Insect Fauna of Korea

Volume 12, Number 25 Eucnemidae

Arthropoda: Insecta: Coleoptera: Elateroidea

Flora and Fauna of Korea

National Institute of Biological Resources Ministry of Environment

Insect Fauna of Korea

Volume 12, Number 25

Eucnemidae

Arthropoda: Insecta: Coleoptera: Elateroidea

2019

National Institute of Biological Resources Ministry of Environment Insect Fauna of Korea Volume 12, Number 25 Eucnemidae Arthropoda: Insecta: Coleoptera: Elateroidea

Copyright © 2019 by the National Institute of Biological Resources Published by the National Institute of Biological Resources Environmental Research Complex, 42, Hwangyeong-ro, Seo-gu Incheon 22689, Republic of Korea www.nibr.go.kr All rights reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the National Institute of Biological Resources. ISBN: 978-89-6811-411-3(96470), 978-89-94555-00-3(Set) Government Publications Registration Number: 11-1480592-001629-01 Printed by Junghaengsa, Inc. in Korea on acid-free paper Publisher: National Institute of Biological Resources Authors: Seunghwan Lee, Jinbae Seung (Seoul National University) Project Staff: Jinwhoa YUM, Junmi Hur, Seon-Yi Kim Published on November 30, 2019 Insect Fauna of Korea

Volume 12, Number 25

Eucnemidae

Arthropoda: Insecta: Coleoptera: Elateroidea

Seunghwan Lee and Jinbae Seung Seoul National University The Flora and Fauna of Korea logo was designed to represent six major target groups of the project including vertebrates, invertebrates, insects, algae, fungi, and bacteria. PREFACE

Biological resources include all organisms and their genetic characteristics. Conservation and utiliza- tion of these resources have the capacity to improve human life and to enhance the world. Therefore it is required that the practical and potential values of these organisms are conserved and used wisely. The first step towards this goal is to document our biological diversity and to investigate them in detail. Biological resources increase the competitiveness of a nation through their use as fundamental resources for making highly valued products, such as new lines of cultivar, materials, and drugs. Many countries in the world are responsible for preserving their biodiversity and using the sustainable ways. The Ministry of Environment (MOE) of Korea has established “CBD-CHM Korea” to share of information of biodiversity. Each nation in the world is investigating and clearing information of native species within its territory. The National Institute of Biological Resources (NIBR) of MOE has published the ‘Flora and Fauna of Korea’ since 2009 to manage biological resources in comprehensive ways and to enhance national compe titiveness by building up the foundation for the sovereignty rights over biological resources. Professional research groups, consisting of professors and other taxonomic experts, have systematically examined 15,545 species of vascular plants, animals and other organisms over the past 10 years and have published their find- ings in 191 volumes in Korean and 196 volumes in English, and two volumes of monographs covering 216 species of invertebrates. This year, 21 volumes of the Flora and Fauna of Korea in both Korean and English versions including 704 species of vascular plants, invertebrates and insects are additionally published. The NIBR will continue to publish and research of flora and fauna of Korea that contribute conservation and sustainable use of biological resources. Finally, I would like to express my sincere appreciation to authors who made efforts in writing the manuscripts of the Flora and Fauna of Korea.

Prof. Dr. Yeon Jae Bae President National Institute of Biological Resources

CONTENTS

List of Taxa 2 Introduction 4 Materials and Methods 6 Taxonomic Notes 9 1. Feaia nipparensis (Hisamatsu) 11 2. Euryptychus vicinus Fleutiaux 13 3. Dromaeolus marginatus Hisamatsu 15 4. Fornax consobrinus Hisamatsu 16 5. Otho sphondyloides (Germar) 18 6. Dirrhagofarsus lewisi (Fleutiaux) 20 7. Dirrhagofarsus modestus (Fleutiaux) 21 8. Dirrhagofarsus unicolor (Hisamatsu) 22 9. Farsus ainu Fleutiaux 24 10. Microrhagus foveolatus (Fleutiaux) 26 11. Microrhagus jejuensis Seung and Lee 27 12. Microrhagus mystagogus (Fleutiaux) 28 13. Microrhagus ramosus Fleutiaux 29 14. Hylis harmandi (Fleutiaux) 31 15. Hylochares harmandi Fleutiaux 33 16. Isorhipis foveata Hisamatsu 35 17. Bioxylus natsumiae Watanabe 37 18. Bioxylus galloisi Fleutiaux 38 19. Bioxylus pilosellus Hisamatsu 39 20. Proxylobius helleri Fleutiaux 41 21. Proxylobius longicornis Hisamatsu 42 22. Xylophilus ainu (Fleutiaux) 43

Literatures Cited 45 Plates 51 Index to Scientific Names 66

1 INSECT FAUNA OF KOREA Eucnemidae

LIST OF TAXA

Class Insecta Linnaeus, 1758 Order Coleoptera Linnaeus, 1758 Suborder Polyphaga Emery, 1886 Superfamily Elateroidea Leach, 1815 Family Eucnemidae Eschscholtz, 1829 Subfamily Eucneminae Eschscholtz, 1829 Tribe Mesogenini Muona, 1993 Genus Feaia Fleutiaux, 1896 Feaia nipparensis (Hisamatsu, 1957) Subfamily Macraulacinae Fleutiaux, 1922 Tribe Euryptychini Mamaev, 1976 Genus Euryptychus LeConte, 1852 Euryptychus vicinus Fleutiaux, 1923 Tribe Macraulacini (Fleutiaux, 1923) Genus Dromaeolus Kiesenwetter, 1858 Dromaeolus marginatus Hisamatsu, 1965 Genus Fornax Laporte, 1835 Fornax consobrinus Hisamatsu, 1963 Subfamily Melasinae Fleming, 1821 Tribe Calyptocerini Muona, 1993 Genus Otho Lacordaire, 1857 Otho sphondyloides (Germar, 1818) Tribe Dirhagini Reitter, 1911 Genus Dirrhagofarsus Fleutiaux, 1935 Dirrhagofarsus lewisi (Fleutiaux, 1900) Dirrhagofarsus modestus (Fleutiaux, 1923) Dirrhagofarsus unicolor (Hisamatsu, 1960) Genus Farsus Jacquelin du Val, 1860 Farsus ainu Fleutiaux, 1923 Genus Microrhagus Dejean, 1833 Microrhagus foveolatus (Fleutiaux, 1923) Microrhagus jejuensis Seung and Lee, 2018

2 LIST OF TAXA

Microrhagus mystagogus (Fleutiaux, 1923) Microrhagus ramosus Fleutiaux, 1902 Tribe Epiphanini Muona, 1993 Genus Hylis Gozis, 1886 Hylis harmandi (Fleutiaux, 1923) Tribe Hylocharini Jacquelin du Val, 1859 Genus Hylochares Latreille, 1834 Hylochares harmandi Fleutiaux, 1900 Tribe Melasini Fleming, 1821 Genus Isorhipis Lacordaire, 1835 Isorhipis foveata Hisamatsu, 1955 Tribe Xylobiini Reitter, 1911 Genus Bioxylus Fleutiaux, 1923 Bioxylus natsumiae Watanabe, 2009 Bioxylus galloisi Fleutiaux, 1923 Bioxylus pilosellus Hisamatsu, 1959 Genus Proxylobius Fleutiaux, 1900 Proxylobius helleri Fleutiaux, 1900 Proxylobius longicornis Hisamatsu, 1958 Genus Xylophilus Mannerheim, 1823 Xylophilus ainu (Fleutiaux, 1923)

3 INSECT FAUNA OF KOREA Eucnemidae

INTRODUCTION

The family Eucnemidae (Coleoptera: Elateroidea) are a cosmopolitan group, especially tend to be more abundant and diverse in tropical areas (Muona, 2010). The family consist of approximately 1,900 species and 200 genera in the world (Otto, 2016). Much of the diversity in tropical regions is still veiled, so the virtual number of species seems to exceed 3,000 (Muona, 2010). They are commonly referred as ‘false click beetles’ to distinguish from the ‘true click beetle’, Elateridae, for a number of reasons. There is a common belief that eucnemids lack ability of ‘clicking’, but several observations of clicking-mechanism in eucnemids have been reported (Ahrens, 1812; Kirk, 1922; Bura- kowski, 1989; Muona, 1993). Clicking-mechanism of the Eucnemidae involved the contraction of muscles within the well-developed prosternal process in the mesosternal cavity allowing the beetle to jump when the connection is released (Otto, 2016). Clicking-mechanism is utilized as defensive strategy, not only escape from dangers, but also making some audible sounds to startle their enemies (Muona, 1993). However, they usually drop off to ground or pretend to be dead when disturbed, because clicking-mechanism is excessive in energy efficiency (Kaschek, 1984; Muona, 1993). The eucnemids are very similar to elaterids in adult morphology, but can be distinguished by following features: body more convex and cylindrical; second antennomere attached subterminally to first one; labrum hidden underneath frontoclypeal region; abdominal ventrite strongly connate (Laterille, 1834; Bonvouloir, 1871; Fleutiaux, 1935; Muona, 1993; Otto, 2016). Most known eucnemids are nocturnal and excellent fliers. They are also very active and rarely encountered as they are running on trunk, especially broken or barkless surface (Muona, 2010). Chemical communication is certainly important in their behavior, like the swarm- ing of Melasis buprestoides (Linnaeus, 1761) and mating behavior of other species of genera Hylochares, Hylis, and Microrhagus (Palmqvist, 1952; Muona, 2010). It is unclear whether they feed or not, however, some identified eucnemid groups lack a functional gut (Dodelin et al., 2005). Larvae are more or less sclerotized and diverse in body form, either elateriform, buprestiform, fusiform, or onisciform (Németh and Otto, 2016; Otto, 2016). They have diagnostic morphological features as following: legs reduced, close to immobile; microtrichial patches and areoles well-developed on most segments of body (Mamaev, 1976; Lucht, 1981; Burakowski, 1989; Muona and Teräväinen, 2008). They were observed to penetrate into rotten wood using their wedge-like head and microtrichial patches, with help of fluid pres- sure (Lucht, 1981). Another characteristic organ, areole is clearly verified to remove excessive water from their body (Muona and Teräväinen, 2008). These known eucnemid larvae are mostly mycetophagous, feed- ing fungal hypae in rotten wood as main food rather than wood particles (Muona and Teräväinen, 2008). They use extra-oral digestion, to break down their food with secreted digestive fluid through their mouthparts, and ingesting nutrition in liquid form (Burakowski and Buchholz, 1991; Muona and Teräväinen, 2008).

4 INTRODUCTION

Since Xylophilus ainu (Fleutiaux, 1923) was initially recorded in Korea by ESK/KSAE (1994), an additional 21 species of Eucnemidae have been recorded in Korean Peninsula to date (Table 1). Suzuki (2012) reported Isorhipis foveata Hisamatsu, 1955, sooner than after, Suzuki (2014) added three species, Dromaeolus marginatus Hisamatsu, 1965, Microrhagus ramosus Fleutiaux, 1902, and Farsus ainu Fleutiaux, 1923. More recently, Seung et al. (2017, 2018) and Seung and Lee (2018a, 2018b, 2018c, 2018d) reported 17 new records, Bioxylus natsumiae Watanabe, 2009, B. galloisi Fleutiaux, 1923, B. pilosellus Hisamatsu, 1959, Dirrha- gofarsus lewisi (Fleutiaux, 1900), D. modestus (Fleutiaux, 1923), D. unicolor (Hisamatsu, 1960), Eurypty- chus vicinus Fleutiaux, 1923, Fornax consobrinus Hisamatsu, 1963, Feaia nipparensis (Hisamatsu, 1957), Hylis harmandi (Fleutiaux, 1923), Hylochares harmandi Fleutiaux, 1900, Microrhagus fovealatus (Fleutiaux, 1923), M. jejuensis Seung and Lee, 2018, M. mystagogus (Fleutiaux, 1923), Otho sphondyloides (Germar, 1818), Proxylobius helleri Fleutiaux, 1900, and P. longicornis Hisamatsu, 1958. The aim of this work is to review of the Korean Eucnemidae according to the latest classification by Muo- na (1993). Diagnoses, descriptions and illustrations for the most species is provided with biological and dis- tributional information. Taxonomic keys to species and higher taxa level are also provided.

Table 1. History of Korean records of the family Eucnemidae

No. Author Year Scientific name Korean name

1 ESK/KSAE 1994 Xylophilus ainu (Fleutiaux) 어리방아벌레 2 Suzuki 2012 Isorhipis foveata Hisamatsu, 등노랑긴어리방아벌레 3 Suzuki 2014 Dromaeolus marginatus Hisamatsu 애기어리방아벌레 4 Farsus ainu Fleutiaux 끝털어리방아벌레 5 Microrhagus ramosus (Fleutiaux) 빗살어리방아벌레 6 Seung et al. 2017 Euryptychus vicinus Fleutiaux 굵은수염어리방아벌레 7 Fornax consobrinus Hisamatsu 넓적어리방아벌레 8 Feaia nipparensis (Hisamatsu) 큰가슴어리방아벌레 9 Seung and Lee 2018a Bioxylus natsumiae Watanabe 붉은등꼬마어리방아벌레 10 Bioxylus galloisi Fleutiaux 갈색날개꼬마어리방아벌레 11 Bioxylus pilosellus Hisamatsu 털꼬마어리방아벌레 12 Proxylobius helleri Fleutiaux 줄수염어리방아벌레 13 Proxylobius longicornis Hisamatsu 긴줄수염어리방아벌레 14 Seung and Lee 2018b Hylochares harmandi Fleutiaux 주름가슴어리방아벌레 15 Otho sphondyloides (Germar) 줄가슴어리방아벌레 16 Seung et al. 2018 Dirrhagofarsus lewisi (Fleutiaux) 큰이마줄어리방아벌레 17 Dirrhagofarsus modestus (Fleutiaux) 이마줄어리방아벌레 18 Dirrhagofarsus unicolor Hisamatsu 갈색이마줄어리방아벌레 19 Seung and Lee 2018c Microrhagus fovealatus (Fleutiaux) 큰빗살어리방아벌레 20 Microrhagus jejuensis Seung and Lee 제주빗살어리방아벌레 21 Microrhagus mystagogus (Fleutiaux) 네모빗살어리방아벌레 22 Seung and Lee 2018d Hylis harmandi (Fleutiaux) 붉은수염어리방아벌레

5 INSECT FAUNA OF KOREA Eucnemidae

MATERIALS AND METHODS

Materials for the present study were mostly collected from 2015 to 2017, from provinces of South Korea as below: Gyeonggi-do (GG), Gangwon-do (GW), Chungcheongnam-do (CN), Jeollabuk-do (JB), Jeollanam-do (JN), Jeju-do (JJ), Seoul (SO). Most specimens are deposited in the insect collection of Seoul National Uni- versity (SNU, Korea). Additional materials were based on specimens in Korea National Arboretum (KNA, Pocheon, Korea) and National Institute of Agricultural Sciences (NIAS, Wanju, Korea). The collection was mainly performed by flight intercept traps (FIT, window traps) installed in the mixed forest. Light trap, malaise trap, and collecting with naked eye were also used for collection. Collected samples were preserved in 95 % ethyl alcohol (ETOH) and later curated for exact identifications. They were pinned with the micro-pins by double-mounted method. Dissection of antennae, hind legs, and aedeagus was conducted as follows: i) Antennae dissection: Specimens were softened in boiling water for 30-60 minutes and dissected the antennae with forceps. Separated antennae were mounted on a slide glass with glycerin and covered with cover glass. ii) Hind legs dissection: Dried specimens were intenerated in boiling water for 30-60 minutes and dissected the structures with forceps. Separated samples were mounted on a slide glass with glycerin and covered with cover glass. iii) Aedeagus dissection: Dried specimens were macerated in water at room temperature for a day and occasionally pierced the side of terminal tergite with micro-pin to catalyze softening. After that, broke terminal tergite, and removed the aedeagal pouch with micro-tools and forceps. Samples were transferred to 5 ml tube with 10 % potassium- hydroxide (KOH) and boiled for a hour at 50-60°C. Dissolved samples were mounted on double cavity microscope slide with glycerin and covered with cover glass. After examination, samples were stored in polyethylene genitalia vi- als with glycerin. Dried specimens were examined under a Leica S8APO microscope (Germany). Photographs of characteristic structures were taken by a Canon EOS-600D digital camera (Japan) through MP-E 65 mm lens. Several layers of images were stacked by the Zerene Stacker version 1.04 software (Zerene Systems, USA) for res- olution improvements. Compensation of images and plates were finished with the Photoshop CC 2014 software (Adobe system, USA). Slide mounted samples were examined under a Leica DM4000B microscope (Germany) and the digital images were taken by the Active Measure version 3.0.3 software (Mitani Co. ltd, Japan) for measurement. All taxa in this study are arranged according to the latest revised classification of Eucnemidae by Muona (1993).

