown in (1) A Santonian-Maastrichtian lineage, which is de- Figures 2-5, rived from Goniocamax lundgreni. It includes B. schmidi, B. propinqua, B, praecursor, B. woodi Christensen, 1995, B. minor I Jeletzky, 1951, B. Genus Belemmnella Nowak, 1913 minor II Christensen, 1995, B. minor III Chri- stensen, 1995, B. ex gr. junior sensu Keutgen & [ICZN 1985, Opinion 1328; name no. 2270] van der Tuuk, 1990 and B. junior. These species are large to very large and stout to slender. In ad- Type species. -Belemmnites lanceolatus Schlotheim, dition, the fissure angle is small to medium-sized, 1813, p. Ill, by subsequent designation by von the bottom of the ventral fissure is generally Bülow-Trümmer (1920: 195); ICZN Opinion 1328 straight and the Schatzky distance is medium-sized (1985); name no. 2980. to large. Diagnosis. - Large belemnitellids (length from apex (2) A late Late Campanian-Early Maastrichtian line- to protoconch up to 110 mm) with a deep alveolus; age, which comprises the late Late Campanian B. anterior part of guard complete calcified and prolonged langei Jeletzky, 1948 andß. najdini Kongiel, 1962, ventrally around ventral fissure in a tongue-like ex- in addition to the Early Maastrichtian B. pulchra tension; guard markedly flattened ventrally; well-de- Schulz, 1982. These species are small and slender veloped dorso-lateral longitudinal depressions and to very slender, Moreover, the fissure angle is large double furrows, which undulate posteriorly, in addi- to very large, the bottom of the ventral fissure is tion to single lateral furrows; vascular imprints weakly irregular and the Schatzky distance is small. The developed or not present in early forms, younger forms origin of this lineage is unknown. with distinct vascular imprints; vascular imprints In addition to the species of the two lineages, other branch off dorso-lateral double furrows posteriorly at species are recorded from the Campanian and basal an angle exceeding 30 degrees; juvenile guard long Maastrichtian. The early Early Campanian B. alpha and slender (needle-shaped); alveolar angle small, 10 Naidin, 1964a is probably an off-shoot derived from to 21 degrees; Schatzky distance short, 0 to 4.5 mm, B. praecursor (see Christensen 1995). Species of un- commonly less than 4 mm; relationship of length from known origin or poorly known species include the late apex to protoconch and dorso-lateral diameter at Late Campanian B. aff. langei of Christensen (1986, protoconch allometric; adult specimens stouter than 1993a), B. langei sensu Schulz (1978), B. cf. najdini juvenile specimens. of Schulz (1978) and B. pauli Christensen, 1995, in addition to the basal Maastrichtian B. minor sensu Discussion. - Three subgenera have been established, Christensen (1975a) and B. langei sensu Birkelund the nominotypical subgenus and B. (Pachybelemnella) (1957). The latter was placed in synonymy with B. Schulz, 1979, which are Early Maastrichtian (Schulz minor III with a query by Christensen (1995), who 1979), in addition to the Late Maastrichtian B. (Nèo- also discussed the species mentioned above. belemnella) Naidin, 1975. These subgenera include The little known late Late Campanian Belemnitella only large species in contrast to the genus Belemnitel- hoeferi (Schloenbach, 1867) from the Northern Cal- la (see above). \ ^ careous Alps of Austria is a valid species, which be- Schulz (1979) studied the Early Maastrichtian sub- longs to the 2?. muewnata group of Christensen (1995). genera in great detail and stressed the importance of Christensen (submitted) redescribed this species, in- the shape of the guard in ventral view compared with cluding biométrie analysis, on the basis of material the length from the apex to the protoconch. Since from the Gschliefgraben area in Austria. It can be dis- growth is allometric in Belemnella he introduced the tinguished from most species of the B. muewnata derived variable 'standardized length from the apex group on the basis of the slender guard (mean Birke- to the protoconch' in order to compare specimens of lund Index about 4.5). Belemnitella sp. from the 'Craie different size. The calculation of this variable is rather marneuse' of Chartreuse in the Sub-Alpine Chain, Sa- laborious. The shape of the guard in ventral view is voie, southeast France (Combémorel 1996) is closely defined by an index, which includes three different similar to B. hoeferi with respect to size, shape and measurements of the lateral diameter. slenderness of the guard, in addition to surface mark- Schulz distinguished two Early Maastrichtian sub- ings. Thus, the two taxa may be conspecific, but a genera of Belemnella: the slenderB. (Belemnella) and specific determination of the specimens from Char- the stout B. (Pachy belemnella). treuse is not possible, because the internal characters are unknown.

