Coleoptera: Elmidae), with Descriptions of Their Larvae

Japanese Journal of Systematic Entomology, 20 (2): 235–244. November 30, 2014.

Taxonomic Treatments of Two Japanese Elmid Beetles, Stenelmis vulgaris Nomura and Leptelmis gracilis Sharp (Coleoptera: Elmidae), with Descriptions of Their Larvae

Masakazu HAYASHI1) and Hiroyuki YOSHITOMI2) 1) Hoshizaki Green Foundation, 1664-2 Sono, Izumo, 691-0076 Japan E-mail: [email protected] 2) Entomological Laboratory, Faculty of Agriculture, Ehime University, 3-5-7 Tarumi, Matsuyama, 790-8566 Japan E-mail: [email protected]

Abstract Two synonymic treatments are made: Stenelmis miyamotoi Nomura et Nakane, 1958 is wingless form of Stenelmis vulgaris Nomura, 1958; Leptelmis parallela Nomura, 1962 is winged form of Leptelmis gracilis Sharp, 1888. The larvae of two species are easily distinguishable by the body shape and coloration from other Japanese elmid larvae. Key words. egg, hind wing venation, new synonymy, riffle beetles, wing dimorphism

Introduction NSMT: National Science Museum, Tsukuba (S. Nomura) The family Elmidae (elmid beetles, riffle beetles) lives in Morphological observation lotic environments, which is one of the most diverse group Living adults and larvae on small water tank were of lotic beetles in Japan. Approximately 50 species of elmid photographed by Canon digital camera, EOS 70D attached beetles, classified into 17 genera are known from Japan (Satô, macro photo lens MP-E 65 mm and macro twin light MT- 1985, 1999; Jeng & Yang, 1998; Yoshitomi & Sato, 2005; 24EX in the laboratory. Kamite et al., 2006; Yoshitomi & Nakajima, 2007, 2012; The external morphology of adults and larvae were Kamite, 2009) but the taxonomic problems exist in several examined under a stereoscopic microscope (Nikon SMZ) species and genera. Hayashi et al. (2013) determined that the and photographed for the external structure under a zoom previously separated winged and wingless species, Stenelmis microscope, Nikon AZ-100 with a CCD camera, Nikon Digital vulgaris and S. miyamotoi, and Leptelmis gracilis and L. Sight (DS-L2). Egg is photographed under a light microscope, parallela, are two forms of the same species by molecular Nikon Eclipse E600 attached phase contrast microscopy with phylogenetic analysis based on mitochondrial cytochrome a CCD camera (DS-L2). About 5 to 10 digital photographs oxidase subunit I and nuclear 28S rRNA gene sequences. In were made by focus stacking, using a digital image processing the dimorphic species, wingless forms have sloping shoulders software, Adobe Photoshop CS4 for Macintosh. (antero-lateral part of the elytra) due to degeneration of the Observation and drawing method of the genitalia and hind hind wings, whereas winged forms have square shoulders wing venation are followed by Yoshitomi (2014). (Hayashi et al., 2013). However, their formal synonymization Terminology is followed after Kodada & Jäch (2005) of these species has not been made. In this paper, we treated for morphological features of elmid larva and adult and two pare of synonymy, described their larvae, and discussed Kukalová-Peck & Lawrence (1993, 2004) for wing venation. their ecology. Taxonomic treatments Materials and methods Stenelmis vulgaris Nomura, 1958 Field collecting (Figs. 1A–B, 2A–C, 3A–F, 4A–C, 5A) From June to August of 2006 and 2014, the first author investigated Elmidae in Hii-kawa Riv., Izumo, Shimane Stenelmis vulgaris Nomura, 1958, 44 Prefecture, Japan. All adults found in one hour during the day Stenelmis vulgaris: Nomura, 1960, 36; Satô, 1977, 2; Satô, were collected from riparian roots and decaying woods under 1985, 437. the running water. Stenelmis miyamotoi Nomura et Nakane, 1958, 81 Syn. nov. Stenelmis miyamotoi: Nomura, 1960, 36; Satô, 1977, 2; Satô, Specimens 1985, 437. All the specimens observed in this paper are collected from Japan: almost adult specimens are dry condition with Type series examined. Paratypes of Stenelmis miyamotoi pinned label; larval specimens are also dry in part, other Nomura et Nakane, 1958, 11 exs. (EUMJ), “Nagara Gifu 19. specimens fixed 70% or 99% ethyl alcohol. Abbreviation of VI. 1957 Masa. Satô”. Paratype of Stenelmis vulgaris Nomura, depositories are as follows: 1958, 1 ex. (EUMJ), “Saeki Oita 24. VI. 1950 K. Kurosa”. EUMJ: Ehime University Museum, Matsuyama (H. Addtional specimens examined (in coll. of HOWP). 7 Yoshitomi) exs. (apterous), Hii-kawa, Shimamura, Izumo-shi, Shimane HOWP: Hoshizaki Institute for Wildlife Protection, Pref., 15. VI. 2014, M. Hayashi leg.; 9 exs. (apterous), Hii- Izumo (M. Hayashi) kawa, Shimamura, Izumo-shi, Shimane Pref., 22. VI. 2014, M.

