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Home , Actinocamax, Aulacoceras, Aulacocerida, Belemnella, Belemnellocamax, Belemnites (genus)

[Palaeontology, Vol. 54, Part 2, 2011, pp. 397–415]

THE CAPSULE: AN ORGANIC SKELETAL STRUCTURE IN THE LATE BELEMNITE GONIOTEUTHIS FROM NORTH-WEST GERMANY by LARISA A. DOGUZHAEVA and STEFAN BENGTSON Department of Palaeozoology, Swedish Museum of Natural History, PO Box 50007, SE-104 05 Stockholm, Sweden; e-mails [email protected]; [email protected]

Typescript received 7 February 2009; accepted in revised form 1 June 2010

Abstract: An unusual, bilaterally symmetrical black struc- A flare along the periphery of the alveolus marks a region ture that embraces the and the where the rostrum was not yet formed, suggesting that the and is overlain by a rostrum has been studied in the Santo- capsule extended beyond the rostrum. Modification of the nian–early Campanian (Late Cretaceous) belemnite genus skeleton in Gonioteuthis comprises a set of supposedly inter- Gonioteuthis from Braunschweig, north-west Germany. The related changes, such as innovation of the organic capsule, structure is here named the capsule. Energy dispersed spec- partial elimination of the calcareous rostrum and a diminish- trometry analyses of the capsule show a co-occurrence of sul- ing of the pro-ostracum, resulting in the appearance of a phur with zinc, barium, iron, lead and titanium, suggesting new type of pro-ostracum that became narrower and shorter their chemical association. The capsule was originally made and lost the spatula-like shape and gently curved growth of organic material that was diagenetically transformed into lines of a median field that are typical for the majority of sulphur-containing matter. The material of the capsule dif- and Cretaceous . The partial replacement fers from the chitin of the connecting rings in the same spec- of a calcareous rostrum with an organic capsule in belemni- imens. The capsule has a complex morphology: (1) ventral tellids may have been an adaptive reaction to an unfavour- and dorsal wing-like projections that are repeated in a brevi- able environmental condition, perhaps related to difficulties conic shape of the alveolus, (2) an with lateral lobes in calcium carbonate secretion during the Late Cretaceous and ventral and dorsal sinuses copied by growth lines and that forced to reduce carbonate production and to (3) a ventral ridge that fits with the position of the fissure in secret an organic capsule around the protoconch and the the rostrum. The alveolus in the most anterior part of the phragmocone. rostrum is crater-like. It is lined with thin, pyritized, lami- nated material, which appears to be the outermost portion Key words: Germany, Late Cretaceous, , cole- of the capsule attached to the inner surface of the rostrum. oids, belemnites, shell ultrastructure, organic skeleton.

T he pioneering Early members of the belemnites, the rostra lack their anterior part (see Chris- can be recognized by the presence of a tensen 1975, 1997a; Christensen and Schulz 1997; Kosˇtˇa´k rostrum (Flower 1945; Flower and Gordon 1959), an ink 2005; Kosˇtˇa´k and Wiese 2008). These belemnites represent sac and arm hooks (Mapes et al. 2007, 2010). From that fewer than ten genera of the family Belemnitellidae time until the Jurassic, the rostrum went through signifi- Pavlow, 1914, which was the only existing family in the cant morphological and ultrastructural transformations, northern hemisphere from the Late Cenomanian to the but without attaining the characteristics of the rostrum in latest Maastrichtian (see Christensen 1997b). Two more Jurassic to Cretaceous belemnoids. The rare peculiar features of the rostrum in the belemnitellids are (earliest Jurassic) small-sized rostra supposedly mark the the ventral fissure and the vascular imprints on the sur- rising of the order (Schwegler 1939; Weis face (see Jeletzky 1950, fig. 1; Christensen 1975, fig. 22, and Delsate 2005; Weis and Mariotti 2008). Through the pl. 12, figs 1–4; Christensen and Schulz 1997, fig. 69A–D, , the rostra became larger, more diverse in pls 1–3). The selective postmortem destruction of the shape and more widely distributed geographically. Most alveolar part of the rostrum in belemnitellids points out Jurassic to Cretaceous belemnites retained the radial-con- an unknown structural peculiarity that made this part of centric structure of the rostra. In some Late Cretaceous the rostrum less resistant to fossilization than the rest of

