The malacologicalsocietymalacological society of Japan

Jeur. Malac,) fi re VENUS {Jap. - Vol, 5z No. 3 ( l99g): 2e9 223

Originand Biogeographic History of

(: )

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 of the ocenebrine muricid gastropod 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 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, The genus Ceratostoma may have arisen in the Atlantic. By the early Miocene, it 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

liye in shallow Recent species of the ocenebrine muricid gastropod genus Ceratostoma 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 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. Our prupose in this paper is to review the species of Ceratostoma, with special emphasis

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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 (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 Cossmann, 1903 Genus Cle]ratostoma Herrmannsen, 1846

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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. Fig. 4. Ceratostoma sp. x2, MFM, Loc. N64 of Itoigawa et aL (1981); Nataki Formation. Fig. 5. Ceratostoma peranguiatum (Nomland). x 1, UCMP12057 (Holotype), Loc. West of Coalinga, California; Jacalitos Formation. Fig. 6. Purpura turris Nomland. x1, UCMP12049 (Holotype), Loc. West of Coalinga, California; Etchegoin Formation. Fig. 7. Ceratostoma nannum (Nomland). x1, UCMPI1315 (Holotype), Loc. North of Coalinga, California; Santa Margarita Formation.

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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.

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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. However, C. maklyamai

has a smaller shell, higheT spire, less welt developed varices, stronger intervarical nodes,

and fewer denticles on the inner side of the outer lip (six instead of seven to nine). Ceratostoma perangulatum (Nomland) (Fig. 5) from the Jaca!itos Formation (Pliocene) of California is another related species. It has a similar number of spiral cords on the body whorl (eleven), but the spire is lower, and C. perangutatum lacks denticles on the inner side of the outer lip, and has weak intervarical nodes as in C. fotiatum, Dimensions of HolotJ{pe specimen: Height 34.9 mm, spire height 7.7 mm, siphonal canal

length 10.5 mm.

Distribution: Early middle Miocene, Heiroku Formation, North Korea; Kadonosawa For- mation, Iwate Prefecture, Japan (T. Matsubara, personal communication).

( leratestoma sp.

(Fig. 4)

211. Ocenebra (.laton?) sp., Itoigawa et al., 1981, pl.36, figs. 2a,b; Itoigawa et ai., 1982, p. Locality: Dan, Toki-machi, Mizunami City, Gifu Prefecture, central Honshu.

Material: One poorly preserved specimen. Description: Shell small, height 13.6mm, with Iow spire (height 4.1 mm); upper three whorls smooth, the remaining two whorls sculptured; penultimate whorl sculptured with five axial ribs and three weak spiral cords; axial scuipture of body whorl consisting of three thin, blade-like varices and a total of four intervarical nodes; varices connecting

across suture; spiral sculpture of body whorl consisting of four cords; suture distinct, wavy,

impressed; aperture ovate; inner side of outer lip crenulated with more than two denticles;

labral tooth discernible on growth lines preceding varix at aperture; siphonal canal broken, mlssmg.

Remarks and comparison: This specimen was described and illustrated by Itoigawa et al, (1981, 1982) from the Nataki Formation (early middle Miocene) in Gifu Prefecture. The "subgenus" authors doubtfully assigned the species to the .laton Pusch, 1837, but gave

no explanation for this assignment, The specimen has nene of the defining attributes of laton, such as the divergent upper cords (see Vermeij & Houart, 1996). Moreover, it is biogeographically far removed from the eastern Atlantic, where species of Jaton have oc-

curred since the late Oligocene. We assign the specimen to Cleratostoma, but there is no

closely similar species in that genus. Distribution: Nataki Formation (early middle Miocene, Gifu Prefecture, Japan).

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Amano & Vermeij: Origin ancl Biogeographic History of Cleratostoma 215

Composition of the genus Ceratostoma

Hall (1959) ecologically divided the genus Cleratostoma into two groups, the warm-water group of C. nuttalli, and the cold- to ternperate-water group of C. fotiatum. Here we distinguish four groups, which we briefly characterize below. The group of C. nuttalli is characterized by a small shell (usually less than 45 mm high), a relatively short siphonal canal (canal length: shell height O.20 to O.32), varices that are rounded on the upper part of the body whorl and more blade-like near the base,

strong intervarical nodes, and always four to five strong denticles on the inner side of

the outer lip. A very small parietal tooth is present near the upper end of the inner lip.