6 MATERIALS AND METHODS

Head Antennae Prosternum Hypomeron Pronotum Posterior angle Scutellum Mesosternum Elytron Metasternum

Abdominal ventrite

A B

Fig. 1. General morphology of Eucnemidae. A, B. Microrhagus foveolatus (Fleutiaux). Male. A. Dorsal view. B. Ventral view.

Frontoclypeal region

Mandible Prosternum Antennal groove Hypomeron

Prosternal process

Mesosternal cavity

Metasternum Metasternal groove

Metepisternum

Metacoxal plate

A B

Fig. 2. General morphology of Eucnemidae. A, B. Dirrhagofarsus modestus (Fleutiaux). A. Ventral view. B. Lateral view.

7 INSECT FAUNA OF KOREA Eucnemidae

Vertex

Frontal keel

Frons

Frontoclypeal region

Compound eye

Antennal socket

Fig. 3. General morphology of Eucnemidae. Otho sphondyloides (Germar). Frontal view.

Apical tooth

Lateral lobe

Secondary lateral lobe

Median lobe

Ventral lobe

Basal strut

Basal piece

A B

Fig. 4. General morphology of Eucnemidae. Proxylobius helleri (Fleutiaux). Aedeagus. A. Dorsal view. B. Lateral view.

8 EUCNEMIDAE

TAXONOMIC NOTES

Class Insecta Linnaeus, 1758

Order Coleoptera Linnaeus, 1758

Suborder Polyphaga Emery, 1886

Superfamily Elateroidea Leach, 1815

Family Eucnemidae Eschscholtz, 1829

Eucnemides Eschscholtz, 1829: 10. Type genus: Eucnemis Ahrens, 1812: 38.

Diagnosis: Body size small to large, 1.5-40 mm in length, variously shaped in body form; head mostly strongly deeply inserted into prothorax, labrum attached underneath frontoclypeal region and mostly in- visible; antennae consist of 11 segments diversely structured and sexually dimorphic; second antennomere smallest and subterminally attached to first; pronotum with well-developed posterior angles; elytra more or less striate, ratio of length to width 1:1.2-4; hypomeron simple or with diverse antennal groove; prosternal process more or less developed between procoxae; abdomen strongly connate, last visible ventrite variously formed at apex; aedeagus mostly trilobed type, variously modified.

Species: About 1,900.

Distribution: Worldwide (except Antarctica).

Korea: Countrywide.

Key to subfamilies of Eucnemidae in Korea

1. Hypomeron‌ with notosternal antennal groove or simple; meso- and metatibiae without row of spines on lateral surface...... Melasinae - Hypomeron‌ with lateral antennal groove or simple; meso- and metatibiae with transverse row of spines on lateral surface...... 2

9 INSECT FAUNA OF KOREA Eucnemidae

2. Hypomeron‌ with basally closed lateral antennal groove; median lobe of aedeagus always free, not fused to lateral lobes...... Eucneminae - Hypomeron‌ with basally opened lateral antennal grooves; median lobe of aedeagus usually fused with lateral lobes...... Macraulacinae

Subfamily Eucneminae Eschscholtz, 1829

Eucnemides Eschscholtz, 1829: 10. Type genus: Eucnemis Ahrens, 1812: 38.

Diagnosis: Antennae frequently increasingly serrate toward apex; hypomeron with basally closed lateral antennal groove or simple; protibiae with one spur; protarsomere 1 simple, without sex-comb in males; meso- and metatibiae with transverse rows of spines.

Tribe Mesogenini Muona, 1993

Mesogenini Muona, 1993: 49. Type genus: Mesogenus Bonvouloir, 1871: 793.

Diagnosis: Antennomeres 4-10 subequal, increasingly serrate toward apex; hypomeron with well-developed lateral antennal groove; median lobe of aedeagus modified, divided near apex.

Remarks: Mesogenini consist of 55 species in 12 genera worldwide and only a single species has been recognized from the Palaearctic region (Muona, 2011; Otto, 2016; Seung et al., 2017).

Genus Feaia Fleutiaux, 1896

Feaia Fleutiaux, 1896a: 540; Muona, 1987: 84; 1993: 50; Otto, 2016; Seung et al., 2017: 573. Type species: Feaia dubia Fleutiaux, 1896b: 603.

Diagnosis: Body oblong, strongly convex; frontoclypeal region trapezoidal, weakly trilobate or simply rounded at anterior edge; antennomere 2 slightly shorter than 3; elytra strongly striate; hypomeral antennal grooves well-developed laterally; metepisternum strongly expanded posterad; metacoxal plate subparal-

10 EUCNEMIDAE: Feaia

lel-sided; abdominal ventrite 5 truncated or strongly pointed at apex.

Distribution: Oriental region, Palaearctic region.

Remarks: This genus consists of five described species worldwide, mostly distributed in Southeast Asia. Fleutiaux (1896) first established Feaia with Feaia singularis, however the genus has remained obscure ever since its description. Muona (1987) established its identity by fixing the type species, Feaia dubia Fleuti- aux, 1896. Later, Muona (1993) included three species in Feaia from another genus Heterotaxis Bonvouloir, 1871. Otto (2016) add a new species from Laos and Seung et al. (2017) transferred a species to Feaia, Feaia nipparensis (Hisamatsu, 1957), originally placed in Heterotaxis.

1. Feaia nipparensis (Hisamatsu, 1957) (Figs. 1; Pls. I, XII)

Heterotaxis nipparensis: Hisamatsu, 1957: 45; Muona, 2007: 87. Feaia nipparensis: Seung et al., 2017: 573.

Adult: Female. Body length 5.7-6.2 mm. Mostly dull black with dark brown antennae, ferruginous pronotal posterior angle and elytral base, yellow-brown tibiae; surface covered with yellow pubescence. Head strongly inserted into prothorax; anterior edge of frontoclypeal region approximately 1.6 times wider than distance between antennal sockets. Antennae strongly serrate; antennomere 3 about 1.3 times longer than 2; 4 about 1.1 times longer than 3. Pronotum strongly convex, as long as wide, abruptly narrowed anterad from basal half; with a medio-longitudinal groove along full length of pronotum. Elytra subparallel-sided, gradually narrowed posterad; ratio of length to combined width 2.1. Prosternum strongly transverse, curved-sided, gradually widened anterad. Hypomeron with well-developed basally closed lateral antennal groove. Metepisternum triangular, distinctly widened posterad. Metacoxal plate parallel-sided, slightly concave medially. Legs slender; metatarsomere 1 as long as 2-4 combined. Abdomen strongly connate; last visible ventrite truncated at apex.

Distribution: Korea, Japan.

Distribution in Korea: GW(Seung et al., 2017).

Material examined: GW: 2♀(Suha, Daegwanryeong, Pyeongchang, 14.vii.2015, J.B. Seung). Remarks: Male was not observed. Female individuals and cadavers of F. nipparensis were observed under bark of dead Betula davurica Pallas, 1784 by authors.

11 INSECT FAUNA OF KOREA Eucnemidae

Subfamily Macraulacinae Fleutiaux, 1922

Macraulacinae Fleutiaux, 1923: 304. Type genus: Macraulacus Bonvouloir, 1871: 508.

Diagnosis: Antennae variably shaped, antennomeres 9-11 originally enlarged; hypomeron with basally opened lateral antennal groove or simple; protibiae with one spur; protarsomere 1 mostly with basal sex-comb in males; meso- and metatibiae with transverse rows of spines.

Key to tribes of Macraulacinae in Korea

1. Antennomeres 9-11 enlarged; hypomeron without antennal groove...... Euryptychini - Antennomeres 9-11 simple; hypomeron with lateral antennal groove...... Macraulacini

Tribe Euryptychini Mamaev, 1976

Euryptychini Mamaev, 1976: 154. Type genus: Euryptychus LeConte, 1852: 46.

Diagnosis: Antennomeres 9-11 distinctly enlarged; hypomeron simple, without antennal groove; protarsomere 1 with lateral sex-comb in males; median lobe of aedeagus bifurcated apically.

Remarks: Euryptychini consist of 20 species in one cosmopolitan genus. Four species have been recognized from the Palaearctic region (Muona, 2007, 2011).

Genus Euryptychus LeConte, 1852

Euryptychus LeConte, 1852: 45; Fleutiaux, 1897: 264; 1902: 661; 1923: 316; 1935: 4; Hisamatsu, 1985: 42; Muona, 2000: 66; Otto, 2016: 287; Seung et al., 2017. Dyscolocerus Bonvouloir, 1875: 729; Fleutiaux, 1902a: 661. Type species: Eucnemis heterocerus Say, 1836: 186.

Diagnosis: Body elongate, weakly convex; frontoclypeal region simply rounded at anterior edge; antennomeres 9-11 strongly enlarged; elytra moderately striate; hypomeron simple, without antennal grooves; metepisternum subparallel-sided; metacoxal plate strongly expanded inward; abdominal ventrite 5 simply

12 EUCNEMIDAE: Euryptychus

rounded at apex.

Distribution: Australian region, Holarctic region, Neotropical region.

Remarks: This genus consists of 20 described species worldwide, among them, four species are recognized from the Palaearctic region (Muona, 2007, 2011). Only a single species, Euryptychus vicinus, present in Korea.

2. Euryptychus vicinus Fleutiaux, 1923 (Figs. 2; Pls. I, XII)

Euryptychus vicinus: Fleutiaux, 1923: 324; 1935: 9; Hisamatsu, 1985: 45; Muona, 2007: 86; Seung et al., 2017: 573.

Adult: Female. Body length 7.7-9.4 mm. Mostly black with brown tarsi; surface covered with yellow-gray pubescence on dorsum, densely pubescent with golden hairs ventrally. Head moderately inserted into prothorax; frons with a short medio-longitudinal groove; anterior edge of frontoclypeal region approximately twice wider than distance between antennal sockets. Antennae with enlarged antennomeres 9-11; antennomere 3 as long as 4-5 combined. Pronotum slightly convex, as long as wide, abruptly narrowed anterad from basal three-fifths; with a weak medio-longitudinal groove at basal half. Elytra parallel-sided, gradually narrowed posterad near apex; ratio of length to combined width 2.3. Prosternum curved-sided, slightly widened anterad. Hypomeron simple, without antennal groove. Metepisternum subparallel-sided, slightly widened posterad. Metacoxal plate strongly expanded inward, medially about five times wider than laterally. Legs slender; metatarsomere 1 about 1.2 times longer than 2-4 combined. Abdomen strongly connate; last visible ventrite simply rounded at apex.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW(Seung et al., 2017).

Material examined: GG: 1♀(Jikdong, Sohol, Pocheon, 29.vi.2007, T.S. Kwon); 1♀(Jikdong, Sohol,

Pocheon, 6.viii.2008, T.S. Kwon). GW: 1♀(Cheonmi, Bangsan, Yanggu, 16.v.2015, S.H. Lee); 1♀(Beo- pheung, Suju, Yeongwol, 2-16.vii.2015, on flight intercept trap, Seung and Lee).

Remarks: Euryptychus vicinus is similar to Euryptychus lewisi Fleutiaux, 1923, but can be distinguished by following character: antennomere 3 as long as antennomeres 4-5 combined, while shorter in E. lewisi. Males were not observed. A female individual was observed wandering on surface of dead tree in May.

13 INSECT FAUNA OF KOREA Eucnemidae

Tribe Macraulacini (Fleutiaux, 1923)

Macraulacinae Fleutiaux, 1923: 304 Type genus: Macraulacus Bonvouloir, 1871: 508.

Diagnosis: Antennomeres 4-10 subequal, mostly filifrom to serrate; hypomeron with well-developed basally opened lateral antennal groove; protarsomere 1 with complete sex-comb in males; median lobe of aedeagus fused with lateral lobes.

Remarks: Macraulacini are the largest tribe in Macraulacinae, consisting of more than 200 species in 80 genera worldwide, among them, 35 species in 15 genera have been recognized from the Palaearctic region (Muona, 1993, 2007, 2011).

Key to genera of Macraulacini in Korea

1. Body‌ mostly black; basal width of metepisternum wider than outer margin of metacoxal plate...... ‌ ...... Dromaeolus - Body‌ mostly chestnut to dark brown; basal width of metepisternum narrower than outer margin of metacoxal plate...... Fornax

Genus Dromaeolus Kiesenwetter, 1858

Dromaeolus Kiesenwetter, 1858: 198; Bonvouloir, 1871: 193; Horn, 1886: 17; Fleutiaux, 1902a: 655; 1923: 299; 1935: 13; Hisamatsu, 1985: 47; Muona, 2000: 86; Otto, 2016: 347; 2017a: 4; 2017b: 16. Megathambus Reitter, 1911: 201. Melanus Broun, 1881: 676. Type species: Eucnemis barnabita A. Villa and J.B. Villa, 1838: 62.