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2010.01027.x 397 398 PALAEONTOLOGY, VOLUME 54 the rostrum. The missing part of the rostrum was trum. This study was aided using scanning electron believed to be originally built up either of some unstable, microscopy and energy dispersive spectrometry. presumably organic material, which was lost during - The genus Gonioteuthis populated the Central European ization (Saemann 1861–1862; Schlu¨ter 1876; Moberg Subprovince in the late middle Coniacian–early Campa- 1885; Crick 1904; Naidin 1964; Christensen 1975) or of nian and rarely reached also the northernmost part of the that was not preserved (Bandel and Spath 1988; Tethyan Realm (Jeletzky 1950; Christensen 1997a, b). The Barskov et al. 1997; Christensen 1997a; Kosˇtˇa´k and Wiese presence of the capsule indicates the partial substitution 2008). Since then, the hypothesis of an organic composi- of calcareous material of the rostrum with an organic tion of the alveolus part of the rostrum has remained substance. This change in skeletal material may have been untested and no observations supporting it have been an adaptive reaction to an environment in the Central reported. The idea of an aragonitic composition of the European Subprovince during the Late Cretaceous that alveolar part of the rostrum is based on stable-isotope was possibly unfavourable to calcium carbonate secretion data from the Turonian in northern Siberia showing con- (see Worsley 1971). sistently high d18O values in the belemnite rostra and lower values in the co-occurring calcitic shells of gastro- pods and inoceramids, as well as different 13C contents in MATERIAL AND METHODS the alveolus and postalveolar parts, respectively (Naidin et al. 1987; Barskov et al. 1997; Dauphin et al. 2007). The belemnites under examination, purchased in Ger- A perishable substance of unknown composition lining many in the beginning of the twentieth century, are now a pseudoalveolus (an unusually shortened alveolus typical housed at the Department of Palaeozoology of the for belemnitellid genera Praeactinocamax Naidin, Actinoc- Swedish Museum of Natural History. Their original labels amax Miller, Goniocamax Naidin, Gonioteuthis Bayle, contain the following information: ‘ granula- Naidin) and continuing a calcareous tus Blainv., Upper Cretaceous, Braunschweig; Collegium rostrum was discovered in the belemnitellid belemnite Coralinum, Braunschweig’. Judging from the distribution Gonioteuthis granulata (de Blainville) and briefly of Gonioteuthis granulata in northern Germany, the speci- described by Ernst (Ernst 1964, pp. 156–157). Based on mens must have come from Santonian to lower Campa- his observations, Ernst concluded that the perishable sub- nian strata (Christensen 1976). stance was associated with the pseudoalveolus, and the Twenty fragmentary rostra are available for study; their latter was an original shell character rather than a post- maximum diameter is 12 mm, and the maximum pre- mortem broadening of the alveolus. Since then, this idea served length is 68 mm; seven rostra preserve the alveolus has attracted little attention. with a fragmentary phragmocone inside. Initially, the cap- Shells of Gonioteuthis preserving what could be referred sule was noticed in the alveolus because of its black col- to as ‘a perishable substance of unknown composition our, which contrasts with the brown hue of the rostrum. lining a pseudoalveolus’ (Ernst 1964, pp. 156–157) have The specimens were split longitudinally to expose the been recovered in collections of the Department of Palae- alveolus and to access the site of the potentially preserved ozoology, Swedish Museum of Natural History. They structure. Because of the ventral fissure of the rostrum, come from Braunschweig, north-west Germany. The pres- the longitudinal fracture usually runs in the sagittal plane. ent article reports our recent study on this enigmatic, In four of the split specimens, a protoconch was exposed rarely observed skeleton structure. This originally organic as well. The portions of rostra that revealed morphologi- skeletal element – here named a capsule – embraces the cal features of the structure under study, such as a black protoconch and phragmocone and is overlain by the ros- material and curved growth lines, were examined in a

EXPLANATION OF PLATE 1 Figs 1–7. Gonioteuthis granulata, Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany; NRM-PZ Mo. 8183, a ventro-dorsally split specimen. 1, general view of the phragmocone surrounded by the capsule showing ventral (left) and dorsal (right) wing-like projections on the longitudinal fracture of the rostrum; scale bar represents 5 mm. 2, view of orad end of the capsule and surrounding alveolus of the rostrum; scale bar represents 5 mm. 3, enlarged detail of 1 to show the ventral portion of the capsule-bearing longitudinal ridges and adjoined part of the fissure; scale bar represents 500 lm. 4, enlarged detail of 3 to show that material of the capsule continues into the material lining the fissure; scale bar represents 250 lm. 5, enlarged detail of 3 showing a cross section of the capsule and its granular ultrastructure; scale bar represents 25 lm. 6, a fragment of the thin conotheca with a mural ring of a ; scale bar represents 100 lm. 7, enlarged detail of 1, a granular ultrastructure of the dorsal wing-like projection of the capsule; scale bar represents 100 lm. PLATE 1

1 2

3

4

5

6

7

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 400 PALAEONTOLOGY, VOLUME 54 scanning electron microscope (Hitachi S-4300) equipped ues along the chamber length and forms the phragmo- with an energy dispersive spectrometer (EDS). Quantita- cone wall. In the first chambers, the phragmocone wall tive analyses were made using an INCA program that (conotheca), which is a continuation of the adoral part of estimates peak-to-background ratios. Elemental analyses septum is c.4lm thick (Pl. 4, figs 3, 4). In the late onto- were performed at an accelerating voltage of 15 kV, and genetic stages, the mural rings are observed to be attached energy calibration as measured on standard minerals of directly to the capsule. The specimen NRM-PZ Mo. 8079 known accuracy was used. All elements were analysed, exhibits the fragmentarily preserved septa attached by and no peaks were omitted. Data obtained for the cap- their mural rings directly to the inner surface of the cap- sule, rostrum, shell wall, septa and chamber filling were sule at the adult stage. The mural rings of the septa are compared. broad (in ratio to length); they are about one- sixth to one-third of the chamber length (Pl. 6, fig. 5). Repository. All material studied is housed at the Department of The line has weak ventral and dorsal saddles (Pl. 7, Palaeozoology, Swedish Museum of Natural History, Stockholm fig. 5). There is an unpaired, narrow, long (about cham- (abbreviation: NRM-PZ Mo.). ber length) mid-dorsal scar (one in each chamber) (Pl. 7, fig. 5). The is narrow and ventral (Pl. 5, figs 1– 3; Pl. 6, fig. 1). The siphuncular tube is almost cylindrical, OBSERVATIONS slightly swollen between the septal necks (Pl. 6, fig. 1). The septal necks are about one-fourth to one-third of the Morphology of the protoconch in Gonioteuthis chamber length. Their apical part merges into the con- necting ring running through the previous chamber and The protoconch is oblate spherical, slightly compressed is attached to the inner surface of the previous septal neck longitudinally (Pl. 1, fig. 1; Pl. 2, figs 4, 6; Pl. 4, figs 1, 2, (Pl. 6, figs 1–9). In the contact between the connecting 5), with the length being less than the width. For exam- ring and septal neck, there is a ring with spherulitic- ple, the length and width of the protoconch is 0.5 and prismatic ultrastructure (Pl. 6, figs 8, 9). The phragmo- 0.7 mm, in specimens NRM-PZ Mo. 8181 and 8183, cone is surrounded by the capsule (Pl. 1, fig. 1; Pl. 4, figs respectively. The aperture is closed by an originally 3, 4; Pl. 5, figs 1, 2). organic membrane. The protoconch is surrounded by the capsule (Pl. 2, figs 4, 6). Morphology of the capsule in Gonioteuthis

