Heterhelus (Heterhelus) scutellaris ( Heer, 1841 ), 1899

Hisamatsu, Sadatomo, 2011, A review of the Japanese Kateretidae fauna (Coleoptera: Cucujoidea), Acta Entomologica Musei Nationalis Pragae 51 (2), pp. 551-585 : 563-567

Heterhelus (Heterhelus) scutellaris ( Heer, 1841)

( Figs. 1G–H , 6 )

Cateretes scutellaris Heer, 1841: 412 . Type locality: Martygni im Wallis [ Switzerland].

Heterhelus scutellaris: GANGLBAUER (1899: 451) [redescription]; REITTER (1911: 13) [in key]; GROUVELLE (1913: 15) [catalogue]; SPORNRAFT (1967: 23) [in key]; AUDISIO (1980:20) [in key]; AUDISIO (1993: 814) [redescription, note]; KIREJTSHUK (1992: 214) (partim) [in key]; JELÍNEK & AUDISIO (2007: 458) (partim) [catalogue].

Heterhelus (Heterhelus) scutellaris : S-T. HISAMATSU & LEE (2007: 384) [list].

Cercus sambuci Erichson, 1843: 229 . Type locality: Germany. Synonymized by GANGBAUER (1889: 451) with Cateretes scutellaris Heer.

Cercus sambuci: ERICHSON (1845: 127) [redescription, note]; STURM (1844: tab.CCLXXXIX, fig. c. C) [figure]; STURM (1845: 9) [redescription, note]; REDTENBACHER (1849: 162) [in key]; REDTENBACHER (1858: 323) [in key].

Cercus (Heterhelus) sambuci: MURRAY (1864: 234) [redescription, note]; MARSEUL (1885: 24) [redescription].

Cercus (Heterhelus) longipennis Murray, 1864: 234 . Type locality: Dauria meridionalis [= Russia, Transbaikalia]. Synonymized by KIREJTSHUK (1989: 146) with Cateretes scutellaris Heer.

Cercus (Heterhelus) longipennis: MARSEUL (1885:25) [redescription]; KIREJTSHUK (1989: 146) [lectotype designation and synonymy].

Cercus longipennis: GEMMINGER & HAROLD (1868: 803) [catalogue].

Heterhelus longipennis: GROUVELLE (1913: 15) [catalogue].

Amartus (Heterhelus) japonicus Reitter, 1878: 166 . Type locality: Japonia [= Japan]. Synonymized by KIREJTSHUK (1989: 146) with Cateretes scutellaris Heer.

Amartus (Heterhelus) japonicus: KIREJTSHUK (1989: 146) [lectotype designation and synonymy].

Heterhelus japonicus: REITTER (1884: 258) [in key]; REITTER (1885: 104) [list]; GROUVELLE (1913: 15) [catalogue]; HAYASHI (1978: 4) [description of larva].

Amartus (Heterhelus) angusticollis Reitter, 1878:167 . Type locality: Japonia [= Japan].Synonymized by HISAMATSU (1979: 247) with Amartus (Heterhelus) japonicus Reitter.

Amartus (Heterhelus) angusticollis: HISAMATSU (1979: 247) [synonymy].

Heterhelus angusticollis: REITTER (1884: 258) [in key]; REITTER (1885:104) [list]; GROUVELLE (1913: 15) [catalogue]; JELÍNEK & AUDISIO (2007: 458) [catalogue].

Type material examined. Amartus (Heterhelus) japonicus : LECTOTYPE: J ( ZMHB): ‘J // Amartus / japonicus n. sp. // Lectotypus Heterhelus / japonicus (Reitt.) / design. Kirejtshuk 1986 // Heterhelus / solani Heer / A. G. Kirejtshuk det. 1999’. PARALECTOTYPES ( ZMHB): 2 JJ, no locality labels.