This group includes C. nuttaUi (Conrad, 1837) (Fig. 9), type of the genus; C. monoceros (Sowerby, 1841) (Fig. 13), and C. notiale Vokes, 1988, The first two species occur in southern California and on the Pacific coast of Baja California, whereas C notiale is from

the Esmeraldas beds (early Pliocene) of Ecuador,

In the group of C. virginiae, the shell is biconic in outline, and is sharply carinated

at the shoulder, with the result that the subsutural area is broad. The siphonal canal is relatively long (canal length: shell height O.30 to O.41). The group consists of two species, C. virginiae (Maury, 1910) from the Chipola Formation (early Miocene) of Florida and C genei (Bellardi & Michelotti, 1841) (Fig. 14) in the sense of Vermeij (1998, in press) from the middle Miocene of Austria and Italy. C. genei, which reaches a height of 76 mm,

has the varices rounded above and b!ade-Iike toward the base as in the C. nuttaUi group.

C. virginiae is a small species (maximum height 22.5 mm), which unlike C. genei lacks

denticles on the inner side of the outer lip.

The C fotiatum group is the most diversified in the genus. It is characterized by a large shell (maximum height 110mm), ten or more strong cords on the body whorl, varices blade-like over their entire Iength, and a relatively long siphonal canal (canal length: shell height usually O.24 to O.36). C. foiiatum (Gmelin, 1791) is distributed in the Recent northeastern Pacific from Prince William Sound (Alaska) to Punta Banda, on the Pacific side of northern Baja California, Mexico (Hall, 1959; Abbott, 1974; Baxter, 1984; Johnson, 1991). In the northern part of its range, this species usually lacks denticles on the inner side of the outer lip, but specimens from California (Fig. 11) typically have

eight denticles and a very small parietal tooth near the upper end of the inner lip. The counterpart of this species in the northwesteTn Pacific is C. burnetti (Adams & Reeve

Fig. 8. Ceratostoma fournieri (Crosse). x 1, Loc. Sakai, Wakayama Prefecture; Recent. Fig. 9. C?ratostoma nuttati (Conrad). x 1, Loc. El Sauzai near Ensenada, Baja California; Recent. Fig. 10. Cleratostoma rorijTuum (Adams & Reeve). x 1, Loc. Fukuma, Fukuoka Prefecture; Recent. Fig. 11. Cleratostoma foiiatum (Gmelin). xO.7, Loc. Pacific Grove, California; Recent. Fig. 12. Cleratostoma burnetti (Adams & Reeve). xO.7, Loc. Oga. Akita Prefecture; Reccnt. Fig. 13. Cleratostoma monoceros (Sowerby). x l, Loc. Guerrero Negro, Baja California Sur, Mexico; Reeent.

Fig. 14. Ceratostoma genei (Bellardl & Michelotti). x 1, NMB no.5S05, Loc. Grund, Austria; middle Miocene.

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216 VENUS: Vo]. S7, No.3(1998)

in Reeve, 1849) (Fig. 12), which occurs in the Japan Sea north to Kurile Islands and

southern Primorye (Pain, 1990; Higo & Goto, 1993). This is the only species of Cle]ratostoma in which the varices are often abaperturally recurved. C fournieri (Crosse, 1861) (Fig. 8) is a warm-temperate Japanese species differing from typical members of the C. foliatum group in having only four principal cords on the body whorl and four denticles on the inner side of the outer lip. Other species in the C. fotiatum group are C. makiyamai (Hatai & Kotaka, 1952) from the Heiroku Formation (early middle Miocene) of North Korea; C. detorae Hall, 1958 from the Oursan Sand (late Miocene) of California;

