Aquatic Insects International Journal of Freshwater Entomology
ISSN: 0165-0424 (Print) 1744-4152 (Online) Journal homepage: http://www.tandfonline.com/loi/naqi20
Review of the Korean Elmidae (Coleoptera: Dryopoidea) with descriptions of three new species
Sang Woo Jung, Manfred A. Jäch & Yeon Jae Bae
To cite this article: Sang Woo Jung, Manfred A. Jäch & Yeon Jae Bae (2014) Review of the Korean Elmidae (Coleoptera: Dryopoidea) with descriptions of three new species, Aquatic Insects, 36:2, 93-124, DOI: 10.1080/01650424.2015.1046457
To link to this article: http://dx.doi.org/10.1080/01650424.2015.1046457
Published online: 15 Jun 2015.
Submit your article to this journal
Article views: 71
View related articles
View Crossmark data
Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=naqi20
Download by: [Korea University] Date: 31 January 2017, At: 01:33 Aquatic Insects, 2014 Vol. 36, No. 2, 93 124, http://dx.doi.org/10.1080/01650424.2015.1046457
Review of the Korean Elmidae (Coleoptera: Dryopoidea) with descriptions of three new species Sang Woo Junga, Manfred A. J€achb and Yeon Jae Baea,c* aDepartment of Life Sciences, Graduate School, Korea University, Seoul 136-713, Korea; bNaturhistorisches Museum Wien, Burgring 7, A-1010 Wien, Austria; cDivision of Environmental Science and Ecological Engineering, College of Life Sciences and Biotechnology, Korea University, Seoul 136-713, Korea (Received 20 July 2014; accepted 21 April 2015; first published online 12 June 2015)
The Korean species of the riffle beetle family Elmidae are revised. Thirteen species belonging to nine genera are recognized including three new species, Grouvellinus aerosus sp. n., Zaitzeviaria kyungseoki sp. n., and Z. obesa sp. n., and one unidentified species of Ordobrevia Sanderson, 1953. Leptelmis coreana Jung and Bae, 2012 and L. ochra Jung and Bae, 2012 are synonymized with L. gracilis gracilis Sharp, 1888. Four species, Macronychus levanidovae Lafer, 1980, Stenelmis koreana Sato,^ 1978, S. cf. montana Zhang and Yang, 1995, and Zaitzevia tsushimana Nomura, 1963, are newly recorded from South Korea. Descriptions, type information, synonymies, illustrations of diagnostic characters and geographical variation, a checklist, and a key to the adults of the Korean species are provided.
http://zoobank.org/urn:lsid:zoobank.org:pub:738109DF-D49D-4398-96B2-FC5863E D76EF Keywords: Coleoptera; Elmidae; new species; new synonym; taxonomy; new faunis- tic records; Korea
Introduction The riffle beetle family Elmidae is a moderately large group of aquatic Coleoptera con- sisting of almost 1500 species in 151 presently recognized genera (J€ach and Balke 2008). They are distributed in all geographical regions with the highest degree of diversity encountered in tropical countries (J€ach and Balke 2008). Adults and larvae are good water quality indicators due to their sensitivity towards environmental changes (Hilsenhoff 1988; Compin and C er eghino 2003;J€ach and Balke 2008). They are collector-gatherers and scrapers feeding on algae, detritus, or wood (Kodada and J€ach 2005; Elliott 2008). Sato(^ 1978) recorded three species of the genus Stenelmis Dufour, 1835 from the Korean Peninsula: S. koreana Sato,^ 1978, S. nipponica Nomura, 1958, and S. vulgaris Nomura, 1958. Since then, five additional species including three new species have been recorded by Yoon (1988), Lee and Lee (1992), Kamite (2009), Jung, Kamite, and Bae (2011), and Jung and Bae (2012): Heterlimnius hasegawai (Nomura, 1958), Leptelmis coreana Jung and Bae, 2012, L. ochra Jung and Bae, 2012, Optioservus gapyeongensis Jung, Kamite, and Bae, 2011, and Zaitzevia nitida Nomura, 1963. Despite these studies, the elmid fauna, regional variations, and their biology are still poorly known in this region.
*Corresponding author. Email: [email protected]
Ó 2015 Taylor & Francis 94 S. W. Jung et al.
The purpose of this study is to describe three new species and to revise the known Korean species of Elmidae using all available material including types, housed in the institutions listed below.
Material and methods Fresh larval and adult material sampled since the 1980s is housed in the Entomological Museum of Korea University (KU), Seoul. They were collected in streams using a hand net (mesh size 0.5 1.0 mm) and a Surber sampler (30£30 cm, mesh size 0.2 mm), as well as with a light trap placed near streams and rivers. Type material of the species previously described from Korea is deposited in the fol- lowing collections: Ehime University Museum, Matsuyama, Japan (EUMJ), Hungarian National History Museum, Budapest, Hungary (HNHM), Natural History Museum, London, UK (NHML), Naturhistorisches Museum Wien, Vienna, Austria (NMW), National Science Museum, Tokyo, Japan (NSMT), and Zoological Institute, Academy of Sciences, St. Petersburg, Russia (ZIAS). Additionally studied non-type material is housed in the NMW and in the following Korean institutions: Museum of Sungshin Women’s University (MSWU), Yeungnam University (YU), and Chungnam National University (CNU). Specimens were examined and photographed under a dissecting microscope with an image analyser (Carl Zeiss Stemi 2000-C with AxioCam MRc5, Zeiss, Germany) and with a scanning electron microscope (SEM S-4700, Hitachi, Japan). Specimens were dehydrated with absolute ethanol and gold-coated for SEM; male and female genitalia were dissected and placed in lactic acid for 24 hours under room temperature before they were examined and illustrated under the dissecting microscope. General terminology follows Kodada and J€ach (2005). Provincial abbreviations are as follows: S (Seoul), GG (Gyeonggi-do), GW (Gangwon-do), CB (Chungcheongbuk-do), CN (Chungcheongnam-do), JB (Jeollabuk-do), JN (Jeollanam-do), GB (Gyeongsangbuk- do), and GN (Gyeongsangnam-do); morphological abbreviations are as follows: MF (macropterous form), BF (brachypterous form), PL (length of pronotum), PW (maximum width of pronotum), EL (length of elytra), EW (maximum width of elytra), and TL (total length of PL and EL).
Taxonomic accounts Genus Grouvellinus Champion, 1923 Type species: Macronychus caucasicus Victor, 1839.
Grouvellinus aerosus sp. n. (Figures 1 12)
Type locality South Korea, Gyeongsangnam-do (Province), Milyang-si, Sanae-myeon, Unmunsan (Mountain).
Type material Holotype. < ‘South Korea, GN, Milyang-si, Sanae-myeon, Unmunsan (Mt.), 18. VI.2009, leg. J.G. Choi’ (KU). Paratypes. 1<,1,, with same label data as holotype (KU). Aquatic Insects 95
Figures 1 12. (Colour online) Grouvellinus aerosus sp. n.: 1 2) dorsal and lateral habitus, female, paratype; 3) aedeagus, dorsal view; 4) aedeagus, lateral view; 5) genital segment, ventral view; 6) ovipositor, ventral view; 7) pronotum; 8) metaventrite; 9) female sternite 8; 10) right hind leg; 11) prosternal process; 12) male sternite 8. Scale bars D 0.1 mm (3 8, 11), 0.2 mm (9, 12), 0.5 mm (10), 1 mm (1 2).
Additional material South Korea: 1, ‘GW, Hwacheon-gun, 12.VI.1987, leg. K.S. Bae’ (KU).
Description Size and shape. Male (holotype): TL D 2.09 mm; PL D 0.59 mm; PW D 0.70 mm; EL D 1.5 mm; EW D 1.0 mm; EL/EW D 1.50; EL/PL D 2.54; EW/PW D 1.42; TL/EW D 2.09. Females (n D 2): TL D 2.14 2.28 mm; PL D 0.60 0.62 mm; PW D 0.74 0.75 mm; EL D 1.54 1.66 mm; EW D 0.98 1.0 mm; EL/EW D 1.57 1.66; EL/PL D 2.56 2.67; EW/PW D 1.30 1.35; TL/EW D 2.18 2.28. Habitus elongate and oval (Figure 1).
Colouration. Head, pronotum, scutellum, and elytra bronze. Pronotum darker than elytra. Antennae, mouthparts, tibiae, tarsi, and elytral setae brown.
Morphology (holotype). Head retractable, granulate dorsally. Frons and clypeus densely pubescent; fronto-clypeal suture distinct and smooth. Labrum transverse, front angles rounded, with several long setae laterally. Antennae 11-segmented; antennomere 2 wide and longer than 1; antennomere 3 long and slender; antennomeres 4 9 shortest; antennomeres 10 wider and longer than 9; antennomere 11 oval, with numerous white setae apically. 96 S. W. Jung et al.
Pronotum (Figure 7) convex at posterior 1/3, wider than long, densely pubescent lat- erally; lateral parts granulate, pubescent; lateral margin smooth, subparallel from basal 1/ 3 to base, gradually narrowed to apex; anterior angle protruding and acute; sublateral cari- nae distinct, sinuate, basal and apical carinae separated from each other; basal carina extending to middle; apical carina extending to basal 1/3; two elongate impressions in front of scutellum. Scutellum triangular with long pubescence. Prosternum with two ele- vated parts at middle; prosternal process (Figure 11) wide and subquadrate. Mesoventrite shiny, convex at middle, mesoventral cavity deep and subquadrate. Metaventrite (Figure 8) flat, shiny at middle; metathoracic discrimen long; large punctures behind mes- ocoxae extending to posterior part; metaventral cavity for reception of abdominal inter- coxal process wide and subquadrate. Legs (Figure 10) long and slender, femora clavate, granulate; tibiae slender, granulate, with apical spur and tomentum on inner surface; hind tibiae with denticulation at apical 1/4. Elytra oblong, wider than pronotum; subparallel-sided, pubescent between striae, con- vex medially; humeri prominent; each elytron with eight punctate striae, strial punctures moderately large and deep from base to middle, becoming gradually finer toward apex; strial interval 3 convex at anterior part; intervals 1 4 wide; intervals 5 6 narrow; inter- vals 7 and 8 carinate; carina of 7th interval longer than carina of 8th interval, extending to apical 1/5; lateral margin smooth from base to apical 1/3, serrate towards apex. Abdomen with plastron, except of median part; admedian keels of ventrite 1 distinct, extending to posterior margin; ventrites 2 and 3 with small lateral process; ventrite 5 long and granulate, with several setae near apex; genital segment and sternite 8 as in Figures 5 and 12. Aedeagus as illustrated (Figures 3 4); penis about 1.5 times as long as phallobasis, slightly longer than parameres, widest at basal part, narrowed toward apex; ventral sac well developed; basolateral apophyses long and slender; parameres wide at anterior part, ventral margin curved bisinuous in lateral view.