Diagnosis: Body oblong, fairly convex; frontoclypeal region weakly trilobate at anterior edge; antennae filiform to serrate, antennomere 2 shortest, as long as 4, antennomeres 3-10 weakly toothed at least in males; elytra indistinctly striate; Hypomeron simple or with well-developed lateral antennal groove; basal width of metepisternum wider than outer margin of metacoxal plate; metacoxal plate expanded inward; abdominal ventrite 5 simply rounded or truncated at apex.

Distribution: Afrotropical region, Australian region, Holarctic region, Neotropical region, Oriental region.

Remarks: Greatest diversity of species are distributed in the tropical and subtropical regions around the globe (Otto, 2017a). This genus consists of 214 described species worldwide, including nine species in the Palaearctic region (Muona, 2007, 2011; Otto, 2017b).

14 EUCNEMIDAE: Fornax

3. Dromaeolus marginatus Hisamatsu, 1965

Dromaeolus marginatus: Hisamatsu, 1965: 134; 1985: 47; Suzuki, 2014: 44.

Adult: Male. Body length 4.8-6.0 mm. Mostly mostly black with brown antennae and red-brown legs; surface covered with yellow-brown pubescence. Head moderately inserted into prothorax; frons without median groove or keel; anterior edge of frontoclypeal region approximately three times wider than distance between antennal sockets. Antennae filiform; antennomere 3 about 1.35 times longer than 2, 1.55 times longer than 4. Pronotum 1.1 times longer than wide, abruptly narrowed anterad from basal three-fourths. Elytra gradually narrowed posterad; ratio of length to combined width 2.1-2.2. Prosternum slightly widened anterad. Hypomeron with well-developed, deep lateral antennal groove. Metepisternum narrow, slightly widened posterad. Metacoxal plate strongly expanded inward. Abdomen strongly connate; last visible ventrite obtusely pointed at apex (Hisamatsu, 1965).

Distribution: Korea, Japan, Taiwan.

Distribution in Korea: GW(Suzuki, 2014).

Material examined: None.

Remarks: Dromaeolus marginatus was firstly reported from Mt. Seorak in Korea (Suzuki, 2014). This species was not available in the present study. For detailed information and figures, see Hisamatsu’s work (1965).

Genus Fornax Laporte, 1835

Fornax Laporte, 1835: 172; Bonvouloir, 1872: 295; Fleutiaux, 1902a: 655; 1923: 293; 1935: 12; Hisamatsu, 1985: 43; Muona, 2000: 75; Otto, 2016: 287; Seung et al., 2017: 572. Filifornax Fleutiaux, 1945: 188. Monilifornax Fleutiaux, 1945: 189. Type species: Fornax ruficollis Laporte, 1835: 172.

Diagnosis: Body elongate, anteriorly and posteriorly narrowed; frontoclypeal region simply rounded or sinuate at anterior edge; antennae filiform, antennomere 3 elongate, antennomeres 4-10 increasingly slenderized toward apex; elytra indistinctly striate; hypomeral antennal grooves well-developed laterally; metepisternum narrow; metacoxal plate strongly expanded inward; abdominal ventrite 5 simply or narrowly rounded at apex.

Distribution: Afrotropical region, Australian region, Holarctic region, Neotropical region, Oriental region.

15 INSECT FAUNA OF KOREA Eucnemidae

Remarks: This genus consists of 317 described species worldwide, including six species in East Asia (Muona, 2007, 2011). Most of the species are known to develop in dead trunk or branch, which are sometimes covered with white rot (Muona, 1993).

4. Fornax consobrinus Hisamatsu, 1963 (Figs. 3; Pls. II, XII, XIII)

Fornax consobrinus: Hisamatsu, 1963: 26; 1985: 48; Muona, 2007: 86; Seung et al., 2017: 572.

Adult: Male. Body length 7.1-8.9 mm. Entirely shiny castaneous; surface covered with dense yellow pubescence. Head moderately inserted into prothorax; anterior edge of frontoclypeal region approximately 2.6 times wider than distance between antennal sockets. Antennae filiform; antennomere 3 about 1.5 times longer than 2, 1.3 times longer than 4; apical segment strongly elongate. Pronotum slightly convex, as long as wide, gradually narrowed anterad from base. Elytra subparallel-sided, gradually narrowed posterad; ratio of length to combined width 2.5. Prosternum quadrate, parallel-sided. Hypomeron with well-developed lateral antennal groove. Metepisternum subparallel-sided, slightly widened posterad. Metacoxal plate strongly expanded inward, medially about eight times wider than laterally. Legs fairly stout; metatarsomere 1 about 1.2 times longer than 2-4 combined. Abdomen strongly connate; last visible ventrite narrowly rounded at apex. Aedeagus elongate; median lobe gradually expanded apically and deeply bifurcate; lateral lobes with apical tooth, with secondary lateral lobe; secondary lateral lobe foliated. Female. Very similar to males, but can be distinguished by following characters: Frontoclypeal region wider, anterior edge approximately 2.2 times wider than distance between antennal sockets. Antennomere 3 longer, about 1.45 times longer than 4; antennomeres 4-10 stouter.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: SO (Seung et al., 2017).

Material examined: SO: 1♂(Mt. Gwanaksan, Daehak, Gwanak, 19.vi.2015, J.B. Seung; 4♂, 7♀(Mt. Gwanaksan, Daehak, Gwanak, 11.iv.2016, in last larval stage, J.B. Seung).

Remarks: Fornax consobrinus is closely related to Fornax nipponicus Fleutiaux, 1923, but can be distinguished by following character: Antennomere 3 about 1.3 times longer than 4, while about twice longer in F. nipponicus. Also, F. consobrinus is differentiated from Fornax victor Fleutiaux, 1923 by its simple claws. Straight-formed fifth instar larvae were collected in sapwood of dead Robinia pseudoacacia Linnaeus, 1753 in April. Adults were observed clicking, well-flying, and active-running.

16 EUCNEMIDAE: Otho

Subfamily Melasinae Fleming, 1821

Melasidae Fleming, 1821: 49. Type genus: Melasis A.G. Olivier, 1790: 30.

Diagnosis: Antennae variably shaped, usually sexually dimorphic; hypomeron simple or with notosternal antennal groove; protibiae with one spur; protarsomere 1 sometimes with apical sex-comb in males.

Key to tribes of Melasinae in Korea

1. Hypomeron with parallel sides...... Melasini - Hypomeron with unparallel sides, more or less narrowed anterad...... 2 2. Metacoxal plate with parallel sides or gradually expanded laterally...... 3 - Metacoxal plate gradually expanded medially...... 5 3. Antennae‌ moniliform; antennomere 3 longer than 4; pronotum always with a medio-longitudinal groove and transverse lateral impressions...... Hylocharini - Antennae‌ filiform, serrate, or pectinate; antennomere 3 distinctly shorter than 4; pronotum sometimes with a medio-longitudinal groove or transverse lateral impressions...... 4 4. Body‌ mostly large sized; antennae strongly serrate or pectinate; lateral lobes of aedeagus simply narrowed apically...... Calyptocerini - Body‌ mostly small sized; antennae filiform or serrate; lateral lobes of aedeagus usually bilobate apically‌ ...... Xylobiini 5. Pronotal lateral carina sinuate or divided; hypomeron usually with notosternal antennal groove..... Dirhagini - Pronotal lateral carina smooth and undivided; hypomeron usually simple...... Epiphanini

Tribe Calyptocerini Muona, 1993

Calyptocerini Muona, 1993: 43. Type genus: Calyptocerus Guérin-Méneville, 1843: 177.

Diagnosis: Antennae strongly serrate or pectinate; antennomere 3 small, shorter than 4; hypomeron without antennal groove; protarsomere 1 simple, without sex-comb in males; median lobe of aedeagus fused with lateral lobes.

Remarks: Calyptocerini consist of 11 species in two genera worldwide and five species in two genera have been recognized from the Palaearctic region (Muona, 2007, 2011; Otto, 2015a).

17 INSECT FAUNA OF KOREA Eucnemidae

Genus Otho Lacordaire, 1857

Otho Lacordaire, 1857: 113; Kiesenwetter, 1858: 209; Jacquelin du Val, 1863: 120; Bonvouloir, 1875: 771; Fleutiaux, 1902a: 663; 1921: 229; 1923: 315; 1935: 5; Reitter, 1911: 203; Hisamatsu, 1959: 29; 1985: 42; Muona, 1993: 61; Otto, 2016: 192; Seung and Lee, 2018b: 308. Type species: Melasis sphondyloides Germar, 1818: 235.

Diagnosis: Body elongate, cylindrical; frontoclypeal region feebly trilobate and sinuate at anterior edge; antennae strongly serrate or pectinate, antennomere 2 subequal to 3; elytra distinctly striate; hypomeron without antennal grooves; metepisternum widened posterad; metacoxal plate subparallel-sided; abdominal ventrite 5 narrowly rounded or produced at apex.

Distribution: Neotropical region, Oriental region, Palaearctic region.

Remarks: This genus consists of seven described species worldwide, among them, four species present in the Palaearctic region (Muona, 2007, 2011; Otto, 2016). Otho sphondyloides was observed on moist branch of Carpinus sp. lying on ground (Muona, 1993).

5. Otho sphondyloides (Germar, 1818) (Figs. 4; Pls. II, XII)

Melasis sphondyloides: Germar, 1818: 235. Otho spondyloides: Lacordaire, 1857: 113; Fleutiaux, 1902a: 663; 1923: 318; 1935: 8; Reitter, 1911: 207; Hisamatsu, 1959: 29; 1985: 44. Otho sphondyloides: Jacquelin du Val, 1863: 120; Bonvouloir, 1875: 772; Lucht, 1989: 182; Muona, 2007: 82; Seung and Lee, 2018b: 309. Hypocoelus sibiricus: Motschulsky, 1845: 34.

Adult: Female. Body length 6.1-8.5 mm. Mostly black with ferruginous antennae, elytral humeri, tibiae and yellow-brown tarsi; surface covered with short yellow-gray pubescnece. Head strongly inserted into prothorax; frons with a medio-longitudinal keel from vertex to anterior edge of frontoclypeal region; anterior edge of frontoclypeal region approximately 2.3 times wider than distance between antennal sockets. Antennae serrate; antennomere 3 shortest, antennomere 4 about 1.5 times longer than wide, about 1.4 times longer than 2-3 combined. Pronotum strongly convex, about 1.25 times wider than long, abruptly narrowed anterad from basal half. Elytra parallel-sided, gradually narrowed posterad near apex; with red-tinged humeri; disc strongly striate; ratio of length to combined width about 2.4. Prosternum strongly transverse, gradually widened anterad. Hypomeron simple, without antennal groove. Metepisternum distinctly widened posterad. Metacoxal plate gradually expanded outward, laterally about 1.5 times wider than medially. Legs fairly

18 EUCNEMIDAE: Otho

slender; metatarsomere 1 about 1.4 times longer than 2-4 combined. Abdomen strongly connate; last visible ventrite gradually narrowed apically, dully produced at apex.

Distribution: Korea, Japan, Russia (Far East), Europe (Austria, Belarus, Bulgaria, Croatia, Germany, Hungary, Latvia, Romania, and Ukraine).

Distribution in Korea: GW(Seung and Lee, 2018b).

Material examined: GW: 2♀(Osaek, Seo, Yangyang, 20.vi.2015, S.H. Lee); 1♀(Wangsan, Wangsan, Gangneung, 5-29.vi.2016 on flight intercept trap, Seung and Jung).

Remarks: Otho sphondyloides shows individual variations in coloration of elytral humeri and mouthparts, dark brown to ferruginous. A female individual was collected in barkless dead broad-leaved tree.

Tribe Dirhagini Reitter, 1911

Dirhagini Reitter, 1911: 202. Type genus: Dirhagus Latreille, 1834: 130.

Diagnosis: Antennae serrate or pectinate; pronotal lateral ridge serrate or divided; hypomeron mostly with notosternal antennal groove; protarsomere 1 with sex-comb in males; median lobe of aedeagus bifurcate apically.

Remarks: Dirhagini are largest group within the subfamily Melasinae with worldwide distribution. This tribe are composed of more than 300 species in 26 genera worldwide, and 34 species in 10 genera present in the Palaearctic region (Muona, 1993, 2007, 2011).

Key to genera of Dirhagini in Korea

1. Elytra‌ simply rounded at apices in lateral view; frontoclypeal region strongly narrow at base, anterior edge of frontoclypeal region more than 3.5 times wider than distance between antennal sockets...... ‌ ...... Microrhagus - Elytra‌ distinctly compressed or protruded at apices in lateral view; frontoclypeal region moderately narrow at base, anterior edge of frontoclypeal region less than three times wider than distance between‌ antennal sockets...... 2 2. Frons‌ with a pair of longitudinal keels near compound eye; antennal grooves parallel-sided; antennomere 2 shorter than 4; abdominal ventrite 5 simple, without dense hairs at middle...... Dirrhagofarsus - Frons‌ without a pair of longitudinal keels near compound eye; antennal grooves triangular, strongly expanded posterad; antennomere 2 as long as 4; abdominal ventrite 5 distinctly depressed and densely‌ pubescent at middle...... Farsus

19 INSECT FAUNA OF KOREA Eucnemidae

Genus Dirrhagofarsus Fleutiaux, 1935

Dirrhagofarsus Fleutiaux, 1935: 15; Ford and Spilman, 1979: 76; Hisamatsu, 1985: 43; Muona, 1987: 82; Muona, 2000: 38; Otto et al., 2016: 179; Seung et al., 2018: 97. Type species: Microrhagus lewisi Fleutiaux, 1900: 358.

Diagnosis: Body elongate, cylindrical; head with vertex with transverse row of vestiture; frons with a pair of longitudinal keel near compound eyes; frontoclypeal region feebly trilobate at anterior edge; antennae filiform or serrate, antennomere 2 significantly shorter than 4; elytra indistinctly striate; hypomeron with well-developed notosternal antennal groove; metepisternum narrow, subparallel-sided; metacoxal plate strongly expanded inward; abdominal ventrite 5 sharply produced at apex.

Distribution: Holarctic region, Oriental region.

Remarks: This genus consists of six species worldwide, distributed widely in the Holarctic and Oriental region, among them, four species are present in the Palaearctic region (Muona, 2007; Otto et al., 2014; Otto, 2016; Kovalev, 2016). Hisamatsu (1960) described what he considered a light-coloured form of Dirrhagus modestus as Dirrhagus modestus f. unicolor, however, this species has been identified as a distinct species (Seung et al., 2018). Immature stages and biology of Dirrhagofarsus lewisi are well-described (Ford and Spilman, 1979).