C nannum (Nomland, 1917) (Fig. 7) from the Santa Margarita Formation (late Miocene) of California; C. perangutatum (Nomland, 1916) (Fig. 5) from the Jacalitos Formation (Pliocene) ef California; and C. petzponderosum (Dall, 1909). known as a probably rewoTked fossil from the Empire Formation (late Miocene) of Oregon and questionably from the Jewett Sand (early Miocene) of southern California. Ceratostoma rorijTuum (Adams & Reeve in Reeve, 1849) (Fig. 10) from the warm- to mild-temperate northwestern Pacific is unique in the genus in having four varices on the

last whorl instead of the usual three. These varices and the intervarical nodes are low,

and the siphonal canal is relatively short (canal length: shell height O.22 to O.26). The small shell (maximum height less than 35 mm) has a very short labral tooth, about nine

low cords on the body whorl, and five to eight denticles on the inner side of the outer ' lip.

We are unable to assign Cleratostoma sp. from the Nataki Formation to any of the

above groups of Ceratostoma. The available material is incomplete and poorly preserved.

Sinelnikova (in Gladenkov & Sinelnikova, 1990) described three species of Ceratostoma from the Etolon Suite (late middle Miocene) of Kamchatka, Among these, C. praeburnettii

has a sealed siphonal canal, four blade-Iike varices en the body whorl, and small denticle

on the adaxial side of the inner lip. From the description and figure, we can discern

neither intervarical nodes nor a labral tooth. Although the shape of the shell and the

presence of four varices might imply that this species is ancestral to C. rorijTuum, the

-]

I I o l' ,S I l i 6a nJae

1

Fig. 15. Distribution ofthe fossil CJeratostoma. t Lf ab"!r red)rkgia)rkrefi95>lii.

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Amano & Vermeij: Origin and Biogeographic History of Ceratostoma 217

poor preservation of the available material makes any definitive statements about this taxon

unwarranted. A second species, C etolonensis, also has four varices. It is said to differ

from C. praeburnettii by having a more strongly recurved siphonal canal and less distinct spiral sculpture. We think it likely that C. etotonensis is a synonym of C. praeburnettii.

The third species, C chetl'sliensis, has six varices and coarse cords on the last whorl. It might be a member of the C. fotiatum group, but poor preservation again makes definitive

assignment impossible, We therefore do not discuss these three species further in this paper. Hali (1959) included Putlpura turris Nornland, 1916, from the Etchegoin Formation (Pliocene) of central California, in the genus Cleratostoma. We disagree with this assign- ment. We have examined the two available specimens, UCMP 12049 (holotype, Fig. 6)

and UCMP 31200. The larger specimen is complete, and measures 42.l mm in height.

Axial sculpture on the last whorl consists of ten blade-like elements, all of which extend

abapically to the base. One of the specimens has two of the ribs enlarged to form varices,

separated by two intervarical nodes. In the other, there arc three varices on the last

whorl, adjacent ones being separated by three intervarica] ribs. The varices appear to be

connected across the suture. Spiral sculpture on the last whorl consists of five very strong

cords. At the base, the body whorl is not constricted on its apertural side. The inner

Table 1.Stratigraphic und distribution of Ceratostoma, gcographic': t'i t)': e v 7v l m o) tum r}y, tele m Ji-"f,

Species Formation name Recent distribution"

C. nuttaUi group C. nuttaUi' San Pedro beds (Pleistecene) S. California and Baja California C, monoceros* San Pedro beds (Pleistocene) Baja California C. notiate Esmeraldas beds (early Pliocene) C. virginiae group C. virginiae Chipola F. (early Miocene) C. genei rniddle Miocene of Austria and Italy C. foliatum group C foliatum' San Pedro beds CPIeistocene) Alaska to northem Baja California Erk River F. (Pleistocene)

C burnetti" Tomikawa F. (early Pleistocene) Primorye, Korea, North Japan, Kurile ls.

