Japanese Journal of Systematic Entomology, 23 (1): 103–112. June 15, 2017.

A Revision of the Genus (Coleoptera: ) from Japan

Koichi SOGOH and Hiroyuki YOSHITOMI Entomological Laboratory, Faculty of Agriculture, Ehime University, Tarumi 3-5-7, Matsuyama, 790-8566 Japan

Abstract The Japanese species of the genus Lycoperdina is revised. Three previously known species are redescribed: Lycoperdina castaneipennis Gorham, 1874, Lycoperdina dux Gorham, 1873 and Lycoperdina mandarinea Gerstaecker, 1858, and one new spe- cies is described: Lycoperdina hiranoi sp. nov. from Honshu and Hokkaido. The distributional pattern of Japanese species is briefly discussed.

Introduction Systematics The genus Lycoperdina Latreille, 1807 belongs to the Genus Lycoperdina Latreille, 1807 subfamily Redtenbacher, 1844 (Tomaszewska, 2005), and contains 28 species: 18 from the Palearctic, 9 from Lycoperdina Latreille, 1807: 73 (Type species: Galleruca the Ethiopian, and 1 from the Nearctic Regions (Shockley et bovistae Fabricius, 1792); Tomaszewska, 2005: 47. Golgia Mulsant, 1846: 20 (Type species: Silpha succincta al., 2009). Linnaeus, 1767). From Japan, three species have been recorded. First report Lycoperdinodes Arrow, 1923: 485 [replacement name for of the genus from Japan was Gorham (1873), he described Lycoperdinella Arrow, 1920]. Lycoperdina dux from Hyogo, and recorded L. mandarinea Lycoperdinella Arrow, 1920: 23 [homonym Lycoperdinella from Nagasaki. Subsequently Gorham (1874) described L. Champion, 1913] (Type species: Lycoperdinella morosa castaneipennis from Hyogo. In addition Gorham (1887) and Arrow, 1920). Hirano (2011) recognized the presence of an undescribed Falsoylaia Pic, 1945: 10 (Type species: Falsoylaia species from Hokkaido and Honshu. obscuresuturalis Pic, 1945). In this paper, we review the Japanese species of the genus Lycoperdina. Diagnosis (modified Tomaszewska, 2005). Body (Fig. 1) This paper is based on the Sogoh’s undergraduate elongate to ovate, convex dorsally, glossy. Coloration of body dissertation for Faculty of Agriculture, Ehime University light to dark reddish-brown, having vague black marking on (February 2017). middle of pronotum and elytra in some species; legs black to reddish brown. Antennae (Fig. 3) shorter than a half of body Materials and Methods length; antennal club 2-segmented, loose. Pronotum (Fig. 4A–D) with stridulatory membrane in anterior margin; basal Materials. The specimens used in this paper were preserved sulcus distinct; lateral sulci deep and subparallel; anterior in Ehime University Museum, Matsuyama (EUMJ); Kyushu angles acute or narrowly rounded; posterior angles right University Museum, Fukuoka (KUMJ); Tochigi Prefectual angle or acute. Mesoventrite (Fig. 4E–H) with intercoxal Museum, Utsunomiya (TPMJ) and private collection of Mr. Y. process short, narrowly separated mesocoxae. Elytra elongate, Hirano (HI). convex, widest at basal 1/3 or 1/2, gently tapering apically, Methods. Dissected parts are cleared in 10% solution blunt at apex, densely and irregularly covered with fine of KOH, and placed in glycerine on slides. Measurements, punctures. Femur (Fig. 2A, C, E, G) widest at 1/3 from edge. general observations, and dissections were made under a Abdomen (Fig. 6A–H) with five freely articulated ventrites; microscope (Leica MZ95). Photographs were taken under a anterior margin of intercoxal process straight; ventrite 1 as Leica MZ95 using a microscopy camera system (Nikon DS- long as ventrites 2–4 combined; ventrites 2–4 subequal in Fi1-L2), and combined with automontage software Combine length. Male genital segment (= tergite and sternite 9; Fig. ZM (Alan Hadley, UK). The figures were prepared using 6I–L) with paired apophyses fused near base or mid length. Adobe Photoshop CS. After examination, dissected parts and Aedeagus (Fig. 5) moderately long, strongly sclerotized genitalia were placed in genitalia tube with glycerine. and stout; median lobe with short thumb-like apical branch; Technical terms refer to Tomaszewska (2005). tegmen placed at base of median lobe, ring-like shaped, fused Morphological abbreviations used in this study are as parameres. follows: AI: aedeagal index (length/basal width); EL: elytral length from anterior margin to elytral apex; EW: maximum elytral width; IOI: interocular index (interocular distance/ Key to the species of Lycoperdina from Japan compound eye width); PML: pronotum length in median line; 1. Elytra (Fig. 1C, F) with large black marking at center; legs PSL: pronotum length in lateral from right anterior angle to fully reddish brown. …...... ……...L. mandarinea posterior margin; PW: maximum width of pronotum; TL: total -. Elytra (Fig. 1A, B, D, E, G, H) uniformly reddish-brown or length (PML + EL). The average is given in parenthesis after infuscate at center; legs black or partially red…...... …..2 the range. 2. Elytra oval, strongly convex, widest at middle (Fig. 1A,

Ⓒ Japanese Society of Systematic Entomology 104 Sogoh, K. and H. Yoshitomi

Fig. 1. Habitus of Lycoperdina spp. from Japan. —Male (A, B, C, G, I) and female (D, E, F, H) and ventral habitus in male (I). — A, D, Lycoperdina castaneipennis Gorham, 1874; B, E, L. dux Gorham, 1873; C, F, L. mandarinea Gerstaecker, 1858; G–I, L. hiranoi sp. nov. (G, I, holotype; H, paratype). Scale bars = 3.0 mm.

