VENUS 67 (3-4): 159-172, 2009

On the Genus Fusinus in Japan V: Further Species, an Unnamed Form and Discussion

Paul Callomon* and Martin Avery Snyder Department of Malacology, Academy of Natural Sciences, 1900 Parkway, Philadelphia PA 19103-1195, USA; *[email protected]

Abstract: In the concluding paper of the series, the authors address a number of names that have been cited erroneously in literature as belonging to the genus Fusinus and/or occurring in Japan. Two forms that until now had not been assigned to named species are identified as further variants of F. perplexus, a third unnamed form is figured and there is brief discussion of some related genera and species. The identity of Fusinus beckii (Reeve, 1848) is confirmed by the rediscovery of its lectotype and a second specimen from the A ndaman Islands. Though originally described as a subgenus of Fusinus, Chryseofusus Hadorn & Fraussen, 2003 is here accorded full generic status. Lectotypes are selected for Fusinus beckii (Reeve, 1848) and F. gracillimus (Adams & Reeve, 1848). Fusus solidulus A. Adams, 1864 is considered a nomen dubium.

Keywords: Fusinus, Fasciolariidae, Japan, unnamed forms, perplexus, beckii, gracillimus, spectru m, Chryseofusus, Pseudolatirus, Trophonofusus, Fusus solidulus

Introduction

The preceding papers in this series (Callomon & Snyder 2004, 2006, 2007, 2008) figured and described all the valid species in the genus Fusinus Rafinesque, 1815 that occur in Japan, together with a number of new ones. All are morphologically quite variable, but fortunately the genus has been largely overlooked by scientists and synonyms are thus relatively few. Identification of Fusinus species was difficult for early Japanese workers, as the types of almost all the Japanese taxa were scattered across the world. Unfortunately, the first major treatment of the western Pacific species to be widely adopted in Japan, in Tryon’s Manual of Conchology (1881), contained many errors. Hirase (1907) summarized the Japanese Fusi nus in a work that replicated Tryon’s with some additions based on local material, though it is likely that Tokubei Kuroda was actually responsible for most or all of the content (Callomon & Tada, 2006). Kuroda (1949) later revised the genus and added several new names that were subsequently invalidated when the journal Yume-Hamaguri was rendered unavailable for nomenclatural purposes in 1985. Authorship of some of Kuroda’s new names thereby fell unintentionally on Kira (1959), and one more that escaped that fate was named in the previous paper in this series (Fusinus amadeus Callomon & Snyder, 2008).

Abbreviations used: ANSP - Academy of Natural Sciences, Philadelphia PA, USA; LMD - Löbbecke Museum und Aquazoo, Düsseldorf, Germany; NC - Kazutaka Noda collection, Gobo, Wakayama Prefecture, Japan; NSMT - National Museum of Nature and Science (formerly National Science Museum), Tokyo; OMNH - Osaka Museum of Natural History, Japan; SL - Shell length; UMUT - University Museum, University of Tokyo. 160 P. Callomon & M. A. Snyder

Systematics

Family Fasciolariidae Genus Fusinus Rafinesque, 1815

Fusinus perplexus (A. Adams, 1864) (Figs. 1-9)

Material examined: 96.4 mm SL (Fig. 1), 90.9 mm SL (Fig. 2), 72.8 mm SL (Fig. 3) & 84.7 mm SL (Fig. 4), Wajima Bay (Japan Sea), Noto Peninsula, Ishikawa Prefecture, Japan (NC); 107.7 mm SL, off Fukui, Japan Sea, 90 m, ANSP 418038 (Fig. 5); 140.0 mm SL (Fig. 6), 128.1 mm SL, off Wajima, 50 m, ANSP 418039; 78.2 mm SL (Fig. 7), 70.0 mm SL (Fig. 8), 61.5 mm SL (Fig. 9), Ariake Sea, Kyushu, ANSP 418040. Remarks: Japan Sea material (Figs. 1-6). Fusinus perplexus is a very variable species, and often can only be identified using a series of intergrades to the nominate form. For a full treatment of the species, see Callomon & Snyder (2004). The authors originally had considered the smaller specimens shown here to belong to a separate species, but latterly examination of larger specimens from the Japan Sea coast of the Noto Peninsula allowed them to be linked to F. perplexus. Coloration often varies in Japanese Fusinus, and tends to be stronger in specimens from the inland seas or the Sea of Japan than in those from the Pacific coasts. The protoconch in the presen t form (Figs. 2c, 2d) has a larger number of whorls than usual for Fusinus, which might imply a longer lecithotrophic stage, or even planktotrophy, though the latter is as yet unrecorded for any fasciolariid. Some as-yet unnamed Fusinus specimens from the Philippines in the authors’ possession bear protoconchs of up to four whorls, which suggests that larval morphology in this genus may not be as st able as hitherto supposed. Ariake Sea material (Figs. 7-9). These specimens are similar to the colored F. perplexus figured by Callomon & Snyder (2004: Fig. 20) from Tokyo Bay, and like them are unusually small for this species. The Ariake Sea in western Kyushu is a shallow, almost entirely landlocked bay with a largely sedimentary bottom. According to the original data, these specimens were collected “at low tide”. This is unusually shallow in comparison with most Fusinus, though F. tuberosus (Reeve, 1847) and F. nigrirostratus (E. A. Smith, 1879) are both known from subtidal depths (Callomon & Snyder, 2007).