74 Bulletin of the Geological Society of Denmark Subgenus Belemnella (Belemnella) Nowak, 1913 (Arkhangelsky, 1912) (oldest), pseudobtusa Schulz, 1979, obtusa, sumensis Jeletzky, 1949a and cimbrica Diagnosis. - Belemnella with a slender guard; spe- Birkelund, 1957 (youngest) (Fig. 2), which is derived cies with a lanceolate guard very slender; species with fromß. (B.) lanceolata. The general trends in this lin- cylindrical to cone-shaped guard slender. eage are (1) the decreasing length from the apex to the protoconch, (2) the shape of the guard in ventral Discussion. - Schulz (1979) recognized two, possi- view changes from lanceolate to cylindrical, (3) the bly three, evolutionary lineages within Belemnella increasing Schatzky Distance and (4) the increasing (Belemnella). A lineage including lanceolata (oldest), alveolar angle. Another lineage includes infläta and gracilis (Arkhangelsky, 1912) and fastigata Schulz, desnensis (Jeletzky, 1941) (Fig. 2). The latter species 1979 (youngest) (Fig. 2). The general trends of this has a guard which is strongly lanceolate in ventral lineage are: (1) the decreasing length from the apex view. to the protoconch, (2) the shape of the guard in ventral view changes from lanceolate to cylindrical, (3) Distribution. — Belemnella (Pachybelemnella) is the increasing Schatzky Distance and (4) the increas- distributed in the Early Maastrichtian (Fig. 1) and ing alveolar angle. Another lineage includes lanceo- occurs in the North European Province and northern lata and longissima Schulz, 1979 (Fig. 2). The latter part of the Tethyan Realm. The stratigraphical ranges taxon is characterized by its very large length from of the species are shown in Figure 2. the apex to the protoconch and its very lanceolate shape in ventral view. Schulz (1979) placedß. praearkhangelskii Naidin, Subgenus Belemnella (Neobelemnella) Naidin, 1975 1964a in subgenus Belemnella with a query (Fig. 2). This species occurs only at a very restricted horizon Type species. - By monotypy Belemnitella kazimiro- at Hemmoor and Kronsmoor, that is in the middle part viensis Skolosdröwna, 1932, p. 117. of the sumensis Zone, 35-38 m above the base of the Maastrichtian. He also recorded four unhorizoned Diagnosis. - Belemnella with a large alveolar angle, specimens of this taxon from Møns Klint, Denmark, 20 to 23 degrees; large Schatzky Distance, 3 to 5 mm; and Rügen, Germany, which may have come from the well developed vascular markings; a short and cone- same zone. Keutgen in Jagt et al. (1995) has subse- shaped juvenile guard; vascular imprints branch off quently reported the species from the middle part of dorso-lateral double furrows posteriorly at an angle the sumensis Zone of northeast Belgium. Schulz less than 40 degrees. (1979) suggested that B. praearkhangelskii probably belongs to another evolutionary lineage. Discussion. -Jeletzky (1951) spelled the species name B. (B.) lanceolata has been widely used as a zonal casimirovensis, as did Birkelund (1957) and some later index fossil of the early Early Maastrichtian (see dis- authors. According to the International Code of Zoo- cussion by Christensen 1996). Schulz' (1979) con- logical Nomenclature, Article 33c, 1985, this is, how- cept of this species is more restricted, however, than ever, an incorrect subsequent spelling. earlier authors, including Jeletzky (1951), Naidin The subgenus B. (Neobelemnella) has several char- (1952) and Birkelund (1957). He recorded B. (B.) acters which otherwise are typical for the genus lanceolata only from the earliest Early Maastrichtian. Belemnitella, namely a short and cone-shaped juvenile guard and well developed vascular imprints, Distribution. - This subgenus is distributed in the Moreover, the Schatzky Distance is relatively large. Early Maastrichtian (Fig. 1) and occurs in the North Naidin (1975) placed the following taxa in syn- European Province and northern part of the Tethyan onymy with B. (N.) kazimiroviensis: Belemnitella Realm. The stratigraphical ranges of the species are americana Arkhangelsky, 1912, Belemnitella ark- shown in Figure 2. hangelskii (Naidin, 1952), Belemnitella pensaensis (Kongiel, 1962) and Belemnitella skolosdrownae (Kongiel, 1962). Subgenus Belemnella (Pachybelemnella) Schulz, 1979 Naidin (1975) suggested a late Early Maastrichtian- Late Maastrichtian evolutionary lineage, which in- Type species. - B. (P.) obtusa Schulz, 1979, p. 106, cludes, in ascending order, Belemnella sumensis PI. 9: 1-8, PI. 12: 9, by original designation. sumensis Naidin 1964a, B. sumensis postsumensis Naidin, 1964a, B. sumensis praearkhangelskiiNaidin, Diagnosis. - Belemnella with a stout guard; species 1964a, B. sumensis kajnarensis Naidin, 1964a and with a very lanceolate guard slender; species with a Belemnella (Neobelemnella) kazimiroviensis. This lin- lanceolate or cylindrical guard stout. eage bridges the Early and Late Maastrichtian boundary in the eastern part of the Russian Platform. Ac- Discussion. - Schulz (1979) distinguished two line- cording to Schulz (1979), however, B. sumensis Nai- ages within this subgenus. A lineage including infläta din, 1964a is not conspecific withZ?. sumensis Jeletzky,

Christensen: Late Cretaceous Belemnitellidae 75 500 km

Central Russian Central European Boundary between North European Subprovince Subprovince Province and Tethyan Realm Fig. 6. Map showing the Central European and Central Russian palaeobiogeographical Subprovinces within the North European Province as defined on belemnites. A third subprovince, the Balto-Scandian Subprovince, is recognized during the Early Coniacian and latest Early Campanian-early Late Campanian. It is characterized by Goniocamax lundgreni in the Early Coniacian and the genus Belemnellocamax in the latest Early Campanian and early Late Campanian. Late Cretaceous land and sea areas represent maximum inundation for all stages. The boundaries are not reliable in detail and the biogeographic units are typically gradational in character. After Christensen (1976).

1949a, and he placed B. s. sumensis and B. sumensis J. A. Jeletzky (unpublished MS 1972) has suggested postsumensis in synonymy withß. (B. ?)praearkhan- that the genus is dubious being based on a possibly gelskii. pathological specimen. I agree.