D); EL/EW 1.29–1.42 (1.34). Distributed in Honshu to margin subparallel-sided; male fore tibia (Fig. 2D) Kyushu…...... L. castaneipennis slightly projecting in inner margin; ventrite 5 rounded -. Elytra oblong, rather flattened, widest at basal 1/3 (Fig. 1B, at apex in male, triangle and slightly pointed at apex in E, G, H); EL/EW 1.38–1.56 (1.46)……...... 3 female. Median lobe (Fig. 5D) slightly curved ventrally, 3. Pronotum (Fig. 4B) transverse, widest at base; lateral convex in ventral 1/3; apex expanded and emarginated at

June 15, 2017, JJSE 23 (1) Revision of the genus Lycoperdina from Japan 105

Fig. 2. Male fore legs of Lycoperdina spp. — Femur, trochanter and coxa (A, C, E, G) and tibia (B, D, F, H). — A, B, Lycoperdina castaneipennis Gorham, 1874; C, D, L. dux Gorham, 1873; E, F, L. mandarinea Gerstaecker, 1858; G, H, L. hiranoi sp. nov. Scale bar = 1.0 mm.

Fig. 3. Male left antennae of Lycoperdina spp. —A, Lycoperdina castaneipennis Gorham, 1874; B, L. dux Gorham, 1873; C, L. mandarinea Gerstaecker, 1858; D, L. hiranoi sp. nov. Scale bar = 1.0 mm.

middle; apical branch moderately long, curved ventrally. Lycoperdina castaneipennis Gorham, 1874, 151. Gorham, TL 4.75–6.48 (5.72) mm. Distributed in Hokkaido to 1887, 642 [note]; Strohecker, 1953, 72 [listed]; Kyushu……...... …L. dux Tomaszewska, 2007, 564 [catalogued]; Shockley et al., -. Pronotum (Fig. 4D) weakly transverse, widest at apical 2009, 54 [listed]; Hirano, 2011, 19 [noted]; Jung, 2014, 1/3; lateral margins slightly tapered in basal 1/3; male fore 39 [note]. tibia (Fig. 2H) with distinct tooth-like projection in inner Lycoperdina (Golgia) castaneipennis: Csiki, 1910, 35, margin; ventrite 5 moderately flattened at apex in male, [catalogued]; Ohta, 1931, 225 [note]; Strohecker, 1970, 245 [lectotype designation, male genitalia]; Sasaji, 1985, triangle and slightly pointed at apex in female. Median 241 [note, photo]. lobe (Fig. 5J) straight; apex uneven, expanded and slanted; apical branch very short, strongly curved ventrally. Specimens examined. [HONSHU] 2♂1♀ TL 4.35–5.69 (5.16) mm. Distributed in Hokkaido and (EUMJ), Yumoto, Nasu-onsen (alt. 850 m), 16. x. 1967, S. Honshu……...... ……...L. hiranoi Hisamatsu leg.; 6♂6♀ (EUMJ), Nasu (alt. 850 m), 7. x. 1948, S. Hisamatsu leg.; 2♂4♀ (TPMJ), Kuriyama-mura, Tashiro- Lycoperdina castaneipennis Gorham, 1874 rindô (alt. 1,000 m), 4. x. 1981, T. Miyamoto leg.; 2♂1♀ (Figs. 1A, D, 2A, B, 3A, 4A, E, 5A–C, 6A, B, I, 7B, E, F) (TPMJ), Himuro-yama, Kuzuu-machi (alt. 1,100 m), 1. iii. [Japanese name: Kuribane-tsuya-tentoudamashi] 1986, K. Kusano leg.; 1♂ (TPMJ), Kuroiso-shi (alt. 330 m), 25. x. 1983, M. Takaoka leg. 1♀ (KUMJ),

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Fig. 4. Pronota (A–D) and meso- and metasterna (E–H) of Lycoperdina spp. — A, E, Lycoperdina castaneipennis Gorham, 1874; B, F, L . dux Gorham, 1873; C, G, L. mandarinea Gerstaecker, 1858; D, H, L. hiranoi sp. nov. Scale bar = 1.0 mm.