Fusinus sp. (Fig. 10)

Material examined: 74.3 mm SL, Shimo-kusui, Nada-cho, Gobo-shi, Wakayama Prefecture. ANSP 413804, ex H. Katori. Remarks: The single specimen to hand cannot be confiden tly assigned to any known Fusinus species. Its broad aspect and dominant spiral cords distinguish it from F. nigrirostratus (E. A. Smith, 1879) (Callomon & Snyder, 2006: figs. 46-52), while its broader canal and reduced axial sculpture set it apart from F. gemmuliferus Kira, 1959 (Callomon & Snyder, 2006: figs. 12-15). It is closest of all to F. amadeus Callomon & Snyder, 2008, of which it might prove to be an unusually broad specimen. On the Genus Fusinus in Japan V 161

1a 1b 2a 2b 2c

2d

3a 2e

4a 4b 7a 7b

3b

8a 8b 2cm

5 6

9a 9b 10a 10b

Figs. 1-9. Fusinus perplexus. 1-4, 6. Wajima Bay (Japan Sea), 96.4 mm SL (1), 90.9 mm SL (2), 72.8 mm SL (3), 84.7 mm SL (4), 140.0 mm SL (6). 5. Off Fukui, Japan Sea, 107.7 mm SL. 7-9. Ariake Sea, Kyushu, 78.2 mm SL (7), 70.0 mm SL (8), 61.5 mm SL (9). Fig. 10. Fusinus sp. Shimo-kusui, Wakayama Prefecture, ANSP 413804, 74.3 mm SL. 162 P. Callomon & M. A. Snyder

Fusinus species erroneously attributed to the Japanese fauna, and species here excluded from Fusinus

Fusinus beckii (Reeve, 1848) (Figs. 11-12)

Fusus ventricosus Reeve, 1847 in 1847-48: pl. 8, fig. 34a. Non Gray, 1839: 117, nec Anton, 1838: 77, nec Lesson, 1842: 214, nec Menke, 1843: 26. Fusus beckii Reeve, 1848 in 1847-48: pl. 17, fig. 34b

Type material: Lectotype, here selected, 151.4 mm SL, BMNH 20041009 (Fig. 11). No locality. Other material examined: 169.5 mm SL, Andaman Islands, ANSP 280045 (Fig. 12). Description: Shell medium sized for genus, thick and heavy. Protoconch absent in material examined. Teleoconch of nine whorls; whorl profile sharply angulate, becoming smoothly rounded from latter part of penultimate whorl. Body whorl rounded, well inflated. Well defined periphery on early whorls, with weak spiral cord forming regularly spaced lobes in crossing axial ribs; cord doubled on early whorls but single by end of sixth. Thick, rounded axial ribs on early whorls decline and become very weak and random by sixth. Body whorl bears scattered thick axial ribs, not reflected on interior face. Weak peripheral cord and lobes present in places on body whorl. Cle ar growth pauses on body whorl; three in lectotype, one in ANSP specimen (Fig. 12d), each with developed dentition. Seven fine, well spaced spiral cords above periphery on teleoconch, four between periphery and suture; thick, broadly flattened spiral cords over entire body whorl and upper half of neck. Aperture quadrate, constricted at terminus of suture. Labral margin thick, with prominent polished, widely-spaced teeth at termini of spiral cords on internal face; internal cords corresponding to troughs between spiral cords on outer body whorl, interspersed in lower two thirds with weaker minor cords that do not reach lip. Interior smooth, glossy; parietal margin strongly protruding and thickened, forming heavy shield that diverts upper part of canal outwards. Neck slender, hardly tapering, gently recurved in distal half, with distinct fasciole in distal third. Canal narrow, parallel; parietal margin sharply produced over entire length. Shell dull white overall with very faint pale brown stains on tips of some early axial ribs. Periostracum present only on ANSP specimen (Fig. 12c, 12d); thin, brown, of typical axial lamellar construction; worn, but retaining some short, sharp tufts. Operculum absent in both specimens. Remarks: Reeve (1847 in 1847-48) gave no locality for Fusinus beckii, and originally cited two specimens. Later in the same work he included a figure of a third specimen (pl. 17, fig. 34b), from the Metcalfe collection, which he termed “a more characteristic one”. The lectotype (Fig. 11) is Reeve’s first figured specimen (pl. 8, fig 34a), now in the BMNH. It is accompanied by a slip of paper reading “Van Diemen’s Land” (usually Tasmania, though occasionally used for northeastern Australia), but there is some doubt whether this is its original data. There is no evidence that any of the specimens cited by Reeve was of Japanese or even western Asian origin. The name was indirectly connected with Japan by Tryon (1881: 53, pl. 33, fig. 99), who treated F. beckii as a variety of F. nicobaricus (Röding, 1798), stating that it “only differs in being denuded of epidermis”, and cited the latter as being from “Japan, Philippines”. He also erroneously synonymized F. nicobaricus with F. oblitus (Reeve, 1847), a Recent species from the Indian Ocean that does not resemble F. beckii. Tryon’s Manual was a major resource for pioneer malacologists in Japan, especially after Kuroda translated various sections of it in the Conchological Magazine, and through it the name On the Genus Fusinus in Japan V 163