Fig.1. Living individuals of Stenelmis vulgaris (A, B, E) and Leptelmis gracilis (C, D, F) collected from Hii-kawa Riv., Shimane Pref., Japan in 2014. A–D, Adults; E–F, larvae; A, C, apterous; B, D, macropterous.

Hayashi leg.; 2 exs. (apterous), Ibo-gawa, Ibo, Hikawa-chô, slightlycurved interiorly in apical portion, projecting dorso- Izumo-shi, Shimane Pref., 24. VI. 2014, M. Hayashi leg.; 40 interiorly and ventro-apically in proximal 1/2 of dorsal and exs. (apterous), Hii-kawa, Shimamura, Izumo-shi, Shimane ventral portions respectively, sparsely punctate, sparsely Pref., 29. VI. 2014, M. Hayashi leg.; 42 exs. (40 apterous and bearing short setae; median lobe long, about 1.1 times as long 2 macropterous), Hii-kawa, Shimamura, Izumo-shi, Shimane as parameres, tapered posteriorly from base to apical 1/3, Pref., 2. VII. 2014, M. Hayashi leg.; slightly expanded in apical part, sparsely punctate in ventral Descriptions. Body size: macropterous 2.6–3.0 mm, portion. apterous 2.5–3.1 mm. Female: Tergite VIII (Fig. 4A) semicircular, densely Male: Tergite VIII (Fig. 3A) semicircular, sparsely bearing bearing short setae in caudal portion, covered with short setae, covered with microtrichia in basal portion. Sternite microtrichia in basal portion. Sternite VIII (Fig. 4B) projecting VIII (Fig. 3B) transverse, bearing short setae and short antero-laterally in lateral portions, bearing long/short setae, spines, with rather short median strut. Sternite IX (Fig. 3C) with long and slender median strut. Ovipositor (Fig. 4C) asymmetrical, straight in caudal margin, with stout median relatively short and stout; approximate ratio of stylus, distal strut. Aedeagus (Fig. 3D–F) atout, about 0.65 mm; phallobase portion of coxite, basal portion of coxite, and valvifer (n = 1) short, about 0.6 times as long as parameres; parameres long, as 1.0 : 3.4 : 1.9 : 5.7.

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Fig. 2. Male adults of Stenelmis vulgaris (A–C) and Leptelmis gracilis (D–F). A, C, D, F, Dorsal view; B, E, veltral view; A, B, D, E, apterous; C, F, macropterous.

Hind wing: The venation as shown in Fig. 5A. 437. Remarks. Stenelmis miyamotoi Nomura et Nakane is Leptelmis parallela Nomura, 1962, 46. Syn. nov. wingless form (apterous) of Stenelmis vulgaris Nomura Leptelmis parallela: Satô, 1977, 3; Satô, 1985, 437. (Hayashi et al., 2013). Type series examined. Holotype of Leptelmis parallela Leptelmis gracilis Sharp, 1888 Nomura, 1962, (NSMT), “♂”, “Ikenobe, Miki-chô Kagawa (Figs. 1C–D, 2D–F, 3G–L, 4D–G, 5B) Pref. 15 Aug. 1954 M. Chûjo”, “HOLOTYPE Leptelmis parallela Nomura (1962)”. Leptelmis gracilis Sharp, 1888, 244. Addtional specimens examined (in coll. of HOWP). Leptelmis gracilis: Satô, 1960, 43; Satô, 1977, 2; Satô, 1985, 4 exs. (apterous) & 3 larvae, Ibo-gawa, Ibo, Hikawa-chô,

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Fig. 3. Male genitalia of Stenelmis vulgaris (A–F) and Leptelmis gracilis (G–L). A, G, Tergite VIII; B, H, sternite VIII; C, I, sternite IX; D–F, J–L, aedeagus in dorsal (D, J) and ventral (E, K) and lateral (F, L) view.

