THE VELIGER © CMS, Inc., 2007 The Veliger 50(4):255-262 (December 16, 2008) A New Genus for Vesicomya inflata Kanie & Nishida, a Lucinid Shell Convergent with that of Vesicomyids, from Cretaceous Strata of Hokkaido, Japan KAZUTAKA AMANO Department ofGeoscience, Joetsu University ofEducation, Joetsu 943-8512, Japan (e-mail: [email protected]) ROBERT G. JENKINS Faculty ofEducation and Human Sciences, Yokohama National University, Yokohama 240-8501, Japan YUKITO KURIHARA Department of Geology and Paleontology, National Museum ofNature and Science, Tokyo 169-0073, Japan AND STEFFEN KIEL Institute ofGeosciences - Paleontology, Christian-Albrechts-University, 24118 Kiel, Germany Abstract. Newly collected specimens of the large bivalve Vesicomya inflata Kanie & Nishida from the lower Cenomanian Tenkaritoge Formation reveal that it is not a vesicomyid but is instead an unusual lucinid. The new monotypic genus Ezolucina is herein proposed for this species, which is characterized by venerid or vesicomyid shell shape, large size, a smooth surface, adeeply impressedlunule, one cardinal and one anteriorlateral tooth in the right valve, and two cardinal teeth in the left valve. Stable carbon isotope analyses and petrographic observations show that the carbonate concretions yielding this species do not represent ancient hydrocarbon-seep deposits as was suggested previously. Rather, Ezolucina inflata and the associated solemyid, lucinid, thyasirid, and manzanellid bivalves lived in organic- and sulfide-rich sediment. INTRODUCTION and belong to a clade informally known as "small" vesicomyids, composed of the genera Vesicomya, Large fossil bivalves fromthe "MiddleYezo Group" in Waisiuconcha, Isorropodon, Callogonia and Pliocardia northern Hokkaido, Japan, were described by Kanie & (cf. Cosel & Salas, 2001; Krylova & Sahling, 2006). In Nishida (2000) as Vesicomya inflata, and listed as being contrast, the Cretaceous ""Vesicomya" inflata reaches a the earliest record for the genus Vesicomya (Campbell, length of up to 157 mm. In a recent revision of fossil 2006; Kiel & Little, 2006). They were found in two North Pacific vesicomyids, Amano & Kiel (2007) large calcareous concretions surrounded by mudstone, pointed out that V. inflata has a deeply impressed and were associated with the solemyid Acharax asymmetrical lunule, a feature unknown in vesico- cretacea Kanie & Nishida, 2000 and the lucinid Miltha myids, and that its hinge structure had neither been sp. Extant members of these bivalve taxa harbor described nor illustrated. Consequently, Amano & Kiel chemoautotrophic endosymbionts, and because car- (2007) excluded V. inflata from the Vesicomyidae and bonate concretions bearing these taxa have repeatedly suggested lucinid affinities instead. been demonstrated to represent ancient hydrocarbon- Newly collected specimens from the type locality of seepdeposits(Majima et al., 2005; Campbell, 2006), the ""Vesicomya" inflata at Sanjussen-zawa Creek in concretions bearing Vesicomya inflata were interpreted northern Hokkaido possess hinge dentition, a pallial & as ancient hydrocarbon-seep deposits (Kanie line, and adductor scarsthat clearlyplace this species in Nishida, 2000; Kanie et al., 2000). anewgenus ofthe Lucinidae. In addition, petrographic Living Vesicomya species have small shells that thin sections and stable carbon isotope analyses ofthe mm & rarely exceed 13 in length (Cosel Salas, 2001) fossil-bearing concretions are used to evaluate the Page 256 The Veliger, Vol. 50, No. 4 Nakagawa area y&JHorokanai area vo Tappu area Figurf Type locality ofEzolucina inflata (Kanie & Nishida). environmental reconstruction ofthis site as an ancient trometer at EPUT. Carbon isotopic composition is hydrocarbon seep. expressed