Additional material examined. CZECH REPUBLIC: BOHEMIA: 2JJ 4♀♀, Čelákovice , 9.v.1959, J. Jelínek leg. AUSTRIA: 1 J, Wolfsberg , vii.1911, Gustav Schaaff leg. RUSSIA: FAR EAST: 1 J 1 ♀, Vladivostok, Sib. [eria] or., Jureček and Hedviga Jurečková leg. JAPAN: HOKKAIDO: 4 JJ 1 ♀, Hitsujigaoka, Sapporo , 11.v.1995, K. Ishida leg. ; 1 J, Sôunkyo , 17.vii.1970, M. Sakai leg. ; 1 ♀, Shakotan, 23.vi.1986, M. Sakai leg. AOMORI: 110 JJ 182 ♀♀, Nakasato, Kitatsugaru-gun , 30.v.1966, A. Abe leg. ; 12 exs., Juniko, Nishitsugaru-gun , 22.vi.1966, A. Abe leg. ; 1 J 1 ♀, Towada, 9.vi.1958, K. Shimoyama leg. YAMAGATA: 2 exs., Sakata, 18.v.1936, K. Suzuki leg. MIYAGI: 11JJ 5♀♀, Kamoshika spa, Mt. Zaô , 29.v.1971, H. Taguchi leg. NIIGATA: 4 exs., Kurokawa, 15.iv.1959, K. Baba leg. ; 3 exs., Noo, 20.iv.1959, K. Baba leg. NAGANO: 1 ex., Kamikôchi, 30.vii.1959, M. Miyatake leg. GUNMA: 1 ex., Numata, 10.v.1950, T. Takei leg. YAMANASHI: 1 ex., Fuji-rindô, 7.–8.vi.1980, Y. Notsu leg. TOKYO: 31 exs., Mt. Takao , 1.v.1960, S. Hisamatsu leg. ; 2 exs., Asakawa , 4.v.1953, S. Hisamatsu leg. ; 10 JJ 8 ♀♀, Izu-ôshima , 3.–4.v.1979, Y. Notsu leg. ; 16 JJ 19 ♀♀, Izu-ôshima, 17.iv.1974, Y. Furuki leg. KANAGAWA: 2 JJ 1 ♀, Mt. Takatori , 14.iv.1951, H. Hattori leg. ; 1 ex., Mt. Ôama, 2.v.1965, R. Kiryu leg. SHIGA: 1 ♀, Mt. Hira, 3.vi.1957, T. Shibata leg. MIE: 1 J, Mt. Oike , 22.iv.1956, M. Satô leg. HYÔGO: 37 exs., Mt. Senzan , Sumoto, 15.iv.1971, M. Tomokuni leg. SHIMANE: 4 JJ 5 ♀♀, Shimoyamasa, Hirose-chô, Nogi-gun, 8.viii.1980, Y. Seiyama leg. YAMAGUCHI: 36 JJ 69 ♀♀, Mt. Shizukisan , Hagi, 23.iv.1970, S. Hisamatsu leg. EHIME: 33 JJ 24 ♀♀, Mt. Akaboshi , 5.v.1966, S. Hisamatsu leg. ; 3 JJ 5 ♀♀, Komenono , 19.–24.iv.1977, A. Oda leg. ; 9 JJ 14 ♀♀, Omogokei , 23.iv.1972, S. Hisamatsu leg. ; 6 exs., Uwajima, 23.iii.1958, F. Takechi leg. TOKUSHIMA: 40 JJ 39 ♀♀, Mt. Tsurigi, 6.vi.1970, S. Hisamatsu leg. FUKUOKA: 3 exs., Inunaki pass, 29.iv.1968, M. Iga leg. NAGASAKI: 1 ♀, Sasuna-Shûshi , Tsushima , 14.iv.1941, T. Shirôzu leg.

Diagnosis. Body coloration fully reddish-yellow to dark brown except reddish brown mouthparts, antennae and legs. Antennae with almost 2-segmented club. Lateral margins of pronotum variable in shape; distinctly projecting at mid-length or uniformly rounded. Abdominal sternite VII rather long, W7/L7 = 2.29, n = 4). Parameres of male genitalia slender and long (LP/WP = 3.10 (n = 1)); apex of median lobe ( Figs. 6C, 6E ) sharply acuminate in lateral aspect. Ovipositor of female genitalia unpigmented in basal part, with long styli.

Redescription. Length 2.2–3.1 mm.

Male. Body ( Fig. 1G ) oval, convex, dully shining, covered with whitish or yellowish setae. Coloration variable, fully reddish-yellow to dark brown except reddish brown mouthparts, antennae and legs.

Head densely punctate, punctures on disc smaller than eye-facet, separated by <1 diameter; interspaces finely reticulate. Frontoclypeal suture incomplete, distinctly visible. Front margin of clypeus with medial arcuate emargination. Labrum broadly arcuate. Mandibles moderately bent inward. Antennae ( Fig. 6O ) 1.12–1.16 times longer than HW (n = 6), with club indistinctly 3-segmented (appearing 2-segmented); approximate ratio of each segment (n = 1) is 2.38: 1.81: 1.69: 1.25: 1.19: 1.06: 1.06: 1.00: 1.19: 1.75: 2.25.

Pronotum ( Figs. 6H–N ) convex, transverse, variable in shape (see ‘Variability’ section below), 1.41–1.54 times as wide as long (n = 10); anterior corner slightly prominent; anterior margin unbordered; basal margin indistinctly bordered, moderately sinuate before obtusely angulate posterior corners; lateral margins narrowly explanate, feebly serrate; punctures on disc similar in size to those on head; interspaces finely reticulate.

Elytra conjointly 1.08–1.18 times as long as wide, 1.96–2.18 times as long as pronotum; punctures on disc larger and shallower than those on pronotum; interspaces smooth.Abdominal tergite VI partially obscured by elytra. Abdominal tergite VII fully exposed, apex truncate. Abdominal tergite VIII externally visible.

Prosternum (excluding prosternal process) 0.37 times as long as mesoventrite (n = 1) 0.28 times as long as metaventrite (n = 1); prosternal process slender, subparallel-sided. Metaventrite convex, shining, densely covered with whitish or yellowish setae; metathoracic discrimen in basal 1/2; punctures on disc distinctly smaller than those on head, separated by one diameter at middle, becoming denser laterally. Inter-mesocoxal distance separated by 1.45 times width of inter-procoxal distance. Inter-metacoxal distance separated by 2.52 times width of inter-procoxal distance. Abdominal sternites shining; approximate ratio of length of abdominal sternites III–VII (n = 1) is 2.64: 1.00: 1.00: 1.79: 2.21; abdominal sternite VII rather long (W7/L7 = 2.29, n = 4). Legs flattened; protibiae rather slender and short, shorter than HW; tarsal claws simple.