Morphology of the phragmocone in Gonioteuthis The capsule is a bilaterally symmetrical structure located between the phragmocone and the rostrum (Pl. 1, figs 1, The phragmocone exhibits both longiconic and breviconic 2). It is thin around the protoconch (Pl. 2, figs 4, 6) and characteristics (Pl. 1, fig. 1; Pl. 2, figs 1, 2; Pl. 3, fig. 1; the first c. 15 chambers of the phragmocone (Pl. 4, figs 3, Pl. 4, figs 1, 2, 5). At some distance from the protoconch, 4); thereafter, it gets abruptly thicker (Pl. 3, figs 1, 2). At the longiconic phragmocone changes shape abruptly and the anteriormost part of the preserved portion of the skel- becomes breviconic. There are 20–25 short chambers eton, the capsule is thicker than the rostrum; the rostrum (Pl. 1, fig. 1; Pl. 4, figs 2, 3; Pl. 6, fig. 1). A septum con- gets thinner and wedges out, but the capsule shows no sists of two more or less distinctly separated adoral and signs of thinning. Thereafter, the capsule becomes a solo adapical portions that are of approximately equal thick- structure surrounding the phragmocone. The length of ness. The adapical part of the septum forms a prominent the anterior portion of the capsule that is not covered by mural ring and wedges out, while the adoral part contin- the rostrum is not yet known.

EXPLANATION OF PLATE 2 Figs 1–6. Gonioteuthis granulata, Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany; A–D-NRM-PZ Mo. 8180. 1, the capsule exposed on the longitudinally split rostrum; scale bar represents 2.5 mm. 2, counter-part of the specimen seen in 1 showing an imprint of the capsule on the inner surface of the rostrum and a ridged anterior part of the rostrum, scale bar represents 2.5 mm. 3, 5, growth lines of the capsule showing a lobe between two sinuses; scale bars represent 2.5 mm and 1 mm, respectively. 4, 6, NRM-PZ Mo. 8181. 4, a spherical protoconch surrounded by a partly preserved capsule; scale bar represents 250 lm. 6, enlarged detail of 4 showing remains of the capsule near the apical portion of the protoconch; scale bar represents 0.5 mm. PLATE 2

1 2

4

3

5 6

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 402 PALAEONTOLOGY, VOLUME 54

The capsule has ventral and dorsal wing-like projec- part, the alveolus has a crater-like shape with a small pit tions (Pl. 1, figs 1, 2) that are finely ornamented with of its longiconic part on the bottom of the crater and growth lines and longitudinal ridges (Pl. 1, fig. 3; Pl. 2, with a narrow rounded marginal flare distinguished figs 1–3, 5). The growth lines show that the capsule had because of its larger angle of expansion than in the pre- a simple aperture in earlier ontogenetic stages and a ceding part (Pl. 3, fig. 1). The marginal edge flare marks complex aperture in later ontogeny with pronounced the real aperture of the rostrum rather than its broken lateral lobes and ventral and dorsal sinuses (Pl. 2, figs end. This flare indicates that the capsule further enlarges 1–3, 5). There is a pronounced ventral ridge (Pl. 5, figs its angle of expansion in this part of the skeleton. The 1, 2) that fixes the capsule to the fissure in the rostrum. internal surface of the crater is roughly sculptured with The specimen NRM-PZ Mo. 8180, cut through the longitudinal ridges and grooves (Pl. 2, fig. 2; Pl. 3, figs 1, anterior part of the shell, shows the siphuncle to lie 4). However, the ridged inner surface of the crater is near the inner surface of the capsule; the wall of the smoothed by a thin layer of the capsule on its surface phragmocone (conotheca) is missing between them (Pl. 3, fig. 5). (Pl. 5, figs 1–3). The rostrum has a ventral fissure with rounded edges near the outer surface. The fissure comes to the surface of the capsule. The sides of the fissure are triangle-shaped Internal morphology of the rostrum in Gonioteuthis with an acute angle near the protoconch. They are flat- tened and roughly ornamented with obliquely oriented The rostrum is typically solid, with the exception of the ribs and grooves. The surface of the fissure is coated with anteriormost part, which is abnormally loosely mineral- a black material like that of the capsule or bears numer- ized and porous and has a roughly ridged inner surface. ous fragments of this material. A thin pyritized laminated material, apparently the outer- most portion of the capsule, is consistently attached to the inner surface of the rostrum. Morphology of the pro-ostracum in Gonioteuthis In the rostrum, the alveolus embraces the capsule fol- lowing the shape of the latter and repeating its ontoge- A pro-ostracum was observed in the specimen NRM-PZ netic changes (compare Pl. 2, figs 1, 2). The initial Mo. 8179 (Pl. 7, figs 1–4). The pro-ostracum is narrow portion of the alveolus is either longiconic tubular or and has an acute pointed triangular median field and moderately breviconic like that of the capsule (Pl. 4, figs longitudinal ridges of the lateral fields on its sides. It 2, 5). Subsequently, the alveolus abruptly broadens and shows longitudinal closely packed bends with a feather- continues to broaden forwards (Pl. 3, fig. 1; Pl. 4, fig. 1). like pattern in each (Pl. 7, figs 1–4). It lacks a spatula-like In the specimen NRM-PZ Mo. 8180 (Pl. 2, figs 1–3), the shape and gently curved growth lines of a median field tubular part of the alveolus is about 2 mm in maximum that are typical for the majority of Jurassic and Creta- diameter, and through an interval of 6 mm, the alveolus ceous belemnites (see Doguzhaeva in press). gets three-and-a-half times wider. In the specimen NRM- PZ Mo. 8181, the alveolus expands by about 70 degrees from the tubular portion (Pl. 3, fig. 1). The changes in EDS data on the chemical composition of the skeleton in alveolus diameter are not reflected on the outer surface Gonioteuthis of the rostrum, because the rostrum gets thinner while the alveolus gets broader. Thus, the external cylindrical A rostrum was analysed with the aid of polished and shape of the rostrum is not broken. In the anteriormost slightly etched longitudinal sections. It shows high peaks