in the conventional 8 notation relative to the Vienna Peedee Belemnite standard (S13C, %o vs. V- MATERIALS and METHODS PDB, standard deviation <0.03%o). The type material was examined at the Yokosuka City MINERALOGY AND ISOTOPE COMPOSI- Museum, and nine new specimens of "Vesicomya" TION OF CARBONATES inflata were collected from four float carbonate concretions with molluscan fossils (HRK A-D) at the Thin-section observation and XRD analysis show that type locality in northern Hokkaido (Figure 1). Strati- the concretions are almost entirely composed of graphically the specimens are from the My 4 Member homogeneous micritic calcite and siliciclastic sediment. of the Tenkaritoge Formation, which is considered Structures typical for methane-induced carbonates, like early Cenomanian (Hashimoto et al., 1965; Nishida et clotted fabrics and stromatolitic laminae (cf. Greinert al., 1998). The figured specimens and additional new et al., 2002; Peckmann & Thiel, 2004), were not seen. material are housed at the Joetsu University of 813C values range from -7.7 to -6.3%o (vs. V-PDB) Education (JUE). The mineralogy ofthe fossil-bearingconcretions was Table identified by thin-section observations and X-ray 1 diffraction (XRD) analysis. Standard thin-section Stable carbon and oxygen isotope composition of the observations were performed by plane- and cross- carbonate concretions from the type locality of polarized and reflected light microscopy. XRD analy- Ezolucina inflata (Kanie & Nishida). ses were carried out on unoriented slurries using a PANalytical X'Pert PRO at the Department of Earth Sample no. Texture Mineralogy 8l3C 5,8 and Planetary Science, the University of Tokyo HRK A micrite calcite -7.4 -1.3 (EPUT). Carbon and oxygen isotopes were analyzed HRK A micrite calcite -7.3 -1.2 using 2 to 10 mg powdered carbonate matrix. Carbon HHRRKKCB micrite calcite -6.3 -3.8 dioxide was produced from each powdered sample by HRKC micrite calcite -6.7 -4.9 micrite calcite -6.5 -4.2 reaction with 100% phosphoric acid in vacuo (25°C), HRK C micrite calcite -7.7 -5.2 and analyzed with a Finnigan MAT252 mass spec- K. Amano et al., 2007 Page 257 (Table 1) and are also not indicative of anaerobic Paratype: Articulated specimen, length 82.6 mm+, methane oxidation. Methane-derivedcarbonateusually height 59.8 mm, width 35.8 mm+, YCM-GP1174. shows 813C values lower than —40%o (cf. Peckmann & Topotypes: Articulated specimen, length 157.5 mm, Thiel, 2004). Thus, neither thin section observations nor stable isotope analyses support the idea ofKanie& height 123.8 mm, width 74.0 mm+, YCM-GP1177; Nishida (2000) and Kanie et al. (2000) that "Vesico- right valve, length 33.7 mm, height 26.2 mm, JUE mya" inflata and associated mollusks lived at an no. 15853; right valve, length 28.9 mm, height ancient hydrocarbon seep. 21.9 mm, JUE no. 15854; articulated specimen, length 15.4 mm, height 13.5 mm, width 5.9 mm, JUE SYSTEMATICS no. 15855. Type locality: Bed of Sanjussen-zawa Creek, 6.5 km Family Lucinidae Fleming, 1828 upstream from where it flows into the Uryu River, Genus Ezolucina Amano, Jenkins, Kurihara & HorokanaiTown, Hokkaido (44°14'24"N, 142°5'26"E); Kiel, gen. nov. My 4 Member ofthe Tenkaritoge Formation (locality R7203 by Nishida et al., 1998). Type species: Vesicomya inflata Kanie & Nishida, 2000. Stratigraphic and geographic distribution: Late Creta- ceous (early Cenomanian); known only from the type Diagnosis: Large, inflated veneriform shell with smooth locality. surface except for rough, low commarginal lamellae; posterior radial sulcus weak, and lunule deeply Original description: "Large shell (length [L] L = impressed. Hinge ofright valve with one stout cardinal 130 mm in holotype and probably gerontic