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Gastropoda:Muricidae The malacologicalsocietymalacological society of Japan Jeur. Malac,) fi re VENUS {Jap. - Vol, 5z No. 3 ( l99g): 2e9 223 Originand Biogeographic History of Ceratostoma (Gastropoda: Muricidae) Kazutaka AMANo and Geerat J. VERMEIJ of943ofGeoscience. Jbetsu Uhiversity oj' EZIucation. i2xmayashiki-1. Jbetsu, MigataDqpartmentPrellercture, -85i2 Japan, and Department of Geolegy and Center for Population Uitiversity Calijbrnia at Davis, One Shields Avenue, Davis, CA 95616 USABiology Abstract: We examined the two Neogene species of the ocenebrine muricid gastropod genus Ceratostoma Herrmannsen, 1846, from Japan and North Korea, namely, C. makiyamai The (Hatai & Kotaka, 1952) and C. sp,, both from the early middle Miecene. genus Ceratostotna is divisible into fouT groups based on C. nuttalli (Conrad), C. virginiae (Maury), C, fotiatum (Gemelin), and C. rorijIuum (Adams & Reeve). Three Miocene species from Kamehatka assigned by Russian workers to Ceratostoma are difficult to evaluate owing to poor preservation. Purpura turris Nomland, from the Pliecene of California, which was assigned to Ceratostoma by earlicr authors, is here tentatively assigned to Crassilabrum Jousseaume, 1880, have arisen the the early Miocene, it The genus Ceratostoma may in Atlantic. By had reached California, and then spTead westward to northeast Asia by earLy middle Mio- Miocene ccne time. This pattern of east to west expansion during the early half of the also characterizes many other north-temperate marine genera, including the ocenebrine NiiceUa. Keywords: origin, biogeography, Ceratostoma, Muricidae Introduction muricid Ceratostoma liye in shallow Recent species of the ocenebrine gastropod genus waters on both sides of the temperate and boreal North Pacific. There are six living species: C nuttaUi (Conrad) from California and Baja California, C. monoceros (Sowerby) C from Baja California, C. fotiatum (Gmelin) from Alaska to northern Baja California, burnetti (Adams & Reeve) in cool-temperate east Asia, and C, fournieri (Crosse) and C, rorijluum (Adams & Reeve) in warm- to mild-temperature east Asia (Fig, 1). C. fotiatum has been reported to feed on barnacles and bivalves (Spight & Lyons, 1974; Kent, 1981; Strathmann, 1987). The latter prey are apparently edge-drilled, and the long labral tooth 1981). of C, foliatum may be used to stabilize the predator during feeding (Kent, Pacific Hall (1959) reviewed the Recent species of Ceratostoma as well as northeastern fossil taxa, but he did not comment on Atlantic or northwestern Pacific fossil members of the genus. This is not surprising, for no species had been described under the generic name Ceratostoma from either region. with special emphasis Our prupose in this paper is to review the species of Ceratostoma, NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 210 VENUS: Vol. 57, No. 3 (1998) Fig. 1. Distribution of the Recent Cerzitostoma (after Habe & Ito, 1965; Abbott, l974. Radwin &D'Attilio, 1976; Baxter, 1984; Pain, 1990; Johnson, 1991; Higo & Goto. 1993); e VJVd tsONtsSa)temp tsSb-Ai. on northeast Asian fossil forms. We also reconstruct the biogeographic history of the group and compare it to that of other Neogene molluscan clades. The biogeographic study was prompted by the observation that ocenebrine genera show a diversity of histories despite the fact that most have a similar, nonplanktotrophic larval development, Thus, Ocinebrellus has remained confined to northeast Asia since the origin of this genus in the late middle Miocene (Amano & Vermeij, in preparation), whereas MiceUa originated in the northeastern Pacific during the early Miocene and then spread to Asia by the early middle Miocene and to the Atlantic during the Pliocene (Amano et aL, 1993; Collins et aL, 1996). Materials and Methods We examined fossil and Recent species at the following institutions: California Academy ofSciences, San Francisco; United States National Museum, Washington; Naturhistorisches Museum, Vienna (NMB); University of California Museum of Paleontology, Berkeley (UCMP); Tohoku University (IGCP); and Mizunami Fossil Museum (MFM). We measured or evaluated the following characters: shell height, spire height, length of siphonal canal, aperture height, number and shape of axial sculptural elements, number of spiral ¢ ords on last whorl, and number of denticles on adaxial (inner) side of outer lip. Description of Northwestern Pacific Fossil Species Family MURICIDAE Rafinesque, 1815 Subfamily Ocenebrinae Cossmann, 1903 Genus Cle]ratostoma Herrmannsen, 1846 NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan Amano & Vermeij: Origin and Biogeographic History of Ceratostoma 211 Fig. 2-3. Ceratostoma maklyamai (Hatai & Kotaka). 