Female. Externally similar to male, on average larger; sternite 8 (Figure 9) long; ovipos- itor (Figure 6) long and slender; stylus long and slender; apical portion of coxite about 6.5 times as long as stylus, outer side concave at middle; valvifer shorter than coxite.
Etymology The specific epithet is derived from the Latin adjective aerosus (bronze coloured) refer- ring to the body colour. The name is an adjective in the nominative singular.
Habitat The new species was collected in mountain streams.
Distribution South Korea (Milyang-si, Hwacheon-gun).
Remarks The new species is morphologically similar to Grouvellinus pilosus Jeng and Yang, 1998 from Taiwan. It can be distinguished from the latter by the relatively longer and wider Aquatic Insects 97 parameres (in lateral view) and by the longer phallobasis. We obtained mitochondrial DNA cytochrome oxidase subunit I (COI) gene sequences from one paratype (voucher specimen: 011-EGA, unpublished) of this species to distinguish it from Grouvellinus niti- dus Nomura, 1963 (GenBank: GU816133.1).
Genus Heterlimnius Hinton, 1935 Type species: Helmis koebelei Martin, 1927.
Heterlimnius hasegawai (Nomura, 1958) (Figures 13 17) Optioservus hasegawai Nomura 1958a: 54 (orig. descr.) (for synonyms see Jung et al. 2011: 182).
Type locality Russia, Sakhalin.
Figures 13 17. (Colour online) Heterlimnius hasegawai (Nomura, 1958): 13 14) dorsal and lateral habitus, female; 15 17) dorsal, ventral, and lateral habitus, larva. Scale bar D 1 mm. 98 S. W. Jung et al.
Type material Holotype. , ‘Russia, Shiritori, Saghalien, 24.VII.1938, leg. H. Hasegawa’ (NSMT).
Additional material See Jung et al. (2011).
Diagnosis The adult (Figure 13)ofHeterlimnius hasegawai can be characterized by the following combination of characters: body length 2.20 2.72 mm, head densely rugose and punc- tate, antennae 10- or 11-segmented, lateral margin of pronotum smooth or feebly serrate, long sublateral carinae on pronotum, prosternal process wide and round at apex, strial intervals flat, male genitalia slender, long, and wide apically. The larva (Figures 15 17) of this species can be recognized by the following characters: body elongate, subtriangu- lar in cross section, mandibles slender and about 1.64 times as long as wide, prothorax without ventral posterior sclerite, mesothorax and metathorax with five ventral sclerites (one anteromedial sclerite and four lateral sclerites), abdomen with seven pleural scler- ites, more or less humped in the middle, terminal abdominal segments moderately long and slightly emarginate apically.
Distribution
South Korea, China (Jilin, Liaoning), Far East of Russia (incl. Sakhalin and Kuril Islands), Japan.
Remarks This species was originally described from Japan by Nomura (1958a), who placed it in the genus Optioservus Sanderson, 1954. However, Kamite (2009) transferred it to the genus Heterlimnius.
Genus Optioservus Sanderson, 1954 Type species: Limnius trivittatus Brown, 1930.
Optioservus gapyeongensis Jung, Kamite, and Bae, 2011 (Figures 18 22) Optioservus gapyeongensis Jung, Kamite, and Bae, 2011: 179 (orig. descr.)
Type locality South Korea, Gyeonggi-do (Province), Gapyeong-gun, Buk-myeon, Jeokmok-ri, Garim Spring, 37 58024.800N, 127 26043.700E, alt. 320 m.
Type material Holotype. < ‘South Korea, GG, Gapyeong-gun, Buk-myeon, Jeokmok-ri, Garim Spring, 10.V.2009, leg. Y.J. Bae & S.W. Jung’ (KU). Aquatic Insects 99
Figures 18 22. (Colour online) Optioservus gapyeongensis Jung et al., 2011: 18 19) dorsal and lateral habitus, female; 20 22) dorsal, ventral, and lateral habitus, larva. Scale bar D 1 mm.
Additional material See Jung et al. (2011).
Diagnosis The adult (Figure 18)ofOptioservus gapyeongensis can be distinguished from congeners by the following combination of characters: body oblong, body length about 2.64 mm; pronotum strongly convex, flat and wide laterally, anterior margin arched and protruding, lateral margin strongly serrate, sublateral carinae long; prosternal process long and pointed apically; penis slightly emarginate apically, strongly curved in lateral view, para- meres subtriangular and wide. The larva (Figures 20 22) of this species can be recog- nized by the following characters: body elongate, subtriangular in cross section, small setigerous tubercles dorsally, elongate tubercles at posterior margins, head subquadrate, with short teeth on anterior margin, mandible with long articulated setose process, protho- rax without ventral posterior sclerite, mesothorax and metathorax with three ventral scler- ites (one anteromedial sclerite and two lateral sclerites), abdomen with seven pleural sclerites, terminal segments of abdomen long and slightly emarginate apically. 100 S. W. Jung et al.
Figures 23 32. (Colour online) Ordobrevia sp.: 23) dorsal habitus, female; 24) head; 25) left antenna; 26) prosternal process; 27) pronotum; 28) abdomen; 29) right elytron; 30) ovipositor, ventral view; 31) metaventrite; 32) left hind leg. Scale bars D 0.2 mm (24 29, 31 32), 0.5 mm (23, 30).
Distribution South Korea, China (Liaoning), Far East of Russia.
Remarks In Korea, this species is commonly found in mosses (Bryophytes) of headwater streams or springs. Larvae and adults were found from early March to late September, while no adults were collected in the winter season (from November to February). We obtained COI gene sequences from one male (voucher specimen: 003-EOG, unpublished) of this species to distinguish it from Optioservus hagai Nomura, 1958a and O. maculatus Nomura, 1958a (GenBank: GU816147.1, GU816138.1).
Genus Ordobrevia Sanderson, 1953 Type species: Stenelmis nubifer Fall, 1901.
Ordobrevia sp. (Figures 23 32)
Material South Korea: 1, ‘GW, Pyeongchang-gun, Daegwallyeong-myeon, Odaesan National Park, 10.VI.2008, leg. Y.C. Jeon’ (KU); 1, ‘GW, Hongcheon-gun, Naechon-myeon, Nae- choncheon (Stream), 11.VI.2004, leg. M.C. Kim’ (KU); 1, ‘GG, Icheon-si, Sindun- myeon, Doam-ri, 14.VI.2005, leg. Y.C. Jeon’ (KU); 1, ‘GN, Miryang-si, Sannae-myeon, Unmansan (Mt.), 9.IX.2008, leg. J.K. Choi’ (KU). Aquatic Insects 101
Description Size and shape. Females (n D 3): TL D 2.20 2.30 (2.24) mm; PL D 0.66 0.70 (0.68) mm; PW D 0.64 0.70 (0.67) mm; EL D 1.50 1.60 (1.55) mm; EW D 0.80 0.85 (0.81) mm; EL/EW D 1.84 2.00 (1. 90); EL/PL D 2.14 2.37 (2.26); EW/PW D 1.14 1.32 (1.22); TL/EW D 2.62 2.87 (2.75). Habitus elongate and subparallel-sided (Figure 23).
Colouration. Body brown to grey. Head dark brown; pronotum grey and brownish; antennae, mouthparts, scutellum, elytra, and legs brown.
Morphology (female). Head (Figure 24) finely punctate and granulate; frons smooth between granules in middle, rugose between granules laterally. Clypeus granulate; fronto- clypeal suture nearly straight. Antennae (Figure 25) 11-segmented; antennomere 1 long; antennomeres 3 6 shortest; antennomere 11 large and oval, with setae. Mandibles slen- der, with three small teeth. Maxillary palp 4-segmented; palpomere 1 short; palpomeres 2 and 4 wide; palpomere 4 long and relatively large; galea 2-segmented. Labial palp 3-seg- mented; palpomere 1 short; palpomere 2 wider and longer than 1; palpomere 3 relatively large and wide. Pronotum (Figure 27) quadrate, densely granulate, widest at basal 1/3; median longi- tudinal impression narrow and shallow; sublateral impressions reaching from base to mid- dle, smooth between granules, shiny, brownish, strongly rugose at lateral parts; anterior angles more or less protruding and blunt, less than 45 in dorsal view; lateral margin strongly serrate from basal 1/3 to apex, more or less smooth from base to basal 1/3; two deep impressions in front of scutellum. Scutellum flat and subcordate, granulate. Proster- num reticulate; prosternal process (Figure 26) longer than wide, reticulate, widest at mid- dle, round at apex. Mesoventrite reticulate, mesoventral cavity deep and subtriangular; round at apex, mesoventral discrimen present, mesoventral process with sinuate margins. Metaventrite (Figure 31) reticulate, longitudinal impression extending to middle, trans- verse suture present near posterior part, metaventral cavity for reception of abdominal intercoxal process long and narrow. Legs (Figure 32) long and slender; femora and tibiae densely granulate; tibiae with thick setae laterally and ventrally. Elytra (Figure 29) sub- parallel-sided, wider than pronotum, densely granulate anteriorly and laterally; lateral margin slightly crenulate; humeri prominent; hind wings well developed; each elytron with eight punctate striae; accessory stria (five punctures) in interval 1, merging with 1st stria; strial punctures deep and large from middle to base, becoming gradually smaller and obsolete toward apex; strial intervals 3, 6, and 7 convex at anterior parts; lateral cari- nae in 6th interval from basal 1/5 to apical 1/3. Abdomen (Figure 28) with five ventrites, distinctly pubescent, finely punctate, densely granulate; abdominal intercoxal process triangular, reticulate, admedian keels absent; ventrites 1 and 2 reticulate laterally; ventrites 3 and 4 with small lateral processes; ventrite 5 long and wide, with thick setae near apex; apex truncate. Ovipositor (Figure 30) long and slender; stylus long, narrow near base; apical portion of coxite about 4.4 times as long as stylus; proximal ventral plates narrow; valvifer about 1.1 times as long as coxite.
Male. Unknown.
Distribution South Korea. 102 S. W. Jung et al.
Remarks Most probably, this is an undescribed species. However, we refrain from a formal descrip- tion, because only females are known so far. This species distinctly differs from Palaearc- tic congeners (Ordobrevia amamiensis Nomura, 1957a, O. foveicollis Schonfeldt,€ 1888, O. gotoi Nomura, 1959, and O. maculata Nomura, 1957b) in the brownish glabrous sub- lateral pronotal impressions reaching from base to middle. It seems to be morphologically related to Ordobrevia gotoi (endemic to Japan), but can be distinguished by the combina- tion of the following characters: antennal segments short and wide; pronotum wide, impressions brownish and smooth between granules; elytra with short accessory stria, and strial interval 3 convex anteriorly. We obtained mitochondrial COI gene sequences from one female (voucher specimen: 016-EOF, unpublished) of this species to distinguish it from Ordobrevia maculate (GenBank: GU816106.1). This species identity should be clarified by collecting males and examining their genitalia.
