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Home , Belemnites (genus), Belemnitida, Belemnitina, Phragmoteuthida, Sepia (cephalopod)

Geology, published online on 9 August 2012 as doi:10.1130/G33402.1 originated in the —A new look at an old group

Yasuhiro Iba1, Shin-ichi Sano2, Jörg Mutterlose3, and Yasuo Kondo4 1Hokkaido University of Education, Kushiro 085-8580, Japan 2Fukui Prefectural Dinosaur Museum, Fukui 911-8601, Japan 3Department of Geology, Mineralogy and Geophysics, Ruhr-University, Bochum 44801, Germany 4Department of Natural Science, Kochi University, Kochi 780-8520, Japan

ABSTRACT standing evolutionary dynamics of nektonic biota (e.g., Mutterlose, 1998; Belemnites ( ), a very successful group of Christensen, 2002; Iba et al., 2011). Mesozoic , provide an important clue for understanding Although the evolutionary history of belemnites through the late Mesozoic marine ecosystems and the origin of modern cephalopods. Early to has been well studied (e.g., Doyle, 1993; Following current hypotheses, belemnites originated in the earliest Mutterlose, 1998; Riegraf et al., 1998; Schlegelmilch, 1998; Christensen, Jurassic (, 201.6–197 Ma) with very small forms. Accord- 2002), their early evolution has not yet been fully understood. Currently ing to this view their paleobiogeographic distribution was restricted the Hettangian Schwegleria (suborder ) is considered as to northern Europe until the (190–183 Ma). The fossil the root of all belemnites (e.g., Doyle, 1994; Schlegelmilch, 1998; Weis record is, however, biased by the fact that all the previous studies on and Delsate, 2006). Belemnites originated in northern Europe as very belemnites focused on Europe. Here we report two belemnite taxa small forms in the earliest Jurassic following the Triassic–Jurassic mass from the Hettangian of Japan: a new species of the Sinobelemnitidae event (Doyle, 1994). Belemnites were restricted to the Euro- and a large taxon of the suborder Belemnitina. The Sinobelemnitidae, pean shelf till the Pliensbachian (Doyle, 1994; Schlegelmilch, 1998). In which may be included in the future in a new suborder, have also been contrast to this hypothesis, we describe here two belemnite taxa from the recorded from the Triassic of China, specimens so far poorly under- Hettangian of Japan: a new species of Sichuanobelus (family Sinobelem- stood. The presence of a very large rostrum attributed to the Belemni- nitidae) and an extremely large belemnite of the suborder Belemnitina. tina suggests in addition that a diverse belemnite fauna evolved earlier Based on these remarkable fi ndings, we provide here a major revision of than previously thought. Our new fi ndings therefore (1) extend the the early evolutionary history of belemnites, including their origin, early origin of the belemnites back by ~33 m.y. into the Triassic, (2) suggest phylogeny, and biogeography. that this group did not necessarily originate in northern Europe, and (3) imply that belemnites survived the Triassic–Jurassic extinction, GEOLOGICAL SETTING AND OCCURRENCE OF FOSSILS one of the fi ve big mass in the Phanerozoic. Since belem- Sediments of the Jurassic Shizugawa Group, which crop out along nites provided a considerable amount of food as prey, the origination the Pacifi c coast of northeastern Japan (Fig. 2), consist of the Nirano- of belemnites is probably an important event also for the evolution of hama and Hosoura Formations. The former has a middle to late Hettan- their predators, such as marine reptiles and sharks. gian age (199–196 Ma), based on the occurrence of the age-diagnostic ammonite Alsatites onoderai (e.g., Matsumoto, 1956; Takahashi, 1969; INTRODUCTION Sato and Westermann, 1991). The Sinemurian– age of the over- The Cephalopoda, which appeared in the , dominated the lying Hosoura Formation is based on a well-established ammonite bio- marine ecosystem for 500 m.y. until now (e.g., Kröger et al., 2011). Bel- stratigraphy (e.g., Takahashi, 1969; Sato and Westermann, 1991). A total emnites (order Belemnitida), a very successful group of Mesozoic cepha- of 60 specimens of Sichuanobelus utatsuensis sp. nov. (specimens UMUT lopods, were jet-propelled swimmers with ten arms like modern [University Museum, University of Tokyo] MM 30943–31002) and one (e.g., Stevens, 1965; Doyle et al., 1994) (Fig. 1). They held an important position, both as predators for smaller and as prey for marine rep- E140° E130° tiles and sharks (Doyle and Macdonald, 1993; Cicimurri and Everhart, N50° 2001; Rexfort and Mutterlose, 2006). Their diversity changes are well cor- related with Earth’s environmental shifts such as extinction events, sea- N N level changes, and climatic changes, thereby providing clues for under- N40° Study area Stomach Proostracum A Rostrum N30° Fossil locality Sedimentary rocks Gonads Gills Mantle Cretaceous Arm hooks Ink sac cavity Heart Dorsal Jurassic BCgroove Rostrum Triassic Mantle Fin muscles epithelium Pre-Permian Growth Fin cartilage rings Igneous rocks Cretaceous Ventral granitoids groove Hikami 10 km granitoids Figure 1. Reconstruction of the belemnite , based on Naef (1922), Stevens (1965), and our own observations. A: Figure 2. Locality of the Hettangian belemnites in north- Dorsoventral section (head in lateral view). B: Transverse eastern Japan. Geological map based on Shiino et al. section. C: Reconstruction. (2011). Broader lines indicate faults.