Distribution. — This subgenus is restricted to the Late Distribution. - This genus is restricted to the latest Maastrichtian (Fig. 1) and occurs in the North Euro- Maastrichtian and occurs in Poland and the Crimea. pean Province and northern part of the Tethyan Realm. B. (N.) kazimiroviensis occurs in the Late Maastrich- tian in the eastern part of the Russian Platform, late Late Maastrichtian in Denmark and latest Late Maas- trichtian in The Netherlands (Christensen 1996). Palaeobiogeography The belemnitellids were distributed in the North American and North European Provinces of the North Genus Fusiteuthis Kongiel, 1962 Temperate Realm, in addition to the northern Euro- pean margin of the Tethyan Realm (Christensen 1976, Type species. - By monotypy F. polonica Kongiel, 1988, 1993b). 1962, p. 28, PI. 1:1-3.

Diagnosis. — Medium-sized belemnitellids with a shallow alveolus; guard lanceolate in ventral and lateral North European Province views; dorso-lateral longitudinal depressions present, This province extends from Ireland in the west to the guard otherwise smooth. Ural Mountains and beyond in the east (Fig. 6). The centre of origin of the belemnitellids lay there for the Discussion. - F. polonica was established on the ba- following reasons. They are common and all known sis of a single specimen from the latest Maastrich- genera and subgenera occur there, the earliest species tian of central Poland (PL 2:6-7). Later, Naidin (1973, of the family, Praeactinocamax primus, appears in the 1975) recorded two specimens of F. sp. from the Late Early Cenomanian, some way above the base the sub- Maastrichtian of Crimea. stage, in this province and the earliest species of the

76 Bulletin of the Geological Society of Denmark Table 2. The occurrence of belemnitellid genera and subgenera in the North European and North American Provinces of the North Temperate Realm and the Tethyan Realm. Fusiteuthis is most likely a nomen dubium. The genera and subgenera are ranked with respect to their palaeobiogeographical distribution. Nine genera and two subgenera occur in the North European Province, five genera and two subgenera in the Tethyan Realm in Europe and essentially two genera in the North American Province. The numbers 1-12 refer to the first appearances of the genera and subgenera in the various palaeobiogeographical units. Detailed data are not available for the subgenera of Belemnella. 1, Early Cenomanian; 2, Late Cenomanian; 3, Middle Turonian; 4, Early Santonian; 5, latest Santonian; 6, latest Santonian-earliest Campanian; 7, Middle Coniacian; 8, Early Campanian; 9, Early Santonian; 10, Late Santonian; 11, Early Turonian; 12, latest Santonian- earliest Campanian.

Genera and subgenera North European Province Tethyan Realm North American Province

Praeactinocamax +' Belemnitella +4 Belemnella (Belemnella) + Belemnella (Pachybelemnella) + Belemnella (Neobelemnella) + Gonioteuthis +7 Belemnellocamax +9 Actinocamax +" (+)12 Goniocamax + Belemnocamax + Fusiteuthis +

other genera either occurs only there or appears ear- nitella mucronata and comprises about 95% of the lier there than elsewhere (Table 2). The last belemnitel- belemnite fauna (Christensen 1976). lids became extinct at the Maastrichtian-Danian At other times during the Late Cretaceous the sub- boundary. In addition to the belemnitellids, the gen- provinces are less distinct, and no subprovinces can era Neohibolites and Parahibolites of the family be recognized in the later part of the Late Campanian Belemnopseidae also occur in this province. and Maastrichtian. The North European Province includes the Central European, Central Russian and Balto-Scandian Sub- provinces (Fig. 6). The Central Russian and Central European Subprovinces are well-defined from the Tethyan Realm Middle Coniacian to the boundary between the Early Belemmnitellids occur sporadically at the northern and Late Campanian, a period of around 7 Ma. The European margin of the Tethyan Realm (Fig. 4). They two subprovinces are characterized by two independ- have been recorded from the Late Cenomanian of the ently evolving belemnite lineages: the Gonioteuthis Vocontian Basin in southeastern France (Gale & Chri- stock inhabited the Central European Subprovince, and stensen 1996), the Late Santonian-basal Early Campa- the Goniocamax-Belemnitella stock inhabited the Cen- nian of the Corbières in the French Pyrénées (Chri- tral Russian Subprovince (Christensen 1975a, 1976, stensen, Bilotte & Melchior 1990, Christensen, Bilotte 1988, 1990b, Christensen & Schulz in press). & Hansotte 1993), the Late Campanian of Bulgaria The belemnite faunas of Balto-Scandia show affin- (Stoyanova-Vergilova & Jolkicev 1993) and Roma- ity to those of the Central Russian Subprovince at cer- nia (Neagu & Georgescu 1991), the late Late Campa- tain times and to those of the Central European Sub- nian of the Northern Calcareous Alps of Austria (Chri- province at other times. However, in the Early Conia- stensen, submitted) and probably the Sub-Alpine cian and latest Early Campanian and early Late Cam- Chain at Chartreuse in Savoie, southeast France (Com- panian a third subprovince can be recognized, named bémorel 1996), the Early Maastrichtian of Bavaria in here the Balto-Scandian Subprovince. It is character- southern Germany (Schmid & Schulz 1979, Schulz ized by Goniocamax lundgreni in the Early Conia- & Schmid 1983a), in addition to the Campanian and cian, Belemnellocamax mammillatus in the latest Early Maastrichtian of Azerbaijan (Ali-Zade 1972). Campanian andß. balsvikensis in the early Late Cam- There are previous records of the Early Campanian panian. B. mammillatus co-occurs with Belemnitella Gonioteuthis quadrata and the Late Campanian Be- mucronata, Gonioteuthis quadrata scaniensis and lemnitella mucronata from the Aquitaine Basin (sum- Belemnellocamax ex gr. grossouvrei and comprises marized by Séronie-Vivien 1972), but these records about 90-95% of the belemnite fauna (Christensen need to be confirmed, as do previous records of B. 1975a, Table 3). B. balsvikensis co-occurs wiihBelem- mucronata from Greece and Turkey.