Key to species of Dirrhagofarsus in Korea

1. Antennomere‌ 3 in males more than 1.5 times longer than 4; elytra ratio length to combined width about 2.5‌ ...... D. unicolor - Antennomere‌ 3 in males less than 1.5 times longer than 4; ratio length to combined width about 2.7...... 2 2. Frons‌ with a weak medio-longitudinal keel; elytral apices simply convex in lateral view...... D. modestus - Frons‌ without medio-longitudinal keel; elytral apices distinctly protruded in lateral view...... D. lewisi

6. Dirrhagofarsus lewisi (Fleutiaux, 1900) (Figs. 5; Pls. III, XII)

Microrhagus lewisi: Fleutiaux, 1900: 358. Dirhagus lewisi: Fleutiaux, 1923: 308. Dirrhagofarsus lewisi: Fleutiaux, 1935: 16; Ford and Spilman, 1979: 75; Hisamatsu, 1960: 101; 1985: 50; Muona, 2007: 83; Otto et al., 2016: 180; Seung et al., 2018: 99.

Adult: Female. Body length 6.1-7.7 mm. Mostly brown to dark brown with ferruginous antennae and

20 EUCNEMIDAE: Dirrhagofarsus

legs; surface covered with yellow-brown pubescence. Head deeply inserted into prothorax; frons without median keel or groove; anterior edge of frontoclypeal region about 2.7 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 about 2.5 times longer than 2, 1.3 times longer than 4. Pronotum convex, as long as wide; anterolateral carina about one-fifth as long as pronotum; posterolateral carina about four-fifths as long as pronotum. Elytra parallel-sided, gradually narrowed posterad near apex; ratio of length to combined width about 2.65; apices distinctly protruded in lateral view. Pros- ternum transverse, wider than long, slightly widened anterad. Hypomeron with well-developed notosternal, parallel-sided antennal groove. Metepisternum narrow, slightly widened posterad. Metacoxal plate strongly expanded inward, medially about four times wider than laterally. Legs slender; metatarsomere 2 about 1.6 times longer than 3, as long as 5. Abdomen strongly connate; last visible ventrite acute at apex.

Distribution: Korea, Japan, Nearctic region (USA).

Distribution in Korea: GG, GW, JN (Seung et al., 2018).

Material examined: GG: 1♀(Hagui, Uiwang, 22.vi.2015, on light trap, J.B. Seung). GW: 1♀(Beopheung,

Suju, Yeongwol, 3-16.vii.2015, on flight intercept trap, Seung and Lee); 1♀ (Seorim, Seo, Yangyang,

9.vii.2016, S.H. Lee). JN: 1♀(Jungdae, Ganjeon, Gurye, 4-15.vii.2016, Seung and Lee). Remarks: Dirrhagofarsus lewisi is differentiated from other Dirrhagofarsus species by its less glossy surface of body, antennal structure, simple frons, distinctly protruded elytral apices.

7. Dirrhagofarsus modestus (Fleutiaux, 1923) (Figs. 6; Pls. III, XII, XIII)

Dirhagus modestus: Fleutiaux, 1923: 311; Muona, 2007: 85. Dirrhagus modestus: Fleutiaux, 1935: 17; Hisamatsu, 1960: 101. Rhacopus modestus: Hisamatsu, 1985: 50. Dirrhagofarsus modestus: Muona, 1993: 46; 2007: 83; Otto et al., 2016: 180; Seung et al., 2018: 102.

Adult: Male. Body length 4.5-5.9 mm. Mostly shiny black with red-brown antennae, mouthparts and legs; surface covered with yellow-gray pubescence. Head deeply inserted into prothorax; frons with a weak medio-longitudinal keel; anterior edge of frontoclypeal region about 2.9 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 about 2.5 times longer than wide, about twice longer than 2, about 1.35 times longer than 4. Pronotum moderately convex, as long as wide; disc with a medio-longitudinal keel at basal half; anterolateral carina about one-sixth as long as pronotum; posterolateral carina about four-fifths as long as pronotum. Elytra parallel-sided, gradually narrowed posterad near apex; ratio of length to combined width about 2.7; apices simply convex in lateral view. Prosternum slightly wider than long. Hypomeron with well-developed notosternal, parallel-sided antennal groove. Metepisternum

21 INSECT FAUNA OF KOREA Eucnemidae

narrow, parallel-sided. Metacoxal plate strongly expanded inward, medially about four times wider than laterally. Legs slender; metatarsomere 2 about 1.6 times longer than 3, as long as 5. Abdomen strongly connate; last visible ventrite acute at apex. Aedeagus elongate, about 5.3 times longer than wide; median lobe gradually narrowed toward apex, bifurcate at apex; lateral lobes as long as median lobe, with basally attached secondary lateral lobes; secondary lateral lobes slender, weakly pointed apically. Female. Very similar to males, but can be distinguished by following characters: Body larger than male, 5.2-6.8 mm. Frontoclypeal region wider, anterior edge about 2.7 times wider than distance between antennal sockets. Antennae relatively shorter, antennomere 3 about 1.7 times longer than 4; antennomeres 4-10 stouter.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW, JN, JJ, SO (Seung et al., 2018).

Material examined: GG: 1♀(Hagui, Uiwang, 22.vi.2015, on light trap, J.B. Seung); 2♂, 1♀(Hagui,

Uiwang, 1.vi.2016, on light trap, J.B. Seung). GW: 1♂(Beopheung, Suju, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 1♀(Deokgu, Sangdong, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 1♀(Beopheung, Suju, Yeongwol, 3-16.vii.2015, on flight intercept trap, Seung and Lee); 1♀(Hoenggye, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and

Jung); 3♀(Suha, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and Jung).

JN: 2♀(Jungdae, Ganjeon, Gurye, 4-15.vii.2016, on flight intercept traps, Seung and Lee). JJ: 2♂(Gyorae,

Jocheon, Jeju, 10.vi-21.vii.2016, on flight intercept trap, Seung and Jung). SO: 7♂, 9♀(Daehak, Gwanak, 18.i.2016, in overwintering larval stage, J.B. Seung).

Remarks: Dirrhagofarsus modestus is distinguished from other Dirrhagofarsus species by its black body coloration, antennal structure, and aedeagal characters. Mature larvae of D. modestus were observed in standing dead Alnus japonica (Thunb.) Steudel in January. They remained in U-form in oval larval cells. Adults emerged with other eucnemid species, D. unicolor and Hylis sp. in April. Also, they were commonly attracted to light trap. They were observed clicking as well as actively flying and running.

8. Dirrhagofarsus unicolor (Hisamatsu, 1960) (Figs. 7; Pls. IV, XII, XIII)

Dirrhagus modestus f. unicolor: Hisamatsu, 1960: 102. Dirrhagofarsus unicolor: Seung et al., 2018: 105.

Adult: Male. Body length 4.3-5.3 mm. Mostly brown with yellow-brown tarsi; surface covered with yellow pubescence. Head deeply inserted into prothorax; frons without median keel or groove; anterior edge of frontoclypeal region about three times wider than distance between antennal sockets. Antennae serrate;

22 EUCNEMIDAE: Farsus

antennomere 3 about 2.3 times longer than wide, about twice longer than 2, about 1.5 times longer than 4. Pronotum moderately convex, as long as wide; anterolateral carina about one-sixth as long as pronotum; posterolateral carina about four-fifths as long as pronotum. Elytra parallel-sided, gradually narrowed posterad near apex; ratio of length to combined width about 2.5; apices simply rounded in lateral view. Proster- num wider than long, parallel-sided. Hypomeron with well-developed notosternal, parallel-sided antennal groove. Metepisternum narrow, slightly widened posterad. Metacoxal plate strongly expanded inward, medially about four times wider than laterally. Legs slender; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.2 times longer than 2. Abdomen strongly connate; last visible ventrite acute at apex. Aedeagus elongate, about 4.5 times longer than wide; median lobe gradually narrowed toward apex, bifurcate at apex; lateral lobes slightly longer than median lobe, with basally attached secondary lateral lobes; secondary lateral lobes slender, subparallel-sided weakly pointed apically. Female. Very similar to males, but can be distinguished by following characters: Body slightly larger than male, 4.5-5.7 mm. Frontoclypeal region wider, anterior edge about 2.8 times wider than distance between antennal sockets. Antennomere 3 about 1.75 times longer than 4; antennomeres 4-10 stouter.

Distribution: Korea, Japan.

Distribution in Korea: GG, SO (Seung et al., 2018).

Material examined: SO: 4♂ (Daehak, Gwanak, 18.i.2016, in overwintering larval stage, J.B. Seung).

GG: 2♂ (Hagui, Uiwang, 22.vi.2015, on light trap, J.B. Seung); 1♂ (Hagui, Uiwang, 1.vi.2016, on light trap, J.B. Seung); 1♂(Hagui, Uiwang, 4.vi.2016, on light trap, M.S. Oh); 2♀(Baekgok, Mado, Hwaseong, 6-28.vi.2016, on flight intercept trap, Seung and Yeom).

Remarks: Dirrhagofarsus unicolor is distinguished from other Dirrhagofarsus species by its antennal structure, relatively short elytra, short metatarsomere 2, and aedeagal characters. Mature larvae of D. unicolor were observed in standing dead Alnus japonica (Thunb.) Steudel in January. Adults emerged with other eucnemid species, D. modestus and Hylis sp. in April. Also, they were commonly attracted to light trap. They were observed clicking as well as flying and running.

Genus Farsus Jacquelin du Val, 1860

Farsus Jacquelin du Val, 1860: 116; Bonvouloir, 1871: 71; Horn, 1890: 213; Fleutiaux, 1921: 170; 1923: 305; Hisamatsu, 1985: 43. Type species: Hylochares unicolor Latreille, 1834: 128.

Diagnosis: Body oblong, subcylindrical; frontoclypeal region feebly trilobate at anterior edge; antennae subfiliform or serrate, antennomere 2 slightly shorter or as long as 4; elytra indistinctly striate; hypomeron

23 INSECT FAUNA OF KOREA Eucnemidae

with well-developed notosternal antennal groove; metepisternum narrow, subparallel-sided; metacoxal plate expanded inward; abdominal ventrite 5 with transverse crest with dense setae.

Distribution: Afrotropical region, Neotropical region, Oriental region, Palaearctic region.

Remarks: This genus consists of 12 described species worldwide, mostly distributed in the Oriental and Palaearctic region. Among them, four species is present in the Palaearctic region (Muona, 2007; Németh and Otto, 2016). Individual adults were observed in pupal cells in fairly solid surface wood of dead log, covered with white rot (Muona, 1993). Detailed morphology of immature stage was described by Németh and Otto (2016).

9. Farsus ainu Fleutiaux, 1923 (Figs. 8; Pls. IV, XII, XIII)

Farsus ainu: Fleutiaux, 1923: 306; 1935: 15; Hisamatsu, 1985: 50; Muona, 2007: 82; Suzuki, 2014: 44.

Adult: Male. Body length 5.7-6.2 mm. Mostly dark brown with orange-brown antennae and legs; surface covered with yellow-brown pubescence. Head strongly inserted into prothorax; frons weakly depressed at middle; anterior edge of frontoclypeal region about 2.4 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 about 2.2 times longer than wide, about 1.9 times longer than 2, about 1.7 times longer than 4. Pronotum quadrate, strongly convex, as long as wide; anterolateral carina about one-fourth as long as pronotum; posterolateral carina about two-thirds as long as pronotum. Elytra parallel-sided, gradually narrowed posterad near apex; ratio of length to combined width about 2.3; apices distincly compressed and protruded in lateral view. Prosternum transverse, gradually expanded anterad. Hypomeron with well-developed notosternal, triangular antennal groove. Metepisternum narrow, parallel-sided. Metacoxal plate expanded inward, medially about 2.7 times wider than laterally. Legs slender; metatarsomere 1 about 1.1 times longer than 2-4 combined; metatarsomere 2 about 1.2 times longer than 3; metatarsomere 5 about 1.2 times longer than 2. Abdomen strongly connate; last visible ventrite with transverse crest with dense hairs at middle, rounded at apex. Aedeagus elongate, about 5.6 times longer than wide; median lobe gradually widened apically, bifurcate at apex; lateral lobes as long as median lobe, with weak apical tooth extended inward near apex.

Distribution: Korea, Japan.

Distribution in Korea: GW(Suzuki, 2014).

Material examined: GW: 1♂(Osaek, Seo, Yangyang, 20.vi.2015, S.H. Lee); 1♂(Micheon, Seo, Yang- yang, 9.vii.2015, S.H. Lee).

Remarks: Farsus ainu is readily distinguished from other Korean eucnemids by crest with dense setae at middle of abdominal ventrite 5. Male individuals were observed under bark of dead tree.

24 EUCNEMIDAE: Microrhagus

Genus Microrhagus Dejean, 1833

Microrhagus Dejean, 1833: 85; Guérin-Méneville, 1843: 171; Leconte, 1852: 46; Lacordaire, 1857: 102; Kiesenwetter, 1858: 199; Bonvouloir, 1871: 71; Horn, 1886: 11; 1890: 231; Muona, 1987: 86; Muona, 2000: 39; Seung and Lee, 2018c: 81. Dirhagus Latreille, 1834: 130; Jacquelin du Val, 1863: 116; Reitter, 1911: 202; Fleutiaux, 1921: 171; 1923: 306; 1935: 15; Hisamatsu, 1960: 92; 1985: 43; Muona, 1987: 82. Aulacostenus Motschulsky, 1870: 117. Emyirhagus Olexa, 1975: 162. Type species: Elater pygmaeus Fabricius, 1792: 234.

Diagnosis: Body oblong, narrowing craniad and caudad; frontoclypeal region strongly narrow at base; antennae serrate or pectinate, antennomere 3 not longer than 4-5 combined; elytra indistinctly striate; hypomeron with well-developed notosternal antennal groove; metepisternum subparallel-sided or widened posterad; metacoxal plate expanded inward; protarsomere 1 with apical sex-comb in males; abdominal ventrite 5 obtusely produced or simply rounded at apex.

Distribution: Worldwide (except Antarctica).

Remarks: This genus consists of 156 described species worldwide, among them, 18 species are present in the Palaearctic region (Muona, 2007, 2011; Kovalev, 2013, 2016; Otto, 2013, 2015b, 2016; Seung and Lee, 2018c). Kovalev (2016) suggested that the genus Microrhagus is diverse and heterogeneous group and needs further revision for them. Beetles were observed on dead Abies, Corylus, Eugenia, Fraxinus, Quercus, Salix (Muona, 1993). Their immature stages were well-described by Otto (2015b). All Korean Microrhagus species show sexual dimorphism with antennal structure, pectinate in males and serrate in females.