2, x1,2, IGPS no. 74368 (Holotype); 3, x 1.2, IGPS no. 74368; Loc. Paiponchon, Myonchon District, Hamukyon-pukuton, North Korea; Heiroku Fermation. x2, MFM, Loc. N64 of Itoigawa et aL Nataki Formation. Fig. 4. Ceratostoma sp. (1981); Fig. Ceratostoma x 1, UCMP12057 Loc. West of 5. peranguiatum (Nomland). (Holotype), Coalinga, California; Jacalitos Formation. Loc. West of Coalinga, Fig. 6. Purpura turris Nomland. x1, UCMP12049 (Holotype), California; Etchegoin Formation. Fig. 7. Ceratostoma nannum (Nomland). x1, UCMPI1315 (Holotype), Loc. North of Coalinga, California; Santa Margarita Formation. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 212 VENUS: Vol. S7, No.3(1998) bpes species: Murex nuttalli Conrad, 1837. Remarks: The genus Ctiratostoma usually has three (rarely four) varices per whorl, adjacent ones being separated by a singLe intervarical node. Typically, the varices are blade-like or rounded. Juvenile specimens have axial sculpture that is not differentiated into varices and intervarical nodes. They typically also lack the labral tooth and have an open siphonal canal. Specimens in which axial sculpture has become differentiated into varices and nodes are characterized by a labral tooth, which is formed as an adaxiar infolding of the outer lip near the latter's abapical end, The tooth apparently forms discontinuously on the adapertural side of each varix (see Vermeij, 1998, in press). Some species of Ceratostoma have smail denticles on the inner (adaxial) side of the outer lip, and a very small parietal tooth near the uppermost part of the inner lip, In the adult shell, the siphonal canal is ventrally sealed. Several genera resemble Ceratostoma in having a trivaricate shell with a labral tooth. Microrhytis Emerson. 1959, from the Miocene of tropical America, differs from Ceratostoma in having small lamellate varices on the early whorls instead of cancellate sculpture, and by having a strong keel at the shoulder connecting adjacent varices (Vermeij & Vokes, 1997). The Pliocene South African genus Namamurex Carrington and Kensley, 1969, differs from Ceratostoma by having obsolete spiral sculpture other than a strong peripheral keel con- necting the varices, and by having the siphonal canal open instead of sealed. laton Pusch, 1837, which in the Recent fauna is confined to West Africa, has a labral tooth formed at the end of a cord, and is characterized by having two broad upper cords arising on each intervarical node and then strongly diverging adaperturally to the next formed varix (Vermeij & Houart, 1996). The late Oligocene genus Odontopurpura Vermeij & Vokes, 1997, from North Carolina, differs from Ceratostoma by having a labral tooth formed at the end of a cord, by having long intervarical ribs instead of short nodes, and by having an erect (partially free-standing) inner lip (see Vermeij & Vokes, 1997). Ceratostoma maklyamai (Hatai & Kotaka, 1952) (Figs, 2, 3) 7)-itonalia (Pterorhytis) makiyanTai Hatai & Kotaka, 1952, p. 79, 82, pr. 7, figs. 26, 27. 71ype tocality: Paiponchon, Shinsoruton, San-u-nanmyon, Myonchon District, Hamukyon- pukuton, North Korea. 71ype specimens: IGPS 74368 (Holotype). Material: Two fossil specimens. Description: Shell of medium size, shell height 34.9 mm, with low spire; protoconch eroded and broken, apparently smooth; teleoconch consisting of five whorls; sculpture of early whorls consisting of eight axial ribs and two strong spiral cords, together forming a cancellate sculpture; axial sculpture on last whorl consisting of three blade-like varices, adjacent ones separated by a single interyarical node; varices connected across suture; spiral sculpture of body wherl consisting of ten principal cords and five weaker inter- calated ones; body whorl basally strongly constricted; suture distinct, wavy, impressed; aperture ovate; inner side of outer lip with six denticles; labral tooth absent; siphonal canal long, narrowly open. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Amano & Vermeij: Origin and Biogeographic History of Ceratostoma 213 Remarks and comparisons: The absence of a spinose labral tooth in C. makiyamai is attributable either to the weathered state of the specimens or to the immature nature of the shells. The siphonal canal is narrowly open, a condition again associated either with poor preservation or with a non-adult condition. In spite of these features, we assign the species to Cbratostoma because of its three varices separated by a single intervarical node on the last whorl, and cancellate sculpture on the early whorls. C. maklyamai closely reserntiles C. foliatum in the number of cords on the last whorl, by having varices that connect across the suture, and by having a long siphonal canal.
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