stage) tooth and an anterior lateral tooth, left valve hinge probably of rounded triangular form (height [H] H/L with two cardinal teeth. Anterior adductor scar = 0.8). Umbo (shell apex) situated almost centrally, quadrate and anteriorly detached from pallial line; 45^46% from anterior margin. Strongly inflated valve pallia] line entire and deeply impressed. ([H]/breadth [B], H/B^0.68). Shells ofright and left are Discussion: Here Gabb, 1866 is similar to Ezolucina etoqumiviadldvleed.grPoowsttehrisotrageenrdepisretsreunntceadtebdy. pIanrtahteypjeuv(eLnil=e lgaetne.ranlovt.oobtyh hianvirnigghtonvealcvaerdainnadl atoodtehe,ployneimapnrteesrsieodr 88.6 mm), the shell height is shortened (H/L = 0.68). Postero-dorsal end is truncated meeting with the lwuintuhle,finbeutvedinftfrearls fcrreonmulEaztioolnusc,inaanbdy iittss smmualclhermsoherlel pasosgteerroinotricmaorngei.n.TheLruenuilseaalunndulheinagteanftoerrmo-disortshaelspaarmte. deeply impressed lunule. Another large Cretaceous lucinid is Nipponothracia, which differs from Ezolucina Tprhoebabvlenyterxatlernmaalrgainnd loisngwoenaktlheybaasricshoefd.morLpihgoalmoegnyt by having an edentulous hinge and a very elongate anterior adductor scar (cf. Kanie & Sakai, 1997; Kase of the hinge amrema at postero-dorsal part. Test is very et al., 2007; Kiel et al., 2008). The medium-sized North tohrincakmaebnotuetd7by coantctehnetrviecntrgarlomwatrhginli.neSsh.elTlhseurfuamcebois American lucinid Nymphalucina Speden, 1970 from the bends strongly inside, where characteristic subumbonal Late Cretaceous Pierre Shale and Fox Hill Formation pit exists at the inside ofthe shell apex." is oval in shape and lacks the sloping posterodorsal margin of Ezolucina. Trinitasia Maury, 1928, which Supplementary description: Shell large in size (up to was questionably placed in Lucinidae by Chavan 157.5 mm in length), thick, well inflated in adult (1969), is comparable in shell form and sculpture, but specimens, veneriform, rather equivalve, slightly in- based on internal shell features Woodring (1982) equilateral. Small specimens less inflated and Fela- showed that Trinitasia is not a lucinid but a mactrid. niella-like in shape. Umbo projecting above dorsal Etymology: A combination of the old name of margin, prosogyrate, situated anteriorly at two-fifths of shell length. Anterodorsal margin broadly arcuate, Hokkaido (Ezo) and the genus Lucina. grading into rounded anterior margin; ventral margin Ezolucina inflata (Kanie & Nishida, 2000) broadly arcuate; posterodorsal margin straight, grad- ually sloping, forming blunt angle with subtruncated (Figures 2-9) posteriormargin. Surface smooth except for rough and Vesicomya inflata Kanie & Nishida, 2000: p. 79-82, vlioswiblecoimnmarrigghitnavlalvleameolflaae;smcaolalrssepeccoinmceenn.trSihcalrliobws .figs. 1, 2. radial sulcus running from beak to posterior end in large specimens. Holotype: Articulated specimen, length 131.8 mm, Hinge plate wide; right valve with one stout and height 105.4 mm, width76.3 mm, YCM-GP1173. triangular central tooth (3b), one elongate anterior Page 258 The Veliger, Vol. 50, No. 4 Figures 2^4. Type material of Ezolucina inflata (Kanie & Nishida). Figure 2. Holotype, length = 131.8 mm, YCM-GP1173. Figure2a. Lateralviewofrightvalve. Figure2b. Dorsalviewshowinga posteriorradialsulcusanddeeplyimpressedlunule. Figure 3. Topotype, length = 157.5 mm+, YCM-GP1177. Figure 3a. Dorsalview. Figure3b. Lateralviewofleftvalve. Figure4. Paratype, length = 82.6 mm, YCM-GP1174. Figure4a. Lateral view ofright valve. Figure4b. Dorsal viewshowinga posteriorradial sulcus and deeply impressed lunule. Figure 4c. Lateral view ofleft valve. K. Amano et al., 2007 Page 259 Figures 5-8. Additional specimens of Ezolucina inflata (Kanie & Nishida). Figure 5. Topotype, length = 28.9 mm, JUE no. 15854. Figure 5a. Hinge ofright valve showing one strong cardinal and distinct anterior lateral tooth, length ofhinge plate = 21.3 mm. Figure5b. Dorsalviewshowingdeeplydepressedlunule. Figure5c. Lateralviewofrightvalve. Figure6. Topotype, length = 39.9 mm+,JUEno. 