Christensen: Late Cretaceous Belemnitellidae 77 generally very rare and belemnopseids do not occur. The belemnite faunas are essentially restricted to species of Praeactinocamax (Turonian to Early San- tonian) and Belemnitella (uppermost Santonian to Maastrichtian) (Fig. 3). Actinocamax is represented only by a single specimen of A. verusl from central East Greenland (Jeletzky in Donovan 1954) and two specimens of A. aff. laevigatus from Kansas (Jeletzky 1961). However, Jeletzky (1961) suggested that the two specimens from Kansas may be juveniles of one of the large species of Praeactinocamax. The belemnite fauna of the middle Turonian seems to be rather diverse, but this may due to excessive subdivision (see above). The earliest belemnitellids from this province are Middle Turonian in age. The belemnitellids of the North American Province were discussed by Chri- stensen (1993b). Seibertz & Spaeth (1995) recorded three fragmen- tary belemnites from the Early Turonian of northern Mexico. The anterior end with the critical characters is missing in these specimens, but, nevertheless, Sei- bertz & Spaeth tentatively assigned them to Praeacti- Fig. 7. Approximate distribution of land and sea in North nocamax cf. manitobensis. P. manitobensis occurs in America and Greenland during Turonian and Coniacian the Western Interior of North America and has not time. The distribution of land and sea is based on Williams been recorded previously south of Kansas (Cobban & Stelck (1975) for North America and Christensen (1993 1991). If the specimens from northern Mexico are b) for Greenland. correctly identified, which is open to discussion, then they are the most southerly belemnitellids recorded to date. Northern Mexico lay at palaeolatitude 15- 20°N. B. mucronata was recorded also from Austria, but B. americana (Morton, 1830) occurs in the Mount this may be a misconception. B. hoeferi occurs in the Laurel Formation and basal part of the Navesink For- Northern Calcareous Alps (Schloenbach 1867, Chri- mation of the Atlantic Coastal Plain (Owens & Sohl stensen, submitted; see above). 1973, Fig. 4). In ammonite terms, the Mount Laurel Species of the following genera and subgenera are Formation is late Late Campanian, but not latest recorded from the Tethyan Realm: Praeactinocamax, Campanian in age (Kennedy & Cobban 1994), and Gonioteuthis, Belemnellocamax, Belemnitella, Belem- the basal part of the Navesink Formation is latest nella (Belemnella), B. (Pachybelemnella) andß. (Neo- Campanian and Early Maastrichtian in age (Kennedy, belemnella) (Fig. 4). The majority of the species oc- Johnson & Cobban 1995). curring in the Tethyan Realm are conspecific with B. bulbosa occurs in the Fox Hills Formation of those from the North European Province and they thus South Dakota, which is probably early Late Maastrich- provide a basis for correlation. tian in age (Waage 1968, Landman & Waage 1993). Local Late Campanian species of Belemnitella oc- A small species of Belemnitella, probably conspecific cur together with species of Belemnitella from the with B. bulbosa, occurs in the upper part of the un- North European Province in Azerbaijan, Bulgaria and derlying Pierre Shale, Bacutites baculus andß. clino- Romania (Ali-Zade 1972, Neagu & Georgescu 1991, lobatus Zones. These zones are Early but not earliest Stoyanova-Vergilova & Jolkicev, 1993). Maastrichtian in age (Kennedy & Cobban 1993). The middle Late Cenomanian Praeactinocamax The majority of the belemnitellids of the North Ame- plenus is the earliest belemnitellid recorded from the rican Province are endemic, have a punctuated, strong- Tethyan Realm (Gale & Christensen 1996) and belem- ly discontinuous stratigraphical distribution and are nitellids are not recorded from the Turonian, Coniacian derived from species from the North European Prov- and Early and Middle Santonian. ince (Christensen 1993b). The endemic taxa include the Middle Turonian-Early Santonian species of Prae- actinocamax and the Late Campanian-Maastrichtian species of Belemnitella. These taxa evolved from Eu- North American Province ropean species by allopatric speciation. Three Euro- This province includes Greenland, the Western Inte- pean taxa, Actinocamax verusl, Belemnitella praecur- rior of North America, in addition to the Atlantic and sor and B. ex gr. alpha/praecursor, occur in the up- Gulf coasts of the USA (Fig. 7). Belemnitellids are permost Santonian-basal Early Campanian.