EXPLANATION OF PLATE 3 Figs 1–7. Gonioteuthis granulata, Santonian–early Campanian, late Cretaceous; Braunschweig, north-west Germany. NRM-PZ Mo. 8181. 1, a polished longitudinal section through the protoconch and alveolus (filled with a granular material) and partly preserved capsule located between the alveolus and rostrum; scale bar is 5 mm. 2, enlarged detail of 1 to show rapid capsule thickening in anterior direction; the alveolus is on its left and rostrum on right; scale bar is 1 mm. 3, enlarged detail of 2 to show an irregular mineralization of the capsule; scale bar represents 50 lm. 4, the heavily ridged inner surface of the rostrum close to its anterior edge that was in contact with the outer surface of the capsule; scale bar represents 1 mm. 5, enlarged detail of 4 showing a film lining of the inner surface of the rostrum that seems to be a remnant of the capsule; scale bar is 100 lm. 6, enlarged detail of 5 showing a microlamination of the film lining the inner surface of the rostrum; scale bar represents 2.5 lm. 7, enlarged detail of 4 showing an irregular patch-like pattern of the film on the inner surface of the rostrum; scale bar represents 1 lm. PLATE 3

1 2

3

4

6

5

7

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 404 PALAEONTOLOGY, VOLUME 54

A 1

23

TEXT-FIG. 1. Energy dispersive spectrometer data on the capsule material and the fill of the adjusted portion of the chamber in Gonioteuthis granulata. NRM-PZ Mo. 8180. Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany. A, cross section of the capsule with the marked points of the spectra 1–3 taken; scale bar represents 1 mm. 1, 2, the spectra of the capsule. 3, the spectrum of the chamber filling near the capsule. Abbreviations: c, chamber; csos, the outer surface of the capsule; cs, cross section of the capsule; 1–3, points where the spectra were taken. of Ca, O and C (Text-fig. 2A). Mainly along the central and the phragmocone wall show high peaks of Ca, P, O line and periphery, but also around the alveolus (NRM- and lower peaks of C, Si, Fe and S (NRM-PZ Mo. 8181; PZ Mo. 8188; Pl. 4, fig. 5), the rostrum is silicified. Text-fig. 2C). The connecting rings of the siphuncle A fissure was analysed in the specimens longitudinally (NRM-PZ Mo. 8180; Text-fig. 2D) reveal high peaks of split in a way that both sides became exposed (NRM-PZ Ca, P, O and lower peaks of C and Fe. The capsule Mo. 8188; Text-fig. 2B). The fissure shows high peaks of (NRM-PZ Mo. 8180; Text-figs 1, 2E) shows a high con- Si, Ca, O and S and lower peaks of C and Fe. The septa tent of S that is associated with a high content of either

EXPLANATION OF PLATE 4 Figs 1–5. Gonioteuthis granulata, Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany. NRM-PZ Mo. 8181. 1, an overall view caused by a longitudinal fracture through the protoconch and several initial chambers of the phragmocone (bottom) with the exposed angle-shaped alveolus (dark) that fits with the paired wing-like projections of the capsule (compare with Pl. 1, fig. 1); scale bar represents 5 mm. 2, enlarged detail of 1 showing a spherical protoconch and gradually expanding initial portion of the phragmocone; scale bar represents 2 mm. 3, enlarged detail of 2 showing the capsule between the rostrum and the conotheca; scale bar represents 100 lm. 4, enlarged detail of 3 showing the capsule is structurally different from the rostrum (left) and conotheca (right); scale bar is 25 lm. 5, Gonioteuthis granulata (NRM-PZ Mo. 8188). Polished longitudinal section of the shell through the protoconch and alveolus surrounded by heavily silicified portions (dark grey) of the rostrum; scale bar represents 1 mm. PLATE 4