Key to species of Microrhagus in Korea

1. Frons‌ without carina or groove; pronotum nearly parallel-sided, without dimples; male antennae pectinate from antennomere 4...... M. mystagogus - Frons with keel or groove at midline; pronotum narrowed anterad, with paired dimples near middle; male antennae pectinate from antennomere 3...... 2 2. Pronotum with a longitudinal groove at midline; elytra 2.7 times longer than combined width; male ‌ antennomeres 3-10 with rami near apex; lateral lobes of aedeagus truncate...... M. jejuensis - Pronotum without groove at midline; elytra less than 2.5 times longer than combined width; male antennomere 3 with ramus at base, antennomere 4 with ramus at mid-length, antennomeres 5-10 with rami at apex; lateral lobes of aedeagus narrowly produced apically...... 3 3. Pronotum less densely punctate, average distance between punctures greater than puncture diameter; ‌

25 INSECT FAUNA OF KOREA Eucnemidae

antennal ramus 3 as long as length of antennomere 3; ventral lobe of aedeagus broadened apically...... ‌ ...... M. foveolatus - Pronotum very densely punctate, average distance between punctures smaller than puncture diameter; antennal ramus 3 about 1.4 times longer than length of antennomere 3; ventral lobe of aedeagus subparallel-sided...... M. ramosus

10. Microrhagus foveolatus (Fleutiaux, 1923) (Figs. 9; Pls. V, XII, XIII)

Dirhagus foveolatus: Fleutiaux, 1923: 308; Hisamatsu, 1985: 49. Dirrhagus foveolatus: Fleutiaux, 1935: 17; Hisamatsu, 1960; Medvedev, 1970: 168. Microrhagus foveolatus: Muona, 2007: 82; Seung and Lee, 2018c: 84.

Adult: Male. Body length 5.1-6.0 mm. Mostly shiny black with yellow-brown tarsi; surface covered with yellow pubescence. Head moderately inserted into prothorax; frons with a weak medio-longitudinal keel; anterior edge of frontoclypeal region about 3.7 times wider than distance between antennal sockets. Antennae pectinate from antennomere 3; rami of antennomeres 3, 4, and 5 about 1.1, 2.1 and 2.1 times as long as corresponding antennomeres; antennomere 3 with ramus near base, 1.7 times longer than 2, and 1.3 times longer than 4; antennomere 4 with ramus at mid-length; antennomeres 5-10 with rami near apex. Pronotum 1.2 times wider than long; average distance between punctures larger than puncture diameter; disc with a pair of dimples near middle; anterolateral carina exceeding half as long as pronotum; posterolateral carina almost reaching half as long as pronotum. Elytra gradually narrowed posterad; ratio of length to combined width about 2.5; apices simply rounded. Prosternum with curved sides. Hypomeron with well-developed notosternal antennal groove. Metepisternum gradually widened posterad. Metacoxal plate expanded inward, medially about 2.3 times wider than laterally. Legs slender; metatarsomere 1 about 1.2 times longer than 2-4 combined; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.7 times longer than 2. Abdomen strongly connate; last visible ventrite narrowly rounded at apex. Aedeagus elongate, about 4.3 times longer than wide; median lobe bifurcate at apex; lateral lobes as long as median lobe, blunt at apex; ventral lobe as long as median lobe, gradually expanded apically, broadly truncate at apex, with dense pubescence. Female. Can be distinguished from males by following characters: Body length 5.3-6.4 mm. Frontoclypeal region wider, anterior edge about 3.5 times wider than distance between antennal sockets. Antennae serrate; antennomere 2 short, as long as 4; antennomere 3 about 1.9 times longer than wide, 1.7 times longer than each length of 2 and 4; antennomeres 4-10 gradually increasing in length; apical antennomere about 3.5 times longer than wide, 2.2 times longer than previous.

26 EUCNEMIDAE: Microrhagus

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW, JJ, SO (Seung and Lee, 2018c).

Material examined: GG: 1♀(Deoksu, Danwol, Yangpyeong, 8-22.v.2016, on flight intercept trap, Seung and Jung). GW: 1♀(Suha, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and Jung); 1♀(Hoenggye, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and

Jung). JJ: 1♂, 1♀(Gyorae, Jocheon, Jeju, 10.vi.2016, J.B. Seung); 2♂ (Gyorae, Jocheon, Jeju, 13.v-10. vi.2016, Seung and Jung). SO: 1♀(Gil, Gangdong, 16.v.2016, B.H. Jung). Remarks: Microrhagus foveolatus is distinguished from M. ramosus by its relatively shorter antennal rami, sparser punctures of pronotum, relatively more elongate elytra, and aedeagal structure. Adults were observed on a standing dead tree with peeling loose bark which were covered with hyphae.

11. Microrhagus jejuensis Seung and Lee, 2018 (Figs. 10; Pls. V, XII, XIII)

Microrhagus jejuensis: Seung and Lee, 2018c: 86.

Adult: Male. Body length 4.9-5.2 mm. Mostly shiny black with red-brown antennal rami, mouthparts, tibiae and yellow-brown tarsi; surface covered with yellow pubescence. Head moderately inserted into prothorax; frons with a short median groove; anterior edge of frontoclypeal region about 3.9 times wider than distance between antennal sockets. Antennae pectinate from antennomere 3; rami of antennomeres 3, 4, and 5 about 1.1, 1.7 and 1.8 times as long as corresponding antennomeres; antennomere 3 about 1.7 times longer than 2, as long as 4; antennomeres 3-10 with rami near apex. Pronotum 1.3 times wider than long; disc with a pair of dimples at middle and a groove along midline; anterolateral carina almost one-third as long as pronotum; posterolateral carina three-fifths as long as pronotum. Elytra parallel-sided, gradually narrowed near apices; ratio of length to combined width about 2.7; apices simply rounded. Prosternum with curved sides, slightly widened anterad. Hypomeron with well-developed notosternal antennal groove. Metepisternum narrow, gradually widened posterad. Metacoxal plate expanded inward, medially about 1.6 times wider than laterally. Legs slender; metatarsomere 1 about 1.5 times longer than 2-4 combined; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.2 times longer than 2. Abdomen strongly connate; last visible ventrite narrowly rounded at apex. Aedeagus elongate, about 4.3 times longer than wide; median lobe broadly bifurcate at apex; lateral lobes slender with inward apical tooth, truncate at apex with long setae. Female. Can be distinguished from males by following characters: Body length 4.8-6.1 mm. Frontoclypeal region wider, anterior edge about 3.7 times wider than distance between antennal sockets. Antennae serrate; antennomere 3 about twice longer than 2, about 1.3 times longer than 4; antennomeres 4-10 gradually increasing in length; apical antennomere about 5.3 times longer than wide, 1.8 times longer than 10.

27 INSECT FAUNA OF KOREA Eucnemidae

Distribution: Korea.

Distribution in Korea: JJ (Seung and Lee, 2018c).

Material examined: JJ: 1♂(Hwasun, Andeok, Seogwipo, 12.v.2016, J.B. Seung); 1♀(Sanghyo, Seogwi- po, 12.v.2016, J.B. Seung); 2♀(Gyorae, Jocheon, Jeju, 12.v-10.vi.2016, on flight intercept trap, Seung and

Jung); 2♂, 1♀(Hwasun, Andeok, Seogwipo, 12.v-10.vi.2016, on flight intercept trap, Seung and Jung). Remarks: Microrhagus jejuensis similar to M. foveolatus, but can be distinguished from M. foveolatus by a median groove on frons, median groove on pronotum, elongate elytra, short lateral lobes of aedeagus. The structure of aedeagus resembles that of Microrhagus pectinicornis Hisamatsu, 1960, but the latter species differs from the new species in longer rami of antennomeres (rami of antennomeres 3, 4, and 5 about 2.2, 3.6, and 4 times as long as corresponding antennomeres). Also, each ramus of antennomeres 3 and 4 is near base in M. pectinicornis, and not in M. jejuensis.

12. Microrhagus mystagogus (Fleutiaux, 1923) (Figs. 11; Pls. VI, XII, XIII)

Dirhagus mystagogus: Fleutiaux, 1923: 308; Hisamatsu, 1985: 49. Dirrhagus mystagogus: Fleutiaux, 1935: 17; Hisamatsu, 1960: 90. Microrhagus mystagogus: Muona, 2007: 83; Kovalev, 2013: 269; Seung and Lee, 2018c: 89.

Adult: Male. Body length 3.2-3.9 mm. Mostly black with red-brown antennae, femur and yellow-brown tibiae and tarsi; surface covered with short yellow pubescence. Head deeply inserted into prothorax; frons without median keel or groove; anterior edge of frontoclypeal region about 3.9 times wider than distance between antennal sockets. Antennae pectinate from antennomere 4; rami of antennomeres 4, 5, and 6 about 1.4, 2.3 and 2.6 times as long as corresponding antennomeres; antennomere 3 about three times longer than 2, 1.8 times longer than 4; antennomeres 4-10 with rami near apex. Pronotum as long as wide; anterolateral carina almost one-third as long as pronotum; posterolateral carina two-thirds as long as pronotum. Elytra parallel-sided, gradually narrowed near apices; ratio of length to combined width about 2.3; apices simply rounded. Prosternum with subparallel-sided, slightly bisinuate at anterior edge. Hypomeron with well- developed notosternal antennal groove. Metepisternum gradually widened posterad. Metacoxal plate expanded inward, medially about 1.8 times wider than laterally. Legs slender; metatarsomere 1 about 1.3 times longer than 2-4 combined; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.2 times longer than 2. Abdomen strongly connate; last visible ventrite simply rounded at apex. Aedeagus about 3.9 times longer than wide; median lobe slightly bent ventrally; lateral lobes strongly bent ventrally, enlarged near apex with dense seta. Female. Can be distinguished from males by following characters: Body length 3.4-3.9 mm. Frontoclypeal region wider, anterior edge about 3.7 times wider than distance between antennal sockets. Antennae serrate;

28 EUCNEMIDAE: Microrhagus

antennomere 3 about 2.3 times longer than 2, 1.6 times longer than 4; antennomeres 4-10 gradually more strongly serrate; apical antennomere about 3.5 times longer than wide, twice longer than previous.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW(Seung and Lee, 2018c).

Material examined: GG: 1♀(Deoksu, Danwol, Yangpyeong, 29.vi-16.vii.2016, on flight intercept trap,

Seung and Jung). GW: 1♀(Yeongheung, Yeongwol, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap,

Seung and Lee); 1♂(Beopheung, Suju, Yeongwol, 3-16.vii.2015, on flight intercept trap, Seung and Lee);

1♂, 2♀(Beopheung, Suju, Yeongwol, 5-29.vi.2016, on flight intercept trap, Seung and Jung). Remarks: Microrhagus mystagogus is readily distinguished from other Korean Microrhagus species by pectinate structure from antennomere 4 in males. Also, convex pronotum and structure of aedeagus is characteristic. Its aedeagal structure is well-illustrated by Kovalev (2013).

13. Microrhagus ramosus Fleutiaux, 1902 (Figs. 12; Pls. VI, XII, XIII)

Microrhagus ramosus: Fleutiaux, 1902b: 24; Muona, 2007: 83; Suzuki, 2014: 44; Seung and Lee, 2018c: 91. Dirhagus ramosus: Fleutiaux, 1923: 308. Dirrhagus ramosus: Fleutiaux, 1935: 17; Hisamatsu, 1960: 95; 1985: 49.

Adult: Male. Body length 3.3-4.8 mm. Mostly black with red-brown antennal rami, tibiae and yellow- brown tarsi; surface covered with yellow pubescence. Head moderately inserted into prothorax; frons with a weak medio-longitudinal keel; anterior edge of frontoclypeal region about 4.2 times wider than distance between antennal sockets. Antennae pectinate from antennomere 3; rami of antennomeres 3, 4, and 5 about 1.4, 2.5 and 2.4 times as long as corresponding antennomeres; antennomere 3 with ramus near base, 1.7 times longer than 2, and 1.3 times longer than 4; antennomere 4 with ramus at mid-length; antennomeres 5-10 with rami near apex. Pronotum 1.1 times wider than long; average distance between punctures smaller than puncture diameter; disc with a pair of dimples at middle; anterolateral carina reaching half as long as pronotum; posterolateral carina almost exceeding half as long as pronotum. Elytra gradually narrowed posterad; ratio of length to combined width about 2.2; apices simply rounded. Prosternum with curved sides. Hypomer- on with well-developed notosternal antennal groove. Metepisternum gradually widened posterad. Metacoxal plate expanded inward, medially about 2.3 times wider than laterally. Legs slender; metatarsomere 1 about 1.3 times longer than 2-4 combined; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.5 times longer than 2. Abdomen strongly connate; last visible ventrite narrowly rounded at apex. Aedeagus elongate, about five times longer than wide; median lobe bifurcate at apex; lateral lobes as long as median lobe; ventral lobe shorter than median lobe, parallel-sided, truncate at apex, with dense seta.

29 INSECT FAUNA OF KOREA Eucnemidae

Female. Can be distinguished from males by following characters: Body length 3.2-4.3 mm. Frontoclypeal region wider, anterior edge about four times wider than distance between antennal sockets. Antennae serrate; antennomere 2 short, as long as 4; antennomere 3 about 1.9 times longer than wide, 1.7 times longer than each length of 2 and 4; antennomeres 4-10 gradually increasing in length; apical antennomere about 3.5 times longer than wide, 2.2 times longer than 10.

Distribution: Korea, Japan.

Distribution in Korea: GW, JN, JJ (Seung and Lee, 2018c).

Material examined: GW: 3♂, 1♀(Beopheung, Suju, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 2♂, 1♀(Deokgu, Sangdong, Yeongwol, 19.vi-2.vii, 2015, on flight intercept trap,

Seung and Lee); 3♂(Deokgu, Sangdong, Yeongwol, 2-16.vii.2015, on flight intercept trap, Seung and Lee);

2♂(Hoenggye, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and Jung); 1♂

(Jindong, Girin, Inje, 13.vii.2016, M.S. Oh). JN: 1♂(Donggok, Ongnyong, Gwangyang, 4-15.vii.2016, on flight intercept trap, Seung and Lee). JJ: 1♂ (Gyorae, Jocheon, Jeju, 13.v-10.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Microrhagus ramosus shows individual morphological variation as below: pronotal anterolateral carina and posterolateral carina obscure in some, appearing fused in some; ventral lobe of aedeagus sub-parallel sided in some, slightly widened near apex.

Tribe Epiphanini Muona, 1993

Epiphanini Muona, 1993: 45. Type genus: Epiphanis Eschscholtz, 1829: 35.

Diagnosis: Hypomeron gradually narrowed anterad, without antennal groove; protarsomere 1 simple, without sex-comb in males; median lobe of aedeagus deeply bifurcate at apex, fused with lateral lobes.

Remarks: Epiphanini consist of 21 species in two worldwide genera, with 12 species in both genera present in the Palaearctic region (Muona, 2007, 2011).

Genus Hylis Gozis, 1886

Hylis Gozis, 1886: 21; Méquignon, 1922: 372; Muona, 1987: 85; 2000: 34; Burakowski, 1991: 63; Otto, 2016: 205; Seung and Lee, 2018d (in press).