78 Bulletin of the Geological Society of Denmark Since most of the belemnitellid species of the North the Cenomanian-Turonian boundary and declined ir- American Province are endemic intercontinental cor- regularly thereafter. It was about 22°C in the Coniacian relation based on belemnites is not possible. and dropped to around 19°C in the Late Santonian. Following Schönfeld & Schulz the palaeotemperature was about 15°C in the early Late Campanian, increased to about 17°C in the earliest Maastrichtian and decreased to around 15°C in the middle Early Maastrich- tian. Mode of life of Late Cretaceous belemnites The South Temperate dimitobelids were also steno- The belemnitellids were neritic animals restricted to thermal shallow-water dwellers (Doyle & Howlett the shelf. They are common in nearshore sediments, 1989), but they appear to be adapted to life in cooler such as biocalcarenites, greensands, marls and shal- waters than the belemnitellids. Pirrie & Marshall low-water chalks, and populations from these sedi- (1990) and Crame, Lomas, Pirrie & Luther (1996) mentary deposits characteristically contain all growth suggested that the maximum surface water tempera- stages. They are less common to virtually absent in ture was about 20°C in the Turonian-Coniacian and deeper water chalks, and populations from these de- declined thereafter. It was about 14°C in the Santonian posits consist mainly of adult specimens (Christen- and Campanian and about 12°C in the Maastrichtian sen 1976). W.J. Kennedy (personal communication, of James Ross Island, Antarctica (palaeolatitude 60- November 1996) noted that he has never seen a single 65°S). specimen in cores from the Central Graben, of which Since both the belemnitellids and dimitobelids were he has examined several kilometres. It therefore ap- shallow-water dwellers, oceans with deep water may pears that the breeding, spawning, hatching, and, pos- have acted as physical barriers and precluded spread sibly for the females at least, dying grounds, were in- of these families. Warm tropical waters in the Tethyan ner neritic, shallow water environments. Adult speci- Realm may also have acted as a physical barrier for mens from offshore, deeper water chalks may be con- these families. sidered as stray fully-grown individuals that died out- The belemnopseids Neohibolites and Parahibolites side their normal habitat (Christensen 1976, Surlyk were widely distributed in the Albian-Cenomanian, & Birkelund 1977). from the Central European Subprovince of the North It is noteworthy that belemnitellids are absent or Temperate Province, across the Tethyan Realm to the extremely rare at some horizons in the Late Campanian South Temperate Province, and have been recorded and Maastrichtian chalks of northwest Europe. At from deep marine and continental slope deposits, as Kronsmoor in northwest Germany, the topmost five well as inner neritic sediments. The habitat of these metres of the Late Campanian have not yielded belem- genera may therefore have been surface oceanic, but nites, and Belemnella and Belemnitella are extremely they probably spent their breeding season inshore rare in the basal three metres of the Maastrichtian (Doyle & Howlett 1989). They were eurythermal, (Schulz 1978,1979,1982). At Hemmoor in northwest adapted to life in warm temperate as well as tropical Germany belemnites are virtually absent in the top- waters. most five metres of the late Early Maastrichtian (only one specimen of Belemnella fastigata is recorded) and they are not recorded from the basal five metres of the early Late Maastrichtian (Schulz and Schmid 1983 b). They are virtually absent in the early Late Maa- Belemnitellid distribution strichtian of Denmark (Surlyk 1970); only two speci- Praeactinocamax and Belemnitella are more widely mens of Belemnella cf. cimbrica are recorded (Schulz distributed than the remaining genera and subgenera and Schmid 1983b). The topmost six metres of the and occur in both the North European and North Ame- chalk of Hemmoor, which equates with the basal part rican Provinces, as well as in northern European part of the late Late Maastrichtian Belemnella kazimiro- of the Tethyan Realm (Table 2). Gonioteuthis, Belem- viensis Zone of Denmark, have not yielded belemnites nellocamax and Belemnella are distributed in both the (Schulz and Schmid 1983b). North European Province and the Tethyan Realm. The belemnitellids were stenothermal animals Actinocamax, Belemnocamax and Goniocamax occurs adapted to life in warm-temperate waters according only in the North European Province, except for a sin- to the palaeotemperature curves by Jenkyns, Gale & gle specimen of A. verusi from East Greenland (see Corfield (1994, Fig. 12; Cenomanian to Santonian of above), but they have different distribution patterns. east Kent) and Schönfeld & Schulz (1996, Fig. 4; Late Actinocamax is widely distributed in the North Euro- Campanian to Early Maastrichtian of northern Ger- pean Province, Goniocamax is mainly restricted to the many). These areas were situated at about palaeola- Central Russian Subprovince and Balto-Scandia and titude 40°N. According to Jenkyns et al. the palaeotem- Belemnocamax occurs only in northwest Germany and perature was about 21 °C in the basal Cenomanian, eastern England. Thus, the genera and subgenera can increased gradually to a maximum of about 28°C at be ranked in the following way with respect to their

Christensen: Late Cretaceous Belemnitellidae 79 ZONAL Fig. 8. Late Cretaceous STA- ZONAL BELEMNITES, BELEMNITE ZONES, BELEMNITES, belemnite stratigraphy of GES RUSSIAN PUTFORM NW EUROPE BALTO-SCANDIA northwest Europe, Balto- Scandia and the Russian S/a kazimiroviensis Bln. kazimiroviensis Platform. In order to ease Bit. junior Bit. junior (W) reading of the diagram the following abbreviations are Bln. fastigata used for the genera. X Bln. cimbriça P. = Praeactinocamax, en Bln. sumensis B. sumensis A. = Actinocamax; Bit. = Belemnitella; Bln. obtusa B. lanceolata Bin. = Belemnella; Bln. pseudobtusa OQ Blc. = Belemnellocamax; B. licharewi Bln. lanceolata Bln. lanceolata Gt. = Gonioteuthis; Gc. = Goniocamax; B.l. najdini 'Bit. langei] N. = Neohibolites. Vertical B.l. langei axis not to scale. Modified from Christensen (1986). S- B/f. minor] g B.I. minor x LU , Bit. mucronata a. L Bit. mucronata Bit. mucronata Q. Blc. balsvikensis/ Bit. mucronata Gt.q. gracilis/ Blc. mammillatus/ Bit. mucronata/ Bit. mucronata/ Gt. q. gracilis/ Bit. mucronata Gt. q. scaniensis Blc. mammillatus Gt. q. gracilis Bit. alpha/Bit, praecursor/ O it.q. quadrata Gt. q. quadrata ce Bit. praecursor/A. laevigatus/ Gt. granulataquadrata/ Gt. granulataquadrata Gt. granulataquadrata Bit. alpha CPteria-beds')

Bit. praecursor/ Gt. granulata Gt. granulata Gt. granulata

< Gt. westfalicagranulata/ ïf. westfalicagranulata P M Bit, propinqua U Bit. propinqua Gt. westfalica Gt. westfalica/ Bit. propinqua