Izumo-shi, Shimane Pref., 24. VI. 2014, M. Hayashi leg.; 2 as parameres; parameres long, straightly projecting, sparsely exs. (apterous), Hii-kawa, Shimamura, Izumo-shi, Shimane punctate, bearing short setae; median lobe long, about 1.3 Pref., 29. VI. 2014, M. Hayashi leg.; 4 exs. (apterous), Hii- times as long as parameres, subparallel-sided near base to kawa, Shimamura, Izumo-shi, Shimane Pref., 2. VII. 2014, apical 1/3, slightly expanded in apical part, sparsely punctate M. Hayashi leg.; 3 exs. (2 apterous and 1 macropterous), Hii- in ventral portion. kawa, Shimamura, Izumo-shi, Shimane Pref., 4. VII. 2014, M. Female: Tergite VIII (Fig. 4D) semicircular, densely Hayashi leg.; bearing short setae in caudal portion, covered with Descriptions. Body size: macropterous 2.6–2.8 mm, microtrichia in basal portion. Sternite VIII (Fig. 4E) projecting apterous 2.4–2.8 mm. antero-laterally, bearing long setae and minute spines, with Male: Tergite VIII (Fig. 3G) semicircular, densely bearing long and stout median strut. Ovipositor (Fig. 4F) relatively short setae in caudal portion, covered with microtrichia in long; distal portion of coxite slender; approximate ratio of basal portion. Sternite VIII (Fig. 3H) transverse, bearing stylus, distal portion of coxite, basal portion of coxite, and rather long setae and short spines, with long median strut. valvifer (n = 1) as 1.0 : 3.5 : 3.0 : 7.5; vagina with about 30 Sternite IX (Fig. 3I) asymmetrical, shallowly concave in small serrae (Fig. 4G). caudal margin, with long and slender median strut. Aedeagus Hind wing: The venation as shown in Fig. 5B. (Fig. 3J–L) long, about 0.69 mm; phallobase long, as long Remarks. Leptelmis parallela Nomura is winged form

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Fig. 4. Female genitalia of Stenelmis vulgaris (A–C) and Leptelmis gracilis (D–G). A, D, Tergite VIII; B, E, sternite VIII; C, F, ovipositor; G, sclerites on vagina.

Fig. 5. Hind wing venation of Stenelmis vulgaris (A) and Leptelmis gracilis (B).

(macropterous) of Leptelmis gracilis Sharp (Hayashi et al., Description of larvae 2013). Stenelmis vulgaris Nomura (Figs. 1E, 6A–C, 7A–B) In some elmid genera/species, bursa copulatrix of female Description. Body elongate, gradually narrowing middle genitalia have some large sclerites or small spines and serrae to apex; convex dorsally and flattened ventrally; slightly (Berthélemy, 1979), and these are vary in form within genera prominent spiracles on mesothrax and abdominal segment I to and species (Kodada & Jäch, 2005). In the observation of VIII; middle suture present on thorax and abdominal segment this species, we found small serrae bearing inner surface of I to VIII; body color entirely cream-colored but dark band vagina (Fig. 4G), but could not observed the other sclerotized on base and apical margin in all segments, and appendages structures in bursa and bursa copulatrix. yellowish brown. Body surface entirely smooth with long setae.

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Fig. 6. Mature larvae of Stenelmis vulgaris (A–C) and Leptelmis gracilis (D–E). A, D, Dorsal view; B, lateral view; C, E, ventral view.