Kuromori, Sudama (alt. 1,175 m), 23. vii. 1987, R. Inagawa A. Amagasu leg. 1♀ (EUMJ), Maya-san (alt. leg. 1♀ (KUMJ), near Matsutomi, Sudama (alt. 1,030 m), 700 m), 20. x. 1974, S. Kinoshita leg.; 2♂5♀ (EUMJ), Awaji- 21–23. vii. 1987, R. Inagawa leg.; 1♀ (KUMJ), Mikuni- shima, Yuzuruha-san (alt. 600 m), 5. x. 1972, M. Sakai leg. yama (alt. 1,300 m), 15. ix. 1968, Y. Hirano leg. 1♀ (EUMJ), Gagyusan, Takahashi-shi (alt. Pref.> 1♂ (EUMJ), Mikuni-tôge, Yamakita-chô (alt. 1,100 450 m), 14. x. 1978, S. Hisamatsu leg. [SHIKOKU] 3♂5♀ (EUMJ), Shiratsue-san, Matsuyama-shi (alt. Tanzawa (500 m), 3. xi. 1981, Y. Hirano leg. 1,150 m), 23. x. 1970, S. Hisamatsu leg.; 8♂15♀ (EUMJ), 1♂ (EUMJ), Komanoyu, Kisofukushima (alt. 1,100 m), 18. ditto, 20. x. 1968, M. Sakai leg.; 2♂1♀ (EUMJ), ditto, 9. xi. x. 1966, S. Hisamatsu leg. 1♀ (EUMJ), 1975, A. Oda leg.; 1♀ (EUMJ), ditto, 23. x. 1970, M. Sakai Kadoketa-yama, Misakubo-chô (alt. 1,350 m), 23. xi. 1994, H. leg.; 8♂16♀ (EUMJ), Mado-tôge (alt. 600 m), 27. xi. 1968, Yoshitomi leg. 2♂8♀ (EUMJ), Mennoki-tôge, M. Takagi & M. Sakai leg.; 8♂14♀ (EUMJ), Narabara-san Inabu-chô (alt. 1,100 m), 17. iv. 1994, H. Yoshitomi leg. 5♂7♀ (EUMJ), Kamihinachi, Nabari-shi (alt. 400 m), 22. x. 1978, M. Kotani leg.; 7♂5♀ (EUMJ), ditto, 3. xi. 1968, 9. x. 1996, K. Aita leg.; 4♂8♀ (KUMJ), Kami-Hinawa, 16. M. Sakai leg.; 1♀ (EUMJ), Komenono, Matsuyama-shi (alt. xi. 1984, A. Amagasu & H. Ichihashi leg.; 4♂10♀ (KUMJ), 800 m), 20. x. 1968, S. Kinoshita leg.; 1♂ (EUMJ), ditto, 23. Kiyohara, Ohmiya-chô, Watarai-gun (alt. 142 m), 11. xi. 1984, x. 1970, M. Tomokuni leg.; 1♀ (EUMJ), ditto, 20. x. 1995, S.

June 15, 2017, JJSE 23 (1) Revision of the genus Lycoperdina from Japan 107

Fig. 5. Male genitalia of Lycoperdina spp. in dorsal (A, D, G, J), lateral (B, E, H, K) and ventral (C, F, I, L) views. — A– C, Lycoperdina castaneipennis Gorham, 1874; D–F, L. dux Gorham, 1873; G–I, L. mandarinea Gerstaecker, 1858; J–L, L. hiranoi sp. nov. Scale bar = 1.0 mm.

Yoshimichi leg.; 1♀ (EUMJ), ditto, 23. x. 1996, H. Nakanishi 1♀ (EUMJ), Koboke, Yamashiro-machi (alt. 400 m), 18. leg.; 11♂18♀ (EUMJ), Ishizuchi-san (alt. 1,450 m), 18–20. xi. 2006, T. Kitano leg.; 1♀ (EUMJ), Kohtsu (alt. 1,000 m), x. 1975, A. Oda leg.; 1♀ (EUMJ), Saragamine (alt. 1,250 31. x. 1965, M. Sakai leg. [KYUSHU] 1♀ m), 23.x.1957, M. Miyatake leg.; 1♀ (EUMJ), ditto, 23. x. (KUMJ), Homan-zan (alt. 790 m), 31. v. 1957, H. Kamiya leg. 1957, S. Hisamatsu leg.; 2♂4♀(EUMJ), Jojusha, Ishizuchi- 1♂ (EUMJ), Yunono, Kirishima (alt. 800 san (alt. 1,450 m), 15. ix. 1988, M. Sakai leg.; 1♂ (EUMJ), m), 25. vii. 1968, S. Kinoshita leg. Jiyoshi-tôge, Yanadani-mura (alt. 500 m), 4. vi. 2001, J. Redescription. Male (Fig. 1A). Body oval, strongly Ogawa leg.; 4♂3♀ (EUMJ), Takanawa-san (alt. 900 m), 6. xi. convex, glossy. Coloration of body reddish-brown and 2004, Y. Sato leg. 1♂1♀ (EUMJ), Otaki-san becoming lighter or darker in variation, but having vague (alt. 850 m), 5.viii.1956, K. Ogawa leg. black marking on middle of pronotum and elytra in some

June 15, 2017, JJSE 23 (1) 108 Sogoh, K. and H. Yoshitomi

Fig. 6. Abdominal ventrites (A–H) and male genital segment (I–L) of Lycoperdina spp. — Male (A, C, E, G) and female (B, D, F, H). — A, B, I, Lycoperdina castaneipennis Gorham, 1874; C, D, J, L. dux Gorham, 1873; E, F, K, L. mandarinea Gerstaecker, 1858; G, H, L, L. hiranoi sp. nov. Scale bars = 1.0 mm. specimens; legs black or partially red. EL/PL 2.55–2.86 (2.69), and curved near base. PW/PML 1.27–1.63 (1.37), PW/ EW/PW 1.29–1.59 (1.47), TL/EW 1.74–1.93 (1.83). PSL 1.18–1.44 (1.26), PML/PSL 0.88–0.94 (0.92). Elytra Head densely and irregularly covered with small glossy, oval, strongly convex, widest at middle, densely and punctures; IOI 2.13. Antennae (Fig. 2A) shorter than half of irregularly covered with definite punctures; EL/EW 1.24–1.39 TL; antennomere 11 fan-shaped; approximate ratio of each (1.34). Intercoxal process of mesosternum moderately antennomere (n = 1) as 2.4 : 1.1 : 1.6 : 1.0 : 1.0 : 1.0 : 1.0 : short, wide and carinate. Metasternum distinctly transverse, 1.0 : 1.1 : 1.2 : 1.6. Pronotum (Fig. 4A) transverse, glossy; punctate, pubescent, 1.53 times as long as metacoxal cavity. definite punctures distributed densely and irregularly; anterior Fore trochanter (Fig. 2A) flattened, having a hairy tuft in male; and posterior angles acute; lateral margins rounded in front fore femur widest at basal 2/3; fore tibia (Fig. 2B) setose at