30 EUCNEMIDAE: Hylis

Hypocoelus Guérin-Méneville, 1843: 165; Lacordaire, 1857: 102; Kiesenwetter, 1858: 108; Bonvouloir, 1875: 694; Fleutiaux, 1902a: 661; 1923: 315; 1935: 4; Reitter, 1911: 202; Hisamatsu, 1985: 42. Hypohylis Reitter, 1911: 203. Elatocoelus Hyslop, 1921: 644. Type species: Eucnemis procerulus Mannerheim, 1823: 32.

Diagnosis: Body oblong, narrowing craniad and caudad; frontoclypeal region mostly rounded at anterior edge; antennae more or less serrate, more slender in males than those of female, antennomeres mostly keeled medially, antennomere 4 shorter than 2-3 combined; elytra indistinctly striate; hypomeral antennal grooves absent; metepisternum subparallel-sided, slightly widened posterad; metacoxal plate expanded inward; abdominal ventrite 5 simply rounded or pointed at apex.

Distribution: Australian region, Holarctic region, Oriental region.

Remarks: This genus consists of 18 described species worldwide. Among them, 11 species are present in the Palaearctic region (Muona, 2007, 2011; Otto, 2016).

14. Hylis harmandi (Fleutiaux, 1923) (Figs. 13; Pls. VII, XII, XIII)

Hypocoelus harmandi: Fleutiaux, 1923: 326; 1935: 10; Hisamatsu, 1985: 45. Hylis harmandi: Muona, 2007: 83; Seung and Lee, 2018d (in press).

Adult: Male. Body length 3.4-3.9 mm. Mostly black with chestnut-coloured antennae, tibiae, tarsi; surface rarely covered with yellow-brown pubescence. Head weakly inserted into prothorax; frons with a distinct crest with dense setae; anterior edge of frontoclypeal region broadly rounded, about 2.5 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 about 1.45 times longer than wide, 1.85 times longer than 2, and 1.2 times longer than 4; antennomeres 4-10 gradually lengthened; apical antennomere 2.9 times longer than wide, 1.65 times longer than 10. Pronotum abruptly narrowed anterad from basal two-thirds, about 1.2 times wider than long; disc with a weak medio-longitudinal groove at basal half. Elytra subparallel-sided, gradually narrowed posterad; ratio of length to combined width about 2.3; apices simply rounded. Prosternum gradually widened anterad. Hypomeron simple, without antennal groove. Metepisternum gradually widened posterad. Metacoxal plate expanded inward, medially about 2.3 times wider than laterally. Legs slender; metatarsomere 1 about 1.4 times longer than 2-4 combined; metatarsomere 2 about 1.4 times longer than 3; metatarsomere 5 about 1.1 times longer than 2. Abdomen strongly connate; last visible ventrite truncate at apex. Aedeagus about 4.6 times longer than wide, pubescent at dorsal surface; median lobe elongate, deeply bifurcate at apex; lateral lobes slender, with inward apical tooth. Female. Can be distinguished from males by following characters: Body length 3.9-4.7 mm. Frons with

31 INSECT FAUNA OF KOREA Eucnemidae

indistinct crest, simply pubescent. Frontoclypeal region wider, anterior edge about 2.2 times wider than distance between antennal sockets. Antennae almost moniliform; antennomere 3 about 1.5 times longer than wide, about 1.7 times longer than 2, 4; antennomeres 4-10 stouter; apical antennomere about twice longer than wide.

Distribution: Korea, China (Northeast), Japan, Russia (Far East).

Distribution in Korea: GW(Seung and Lee, 2018d).

Material examined: GW: 1♂, 1♀(Yeongheung, Yeongwol, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 1♀(Yeongheung, Yeongwol, Yeongwol, 2-16.vii.2015, on flight intercept trap,

Seung and Lee); 1♂ (Yeongheung, Yeongwol, Yeongwol, 16-30.vii.2015, on flight intercept trap, Seung and Lee).

Remarks: Hylis harmandi is characteristic by distinct crest with dense setae on the frons in males. Hisa- matsu (1985) commented that length of antennomeres 4-5 combined is about 1.7-1.9 times longer than 3. As a result of the present study, ratio of 1.7 is corresponded to males and 1.9 to females.

Tribe Hylocharini Jacquelin du Val, 1859

Hylocharini Jacquelin du Val, 1859: 119. Type genus: Hylochares Latreille, 1834: 127.

Diagnosis: Antennae weakly serrate or moniliform; pronotum with transverse impression; hypomeron without antennal groove; protarsomere 1 simple, without sex-comb in males; aedeagus bulbous.

Remarks: Hylocharini consist of five species, including two undescribed species, in only one genus worldwide. Among them, three species is present in the Palaearctic region (Muona, 2007, 2011; Brüstle and Muona, 2009).

Genus Hylochares Latreille, 1834

Hylochares Latreille, 1834: 127; Guérin-Méneville, 1843: 171; LeConte, 1852: 46; Lacordaire, 1857: 102; Kiesenwetter, 1858: 209; Jacquelin du Val, 1863: 120; Bonvouloir, 1875: 766; Horn, 1886: 11; Fleutiaux, 1921: 229; 1923: 316; 1935: 5; Hisamatsu, 1985: 42; Seung and Lee, 2018b: 308. Xylophilus Mannerheim, 1823: 14; Eschscholtz, 1829: 35; Reitter, 1911: 203. Type species: Elater cruentatus Gyllenhal, 1808: 435.

32 EUCNEMIDAE: Hylochares

Diagnosis: Body oblong, cylindrical, with glossy surface; frontoclypeal region trilobate at anterior edge; antennae weakly serrate or moniliform, antennomere 3 about twice longer than 2; pronotum with transverse impression at middle; elytra distinctly striate; hypomeral antennal grooves absent; metepisternum gradually widened posterad; metacoxal plate subparallel-sided; abdominal ventrite 5 narrowed or beaked at apex.

Distribution: Holarctic region, Oriental region.

Remarks: This genus, consisting of five described species worldwide, is distributed largely in Holarctic region and Oriental region. Among them, three species are present in the Palaearctic region (Muona, 2007, 2011; Brüstle and Muona, 2009). Hylochares cruentatus (Gyllenhal, 1808) has been known to be a specialist of large standing dead wood with white rot, such as willow (Mannerheim, 1823; Kangas and Kangas, 1944). Detailed biology of Hylochares cruentatus was described by Muona and Brüstle (2008).

15. Hylochares harmandi Fleutiaux, 1900 (Figs. 14; Pls. VII, XII, XIII)

Hylochares harmandi: Fleutiaux, 1900: 360; 1935: 11; Hisamatsu, 1985: 45; Muona, 2007: 84. Seung and Lee, 2018b: 310. Xylophilus harmandi: Medvedev, 1970: 168.

Adult: Male. Body length 4.8-6.6 mm. Mostly glossy black with ferruginous antennae and dark brown legs; surface rarely covered with short yellow-brown pubescence. Head deeply inserted into prothorax; frons with a distinct longitudinal impression; anterior edge of frontoclypeal region distinctly trilobate, about 2.3 times wider than distance between antennal sockets. Antennae moniliform; antennomere 3 about 1.65 times longer than wide, 1.6 times longer than 2, and 1.4 times longer than 4; antennomeres 4-10 subequal, as long as wide; apical antennomere 1.7 times longer than wide, 1.5 times longer than 10. Pronotum strongly convex, 1.1 times wider than long; disc with transverse depressions longitudinally and transversely at middle. Elytra parallel-sided, gradually narrowed near apices; disc strongly striate with rows of regular and deep punctures; ratio of length to combined width about 2.3; apices simply rounded. Prosternum transverse, slightly widened anterad. Hypomeron simple, without antennal groove. Metepisternum gradually widened posterad. Metacoxal plate slightly expanded inward, medially about 1.2 times wider than laterally. Legs fairly robust; length of metatarsomeres 2-4 combined about 1.2 times longer than 1; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.8 times longer than 2. Abdomen strongly connate; last visible ventrite distinctly beaked at apex. Aedeagus bulbous, about 3.5 times longer than wide; median lobe stout, simply rounded at apex; lateral lobes shorter than median lobe, rarely setose apically. Female. Very similar to males, but can be distinguished by following characters: Body length 6.3-8.8 mm. Antennomere 3 more elongate, about 1.9 times longer than wide, 1.7 times longer than 4; apical segment about twice longer than wide, 1.7 times longer than previous. Metatarsomere 1 as long as 2-4 combined;

33 INSECT FAUNA OF KOREA Eucnemidae

metatarsomere 2 about 1.6 times longer than 3.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW, CN, JJ (Seung and Lee, 2018b).

Material examined: GG: 2♂ (Seungan, Gapyeong, Gapyeong, 8.vi.2009, H.C. Park); 1♀ (Samhoe,

Cheongpyeong, Gapyeong, 27.v.2016, S.H. Lee). GW: 1♀(Beopheung, Suju, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 1♀(Wangsan, Wangsan, Gangneung, 5-29.vi.2016, on flight intercept trap, Seung and Jung). CN: 1♂(Bunam, Sindoan, Gyeryong, 30.v.1987, U.H. Baek); 1♂(Haso, Dong,

Daejeon, 6.vi.2016, M.S. Oh). JJ: 1♂ (Gyorae, Jocheon, Jeju, 12.v-10.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Hylochares harmandi can be easily distinguished from other Korean eucnemid species by characteristic frons, antennae, pronotum, elytral striae, and apical structure of abdominal ventrite 5. Adults were observed wandering on fallen timbers in May and June. Individual variation in body size is apparent between male and female (smallest male: 4.8 mm; largest female: 8.8 mm).

Tribe Melasini Fleming, 1821

Melasidae Fleming, 1821: 49. Type genus: Melasis A. G. Olivier, 1790: 30.

Diagnosis: labrum partly visible; antennae pectinate or flabellate in males, serrate in females; hypomeron without antennal groove; protarsomere 1 simple, without sex-comb in males; aedeagus bulbous, median lobe narrowly pointed at apex.

Remarks: Melasini consist of 44 species in three genera worldwide, and 13 species in two genera are present in the Palaearctic region (Muona, 2007, 2011).

Genus Isorhipis Lacordaire, 1835

Isorhipis Lacordaire, 1835: 620; Horn, 1890: 211; Fleutiaux, 1921: 225; 1923: 315; 1935: 3; Hisamatsu, 1985: 42; Muona, 2000: 24. Compsocnemis Bonvouloir, 1871: 72; Fleutiaux, 1902a: 659. Tharops Laporte, 1835: 168; Guérin-Méneville, 1843: 171; LeConte, 1852: 45; Lacordaire, 1857: 99; Kiesen- wetter, 1858: 193; Jacquelin du Val, 1863: 114; Bonvouloir, 1871: 64; Horn, 1886: 6; Fleutiaux, 1902a: 659. Type species: Isorhipis lepaigei Lacordaire, 1835: 623.

34 EUCNEMIDAE: Isorhipis

Diagnosis: Body elongate, cylindrical; frontoclypeal region mostly trapezoidal, broadly rounded at anterior edge; antennae flabellate in males, serrate in females; elytra indistinctly striate; hypomeron mostly parallel- sided, antennal grooves absent; metepisternum gradually widened posterad; metacoxal plate strongly expanded inward; legs long and slender; abdominal ventrite 5 narrowly rounded or pointed at apex.

Distribution: Holarctic region, Neotropical region, Oriental region.

Remarks: This genus consists of 21 described species worldwide, mostly distributed widely in Holarctic region. Among them, seven species are present in the Palaearctic region (Muona, 2007, 2011). Several larvae and adults Isorhipis species were observed in standing dead, barkless trunk of Abies, Acer, Betula, Carpinus, Cyrilla, Fagus, Quercus, Tilia, Ulmus (Muona, 1993).

16. Isorhipis foveata Hisamatsu, 1955 (Figs. 15; Pls. VIII, XII, XIII)

Isorhipis foveata: Hisamatsu, 1955: 96; 1985: 43; Muona, 2007: 84; Suzuki, 2012: 47.

Adult: Male. Body length 3.4-5.6 mm. Mostly black with yellow-brown antennae, mouthparts, basal half of elytra, and legs; surface covered with yellow-gray pubescence. Head moderately inserted into prothorax; frons slightly depressed; frontoclypeal region trapezoidal, weakly trilobate at anterior edge, anterior edge about 1.85 times wider than distance between antennal sockets. Antennae flabellate from antennomere 4; antennomere 3 gradually widened apically, 1.4 times longer than 2; antennomeres 4-10 subequal, branched near apex. Pronotum parallel-sided, about 1.15 times wider than long; disc with paired weak depression near middle. Elytra parallel-sided, gradually narrowed near apices; yellow-tinged at basal half; ratio of length to combined width about 3.05; apices simply rounded. Prosternum parallel-sided. Hypomeron simple, without antennal groove. Metepisternum narrow, subparallel-sided. Metacoxal plate strongly expanded inward, medially about 4.6 times wider than laterally. Legs long and slender; metatarsomere 1 about 1.1 times longer than 2-4 combined; metatarsomere 2 about 1.6 times longer than 3; metatarsomere 5 about 1.4 times longer than 2. Abdomen strongly connate; last visible ventrite weakly pointed at apex. Aedeagus bulbous, about 3.2 times longer than wide; median lobe elongate, strongly bent ventrally near apex; lateral lobes slender, with apical tooth. Female. Can be distinguished from males by following characters: Body length 6.7-7.9 mm. Antennae serrate; antennomere 3 about twice longer than wide, about 2.1 times longer than 2, about 1.65 times longer than 4; antennomeres 4-10 subequal; apical antennomere about 1.6 times longer than wide, about 1.7 times longer than 10.

Distribution: Korea, Japan.

Distribution in Korea: GG, GW, SO (Suzuki, 2012).

35 INSECT FAUNA OF KOREA Eucnemidae

Material examined: GG: 1♀(Jikdong, Sohol, Pocheon, 21.v.1992, G.J. Weon); 1♀(Jikdong, Sohol,

Pocheon, 29.v.1992, G.J. Weon); 1♀(Jikdong, Sohol, Pocheon, 21.v.1995, G.J. Weon); 1♂, 1♀(Deoksu,

Danwol, Yangpyeong, 8-22.v.2016, on flight intercept trap, Seung and Jung). GW: 1♀(Cheonmi, Bangsan,

Yanggu, 16.v.2015, S.H. Lee). SO: 1♂(Cheongnyangri, Dongdaemun, 3.v.2016, J.K. Jung). Remarks: Isorhipis foveata is distinguished from other Isorhipis species by weakly serrated antennae of female, yellow-tinged elytra, and aedeagal structure. A female individual was observed wandering on fallen timbers in May. Individual variations were observed in coloration of elytra, completely tinged with yellow wholly or at basal half. Also, lateral lobes of male genitalia were more or less extended laterally.

Tribe Xylobiini Reitter, 1911

Xylobiini Reitter, 1911: 203. Type genus: Xylobius Latreille, 1834: 124.