1 2

5 3

4

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 406 PALAEONTOLOGY, VOLUME 54

Fe or Mn (Text-fig. 1A, 1), or Ba, Pb and Ti (Text- The morphology of the skeleton in Gonioteuthis and its fig. 1A, 2), or Zn and Na (Text-fig. 1A, 3). The elements significance for the remaining members of the family Ca, C, K, Si and Sr constitute a minor part. However, the Belemnitellidae external portion of capsule material that is in close con- tact with the inner surface of the rostrum shows high It is revealed that in Gonioteuthis the skeleton comprises peaks of Ca but lower peak of S (NRM-PZ Mo. 8179; (1) a comparatively large protoconch with a maximum Text-fig. 2E). The EDS data favour the presence of such diameter of 0.7 mm; (2) a phragmocone with c. 20–25 minerals as barite, pyrite, sphalerite and galena in this chambers or slightly more, a wall formed by continua- structure of the shell. Barite and pyrite form curved inter- tions of the adoral portions of the septa, a ventral siphun- connected bands. The crystals of pyrite form tiny spheres cle, long septal necks and a mid-dorsal scar; (3) a that are c.10lm in diameter or irregularly shaped mass rostrum with a loosely mineralized anterior part, a well- of particles smaller than the balls (Pl. 1, fig. 7). Sphalerite developed fissure and an alveolus that is longiconic or forms perfectly shaped, highly lustrous, black and com- moderately breviconic in early ontogeny and broadly cra- paratively large dispersed crystals (Pl. 2, fig. 1). ter-like, with a narrow marginal flare in late ontogeny; (4) a small triangular pro-ostracum consisting of closely spaced bands with a feather-like arrangement of the crys- DISCUSSION tallites and (5) a bilaterally symmetrical organic capsule embracing the protoconch and the phragmocone and rep- Fossilization and original composition of the capsule in resenting a separate structure surrounding a phragmocone Gonioteuthis in front of the place where the rostrum wedges out. The originally organic capsule has a complex morphology and The material of the capsule shows a co-occurrence of sul- possesses ventral and dorsal wings, a ventral ridge, two phur with zinc, barium, iron, lead and titanium, thus lateral lobes and a dorsal sinus. The ventral ridge pene- suggesting their chemical association. High concentrations trates the fissure. Based on growth lines, the aperture of of Zn, Fe, Ba and Pb are known to be associated with the capsule has lateral lobes. These lobes are additional substances high in organic matter and sulphur. Therefore, evidence that the capsule was not coated anteriorly by the the capsule is considered to have originally consisted of rostrum. Therefore, the modification of the skeleton in organic material that has been transformed diagenetically Gonioteuthis comprises a set of supposedly interrelated into the sulphur-containing minerals barite, pyrite, sphal- changes, such as the innovation of the organic capsule, a erite and galena. The material of the capsule was appar- partial elimination of the calcareous rostrum; a diminish- ently different from chitin, because the chemical ing of the pro-ostracum resulting in the appearance of a composition differs from that of the originally chitinous new type of pro-ostracum. connecting rings in the same specimens. The connecting Although the morphology of the skeleton, with the rings are composed of calcium phosphate and lack S, Zn, exception of the rostrum, is yet poorly known in belem- Fe, Ba and Pb. The pathway of fossilization of the coleoid nitellids, the ridge-like structure on the ventral side of the structures that are known to have been originally chitin- phragmocone in (Gustomesov 1980, fig. 1c, d) ous in living forms, such as arm hooks or mandibles, was is comparable with the ventral ridge of the capsule in Go- phosphatization, pyritization or carbonization, depending nioteuthis. This is indirect evidence that the capsule is a on environments in places of preservation (see Doguzha- characteristic element of the skeleton of the belemnitel- eva and Mutvei 2006; Doguzhaeva et al. 2006a–c, 2007c). lids. It is of great importance that a pro-ostracum similar In the studied belemnite, the chitinous connecting rings to that of Gonioteuthis, much narrower than that of the were preserved through phosphatization. was observed in the late Maastrichtian Belem-

EXPLANATION OF PLATE 5 Figs 1–6. Gonioteuthis granulata, Santonian–late Campanian, Late Cretaceous; Braunschweig, north-west Germany. NRM-PZ Mo. 8180. 1, cross section of the capsule showing a ventral ridge (left bottom corner) and a siphuncle at a short distance from the ridge, scale bar represents 2.5 mm. 2, enlarged detail of 1 showing the position of a siphuncle, scale bar represents 1 mm. 3, enlarged detail of 1 showing irregular mineralization of the ventral ridge and a septum with a septal neck adjusted to the ridge, scale bar represents 0. 5 mm. 4, enlarged detail of 1 showing the capsule wall and its surface, scale bar represents 1 mm. 5, uneven surface of the capsule, scale bar represents 1 mm. 6, enlarged detail of 5 showing regular numerous hollows on the outer surface of the capsule, scale bar represents 250 lm. PLATE 5