June 15, 2017, JJSE 23 (1) Revision of the genus Lycoperdina from Japan 109 apical 1/3, strongly rounded. Ventrite 5 (Fig. 6A) emarginate H. Yoshitomi leg. 2♂ (KUMJ), Kami-Hinawa, at apex. Male genital segment (Fig. 6I) formed by segment 9 Nabari-shi, 16. xi. 1984, A. Amagasu & H. Ichihashi leg.; 1♂ (tergite and sternite 9) with paired apophyses fused near base; (KUMJ), Mizusawa-dani, Yokkaichi (alt. 260 m), 23. xi. 1984, right apophyse with trapezoidal plate at base. A. Amagasu leg. [SHIKOKU] 1♂ (EUMJ), Aedeagus (Fig. 5A–C) strongly sclerotized and stout; AI Matsuyama (alt. 50 m), 4. vi. 1954, S. Hisamatsu leg.; 1♀ 2.25. Median lobe slightly curved ventrally, strongly convex (EUMJ), Narabara-san (alt. 1,050 m), 3. xi. 1968, M. Takagi in ventral 1/3; apex expanded and flattened; apical branch leg.; 1♀ (EUMJ), Shiratsue-san (alt. 1,150 m), 20. x. 1968, M. moderately long. Sakai leg.; 1♂2♀ (EUMJ), Sugitate, Matsuyama-shi (alt. 450 Female (Fig. 1D). Sexual dimorphism distinct in the m), 29. x. 2016, K. Sogoh leg.; 1♀ (EUMJ), ditto, 2. xi. 1967, following characteristics: fore legs slender and without K. Hatta et al. leg.; 1♂ (EUMJ), Takanawa-san, Matsuyama- projection; ventrite 5 (Fig. 6B) triangular, pointed at apex. PW/ shi (alt. 900 m), 24. ix. 2016, K. Kuroda leg. [KYUSHU] PML 1.36–1.50 (1.41), PW/PSL 1.22–1.37 (1.27), PML/PSL 1♂ (EUMJ), Tsushima Isl., Uchiyama 0.87–0.94 (0.90), EL/EW 1.24–1.39 (1.34), EL/PL 2.52–2.89 (alt. 150 m), 4. v. 1978, A. Oda leg. [TAIWAN] 1♀ (EUMJ), (2.73), EW/PW 1.34–1.60 (1.45), TL/EW 1.70–1.90 (1.83). Siangyang, Haiduan Township, Taitung County, 8. vi. 2013, J. Measurements. Male (n = 20). TL 5.01–6.43 (5.85) mm, PW Yamasako leg. 1.93–2.48 (2.18) mm, PML 1.41–1.78 (1.59) mm, PSL 1.50–2.00 Redescription. Male (Fig. 1B). Body oblong, glossy. (1.73) mm, EL 3.60–4.70 (4.26) mm, EW 2.75–3.50 (3.19) mm. Coloration of body reddish-brown and becoming lighter or Female (n = 20). TL 4.15–6.14 (5.61) mm, PW 1.68–2.30 (2.12) darker in variation, but having vague black marking on middle mm, PL 1.12–1.68 (1.50) mm, PL 1.23–1.83 (1.67) mm, EL of pronotum and elytra in some specimens; legs black or 3.03–4.49 (4.10) mm, EW 2.30–3.40 (3.07) mm. partially red. EL/PML 2.69–3.30 (3.06), EW/PW 1.37–1.57 Distribution. Japan (Hokkaido, Honshu, Sado-ga-shima, (1.48), TL/EW 1.85–1.96 (1.90). Awaji-shima, Shikoku, Kyushu), Korea, Russia. Head covered with small punctures densely and irregularly; Biological notes. This is common species in Japan, and IOI 1.62. Antennae (Fig. 3B) shorter than 1/2 TL; antennomere is collected from Lycoperdon gemmatum (Agaricaceae). This 11 fan-shaped; approximate ratio of each antennomere (n = species is mainly distributed in mountain zone (ca. 800–1,150 1) as 2.0 : 1.1 : 1.4 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.2 : 1.2 : 1.6. m in altitude). Taniguchi (1942) noted that this species was Pronotum (Fig. 4B) distinctly transverse, glossy, widest at collected from Calvatia craniformis (Agaricaceae) in Kobe, base, densely and irregularly covered with fine punctures; Japan. Hibernation is occurred in both larvae and