Head visible from above, well exposed from prothorax; with robust short setae and long setae on outer side. width as long as length; clypeus transverse, with branched Abdomen 9-segmented; segments I–VIII transverse, setae on front; a pair of spine on both sides of clypeus; vertex dorsum entirely smooth with simple setae; pleural sclerites with granules and spiky setae; posterior edge emarginate; on segments I–VII; last segment (segment IX) longer than epicranial stem present but short; frontoclypeal suture distinct. width, gradually narrowing to apex, dorsum smooth without Anttena 3-segmented; 1st shorter than 2nd; 2nd with sensorial longitudinal ridge, apex with a pair of spines; ventral appendage; 3rd slender. Labrum transverse and short, with operculum, opercular claws, and anal gills present. long, simple setae along frontal margin; epipharynx with Body length. Mature larvae is ca. 6.0 mm in expanded dense setae. Mandible tridentate at apex, articulated pubescent specimen preserved in 70 % ethanol; ca. 5.0 mm in dry process in inner margin, long, dense setae in outer margin. condition. Maxilla and labium forming a unit (maxillolabial unit). Identification of larvae. The larva is indentified by Maxilla with 4-segmented palpus; cardo small; stipes large; mitochondrial DNA sequences (Hayashi and Sota, 2010). galea and lacinia separate, setose apically. Labium narrow with 2segmented palpus; ligula short and transverse; mentum Leptelmis gracilis Sharp (Figs. 1F, 6D–E, 7C–D) long; submentum short and transverse. Description. Body elongate and flat, gradually narrowing Thorax not serrated on lateral sides; dorsum entirely middle to apex; slightly convex dorsally and flattened smooth with simple setae. Prothorax longer than wideth, 2.5 ventrally; meso- and metathorax and abdominal segment times as long as mesothorax; with 1 ventral sclerite, procoxae I to VIII with blade-like lateral projections on both sides; closed postriorly; scale-like setae on hind margins. Meso- and remarkably prominent spiracles on mesothorax and abdominal metathoracies trasnsverse; metathorax longer than mesothorax. segment I to VIII; middle suture present on thorax and Mesothrax with five ventral sclerites. Legs 5-segmented, short abdominal segment I to VII; body color entirely cream- and stout; 2nd segment with robust short setae, 3rd segments colored but head and apical half of abdominal segment IX

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Fig. 7. Larvae of Stenelmis vulgaris (A, B) and Leptelmis gracilis (C, D): op, ventral operculum; ps, pleural sclerites on abdominal segment VIII; pms, postero-medial sclerite on prothorax; i to ix showing abdominal segment I to IX; arrows indicate position of spiracles. A, C, Dorsal view; B, D, ventral view. Scale bar = 1.0 mm.

Fig. 8. Egg of Stenelmis vulgaris. A, Egg on decaying wood; B, same egg of A (photo by phase contrast microscopy, phase condenser “Ph3”). dark brown. Two types of granules on body surface and large setae in outer margin. Maxilla and labium forming a unit granule with a pectinate seta. (maxillolabial unit). Maxilla with 4-segmented palpus; cardo Head visible from above, well exposed from prothorax; small; stipes large; galea and lacinia separate, setose apically. width as long as length; clypeus transverse, with setae on Labium narrow with 2-segmented palpus; ligula short and front; frons and vertex with granules and setae; posterior edge transverse; mentum long; submentum short and transverse. emarginate; epicranial stem present but short; frontoclypeal Thorax dentate on lateral sides and hind margin; dorsum suture distinct. Anttena 3-segmented; 1st shorter than 2nd; on two types of granules. Prothorax longer than wideth, 1.9 2nd with sensorial appendage; 3rd slender and short. Labrum times as long as mesothorax; with 2 ventral sclerites, procoxae transverse and short, with setae along frontal margin; closed postriorly by a postero-medial sclerite; pectinate epipharynx with dense setae. Mandible tridentate at apex, setae on front and hind margins. Meso- and metathoracies articulated pubescent process in inner margin, long, dense trasnsverse. Mesothrax with five ventral sclerites. Legs

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Table 1. Collecting records of S. vulgaris found in one hour during the day from Hii-kawa Riv (Mizuho-Ôhashi, Izumo: 35°25´22˝N, 132°49´50˝E; alt. 4 m) in June to August, 2006.

June July August Date Total 13 16 21 7 14 15 3 6 10 11 19 26 Apterous 8 27 24 3 17 14 27 56 30 10 23 40 279 Macropterous 0 0 0 11 42 21 14 25 6 4 9 7 139 % macropterous 0 0 0 78.6 71.2 60 34.1 30.9 16.7 28.6 28.1 14.9 33.3

Table 2. Collecting records of S. vulgaris found in one hour during the day from Hii-kawa Riv (Shimamura-bashi, Izumo: 35°25´44˝N, 132°50´58˝E; alt. 3 m) in June to August, 2014. June July August Date Total 15 22 29 2 17 25 5 22 30 Apterous 7 9 40 40 89 201 92 38 64 580 Macropterous 00021823382276 % macropterous 0 0 0 4.8 16.8 10.3 3.2 17.4 25.6 11.6

Tabale 3. Monthly precipitation (mm) in Izumo Weather Station by AMeDAS* Year / month Jan Feb Mar Apr May Jun Jul Aug 2006 95 85 137 139 166 146 452 33 2014 80 71 162 72 63 102 157 363 mean (30 yrs) 119 98 122 110 135 204 250 123 *Automated Meteorological Data Acquisition System by Japan Meteorological Agency (2000–2014).