1 2

3

4

5 6

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 408 PALAEONTOLOGY, VOLUME 54 nitella bulbosa (Jeletzky 1966, p. 48, pl. 1, fig. 1A, D–E). (Kosˇtˇa´k 2005) and the middle Coniacian–early Campa- Such kind of innovation like a new type of pro-ostracum nian lineage of the genus Gonioteuthis (Ernst 1964; Ernst observed in two noticed above genera must have and Schulz 1974). Later, it was shown that the process of requested essential reconstruction of the anterior part of increasing the depth of the alveolus in the Gonioteuthis the skeleton in belemnitellids. Substitution of the calcare- lineage was asynchronous and that the pseudoalveolus ous rostrum with the organic capsule is considered to be depth first decreases but then increases from the late early sufficient initiation for this. The shell of belemnitellids Santonian (Christensen and Schulz 1997). However, Be- exhibits no evidence of any other significant transforma- lemnitella d’Orbigny, Belemnella Nowak and Fusiteuthis tion that could result in origin of a new type of the pro- Kongiel are known to have no pseudoalveolus (see Kong- ostracum. iel 1962; Naidin 1969; Christensen 1975, 1997a, b, 2002). Morphology of the pseudoalveolus can help to clarify Besides, in Belemnitella, imprints of the phragmocone can whether the capsule was a characteristic skeleton structure be observed on the inner surface of the calcitic rostrum. of the family Belemnitellidae. Initially, the term pseudoal- These observations question secretion of the capsule in veolus has been applied to an unusually short alveolus of these genera. belemnitellids, supposing that it was a postmortem arte- Based on observations of the capsule in Gonioteuthis fact resulting from diagenetic distraction or dissolution of and presence of the pseudoalveolus in four further the rostrum around the alveolus (see Krymgolts 1958, p. belemnitellid genera (Praeactinocamax, Actinocamax, Gon- 150, fig. 114; Bandel and Spath 1988; Christensen 1997a, iocamax, Belemnellocamax), we suggest that the partial 1982). Ernst (1964) found an earlier unknown perishable replacement of calcareous rostrum with organic capsule substance in Gonioteuthis that was intercalated between was a Late Cretaceous evolutionary event in Belemnoidea. the phragmocone and rostrum. Hence, the pseudoalveolus was interpreted by him as a primary shell structure that housed, additionally to a phragmocone, the perishable The evolution of the rostrum in the Belemnoidea material around it. Ernst’s idea is supported by Kosˇtˇa´k and Wiese (2008), who considered the shape of the The earliest rostrum-bearing genera, Flower pseudoalveolus to be a taxonomic feature at the generic and Gordon, Palaeoconus Flower and Gordon, Bactritimi- level. Our study on Gonioteuthis confirms Ernst’s opinion mus Flower and Gordon and Gordoniconus Mapes, Weller as well. It is herein demonstrated that the pseudoalveolus and Doguzhaeva are from the Eumorphoceras Zone of accommodates a well-developed capsule of complex mor- the Early Carboniferous, which means that a rostrum phology, the shape of the latter being repeated in the appeared at an early stage of coleoid evolution (Flower shape of the former. These two structures were apparently 1945; Flower and Gordon 1959). The evolution of the interrelated in their origin. If the capsule is not preserved rostrum is briefly exemplified with the data given later. because of its organic composition and little potential for Order Hematitida (Early Carboniferous): in Hematites, fossilization, the pseudoalveolus evidences its original a prominent rostrum, about 50 mm long, has numerous presence. The gradually increasing depth of the pseudoal- broad longitudinal ridges. The flat crests of the ridges veolus was considered to be an evolutionary trend of bear a row of pits that continue into pronounced narrow the Cenomanian–Santonian lineage of Praeactinocamax channels. The central zone of the postalveolar part of the

EXPLANATION OF PLATE 6 Figs 1–9. Morphology of the siphuncular tube of Gonioteuthis granulata, Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany. NRM-PZ Mo. 8079. 1, two septal necks on the ventral side of the siphuncular tube, scale bar represents 250 lm. 2, 3, enlarged details of 1 showing a contact between the septal neck and the connecting ring attached to its inner surface; scale bar represents 100 lm. 4, enlarged details of 3 showing wedging out of the connecting ring near the apical tip of the previous septal neck that transits into the connecting rings of the previous chamber, scale bar represents 20 lm. 5, enlarged details of 2 showing wedging out of the connecting ring near the apical tip of the previous septal neck; scale bar represents 25 lm. 6, isolated fragment of siphuncular tube crusted by calcium phosphate with a partly preserved narrowing apical segment (top) that was housed in the previous septal neck, scale bar represents 200 lm. 7, isolated fragment of the siphuncular tube showing that a succeeding connecting ring (bottom) is inserted into the previous septal neck (top), scale bar represents 100 lm. 8, isolated fragment of siphuncular tube crusted by calcium phosphate with a partly preserved apical segment (top) showing a narrow band around its apical end (top), scale bar represents 200 lm. 9, Enlarged detail of 8 showing loosely mineralized porous spherulitic-prismatic deposits in the contact between the septal neck and connecting ring, scale bar represents 25 lm. PLATE 6