adults stages. anterior and posterior angles acute; lateral margins subparallel, Remarks. The larva of this species (Fig. 7B, E, F) is rounded in front, weakly curved near base; PW/PML 1.31–1.54 similar to that of L. dux (Hayashi et al., 1959 and Fig. 7C, D), (1.45), PW/PSL 1.18–1.43 (1.37), PML/PSL 0.90–0.98 but differs from it by the tubercles on abdominal segments (0.95). Elytra glossy, oblong, widest at basal 1/3; densely relatively small and covered with short setae. and irregularly covered with definite punctures; EL/EW 1.38–1.51 (1.43). Intercoxal process of mesosternum (Fig. 4F) Lycoperdina dux Gorham, 1873 moderately long, narrow and carinate. Metasternum transverse, (Figs. 1B, E, 2C, D, 3B, 4B, F, 5D–F, 6C, D, J, 7 A, C, D) punctate, pubescent, 2.36 times as long as metacoxal cavity. [Japanese name: Fuchitori-tsuya-tentoudamashi] Fore trochanter (Fig. 2C) quadrate, having a hairy tuft; fore femur widest at basal 2/3, 2.00 times as wide as tibiae; Lycoperdina dux Gorham, 1873, 205. Gorham, 1887, 642 fore tibiae (Fig. 2D) setose in apical 1/3, with obscure [note]; Strohecker, 1953, 72 [listed]; Tomaszewska, projection in inner margin. Ventrite 5 (Fig. 6C) rounded at 2007, 564 [catalogued]; Shockley et al., 2009, 54 [listed]; apex. Male genital segment (Fig. 6J) formed by segment 9 Hirano, 2011, 19 [note]. with paired apophyses fused at mid length. Lycoperdina (Golgia) dux: Csiki, 1910, 35 [catalogued]; Aedeagus (Fig. 5D–F) strongly sclerotized and stout; AI Ohta, 1931, 225 [note]; Strohecker, 1970, 245 [lectotype 2.33. Median lobe slightly curved ventrally, convex in ventral designation, male genitalia]; Sasaji, 1985, 241 [note, 1/3; apex expanded and emarginate at middle; apical branch photo]. moderately long, slightly curved ventrally. Female (Fig. 1E). Sexual dimorphism distinct in the Specimens examined. [HOKKAIDO] 2♂5♀ (EUMJ), following characteristics: fore legs slender and without Eniwa-dake, lake side of Shikotsu-ko (alt. 1,200 m), 4. x. projection in tibiae; ventrite 5 (Fig. 6D) triangular and slightly 1968, S. Hisamatsu leg.; 1♂ (EUMJ), Kawayu (alt. 150 m), pointed at apex. PW/PML 1.45–1.58 (1.51), PW/PSL 1.33–1.45 21–22. vii. 1970, M. Sakai leg.; 2♀ (EUMJ), Sapporo-dake (1.39), PML/PSL 0.88–0.94 (0.92), EL/EW 1.40–1.51 (1.47), (alt. 1,200 m), 1. x. 1968, S. Hisamatsu leg. [HONSHU] EL/PML 3.19–3.47 (3.30), EW/PW 1.40–1.60 (1.49), TL/EW 1♂1♀ (EUMJ), Sanbongi (alt. 100 m), 10. vi. 1.84–1.98 (1.91). 1953, A. Fukuda leg.; 1♂1♀ (EUMJ), Tsuna-onsen (alt. 400 Measurements. Male (n = 14). TL 4.75–6.48 (5.58) mm, m), 23. ix. 1971, M. Tomokuni leg.; 1♂ (KUMJ), Zenkojitaira, PW 1.69–2.30 (1.98) mm, PML 1.15–1.75 (1.35) mm, PSL Nishi-gun, Hiraka-machi (alt. 360 m), 24. ix. 1978, A. Abe 1.18–1.95 (1.43) mm, EL 3.60–4.73 (4.23) mm, EW 2.58–3.43 leg. 1♂1♀ (EUMJ), Tsunagi-onsen (alt. 300 (2.91) mm. Female (n = 18). TL 4.96–6.34 (5.83) mm, PW m), 25. ix. 1971, M. Tomokuni leg. 2♂1♀ (EUMJ), 1.80–2.20 (2.04) mm, PML 1.18–1.49 (1.36) mm, PSL Toshima, Miyatsuka-yama, Izu Isls. (alt. 500 m), 4. xi. 2013, 1.28–1.55 (1.47) mm, EL 3.78–4.85 (4.47) mm, EW 2.55–3.35

