Short Notes ©Malacological Society of Japan75

Lectotype Designation and Redescription of exaratoides Kawabe, 1992 (: : )

Hiroshi Fukuda1* and Kunitsugu Kawabe2 1Conservation of Aquatic Biodiversity, Faculty of Agriculture, Okayama University, Tsushima-naka 1-1-1, Kita-ku, Okayama 700-8530, Japan; *[email protected] 2Wakamiyadai 15-18, Yokosuka 239-0829, Japan

Laemodonta exaratoides is an ellobiid that is species. known from Japan and South Korea (Higo et al., We designate the lectotype of L. exaratoides and 1999; Kurozumi, 2000; Min et al., 2008). Kuroda redescribe it in the following lines. (1953, 1958, 1963) cited the unavailable Latin name as “Kuroda, MS.” in his lists of non-marine Family Ellobiidae Pfeiffer, 1854 molluscs of Japan, with no morphological Subfamily Pythiinae Odhner, 1925 descriptions for the species. Kurozumi (2000) cited Laemodonta Philippi, 1846 the author and year of L. exaratoides as “Kuroda, 1957” [the correct year of publication is 1958; see Kawabe, 1992 Malacological Society of Japan (1958: index 6; 1979: 115)]. This was followed by Masuda & Laemodonta (Plecotrema) exaratoides Kuroda, Uchiyama (2004), Nakamoto et al. (2007), Hamada 1953: 11 [nomen nudum]; Kuroda, 1963: 19 (2008), Hayase et al. (2009) and Noda et al. (2009). [nomen nudum]; Kawabe, 1992: 6–8, fig. 1. However, this is not adequate because Kuroda’s Laemodonta exaratoides Kuroda, 1958: 274 [nomen (1958) text constituted a brief mention of the nudum]; Hosaka & Fukuda, 1996: 86, 95, figs distribution range alone and lacked any description. 74, 75; Fukuda, 1996: 43–44, pl. 6, fig. 13; Toki One of us (Kawabe, 1992) gave the first et al., 1998: 183–184, figs 1, 2; Fukuda et al., description of the species with a photograph of the 1999: 53, fig. 27; Yamashita, 1999: 12, 21, fig. shell and L. exaratoides thereby became available 32; Kimura & Kimura, 1999: 50, 51, 55, fig. 6T; under the International Code of Zoological Higo et al., 1999: 405; Fukuda et al., 2000: 145, Nomenclature. Unfortunately the original 189, fig. 18t; Fukuda, 2000: 129, 131, 137, fig. description was published only in Japanese and a 5-20d; Kurozumi, 2000: 819, pl. 406, fig. 10; holotype and paratypes were not designated therein. Matsukuma, 2001: 412, pl. 14, unnumbered fig.; Since Kawabe (1992), the species has often been Aichi Prefectural Government, 2002: 536, referred by Japanese and Korean authors, but an textfig.; Ishikawa, 2002: 131, textfig.; Hyogo English description has been given only by Prefectural Government, 2003: 190–191, pl. 4, Kurozumi (2000) up to the present time. unnumbered fig.; Masuda & Uchiyama, 2004: As shown in the synonymy below, the species 127, textfigs; Min et al., 2004: 340, 341, fig. has often been confused with other congeners, i.e. L. 1071; Nakano, 2006: 406, textfig.; Nakamoto et exarata (H. & A. Adams, 1854) [possible synonym al., 2007: 76, 77, pl. 3, fig. 4; Hamada, 2008: 60, of L. octanflacta (Jonas, 1845) according to pl. 60, fig. G5006; Aichi Environmental Research Hubendick (1956)], L. monilifera (H. & A. Adams, Center, 2009: 546, textfigs.; Suzuki et al., 2009: 1854), L. octanflacta, and L. siamensis (Morelet, 16, 66, fig. 67a, b; Hayase et al., 2009: 30–32, 1875), because of their similar shell characters. L. fig. 5; Noda et al., 2009: 87, 88, pl. 3, fig. A. exaratoides is regarded as threatened in modern Laemodonta (Plecotrema) monilifera — Takahashi Japan and is listed in many regional Red Data & Okamoto, 1969: 58, pl. 13, fig. 6 Books (e.g. Fukuda, 1996; Matsukuma, 2001; Aichi [misidentification]. Prefectural Government, 2002; Ishikawa, 2002; Laemodonta (Plecotrema) exarata — Abe, 1981: Hyogo Prefectural Government, 2003; Nakano, 50–51, pl. 3, fig. 7, pl. 6, figs 6, 7 2006) as an important species in the context of [misidentification]. biodiversity conservation. A lectotype designation is Laemodonta sp. — Fukuda et al., 1990: 22, pl. 2, therefore needed for the precise identification of the fig. 9; Fukuda, 1992: 4, 76, textfig. 2 [epithet 76 VENUS 69 (1–2), 2010

was hidden by a dot line]; Kimura & Masuda, (type locality), 30 May 1976, by K. Kawabe. 1996: 94–97, textfig. 1, pl. 1, figs 1–3. Deposited in the National Museum of Nature and Laemodonta (Plecotrema) octanflata [sic] — Science, Tokyo, with a catalog number NSMT-Mo Yamashita, 1994: 19, fig. 3 [misidentification]. 77001. Laemodenta [sic] exaratoides — Kawai, 1999: 12, Paralectotypes (Figs. 2, 3): 4 shells, all collected 13, fig. 1. alive from the same locality as the lectotype. Laemodonata [sic] octanfracta [sic] — Kwon et al., Paralectotype 1: same data as the lectotype, NSMT- 2001: 188, 189, fig. 716 [misidentification]. Mo 77002. Paralectotypes 2–4: several meters east Laemodonta siamensis — Hamada, 2008: 60, pl. 60, of the seawall on which the lectotype and fig. G5004 [misidentification]. paralectotype 1 were found, under rocks scattered on wet mud in a small reed field, 31 May 1971, by Lectotype (Fig. 1): The syntype figured by K. Kawabe. NSMT-Mo 77003, 77004, 77005. Kawabe (1992: 6, fig. 1). Collected alive from Measurements for the lectotype and crevices in a broken, old seawall in the upper paralectotypes are shown in Table 1. intertidal zone on the northern shore of the Redescription (based on lectotype): Shell small, innermost part of Koajiro Bay, Misaki-cho-Koajiro, ovate fusiform, solid, light yellowish brown. Apex Miura, Kanagawa Prefecture, Pacific coast of pointed. Base with distinct spiral keel surrounding central Honshu, Japan, 35°09´49˝N, 139°37´37˝E wide umbilical depression. moderately high,

Figs. 1–3. Laemodonta exaratoides Kawabe, 1992. 1. Lectotype. 2. Paralectotype 1. 3. Paralectotype 4. Measurements: see Table 1. Short Notes 77

Table 1. Shell measurements (mm) for Laemodonta exaratoides Kawabe, 1992. Lectotype Paralectotype Paralectotype Paralectotype Paralectotype (Fig. 1) 1 (Fig. 2) 2 3 4 (Fig. 3) No. ws 8.5 6.5 8.0 7.5 7.5 L 8.7 7.9 6.5 6.4 6.0 W 5.1 4.7 4.0 4.0 4.0 Lbw 6.5 6.0 5.1 4.9 4.7 Lpw 2.2 2.2 1.6 1.6 1.3 Wpw 3.8 3.5 2.9 2.9 2.7 Ll3ws 7.7 7.2 5.9 5.7 5.4 W3w 2.7 2.5 2.0 2.0 1.8 Lap 5.0 4.6 4.1 3.8 3.9 Wap 2.8 2.9 2.5 2.4 2.5 Wu 1.2 1.1 0.7 0.8 0.7 No. scbw 38 35 47 41 47 No. scpw 8 7 10 11 8 Abbreviations: number of whorls (No. ws), shell length (L), shell width (W), length of body whorl (Lbw), length and width of penultimate whorl (Lpw, Wpw), length of last three whorls (Ll3ws), width of third whorl (W3w), length and width of aperture (Lap, Wap), width of umbilical depression (Wu), and number of spiral cords on body whorl and penultimate whorl (No. scbw, No. scpw). consisting of 8.5 whorls, weakly convex, suture specimens, and therefore the size of the posterior shallow, surface sculptured with many (38 on body parietal