GEOLOGY, October 2012; v. 40; no. 10; p. 1–4; Data Repository item 2012257. | doi:10.1130/G33402.1 | Published online XX Month 2012. ©GEOLOGY 2012 Geological | October Society 2012 of America.| www.gsapubs.org For permission to copy, contact Copyright Permissions, GSA, or [email protected]. 1 Geology, published online on 9 August 2012 as doi:10.1130/G33402.1 specimen belonging to the suborder Belemnitina (UMUT MM 31003) The family Pseudodicoelitidae (suborder Belemnitina) and the families have been collected from the lower shoreface sandstone of the upper Dicoelitidae and Duvaliidae (suborder Pachybelemnopseina) have a dorsal part of the Niranohama Formation in the Niranohama harbor (Utatsu alveolar groove (Jeletzky, 1966, 1980; Riegraf et al., 1998; Schlegelmilch, area; 38°41′24.89″N, 141°30′13.53″E) (Fig. 2). We also investigated 1998). The Pseudodicoelitidae have an apical groove or apical striae in three belemnite specimens (UMUT MM 07078, 07079a, and 07079b), addition to the alveolar groove. The Dicoelitidae are characterized by two including one large form, collected from the present Niranohama Forma- alveolar grooves, a ventral and a shorter dorsal one, whereas the Duvaliidae tion (Yokoyama, 1904), and forgotten for a long time. All specimens are have only one dorsal groove (Jeletzky, 1966, 1980; Riegraf et al., 1998; deposited in the University Museum, University of Tokyo, Japan. Details Schlegelmilch, 1998). The Duvaliidae therefore share the diagnostic feature of the mode of occurrences of the belemnites are shown in Figure DR1 in (dorsal alveolar groove) with the Sinobelemnitidae. The phylogenetic rela- the GSA Data Repository1. tionship of the Duvaliidae, which appeared in the Middle Jurassic (Batho- nian, 168–165 Ma) and were common in low-latitude oceans until the Early SYSTEMATIC PALEONTOLOGY Cretaceous, has not yet been fully understood (e.g., Jeletzky, 1966; Schle- gelmilch, 1998). Our fi ndings suggest that the Duvaliidae are related to the Order: Belemnitida MacGillivray, 1840. Sinobelemnitidae, and that both families may form a new suborder within Suborder: uncertain. the Belemnitida, though more detailed studies are needed. Remarks: The order Belemnitida has been subdivided into two suborders Family: Sinobelemnitidae Zhu and Bian, 1984. based on the presence of apical grooves (Belemnitina) or an alveolar Genus: Sichuanobelus Zhu and Bian, 1984. groove (Pachybelemnopseina), which have been interpreted as a blood Type species: Sichuanobelus longmenshanensis Zhu and Bian, 1984. vessel attachment (e.g., Stevens, 1965; Jeletzky, 1966; Riegraf et al., Sichuanobelus utatsuensis sp. nov. (Figs. 3A–3J). 1998). The Hettangian specimens of the Sinobelemnitidae described Type locality and horizon: upper part of the Niranohama Formation in here have medium-sized rostra with a dorsal alveolar groove lacking a the Niranohama harbor, Utatsu, northeastern Japan. ventral one, like the Triassic material from China (Zhu and Bian, 1984). Material: 62 specimens (UMUT MM 07079a, 07079b, 30943–31002). These features are unknown from the Pachybelemnopseina, which have a Holotype: UMUT MM 30943. ventral alveolar groove. Belemnite rostra morphologically similar to the Paratypes: UMUT MM 30944–30947. Hettangian material have also been recovered from the Hosoura Formation Diagnosis: Rostrum is of medium size with a single alveolar groove on (Sinemurian–Aalenian). These specimens are, however, weathered and the dorsal side. Rostrum is laterally compressed and its lateral sides are preserved as external molds; the dorsoventral position of the groove can slightly fl attened. There are no lateral lines. Groove is deep and V-shaped. therefore not be confi rmed at the moment. The Sinobelemnitidae may Etymology: from the local name of the study area. therefore represent a separate group (suborder?) within the Belemnitida, Description: Rostrum is of medium size (Figs. 3A–3J). Lengths of the characterized by a deep dorsal alveolar groove. completely preserved rostra are 35.3–45.0 mm (Figs. 3A, 3C, 3G, and 3H).