Genus Leptelmis Sharp, 1888 Type species: Leptelmis gracilis Sharp, 1888.
Leptelmis gracilis gracilis Sharp, 1888 (Figures 33 34)
Leptelmis gracilis gracilis Sharp 1888: 244; Bollow 1941: 85 (redescr.); Sato^ 1960:43 (redescr.); Yoshitomi, Shragane, and Hikida 1999: 101 (note); Hayashi and Shimada 2006: 133 (note); Hayashi 2007: 93 (note); Hayashi 2011: 102 (note); Hayashi, Song, and Sota 2013 (syn.); Hayashi and Yoshitomi 2014: 240 (larv. descr., syn.).
Leptelmis parallela Nomura 1962: 46 (desc.); Yoshitomi et al. 1999: 101 (note); Hayashi 2007: 93 (note); Hayashi 2011: 103 (note); Hayashi et al. 2013 (syn.); Hayashi and Yoshi- tomi 2014 (syn.).
Figures 33 34. (Colour online) Leptelmis gracilis gracilis Sharp, 1888, female: 33) brachypter- ous; 34) macropterous. Scale bar D 1 mm. Aquatic Insects 103
Leptelmis coreana Jung and Bae 2012: 255 (desc.). syn. n. Leptelmis ochra Jung and Bae 2012: 256 (desc.). syn. n.
Type locality Japan, Honshu, Yokohama.
Type material Holotype. (gender unknown) ‘Japan, Yokohama, 1910, leg. G. Lewis’ (NHML).
Additional material See Jung and Bae (2012).
Diagnosis Adults of this species can be distinguished from congeners by the body length (2.5 2.6 mm), uniform colour (brown or blackish brown), slender maxillary palpomere 4, distinct transverse impression with long longitudinal impression at middle part of pronotum, wing dimorphism (brachy- or macropterous), convex and paler on strial interval 3 anteri- orly, relatively long legs, large claws with basal teeth, and elongate penis with slightly convex apex.
Distribution South Korea, Japan.
Remarks Leptelmis gracilis gracilis is polymorphic. Winged specimens (Figure 34) have distinct elytral humeri, and the body shape is parallel-sided. Wingless specimens (Figure 33) have no humeri, and the elytra are distinctly oval. We obtained COI gene sequences from both brachypterous and macropterous speci- mens of Leptelmis in Korea (voucher specimens: 009-ELG (BF) and 010-ELG (MF), unpublished) and compared those with Japanese specimens of L. gracilis gracilis. The sequence data (COI gene) showed that the genetic distance (1.1 1.2%) between the Japa- nese and Korean specimens lay within an intra-specific divergence range (< 2%) (Hebert, Ratnasingham, and de Waard 2003). All sequences from Japan (L. gracilis gracilis) were deposited in NCBI (GenBank: AB764141 AB764142). Some variations such as absence or presence of indistinct apical spurs on tibiae and unexpanded apex in male genitalia are also found in local populations of this species (for detailed comparisons, see Jung and Bae 2012). The larva and the ecology of this species were recently described by Hayashi and Yoshitomi (2014). Unfortunately, we were not able to examine Leptelmis gracilis impubis Zhang and Ding, 1995 and L. guangxiana Zhang, Su, and Yang, 2003, both deposited at Insect Col- lections of China Agricultural University, Beijing, China. According to their descriptions (Zhang and Ding 1995; Zhang, Su, and Yang 2003a), they might be junior synonyms of L. gracilis gracilis. 104 S. W. Jung et al.
Figures 35 41. (Colour online) Stenelmis koreana Sato,^ 1978: 35 36) dorsal and lateral habitus, male; 37) aedeagus, ventral view; 38) aedeagus, lateral view; 39) ovipositor, ventral view; 40) pronotum; 41) left middle leg. Scale bars D 0.1 mm (37 39), 1 mm (35 36, 40 41).
Genus Stenelmis Dufour, 1835 Type species: Limnius canaliculatus Gyllenhal, 1808.
Stenelmis koreana Sato,^ 1978 (Figures 35 41) Stenelmis koreana Sato^ 1978: 147 (orig. descr.).
Type locality North Korea, Pyongyang, Hotel garden.
Type material Holotype. < ‘North Korea, Pyongyang, Hotel garden, Prov. South Pyongan, 5. VIII.1971, leg. S. Horvatovich & J. Papp, det. M.Sato’^ (HNHM). Paratypes. 39<,52,, with same label data as holotype (HNHM); 1<,2,, ditto but deposited in EUMJ; 1<, ditto but deposited in NMW; 1<, ditto but 10.VIII.1971, leg. S. Horvatovich & J. Papp, det. M. Sato^ (EUMJ).
Additional material South Korea: 9<,6, ‘GG, Namyangju-si, Wabu-eup, Songchon-ri, Sujongsa (light trap), 14.VII.1984, leg. T.Y. Moon’ (KU); 2, ‘GG, Yeoncheon-gun, Gunnam-myeon, Buksam- ri, Buksamgyo (Br.), 10.VI.2010, leg. S.W. Jung’ (KU); 3,, ditto but 28.VI.2014 (KU); 5, ‘CB, Danyang-gun, Gagok-myeon, Sapyeong-ri, 18.V.1987, leg. K.S. Bae’ (KU).
Diagnosis This species can be distinguished by the presence of four distinct gibbosities on the pro- notum (Figure 40), by the wide pronotal impression, and by the densely granulate ventral surface. The male can be recognized by the middle tibiae (Figure 41), which are dilated at apical 1/3. Aedeagus and ovipositor as in Figures 37 39. Aquatic Insects 105
Distribution South Korea (new record), North Korea.
Remarks In northern South Korea, this species is known only from one locality near the border to North Korea. No specimens were collected in the southern part of South Korea since 1987. We obtained COI gene sequences from two female adults (voucher specimen: 044- ESK, 045-ESK, unpublished) to distinguish this species from Stenelmis vulgaris (Gen- Bank: GU816122.1).
Stenelmis cf. montana Zhang and Yang, 1995 (Figures 42 50) Stenelmis montana Zhang and Yang 1995: 102 (orig. descr.); Zhang, Su, and Yang 2003b: 108 (descr.).
Type locality China, Guangxi, Guilin, Yanshan.
Figures 42 50. (Colour online) Stenelmis cf. montana Zhang and Yang, 1995: 42 43) dorsal and lateral habitus, male; 44) aedeagus, ventral view; 45) aedeagus, lateral view; 46) ovipositor, ventral view; 47) head; 48) pronotum; 49) prosternal process; 50) metaventrite and abdomen. Scale bars D 0.1 mm (44 46), 0.2 mm (47 50), 1 mm (42 43). 106 S. W. Jung et al.
Type material Holotype. < ‘China, Guangxi, Guilin, Yanshan, 240 m, 27.V.1963, leg. C.K. Yang’ (Insect Collections of China Agricultural University, Beijing, China). [not examined]
Material South Korea: 4<,5,, 14 exs. ‘JN, Gokseong-gun, Boseong river, 23.VIII.1988, leg. K.S. Bae’ (KU); 9 exs. ‘JB, Sunchang-gun, Seomjin river, 21.V.1989, leg. K.S. Bae’ (KU); 1<,4,, 1 ex. ‘GB, Andong-si, Imha-myeon, Imha dam (downstream), 23.III.2009, leg. S. W. Jung’ (KU); 2,, 1 ex. ‘GB, Cheongdo-gun, Unmun-myeon, Sinwolgyo (Br.), 14. V.2010, leg. S.W. Jung & I. K. Shin’ (KU); 1<,1, ‘GB, Sangju-si, Nakdong river, 8. X.2012, leg. Y.J. Park’ (KU); 1< ‘CB, Chungju-si, Geumga-myeon, Saam-ri, Masagyo (Br.), 5.X.2012, leg. S.W. Jung’ (KU).
Description Size and shape. Males (n D 5): TL 2.45 2.60 (2.50) mm; PL 0.65 0.75 (0.72) mm; PW 0.72 0.80 (0.77) mm; EL 1.70 1.85 (1.78) mm; EW 1.00 mm; EL/EW 1.70 1.85 (1.78); EL/PL 2.26 2.76 (2.48); EW/PW 1.25 1.38 (1.29); TL/EW 2.45 2.60 (2.50). Females (n D 5): TL 2.45 2.65 (2.57) mm; PL 0.70 0.75 (0.74) mm; PW 0.75 0.80 (0.80) mm; EL 1.75 1.90 (1.83) mm; EW 1.00 mm; EL/EW 1.75 1.90 (1.83); EL/PL 2.40 2.53 (2.47); EW/PW 1.25 1.33 (1.26); TL/EW 2.45 2.65 (2.57).
Colouration. Head black; pronotum, femora and tibiae dark brown; mouth parts, antennae, tarsi, and claws brown; ventral surface granulate and grey.
Morphology. Head (Figure 47) granulate; distance between eyes 0.32 mm; clypeus wider than labrum, granulate. Antennae 11-segmented; antennomere 1 long and thick; antennomere 2 shorter than antennomere 1; antennomeres 3 7 slender; antennomeres 8 10 wide apically; antennomere 11 long and oval, with several setae; approximate ratio of antennal segments (n D 1): 1.6 : 1.1 : 1.1 : 1.0 : 1.1 : 1.1 : 1.1 : 1.3: 1.3: 1.3: 2.3. Pronotum (Figure 48) slightly wider than long, widest at posterior 2/5; densely granu- late; lateral margin serrate; anterior angles protruding and acute; posterior angles sub- acute; median longitudinal impression wide near middle, becoming narrow at base; sublateral carinae extending from base to apical 1/5; two conspicuous deep impressions in front of scutellum. Scutellum subpentagonal, with granules. Prosternum densely granu- late; prosternal process (Figure 49) longer than wide, concave at apex. Meso- and meta- ventrite densely granulate; metathoracic discrimen long, 0.37 mm in length, metaventral cavity for reception of abdominal intercoxal process subquadrate. Legs long and slender, densely granulate except on tarsi; tibiae slender with apical spurs; tarsomere 5 as long as 1 4 combined; elytra subparallel-sided, wider than pronotum, lateral margin crenulate; humeri prominent, hind wings well developed; each elytron with eight punctate striae; strial punctures deep from base to apical 1/3; 2nd stria obsolete at apical 1/4; 4th stria merging with 3rd stria at apical 1/4; interval 6 carinate. Abdomen (Figure 50) with five ventrites, densely granulate laterally; abdominal inter- coxal process wide, ventrite 1 with more or less large tubercle with densely set setae at posteromedial margin, middle impressed; ventrite 2 impressed anteriorly; ventrite 4 densely pubescent near posteromedial margin; ventrite 5 densely granulate, concave Aquatic Insects 107 apically. Aedeagus as in Figures 44 45; penis longer than parameres, slightly narrowed at basal 1/3; corona and fibula present; parameres slender, dorsal margin with distinct setose projection at apical 1/4, about 1.08 times as long as phallobase in lateral view.