C. fournieri* Kume F. (Pliocene) Kyushu to Boso Peninsula. Noto Umegase F. (early Pleistocene) Peninsula C. maktyamai HeiToku F. (early middLe Miocene) C. delorae Oursan Sand (late Miocene) C. nannum Santa Margarita F. (late Miocene) C, perangutaturn Jacalitos F. (Pliocene) C, perponcierosum Empire F. (late Miocenc) ?Jewett Sand (early Miocene) C. rorijZuum group

C, ror(fluum' China, Korea, Kyushu to S. Hokkaido Group unknown

C. sp. Nataki F. middle Miocene) . (early ' Recent species" mainly by Abbott (1974) andHabe & lto(1965), also see text.

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side of the outer lip is strongly thicknened and bears nine denticles. There is no evidence

of a labral tooth. The siphonal canal is sealed. Put:pura turris differs from deratostoma by having an unconstricted base, by lacking

a labral tooth even as an adult, and by having blade-like axial sculptural elements all extend-

ing to the base. Varices are apparently variable in number and are separated from each

other by two to three intervarical ribs rather than by a single intervarical node.

We are uncertain about where to assign Pur:pura turris, but the species appears most similar to C?'assitabrum crassitabrum (Gray in Sowerby, 1834), a temperate Recent species from Peru and Chile, and type of orassitabrum Jousseaume, 1880. Crassilabrum is like

P. turris in having an unconstricted base, lacking a labral tooth, having a sealed siphQnal canal, and having axial sculpture consisting of ten sharp, low. blade-Iike ribs extending

to the base of the body whorl. In several specimens, there is a second, haphazardly placed, varix in addition to the broad terminal varix. Crassilabrum crassitabrun differs

from Purpura turris by having only three instead of fiye very strong spiral cords on the

body whorl, by having six instead of nine denticles on the inner side of the strongly

thickened outer lip, and by having a relatively shorter siphonal canal. Despite these

differences, we tentatively assign P. turris to Crassitabrum. If further research confirms

this assignment, P. turris would be only the second known member of the genus, and the range of the genus would be extended back in time (to the Pliocene) as well as northward (to California).

History of Ceratostoma

The earliest record of Cleratostoma in the fossil record is C? aff. perponclerosum. "Vaqueros'' from the early Miocene () Jewett Sand in the stage of central California (Addicott, l970; Fig. I5; Table 1). Unfortunately, the specimen on which this record is based is very worn. Judging from its large size (height 83 mm) and the presence of broad spiral ribbing, we assign this species to the C. foliatum group. The next species to appear in the fossil record is C. virginiae, from the Chipola Formation (early Miocene, ) of Florida. The Chipola Formation, dated at 18.9Ma (Bryant et al., 1992), is probably slightly younger than the earliest Miocene "Vaqueros" stage of California (see Smith, 1991). C. genei, the only other Atlantic species besides C. virginiae, is a large species from the middle Miocene (?) of

Austria and Italy. Like C. virzginiae in the western Atlantic, C. genei in Europe has no clear predecessors. The origin of the group of C. virginiae therefore remains obscure. The keel connecting the varices in the C, virginiae group resembles that of Microrhytis, whose earliest species, M. christopheri (Gibson-Smith & Gibson-Smith in Vermeij & Vokcs, 1997), from the Cantaure Formation of Venezuela respectively, is of the same early Miocene (Bur-

digalian) age as is C. virginiae (see Verrneij & Vokes, 1997). The keel also occurs in Pterorytis Conrad, 1863 (late Miocene to Pliocene, Atlantic coastal Plain of the United States, and Recent of Peru) and in the Pliocene South African Namamurex. It is therefore possible that the C. virginiae group is phylogenetically connected to Microrhrtis. Pterorytis, and perhaps Namamurex, all genera with a similar labral tooth, and all known mainly

from and originating in the Atlantic.

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Amano & Vermeij: Origin and Biogeographic History of Ceratostoma 219

A relationship between the C virginiae group and the C, nuttaUi group of the eastern Pacific is suggested by the structure of the varices, which in both groups are rounded on the upper part of the whorls and blade-like toward the base. Like the warm-water Atlantic C. vit:giniae group, the C. nuttaUi group has a warm-temperate to tropica] distribution. The oldest species in the C. nuttaUi group is C. notiale, a small species from the early Pliocene of Ecuador (see Vokes, l988). The other two members of the group, C. nuttatti and C, monoceros, are recorded from the San Pedro beds (early Pleistocene) of southern California (Grant & Gale, 1931), and still live in southern California and the Pacific side of Baja California. Evidently, the ancestor of the C. nuttalli group reached the eastern Pacific from the Atlantic before the middle Pliocene emergence of the Central