Figure 3. Hettangian bel- emnites from Japan in comparison with north Europe species. A–J: Si- chuanobelus utatsuensis sp. nov. from Japan. A: Holotype, UMUT (Univer- sity Museum, University of Tokyo) MM 30943. A1: Dorsal view. A2: Lateral view. B: Paratype, UMUT MM 30944. B1: Dorsal view. B2: Lateral view. C: Paratype, UMUT MM 30945, dorsal view. D: UMUT MM 07079b. D1: Dorsal view. D2: Lateral view. D3: Ventral view. E: UMUT MM 30948, dorsal view. F: UMUT MM 30971. F1: Dorsal view. F2: Lat- eral view. F3: Ventral view. G: Paratype, UMUT MM 30946, longitudinal section. H: UMUT MM 30985, longitudinal sec- tion. I: UMUT MM 30978, transverse section. J: Paratype, UMUT MM 30947. J1: Transverse section. J2: Siphuncle (arrow). K and L: Extremely large Belemnitina from Japan. K: UMUT MM 31003. K1: Transverse section. K2: Siphuncle (arrow). L: UMUT MM 07078. L1: Transverse section. L2: Lateral view. L3: Longitudinal section. M and N: Hettan- gian belemnites from Belgium (Weis and Delsate, 2006). M: Schwegleria feifeli, MNHNL (Musée national d’histoire naturelle, Luxembourg) QB268a. M1: Ventral view. M2: Transverse section. N: aff. Subhastites sp., MNHNL HE366. N1: Lateral view. N2: Transverse section. A–C, E, F, and H are silicone rubber casts. One-centimeter scale bar for all specimens except for J2 and K2.

1GSA Data Repository item 2012257, photographs of mode of occurrences of belemnites, is available online at www.geosociety.org/pubs/ft2012.htm, or on request from [email protected] or Documents Secretary, GSA, P.O. Box 9140, Boulder, CO 80301, USA.

2 www.gsapubs.org | October 2012 | GEOLOGY Geology, published online on 9 August 2012 as doi:10.1130/G33402.1