tooth may depend on age. whorl, 8 on penultimate whorl) narrow, incised Miura (2008: 144, text figs) illustrated “L. spiral cords. Body whorl about 76% shell length. exaratoides”, but his specimens are identifiable as Aperture elongate oval; inner with three evenly L. siamensis because they have a heavily eroded spaced, strong teeth; columellar tooth oblique apex, wavy spiral cords and a less developed toward base, strong; anterior parietal tooth longest; umbilical keel, and lack periostracal hairs, all of posterior parietal tooth small; outer lip sharp, inside which are diagnostic characters of the latter species. heavily thickened, with one low, ridge-like tooth Although Toki et al. (1998) illustrated the head- opposite parietal teeth. Protoconch eroded, details foot of L. exaratoides, other characters of the soft- unknown. parts of the species have never been published, as Remarks: The lectotype is quite large in size for with all other Japanese species of Laemodonta. To the species. The paralectotypes 1–3 (6.0 to 6.5 mm confirm the relationships of these species, in length) are of average size for the species by anatomical and molecular work is needed. comparison with other specimens recorded in the references cited in the above synonymy. References The lectotype and four paralectotypes are fully Abe, C. 1981. Fauna of the Land and Fresh Water mature, with a thickened outer lip, and they lack of Tokushima Prefecture. 88 pp., periostracal hair on the shell surface. However, it is 10+19 pls. Kyoiku-shuppan Center, Tokushima. known that this species has many short hairs with (in Japanese) regular intervals along the spiral cords (well Aichi Environmental Research Center (ed.) 2009. illustrated by Abe, 1981: pl. 6, fig. 7). The hairs are Threatened Wildlife of Aichi Prefecture. Red usually observed in juveniles but lost in old Data Book Aichi 2009 – . 651 pp. specimens. Department of the Environment, Aichi Prefectural Government, Nagoya. (in The posterior parietal tooth is variable in size Japanese) and shape. It is large and strong in the lectotype and Aichi Prefectural Government (ed.) 2002. Threatened paralectotypes 2, 3 and 4 (Fig. 3), but very weak and Wildlife of Aichi Prefecture. Red Data Book almost invisible in paralectotype 1 (Fig. 2). The Aichi – Animals. 597 pp. Department of the outer lip of the last specimen is thinner than all other Environment, Aichi, Nagoya. (in Japanese) 78 VENUS 69 (1–2), 2010

Fukuda, H. 1992. Representative molluscan species for Hyogo Prefecture 2003. 2+2+382 pp., 9 occurring in Yamaguchi Prefecture. In: Mitoki, pls. Hyogo Environmental Advancement T. (ed.), A Review of the Molluscan Fauna of Association, Kobe. (in Japanese) Yamaguchi Prefecture, Western Japan, pp. 44– Ishikawa, H. 2002. Laemodonta exaratoides. In: 99, 50 pls. Yamaguchi Museum, Yamaguchi. Association for the Investigation of Natural (in Japanese) Environment of Matsuyama (ed.), Red Data Fukuda, H. 1996. Gastropoda. In: Hanawa, S. & Book Matsuyama 2002, 131 pp. Department of Sakuma, H. (eds.), Present status of estuarine Environment, Matsuyama City, Matsuyama. locales and benthic invertebrates occurring in (in Japanese) estuarine environment in Japan. WWF Japan Kawabe, K. 1992. On ellobiids of Japan – some Science Report 3: 11–63. (in Japanese) species and their habitats and distribution Fukuda, H. 