11a 11b 13

12a 12b

3cm

12c 12d

Fig. 11. Fusinus beckii. Lectotype, no locality, BMNH 20041009, 151.4 mm SL. Fig. 12. Fusinus beckii. Andaman Islands, ANSP 280045, 169.5 mm SL. Fig. 13. Fusinus nicki. South of Zavora, Mozambique, 100 m, ANSP 418680, 156.5 mm SL. 164 P. Callomon & M. A. Snyder

F. beckii came into common use in the Japanese literature. The species itself, however, was known definitively only from the original figures until the lectotype was rediscovered in the BMNH general collection by the senior author in 2004 and the second specimen examined here was found cataloged in the ANSP as “Fusinus sp”. The latter was collected during the 1920s by the entomologist Lt. Col. M. L. Ferrar, and donated by his distant relative Dr. Robert Robertson of the ANSP. It differs from the lectotype in having larger and more prominent shoulder knobs crossed by two flattened spiral cords, but is otherwise identical. More specimens will be needed to confirm its distinctive characters, but these might include the unusual recurvature of the distal third of the n eck, the strong labral dentition and the well developed parietal shield of the aperture. The closest species morphologically to F. beckii is F. similis (Baird, 1873) from Japan, the Philippines and New Caledonia (Callomon & Snyder, 2006: figs. 17-23). The latter differs from F. beckii in having a taller, more slender spire with broader, rounded axial ribs on the early whorls, a shorter and thicker neck, and more prominent shoulder knobs on the body whorl that often form weak axial buttresses resembling those of F. undatus (Gmelin, 1791). The parietal shield of F. similis, while developed, is not as produced as that of F. beckii, and its labral dentition is somewhat finer. Its surface sculpture tends to be smoother in the later whorls, with some specimens bearing the fine gloss more typical of F. undatus. Fusinus beckii also resembles F. nicki Snyder, 2002 (Fig. 13) from eastern Africa. In the latter, however, two strong spiral cords dominate the axial ribs in the early whorls, forming carinate flanges at their summits; these carinae then persist, often with colored interstices, almost to the body whorl. The cords below the periphery and on the neck in F. nickii are much coarser and fewer in number than in F. beckii, suggesting an affinity with F. nicobaricus (Callomon & Snyder, 2007: figs. 53-58), and the labral margin of the aperture bears only scattered, weak teeth. Coloration in the two specimens of F. beckii to hand is limited to very sparse and faint brown stains on some early axial ribs. Fusinus beckii also shares some of the characters of the Japanese Fusinus perplexus (A. Adams, 1864) (Callomon & Snyder, 2004: figs. 2-10, 20, 21), but differs in its more angular whorl profile, ventricose aperture and highly developed parietal shield.

Incorrect identifications and nomina dubia

Fusinus loebbeckei (Kobelt, 1880) (Fig. 14)

Fusus loebbeckei Kobelt, 1880 in 1844-80: 154, pl. 48, fig. 1 Fusus lacteus Dunker, 1882: 12, pl. III, figs. 11, 12.

Remarks: This species was described with no type locality. Dunker (1882) described it again as Fusus lacteus based on the same specimen as Kobelt had used, making his name an objective synonym. Dunker, however, gave “mare Japonicum” as a locality in his description. The holotype (Fig. 14) has no locality on its original label, so it is not clear why Dunker thought this was a Japanese species. Kuroda (1949: 58) also remarked that t he species is known only from the type, and that there is no evidence for its presence in Japan. Odhner (1923: 12) synonymized F. loebbeckei with Fusinus albinus (A. Adams, 1856), whose type locality is “Ichaboe, West Africa” (Ichaboe Island, off Lüderitz District, Namibia). The present authors concur with this. Snyder (2003: 130) cited its distribution as “Recent, West Africa”. On the Genus Fusinus in Japan V 165

15a 14a 16a

14b

2cm 15b 16b

Fig. 14. Fusus loebbeckei. Holotype, no locality, LMD 105705a, 105.1 mm SL. Fig. 15. Fusinus spectrum. Holotype, “Eastern Seas”, BMNH 1874.12.11.158, 79.7 mm SL. Fig. 16. Fusinus gracillimus. Lectotype, “Eastern Seas”, BMNH 74.12.11.157, 74. 3 mm SL.