Female. On average larger than male; ovipositor (Figure 46) long and slender; apical portion of coxite about 5.37 times as long as stylus; valvifer about 4.83 times as long as basal portion of coxite.
Distribution South Korea.
Remarks Unfortunately, we were not able to examine the type material of Stenelmis montana, deposited at Insect Collections of China Agricultural University, Beijing, China. The original description of S. montana (so far recorded from Guangxi, Fujian, and Zhejiang in China) unfortunately does not allow unambiguous identification of the specimens from Korea. In Korea, Stenelmis cf. montana normally occurs together with S. nipponica in streams with a substrate composition of cobbles (40%), pebbles (30%), gravel (10%), and coarse sand (20%). We obtained COI gene sequences from two male and female adults of this species (Voucher specimens: 026-ESM, 027-ESM, 028-ESM, 029-ESM, unpub- lished). The genetic distance between S. cf. montana and S. nipponica ranged 1.9 2.4%.
Stenelmis nipponica Nomura, 1958 (Figures 51 56) Stenelmis nipponica Nomura 1958b: 41 (orig. descr.); Kim et al. 1994: 155 (list); Yoshi- tomi et al. 1999: 99 (note); Hayashi and Shimada 2006: 129 (note); Hayashi 2007:91 (note); Hayashi 2011: 100 (note).
Figures 51 56. (Colour online) Stenelmis nipponica Nomura, 1958: 51) dorsal habitus, male; 52) aedeagus, ventral view; 53) aedeagus, lateral view; 54) ovipositor, ventral view; 55) pronotum; 56) prosternal process. Scale bars D 0.1 mm (52 53), 0.2 mm (54), 0.5 mm (55 56), 1 mm (51). 108 S. W. Jung et al.
Type locality Japan, Shikoku, Ehime Prefecture, Matsuyama.
Type material Holotype. < ‘Japan, Matsuyama, Ehime pref., 12.IX.1948, leg. M. Miyatake’ (NSMT).
Additional material South Korea: 5<,7, ‘JN, Gure-gun, Guryo-eup, Gyesan-ri, Seomjin river, 9.VI.2005, leg. Y.C. Jeon’ (KU); 18<,24, ‘JN, Gokseong-gun, Jukgok-myeon, Namnyang-ri, Taeangyo (Br.), 9.VI.2009, leg. Y.C. Jeon’ (KU); 4<,6, ‘JB, Iksan-si, Chunpo-myeon, Mangyeong river, 3.VI.2010, leg. S.W. Jung’ (KU); 11<,13, ‘CN, Geumsan-gun, Nami-myeon, Guseok-ri, 23.VI.2010, leg. S.W. Jung’ (KU); 17<,12, ‘GB, Uljin-gun, Seo-myeon, Wangpi-ri, Soksagyo (Br.), 31.VII.2012, leg. J.K. Choi’ (KU). Japan: 1<, 1 ex. ‘Japan, Ehime-pref., Odamiyama (alt. 1000 m), Uchiko Town, 26.VII.2006, leg. T. Kurihara, det. Y. Kamite’ (KU).
Diagnosis This species can be distinguished by the combination of the following characters: body size 2.69 2.84 mm; pronotal longitudinal impression (Figure 55) slightly wider at mid- dle part, two conspicuous deep impressions in front of scutellum; prosternal process
(Figure 56) longer than wide and slightly concave apically; elytra without accessory stria, interval 6 carinate; abdominal ventrite 1 with impression near middle, in male with small tubercle at posteromedial margin. Aedeagus and ovipositor as in Figures 52 54.
Variation This species shows considerable geographical variations in the following body parts: abdominal ventrite 1 with small tubercle (Korea) or without tubercle and impression (Japan); median longitudinal impression of pronotum more or less wide (Korea) or nar- row (Japan) at posterior part; male genitalia expanded (Korea) or not expanded (Japan) at anterior part.
Distribution North Korea, South Korea, Japan.
Remarks We obtained COI gene sequences from male and female adults of this species (Voucher specimens: 005-ESN, 006-ESN, unpublished). The genetic distance between Korean and Japanese specimen ranged 0.7 1.5%. Unfortunately, we were not able to examine Stenelmis ventiplana Zhang and Yang, 2003 , described from Guangxi in China. According to its description (Zhang and Yang 2003), it might be junior synonym of S. nipponica. Aquatic Insects 109
Figures 57 61. (Colour online) Stenelmis vulgaris Nomura, 1958: 57) macropterous, female; 58) brachypterous, female; 59) aedeagus, ventral view; 60) aedagus, lateral view; 61) ovipositor, ventral view. Scale bars D 0.1 mm (59 61), 1 mm (57 58).
Stenelmis vulgaris Nomura, 1958 (Figures 57 61) Stenelmis vulgaris Nomura 1958b: 44 (orig. descr.); Kim et al. 1994: 155 (list); Yoshi- tomi et al. 1999: 99 (note); Hayashi and Shimada 2006: 131 (note); Hayashi 2007:91 (note); Hayashi 2011: 100 (note); Hayashi et al. 2013 (note, syn.); Hayashi and Yoshitomi 2014: 239 (larv. descr., syn.). Stenelmis miyamotoi Nomura and Nakane 1958: 81 (descr.); Yoshitomi et al. 1999: 100 (note); Hayashi and Shimada 2006: 132 (note); Hayashi 2007: 92 (note); Hayashi 2011: 101 (note); Hayashi et al. 2013 (syn.); Hayashi and Yoshitomi 2014 (syn.).
Type locality Japan, Honshu, Fukushima Prefecture, Wakamatsu.
Type material Holotype. < ‘Japan, Wakamatsu, Fukushima Pref., 4.VIII.1948, leg. Y. Kurosawa’ (NSMT).
Additional material North Korea: 1<,1, ‘North Korea, Sokdamguqok, 20.VIII.1989, leg. Koz anek’ (NMW). South Korea: 14<,12, ‘GN, Hamyan-gun, 23.VII.1985, leg. K.S. Bae’ (KU); 7<,5, ‘GN, Changwon-si, Pallyong-dong, 15.VIII.2010, leg. S.W. Jung & I.K. Shin’ (KU); 1<,3, ‘GG, Yongin-si, Cheoin-gu, Unhak-dong, 2.VIII.2007, leg. Y.C, Jeon’ (KU); 6<,3, ‘GB, Pohang-si, Buk-gu, Gigye-myeon, 6.VIII.2011, leg. S.W. Jung’ (KU); 14<,17, ‘GB, Pohang-si, Buk-gu, Goki-myeon, Goji-ri, Gokigyo (Br.), 6.VIII.2011, leg. S.W. Jung’ (KU). Japan: 1 ex. ‘Nagaragwa, Gifu-Pref., 19.VI.1957, leg. M. Sato,^ det. M. Sato,^ 1987’ (NMW); 1 ex. ‘Shiga, Yasu, Yasu-cho, 16.VIII.2003, leg. Y. Kamite, det. Y. Kamite’ (KU); 2 exs. ‘Shimane, Hii-gawa, Takeshi-cho, 20.IX.2006, leg. Y. Kamite, det. Y. Kamite’ (KU).
Redescription Size and shape. MF (n D 5): TL D 2.80 3.30 (3.06) mm; PL D 0.75 0.90 (0.83) mm; PW D 0.70 0.82 (0.77) mm; EL D 2.05 2.40 (2.23) mm; EW D 1.00 1.10 (1.06) mm; 110 S. W. Jung et al.
EL/EW D 2.04 2.20 (2.09); EL/PL D 2.64 2.73 (2.68); EW/PW D 1.33 1.42 (1.38); TL/EW D 2.80 3.05 (2.87). BF (n D 5): TL D 2.70 3.20 (3.00) mm; PL D 0.80 0.90 (0.86) mm; PW D 0.70 0.82 (0.76) mm; EL D 1.90 2.30 (2.13) mm; EW D 1.00 1.15 (1.01) mm; EL/EW D 1.90 2.20 (2.10); EL/PL D 2.33 2.62 (2.46); EW/PW D 1.25 1.42 (1.33); TL/EW D 2.70 3.10 (2.95). Habitus (Figures 57 58) elongate, sub- parallel-sided (MF) or oblong (BF).
Colouration. Head black; antennae, pronotum, femora and tibiae reddish brown; elytra, tarsi, and claws brown; ventral surface granulate and brown.
Morphology. Head finely punctate and granulate; distance between eyes 0.35 mm; clyp- eus wider than labrum, granulate. Antennae 11-segmented; antennomere 1 long and wide; antennomere 2 long, more slender than antennomere 1; antennomere 3 long and slender; antennomeres 4 and 5 slender, equal in length; antennomeres 6 10 wide; anten- nomere 11 long and oval, with several setae; approximate ratio of antennal segments (n D 1): 2.0 : 1.3 : 1.5 : 1.0 : 1.0 : 1.1 : 1.3 : 1.5: 1.3: 1.3: 2.0. Pronotum widest at base (MF), respectively at posterior 1/3 (BF); densely granulate; anterior angles protruding and blunt; posterior angles subacute; median longitudinal impression wide becoming abruptly narrower at base; four distinct gibbosities in lateral parts, two prebasal gibbosities coriaceous, more or less convex; oblique impressions between gibbosities; sublateral carinae distinct and comparatively long; two conspicuous deep impressions in front of scutellum. Scutellum flat, granulate. Prosternum densely granulate with small tubercle at anteromedial margin; prosternal process longer than wide, narrow at apical 1/4, slightly concave apically. Mesoventrite densely granulate, mesoventral cavity deep; mesoventral discrimen short. Metaventrite with large punctures and granules; metathoracic discrimen 0.30 mm long, metaventral cavity for reception of abdominal intercoxal process subquadrate. Legs long and slender, densely granulate except on tarsi; tibiae slender with apical spurs; claws shiny with a basal tooth. MF: elytra subparallel-sided, wider than pronotum, lateral margin slightly crenulate; humeri promi- nent, hind wings well developed; each elytron with eight punctate striae; strial punctures deep and large from base to middle becoming smaller at apex; 1st and 2nd and 3rd and 4th stria starting from same puncture near base; 2nd stria obsolete at apical 1/4; 3rd and 4th stria merging at apical 1/3; accessory stria (11 punctures) between stria 4 and stria 5 at apical 1/3; strial interval 3 distinctly convex and paler at anterior part; interval 5 paler; lateral carinae in interval 6. BF: elytra oblong, widest at posterior 1/3; humeri narrow, hind wings very short; accessory stria (4 punctures) between stria 4 and stria 5 at apical 1/3. Abdomen with five ventrites, densely granulate; abdominal intercoxal process wide, without admedian keels; ventrite 1 with large punctures, coarsely granulate; ventrite 2 sparsely granulate, with smaller punctures than on ventrite 1; ventrite 5 long, with several long setae on posterolateral part, apex truncate; lateral margin slightly crenulate. Aedea- gus as in Figures 59 60; penis longer than parameres, slightly concave at apical 1/3 (lat- eral view); lateral margin with longitudinal setose band between basal 1/3 and apical 1/3; corona present; parameres about 1.92 times as long as phallobase in lateral view, furcate in lateral view, with short dorsal branch and long ventral branch.