Diagnosis: Antennomere 3 mostly small; hypomeron mostly without antennal groove; protarsomere 1 simple, without sex-comb in males; median lobe of aedeagus fused with lateral lobes.

Remarks: Xylobiini consist of 53 species in eight genera worldwide and nineteen species in three genera are present in the Palaearctic region (Muona, 2007, 2011; Watanabe, 2009; Otto, 2016).

Key to genera of Xylobiini in Korea

1. Antennomeres 4-11 with medio-longitudinal keel; hypomeral antennal grooves more or less developed...‌ ...... Proxylobius - Antennomeres 4-11 without medio-longitudinal keel; hypomeral antennal grooves absent...... 2 2. Body‌ mostly with dull surface; antennae filiform or serrate; pronotum and scutellum densely and roughly ‌ punctate with pubescence...... Bioxylus - Body with strongly glossy surface; antennae moniliform; pronotum sparsely and regularly punctate; scutellum non-punctate, glabrous...... Xylophilus

Genus Bioxylus Fleutiaux, 1923

Bioxylus Fleutiaux, 1923: 320; 1935: 4; Hisamatsu, 1959: 23; 1985: 42; Muona, 1987: 81; Otto, 2016: 198; Seung and Lee, 2018a: 116. Type species: Xylobius japonensis Fleutiaux, 1900: 359.

36 EUCNEMIDAE: Bioxylus

Diagnosis: Body oblong, subcylindrical; antennae filiform or serrate, antennomere 2 shorter or as long as 3, antennomere 4 as long as 2-3 combined; elytra indistinctly striate; hypomeron simple, without antennal groove; metepisternum mostly subparallel-sided; metacoxal plate subparallel-sided; abdominal ventrite 5 simply rounded or pointed at apex.

Distribution: Afrotropical region, Oriental region, Palaearctic region.

Remarks: This genus consists of 13 described species worldwide, among them, nine species are present in the Palaearctic region (Muona, 2007, 2011; Watanabe, 2009; Otto, 2016). Most species were recorded in Japanese fauna. Recently, three species were recognized in Korean Peninsula (Seung and Lee, 2018a). Bioxylus personatus was observed on fallen trunk of Ulmus (Mamaev, 1976).

Key to species of Bioxylus in Korea

1. Metepisternum triangular, strongly widened posterad; abdominal ventrite 5 distinctly produced apically...‌ ...... B. galloisi - Metepisternum with subparallel sides; abdominal ventrite 5 rounded apically...... 2 2. Body‌ covered with moderately dense pubescences; elytra bicolor, more or less ferruginous at base;‌ abdominal ventrite 5 strongly serrate at apex in males...... B. natsumiae - Body‌ covered with strongly dense pubescences; elytra entirely unicolor, chestnut to black; abdominal ventrite 5 simply rounded at apex in both sexes...... B. pilosellus

17. Bioxylus natsumiae Watanabe, 2009 (Figs. 16; Pls. VIII, XII, XIII)

Bioxylus natsumiae: Watanabe, 2009: 135; Seung and Lee, 2018a: 116.

Adult: Male. Body length 3.3-4.3 mm. Mostly black with red-brown mouthparts, antennae, pronotal margin, elytral base, femur and orange-brown tibiae, tarsi; surface covered with yellow pubescence. Head deeply inserted into prothorax; frons with medio-longitudinal keel from vertex to anterior edge of frontoclypeal region; frontoclypeal region weakly bifurcate at anterior edge, anterior edge about 2.35 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 shortest, closely connected to 4; antennomere 4 about 1.2 times longer than wide, about 1.2 times longer than 2-3 combined; antennomeres 5-10 gradually lengthened and slenderized; apical segment about five times longer than wide, about 1.6 times longer than 10. Pronotum strongly convex, about 1.2 times wider than long. Elytra parallel-sided, gradually narrowed near apices; red-tinged at base; ratio of length to combined width about 2.1; apices simply rounded. Prosternum transverse, gradually widened anterad. Hypomeron simple, without antennal groove. Metepisternum subparallel-sided. Metacoxal plate expanded inward, medially about 1.5 times wider

37 INSECT FAUNA OF KOREA Eucnemidae

than laterally. Legs slender; metatarsomere 1 about 1.3 times longer than 2-4 combined; metatarsomere 2 about 1.15 times longer than 3; metatarsomere 5 about 1.5 times longer than 2. Abdomen strongly connate; last visible ventrite with distinctly serrate at apex. Aedeagus about 3.2 times longer than wide; median lobe elongate, gradually narrowed apically; lateral lobes slender, slightly longer than median lobe, with basally attached secondary lateral lobes; secondary lateral lobes stout, as long as lateral lobes, with short seta. Female. Can be distinguished from males by following characters: Body length 3.4-4.7 mm. Fronto- clypeal region wider, anterior edge about 2.15 times wider than distance between antennal sockets. Apical antennomere about 3.7 times longer than wide. Abdominal ventrite 5 not serrate, simply rounded at apex.

Distribution: Korea, Japan.

Distribution in Korea: GG, GW(Seung and Lee, 2018a).

Material examined: GG: 3♂ (Deoksu, Danwol, Yangpyeong, 16-26.vii.2015, on flight intercept trap,

Seung and Jung); 2♂(Deoksu, Danwol, Yangpyeong, 26.vii-3.viii.2015, on flight intercept trap, Seung and

Jung); 1♂(Deoksu, Danwol, Yangpyeong, 29.vi-16.vii.2016, on flight intercept trap, Seung and Jung); 5♀

(Jikdong, Sohol, Pocheon, 9.vii.2007, T.S. Kwon); 3♀(Jikdong, Sohol, Pocheon, 26.vii.2007, T.S. Kwon).

GW: 2♂ (Yeongheung, Yeongwol, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee);

1♂(Yeongheung, Yeongwol, Yeongwol, 2-16.vii.2015, on flight intercept trap, Seung and Lee); 1♂(Yeo- ngheung, Yeongwol, Yeongwol, 16-30.vii.2015, on flight intercept trap, Seung and Lee); 2♂(Beopheung,

Suju, Yeongwol, 16-30.vii.2015, on flight intercept trap, Seung and Lee); 1♂(Suha, Daegwanryeong, Py- eongchang, 5-29.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Bioxylus natsumiae is easily distinguished from other Bioxylus species by long antennae, red- tinged elytral base, distinctly serrated apex of ventrite 5 in males.

18. Bioxylus galloisi Fleutiaux, 1923 (Figs. 17; Pls. IX, XII)

Bioxylus galloisi: Fleutiaux, 1923: 320; 1935: 11; Hisamatsu, 1958b: 119; 1959: 23; 1985: 44; Muona, 2007: 85; Seung and Lee, 2018a: 117.

Adult: Female. Body length 4.8-5.1 mm. Mostly black with dark brown antennae, pronotal margin, elytra, tibiae, and tarsi; surface covered with short yellow-brown pubescence. Head deeply inserted into prothorax; frons with medio-longitudinal keel from vertex to base of frontoclypeal region; frontoclypeal region weakly bifurcate at anterior edge, anterior edge about 2.6 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 3 shortest, closely connected to 4; antennomere 4 about 1.1 times longer than wide, as long as 2-3 combined, 1.1 times longer than 5; antennomeres 5-10 subequal; apical antennomere about 1.7 times longer than wide, about 1.8 times longer than 10. Pronotum as long as wide,

38 EUCNEMIDAE: Bioxylus

parallel-sided. Elytra parallel-sided, gradually narrowed near apices; ratio of length to combined width about 2.2; apices simply rounded. Prosternum subparallel-sided, slightly widened anterad. Hypomeron simple, without antennal groove. Metepisternum triangular, distinctly widened posterad. Metacoxal plate slightly expanded inward, medially about 1.1 times wider than laterally. Legs slender; metatarsomere 1 about 1.15 times longer than 2-4 combined; metatarsomere 2 about 1.5 times longer than 3; metatarsomere 5 about 1.2 times longer than 2. Abdomen strongly connate; last visible ventrite distinctly beaked at apex.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GW, JJ (Seung and Lee, 2018a).

Material examined: GW: 1♀(Wangsan, Wangsan, Gangneung, 3-29.vi.2016, on flight intercept trap,

Seung and Jung). JJ: 1♀(Gyorae, Jocheon, Jeju, 12.v-10.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Bioxylus galloisi is similar to Bioxylus japonensis (Fleutiaux, 1900) and Bioxylus personatus Mamaev, 1976, but can be distinguished by posteriorly expanded metepisternum and apically beaked abdominal ventrite 5. Individual variations were observed with elytra color, more or less brown, occasionally tinged with black.

19. Bioxylus pilosellus Hisamatsu, 1959 (Figs. 18; Pls. IX, XII, XIII)

Bioxylus pilosellus: Hisamatsu, 1959: 23; 1985: 44; Muona, 2007: 85; Seung and Lee, 2018a: 118.

Adult: Male. Body length 4.7-5.1 mm. Mostly black to dark brown with red-brown tibiae, tarsi; surface very densely covered with yellow pubescence. Head deeply inserted into prothorax; frons with medio-longitudinal keel from vertex to base of frontoclypeal region; frontoclypeal region weakly bifurcate at anterior edge, anterior edge about 2.3 times wider than distance between antennal sockets. Antennae serrate; antennomere 3 shortest, closely connected to 4; antennomere 4 about 1.7 times longer than wide, about 1.3 times longer than 2-3 combined; antennomeres 5-10 slightly slenderized toward apex; apical segment about 3.2 times longer than wide, about 1.7 times longer than 10. Pronotum as long as wide. Elytra subparallel-sided, gradually narrowed posterad; ratio of length to combined width about 2.3; apices simply rounded. Proster- num transverse, slightly widened anterad. Hypomeron simple, without antennal groove. Metepisternum subparallel-sided. Metacoxal plate expanded inward, medially about 1.5 times wider than laterally. Legs slender; metatarsomere 1 about nine-tenths as long as 2-4 combined; metatarsomere 2 about 1.25 times longer than 3; metatarsomere 5 about 1.5 times longer than 2. Abdomen strongly connate; last visible ventrite simply rounded at apex. Aedeagus about 3.55 times longer than wide; median lobe stout, gradually widened from base and abruptly narrowed apically from basal two-thirds; lateral lobes longer than median lobe, with basally attached secondary lateral lobes; secondary lateral lobes as long as lateral lobes, with several long

39 INSECT FAUNA OF KOREA Eucnemidae

hairs from basal two-thirds. Female. Very similar to males, but can be distinguished by following characters: Body length 4.9-6.1 mm. Frontoclypeal region wider, anterior edge about 1.9 times wider than distance between antennal sockets. Antennae relatively shorter; antennomeres 5-10 stouter.

Distribution: Korea, Japan.

Distribution in Korea: GG, GW, JN, JJ (Seung and Lee, 2018a).

Material examined: GG: 2♀(Jikdong, Sohol, Pocheon, 9.vii.2007, T.S. Kwon); 3♀(Jikdong, Sohol,

Pocheon, 26.vii.2007, T.S. Kwon). GW: 2♂(Suha, Daegwanryeong, Pyeongchang, 5-29.vi.2016, on flight intercept trap, Seung and Jung). JN: 1♂(Jungdae, Ganjeon, Gurye, 4-15.vii.2016, on flight intercept trap,

Seung and Lee); 1♀(Pyeonghwal, Samsan, Haenam, 22.viii.2009, T.S. Kwon). JJ: 1♂(Daepo, Seogwipo, 22.vii.2016, J.B. Seung).

Remarks: Bioxylus pilosellus can be easily distinguished from other Bioxylus species by its highly dense pubescence, especially on dorsal surface. A male individual was observed wandering on dead barkless tree with mycelium in July. Individual variation was observed in body coloration, dark brown to dull black.

Genus Proxylobius Fleutiaux, 1900

Proxylobius Fleutiaux, 1900: 360; 1902a: 650; 1921: 144; 1923: 304; Hisamatsu, 1958a: 117; 1985: 42; Otto, 2016: 203; Seung and Lee, 2018a: 119. Type species: Proxylobius helleri Fleutiaux, 1900: 361.

Diagnosis: Body oblong, cylindrical; antennae serrate, antennomeres 4-11 with medio-longitudinal keel; elytra indistinctly striate; hypomeral antennal grooves well-defined or indistinct; metepisternum slightly widened posterad; metacoxal plate subparallel-sided; abdominal ventrite 5 simply rounded at apex.

Distribution: Afrotropical region, Palaearctic region, Oriental region.

Remarks: This genus consists of six described species worldwide, among them, three species are present in the Palaearctic region (Muona, 2007, 2011; Otto, 2016). Recently, two species were recognized from Korea (Seung and Lee, 2018a). An Oriental species, Proxylobius gardneri (Fleutiaux, 1930) was observed in rotten wood of Machilus odoratissima Nees in immature stage (Gardner, 1930).

Key to species of Proxylobius in Korea

1. Antennae‌ moderate in length, almost reaching metacoxal plate; hypomeral antennal grooves well-developed, with outer marginal keel; secondary lateral lobes of aedeagus elongate and slender...... P. helleri

40 EUCNEMIDAE: Proxylobius

- Antennae‌ long, obviously exceeding abdominal ventrite 3; hypomeral antennal grooves indistinct; secondary lateral lobes of aedeagus short and stout...... P. longicornis

20. Proxylobius helleri Fleutiaux, 1900 (Figs. 19; Pls. X, XII, XIII)

Proxylobius helleri: Fleutiaux, 1900: 361; 1921: 144; 1923: 304; 1935: 12; Hisamatsu 1958a: 117; 1985: 46; Muona, 2007: 85; Seung and Lee, 2018a: 119.

Adult: Male. Body length 2.6-4.3 mm. Mostly black with ferruginous mouthparts, antennomeres 2-3, elytral humeri and yellow-brown tibiae, tarsi; surface covered with yellow-brown pubescence. Head deeply inserted into prothorax; frons with medio-longitudinal keel from vertex to anterior edge of frontoclypeal region; frontoclypeal region weakly bifurcate at anterior edge, anterior edge about 2.7 times wider than distance between antennal sockets. Antennae serrate; antennomere 2 about 1.5 times longer than 3; antennomere 3 shortest, closely connected to 4; antennomere 4 about 1.5 times longer than wide, about 1.3 times longer than 2-3 combined; antennomeres 5-8 gradually lengthened and slenderized; antennomeres 9 and 10 subequal, slightly shorter than 8; apical antennomere about 3.4 times longer than wide, about 1.5 times longer than 10. Pronotum convex, about 1.15 times wider than long. Elytra subparallel-sided, gradually narrowed posterad; red-tinged at humeri; ratio of length to combined width about 2.1; apices simply rounded. Prosternum transverse, parallel-sided. Hypomeron with well-defined wide antennal groove. Metepisternum slightly widened posterad. Metacoxal plate expanded inward, medially about 1.35 times wider than laterally. Legs slender; metatarsomere 1 about 1.2 times longer than 2-4 combined; metatarsomere 2 about 1.1 times longer than 3; metatarsomere 5 about 1.5 times longer than 2. Abdomen strongly connate; last visible ventrite simply rounded at apex. Aedeagus about 3.2 times longer than wide; median lobe gradually narrowed apically, slightly bent ventrally; lateral lobes slightly bent ventrally as median lobe, with dull apical tooth, with basally attached secondary lateral lobes; secondary lateral lobes slender, slightly shorter than median lobe, rounded at apex. Female. Very similar to males, but can be distinguished by following characters: Body length 4.8-5.9 mm. Frontoclypeal region wider, anterior edge about 2.15 times wider than distance between antennal sockets. An- tennae relatively shorter; antennomeres 5-10 stouter; apical antennomere about 2.9 times longer than wide.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW, JJ (Seung and Lee, 2018a).