5-segmented, short and stout; 2nd segment with robust short 2006 is 33.3 % (n = 418) but that of 2014 is 11.6 % (n = 656) setae, 3rd segments with robust short setae and long setae. (Tables 1, 2). The difference of ratio between 2006 and 2014 Abdomen 9-segmented. Segments I to VIII transverse, is probably caused by precipitation and/or river flow during granules with branched setae on dorsum and lateral parts; the late stage of mature larva. The precipitation of the three pleural sclerites on segments I to VII. Last segment (segment months, April to June observed by Izumo weather station is IX) with longitudinal ridge on dorsum, longer than width, 451 mm in 2006, 237 mm in 2014, and 30 yrs mean is 448.7 gradually narrowing to apex; apex with a pair of spines; mm (Table 3). Increase of precipitation in a particular period ventral operculum, opercular claws, and anal gills present. might increase the incidence of the apterous adults. In the Body length. Mature larvae is ca. 5.0 mm in expanded laboratory rearing, a few scattered eggs are laid on surface of specimen preserved in 70 % ethanol; ca. 4.0 mm in dry submerged wood. The eggs are nearly round, white colored, condition. about 0.25 mm in diameter, the surface smooth, attached a Identification of larvae. The larva is indentified by short stem-like projection. In the water tank, the adults eat mitochondrial DNA sequences (Hayashi and Sota, 2010). organic matter, probably algae on surface of decaying woods and leafs collected with the habitat. Discussion Leptelmis gracilis is relatively rare species and usually live in middle to lower stream but apterous adults also live Ecological notes in still water (e. g. Satô, 1985; Matsui, 1992). Adults and The ecological observations of S. vulgaris and L. gracilis larvae are usually collected from riparian roots of reeds, are reported by Hayashi (2007) based on field survey in Hii- willows and decaying woods under the water (Hayashi, 2007). kawa River System, Shimane Prefecture, Honshu, Japan. In Macropterous adults attack to light at night (e.g. Yamaji, 2008) 2014, the first author reexamined their ecology based on field but its records are rare. Apterous adults appear May to October, observations in Hii-kawa and the laboratory rearing. The but macropterous adults appear June to October. The apterous knowledge of their ecology is noted blow. adults probably pass the winter because the adults collected in Stenelmis vulgaris is common species and usually live spring are old which body covered by mud-like matter densely. in middle to lower stream. Adults and larvae are collected from riparian roots of reeds, willows and decaying woods Distribution (drift woods) under the running water (Hayashi, 2007). Stenelmis vulgaris (incl. S. miyamotoi) is distributed in Macropterous adults attack to light at night. Adults appear Japan (Honshu, Shikoku, Kyushu) and Korea (Satô, 1985). every summer. The apterous adults appear mid June to It has not been recorded from Hokkaido, Tsushima Island, October but macropterous adults appear late June or early Ryukyu Islands and the other islands around the main islands July to October. The field observations suggest that all adults (Hondo) of Japan. The northern limits are known from Iwate dead until late autumn and the larvae pass the winter. The total and Akita Prefectures, near N 40° latitude. ratio of apterous appeared three months, June to August in Leptelmis gracilis (incl. L. parallela) is distributed in

November 30, 2014, JJSE 20 (2) Wing dimorphism and larvae of Stenelmis vulgaris and Leptelmis gracilis 243

Fig. 9. Map showing the records from each prefecture in Japan. A, Stenelmis vulgaris; B, Leptelmis gracilis. Data from Nomura (1958a, b, 1962), Mori- ya (1997), Ishikawa-mushinokai (1998), Yoshitomi et al. (1999), Takano & Ohmomo (2000), Satoh & Ohmomo (2003), Hirano (2004), Ogata & Nakajima (2006), Arai (2007), Naru- kawa (2007), Hayashi & Kadowaki (2007, 2008), Akiyama (2008), Yamaji (2008), Fujiwara (2009), Tominaga et al (2012), Nakamura (2014), and Ministry of Land, Infrastructure, Transport and Tourism, (2007–2014).