Female. On average larger than male; prosternum without tubercle at anterior margin; abdominal ventrite 5 concave at apex; ovipositor (Figure 61) long, more or less thick; Aquatic Insects 111 stylus long and slender; apical portion of coxite about 4.0 times as long as stylus; basal portion of coxite subquadrate; valvifer about 1.04 times as long as coxite.
Larva. See Hayashi and Yoshitomi (2014) for full description.
Distribution South Korea, North Korea, China (Liaoning), Japan.
Remarks As in many other elmid species, Stenelmis vulgaris may appear in macropterous (Figure 57) and brachypterous (Figure 58) forms (see also Hayashi et al. 2013). This spe- cies is quite common and widely distributed in Korea. According to Hayashi and Yoshi- tomi (2014), the eggs of this species are round, white, and smooth; they were found on submerged wood.
Genus Macronychus Muller,€ 1806 Type species: Macronychus quadrituberculatus Muller,€ 1806.
Macronychus levanidovae Lafer, 1980 (Figures 62 73) Macronychus levanidovae Lafer 1980: 50 (orig. descr.); Lafer 1989: 450 (key); Ciampor and Kodada 1998: 234 (redescr.).
Type locality Russia, Primorskiy Kray, Ussuriyskiy Reserve, Kamenka River.
Type material Holotype. < (ZIAS). For exact label data, see Ciampor and Kodada (1998: 234).
Additional material South Korea: 2, ‘GG, Yeoncheon-gun, Baekhak-myeon, Jeondong-ri, Samicheon (stream), 27.VI.1987, leg. K. S. Bae’ (KU).
Description Size and shape. Females (n D 2): TL D 3.09 3.17 mm; PL D 0.97 1.05 mm; PW D 0.95 1.05 mm; EL D 2.12 mm; EW D 1.35 1.36 mm; EL/EW D 1.56; EL/PL D 2.10; EW/PW D 1.35; TL/EW D 2.31. Body large and elongate. Habitus as in Figure 62.
Colouration. Body generally black. Pronotum back with paler anterior margin. Mouth parts, tarsi, claws brown.
Morphology (female). Head (Figure 64) with long pubescence. Frons with plastron; clypeus and labrum with long pubescence, anterior angles rounded. Antennae, short, 112 S. W. Jung et al.
Figures 62 73. (Colour online) Macronychus levanidovae Lafer, 1980: 62 63) dorsal and lateral habitus, female; 64) head; 65) mandible, ventral view; 66) maxilla, ventral view; 67) tarsus of hind leg; 68) metaventrite; 69) ovipositor, ventral view; 70) left elytron; 71) pronotum; 72) prosternal process; 73) abdomen. Scale bars D 0.1 mm (65 66), 0.2 mm (64, 68 69, 71 72), 0.5 mm (67,
70, 73), 1 mm (62 63).
7-segmented; antennomeres 4 6 shortest; antennomere 7 large and oval with dense setae. Mandibles (Figure 65) with three apical teeth. Maxillary palp (Figure 66) 4-segmented; palpomere 4 elongate and large, as long as palpomeres 1 3 combined. Pronotum (Figure 71) convex, widest at posterior 1/3, with two admedian gibbosities; transverse pronotal depression present; anterior angles protruding and acute; prosternal process (Figure 72) wide, lateral sides raised. Mesoventrite short, mesoventral cavity deep and square. Metaventrite (Figure 68) medially shiny, laterally granulate and with plastron, with several large punctures behind mesocoxae; metaventral cavity for reception of abdominal intercoxal process wide (0.11 mm in width). Legs (Figure 67) very long and slender; femora and tibiae granulate; tibiae with tomentum on inner surface; tarsomere 5 longer than tarsomeres 1 4 combined; approximate ratio of tarsal segments (n D 1): 2.5 : 1.0 : 2.0 : 3.5 : 11.0. Elytra (Figure 70) oblong, subparallel-sided, lateral margin serrate from posterior 1/3 to apex; each elytron with nine punctate striae; punctures moderately large and deep ante- riorly, becoming gradually smaller and shallower toward apex; strial intervals 1 5 wide; intervals 6 8 narrow; interval 3 moderately convex at anterior 1/7, with setae; interval 9 carinate and covered with plastron. Abdomen (Figure 73) with five ventrites, covered with plastron and granules laterally; abdominal intercoxal process 0.54 mm wide, raised at anterior margin; admedian keels of ventrite 1 distinct; ventrites 3 and 4 with small lateral processes; ventrite 5 as long as ven- trites 3 and 4 combined, granulate and pubescent. Aquatic Insects 113
Ovipositor (Figure 69) very long and slender; stylus elongate, narrow and curved; api- cal portion of coxite about 10 times as long as stylus, more or less wide apically; valvifer long and about 1.8 times as long as coxite.
Distribution South Korea (new record), Far East of Russia.
Remarks This species was hitherto unknown from South Korea. The specimens were collected from the demilitarized zone (DMZ) near the border to North Korea. Externally, this spe- cies resembles Macronychus vietnamensis Del eve, 1968; however, the ovipositor of M. levanidovae distinctly differs in the following characters: apical portion of coxite longer, lateral margin more concave; valvifer very long, narrow, about 1.8 times as long as coxite.
Genus Zaitzevia Champion, 1923
Type species: Zaitzevia solidicornis Champion, 1923.
Zaitzevia tsushimana Nomura, 1963 (Figures 74 81) Zaitzevia tsushimana Nomura 1963: 47 (orig. descr.). Zaitzevia nitida Nomura: Yoon 1988: 649 (misid.).
Type locality Japan, Nagasaki Prefecture, Tshushima Island, Uchiyama.
Type material Holotype. < ‘Japan, Uchiyama, Tshushima, Nagasaki Pref., 18.VIII.1960, leg. Masataka Sato’^ (NSMT).
Additional material South Korea: 6<,6,, 2 exs. ‘GW, Jeongseon-gun, Jeongsoen-eup, Shinwal-ri, 28. IV.1987, leg. K.S. Bae’ (KU); 2<,2, ‘GW, Jeongseon-gun, Gasu-ri, 29.IV.1987, leg. K. S. Bae’ (KU); 6exs. ‘GW, Cheorwon-gun, Gimhwa-eup, Amjeong-ri, 18.VI.1987, leg. K. S. Bae’ (KU); 2 exs. ‘GW, Pyeongchang-gun, Odaesan (Mt.), 12.VI.2007, leg. S.W. Jung’ (KU); 1 ex. ‘GW, Samcheok-si, Gagok-myeon, Punggok-ri, Eungbongsan (Mt.), 31. VIII.2011, leg. S.W. Jung’ (KU); 1 ex. ‘GG, Pocheon-si, Idong-myeon, Dopyeong-ri, Baegun valley, 5.VIII.2008, leg. S.W. Jung’ (KU). China: 1<, 6 exs. ‘Jilin, Changbaishan (1400m), 16.VIII.1994, leg. Ji & Wang’ (NHMW); 2, ‘Jilin, near Baihe City, 700m Erdao Baihe, 19.VIII.1994, leg. J€ach’ (NHMW). Japan: 2 exs. ‘Is. Tsushima, Uchiyama, 18»19.VII.1960, leg. M.Sato’^ (EUMJ); 2<, 2 exs. ‘Tsushima, Sasuna, 20.V.1981, leg. Y. Hori & K. Ishida’ (EUMJ); 1 ex. ‘Tsuhima, Mt.Mitake, 21.V.1981, leg. Y. Hori & K. Ishida’ (EUMJ); 1, ‘Nagasaki, Nita-gawa, Kamiagata-machi, Tsushima-shi, 10.XI.2005, 114 S. W. Jung et al.
Figures 74 81. (Colour online) Zaitzevia tsushimana Nomura, 1963: 74) dorsal habitus, male; 75) pronotum; 76) elytron, left; 77) aedeagus, ventral view; 78) ovipositor, ventral view; 79) prosternal process; 80) abdomen; 81) metaventrite. Scale bars D 0.1 mm (79, 81), 0.2 mm (75, 77 78, 80), 0.5 mm (74, 76).
Y. Kamite’ (KU). Russia: 1<,1, ‘Primorskiy Kray, Ussuriyskiy Res., Komarvoka R. by Kaminushka, 18.VII.1992, leg. A.N. Nilsson’ (NHMW); 1<, 2 exs. ‘Primorskiy Kray, Novochuguievka, 26»31.VII.1992, leg. Boukal’ (NHMW); 6 exs. ‘Primorskiy Kray, Novochuguievka, Ussuri River, 1992, leg. D. Boukal’ (NHMW).
Redescription Size and shape. Males (n D 3): TL 1.85 1.90 (1.88) mm; PL 0.42 0.45 (0.44) mm; PW 0.55 mm; EL 1.40 1.47 (1.44) mm; EW 0.75 mm; EL/EW 1.86 1.96 (1.92); EL/PL 3.11 3.50 (3.27); EW/PW 1.36; TL/EW 2.46 2.53 (2.50). Females (n D 6): TL 2.00 2.10 (2.05) mm; PL 0.50 0.55 (0.53) mm; PW 0.60 0.62 (0.60) mm; EL 1.50 1.55 (1.51) mm; EW 0.75 0.85 (0.77) mm; EL/EW 1.76 2.00 (1.95); EL/PL 2.72 3.00 (2.84); EW/PW 1.25 1.37 (1.29); TL/EW 2.41 2.73 (2.64). Habitus elongate and shiny (Figure 74).