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(3.03) mm, Fore trochanter (Fig. 2E) quadrate; fore femur widest at basal Distribution. Japan (Hokkaido, Honshu, Izu Isls. 2/3, 2.00 times as wide as tibiae; fore tibiae (Fig. 2F) setose (Toshima), Shikoku, Kyushu, Tsushima), Taiwan. at apical 1/3, with tooth in inner margin. Ventrite 5 (Fig. 6E) Biological notes. This is common species in Japan, and rounded at apex. Male genital segment (Fig. 6K) formed by is collected from Lycoperdon gemmatum (Agaricaceae). segment 9 with paired apophyses fused at mid length. This species is distributed in low to middle mountain zone Aedeagus (Fig. 5G–I) strongly sclerotized, stout; AI 2.67. (ca. 250–950 m in altitude). Taniguchi (1942) noted that this Median lobe straight; apex rounded; apical branch short, wide, species was collected from Calvatia craniformis (Agaricaceae) projecting posteriorly. in Kobe, Japan. Hashimoto & Hayashi (2014) reported Female (Fig. 1F). Sexual dimorphism distinct in the that this species was attracted by cantharidin-baited trap. following characteristics: fore legs slender, without tooth in Hayashi et al. (1959: 447) showed the illustrations and short inner margin; ventrite 5 (Fig. 6F) triangular, pointed at apex. description of the larva of this species. Hibernation is occurred PW/PML 1.40–1.504 (1.46), PW/PSL 1.30–1.37 (1.34), in both larvae and adults stages. PML/PSL 0.90–0.98 (0.92), EL/EW 1.43–1.65 (1.53), EL/PL Remarks. This is the first record of this species from Izu 2.94–3.12 (3.01), EW/PW 1.29–1.39 (1.35), TL/EW 1.91–2.18 Isls. (Toshima) and Tsushima. (2.04). Measurements. Male (n = 4). TL 4.01–5.18 (4.60) mm, Lycoperdina mandarinea Gerstaecker, 1858 PW 1.50–1.84 (1.67) mm, PML 1.10–1.33 (1.21) mm, PSL (Figs. 1C, F, 2E, F, 3C, 4C, G, 5G–I, 6E, F, K) 1.16–1.39 (1.26) mm, EL 2.91–3.85 (3.39) mm, EW 2.08–2.56 [Japanese name: Seguro-tsuya-tentoudamashi] (2.28) mm. Female (n = 9). TL 3.98–5.08 (4.56) mm, PW 1.48–1.88 (1.67) mm, PML 1.00–1.25 (1.13) mm, PSL Lycoperdina mandarinea Gerstaecker, 1858, 212. 1.10–1.38 (1.23) mm, EL 2.98–3.83 (3.43) mm, EW 2.05–2.50 Gorham, 1887, 642 [note]; Strohecker, 1953, 72 [list]; (2.24) mm. Tomaszewska, 2007, 564 [catalogued]; Shockley et al., Distribution. Japan (Hokkaido, Honshu, Izu Isls. (Hachijô- 2009, 54 [list]; Hirano, 2011, 19 [note]; Jung, 2014, 39 jima), Shikoku, Kyushu, Tokara Isls., Amami-Ôshima, [note]. Ishigaki-jima, Iriomote-jima), Korea, China, Hong Kong, Lycoperdina (Golgia) mandarinea: Csiki, 1910, 35, Taiwan, Cambodia, Laos, Mongolia, Russia, Vietnam. [catalogued]; Ohta, 1931, 225 [noted]; Strohecker, 1970, Biological notes. This species is mainly distributed in 245 [male genitalia]; Sasaji, 1985, 241 [note, photo]. low altitude area (ca 50–150 m in altitude) in Japan, and frequently collected by light trap. Taniguchi (1942) noted Specimens examined. [HONSHU] 3♀ that this species was collected from Lycoperdon gemmatum (EUMJ), Sakata-shi (alt. 100 m), 15. viii. 1952, K. Shirahata (Agaricaceae) in Kobe, Japan. leg. 1♂ (KUMJ), Nagoya (14 m), 7. vii. 1959, Remarks. Hayashi et al. (1959: 448) showed the K. Ohbayashi leg. [SHIKOKU] 2♀ (EUMJ), illustrations and short description of the larva of this species, Shigenobu-chô (alt. 100 m), 25. viii. 1981, M. Sakai leg. however, Hayashi (1978) corrected this larva was not 1♂ (EUMJ), Kawahara, Hikami-mura, Lycoperdina mandarinea but Pocadius sp. (Nitidulidae). Kida-gun (alt. 50 m), 2. ix. 1950, M. Chûjô leg. [RYUKYU] 1♂ (EUMJ), Ohtake (alt. 150 m), 23–29. iii. Lycoperdina hiranoi sp. nov. 1996, H. Yoshitomi leg.; 1♂ (EUMJ), Omoto-dake (alt. 400 (Figs. 1G–I, 2G, H, 3D, 4D, H, 5J–L, 6G, H, L) m), 9. iv. 1975, R. Yano leg. [Japanese name: Hosomune-tsuya-tentoudamashi] Redescription. Male (Fig. 1C). Body oblong, glossy. Coloration of body reddish-brown, becoming lighter or Lycoperdina sp.: Hirano, 2011, 19 [note]. darker in variation and distinct large black marking on elytra; legs fully reddish brown. EL/PL 2.65–2.89 (2.77), EW/PW Type series. Holotype (EUMJ): 1♂, Yumoto, Nasu- 1.31–1.39 (1.36), TL/EW 1.93–2.02 (1.99). onsen, Tochigi Pref. (alt. 850 m), 16. x. 1967, S. Hisamatsu Head densely and irregularly covered with small leg. Paratypes: [HOKKAIDO] 1♀ (EUMJ), Shikotsu-ko, punctures; IOI 1.57. Antennae (Fig. 3C) shorter than 1/2 near Chitose (alt. 650 m), 14. viii. 1979, Y. Seiyama leg.; TL; antennomere 11 fan-shaped; approximate ratio of each 1♀ (TPMJ), Tokachimitsumata, Kamishihoro-chô (alt. 650 antennomere (n = 1) as 3.0 : 1.3 : 1.8 : 1.2 : 1.0 : 1.0 : 1.0 : 1.2 m), 28. vii. 1986, T. Miyamoto leg. [HONSHU] 1♀ (EUMJ), Tsuna-onsen (alt. 400 m), 23. ix. 1971, and irregularly covered with definite punctures; anterior and M. Tomokuni leg.; 1♂, Shimokita-hantô, Yunomata, 17. viii. posterior angles right angled; lateral margins rounded in front, 1957, T. Saigusa leg. 2♂2♀ (EUMJ), Tsunagi- slightly curved near base; PW/PML 1.36–1.42 (1.39), PW/ onsen (alt. 300 m), 25. ix. 1971, M. Tomokuni leg.; 1♂ (HI), PSL 1.29–1.37 (1.33), PML/PSL 0.95–0.97 (0.96). Elytra Kujikeiryu (alt. 300 m), 2. viii. 1986, Y. Hirano leg. 1♂, Fujiwarako (alt. 700 m), 5. x. 1969, Y. Hirano leg. covered with definite punctures; EL/EW 1.40–1.50 (1.46). 3♂8♀ (EUMJ), Nasu (alt. 850 m), 1. x. 1948, Intercoxal process of mesosternum (Fig. 4G) moderately long, S. Hisamatsu leg.; 3♀ (TPMJ), Nishinasuno-chô, Northern wide and carinate, subparallel-sided. Metasternum transverse, part (alt. 1,000 m), 10. x. 1990, S. Ohmomo leg. 1♂2♀ (HI), Miurafuji (alt. 150 m), 10. x. 1988, Y.