American isthmus. A significant gap in time exists between the last known occurrence

of the C, virginiae group (middle Miocene) and the earliest occurrence of the C, nuttaUi

group (early Pliocene). Therefore, we cannot constrain the time when the expansion into the Pacific took place. It also remains unclear whether the C. vir:giniae and C. fotiatum groups are phylogeneti- cally directly connected. A literal reading of the fossil record, including acceptance of C? aff. pet:ponderosum from the Jewett Sand as a true Ceratostoma of the C. fbliatum group, would imply that Ceratostoma originated in the warm-temperate northeastern Pacific

and then expanded eastward into the warm-water Atlantic. Such eastward expansion is net known for any other molluscan group, A more likely scenario is that an as yet un- discovered ancestor of the C virzginiae group expanded westward from the Atlantic to the northeastern Pacific, where it would have given rise to the C. fotiatum group. During the Miocene, the C, fotiatum group diversified in both the eastern and western

parts of the North Pacific. In the northeastern Pacific, three late Miocene species are

known: C. deiorae, C. nannum, and C. per;ponderosum. The earliest record of the C. foliatum group in northeast Asia is C. maklyamai from the early middle Miocene. The available evidence therefore implies that the C. foliatum group spread westward from California to northeast Asia by early middle Miocene time.

Subsequently, in the northeastern Pacific, C foliatum is known from the San Pedro beds (early Pleistocene) of southern California (Grant & Gale, 1931) and the Elk River Formation (Pleistocene) of Oregon. It is the only member of the group Iiving in the

northeastern Pacific today, where it is distributed from Alaska to northernmest Baja

California. In Japan, there is no record of Ceratostoma between the early middle Miocene

and the Pliocene. C. fournieri has been described from the Kume Formation (Pliocene) of Ibaraki Prefecture (as Ocenebra nodosa Noda, Kikuchi, & Nikaido, 1993) and from the Umegase Formation (early Pleistocene) in Chiba Prefecture (Noda et aL, 1993; Buba, ``transitional 1990). These occurrences are confined to the so-called zone" (Noda & Amano, 1977), where the warm Kakegawa and the cold-water Takikawa-Tatsunokuchi faunas intermingle. Biogeographically, C. fournieri is therefore similar to the pectinid bivalve M)'zuhQpecten ptanieostulatus (Nomura & Niino) (see Tanaka & Amano, 1997). In the Recent fauna, C. fournieri is distributed in the mild- to warm-temperate waters of Japan, Korea and China (Habe & Ito, 1965; Radwin & D'Attilio, 1976; Higo & Goto, l993). We suspect that C. fournieri is a southern derivative of the Asian branch of the C. foliatum group. C burnetti was illustrated from the Tomikawa Formation (early

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Pleistocene) of southwestern Hokkaido by Sakagami et aL (1996). The two Recent species of the C. foliatum group in northeast Asia may have arisen from an ancestor that spread from the northeastern Pacific, perhaps during the latest Miocene or ear]y Pliocene. The large geographical gap between the northern limits of C. foliatum in the east and C. burnetti in the west implies that geographically intermediate populations have becQme extinct. A similar east to west expansion of range is inferred to have occurred in the group of Micelta canaliculata (Duclos). Nueella shiwa (Chinzei), which is'closely related to the late Miocene to Recent northeastern Pacific NL canalieutata, was described under the genus Polytropa

by Chinzei (1961) from the Togawa Formation (Pliocene) of Aomori Prefecture. N, canaticulata today is distributed from Attu in the westernmost Aleutian Islands of Alaska to central California (Abbott, 1974; Vermeij et al.. 1990). Unfortunately, there is no fossil record of Ceratostoma rorijluum, Therefore, we cannot