Colouration. Head, pronotum, femora and tibiae black, elytra black or brownish-pur- plish; tarsi, claws and mouth parts brown; ventral surface dark brown. Aquatic Insects 115
Morphology. Head pubescent medially, finely punctate. Antennae short, 8-segmented; antennomere 1 short; antennomeres 2 wide and longer than 1; antenomere 3 as long as antennomere 1; antennomeres 4 7 shortest; antennomere 8 elongate with several setae. Pronotum (Figure 75) finely punctate, with long pubescence; densely granulate antero- and posterolaterally; slightly larger granules in front of scutellum; anterior margin broad; median longitudinal impression widest at middle, extending from base to apical 1/ 4; sublateral carinae sinuate smoothly, extending from base to apical 1/3; anterior angles protruding, acute; posterior angles subrectangular; lateral margin smooth. Scutellum flat, smooth, and triangular. Prosternal process (Figure 79) longer than wide, gradually taper- ing toward apex. Mesoventrite shiny, mesoventral cavity (0.06£0.10 mm) deep, broad apically, mesoventral discrimen present, mesoventral process 0.22 mm wide. Metaven- trite (Figure 81) flat and shiny medially, covered with plastron laterally, with large punc- tures behind mesocoxae and in posterior parts; metathoracic discrimen 0.26 mm long, metaventral cavity for reception of abdominal intercoxal process longer than wide (0.03 mm wide). Legs long and slender; femora granulate; tibiae slender, with tomentum on mesal surface. Elytra (Figure 76) subparallel-sided, wider than pronotum, partly cov- ered with plastron, lateral margin crenulate from middle to apex; densely granulate poste- riorly; humeri prominent, hind wings well developed; each elytron with seven punctate striae; 1st to 4th striae deeply punctate from base to middle; 5th to 7th striae with small punctures; intervals 5 7 carinate. Abdomen (Figure 80) with five ventrites, covered with plastron laterally; abdominal intercoxal process 0.21 mm wide, admedian keels of ventrite 1 distinct, parallel, extend- ing to near posterior margin; ventrite 1 with large punctures anteriorly; ventrite 3 nar- rower than ventrite 2, with several long setae posteromedially; ventrite 5 long and densely granulate, with long setae, concave at apex; lateral margin crenulate. Aedeagus (Figure 77) elongate, moderately sclerotized; ventral sac well-developed; penis more or less slender, acuminate apically, slightly curved ventrad apically (lateral view), about 2.7 times as long as phallobase; endophallus in apical 1/3 of penis, with conspicuous admedian sclerotized subapical teeth; ejaculatory duct indistinctly sclerotized.
Female. Externally similar to male; on average larger; abdominal ventrite 5 truncate apically; ovipositor (Figure 78) long and slender; stylus long and slender; apical portion of coxite about 6.0 times as long as stylus, with apicolateral parts produced lateral; valvi- fer about 1.3 times as long as coxite.
Distribution South Korea (new record), China (new record), Japan, Far East of Russia (new record).
Remarks This species is newly recorded from South Korea, China (Jilin), and Russia. It is widely distributed in mountain streams in South Korea. Two females collected from Jeju Island are slightly larger than typical specimens. Zaitzevia tsushimana is most similar to Z. riva- lis Nomura, 1963 but can be distinguished by the following morphological characters: body smaller; pronotum with median longitudinal impression widest at middle and sublat- eral carinae gently curved; scutellum triangular; elytra sparsely serrate at apical margin; penis slightly narrowed at posterior part; endophallus of male genitalia situated in apical 1/3 of penis; approximate ratio of penis and phallobase as 2.7 : 1.0. We obtained COI 116 S. W. Jung et al.
Figures 82 89. (Colour online) Zaitzeviaria kyungseoki sp. n.: 82) dorsal habitus, male; 83) aedeagus, ventral view; 84) aedeagus, lateral view; 85) apex of aedeagus; 86) pronotum; 87) pros- ternal process; 88) abdomen; 89) right hind leg. Scale bars D 0.1 mm (85), 0.2 mm (83 84, 86 89), 0.5 mm (82). gene sequences from one male and two female adults (Voucher specimens: 008-EZT,
046-EZT, 047-EZT, unpublished) to distinguish them from Z. rivalis (GenBank: GU816162.1).
Genus Zaitzeviaria Nomura, 1959 Type species: Zaitzevia brevis Nomura, 1958a.
Zaitzeviaria kyungseoki sp. n. (Figures 82 89)
Type locality South Korea, Gyeongsangsam-do (Province), Sancheong-gun.
Type material Holotype. < ‘Korea, GN, Sancheong-gun, 14.VII.1985, leg. K.S. Bae’ (KU). Dissected genitalia and other abdominal segments were glued together with the beetle on a small white card.
Description Size and shape. Male (holotype): TL D 1.60 mm; PL D 0.55 mm; PW D 0.55 mm; EL D 1.05 mm; EW D 0.65 mm; EL/EW D 1.65; EL/PL D 1.90; EW/PW D 1.18; TL/EW D 2.46. Habitus elongate and more or less stout (Figure 82).
Colouration. Body generally brown. Aquatic Insects 117
Morphology (holotype). Head with long and dense pubescence. Antennae short, 8-seg- mented; antennomere 1 short; antennomeres 2 and 3 long; antennomeres 4 7 short; antennomere 8 large and oval, with setae. Pronotum (Figure 86) quadrate with long dense pubescence; median longitudinal impression broad medially, extending to apical 1/6; sublateral carinae sinuate, extending to apical 1/3; anterior angles more or less protruding and acute; lateral margin slightly serrate. Scutellum oval, subtriangular. Prosternal process (Figure 87) longer than wide, gradually tapering posteriorly, truncate apically. Mesoventrite shiny, mesoventral cavity (0.1£0.12 mm) deep, round at apex, mesoventral discrimen short, mesoventral process 0.22 mm wide. Metaventrite flat and shiny medially, covered with plastron laterally, with large and coarse punctures behind mesocoxae and along posterior margin; metathoracic discrimen 0.16 mm long, metaventral cavity for reception of abdominal intercoxal process long and narrow. Legs (Figure 89) long and slender; femora granulate; tibiae slender, granulate, with tomen- tum on mesial surface, strong thin setae on mesial, internal, and external face. Elytra oblong, with long pubescence, slightly wider than pronotum, anterior margin dark, lateral margin crenulate, widest at posterior 2/3; hind wings absent; each elytron with eight punctate striae; 1st to 6th striae with deep and large punctures; 7th stria with small punctures; 8th stria with very small punctures; 7th and 8th intervals carinate, cari- nae almost extending to apex. Abdomen (Figure 88) with five ventrites, sparsely pubescent; abdominal intercoxal process 0.2 mm wide, admedian keels of ventrite 1 distinct, extending to posterior mar- gin; ventrites 3 and 4 with small lateral processes. Aedeagus (Figures 83 85) long and slender; ventral sac well-developed; penis about five times as long as phallobasis, wide and diamond-shaped at apex, widest sub-basally, distinctly sinuous in lateral view.
Female. Unknown.
Etymology This species is named in honour of Dr Kyung Seok Bae, who contributed a large collec- tion of Korean elmids including type material of this species.
Distribution South Korea (Sancheong-gun). The species is known only from the type locality.
Remarks This new species is similar to Zaitzeviaria kuriharai Kamite, Ogata, and Sato,^ 2006 (Kamite et al. 2006, p. 152, Figure 14), but can be separated from the latter by the dia- mond-shaped apex of the penis.
Zaitzeviaria obesa sp. n. (Figures 90 96)
Type locality South Korea, Gyeonggi-do (Province), Yeoncheon-gun, Jangnam-myeon, Wondang-ri, Samicheongyo (Br.), Imjin Stream, 38 05024.6500N, 127 00035.7700E. 118 S. W. Jung et al.
Figures 90 96. (Colour online) Zaitzeviaria obesa sp. n.: 90) dorsal habitus, female, paratype; 91) aedeagus, ventral view; 92) aedeagus, lateral view; 93) genital segment, ventral view; 94) oviposi- tor, ventral view; 95) pronotum; 96) female sternite 8. Scale bars D 0.1 mm (91 94, 96), 0.2 mm (95), 0.5 mm (90).
Type material Holotype. < ‘South Korea, GG, Yeoncheon-gun, Jangnam-myeon, Wondang-ri, Sami- cheongyo (Br.), Imjin Stream, 38 05024.6500N, 127 00035.7700E, 05.VII.2014, leg. S.W. Jung’ (KU). Paratypes.5<,5,, 25 exs., with same label data as holotype (KU); 5 exs.,
dito but deposited in NMW.
Additional material South Korea: 1, ‘Daejeon, Daedeok-gu, Miho-dong, 06.III.2008, leg. S.W. Jung & I.K. Shin’ (KU); 7<,10,, 33 exs. ‘GB, Sangju-si, Hamchang-eup, Sinheung-ri, Inangyo (Br.), 06.VII.2014, leg. S.W. Jung’ (KU).
Description Size and shape. Male (holotype): TL D 1.14 mm; PL D 0.37 mm; PW D 0.42 mm; EL D 0.77 mm; EW D 0.52 mm; EL/EW D 1.48; EL/PL D 2.08; EW/PW D 1.23; TL/EW D 2.19. Males (paratypes) (n D 5): TL D 1.14 1.20 (1.17) mm; PL D 0.34 0.37 (0.35) mm; PW D 0.40 0.46 (0.42) mm; EL D 0.80 0.85 (0.82) mm; EW D 0.50 0.54 (0.51) mm; EL/EW D 1.50 1.66 (1. 60); EL/PL D 2.18 2.50 (2.35); EW/PW D 1.13 1.27 (1.20); TL/EW D 2.18 2.33 (2.28). Females (paratypes) (n D 5): TL D 1.16 1.32 (1.24) mm; PL D 0.36 0.42 (0.38) mm; PW D 0.43 0.46 (0.50) mm; EL D 0.80 0.91 (0.86) mm; EW D 0.52 0.59 (0.56) mm; EL/EW D 1.49 1.56 (1. 54); EL/PL D 2.14 2.45 (2.27); EW/PW D 1.17 1.31 (1.24); TL/EW D 2.13 2.27 (2.22). Habitus elongate and more or less stout (Figure 90).
Colouration. Body generally black.