Gonioteuthis Gc. lundgreni/ Gc. lundgreni uilicus extremely rare Gt. praewestfalica

Gt praewestfalica Gc. lundgreni M Gc. lundgreni

g M ce P. plenus triangulus

P. plenus P. plenus P. plenus

M P. primus (NE) P. primus P. primus LU Ü

N. ultimus N. ultimus (SW)

80 Bulletin of the Geological Society of Denmark palaeogeographical distribution, from the largest to invade the Tethyan Realm again until the Late Santo- the smallest areal extent (Table 2): nian. The next extinction event occurred in the latest Early (1) Praeactinocamax and Belemnitella; (2) Gonio- Campanian and early Late Campanian. Three genera teuthis, Belemnellocamax and Belemnella; (3) Acti- disappeared, Actinocamax, Gonioteuthis and Belem- nocamax; (4) Goniocamax; (5) Belemnocamax. nellocamax, and only one genus, Belemnitella, survived. After this extinction, Belemnitella expanded The palaeogeographical distribution of the belemnitel- its area of distribution to cover the entire North Euro- lids was controlled by a number of factors, including pean Province, where it is common. It also invaded sea-level changes, temperature, palaeocurrents, eco- the Tethyan Realm. However, only one species, B. logical tolerance and competition. Gale & Christensen mucronata, occurs in the lower part of the early Late (1996) suggested that the southwards migration of Campanian (Fig. 2), that is a period of about 1.5 Ma. Praeactinocamax plenus into the Tethyan Realm in The last extinction occurred at the Cretaceous- the Late Cenomanian was due to a significant fall of Palaeocene boundary and no belemnites survived. sea temperature, the so-called Plenus Cold Event, Paleogene belemnites have been recorded earlier. working in concert with the development of suitable However, the Eocene Bayanoteuthis rugifer Schloen- shallow water habitats. In addition, lack of competi- bach, 1868, which occurs in southern Europe, has been tion of the Tethyan belemnopseids, which disappeared reinterpreted as a pennetulacean coral (Riegraf 1991) in the Middle Cenomanian, may also have been of and others are considered to be sepiids with a belem- importance. Christensen (1976) suggested that the nite-like guard (Doyle, Donovan & Nixon 1994). parallel evolution of the Gonioteuthis and Gonioca- max-Belemnitella stocks from the Middle Coniacian to the boundary between the Early and Late Campanian in the adjacent Central European and Cen- Late Cretaceous belemnite biostratigraphy tral Russian Subprovinces, respectively, was due to mutual competition. The causes of the palaeogeogra- of Europe phical distribution and migrations of the belemnitellids The belemnitellids are of fundamental importance in will be discussed in more detail in a forthcoming pa- biostratigraphy and correlation of the Late Cretaceous per. of the North European Province, particularly during the Coniacian to Maastrichtian stages. They are common, widely distributed there and the fossilization potential is high (Christensen 1990b, 1996). 25 zones have been established in northwest Europe and a little Late Cretaceous belemnite extinctions less on the Russian Platform (Fig. 8). The last genera of the Tethyan belemnopseids, Parahi- The zonation of the west Europe and the Russian bolites and Neohibolites, became extinct worldwide Platform is based upon the bélemnopseidNeohibolites in the Middle Cenomanian. The last dimitobelid ultimus and species of Praeactinocamax for the Ceno- belemnites in the South Temperate Realm, Dimitobelus manian, species of Belemnitella for the Late Campa- (Dimitocamax) seymourensisDoyle, 1988 andD. (D.) nian, species of Belemnella for the Early Maastrichtian hectori Stevens, 1965, disappeared in the early and species of Belemnitella and Belemnella for the Maastrichtian (Doyle & Zinsmeister 1988, Doyle & Late Maastrichtian. In the Middle Coniacian to Early Howlett 1989, Crame, Lomas, Pirrie & Luther 1996). Campanian the zonation of northwest Europe is based The belemnitellids suffered three extinctions dur- upon species of Gonioteuthis, while it is based upon ing the Late Cretaceous: (1) in the Middle Cenoma- species of Goniocamax and Belemnitella on the Rus- nian-earliest Early Turonian, (2) around the Early and sian Platform (Fig. 8). Late Campanian boundary and (3) at the Maastrich- The Middle Coniacian to Early Campanian belem- tian-Palaeocene boundary. At the first extinction event, nite faunas of Balto-Scandia are of great importance, the belemnitellids Belemnocamax boweri and Praeac- because they include species of both the Gonioteuthis tinocamax plenus of the P. primus-plenus lineage dis- and Goniocamax-Belemnitella stocks, and thus pro- appeared along with the belemnopseids. After this vide a basis for correlation between the two subpro- extinction the belemnitellids are extremely rare in the vinces (Fig. 8). Turonian and early Coniacian of the Central European Belemmnitellids are extremely rare in the Turonian Subprovince and have a very restricted distribution in and early Coniacian of northwest Europe (Christensen the Central Russian Subprovince (Christensen 1982). 1982) and, consequently, no zones have been estab- It appears that the belemnitellids retreated into réfugia lished. Christensen (1995,1996) argued that the con- in the Turonian, which may be situated in the Central ventional late Late Campanian Belemnitella minorand Russian Subprovince. The Central European Sub- B. langet Zones of Jeletzky (1951)shouldnotbemain- province was repopulated by Gonioteuthis praewest- tained. These zones are therefore placed in square falica in the late Coniacian and belemnitellids did not brackets in Figure 8.