Maximum length is estimated at >60 mm based on fragments. Maximum diameter of rostra is located in the alveolar region. Maximum dorsoventral Northern Europe diameter (Dv) and lateral diameter (Dl) of the paratype (Fig. 3B) are 15.0 China North and 12.6 mm, respectively. Dv/Dl is 1.14–1.19. Outline is symmetrical. America ▲ Japan Profi le is slightly asymmetrical (Fig. 3A). Apex is moderately acute (Figs. 3A, 3C, 3E, and 3F). Transverse section is laterally compressed (Figs. 3I and 3J). Lateral side is slightly fl attened (Figs. 3I and 3J). The Africa phragmocone extends over half the length of the rostrum (Figs. 3G and 3H). Apical angle of phragmocone is ~30° (Fig. 3G). Dorsal groove is Belemnitina (earliest Jurassic) long (Figs. 3A, 3C, 3E, and 3F), and starts at the alveolar border, and Sinobelemnitidae (earliest Jurassic) extends toward the apex without reaching it (Figs. 3C–3F). The groove ▲ Sinobelemnitidae (LateTriassic) is very deep and V-shaped (Figs. 3I and 3J), and underlain by a clearly defi ned splitting surface (Figs. 3I and 3J). There are no lateral lines, no Figure 4. Global distributions of belemnites in the early phase of apical grooves, nor an alveolar ventral groove. their evolution. Paleomap (200 Ma) based on Blakey (2010). Comparison: Sichuanobelus includes two other species, S. long men- shanensis and S. yangi, both from the () of China (Zhu and Bian, 1984). S. longmenshanensis has a slender rostrum with a Sinobelemnitidae, described by Zhu and Bian (1984) from the lower part relatively short and shallow dorsal groove; its lateral surface is rounded of Ma’atang Formation in the Longmen Mountains region, China. The in the apical region. These features are missing in S. utatsuensis. S. Chinese sinobelemnitids are well preserved and co-occur with ammonites utatsuensis differs from S. yangi by its shorter rostrum with a very deep indicating a Carnian age such as Protrachyceras, Discotropites, Tropites, groove, and a longer phragmocone. and Paratropites (Zhu and Bian, 1984; Gou, 1993). The Sinobelemnitidae can be clearly distinguished from the by showing the diag- Suborder: Belemnitina MacGillivray, 1840. nostic features of the Belemnitida: (1) a calcitic rostrum with concentric Genus and species: indeterminate (Figs. 3K and 3L). growth line, and (2) a high apical angle of phragmocone (>12°). Previous Material: UMUT MM 07078, 31003. studies (e.g., Doyle, 1993, 1994; Doyle et al., 1994) also considered the Description: Rostrum is extremely large and laterally compressed Chinese Sinobelemnitidae as belemnites s.s. (order Belemnitida). At the (Figs. 3K and 3L). Dv and Dl in the alveolar region are 33.1 and 28.5 mm, same time, the Triassic age assignment of these forms has been questioned respectively (Fig. 3K), and near the protoconch (Fig. 3L) are estimated at (Doyle, 1993, 1994; Weis and Delsate, 2006) because the typical belem- 26.4 and 20.9 mm, respectively. Dv/Dl in the alveolar region and near the nite morphology (long rostra with a well-developed alveolar groove), protoconch are 1.16 and 1.26, respectively. The protoconch is located at superfi cially exposed by the Sinobelemnitidae, had not been recorded the center of rostrum. Apical angle of phragmocone is ~30°. Concentrated from pre–Middle Jurassic strata in Europe (Doyle, 1993, 1994; Doyle growth lines can be observed in the transverse section of rostra. Dorsal, et al., 1994). The Sinobelemnitidae have a long belemnite rostrum with, ventral, and lateral sides are rounded. No alveolar groove or lateral lines however, one well-developed deep groove on the dorsal side. This dorsal have been observed (Figs. 3K and 3L). groove, which is only present in the Japanese Hettangian belemnites, is Comparison: The absence of an alveolar groove indicates an affi liation a morphological feature that has been overlooked in previous European