June 15, 2017, JJSE 23 (1) Revision of the genus Lycoperdina from Japan 111

Fig. 7. Adult (A) and larvae (B–F) of Lycoperdina spp. A, C, D, Lycoperdina dux; B, E, F, L. castaneipennis. C– F, Fully expanded specimens of final instar larvae in dorsal (C, E) and lateral (D, F) aspects.

Hirano leg. 1♂ (HI), Ohmiya (alt. 50 m), iii. Aedeagus (Fig. 5J–L) strongly sclerotized and stout; 1971, Y. Hirano leg. 1♀ (KUMJ), Mizusawa- AI 1.94. Median lobe straight; apex uneven, expanded and dani, Yokkaichi (alt. 260 m), 23. iv. 1984, A. Amagasu leg.; slanted; apical branch very short, strongly curved ventrally. 1♂ (KUMJ), Kami-Hinawa, Nabari-shi, 16. xi. 1984, H. Female (Fig. 1H). Sexual dimorphism distinct in the Ichihashi leg. following characteristics: fore legs slender, without tooth Description. Male (Fig. 1G). Body oblong, glossy. in inner margin; ventrite 5 (Fig. 6H) triangular and slightly Coloration of body reddish-brown and becoming lighter or pointed at apex. PW/PML 1.37–1.58 (1.47), PW/PSL 1.28– darker in variation, but having vague black marking on middle 1.48 (1.35), PML/PSL 0.89–0.95 (0.92), EL/EW 1.39–1.56 of pronotum and elytra in some specimens; legs black or (1.47), EL/PML 2.84–3.28 (3.09), EW/PW 1.37–1.55 (1.43), partially red. EL/PML 3.04–3.26 (3.11), EW/PW 1.43–1.53 TL/EW 1.88–2.05 (1.95). (1.46), TL/EW 1.88–1.94 (1.91). Measurements. Male (n = 8). TL 5.00–5.65 (5.25) mm, Head densely and irregularly covered with small punctures; PW 1.76–2.00 (1.88) mm, PML 1.21–1.45 (1.29) mm, PSL IOI 1.35. Antennae (Fig. 3D) shorter than 1/2 TL; antennomere 1.30–1.55 (1.43) mm EL 3.75–4.20 (3.96) mm, EW 2.57–2.99 11 fan-shaped; approximate ratio of each antennomere (n (2.74) mm. Female (n =17). TL 4.35–5.69 (5.13) mm, PW = 1) as 2.0 : 1.0 : 1.4 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.0 : 1.58–2.03 (1.84) mm, PML 1.10–1.38 (1.26) mm, PSL 1.5. Pronotum (Fig. 4D) transverse, glossy, widest at apical 1.19–1.50 (1.37) mm, EL 3.25–4.36 (3.87) mm, EW 2.24–2.88 1/3; densely and irregularly covered with definite punctures; (2.64) mm. anterior and posterior angles acute; lateral margins rounded in Distribution. Japan (Hokkaido, Honshu: Tohoku, Kanto, front and strongly curved near base. PW/PML 1.44–1.51 (1.47), and Chubu districts). PW/PSL 1.28–1.34 (1.31); PML/PSL 0.88–0.90 (0.89). Elytra Biological notes. This is rare species in Japan, and glossy, oblong, widest at basal 1/3, densely and irregularly is collected from Lycoperdon gemmatum (Agaricaceae) covered with definite punctures; EL/EW 1.44–1.46 (1.45). (Hirano, 2011). This species is mainly distributed in low Intercoxal process of mesosternum (Fig. 4H) moderately to mountainous altitude area (ca. 300–750 m in altitude) in long, narrow and carinate. Metasternum transverse, punctate, northern part of Japan. pubescent, 2.09 times as long as metacoxal cavity. Fore Etymology. The species name is dedicated for Mr. trochanter (Fig. 2G) quadrate; fore femur widest at basal Yukihiko Hirano, who recognized this species firstly. 2/3, 2.00 times as wide as tibiae; fore tibia (Fig. 2H) setose Remarks. This species is similar to L. koltzei and L. at apical 1/3, with tooth in inner margin. Ventrite 5 (Fig. 6G) ferruginea in the shapes of male genitalia (Strohecker, 1970). moderately flattened at apex. Male genital segment (Fig. 6L) formed by segment 9 (tergite and sternite 9) with paired Discussion apophyses fused near base; right apophyse slightly curved at In the present paper, we recognized four species of the base. genus Lycoperdina from Japan. Based on the label data,