Christensen: Late Cretaceous Belemnitellidae 81 Belemnitella, B.langei,najdiniandexgr.langet/ The B.minorzoneswersubdividedintofivinfor- mal subzones,mainlyonthebasisfsmalspecie mucronata,B. woodi,B.minorlandB.minorIzones. nitella (Fig.9)Thesearinascendingorder:thB. Late CampanianofNorfolkintfourinformalzones For instance,Christensen(1995,1996)subdividedthe praewestfalica, G.westfalica andG.ernsti), blage zonesarapplicablonlyinBalto-Scandiafor mark intothreezones:thConiacianGoniocamax Coniacian andLowerSantoniaofBornholm,Den- nation ofNorfolkisapplicablenortheastBelgium. Keutgen (1995)hasshowsubsequentlythatthezo- najdini. Furtherstudiesarenecessarytoseeifthes on thebasisflargtveryspecieBelem- tonian GonioteuthiswestfalicaZone(Fig.10)Thes teuthis praewestfalicaZoneandthlatEarlySan- lundgreni Zone,theearlyEarlySantonianGonio- zones canbeextendedtootherareaiEurope. from theCentralRussianSubprovinc(Gonioteuthis schmidi andB.propinqua),othersarenotrecorde very rareornotrecordedfromthCentralEuropean the followingreasons.Somofindexspeciesar However, itsverylikelthathezoneandassem- zones weresubdividedintoeightassemblagzones. the CentralRussianSubprovinc e (MiddlConiacian), Subprovince (Goniocamaxlundgreni,Belemnitella occurs earlieronBornholm(Earl y Coniacian)thani corded outsideBornholm.Moreover , G.lundgreni Goniocamax birkelundaeand G.striatusarenotre- 82 1: Christensen &Schulz(ipress)subdividedthe Some ofthbelemnitzoneshavbeensubdivided. UPPER CAMPANIAN SUB- STAGES

| Belemnitella Belemnella NW Europe Belemnite zones, pseudobtusa mucronata minor langei lanceolata sumensis obtusa cimbrica fastigata Bit woodi Bit minor] BltminorW BU. mucronata Bit minorIII Belemniteila zones not exposed not exposed Norfolk Actinocamax, Belemnocamax,Gonioteuthis,Belem- I thankthejournalreferees,Prof.WJKennedyOx- Acknowledgments Central EuropeanSubprovince,whileitsearlyEarly Belemnit familienBelemnitellidaeforekommerkui Dansk sammendrag port. Geological Museum,Copenhagenfortechnicalsup- ford, andDr.PDoyleLondonforconstructivecriti- Santonian iageoBornholm. and G.praewestfalicaislateConiacianagth nellocamax, Goniocamax,Belemnitella,Belemnella følgende slægterogunderslægter:Praeactinocamax, cism ofthemanuscript.Ialsthankstaf nella (Neobelemnella)ogdendubiøseFusiteuthis. Cenomanien tilØvreMaastrichtien.Deinkluderer Øvre Kridtpådennordlighalvkugle,fraNedr ter gradvisttilengennemde t mesteafØvreCam- gradvist tilseksiNedreSantonien . Detaftagerderef- Øvre Kridt.Detrntiloi Cenomanienogstiger (Belemnella), Belemnella(Pachybelemnella),Belem- panien ogtiltagertilttrei Maastrichtien. Belemnitella subzones ex gr.langei/najdini Bit. minorWBIt.langei Bit. najdini/Bit.pauli Bit. minor\/Blt.langei/ Bit. minor\ BltminorW Bit. minorW Antallet afslægterogunderslægtevarieregennem Det evolutionærecenterfobelemnitellidern e låi Bulletin oftheGeological SocietyofDenmar k Bit. =Belemnitella.Vertical zones (Schulz1979),in Maastrichtian Belemnella Belemnitella zones correlation diagram Christensen (1996). Norfolk (Christensen1995). zones andsubzoneof addition toBelemnitella (Jeletzky 1951)andEarl Fig. 9Stratigraphical axis nottoscale.After showing LateCampanian Ernst & Schulz (1974) Schulz ef al. (1984) Chrstensen & Schulz to (In press) LU Schulz (1996) (3 < Lägerdorf, NW Germany Bornholm, Denmark te Faunal zones Gonioteuthis Zones Belemnite zones Belemnite assemblage zones rogalae/ westfalicagranulata ou westfalicagranulata (Rl: 11.0-12.5) westfalica rogalae/westfalica (Rl: 8.5-11.0) not exposed

Gt. westfalica/A. v. verus/Blt. westfalica propinqua/Gc. striatus/Gt ernsti SJIA N coranguinum/ Gonioteuthis OU westfalica (Rl < 8.5) westfalica Gt. westfalica/A. v. verus/ (Rl < 8.5) Bit. propinqua/Gc. striatus < — — _ — Gt. westfalica/Gc. lundgreni/ w Bit. propinqua LT LU pachti/ Gonioteuthis Ge. lundgreni/Gt. praewestfalica/ ^L undulatoplicatus extremely rare Gonioteuthis Bit. propinqua —i praewestfalica Ge. lundgreni/Gt. praewestfalica/ Ge. birkelundae/Blt. schmidi

bucailli/ Goniocamax lundgreni/ U praewestfalica praewestfalica Goniocamax birkelundae