Male genitalia with parameres ( Figs. 6A, 6D ) asymmetrical, slender and long (LP/WP = 3.10 (n = 1)); parameres regularly arcuate inward, bearing long setae; apex of median lobe ( Figs. 6C, 6E ) sharply acuminate in lateral aspect; interparameral lobe broadly rounded.

Female. Coloration rarely becoming fully black (see ‘Coloration’ subsection below within ‘Variability’ section). Apical margin of abdominal tergite VII rounded. Ovipositor deeply notched at apex, with long styli.

Variability. This species expresses variation in body coloration and shape of pronotal sides. A total of 849 Japanese specimens were examined to verify species limits and phenotypic plasticity.

Coloration. Dorsal body surface of males (n = 361) appears reddish brown. Females (n = 488) appear largely reddish brown to rarely fully black.

Lateral margins of pronotum ( Figs. 6H–N ). Margins distinctly projecting at mid-length in specimens from northern Japan (Hokkaido and Tohoku regions), becoming more rounded in specimens from western Japan (western areas of Honshû, Shikoku, and Kyûshû regions).

Bionomics. This species is dependent on Sambucus (Caprifoliaceae) plants for larval development. Adults feed on pollen and petals, and larvae develop inside the ovaries where they feed on seeds. Adults also aggregate on flowers of Spiraea (Rosaceae) , Fagaceae ( AUDISIO 1993) and Magnolia kobus DC. var. borealis Sarg. (Magnoliaceae) in Japan ( ISHIDA 1996).

Their life cycle was reported by HAYASHI (1977). According to his observations in Kanagawa Prefecture, Japan, H. scutellaris was common in lowlands, while many populations of H. morio were observed in highland areas. In April, adults gather at flowers of Sambucus racemosa L. subsp. sieboldiana (Miq.) H. Hara (Caprifoliaceae) in synchrony with flowering events. Subsequently, they mate and lay eggs on the flowers. Hatching larvae burrow into the ovaries until seeds mature a few days later, and then they burrow into the seed. Larvae continue to grow until the last instar larvae (probably 3rd instar) within a seed. Although, Sambucus racemosa subsp. sieboldiana had three seeds in an ovary, the larva burrowed into only one seed, and never burrowed into the other two seeds. In mid May, when ovaries ripen into fruit, the larvae bore an exit hole, drop to the ground, and burrow into the soil where they pupate. Adults eclose around mid June, but remain in earthen chambers until the following April. Of the three hundred fruits that were investigated, only forty-two possessed emerging larvae ( HAYASHI 1977).

Distribution. Japan (Hokkaido, Honshû, Shikoku, Kyûshû); Europe, Russia (East Siberia, Far East) and Mongolia ( HISAMATSU 1985, JELÍNEK & AUDISIO 2007).

Note. The holotype of Heterhelus scutellaris was not examined, however, a series of Palaearctic specimens identified by European specialists was utilized for this revision. Herein, I follow Kirejtshuk’s opinion that Heterhelus japonicus is a junior synonym of Heterhelus scutellaris . Heterhelus angusticollis was synonymized with H. scutellaris by HISAMATSU (1979), but no reason for the synonomy was given. JELÍNEK & AUDISIO (2007) listed Heterhelus angusticollis in their catalogue as a valid species. The holotype of H. angusticollis could not be examined for this study, therefore the species remains enigmatic for the author. However, based on the original description, this species seems to be junior synonym of H. scutellaris . Location and study of the types for both species is needed to clarify the taxonomic position of H. angusticollis .

Figures

Fig. 1. Habitus and labels of Kateretidae spp. A–B – Kateretes japonicus Hisamatsu, 1985: A – holotype, male; B – ditto, labels; C–D – Kateretes takagii S-T.Hisamatsu, 2006:C – holotype, male; D – ditto, labels; E–F – Platamartus jakowlewi Reitter, 1892: E – male and F – female (both Tochigi Pref.); G–H – Heterhelus (Heterhelus) japonicus (Reitter, 1878): G – lectotype, male; H – ditto, labels. Scale bars = 1.0 mm.

Fig. 6. Heterhelus (Heterhelus) scutellaris (Heer, 1841).A – tegmen (ventral view); B – median lobe (ventral view); C – median lobe (lateral view); D – parameres of the lectotype of H. japonicus; E – median lobe of the lectotype of H. japonicus (lateral view); F – ovipositor; G – male abdominal sternite VII; H–N – variation of pronotum (H – Hokkaido; I – Aomori; J – Niigata; K – Tokyo; L – Yamaguchi; M – Ehime; N – Fukuoka); O – male right antenna. A, B, C, G, O = Hokkaido, Hitsujigaoka; F = Ehime Pref., Omogokei.