June 15, 2017, JJSE 23 (1) 112 Sogoh, K. and H. Yoshitomi

Entomologischen Sammlung zu Berlin. W. Engelmann, Leipzig. Gorham, H. S., 1873. Descriptions of a new genus and two new species of Coleoptera from Japan. Entomologist's Monthly Magazine, 9: 257–258. Gorham, H. S., 1874. Description of a new Japanese Lycoperdina. Entomologist's Monthly Magazine, 11: 151. Gorham, H. S., 1887. Revision of the Japanese species of the coleopterous family Endomychidae. Proceedings of the General Meetings for Scientific Business of the Zoological Society of London, 1887: 642–653. Hashimoto, K. & Hayashi, F., 2014. Cantharidin world in nature: a concealed assemblage with interactions via the terpenoid cantharidin. Entomological Science, 17: 388–395. Hayashi, N., 1978. [Bibliography of larvae of (Coleoptera) in Japan]. Kenkyu to Hyoron, (21): 30–45. (In Japanese.) Hayashi, N., Fukuda, A., & Kurosa, K., 1959. Coleoptera. Pp. 447–449. Illustrated larvae of Japan. Hokuryukan. (In Japanese.) Hirano, Y., 2011. [Notes on the genus Lycoperdina from Japan]. Kanagawa-Chûhô, (175): 19–22. (In Japanese.) Jung, B. H., 2014. of Endomychidae Leach (Coleoptera: Cucujoidea) in Korea. Korean Journal of applied entomology, 53 (1): 39–49 Latreille, P. A., 1807. Genera crustaceorum et insectorum secundum Fig. 8. Box plot of the altitude of collecting site of Lycoperdina ordinem naturalem in familia disposita, iconobis exemplisque spp. in Japan. The altitude data were traced from the specimen plurimis explicta. Tomus tertius. Amand Koenig, Parisii et labels. Argentorati. 258 pp. two or three species were collected sympatrically (e.g. L. Mulsant, E., 1846. Histoire Naturelle des Coléoptères de France. Sulcicolles-Securipalpes. Maison, Paris. xxiv + 280 pp., 1 pl. castaneipennis and L. dux in “Shiratsue-san” and “Takanawa- Ohta, Y., 1931. Beiträge zur Kenntnis der Endomychiden Japans. san”, L. castaneipennis and L. hiranoi in “Yumoto”, L. dux Journal of the Faculty of Agriculture, Hokkaido Imperial and L. hiranoi in “Tsuna-onsen” and “Tsunagi-onsen”, L. University, 30: 205–244. castaneipennis, L. dux and L. hiranoi in “Kami-Hinawa”). On Pic, M., 1945. Coléoptères du globe (suite). L’Échange, Revue the other hand, Fig. 8 shows that the distribution of four species Linnéenne, 61: 10–16. is weakly associated with the altitude. However, we cannot find Sasaji, H., 1985. Endomychidae. Pp. 239–242, pl. 39. The that two or more species live in a same mycelium or not. Thus, Coleoptera of Japan in Color, 3. Hoikusha, Osaka. (In Japanese.) a full understanding of the distributional pattern of this genus Shockley, F. W., Tomaszewska, K. W. & McHugh, J. V., 2009. An awaits further sampling and investigation in the field. annotated checklist of the handsome fungus of the world (Coleoptera: Cucujoidea: Endomychidae). Zootaxa, Acknowledgements 1999: 1–113 Strohecker, H. F., 1953. Coleoptera, Endomychidae. pp. 1–145. We wish to express our deep gratitude to Mr. Y. Hirano In: Wytsman, P. (ed.), Genera Insectorum. Louis Desmet- (Kanagawa), Dr. M. Maruyama (KUMJ), and Dr. T. Kurihara Verteneuil, Bruxelles. (TPMJ) for giving us the chance to examine the precious Strohecker, H. F., 1970. Ergebnisse der zoologischen Forschungen materials, and to Mr. H. Kamezawa (Saitama), Dr. K. Konishi von Dr. Z. Kaszab in der Mongolei. 225. Endomychidae and Dr. M. Sakai (both EUMJ) for their useful suggestion. (Coleoptera). Annales Historico-Naturales Musei Nationalis Hungarici, 62: 245–247. Taniguchi, K., 1942. Ueber 21 Endomychiden aus Japan, Korea References und Formosa. The Entomological world, 10 (97): 152–163. (In Arrow, G. J., 1920. A contribution to the classification of the Japanese, with German title.) coleopterous family Endomychidae. Transactions of the Tomaszewska, W. K., 2005. Phylogeny and generic classification of Entomological Society of London, 1920: 1–83. the subfamily Lycoperdininae with a re-analysis of the family Arrow, G. J., 1923. Notes on endomychid Coleoptera and descriptions Endomychidae (Coleoptera: Cucujoidea). Annales Zoologici, of new species in the British Museum. Transactions of the 55: 1–172. Entomological Society of London, 1923: 484–500. Tomaszewska, W. K., 2007. Endomychidae (the remaining Csiki, E., 1910. Pars 12: Fam. Endomychidae. pp. 1–68. In: subfamilies). pp. 559–568. In: Löbl, I. & Smetana, A. (eds), Schenkling, S. (ed.), Coleopterorum Catalogus. W. Junk, Berlin. Catalogue of Palaearctic Coleoptera. Volume 4. Elateroidea – Gerstaecker, A., 1858. Monographie der Endomychidae, einer Derodontoidea – Bostrichoidea – Lymexyloidea – Cleroidea – Familie der Coleopteren. XIV + 433 p., 3 pls. In: Gerstaecker, Cucujoidea. Apollo Books, Stenstrup. A. (ed.), Entomographien. Abhandlungen im Bereich der Gliederthiere, mit besonderer Benutzung der Koenigl. [Received: April 14, 2017; accepted: May 24, 2017]

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