1 6

2

7

3

8

4

9 5

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 410 PALAEONTOLOGY, VOLUME 54 rostrum is built of loosely packed calcareous rods and Order Belemnotheutida (–Early Creta- seems to contain originally organic material. The rostrum ceous): in Pearce, the sheath-like rostrum has little similarity with any other known rostra (see has a short postalveolus part that consists of a solid pris- Doguzhaeva in press, fig. 1A; Doguzhaeva et al. 2002, matic outer layer and a loosely mineralized inner layer 2007b). (Makowski 1952; Bandel and Kulicki 1988). In the adoral Order Donovaniconida (Early–Late Carboniferous): in direction, the rostrum loses the inner layer, and the outer the Early Carboniferous Gordoniconus (Mapes et al. 2007, layer becomes spherulitic-prismatic. The numerous inter- 2010) and Late Carboniferous Saundersites Doguzhaeva, spaces between the crystalline units of the rostrum indi- Mapes and Mutvei (Doguzhaeva et al. 2007a, fig. 6.1A, cate a higher organic content (Doguzhaeva et al. 2006b). B), the rostrum is sheath-like with a comparatively short, Order Diplobelida (–Late Cretaceous): loosely calcified, pointed, longiconic postalveolus part in Tauriconites Kabanov (in Drushchits, Kabanov and which is about half the body chamber in length Nerodenko 1984), the rostrum is stout, short, with a deep (Doguzhaeva et al. 2010, fig. 1C). In Late Carboniferous alveolus that comes close to the of the rostrum. The Donovaniconus Doguzhaeva, Mapes and Mutvei, the ros- rostrum possesses sectors with a feather-like arrangement trum is sheath-like; the postalveolus part is unknown of crystallites similar to that in the Tertian sepiids (Doguzhaeva et al. 2003a). (Doguzhaeva 2008). Order (Late Carboniferous–Jurassic): the Order Belemnitida (Early Jurassic–Late Cretaceous): in Late Carboniferous Mutveiconites Doguzhaeva has a the Early Jurassic Lissajous, the primordial loosely calcified sheath-like rostrum with a short rounded rostrum was loosely calcified and apparently contained postalveolus portion that is about the length of a proto- much organic material (Doguzhaeva et al. 2003b, text-fig. conch in juvenile individuals (Doguzhaeva et al. 2006c, 9). The rostrum has a radial-concentric structure. The pl. 1, figs A, B, pl. 5, figs A, B, 2010, fig. 1B). The ros- Late Cretaceous Gonioteuthis has an organic capsule in trum extends forwards to a distance of about the first ten addition to the calcareous rostrum. The loss of the ante- chambers of the longiconic phragmocone. In the Late rior part of the rostrum and development of pseudoalveo- Stenoconites Gordon, the rostrum is about 8 cm lus in several other genera of the Late Cretaceous long and is characteristically ribbed (Gordon 1966, figs belemnites apparently evidences the development of the 2a, c; Chen 1982, text-fig. 5). In the Late Aulacoc- capsule in them as well. The Maastrichtian Belemnitella, eras von Hauer, Dictyoconites Mojsisovics (Mojsisovics Belemnella and Fusiteuthis are the last survivors of the 1902, pls 8–13) and Austroteuthis Jeletzky and Zapfe Belemnoidea, coming near the K ⁄ Pg boundary (Kongiel (1967, pl. 2, figs 1A–D), a large, c. 15 cm long rostrum 1962; Christensen 2002). These forms exhibit a deep has a regularly cylindrical oral half and gradually tapers breviconic alveolus and lack the pseudoalveolus. to a point in the apical half; the apical angle is about Thus, in the earliest Carboniferous belemnoids, the ros- 14–17 degrees. In Atractites Gu¨mbel, the rostrum has an trum morphology and the degree of mineralization were elongate fusiform shape (Mariotti and Pignatti 1999, fig. highly variable, with the exception of a constantly high 1). Rostra are typical ribbed, but some are smooth. The content of organic material (see Doguzhaeva et al. 2002, rostrum has a radial-concentric structure with an alterna- 2010; Mapes et al. 2010). It is remarkable that in the four tion of colourless and black layers; the latter were suppos- Early Carboniferous genera so far described the rostrum edly enriched in organic material (Krymgolts 1958). is more pronounced and better developed than in the Order (Late Permian–Early Jurassic): Late Carboniferous belemnoids. In the Late Permian be- at this time, the rostrum is unknown in this order. lemnoids, which are still extremely rare and only known

EXPLANATION OF PLATE 7 Figs 1–6. Pro-ostracum and mid-dorsal scar in Gonioteuthis granulata, Santonian– early Campanian, Late Cretaceous; Braunschweig, north-west Germany. A–D, NRM-PZ Mo. 8179. 1, a dorsal view on the phragmocone showing converging ribs forming a narrow, acutely pointed median field of the pro-ostracum and longitudinal ridges of the lateral fields on each side of the median field; scale bar represents 0.5 mm. 2, a dorso-lateral view on the phragmocone showing converging ribs of the median field and longitudinal ribs of the lateral fields, scale bar represents 200 lm. 3, enlarged detail of 2 showing longitudinal closely packed bands with a feather-like pattern in each and that the pro-ostracum is built from; scale bar represents 50 lm. 4, enlarged detail of 2 showing that the pro-ostracum wall differs from the underlying material; scale bar represents 50 lm. 5, NRM-PZ Mo. 8181; a mid-dorsal scar on the inner surface of the conotheca and a suture line with a weak dorsal saddle; scale bar represents 0.5 mm. 6, enlarged detail of 5 showing that the rostrum (left) and the conotheca (right) are separated with a structure located between them; this may be a pro-ostracum; scale bar represents 20 lm. PLATE 7