Fusinus spectrum (Adams & Reeve, 1848) (Fig. 15)

Fusus spectrum Adams & Reeve in Reeve, 1848 in 1847-8: pl. 18, fig. 68. Remarks: Hadorn (1996) showed this to be an earlier available name for the eastern Pacific species Fusinus panamensis Dall, 1908. Having examined the holotype of F. spectrum (Fig. 15), the present authors agree. This is despite Reeve’s giving the type locality as “Eastern Seas”. 166 P. Callomon & M. A. Snyder

Tryon (1881: 58, pl. 36, fig. 135) added “Japan” to the locality. His figure is apparently based on a specimen of F. spectrum. He also stated that Fusus reeveanus Philippi, 1860 (= Fusus multicarinatus Lamarck sensu Reeve, 1847, pl. 6, fig. 2, non Lamarck, 1822) “appears to be the same species”, a verdict that Hirase (1907: 293) reiterated. The name Fusinus spectrum was subsequently used in Japanese collections and literature. Hirase (1907: 294, fig. 86) acknowledged copying Tryon’s figure and localities. He added records from “Nagasaki” and “Hakodate” with provenance given as “Mr. Schrenck” and “Mr. Pilsbry”, but with no figures. Hirase’s figure is a poor reproduction of Tryon’s that omits the characteristic keeled spiral cord on the shoulder of the body whorl. F. spec trum is not known from the western Pacific.

Fusinus gracillimus (Adams & Reeve, 1848) (Fig. 16)

Fusus gracillimus Adams & Reeve in Reeve, 1848 (Jan), pl. 18, fig. 69. Fusus gracillimus – Adams & Reeve, 1848 in 1848-50: 41, pl. 7, fig. 1; Adams, 1863: 106; Sowerby, 1880: pl 2, fig. 10; Kobelt, 1880 in 1844-80: 198, pl. 64, fig. 1; Tryon, 1881: 63, pl. 38, fig. 159. Fusinus gracillinus [sic] (Adams & Ree ve) – Nomura, 1935: 142, fig. 6; Hu, 1991: 204, pl. 34, figs. 18, 19.

Type material: Lectotype, here selected: 74. 3 mm SL, “Eastern Seas”, BMNH 74.12.11.157 ex Lombe Taylor (Fig. 16). A second specimen is cataloged as “type” in the BMNH, no. 1907.10.28.139 (Kaicher, 1986, card 4664), but this is unlikely to be part of the original Samarang material (K. Way, BMNH: notes on label with lectotype). Remark s: The type locality of Fusus gracillimus was given as “Eastern Seas” and most subsequent accounts have simply reiterated this. Tryon copied the original figure, and stated: “light chestnut-brown. Very like F. turricula, Kiener, but more slender”. Fusus turriculus Kiener, 1840 has long been considered a synonym of Fusinus forceps (Perry, 1811) by most authors (see Callomon & Snyder, 2004: 18). Kobelt cited the locality as “Indochinesischen Meeren” (Indo- Chinese Seas) but admitted copying both the figure and description from Reeve. Adams (1864: 106) reported collecting this species from 6 fathoms at “Tsaulian” (possibly Dalian, China, on the Liaotung Peninsula in the Yellow Sea), but gave no figure or description. Kuroda (1949: 3) observed: “An extremely elongate shell with a red-brown periostracum. The cords bear red spots. Mr. Teramachi has collected a similar shell...but that may be a local form”. Nomura (1935) cited F. gracillimus as a fossil from Taiwan, and added “Living: Japan. Eastern Seas”, but without citing any Recent material. His figure quite closely resembles that of Adams and Reeve. Hu (1991) cited and figured under this name two badly broken fossil specimens, neither of whic h is complete enough to be confidently assigned to F. gracillimus or any similar species. Extensive reviews of Japanese collections have yielded no evidence for the presence of F. gracillimus in Japan or any adjacent area. This name is used consistently for a relatively common species trawled off the coasts of Mozambique and Madagascar that resembles the type of F. gracillimus in most respects except for its considerably greater size. Discussion: Tryon’s work apart, Arthur Adams’s 1864 paper on the “Fusidae” of Japan has also been the root of considerable confusion. In it he recorded Fusus fragosus Reeve, 1848 (in 1847-48: sp. 78) from the Tsushima Strait off northern Kyushu, but this identification must have been an error, a conclusion also reached by Kuroda (1949: 3). F. fragosus is a later name for F. sanctaeluciae (Von Salis, 1793), a Mediterranean species. Adams also cited F. novaehollandiae (Reeve, 1848: Conch. Icon, sp. 70) from Tateyama and Okinoshima, and F. crebriliratus (Reeve, 1848: Conch. Icon., sp. 20) from Tateyama and Hakodate. Neither species occurs in Japan; F. On the Genus Fusinus in Japan V 167 novaehollandiae is known from Australia and New Zealand, while F. crebriliratus is a junior synonym of another Australian species, F. australis (Quoy & Gaimard, 1833). In the same paper Adams described Fusus perplexus, for which a type exists (Callomon & Snyder, 2004: 14, figs 2-10, 20, 21; present work, Figs. 1-9) and Fusus solidulus, whose type is lost. Kuroda (1949: 3-4) translated Adams’s description of the latter, but added that without any indication of size it would be difficult to understand which species Adams had to hand. Higo & Goto (1993: 242, no. 3529) treated F. solidulus as a valid species, but only provisionally placed it in Fusinus; Higo et al. (1999: 263, no. G3039) synonymized it with F. tuberosus (Reeve, 1847). However, a review of Adams’s figureless description shows it to be insufficient to identify any of the known Japanese taxa and F. solidulus is thus here considered a nomen dubium.