Morphology (holotype). Head with long pubescence. Antennae short, 8-segmented; antennomere 1 short; antennomere 2 long and wider than antenomere 1; antenomere 3 Aquatic Insects 119 shorter than antenomere 2; antennomeres 4 7 shortest; antennomere 8 large and oval, with setae. Pronotum (Figure 95) quadrate with long pubescence; anterior margin broad and paler; median longitudinal impression narrow, extending to apical 1/4; sublateral carinae gently curved, extending to middle; anterior angles more or less protruding, less than 45 wide (dorsal view); posterior angles more or less protruding and acute; lateral margin more or less smooth and subparallel-sided. Scutellum flat and triangular. Prosternal pro- cess longer than wide, gradually narrowed posteriorly, rounded apically. Mesoventrite shiny, mesoventral cavity deep, rounded apically, mesoventral discrimen short. Metaven- trite flat and shiny medially, covered with plastron laterally, with one large puncture behind mesocoxae; metathoracic discrimen long, extending near mesoventral process, metaventral cavity for reception of abdominal intercoxal process wide. Legs long and slender; tibiae slender, with apical spurs, tomentum on mesal surface. Elytra subparallel- sided, slightly wider than pronotum, with long pubescence, lateral margin slightly crenu- late; hind wings absent; each elytron with eight punctate striae; 1st to 6th striae with deep and large punctures; 7th stria with small punctures; 8th stria with large punctures anteri- orly and in the middle; 3rd stria merging with 4th at posterior 1/4.5; 5th stria merging with 6th at posterior 1/4; 7th and 8th intervals carinate. Abdomen with five ventrites, covered with plastron laterally; abdominal intercoxal process wide, admedian keels of ventrite 1 distinct; ventrites 3 and 4 with small lateral processes; ventrite 5 long, with several median and lateral setae posteriorly; lateral mar- gin crenulate; genital segment as in Figure 93. Aedeagus as illustrated (Figures 91 92); penis slender, five times as long as phalloba- sis, more or less straight in ventral view, slightly curved in lateral view, apex acute; ven- tral sac well-developed.
Female. Externally similar to male, on average larger; sternite 8 as in Figure 93 . Ovi- positor (Figure 94) long and slender; stylus more or less thick and curved; apical portion of coxite about 5.5 times as long as stylus, with apico-lateral portion produced laterad; valvifer as long as coxite.
Etymology The specific epithet is derived from the Latin adjective obesus (D obese). The name refers to the stout body form.
Distribution South Korea (Daedeok-gu, Sangju-si, Yeoncheon-gun).
Remarks This new species is allied to Zaitzeviaria gotoi from Japan. It can be distinguished from the latter by the combination of the following characters: body more stout, pubescence long; pronotum broad anteriorly; anterior pronotal angles smaller than 45 in dorsal view; sublateral carinae gently curved; prosternal process wide; elytra more deeply punctate, about 1.5 times as long as wide; metaventral cavity for reception of abdominal intercoxal process wide; penis more straight in lateral view (distinctly curved in Z. gotoi); apex slightly wider than in Z. gotoi (especially in lateral view). We obtained COI gene 120 S. W. Jung et al. sequences from both male and female adults of this species (Voucher specimens: 054- EZO, 055-EZO, unpublished). The genetic distance between Z. obesa and Z. gotoi ranged 15.5 16.0%.
Checklist of the species of Korean Elmidae Grouvellinus aerosus Jung, J€ach, and Bae, sp. n. (Korea) Heterlimnius hasegawai (Nomura, 1958) (China, Far East of Russia, Japan, Korea) Leptelmis gracilis gracilis Sharp, 1888 (Japan, Korea) Macronychus levanidovae Lafer, 1980 (Far East of Russia, Korea) Optioservus gapyeongensis Jung, Kamite, and Bae, 2011 (China, Far East of Russia, Korea) Ordobrevia sp. (Korea) Stenelmis koreana Sato,^ 1978 (Korea) Stenelmis cf. montana Zhang and Yang, 1995 (Korea) Stenelmis nipponica Nomura, 1958 (Japan, Korea) Stenelmis vulgaris Nomura, 1958 (Japan, Korea) Zaitzevia tsushimana Nomura, 1963 (China, Far East of Russia, Japan, Korea) Zaitzeviaria kyungseoki Jung, J€ach, and Bae, sp. n. (Korea) Zaitzeviaria obesa Jung, J€ach, and Bae, sp. n. (Korea)
Key to the adults of Korean species of Elmidae
1 Antennae with ten or eleven segments (Figure 25) ...... 2 - Antennae with seven or eight segments (Figure 64) ...... 10 2 Protibia with tomentum (Figure 13) on the inner side ...... 3 - Protibia without tomentum (Figure 34) on the inner side ...... 5 3 Body elongate, elytra unicoloured, without yellow markings (Figure 23) ...... Grouvellinus aerosus - Body oval, elytra with yellow markings (Figures 13, 18) ...... 4 4 Lateral pronotal margin weakly serrate, yellow spots on elytra smaller (Figures 18, 19) ...... Optioservus gapyeongensis - Lateral pronotal margin not serrate, yellow spots on elytra (if present) larger (Figures 13, 14) ...... Heterlimnius hasegawai 5 Pronotum with transverse impression and three gibbosities (Figures 33 34) ...... Leptelmis gracilis gracilis - Pronotum with median longitudinal impression (Figure 55)...... 6 6 Body length 2.2 2.3 mm, pronotum with narrow median longitudinal depression (Figure 27)...... Ordobrevia sp. - Body length 2.5 3.3 mm, pronotum with wide median longitudinal depression (Figure 48)...... 7 7 Pronotum with four distinct gibbosities (Figure 40) ...... Stenelmis koreana - Pronotum with two or without gibbosities ...... 8 Aquatic Insects 121
8 Body brown (Figures 57 58), legs about 3 mm long ...... Stenelmis vulgaris - Body black (Figure 42), legs about 2 mm long ...... 9 9 Body at least 2.7 mm long, tubercle on abdominal ventrite 1 small or absent ...... Stenelmis nipponica - Body less than 2.5 mm long, tubercle on abdominal ventrite 1 large (Figure 50) ...... Stenelmis cf. montana 10 Body at least 2.7 mm long (Figure 62), antennae with seven segments (Figure 64)...... Macronychus levanidovae - Body less than 2.2 mm long (Figure 82), antennae with eight segments ...... 11 11 Elytra with three sublateral carinae (5th to 7th intervals) (Figure 76) ...... Zaitzevia tsushimana - Elytra with two sublateral carinae (7th to 8th intervals) (Figure 90) ...... 12 12 Body about 1.2 mm long, pronotum wider than long (Figure 95) ...... Zaitzeviaria obesa - Body about 1.6 mm long, pronotum as long as wide (Figure 86) ...... Zaitzeviaria kyungseoki
Discussion In this study, 13 species of Elmidae are recognized from the Korean Peninsula, six of which are supposed to be endemic. All 13 species occur in South Korea, while only three species, Stenelmis koreana, S. nipponica, and S. vulgaris, were so far recorded from North Korea. In six Korean genera (Heterlimnius, Leptelmis, Macronychus, Optioservus, Stenelmis, and Zaitzeviaria), hind wing polymorphism occurs (Ciampor and Kodada 1998; Ogata and Nakajima 2006; Kamite 2009, 2013; Hayashi et al. 2013). In brachypterous speci- mens, the eyes are normally smaller, the pronotum is more or less longer than wide, the elytra are expanded posteriorly, the humeri are not prominent, and the patterns of the ely- tral striae are different. According to Den Boer, Van Huizen, Den Boer-Daanje, Aukema, and Den Bieman (1980), wing dimorphism may result from a process of diversifying selection depending on the stability of their habitats. The species of Stenelmis and Ordobrevia have never been revised on a global scale. According to the current definition, they can be differentiated by the larval morphology only: The prepleurite is divided in Stenelmis, while undivided in Ordobrevia, and the ter- minal abdominal segment is distinctly keeled in Stenelmis, while rounded or slightly keeled in Ordobrevia. Based on these characteristics, some species of Stenelmis might be placed in Ordobrevia. However, these characters have not yet been tested for a larger number of species. Further phylogenetic analysis using ultrastructure and molecular markers (mitochondrial COI, 28S rRNA, and Histone H3 genes) could be helpful in resolving numerous taxonomic problems in the generic classification of Elmidae. Simi- larly, the species of the genera Heterlimnius and Optioservus can hardly be distinguished based on morphological characters of adults. Very little is known about Korean elmid larvae at the species level. The larvae of four Korean species, Heterlimnius hasegawai, Leptelmis gracilis, Optioservus gapyeongensis, and Stenelmis vulgaris, were described by Kamite (2009), Jung et al. (2011), and Hayashi and Yoshitomi (2014). The larvae of the remaining nine species shall be identified by molecular (COI gene) and morphological studies in the near future. 122 S. W. Jung et al.
Acknowledgements We thank Dr Yuuki Kamite (Nagoya City Public Health Research Institute, Japan), Dr Hiroyuki Yoshitomi (EUMJ), Dr Otto Merkl (HNHM), Dr Ah Yong Kim (MSWU), Dr Jin Kyeong Choi (YU), and Mr Dae Hyen Lee (CNU) for providing type material and other valuable specimens; Dr William D. Shepard (Essig Museum of Entomology, USA), Dr Fedor Ciampor (Institute of Zoology SAV, Slovakia), and Dr German S. Lafer (Institute of Biology and Pedology, Russia) for providing useful information and references. This work was supported by a grant from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR 201401203).
Disclosure statement No potential conflict of interest was reported by the authors.