1 2

3

4

5

6

DOGUZHAEVA and BENGTSON, Gonioteuthis granulata 412 PALAEONTOLOGY, VOLUME 54

AB

C D

E F

TEXT-FIG. 2. A–F, Energy dispersive spectrometer data on the shell of Gonioteuthis granulata; Santonian–early Campanian, Late Cretaceous; Braunschweig, north-west Germany. A, NRM-PZ Mo. 8081, rostrum showing peaks of Ca, O and C. B, NRM-PZ Mo. 8188, fissure showing high peaks of Sn, Si, Ca, O and S, and lower peaks of C and Fe. C, NRM-PZ Mo. 8181, conotheca showing high peaks of Ca, P, O and lower peaks of C, Si, Fe and S. D, NRM-PZ Mo. 8180; connecting ring of the siphuncle showing high peaks of Ca, P, O and lower peaks of C and Fe. E, NRM-PZ Mo. 8180; capsule showing high peaks of Ba, Fe, C, Ca, Te, O and minor peaks of Sr, Si, S, Mg and Zn. F, NRM-PZ Mo. 8179; capsule showing high peaks of Fe, Ca, K, C, O, Mn and minor peaks of Si, Sr, S, Mg, Na and Zn. DOGUZHAEVA AND BENGTSON: ORGANIC SHELL STRUCTURE IN BELEMNITES 413 from the USA (Gordon 1966) and South China (Chen and KULLMANN, J. (eds). Cephalopods – present and past. 1982), the rostra are essentially different from Carbonifer- Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, 765 pp. ous rostra but similar to ones. They are well BARSKOV, I. S., KIYASHKO, S. I., DAUPHIN, Y. and mineralized, have a radial-concentric structure and lack DENIS, A. 1997. Microstructures des zones calcitiques et indications of a significant content of organic material aragonitiques des rostres de Goniocamax (Cephalopoda, Belemnitida), de Turonian de Sibe´rie du Nord. Geodiversitas, (Chen 1982, pl. 1, figs 1–20, text-figs 2–3). In Triassic 19, 669–680. belemnoids, which are less restricted in geographical dis- BAYLE, E. 1878. Explication de la carte ge´ologique de la France, tribution than their Carboniferous and Permian precur- Vol. 4. Fossiles principaux des terrains. Imprimerie Nationale, sors, the rostra can achieve large sizes about 15–20 cm in Paris. length and 2–3 cm in diameter, are strongly mineralized BLAINVILLE, M. H. D. de 1827. Me´moire sur les Belemnites and are smooth or have longitudinal ribs and grooves considere´es zoologiquement et ge´ologiquement. F.-G. Levrault, (Mojsisovics 1871, 1902; Bu¨low 1915; Smith 1927; Krym- Paris, 136 pp. golts 1958; Mariotti and Pignatti 1999). In the first Early BU¨ LOW, E. von 1915. Orthoceren und Belemniten der Trias Jurassic (Hettangian) belemnitids, the rostrum is small von Timor. Palaeontologie von Timor, 4, 1–72. and has a typical radial-concentric structure. This type of CHEN, T. 1982. Discovery of Permian belemnoids in South rostrum morphology and structure was developed in the China, with comments on the origin of . Acta Palae- ontologica Sinica, 21, 181–190. majority of Jurassic to Cretaceous belemnitids. In Jurassic CHRISTENSEN, W. K. 1975. Upper Cretaceous belemnites belemnotheutids, such as Belemnotheutis, the rostrum has from the Kristianstad area in Scania. Fossil and Strata, 7, a short postalveolus part (Makowski 1952). In the Early 1–69. Cretaceous belemnites, the rostrum is variable in size and —— 1976. Palaeobiogeography of Late Cretaceous belemnites of morphology; typically, they have a radial-concentric struc- . Pala¨ontologische Zeitschrift, 50, 113–129. ture. In some Late Cretaceous belemnites, the rostrum is —— 1982. Late Turonian – early Coniacian belemnites from partly replaced with an organic capsule. No similar phe- western and central Europe. Bulletin of the Geological Society of nomenon of fossilization is known in any other group of Denmark, 31, 63–79. belemnoids. —— 1997a. The Late Cretaceous belemnite family Belemnitelli- The partial substitution of a calcareous rostrum with dae: and evolutionary history. Bulletin of the an organic capsule may have been an adaptive reaction to Geological Society of Denmark, 44, 59–88. —— 1997b. Palaeobiography and migration in the Late Creta- an environment unfavourable to carbonate secretion ceous belemnite family Belemnitellidae. Acta Palaeontologica during the Late Cretaceous (Worsley 1971). This may Polonica, 42, 457–495. have induced belemnitellids to reduce carbonate secretion —— 2002. Fusiteuthis polonica, a rare and unusual belemnite and to secret an organic capsule for protecting the from the Maastrichtian. Acta Palaeontologica Polonica, 47, phragmocone. 679–683. —— and SCHULZ, M.-G. 1997. Coniacian and Santonian bel- Acknowledgements. We thank the Swedish Academy of Sciences emnite faunas from Bornholm, Denmark. Fossil and Strata, for supporting a ten-month visit of the first author as guest 44, 1–128. researcher during 2008–2010 at the Department of Palaeozoolo- CRICK, G. C. 1904. Notes on Actinocamax Mill., its identity gy, Swedish Museum of Natural History, Stockholm. We extend with Atractilites Link. Geological Magazine, 1, 407–410. our thanks to Royal H. Mapes (Ohio University, USA), an anon- DAUPHIN, Y., WILLIAMS, T. C. and BARSKOV, I. 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