involutus/bucailli Goniocamax Goniocamax lundgreni/ no belemnites lundgreni koeneni Actinocamax verus ef. antefragilis CONIACIA N not exposed Goniocamax lundgreni 1 Fig. 10. Stratigraphical correlation diagram of the Coniacian and Santonian, showing faunal and Gonioteuthis zones of Lägerdorf and belemnite zones and assemblage zones of Bornholm. Ge. = Goniocamax; Gt. = Gonioteuthis; Bit. = Belemnitella; A. = Actinocamax. Vertical axis not to scale. Modified from Christensen & Schulz (in press). den Nordeuropæiske Provins, hvor alle kendte slæg- References ter og underslægter forekommer. Den første art inden Andreae, A. 1895. Ein neuer Actinocamax aus der Quadra- for familien, Praeactinocamax primus, er fra Nedre tenkreide von Braunschweig. Mitteilungen aus dem Cenomanien og kendes kun fra denne provins. Roemer-Museum Hildesheim 2, 4 pp. Belemnitelliderne udvandrede fra den Nordeuro- Ali-Zade, A. A. 1972. Cretaceous belemnites of Azerbaijan. pæiske Provins til henholdsvis den Nordamerikanske 279 pp. Moscow. 'Nedra' Publishing House. [In Rus- Provins og Tethys Området. Arter af fem slægter og sian.] to underslægter findes i Tethys Området, og hoved- Arkhangelsky, A. D. 1912. The Upper Cretaceous depos- parten af disse er identiske med arter fra den Nord- its in the eastern part of the Europen Russia. Materialy europæiske Provins. Den tidligste art er fra Øvre dlya Geologii Rossii 25, 631 pp. [In Russian.] Cenomanien. Arter af hovedsagelig to slægter fore- Bändel, K. & Spaeth, C. 1988. Structural differences in kommer i den Nordamerikanske Provins og hoved- the ontogeny of some belemnite rostra. In Wiedmann, J. & Kullmann, J. (eds) Cephalopods - Present and Past, parten af disse er endemiske. De tidligste arter er fra 247-271. Stuttgart: Schweizerbart'sche Verlagsbuch- Mellem Turonien. handlung. Bather, F. A. 1888. Shell-growth in Cephalopoda (Siphono- poda). Annals and Magazine of Natural History (6), 1, 298-310. Bayle, E. 1878. Fossiles principaux des terrains de la Fran- ce. Explication de la Carte Géologique de la France 4(1), Atlas, 79 plates.

Christensen: Late Cretaceous Belemnitellidae 83 Birkelund, T. 1956. Upper Cretaceous belemnites from Christensen, W. K. 1993a. Upper Cretaceous belemnitellids West Greenland. Meddelelser om Grønland 137, 30 pp. from the Båstad Basin, southern Sweden. Geologiska Birkelund, T. 1957. Upper Cretaceous belemnites of Den- Föreningens i Stockholm Förhandlingar 115, 39-57. mark. Biologiske Skrifter. Det Kongelige Danske Viden- Christensen, W. K. 1993b. Actinocamax cobbanin. sp. from skabernes Selskab 9,69 pp. the Coniacian of Montana and Wyoming and the occur- Blainville, H. M. D. de 1825-1827. Manual de Malacologie rence of Late Cretaceous belemnites in North America et de Conchyliologie. 664 pp. (1825), 87 plates (1827). and Greenland. Journal of Paleontology 67, 434-446. Paris: Levrault. Christensen, W. K. 1993c. Belemnocamax boweri Crick, Blainville, H. M. 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Christensen: Late Cretaceous Belemnitellidae 85 Jeletzky, J. A. 1965. Taxonomy and phylogeny of fossil Naef, A. 1922. Die fossilen Tintenfische. 322 pp. Jena: Coleoidea (= Dibranchiata). Geological Survey of Cana- Verlag von Gustav Fischer. da, Paper 65-2, 42, 72-76. Naidin, D. P. 1952. The Upper Cretaceous belemnites of Jenkyns, H. H., Gale, A. S. & Corfield, R. M. 1994. Car- western Ukraine. Trudy Moskovskogo Geologo-Razve- bon- and oxygen-isotope stratigraphy of the English dochnogo Instituta imeni S. Ordzhinikidze 27, 170 pp. Chalk and Italian Scaglia and its palaeoclimatic signifi- [In Russian.] cance. Geological Magazine 131, 1-34. Naidin, D. P. 1953. A new belemnite from the Upper Cre- Kennedy, W. J. & Cobban, W. A. 1993. Ammonites from taceous deposits of Crimea. Byulletin Moskovskogo the Saratoga Chalk (Upper Cretaceous), Arkansas. Jour- Obschestra Ispytateli Prirody, Otdel Geologicheskii 28, nal of Paleontology 67, 404^34. 64-65. [In Russian.] Kennedy, W. J. & Cobban, W. 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86 Bulletin of the Geological Society of Denmark Nauk, St Petersburg, Zapiski Seriia 8, Fizino-mathema- Dänemark un Limburg. Newsletters in Stratigraphy 13, ticheskii 21, 68 pp. [In Russian.] 21-39. Pirrie, D. & Marshal, J. D. 1990. High-paleolatitude Late Schönfeld, J. & Schulz, M.-G. (Coordinators) 1996. New Cretaceous paleotemperatures: New data from James results on biostratigraphy, palaeomagnetism, geoche- Ross Island, Antarctica. Geology 18, 31-34. mistry and correlation from the standard section for the Riegraf, W. 1991. Triassic belemnoids (Cephalopoda, Upper Cretaceous white chalk of northern Germany Coleoidea) formerly described as corals, versus Tertiary (Lägerdorf - Kronsmoor - Hemmoor). Mitteilungen aus corals described as belemnites (or the fairy tale of Terti- dem Geologisch-Paläontolischen Institut der Universität ary "belemnites"). Fossil Cnidaria 20, 40-45. Hamburg 77, 545-575. Schloenbach, U. 1867. Ueber einen Belemniten aus der Seibertz, E. & Spaeth, C. 1995. 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