References Bollow, H. (1941), ‘Monographie der palaearktschen Dryopidae, mit Berucksichtigung€ der even- tuell transgredierenden Arten. (Col.)’, Mitteilungen der Munchner€ Entomologischen Gesell- schaft, 31, 1 88. Brown, W.J. (1930), ‘New Species of Coleoptera. I’, The Canadian Entomologist, LXII, 87 92. Champion, G.C. (1923), ‘Some Indian Coleoptera (11)’, The Entomologist’s Monthly Magazine, 59, 165 179. Ciampor, F. Jr., and Kodada, J. (1998), ‘Elmidae: I. Taxonomic Revision of the genus Macronychus Muller€ (Coleoptera)’, in Water Beetles of China. Vol. II, eds. M.A. J€ach and L. Ji, Wien: Zoo- logisch-Botanische Gesellschaft in Osterreich€ and Wiener Coleopterologenverein, pp. 219 287. Compin, A., and C er eghino, R. (2003), ‘Sensitivity of Aquatic Insect Species Richness to Distur- bance in the Adour-Garonne Stream System’, Ecological Indicators, 3, 135 142. Del eve, J. (1968), ‘Dryopidae et Elminthidae (Coleoptera) du Vietnam’, Annales Historico-Natu- rales Musei Nationalis Hungarici, 60, 149 181. Den Boer, P.J., Van Huizen, T.H.P., Den Boer-Daanje, W., Aukema, B., and Den Bieman, C.F.M. (1980), ‘Wing Polymorphism and Dimorphism in Ground Beetles as Stages in an Evolutionary Process (Coleoptera: Carabidae)’, Entomologia Generalis, 6, 107 134. Dufour, L. (1835), ‘Recherches anatomiques et consid erations entomologiques sur les insects col eopt eres des genres Macronique et Elmis’, Annales des Sciences Naturelles, 3, 151 174. Elliott, J.M. (2008), ‘The Ecology of Riffle Beetles (Coleoptera: Elmidae)’, Freshwater Reviews,1, 189 203. Fall, H.C. (1901), ‘List of the Coleoptera of Southern California, with Notes on Habits and Distribu- tion and Descriptions of new Species’, Occasional papers of the California Academy of Scien- ces,8,1 282. Gyllenhal, L. (1808), ‘Insecta Suecica descripta’, Classis I. Coleoptera sive Eleuterata 1. Scaris: Leverentz, pp. VIII C 572. Hayashi, M. (2007), ‘Distributional Records and Ecological Notes on Aquatic Coleoptera of Shi- mane Prefecture’, Bulletin of the Hoshizaki Green Foundation, 10, 77 113. (In Japanese) Hayashi, M. (2011), ‘Aquatic Coleoptera of Shimane Prefecture’, Special Bulletin of the Hoshizaki Green Foundation,1,1 117. (In Japanese) Hayashi, M., and Shimada, T. (2006), ‘Records on Elmidae and Dryopidae from Eastern Area of Shimane Prefecture and Oki Islands, Japan’, Bulletin of the Hoshizaki Green Foundation,9, 127 143. (In Japanese) Hayashi, M., Song, S.D., and Sota, T. (2013), ‘Patterns of Hind-Wing Degeneration in Japanese Rif- fle Beetles (Coleoptera: Elmidae)’, European Journal of Entomology, 110, 689 697. Hayashi, M., and Yoshitomi, H. (2014), ‘Taxonomic Treatments of two Japanese Elmid Beetles, Stenelmis vulgaris Nomura and Leptelmis gracilis Sharp (Coleoptera: Elmidae), with Descrip- tions of their Larvae’, Japanese Journal of Systematic Entomology, 20, 235 244. Hebert, P.D.N., Ratnasingham, S., and deWaard, J.R. (2003), ‘Barcoding Animal Life: Cytochrome c Oxidase Subunit 1 Divergences Among Closely Related Species’, Proceedings of the Royal Society of London, Series B-Biological Sciences (supplement), 270, 96 99. Aquatic Insects 123
Hilsenhoff, W.L. (1988), ‘Rapid Field Assessment of Organic Pollution with a Family-Level Biotic index’, Journal of the North American Benthological Society,7,65 68. Hinton, H.E. (1935), ‘Notes on the Dryopoidea (Col.)’, Stylops, 4, 169 179. J€ach, M.A., and Balke, M. (2008), ‘Global Diversity of Water Beetles (Coleoptera) in Freshwater’, Hydrobiologia, 595, 419 442. Jeng, M.L., and Yang, P.S. (1998), ‘Taxonomic Review of the genus Grouvellinus Champion (Cole- optera: Elmidae) from Taiwan and Japan’, Proceedings of the Entomological Society of Wash- ington, 100, 526 544. Jung, S.W., and Bae, Y.J. (2012), ‘Riffle Beetle genus Leptelmis (Coleoptera: Elmidae) in Korea with Description of Two new Species’, Animal Cells and systems, 16, 254 259. Jung, S.W., Kamite, Y., and Bae, Y.J. (2011), ‘Description of Optioservus gapyeongensis new Spe- cies and Heterlimnius hasegawai (Nomura) (Coleoptera: Elmidae) new to Korea’, Entomologi- cal Research, 41, 178 184. Kamite, Y. (2009), ‘A Revision of the genus Heterlimnius Hinton (Coleoptera, Elmidae)’, Japanese Journal of Systematic Entomology, 15, 199 226. Kamite, Y. (2013), ‘Revision of the genus Optioservus Sanderson, Part 1: Generic Redescription and Revision of the Nearctic species (O. fastiditus and O. quadrimaculatus Species Groups) (Coleoptera: Elmidae)’, Koleopterologische Rundschau, 83, 133 164. Kamite, Y., Ogata, T., and Sato,^ M. (2006), ‘A new Species of the genus Zaitzeviaria (Coleoptera, Elmidae) from Tsushima Islands, Japan’, The Japanese Journal of Systematic Entomology, 12, 149 153. Kim, J.I., Kwon, Y.J., Paek, J.C., Lee, S.M., An, S.R., Park, H.C., and Chu, H.Y. (1994), ‘Check List of Insects from Korea’, Kon-Kuk University Press, 23, 117 214. Kodada, J., and J€ach, M.A. (2005), ‘Chapter 18.2. Elmidae Curtis, 1830’, in Handbook of Zoology, Volume IV, Arthropoda: Insecta, Part 38, Coleoptera, Beetles, Volume 1: Morphology and Sys- tematics (Achostemata, Adephaga, Myxophaga, Polyphaga partim), eds. R.G. Beutel and R.A. B. Leschen, Berlin and New York: Walter de Gruyter, pp. 471 492. Lafer, G.S. (1980), ‘Zhuki-Pritsepyshi (Coleoptera, Dryopidae) Dal’nego Vostoka SSSR’, Fauna
presnykh vod Dal’nego Vostoka,44 53. Lafer, G.S. (1989), ‘36. Elmidae’, in Opredelitel’ nasekomykh Dal’nego Vostoka SSSR v shesti tom- akh (Tom. 3), Zhestkokrylye, ili zhuki (Chast’ 1), ed. P.A. Ler, Leningrad: Nauka, pp. 449 451. Lee, S.H., and Lee, C.E. (1992), ‘Notes on Optioservus kubotai Nomura from Korea (Coleoptera, Elmidae)’, Nature and Life, 22, 63 64. (In Korean) Martin, J.O. (1927), ‘A new Helmis (Coleoptera: Helmidae) from the Northwest’, The Pan-Pacific Entomologist, 4, 68. Muller,€ P.W.I. (1806), ‘V. Macronychus, Krallenk€afer. Eine neue K€afergattung. Mit der Beschrei- bung einer neuen Art von Hakenk€afer, Parnus’, Magazin fur€ Insektenkunde, 5, 207 220. Nomura, S. (1957a), ‘Drei neue Dryopiden-Arten aus Japan’, Akitu, Transactiions of the Kyoto Entomological Society,6,1 5. Nomura, S. (1957b), ‘Mordellid- and Elmid-Beetles of Yakushima (Coleoptera)’, Entomological Review of Japan,8,40 44. Nomura, S. (1958a), ‘Notes on Japanese Dryopoidea (Coleoptera), with two Species from Saghalien’, Toh^ o-Gakuh^ o^,8,45 59. Nomura, S. (1958b), ‘Drei neue Stenelmis-Arten aus Japan (Coleoptera, Elmidae)’, Entomological Review of Japan,9,41 45. Nomura, S. (1959), ‘Notes on the Japanese Dryopoidea (Coleoptera)’, Toh^ o-Gakuh^ o^,9,33 38. Nomura, S. (1962), ‘Some new and Remarkable Species of the Coleoptera from Japan and its Adja- cent Regions’, Toh^ o-Gakuh^ o^, 12, 35 51. Nomura, S. (1963), ‘Notes on the Dryopoidea (Coleoptera) IV’, Toh^ o-Gakuh^ o^, 13, 41 56. Nomura, S., and Nakane, T. (1958), ‘A new Species of the genus Stenelmis from Japan’, Akitu,7, 81 82. Ogata, T., and Nakajima, J. (2006), ‘Elmid Beetles of Fukuoka Prefecture, Northern Kyushu, Japan’, Bulletin of the Hoshizaki Green Foundation, 9, 227 243. Sanderson, M.W. (1953), ‘A Revision of the Nearctic Genera of Elmidae (Coleoptera)’, Journal of the Kansas Entomological Society, 26, 148 163. Sanderson, M.W. (1954), ‘A Revision of the Nearctic Genera of Elmidae (Coleoptera)’, Journal of the Kansas Entomological Society, 27, 1 13. 124 S. W. Jung et al.
Sato,^ M. (1960), ‘On the genus Leptelmis Sharp from Japan with Checklist of the World (Coleop- tera: Elmidae)’, Akitu,9,43 46. Sato,^ M. (1978), ‘Stenelmis Dufour Species of Korea (Coleoptera, Elminthidae)’, Annales Histor- ico-Naturales Musei Nationalis Hungarici, 70, 147 149. Schonfeldt,€ H.G. (1888), ‘Stenelmis foveicollis, ein neuer japanischer K€afer’, Entomologische Nachrichten, 14, 193 194. Sharp, D. (1888), ‘XXXI. Description of Some new Coleoptera from Japan’, Annals and Magazine of Natural History, 6, 242 245. Victor, T. (1839), ‘Col eopt eres du Caucase et des provinces Transcaucasiennes. (Continuation)’, Bulletin de la Societ e Imperiale des Naturalistes de Moscou, 12, 68 93. Yoon, I.B. (1988), ‘Order Coleoptera’, in Illustrated Encyclopedia of Fauna and Flora of Korea. Aquatic Insects (Vol. 30), Seoul: Ministry of Education of Korea, pp. 646 469. (In Korean) Yoshitomi, H., Shragane, A., and Hikida, N. (1999), ‘The Fauna of Elmidae and Dryopidae in the Yahagi River System’, Yahagigawa-Kenkyu,3,95 116. (In Japanese) Zhang, Z.H., and Ding, W. (1995), ‘Two new Species and a new Subspecies of Elmidae (Coleop- tera: Dryopoidea) from China’, Entomotaxonomia, 17, 15 19. (In Chinese) Zhang, Z.H., Su, H.T., and Yang, C.K. (2003a), ‘Three new Species and one new Record of Leptel- mis (Coleoptera: Dryopoidea: Elmididae)’, Entomotaxonomia, 25, 189 194. (In Chinese) Zhang, Z.H., Su, H.T., and Yang, C.K. (2003b), ‘Four new Species of Stenelmis (Coleoptera: Dryo- poidea, Elmididae) from China’, Journal of China Agricultural University, 8, 106 108. (In Chinese) Zhang, Z., and Yang, J.C. (1995), ‘Coleoptera: Dryopoidea’, in Insects and Macrofungi of Gutian- shan, Zhejiang, ed. T. Zhu, Hangzhou: Zhejiang Science and Technology Publishing House, pp. 102 110. (In Chinese) Zhang, Z., and Yang, C. (2003), ‘Four new Species of Stenelmis of Elmidae (Coleoptera, Dryopoidea)’, Acta Zootaxonomica Sinica, 28 (2), 275 281. (In Chinese)