15856. Figure6a. Dorsalviewofleftvalveshowingdeeplyimpressed lunule. 6b. Hingeofleftvalveshowing two strongcardinal teeth, length ofhinge illustrated here = 21.8 mm. 6c. Lateral view ofleft valve. Figure 7. Topotype, length = 15.4 mm,JUEno. 15855. Figure7a. Lateralviewofleftvalve. Figure7b. Lateralviewofrightvalve havingrudeconcentric ridges. Figure 8. Topotype, length = 33.7 mm, JUE no. 15853. Figure 8a. Lateral view ofright valve stressing anterior adductor scar. Figure 8b. Normal lateral view ofright valve. lateral tooth (AI) parallel to hinge base, and a weak developed, entire, and quite distant from shell margin. blunt node just below deeply impressed lunule; left Inner ventral margin smooth. valve with strong anteriorcardinal (2), touching deeply impressed lunule, posterior tooth (4b) rather thin. Remarks: Very similar to Ezolucina inflata in shell Ligament occupying two-thirds of posterodorsal mar- shape and size is 'Lucina' colusaensis Stanton, 1895, a gin. Anterior adductor scar elongate quadrate, moder- species that is apparently restricted to Upper Jurassic ate in size (adductor length = 9.1 mm in JUE (Tithonian) to Lower Cretaceous (Hauterivian) cold- no. 15853; adductor length/shell length = 0.27) and seep carbonates in northern California, USA (Stanton, a(bmoauxtim7u5m%diosftaintcselfernogmthpadlleitaalclhiende =fr1o.m9 mpamlliianlJlUinEe 21080985),.p.Co60m,ppair.e1d1,tfoigso.u4r, 5m;atCearmipablelolf,E2z0o0l6u;ciKniaelientflaalt.a, no. 15853); posterior adductor scar pear-shaped. Inner the thickness of 'Lucina' colusaensis resembles that of surface covered by coarse and distinct radial striations the less inflated specimens of Ezolucina inflata. (Figure 9). Lunule broadly lanceolate, well defined, Unfortunately, 'Lucina" colusaensis is usually poorly deeply impressed, slightly asymmetrical, slightly larger preserved and features of the interior of this shell are in right valve than in left valve, and occupying one- unknown (Stanton, 1895; SK, pers. observation; K.A. third ofanterodorsal margin. Pallial line narrow, well- Campbell, personal communication 2007); thus a Page 260 The Veliger, Vol. 50, No. 4 Figure9. Musclescarsand palliallineofEzolucinain/Jata(Kanie&Nishida)basedonJUEno. 15853.Abbreviations: aa, anterior adductor scar; pa, posterior adductor scar; pi, pallial line. confirmation of its generic position must await the p. 161. pi. 14: 5) from a potential Hauterivian seep site availability ofwell-preserved specimens. in eastern Czech Republic lacks the strongly sloping Another large lucinid with prominent umbos is posterodorsal margin ofEzolucina inflata. Saxolucina (Megaxinus) matsushitai Matsumoto (1971, p. 663-665, pi. 1, fig. 1, pi. 2, figs. 1-3) from DISCUSSION the Oligocene Setogawa Group [now considered as Miocene in age; see Watanabe (1988)] in central Japan, Examination of the type and additional specimens but this species is clearly distinct from Ezolucina inflata clearly shows that Vesicomya inflata is a member of by having a less inflated shell with an edentulous Lucinidae because it has the hinge structure and the dentition. adductor muscle scarand pallial line patterns typical of Ezolucina inflata resembles Here excavata (Carpen- Lucinidae (i.e., a lucinoid hinge dentition, a broadly ter, 1857) in having a deeply depressed lunule, one lanceolate, asymmetric, sunken lunule, and an elongate cardinal tooth and anterior lateral tooth