to the Belemnitina. A detailed taxonomic assignment is currently not studies. The presence of Sichuanobelus in the Hettangian of Japan indi- possible due to the fragmental nature of the specimen. cates that the Sinobelemnitidae are part of a distinctive belemnite lineage (new suborder?), which fl ourished throughout the Late Triassic to earliest DISCUSSION AND CONCLUSIONS Jurassic in the eastern Tethys and western Panthalassa (Fig. 4). The fi ndings of Hettangian belemnites from Japan request a revision The fi nding of large rostra of the Belemnitina from the Hettangian of of the early evolutionary history of belemnites. Current hypotheses of the Japan indicates that this suborder already occurred in the earliest Jurassic early evolution of belemnites are summarized as follows. The suborder with well-developed large rostra (Fig. 4). Size and morphology of the Jap- Belemnitina fi rst appeared in the Hettangian, while the suborder Pachy- anese Hettangian Belemnitina are completely different from those of the belemnopseina has not been documented before the Middle Jurassic coeval European genera Schwegleria and aff. Subhastites (Figs. 3K–3N). (Doyle, 1993, 1994; Riegraf et al., 1998). Two genera of the Belemnitina, These European genera are characterized by very short and small rostra Schwegleria and aff. Subhastites, which are characterized by small and (Figs. 3M and 3N) (Doyle, 1994; Schlegelmilch, 1998; Weis and Delsate, short rostra, occur in the Hettangian of Belgium, southwest Germany, and 2006). The diameter and length of the rostrum of Schwegleria are ~4 mm northern Ireland (Doyle, 1994; Schlegelmilch, 1998; Weis and Delsate, and 10 mm, respectively (Weis and Delsate, 2006) (Fig. 3M), whereas 2006) (Figs. 3M, 3N, and 4). These are the oldest belemnites so far known, the rostrum of the Japanese Belemnitina is 33 mm in diameter (Fig. 3K). and Schwegleria (Fig. 3M) was considered as a possible stockgroup of This large diameter can be compared with that of Megateuthis (Middle all belemnites (e.g., Doyle, 1994; Weis and Delsate, 2006). Occurrences Jurassic), the largest belemnite (up to 40 cm in length) ever observed (e.g., of Hettangian proostraca and arm hooks of belemnites are also known Schlegelmilch, 1998). The fi ndings of the Japanese specimens in the earli- from northern Europe, southern England, southwest Germany, and Lux- est Jurassic imply an origin of the Belemnitina in pre-Jurassic time. embourg (e.g., Weis and Delsate, 2006; Klug and Fuchs, 2010). It has It is concluded that the earliest Jurassic Belemnitida had a much been concluded that the belemnites evolved in Europe as small forms in higher diversity than previously thought, including large taxa. The Belem- the Hettangian; their distribution was restricted to the European shelf seas nitida did not necessarily originate in northern Europe, contrary to previ- until the Pliensbachian for 18 m.y. (Doyle, 1994). They expanded their ous hypotheses. The Sinobelemnitidae can be considered as a possible distribution worldwide in the (183–176 Ma). rootstock of all belemnites; the small European belemnites of the earliest Findings of pre-Jurassic “belemnites” (e.g., Jeletzkya from the Car- Jurassic are here seen as an endemic offshoot. It is postulated here that boniferous of North America, Palaeobelemnopsidae from the Permian of the belemnites did not originate in the earliest Jurassic, but in the Late China, Sinobelemnitidae from the Triassic of China) challenged this view, Triassic. The fossil record of the order Belemnitida is therefore extended but were reclassifi ed later mostly as aulacocerid or phragmoteuthid cole- by ~33 m.y. before the Triassic-Jurassic boundary, where one of the fi ve oids (Doyle, 1993, 1994; Doyle et al., 1994). The only exceptions are the largest mass extinctions in the Phanerozoic occurred. The Prototeuthina,