Other fusinine groups and species

The genus Chryseofusus Hadorn & Fraussen (2003: 210) has as its type species Fusus chrysodomoides Schepman, 1911, which is known from Madagascar to Fiji. The genus is represented in Japan by two species: Chryseofusus graciliformis (Sowerby, 1880) (Fig. 17) and C. satsumaensis Hadorn & Chino, 2005 (Fig. 18). Although Chryseofusus was originally described as a subgenus of Fusinus, the present authors consider that it warrants full generic status as the shell characters used to differentiate it from Fusinus are of at least the same consistency and significance as those used to distinguish other related genera such as Granulifusus. These are: “...the slightly convex, usually unkeeled whorls with subsutural concavity, the presence of axial ribs usually only on upper whorls, the weak, close-set, regular spiral sculpture crossed by strong, close-set, curved axial growth lines...Outer lip always simple; inner lip completely smooth, the never detached parietal callus consists of an extended, adherent thin la yer of callus” (Hadorn & Fraussen, 2003: 211). The radula of Chryseofusus is more or less identical to that of Fusinus, and Hadorn & Fraussen (2006) were latterly unable to distinguish them convincingly in new species of both genera described alongside each other. However, Habe (1945, 1946, 1958), Barnard (1959) and Maes (1967) among others have shown that the radulae of all fasciolariid genera feature a similar basic morphology, with a normally tricuspid central tooth flanked by a pair of wing- like laterals bearing a variable number of cusps. At extremes, the laterals can be very broad with numerous cusps (Pleuropoca) or narrower with fewer (Fusinus), but basically similar radulae can be found across other genera (Benimakia, Granulifusus etc.) that are otherwise morphologically very disti nct. Characters such as the number of cusps on the lateral teeth can also vary with the age of the specimen (e.g. Barnard, 1959: fig. 19j). Fraussen et al. (2007: 81) ably summarized the anatomical distinctions between the Fasciolariidae and Buccinidae and included some discussion of the radulae. A species that is certainly Fusinine and may ultimately prove to belong in Fusinus is Trophonofusus muricatoides (Yokoyama, 1920), described as a fossil from the Miura Peninsula west of Tokyo. This was recorded and figured as Recent by Kuroda et al. (1971: 186, pl. 50, figs. 9, 10) and latterly by Okutani & Tsuchiya (2000: 517, pl. 257, fig. 58). Examination of figures of the holotype (Fig. 19) suggest that Taki & Oyama (1954: pl. 4, fig. 17) may have been correct in assigning this species to Fusi nus, based on its inflated whorls and dominant early axial sculpture. Further investigation of anatomy will be required to definitively answer this question. Pseudolatirus pallidus Kuroda & Habe in Habe, 1961 is another Japanese species that may well prove too closely related to Fusinus to exclude from that genus. Habe (1961: app. 25) measured a specimen of 56.8 mm SL and cited it as the “type” (moshiki-hyohon). The figured specimen (Habe, 1961: pl. 33, fig. 9) shown here (Fig. 20) seems too small (52.2 mm SL) to be the same specimen, in which case it is a figured paratype. It is shown together with three other 168 P. Callomon & M. A. Snyder

19a 19b 18

21a

19c

21b

22a

17

22b 20a 20b

1cm 23a 23b

Fig. 17. Chryseofusus graciliformis. “Chiba, Japan”, ANSP 289672, ex A. R. Cahn, 51.9 mm SL. Fig. 18. Chryseofusus satsumaensis. Paratype, southwest off Cape Noma, Kagoshima, Kyushu, ANSP 412950, 73.5 mm SL. Fig. 19. Trophonofusus muricatoides. Holotype, Miura City, Kanagawa Pref. (fossil), UMUT CM20162, 12.4 mm SL. 19a, b, to scale with plate; 19c, enlarged. Fig. 20. Pseudolatirus pallidus. Figured paratype (see text), off Cape Ashizuri, Kochi Pref, NSMT-Mo 40378, 52.2 mm SL. Figs. 21-23. Pseudolatirus sp. cf. pallidus. “Tosa”, OMNH 7925, ex T. Kira, 33.2 mm SL (21), 31.6 mm SL (22), 25.8 mm SL (23). On the Genus Fusinus in Japan V 169 specimens for comparison (Figs. 21-23) from Tetsuaki Kira’s collection in the OMNH. The latter were identified as P. pallidus, probably by Kuroda, but they differ somewhat from the type in having finer, more broadly spaced axial sculpture and a more slender profile. Examination of other “Pseudolatirus” material from Taiwan, the Philippines and Australia suggests tha t this genus too requires some attention.