in the right anterioradductor muscle scardetached from thepallial valve, and twocardinal teeth in the leftvalve. However, line). The previous assignment of this species to the Here excavata differs from Ezolucina inflata by its Vesicomyidae was due to the superficial resemblance in subcircular shell with concentric lamellae, many fine shell outline and the lack of the information on the ventralcrenulations and more deeply sunken lunule. Its shell interior. The new monotypic genus Ezolucina veneriform shape with the strongly sloping postero- based on V. inflata is more elongate than most lucinids dorsal margin sets Ezolucina inflata apart from most and has moderately prominent umbones somewhat other Cretaceous lucinids, which mostly have a nearly suggestive of a vesicomyid or an eomiodontid. How- round outline (e.g., Discoloripes septentrionalis Kelly, ever, shells ofthese families can easily be distinguished 1992; "Lucina" spp. in Stephenson, 1952; Callucina from those ofthe Lucinidae by their hinge and muscle olea Vokes, 1946; "Lucina" aquensis Holzapfel, 1889, scar patterns. A p. 188, pi. 19, fig. 4). lucinid with similar hinge In their compilation of the stratigraphic ranges of dentition was described and figured as Lucina sub- mollusks at cold seeps, Kiel & Little (2006) listed V. nummismalis d'Orbigny, 1850 from the Campanian inflata as the oldest fossil occurrence of Vesicomya, VaalsGreensand ofGermany (Holzapfel, 1889, p. 187— based on the available literature (Kanie & Nishida, 8. pi. 19. figs. 1-3). This species is distinct from 2000). When Amano & Kiel (2007) subsequently Ezolucina inflata because it is very flat, has distinct revised the North Pacific fossil record of the Vesico- commarginal ribs, and lacks the strongly sloping myidae, they could not confirm any of the previous posterodorsal margin of Ezolucina inflata. Also "Luci- records of this genus. One of the oldest "small" na aff. valentula de Lor." described by Ascher (1906, vesicomyids related to Vesicomya (cf. Cosel & Salas, K. Amano et al., 2007 Page 261 deep-water environments of the Late Cretaceous were Table 2 inhabited by a number ofexceptionally large taxa like Faunal list based on fossils that we collected from the bivalve Inoceramus or the capulid limpet Gigantocapu- type locality ofEzolucina inflata (Kanie & Nishida). All lus giganteus (cf. Takahashi et al., 2007). Thus, the of these species possibly harbor symbiotic bacteria. paleoenvironment of Ezolucina inflata remains enig- Numbers indicate the number ofrecovered individuals, matic. capital letters are sample designations. Acknowledgments. We are very grateful to James L. Goedert Species Carbonate no. (HRK -) B C D (Burke Museum, Seattle) for his critical reading of this Acharax cretacea Kanie and Nishida 2 manuscript. We thanks John D. Taylor (Natural History ENzucoilnueclilnaa?insfpl.ata (Kanie and Nishida) 2 41 MDauvsies)umf)oratnhdeirGereevriaetw.J.WVeeramlesiojt(hUannikvserTsaimtyioofNiCaslhiifdoarn(iSaagaat Milthal sp. University) for his offering some fossil specimens. Thyasira sp. 1 LITERATURE CITED 2001; Krylova & Sahling, 2006) is "Vesicomya" Amanthoe,NKo.rt&hSP.acKiifeicl.re2g0i0o7n.. FVoeslsiiglerve4s9i:c2o7m0y-i2d93b.ivalves from kawadai (Aoki, 1954) from the lower Miocene Taira Aoki, S. 1954. Mollusca from the Miocene Kabeya Forma- Formation in Fukushima Prefecture, Japan (Aoki, tion, Joban coal-field, Fukushima Prefecture, Japan. 1954; Kamada, 1962), but more material is required to Science Reports of the Tokyo Kyoiku Daigaku, confirm its generic position. Another certain record of Section C 3:23-41. a "smair' vesicomyid is Isorropodon frankfortensis Ascher, E. 1906. Die Gastropoden, Bivalven und Brachio- Amano & Kiel, 2007 from the lower Miocene Astoria poden der Grodischter Schichten. Beitrage zur Palaonto- logie und Geologie Osterreich-Ungarns und des Orients Formation in Washington State, USA. 19:135-172. Kanie et al. (2000) suggested that the carbonate Bouchet, P. & R. Von Cosel. 