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Belemnites Originated in the Triassic—A New Look at an Old Group

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Belemnites Originated in the Triassic—A New Look at an Old Group Geology, published online on 9 August 2012 as doi:10.1130/G33402.1 Belemnites 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 Science, Kochi University, Kochi 780-8520, Japan ABSTRACT standing evolutionary dynamics of nektonic biota (e.g., Mutterlose, 1998; Belemnites (order Belemnitida), a very successful group of Christensen, 2002; Iba et al., 2011). Mesozoic cephalopods, 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 Jurassic to Cretaceous 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 (Hettangian, 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 genus Schwegleria (suborder Belemnitina) is considered as to northern Europe until the Pliensbachian (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 extinction 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 extinctions 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–Aalenian age of the over- The Cephalopoda, which appeared in the Cambrian, 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 squids [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 animals 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 Phragmocone Rostrum Siphuncle N30° Fossil locality Protoconch Sedimentary rocks Gonads Gills Mantle Mantle Cretaceous Arm hooks Ink sac cavity Heart Dorsal Jurassic BCgroove Rostrum Triassic Mantle Fin muscles epithelium Permian Pre-Permian Growth Fin cartilage rings Igneous rocks Cretaceous Ventral granitoids groove Hikami 10 km granitoids Figure 1. Reconstruction of the belemnite animal, 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
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