Acknowledgements

The authors would like to thank Drs. Kazunori Hasegawa and Hiroshi Saito (NSMT), Dr. Takenori Sasaki, University Museum, University of Tokyo, and Mr. Kazutaka Noda of Gobo, Wakayama Prefecture, for their collaboration and for the loan of material. Drs. Robert Robertson and Gary Rosenberg (ANSP) provided helpful input.

References

Adams, A. 1864. On the species of Fusidae which inhabit the Seas of Japan. Proceedings of the Linnean Society, Zoology 7: 105-108. Adams, A. & Reeve, L. A. 1848-50. Mollusca. In: A. Adams (ed.), The Zoology of the Voyage of H. M. S. Samarang ... 1843-1846. (x) + 87 pp., 24 pls. 1-24, 1848; 25-44, 45-87, 1850. Dates of plates not known. London. Anton, H. E. 1838. Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Edouard Anton befinden. xvi + 110 pp. Halle. Barnard, K. H. 1959. Contributions to the knowledge of the South African marine Mollusca. Part II. Gastropoda: Prosobranchiata: Rhachiglossa. Annals of the South African Museum 45 (1): 1-237. Callomon, P. & Snyder, M. A. 2004. On some Fusinus (Gastropoda: Fasciolariidae) from Japan, with type selections. Venus 63: 13-27. Callomon, P. & Snyder, M. A. 2006. On the genus Fusinus in Japan II: F. undatus, F. similis and related Pacific taxa, with the description of F. mauiensis n.sp. (Gastropoda: Fasiolariidae). Venus 65: 177-191. Callomon, P. & Snyder, M. A. 2007. On the genus Fusinus in Japan III: nine further species, with type selections. Venus 66: 19-47. Callomon, P. & Snyder, M. A. 2008. On the genus Fusinus in Japan IV: F. longissimus (Gmelin, 1798) and two new species (Gastropoda: Fasciolariidae). Venus 67: 1-13. Callomon, P. & Tada, A. 2006. Yoichiro Hirase and his role in Japanese Malacology. Bulletin of the Nishinomiya Shell Museum 4: 1-30 (English text), 1-22 (Japanese text). Dunker, W. 1882. Index molluscorum maris Japonici conscriptus et tabulis iconum xvi illustratus. 301 pp., 16 pls. T. Fischer, Cassel. Fraussen, K., Kantor, Y. & Hadorn, R. 2007. Amiantofusus gen. nov. for Fusus amiantus Dall, 1889 (Mollusca: Gastropoda: Fasciolariidae) with description of a new and extensive Indo-West Pacific radiation. Novapex 8: 79-101. Gray, J. E. 1839. Molluscuous Animals and their shells. In: J. Richardson et al. (ed.), The Zoology of Captain Beechey’s Voyage ... performed in His Majesty’s Ship Blossom, pp. 104-155 , pls. 33-65. H. G. Bohn, London. Habe, T. 1945. On the radulae of Japanese marine gastropods (2). Japanese Journal of Malacology 14: 1-9. Habe, T. 1946. On the radulae of Japanese marine gastropods (3). Japanese Journal of Malacology 14: 190-199. Habe, T. 1958. On the radulae of Japanese marine gastropods (4). Venus (Japanese Journal of Malacology) 20: 43-60. Habe, T. 1961. Coloured Illustrations of the Shells of Japan, vol. II. xii + 183 + appendix 42 pp., 66 pls. Hoikusha, Osaka. Hadorn, R. 1996. Beiträge zur Kenntnis der Gattung Fusinus Rafinesque, 1815 (Gastropoda: Fasciolariidae), II. Vergleich der Taxa Fusinus panamensis Dall, 1908 und Fusinus spectrum (Adams & Reeve, 1848). Club Conchylia Informationen 28: 66-69. Hadorn, R. & Chino, M. 2005. A new Fusinus (Gastropoda: Fasciolariidae) fr om Japan. Iberus 23: 157-163. Hadorn, R. & Fraussen, K. 2003. The deep-water Indo-Pacific radiation of Fusinus (Chryseofusus 170 P. Callomon & M. A. Snyder