Honshu, Shikoku and Kyushu (Satô, 1985). It has not been Hayashi, M. & T. Sota, 2010. Identification of elmid larvae recorded from mall islands around the Japanese main islands, (Coleoptera: Elmidae) from Sanin District of Honshu, Japan, Hokkaido, Tsushima Island, Ryukyu Islands and Korea. The based on mitochondrial DNA sequences. Entomological Science, 13: 417–424. northern limit is known from Iwate Prefecture, near N 40° Hayashi, M., S. D. Song & T. Sota, 2013. Patterns of hind-wing latitude. Two allied species, L. coreana and L. ochra were degeneration in Japanese riffle beetles (Coleoptera: Elmidae). described from Korea, the former is wingless form and the European Journal of Entomology, 110 (4): 689–697. latter is winged form (Jung & Bae, 2012). The pair of Korean Hayashi, M. & H. Kadowaki, 2007. Records on aquatic Coleoptera species is similar to the wing dimorphism of L. gracilis. From from rivers in and around Mt. Daisen, Tottori Prefcture, Japan. Taiwan, Leptelmis formosana was described (Nomura, 1962), Bulletin of the Hoshizaki Green Foundation, 10: 149–168. (In and this species is also allied to L. gracilis. Japanese.) Yoshitomi et al. (1999) reported records of L. gracilis Hayashi, M. & H. Kadowaki, 2008. Records on aquatic Coleoptera from rivers in and around Mt. Daisen, Tottori Prefcture, from Yahagi River System, Aichi Prefecture, Honshu and Japan, part II. Bulletin of the Hoshizaki Green Foundation, 11: figured all known localities on a map until 1999. We made a 269–286. (In Japanese.) new map based on the additional records from each prefecture Hirano, Y., 2004. Coleoptera. In: Insect fauna of Kanagawa, in Japan (Fig. 9). II: 335–835. Kanagawa Konchu Danwakai, Odawara. (In Japanese.) Acknowledgments Ishikawa-mushinokai ed., 1998. Insects of Ishikawa Prefecture. 537 pp. Ishikawa Prefectural Goverment, Kanazawa. (In Japanese.) We thank S. Imasaka, J. Nakajima, N. Ogawa and T. Japan Meteorological Agency, 2000–2014. Website of Japan Tsuduki who informed us about records of Leptelmis gracilis Meteorological Agency (http://www.jma.go.jp/jma/index. from Japan. html) (sited Sep 5, 2014) Jeng, M. L. & P. S. Yang, 1998. Taxonomic review of the genus Grouvellinus Champion (Coleoptera: Elmidae) from Taiwan References and Japan. Proceeding of the Entomological Society of Akiyama, Y., 2008. The correction of distributional records and the Washington, 100: 526–544. supplement of the riffle beetles (Elmidae, Coleoptera) from Jung, S.-W. & Y.-J. Bae, 2012. Riffle beetle genus Leptelmis Hiroshima Prefecture, Southwestern Japan. Miscellaneous (Coleoptera Elmidae) in Korea with descriptions of two new Reports of Hiwa Museum for Natural History, 49: 113–117. (In species. Animal Cells and Systems, 16 (3): 254–259. Japanese.) Kamite, Y., 2009. A revision of the genus Heterlimnius Hinton Arai, K., 2007. Elmidae and Dryopidae of Saitama Prefecture. (Coleoptera, Elmidae). Japanese Journal of Systematic Yosegaki, 125: 1–21. (In Japanese.) Entomology, 15: 199–226. Berthélemy, C., 1979. Elmidae de la region paléarctique occidentale: Kamite, Y., T. Ogata & M. Satô, 2006. A new species of the genus systématique et repartition (Coleoptera Dryopoidea). Annales Zaitzeviaria (Coleoptera, Elmidae) from Tsushima Islands, de Limmnologie, 15 (1): 1–102. Japan. Japanese Journal of Systematic Entomology, 12: Fujiwara, J., 2009. Notes on Elmidae of Hyogo Prefecture, Japan. 149–153. Kiberihamushi, 32 (1): 52–54. (In Japanese.) Kodada, J., & M. A. Jäch, 2005. Elmidae. p. 471–496. In Beutel, Hayashi, M., 2007. Distributional records and ecological notes R. G., & R. A. B. Leschen (eds.), Handbook of Zoology, Vol. on aquatic Coleoptera of Shimane Prefecture. Bulletin of the IV (Part 38), Coleoptera, Beetles, Vol. 1: Morphology and Hoshizaki Green Foundation, 10: 77–113. (In Japanese.) Systematics Archostemata, Adephaga, Myxophaga, Polyphaga

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