2000. Gastropods I – General remarks. ranges. Kakitsubata (18): 6–12. (in Japanese) In: Sato, M. (ed.), Life in Ariake Sea: Kawai, H. 1999. Ellobiids collected from Chita Biodiversity in Tidal Flats and Estuaries, pp. Peninsula. Kakitsubata (25): 12–13. (in 100–137. Kaiyusha, Tokyo. (in Japanese) Japanese) Fukuda, H., Asami, T., Yamashita, H., Sato, M., Kimura, S. & Kimura, T. 1999. The gastropod fauna Hori, S. & Nakamura, Y. 2000. Marine of the marshes of the reed (Phragmites molluscan and brachiopod fauna of Tanoura, australis (Cav.)) in the estuaries in Mikawa Nagashima Island, Kaminoseki-cho, Bay and Ise Bay, Japan. Japanese Journal of Yamaguchi Prefecture, Japan. Yuriyagai 7: Benthology 54: 44–56. (in Japanese) 115–196. Kimura, S. & Masuda, O. 1996. The first occurrence Fukuda, H., Tsuchida, E., Hori, S., Shikano, Y. & of Laemodonta sp. in Hyogo Prefecture (Seto Mitoki, T. 1990. Study on the Mollusca of Inland Sea). Chiribotan 26: 94–97. (in Yamaguchi Pref. – 1. Revision of the Japanese) remarkable molluscan shells from Yamaguchi Kuroda, T. 1953. A Catalogue of Land Shells of Pref. in Mr. T. Kawamoto’s Collection. (1) Japan Including the Okinawa and Ogasawara Gastropoda. Bulletin of the Yamaguchi Islands. 4+55+15+1 pp., 4 pls. [Published by Museum (16): 1–46. (in Japanese) T. Kira, Shijonawate] (in Japanese) Fukuda, H., Yamashita, H. & Fujii, A. 1999. Kuroda, T. 1958. Land shell fauna of Japan and her Molluscan fauna of the estuary of the Tagori adjacent regions (3). Venus 19: 273–294. (in River, Tara, Saga Prefecture, western Japan. Japanese) Saga Nature Study (5): 45–57. (in Japanese) Kuroda, T. 1963. A Catalogue of the Non-Marine Hamada, T. 2008. The Shells of Oita Prefecture. 136 Mollusca of Japan, Including the Okinawa and pp., 100 pls. Published by the author, Oita. (in Ogasawara Islands. v+71 pp. Malacological Japanese) Society of Japan, Tokyo. (in Japanese) Hayase, Y., Kimura, S., Kawabe, K. & Yahashi, M. Kurozumi, T. 2000. Family Ellobiidae. In: Okutani, 2009. Additional records of land and T. (ed.), Marine Mollusks in Japan, pp. 816– freshwater molluscs of Sugashima Island, Toba 823. Tokai University Press, Tokyo. City, Mie Prefecture. Kakitsubata (34): 30–34. Kwon, O.-K., Min, D.-K., Lee, J.-R., Lee, J.-S., Je, (in Japanese) J.-G. & Choe, B.-L. 2001. Korean Mollusks Higo, S., Callomon, P. & Goto, Y. 1999. Catalogue with Color Illustration. 335 pp. Min Shell and Bibliography of the Marine Shell-bearing Museum, Seoul. (in Korean) Mollusca of Japan. 749 pp. Elle Scientific Malacological Society of Japan (ed.) 1958. [Index Publications, Yao. of] Venus, Japanese Journal of Malacology, Hosaka, K. & Fukuda, H. 1996. Discovery of a Volume 19 (1956–1958). Published together population of an endangered brackish water with Venus 20(1), 6 pp. Malacological Society Stenothyra japonica (Gastropoda: of Japan, Tokyo. Neotaenioglossa: Stenothyridae) and Malacological Society of Japan (ed.) 1979. Special characteristics of the accompanied molluscan Publication for Commemoration of the 50th fauna in Onoda City, Yamaguchi Prefecture, Anniversary of the Malacological Society of western Japan, with a comment on the Japan. Venus: Japanese Journal of conservation value. Yuriyagai 4: 65–96. Malacology, Volumes 1–36 (1928–1977). 115 Hubendick, B. 1956. A conchological survey of the pp. Malacological Society of Japan, Tokyo. genus Plecotrema (Gastropoda, Ellobiidae). Masuda, O. & Uchiyama, R. 2004. Freshwater Proceedings of the Malacological Society of Molluscs of Japan, 2. Freshwater Molluscs of London 32: 110–126, pl. 23. Japan, Including Brackish Water Species. Hyogo Prefectural Government (ed.) 2003. Precious Pisces Ecological Field Guide Series 2. 240 Nature of Hyogo Prefecture – Red Data Book pp. Pisces Publishers, Yokohama. (in Short Notes 79