2004. The world's largest concretions from Sanjussen-zawa Creek were ancient lucinid is an undescribed species from Taiwan (Mollusca: cold-seep deposits because of the presence of chemo- Bivalvia). Zoological Studies 43:704-711. symbiotic species like solemyids, vesicomyids, and Bretsky, S. S. 1976. Evolution and classification of the lucinids. Even when the vesicomyids are removed from Lucinidae(Mollusca; Bivalvia). PalaeontographicaAmer- icana 8:219-337. this list, most taxa from these concretions (see Table 2) Campbell, K. A. 2006. Hydrocarbon seep and hydrothermal rely at least partly on symbiotic sulfur-oxidizing vent paleoenvironments and paleontology: Past develop- bacteria. However, the isotopedata and the lithological ments and future research directions. Palaeogeography, observations presented here do not support a recon- Palaeoclimatology, Palaeoecology 232:362^107. struction as ancient cold-seep deposit. The data only Carpenter, P. P. 1857. Report on the present state of our suggests that these taxa lived in organic- and sulfide- knowledge with regard to the Mollusca ofthe west coast ofNorth America. British Association for the Advance- rich sediment favorable for species with sulphophilic ment ofScience, Report [for 1856], 159-368. symbionts. Three ofthe five taxa from the concretions Chavan, A. 1969. Superfamily Lucinacea Fleming, 1828. at Sanjussen-zawa Creek (Acharax, Thyasira, and Pp. N491-N518 in R. C. Moore (ed.), Treatise on Miltha) can also be found in the mudstone and InvertebratePaleontology. PartN,Vol. 2(of3), Mollusca siltstone of the Yezo Group (Tashiro, 2004) and their 6, Bivalvia. Geological Society ofAmerica and University relative abundance at this site might be due their early ofKansas Press: Lawrence, Kansas. Cosel, R. von. 2006. Taxonomy of tropical West African diagenetic preservation in the concretions. bivalves. VI. Remarks on Lucinidae (Mollusca, Bivalvia), The exceptionally large size of Ezolucina inflata is with description ofsix new genera and eight new species. remarkable; among Cretaceous lucinids it even exceeds Zoosystema 28:805-851. that ofthe cold-seep restricted Nipponothracia. Among Cosel, R. von. & C. Salas. 2001. Vesicomyidae (Mollusca: Recent species, it matches the size of Meganodontia Bivalvia) of the genera Vesicomya, Waisiuconcha, Issor- acetabulum Bouchet & Cosel, 2004, recently described opodon and Callogonia in the eastern Atlantic and the Mediterranean. Sarsia 86:333-366. Tashethespleacrigeesstwalisvifngoulnudciniindafnroamreaadoefptphreosfu2m5e6d-4d7if2fums.e FlemtihnegD,esJc.ri1p8t2i8v.eAChaHriascttoerrysoafndBrSiytsisthemAantiicmaallsA,rrEaxnhgibeimteinntg gas seepage and was associated with other bivalves oftheGeneraand SpeciesofQuadrupeds, Birds, Reptiles, bearing chemotrophic endosymbionts, like solemyids, Fishes, Mollusca and Radiata of the United Kingdom; thyasirids, and other lucinids. This set oftaxa is similar Including the Indigenous, Extirpated, and Extinct Kinds; to that associated with Ezolucina inflata, but our Togetherwith Periodical and OccasionalVisitants. Bell & cenavribroonnmiesnotto.peModsamttamodtohenromtoidnedircnatdeegeaps-wsaeteepragleuciinniidtss GabbCB,rraedtWfa.ucteeo:Mu.sEdain1n8db6u6Tr.egrhtP.iaalr5ey5o4nfotpsopsil.losg.ySecotf. C1a.l1i:fo1r-n3i8a., Vol. 2. rarely exceed 50 in length (Cosel, 2006) and are Greinert, J., G. Bohrmann & M. Elvert. 2002. Stromat- thus much smaller than Ezolucina inflata. 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