Material examined: GG: 2♀(Jikdong, Sohol, Pocheon, 29.vi.2007, T.S. Kwon); 8♀(Jikdong, Sohol,

Pocheon, 9.vii.2007, T.S. Kwon); 3♀(Jikdong, Sohol, Pocheon, 26.vii.2007, T.S. Kwon). GW: 2♂(Deok- gu, Sangdong, Yeongwol, 19.vi-2.vii.2015, on flight intercept trap, Seung and Lee); 3♂ (Deokgu, Sang-

41 INSECT FAUNA OF KOREA Eucnemidae

dong, Yeongwol, 3-16.vii.2015, on flight intercept trap, Seung and Lee); 2♂(Beopheung, Suju, Yeongwol,

3-16.vii.2015, on flight intercept trap, Seung and Lee); 2♂(Beopheung, Suju, Yeongwol, 16-30.vii.2015, on flight intercept trap, Seung and Lee). JJ: 1♀(Gyorae, Jocheon, Jeju, 12.v-10.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Individual variation was observed in coloration of elytral humeri, ferruginous to blackish brown.

21. Proxylobius longicornis Hisamatsu, 1958 (Figs. 20; Pls. X, XII, XIII)

Proxylobius longicornis: Hisamatsu, 1958a: 117; 1985: 46; Muona, 2007: 85; Seung and Lee 2018a: 120.

Adult: Male. Body length 4.5 mm. Mostly black with ferruginous mouthparts, antennomeres 2-3, elytral humeri and yellow-brown tibiae, tarsi; surface covered with yellow-brown pubescence. Head deeply inserted into prothorax; frons with medio-longitudinal keel from vertex to anterior edge of frontoclypeal region; frontoclypeal region distinctly bifurcate at anterior edge, anterior edge about 2.7 times wider than distance between antennal sockets. Antennae weakly serrate; antennomere 2 about 1.6 times longer than 3; antennomere 3 shortest, closely connected to 4, about two-thirds of 2; antennomere 4 about 1.8 times longer than wide, about twice longer than 2-3 combined; antennomeres 5-8 gradually lengthened and slenderized; antennomeres 9 and 10 subequal, slightly shorter than 8; apical antennomere about six times longer than wide, about 1.7 times longer than 10. Pronotum convex, about 1.15 times wider than long. Elytra subparallel- sided, gradually narrowed posterad; red-tinged at humeri; ratio of length to combined width about 2.2; apices simply rounded. Prosternum transverse, subparallel-sided, slightly widened anterad. Hypomeron with indistinct antennal groove. Metepisternum gradually widened posterad. Metacoxal plate with parallel sides. Legs slender; metatarsomere 1 about 1.45 times longer than 2-4 combined; metatarsomere 2 about 1.3 times longer than 3; metatarsomere 5 about 1.7 times longer than 2. Abdomen strongly connate; last visible ventrite simply rounded at apex. Aedeagus about 3.45 times longer than wide; median lobe stout, gradually narrowed apically, slightly bent ventrally; lateral lobes slightly bent ventrally as median lobe, with weakly- developed apical tooth, with basally attached secondary lateral lobes; secondary lateral lobes stout, distinctly shorter than median lobe, rounded at apex.

Distribution: Korea, Japan.

Distribution in Korea: JN (Seung and Lee, 2018a).

Material examined: JN: 1♂ (Jungdae, Ganjeon, Gurye, 4-15.vii.2016, on flight intercept trap, Seung and Lee).

42 EUCNEMIDAE: Xylophilus

Remarks: Proxylobius longicornis is very similar to P. helleri, but can be distinguished by its long antennae, indistinct hypomeral antennal grooves, short and stout secondary lateral lobes of aedeagus.

Genus Xylophilus Mannerheim, 1823

Xylophilus Mannerheim, 1823: 14; Eschscholtz, 1829: 35; Guérin-Méneville, 1843: 165; Méquignon, 1922: 370; Fleutiaux, 1935: 10; Muona, 2000: 30; Otto, 2016: 194; Seung and Lee, 2018a: 121. Xylobius Latreille, 1834: 124; Guérin-Méneville, 1843: 171; Bonvouloir, 1875: 761; Horn, 1886: 11; Fleuti- aux, 1902a: 663; Hisamatsu, 1959: 29; 1985: 42. Xyloecus Dejean, 1833: 85. Type species: Elater alni Fabricius, 1801: 246.

Diagnosis: Body oblong, subcylindrical, with strongly glossy surface; frontoclypeal region feebly trilobate at anterior edge; antennae moniliform, antennomere 2 subequal or slightly shorter than 3; elytra indistinctly striate; hypomeral antennal grooves absent; metepisternum subparallel-sided; metacoxal plate subparallel-sided; abdominal ventrite 5 simply rounded at apex.

Distribution: Afrotropical region, Holarctic region, Neotropical region, Oriental region.

Remarks: This genus consists of 17 described species worldwide, widely distributed in the Afrotropical, Holarctic, Neotropical, and Oriental region. Among them, eight species are present in the Palaearctic region (Muona, 2007, 2011; Otto, 2016). Although their biology for these beetles is mostly veiled yet, their larvae are usually known as developing in standing or fallen dead wood with white-rotten surfaces (Gardner, 1935; Muona, 1993). Also, some study indicated that Xylophilus corticalis (Paykull) may lack a functional gut (Dodelin et al., 2005).

22. Xylophilus ainu (Fleutiaux, 1923) (Figs. 21; Pls. XI, XII, XIII)

Xylophilus ainu: Fleutiaux, 1935: 11; Muona, 2007: 85; Seung and Lee, 2018a: 122. Xylobius ainu: Fleutiaux, 1923: 322; Hisamatsu, 1985: 44; ESK/KSAE: 161.

Adult: Male. Body length 3.5-4.7 mm. Mostly shiny black with ferruginous antennae, elytral humeri, tibiae and yellow tarsi; surface rarely covered with short yellow pubescence. Head moderately inserted into prothorax; frons with a weak medio-longitudinal groove; frontoclypeal region weakly trilobate at anteri- or edge, anterior edge about 2.5 times wider than distance between antennal sockets. Antennae moniliform;

43 INSECT FAUNA OF KOREA Eucnemidae

antennomere 3 as long as wide, subequal to 2. Pronotum about 1.15 times longer than wide, abruptly narrowed anterad from basal two-thirds. Elytra parallel-sided, gradually narrowed near apices; ratio of length to combined width about 2.4, apices simply rounded. Prosternum transverse, subparallel-sided, slightly widened anterad. Hypomeron simple, without antennal groove. Metepisternum parallel-sided, same width with elytral epipleuron. Metacoxal plate expanded inward, medially about 1.5 times wider than laterally. Legs slender; metatarsomere 1 as long as 2-4 combined; metatarsomere 2 about 1.1 times longer than 3; metatarsomere 5 about twice longer than 2. Abdomen strongly connate; last ventrite narrowly rounded at apex. Aedeagus elongate, about 3.6 times as long as wide; median lobe gradually narrowed toward apex, strongly bent ventrally in lateral view; lateral lobes longer than median lobe, with subapical tooth. Female. Very similar to males, but can be distinguished by following characters: Body larger and stouter, body length 4.8-5.9 mm. Antennae relatively shorter, antennomere 2 about 1.2 times longer than 3, antennomeres 4-10 stouter and transverse.

Distribution: Korea, Japan, Russia (Far East).

Distribution in Korea: GG, GW, JB, JJ (Seung and Lee, 2018a).

Material examined: GG: 2♀(Jikdong, Sohol, Pocheon, 29.vi.2007, T.S. Kwon). GW: 1♂, 1♀(Osaek,

Seo, Yangyang, 20-22.vi.2015, S.H. Lee); 1♀(Beopheung, Suju, Yeongwol, 2-16.vii.2015, on flight intercept trap, Seung and Lee). JB: 1♂(Bongdeok, Bokheung, Sunchang, 27.v-3.vi.2016, on flight intercept trap, Seung and Jung). JJ: 1♂, 2♀(Mt. Hallasan, Gyorae, Jocheon, Jeju, 10.vi.2016, J.B. Seung); 3♂, 2♀

(Gyorae, Jocheon, Jeju, 10.vi.2016, J.B. Seung; 4♂, 3♀(Daepo, Seogwipo, 11.vi.2016, J.B. Seung; 1♂

(Gyorae, Jocheon, Jeju, 13.v-10.vi.2016, on flight intercept trap, Seung and Jung); 1♀(Hwasun, Andeok, Seogwipo, 13.v-10.vi.2016, on flight intercept trap, Seung and Jung).

Remarks: Xylophilus ainu is very similar to the Japanese species, Xylophilus rufomarginatus, Fleutiaux, 1923, but can be distinguished by following characters: antennomere 4 as long as 3, while twice longer in X. rufomarginatus; metepisternum as wide as elytral epipleuron, while wider in X. rufomarginatus. Several individuals were observed wandering and mating on surface of dead barkless trees.

44 LITERATURES CITED

LITERATURES CITED

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도판 PLATES

PLATES

51 109 INSECT FAUNA OF KOREA Eucnemidae

PLATES

1. Feaia nipparensis (Hisamatsu, 1957) (a-d, female) 2. Euryptychus vicinus Fleutiaux, 1923 (a-d, female) 3. Fornax consobrinus Hisamatsu, 1963 (a, c-e, g, male; b, f, female) 4. Otho sphondyloides (Germar, 1818) (a-d, female) 5. Dirrhagofarsus lewisi (Fleutiaux, 1900) (a-d, female) 6. Dirrhagofarsus modestus (Fleutiaux, 1923) (a, c-e, g, male; b, f, female) 7. Dirrhagofarsus unicolor (Hisamatsu, 1960) (a, c-e, g, male; b, f, female) 8. Farsus ainu Fleutiaux, 1923 (a-e, male) 9. Microrhagus foveolatus (Fleutiaux, 1923) (a, c-e, g, male; b, f, female) 10. Microrhagus jejuensis Seung and Lee, 2018 (a, c-e, g, male; b, f, female) 11. Microrhagus mystagogus (Fleutiaux, 1923) (a, c-e, g, male; b, f, female) 12. Microrhagus ramosus Fleutiaux, 1902 (a, c-e, g, male; b, f, female) 13. Hylis harmandi (Fleutiaux, 1923) (a, c-e, g, male; b, f, female) 14. Hylochares harmandi Fleutiaux, 1900 (a, c-e, g, male; b, f, female) 15. Isorhipis foveata Hisamatsu, 1955 (a, c-e, g, male; b, f, female) 16. Bioxylus natsumiae Watanabe, 2009 (a, c-e, g, male; b, f, female) 17. Bioxylus galloisi Fleutiaux, 1923 (a-d, female) 18. Bioxylus pilosellus Hisamatsu, 1959 (a, c-e, g, male; b, f, female) 19. Proxylobius helleri Fleutiaux. 1900 (a, c-e, g, male; b, f, female) 20. Proxylobius longicornis Hisamatsu, 1958 (a-e, male) 21. Xylophilus ainu (Fleutiaux, 1923) (a, c-e, g, male; b, f, female) Scale bar: 1 mm (Plates I-XII); 0.5 mm (Plate XIII).

52 PLATES

PLATE I

1a 1b 1c

2a 2b 2c

53 INSECT FAUNA OF KOREA Eucnemidae

PLATE II

3a 3b 3c 3d

4a 4b 4c

54 PLATES

PLATE III

5a 5b 5c

6a 6b 6c 6d

55 INSECT FAUNA OF KOREA Eucnemidae

PLATE IV

7a 7b 7c 7d

8a 8b 8c

56 PLATES

PLATE V

9a 9b 9c 9d

10a 10b 10c 10d

57 INSECT FAUNA OF KOREA Eucnemidae

PLATE VI

11a 11b 11c 11d

12a 12b 12c 12d

58 PLATES

PLATE VII

13a 13b 13c 13d

14a 14b 14c 14d

59 INSECT FAUNA OF KOREA Eucnemidae

PLATE VIII

15a 15b 15c 15d

16a 16b 16c 16d

60 PLATES

PLATE IX

17a 17b 17c

18a 18b 18c 18d

61 INSECT FAUNA OF KOREA Eucnemidae

PLATE X

19a 19b 19c 19d

20a 20b 20c

62 PLATES

PLATE XI

21a 21b 21c 21d

63 INSECT FAUNA OF KOREA Eucnemidae

PLATE XII

1d 2d 3e 3f 4d 5d 6e 6f 7e 7f 8d

9e 9f 10e 10f 11e 11f 12e 12f 13e 13f 14e 14f

15e 15f 16e 16f 17d 18e 18f 19e 19f 20d 21e 21f

64 PLATES

PLATE XIII

3g 6g 7g 8e

9g 10g 11g 12g

13g 14g 15g 16g

18g 19g 20e 21g

65 INSECT FAUNA OF KOREA Eucnemidae

INDEX TO SCIENTIFIC NAMES

B F

Bioxylys 36 Farsus 23 natsumiae 37 ainu 24 galloisi 38 Feaia 10 pilosellus 39 nipparensis 11 Fornax 15 consobrinus 16 C

Calyptocerini 17 H

Hylis 30 D harmandi 31 Hylocharini 32 Dirhagini 19 Hylochares 32 Dirrhagofarsus 20 harmandi 33 lewisi 20 modestus 21 unicolor 22 I Dromaeolus 14 marginatus 14 Isorhipis 34 foveata 35

E M Epiphanini 30 Eucnemidae 9 Macraulacinae 11 Eucneminae 10 Macraulacini 13 Euryptychini 12 Melasinae 16 Euryptychus 12 Melasini 34 vicinus 13 Mesogenini 10

66 INDEX TO SCIENTIFIC NAMES

Microrhagus 25 P foveolatus 26 jejuensis 27 Proxylobius 40 mystagogus 28 helleri 41 ramosus 29 longicornis 42

O X

Otho 17 Xylobiini 36 sphondyloides 18 Xylophilus 43 ainu 43

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