Japanese) ウスコミミガイ（腹足綱：有肺目： Matsukuma, A. 2001. Laemodonta exaratoides Kawabe, 1992. In: Fukuoka Prefectural オカミミガイ科）のレクト Government (ed.), Red Data Book 2001 タイプ選定とその再記載 Fukuoka, 412 pp., pl. 14. Fukuoka Prefectural Government, Fukuoka. (in Japanese) Min, D.-K., Lee, J.-S., Koh, D.-B. & Je, J.-G. 2004. 福田 宏・河辺訓受 Mollusks in Korea (Revised Supplementary Edition). 569 pp. Min Molluscan Research Institute, Seoul. (in Korean) ウスコミミガイの学名 Laemodonta exaratoides は Miura, T. 2008. Pictorial Guidebook of Organisms 黒田（ 1953, 1958, 1963）によって MS名として公 of Tidal Flats. 198 pp. Nanpou-shinsha, 刊されたが，形態の記載文を伴わなかったため長 Kagoshima. (in Japanese) Nakamoto, H., Toki, Y., Noda, K. & Ekawa, K. らく不適格名のままであった。黒住（ 2000）は 2007. A molluscan fauna of “Nagusa-no- 本種の学名 L. exaratoides の命名者・命名年を Hama”, Wakayama-city, Wakayama 「Kuroda, 1957」［正しくは 1958］としており，増 Prefecture, Japan, II. Nanki Seibutu 51: 83–88. (in Japanese) 田・内山（2004），中本他（2007），濱田（2008）， Nakano, T. 2006. Laemodonta exaratoides Kawabe, 早瀬他（ 2009），野田他（ 2009）もこれを踏襲して 1992. In: Mie Prefectural Government (ed.), いるが，黒田（ 1958）は分布域に言及しているの Mie Prefecture Red Data Book 2005, Animals, 406 pp. Mie Prefecture Environmental みで形態の記載は行っていないため，これを本種 Conservation Agency, Tsu. (in Japanese) の原記載と見なすのは適切でない。 Noda, K., Nakamoto, H., Toki, Y. & Ekawa, K. 河辺（ 1992）はこの学名とともに形態の記載文 2009. Molluscan fauna of the Wakagawa River estuary, Wakayama Prefecture, Japan, 3. Nanki 及び殻の写真を公表したため，その時点で国際動 Seibutu 49: 73–78. (in Japanese) 物命名規約上の適格名となった。しかし河辺の原 Suzuki, T., Kimura, S. & Kimura, T. 2009. Guide 記載は簡素な日本語のみで，ホロタイプやパラタ Book for Investigation of Organisms on Tidal Flats. 120 pp. Wetlands International Japan, イプの指定も行っていない。これ以後本種は多く Tokyo. (in Japanese) の著者によって言及されてきたが，英文による記 Takahashi, G. & Okamoto, M. 1969. A Catalogue of 載は黒住（2000）を除いていまだ存在しない。 Molluscan Shells of Fukuoka Prefecture. 本種は同属の L. exarata （H. & A. Adams, 1854） ii+155 pp. Published by the authors, Fukuoka. (in Japanese) （恐らくイササコミミガイ）， L. monilifera（ H. & A. Toki, Y., Nakamoto, H. & Noda, K. 1998. Notes on Adams, 1854） マキスジコミミガイ， L. octanflacta Laemodonta exaratoides (Gastropoda, （Jonas, 1845） イササコミミガイ， L. siamensis Archaeopulmonata, Ellobiidae) from Wakayama City, central Japan. Nanki Seibutu （Morelet, 1875） クリイロコミミガイと殻の形態が 40: 183–184. (in Japanese) 近似するため，従来よりしばしば同定が混乱して Yamashita, H. 1999. Molluscan fauna of salt marsh いた。その一方で，ウスコミミガイは近年絶滅が in Shioya, Kawachi-machi, Kumamoto City, Kumamoto Prefecture, with notes on the 危惧され，生物多様性保全上の重要種として多数 conservation. Kyusyu no Kai (52): 7–22 (in のレッドデータブックに登載されるようになった。 Japanese) このため，本種の同定を今後より確実なものとす Yamashita, K. 1994. An investigation of non-marine molluscs of Ikushima Island. Kainakama 28: るために，河辺（ 1992: 6, fig. 1）で図示された個体 15–24. (in Japanese) をレクトタイプ（国立科学博物館所蔵 NSMT-Mo 77001）に選定する。タイプ産地は神奈川県三浦市 (Accepted August 8, 2010) 三崎町小網代・小網代湾最奥部北岸（北緯 35°0949, 東経 139°3737）である。レクトタイプ はこの種としては著しく大型（殻長 8.7 mm，殻径 5.1 mm）の老成個体で，体層上に 38本，次体層上 に 8本の螺肋を巡らし，本種の幼若個体に見られ る殻表の微細な殻毛は失われている。