subgen. nov.) (Gastropoda: Fasciolariidae). Iberus 21: 207-240. Hadorn, R. & Fraussen, K. 2006. Five new species of Fusinus (Gastropoda: Fasciolariidae) from western Pacific and Arafura Sea. Novapex 7: 91-102. Higo, S. & Goto, Y. 1993. A systematic List of Molluscan Shells from the Japanese Is. and the Adjacent Area. (3) + xiii + 693 + 13 + 149 pp. Elle Corporation, Osaka. Higo, S., Callomon, P. & Goto, Y. 1999. Catalogue and Bibliography of the Marine Shell-bearing Mollusca of Japan. 749 pp., maps. Elle Scientific Publications, Yao. Hirase, Y. 1907. Honpo kaisan kairui zusetsu [On Japanese marine Mollusca] part 9. Conchological Magazine 1: 281-296, pls. 15-16. Hu, C. H. 1991. Pleistocene fossil mollusks from the Touwo and Baishatun Tongxiao formations of Tongxiao Village, Miaoli County. Taiwan bei lei hua shi zhi [Fauna of Taiwan, Molluscan Fossils] 1 (3): 175-314, pls. 29-53. Kaicher, S. D. 1986. Fasciolariidae II. In: Card Catalogue of World-wide Shells, cards 4624-4729. S. Kaicher, Florida. Kira, T. 1959. Coloured Illustrations of the Shells of Japan, Enlarged and Revised Edition. (5) + VII + (2) + 240 pp., 71 pls. Hoikusha, Osaka. Kobelt, W. 1844-80. Die Gattungen Pyrula und Fusus nebst Ficula, Bulbus, Tudicla, Busycon, Neptunea und Euthria. In: Küster, H. C. (ed.), Systematisches Conchylien-Cabinet von Martini und Chemnitz. 2nd edition, vol. 3 (3b): 1-247, pls. 1-68, 14a. Bauer & Raspe, Nuremberg. Kuroda, T. 1949. Naga-nishi-ko. [On Fusinus]. Yume-Hamaguri (37): 1-8; (39): 57-60. (in Japanese) Kuroda, T., Habe, T. & Oyama, K. 1971. The Sea Shells of Sagam i Bay. xix, 741 pp. [Japanese text], 121 pls., 489 pp. [English text], 51 pp., index, map. Maruzen, Tokyo. Lesson, R. P. 1842. Mollusques recueillis dans la mer du Sud. Genre Turbinella Lamk. Revue Zoologique 5: 210-214. Maes, V. O. 1967. Radulae of two species of Pleuroploca (Fasciolariidae) from the Indo-Pacific. Nautilus 81: 48-54. Menke, C. T. 1843. Molluscorum Novae Hollandiae. 46 pp., Hanover. Nomura, S. 1935. Catalogue of the tertiary and quaternary mollusca from the island of Taiwan (Formosa) in the Institute of Geology and Palaeontology, Tohoku Imperial University, Sendai, Japan. Part 2. Scaphopoda and Gastropoda. Science Reports of the Tohoku Imperial University, Sendai, Japan, Second Series (Geology) 18: 53-228, pls. 6-10. Odhner, N. H. J. 1923. Contributions to the marine molluscan f aunas of South and west Africa. Göteborgs Kungliga Vetenskaps-och Vitterhetssamhället 26 (7): 3-29, pl. 1. Okutani, T. & Tsuchiya, K. 2000. Fasciolariidae. In: Okutani, T. (ed.), Marine Mollusks in Japan, pp. 504-517. Tokai University Press, Tokyo. Reeve, L. A. 1847-48. Monograph of the genus Fusus. Conchologia Iconica 4: pls. 1-14 (1847), pls. 15-21 (1848). London. Snyder, M. A. 2003. Catalogue of the marine gastropod family Fasciolariidae. Academy of Natural Sciences Special Publication 21: iv + 431 pp. Sowerby, G. B. 1880. Monograph of the Genus Fusus. Thesaurus Conchyliorum, or Monographs of Genera of Shells 4: 69-97, pls. 406-417. Taki, Is. & Oyama, K. 1954. Matajiro Yokoyama’s The Pliocene and Later Faunas from the Kwanto Region in Japan. Paleontological Society of Japan Special Paper (2): 1-68, pls. 1-49. Tryon, G. W. 1881. Family Fusidae. In: Manual of Conchology, vol. III. Tritonidae, Fusidae, Buccinidae, 46-97, pls. 28-70. Yokoyama, M. 1920. Fossils from the Miura Peninsula and its immediate north. Journal of the College of Science, Imperial University of Tokyo 39 (6): 1-193, pls. 1-20.