GEOLOGY | October 2012 | www.gsapubs.org 3 Geology, published online on 9 August 2012 as doi:10.1130/G33402.1 the fi rst -bearing vampyropods, appeared in the Late Triassic Jeletzky, J.A., 1966, Comparative morphology, phylogeny, and classifi cation of (Fuchs and Larson, 2011a). The order , an extinct lin- fossil : University of Kansas Paleontological Contributions, Mol- eage related to modern eight-armed cephalopods (e.g., ), also lusca, v. 7, 162 p. Jeletzky, J.A., 1980, Dicoelitid belemnites from the Toarcian–middle of radiated in the Late Triassic (Fuchs and Larson, 2011b). The Late Trias- western and Arctic Canada: Geological Survey of Canada Bulletin, no. 338, sic can therefore be considered as a key period for the early evolution of p. 1–71. Mesozoic–Cenozoic coleoid cephalopods. Since belemnites provided a Klug, C., and Fuchs, D., 2010, An earliest Hettangian (Jurassic) belemnitid from considerable amount of food as prey for marine reptiles and sharks (Doyle Great Britain with a preserved proostracum, in Tanabe, K., et al., eds., Ceph- alopods—Present and past: Tokyo, Tokai University Press, p. 181–185. and Macdonald, 1993; Cicimurri and Everhart, 2001), the origination of Kröger, B., Vinther, J., and Fuchs, D., 2011, origin and evolution: Mesozoic coleoid cephalopods is probably an important event also for the A congruent picture emerging from fossils, development and molecules: evolution of their predators. Such views of the Triassic marine revolution BioEssays, v. 33, p. 602–613, doi:10.1002/bies.201100001. would contribute to reconstructing the evolution of the Mesozoic marine MacGillivray, W., 1840, A manual of geology: London, Scott, Webster and Geary, ecosystem in the future. 239 p. Matsumoto, T., 1956, Yebisites, a new Lower Jurassic ammonite from Japan: Palaeontological Society of Japan Transactions and Proceedings, no. 31, ACKNOWLEDGMENTS p. 205–212. We would like to thank the people in Niranohama, who were suffered from Mutterlose, J., 1998, The Barremian– turnover of biota in northwestern the 2011 Tohoku earthquake and tsunami, for their help during our fi eld work be- Europe: Evidence from belemnites: Palaeogeography, Palaeoclimatology, fore and also after the disaster. We thank N. Mariotti for discussions, and R. Weis, Palaeoecology, v. 144, p. 161–173, doi:10.1016/S0031-0182(98)00081-9. T. Sasaki, and Y. Ito for help in observing fossil collection in their institution. We Naef, A., 1922, Die fossilen tintenfi sche: Stuttgart, Gustav Fisher, 322 p. also thank K. Sugawara, Y. Kashiyama, and T. Miura for fi eld assistance. This re- Rexfort, A., and Mutterlose, J., 2006, Stable isotope records from offi cina- search was fi nancially supported by the Japan Society for the Promotion of Science lis—A key to understanding the ecology of belemnites?: Earth and Plan- (grant 23840002 to Iba), the German Research Foundation (grant Mu 667/40-1 to etary Science Letters, v. 247, p. 212–221, doi:10.1016/j.epsl.2006.04.025. Mutterlose), and the Researcher Overseas Visit Program of Hokkaido University of Riegraf, W., Janssen, N., and Schmitt-Riegraf, C., 1998, Cephalopoda dibranchi- Education (in 2011 for Iba). ate fossils (Coleoidea) II, in Westphal, F., ed., Fossilium catalogus animalia, Pars 135: Leiden, Backhuys Publishers, p. 5–512. 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Gou, Z., 1993, Bivalve fauna of Upper Triassic in Maatang area, Jiangyou, Sich- uan: Acta Palaeontologica Sinica, v. 32, p. 13–30. Manuscript received 11 March 2012 Iba, Y., Mutterlose, J., Tanabe, K., Sano, S., Misaki, A., and Terabe, K., 2011, Revised manuscript received 16 April 2012 Belemnite extinction and the origin of modern cephalopods 35 m.y. prior to Manuscript accepted 17 April 2012 the Cretaceous–Paleogene event: Geology, v. 39, p. 483–486, doi:10.1130 /G31724.1. Printed in USA

4 www.gsapubs.org | October 2012 | GEOLOGY GSA DATA REPOSITORY 

Supplementary material for:

Belemnites (Cephalopoda) originated in the Triassic – a new look at an old group

Yasuhiro Iba1, Shin-ichi Sano2, Jörg Mutterlose3, and Yasuo Kondo4 1Hokkaido University of Education, Kushiro 085-8580, Japan 2Fukui Prefectural Dinosaur Museum, Fukui 911-8601, Japan 3Department of Geology, Mineralogy and Geophysics, Ruhr-University Bochum 44801, Germany 4Department of Natural Environmental Science, Kochi University, Kochi 780-8520, Japan

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