make inferences about the history of this enigmatic Japanese species. Many melluscan groups in addition to ([]keratostoma originated on the American side of the North Pacific during or before the early Miocene and then spread westward to northeast Asia by early middle Miocene time. Examples include the ocenebrine Nucetla (Amano et at,, 1993; Collins et aL, 1996), the littorinid Littorina (Reid, 1996), the buc- cinid Lirabuccinum (Vermeij, 1991), the naticid Crlyptonatica and Glossaulax (Marincovich, 1977; Majima, 1989), large radially plicate mytilids (Kafanov, 1987), the tellinid Rexithaerus (Matsubara, 1994) and the rock-boring myid Platyodon (Uozumi & Fujie, 1956; Adegoke, 1967). We suspect a similar and contemporaneous pattern of westward geographical expan- sion in the North Pacific clade of Haliotidae (see Itoigawa & Tomida, 1982; Lindberg, ``Mercenaria", 1992; Brown, 1993), the venerid genus and the ostreid Crassostrea. All these groups originated in warm-temperate waters, and their westward spread to Asia coin- "Climatic cided with the so-called Optimum" of the early middle Miocene, a time when warm conditions reached their greatest latitudinal extent in the North Pacific. Cleratostoma and the other westward-spreading Neogene groups extended into cool- temperate waters, perhaps by the late Miocene, but only Nucella and Littorina became sufficiently cold-adapted to penetrate through the newly opened Bering Strait into the Arctic- Atlantic basin near the beginning of the Pliocene. Those groups that did not achieve such

cold adaptation haye since contracted in range or, in the case of the mytilid Pticatomytiius,

became extinct. Many now have a discontinuous distribution in the northeastern and

northwestern Pacific, the living species being separated by a wide geographical gap

extending from south-central Alaska to the Aleutian and Kurile Islands, Kamchatka, and the northern Okhotsk Sea (see also Vermeij, 1989). It remains unclear why Nucetta and Littorina, but not Cle?ratostoma and many other groups, adapted tQ the cold sufficiently to extend their geographical ranges beyond the North Pacific.

Acknowlectgments: - We thank Dr. Louie Marincovich (California Academy of Sciences), Prof', David Lindberg (Univ. California, Berkeley), Dr. Masanori Shimamoto (Toheku Univ.), Dr. Hiroaki Karasawa (Mizunami Fossi] Mus.) for their help in examining some fossil and Recent specimens. We also thank Dr. Takashi Matsubara (Mus. Nature and Human Activitie$, Hyogo) for his information on the fossil Ceratostoma. Finally we thank Mrs. Janiee Cooper and Mrs. Edith Zipser (Univ. California, Davis)

for their technica] assistance.

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Amano ＆ Vermeij ： Origin and Biogeographic History of Ce 厂atostofina 221

ヒ レ ガ イ 属 の 起 源 と 生 物 地 理 学 的 変 遷

・ 天 野 和 孝 G ，J．フ ェ ル メ イ ・ ー （ヒ越 教 育大 学地 学 教 室 カ リ フ ォ ル ニ ア 大 学 デ ビ ス 校 地 質 学 部）

要 約

II ’ ・ 冂 本 お よ び 北 の の ヒ レ ガ ． ／ 朝鮮 巾 期 噺 統 初 期 イ風 CeratO，sroln a m ‘iki＞amai ， C ． sp ．を検 討 し た 。 こ れ ら を 加 え ヒ レ ガ イ は ．nutta 〃i Conrad virgilliae ． − ， 属 C （ ）， C ． （Maury ）． CJ．．／bliatum （Gmelin ）， C ． rori 〃 ＆ の ・グ ル ープ ，伽 f z （Adanis Reeve ） 1 に細 分 され る 。 本属 は お そ ら く大 西 洋起源 で ，中新 111：前 期 ま で に カ リ フ ォ ル ニ ア に ，中 新 世 中期 の 初 期 ま で に 北 東 ア ジ ア に 分 布 を 広 げ た 。 こ の よ うな 中 新 1［lr前 半 に 」 え へ ー 北 東 ．太平 洋 か ら 北 西 太 1洋 と 分 布 を ひ ろ げ る 移 動 パ タ ン は チ ヂ ミ ボ ラ 属 な ど 他 の 多 くの 北 方系 種 に も特 徴 的 に 見 ら れ る 。

Refe 『ences

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[Received: May 30, 1998; Accepted July 15, 1998]

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