(Received June 9, 2008 / Accepted October 16, 2008) On the Genus Fusinus in Japan V 171

日本産ナガニシ属の研究 V：追加種，未記載種と考察

P. カロモン・M. A. スナイダー

要 約

このシリーズの締めくくりとして，今日まで既知の種類に同定されていなかった 2 つの型について検 討した結果，ナガニシの種内変異と結論づけた。また，未記載と考えられる 1 種について図示し，その 類縁について考察した。さらに，過去の文献で誤ってナガニシ属に含められた，もしくは日本から報告 されたタクサについて，タイプ標本の検討に基づいて考察を行った。

Fusinus perplexus A. Adams, 1864 ナガニシ 本論文で示した日本海産の小型の個体は，当初独立した種類であると考えられたが，同じ海域から得 られた大型の個体と合わせて比較した結果，典型的なナガニシと連続的であることが明らかになった。 本種の色彩に関して，内湾や日本海に産するものは太平洋の外洋の個体に比べて濃色となる傾向があ る。一方，有明海産の個体は，第 1 報で図示した東京湾産の濃色個体に近似するが，この種類としては 異例に小型である。また，多くの日本産ナガニシ類が潮下帯に生息するのに対して，添付データによれ ばこれら有明海産の個体は低潮線付近から採集されていることでも特異である。

Fusinus sp. 和歌山県御坊沖から得られた 1 個体は，これまで知られているいずれの種類にも確実に同定できず， 未知の種類と考えられる。しかし既知の種類の中では F. gemmuliferus Kira, 1959 コブシナガニシに最も近 似し，その著しく幅広い個体の可能性もある。

日本産から除外される種類

Fusinus beckii (Reeve, 1848) 原記載にはタイプ産地は示されていない。図示された 2 個体のうちの最初の個体をここでレクトタイ プに指定した。この標本に添付されていた紙片には「Van Die men’s Land（通常タスマニア，時に北オー ストラリアの地名として用いられる）」と記されていたが，標本の取り違いの可能性がある。 Tryon (1881) は本タクソンを F. nicobaricus (Röding, 1798) の異名とみなし，後者の分布を「日本，フィリピン」とし た。これによって， F. beckii の名前は日本の文献でもしばしば用いられるようになったが，実際のとこ ろ著者らが 2004 年にロンドン自然史博物館からタイプ標本を再発見し，本論文でアンダマン諸島から得 られた別の個体を報告するまでその実体は不明であった。F. beckii は国内の文献ではサイヅチナガニシ の和名とともに用いられることが多いが，例えば木村（ 1997）がこの和名と学名の組み合わせで図示し ているものは Fusinus teretron Callomon & Snyder, 2008 イボウネナガニシである（第 4 報参照）。

Fusinus loebbeckei (Kobelt, 1880) 本種も原記載にタイプ産地が示されていない。後に Dunker (1882) は本タクソンと同一のタイプ標本 に基づいて Fusinus lacteus を記載したが，その際に産地を「mare Japonicum」としている。ホロタイプの 原ラベルには産地は記されておらず，Dunker が何故これを日本産とみなしたのか明らかでない。一方， Odhner (1923) は本タクソンを西アフリカ産の Fusinus albinus (A. Adams, 1856) の異名としており，我々 もこの見解に同意する。 172 P. Callomon & M. A. Snyder

Fusinus spectrum (A. Adams & Reeve, 1848) 本種のタイプ産地は「Eastern Seas」とされ，Tryon (1881) は産地に日本を加えている。これに従って， 平瀬 (1907) をはじめとして様々な文献やコレクションで日本産のナガニシ類にカドバリナガニシの和名 と共にこの学名が用いられてきた。しかし，Hadorn (1996) は F. spectrum を東太平洋産の F. panamensis Dall, 1908 の古参シノニムとみなし，我々のタイプ標本の検討もこれを裏付けた。F. beckii 同様，日本産 の個体で本種に同定されたものは，実際にはイボウネナガニシなどであった（第 4 報参照）。

Fusinus gracillimus (Adams & Reeve, 1838) 本種のタイプ産地も「Eastern Seas」とされ，後に Adams (1864) は中国沿岸の黄海から，また黒田 （1949）は日向灘から本種を報告している。しかし，今回の我々による日本および周辺海域産のナガニシ の詳細な調査によって本種の存在は確認されていない。一方で，これに最も近似した種類はモザンビー クとマダガスカルの沖合から得られている。

その他の関連する種類について

Chryseofusus Hadorn & Fraussen (2003) は Fusus chrysodomoides Schepman, 1911 をタイプ種として創設さ れ，日本産の C. graciliformis (Sowerby, 1880) と C. satsumaensis Hadorn & Chino, 2005 もこれに含められて いる。Chryseofusus は当初ナガニシ属の亜属とされたが，我々はこれを Granulifusus などと同様にナガニ シ亜科の独立の属とみなす。一方で，三浦半島から化石種として記載され，その後現生が確認されてい る Trophonofusus muricatoides (Yokoyama, 1920) フツツカナガニシは，少なくとも貝殻の特徴からはナガ ニシ属の種類である可能性が高い。また，Pseudolatirus pallidus Kuroda & Habe, 1961 シロヒメナガニシ モドキも同様にナガニシ属に移される可能性が高いが，これらの正しい属位の決定のためには今後の詳 しい解剖学的研究が必要である。