Russia: the Russian Far East (Primorskiy Territory).

Material examined. 38 (17♂ 17♀ 4 exs).

Host Carpinus cordata: 1♀, Oshirakawa, Azumi, Matsumoto, Nagano Pref., 29.vii.1990, N. Hirano leg., 7.vii.1990(larva), (genitalia slide no. OPU-SK568)

Host C. japonica: 1♂ 1♀, Oshirakawa, Azumi, Matsumoto, Nagano Pref., 26&29.iv.1991, N. Hirano leg., 27.x.1990(larva), SK563, 564; [Soni, Uda, Nara Pref., S. Kobayashi leg.]: 1♀, Konagao, 18.vii.2012em., 14.vii.2012(larva); 1♀, Kameyama, Taroji, 23.vii.2011em., 24.vi.2011(larva). 3♂, Kabuto-dake climb point, 9&16.ix.2010em., 17.x.2010(larva), SK406.

Host C. laxiflora, S. Kobayashi leg.: 1♂, Mitsuigatani, Konagao, Soni, Uda, Nara Pref., 16.ii.2011em., 14.ix.2011(larva), SK407; 1♂ 1♀, Ohshirakawa, Nagawa, Matsumoto, Nagano Pref., 7.viii.2012em., 3.viii.2012(larva).

Host C. tschonoskii: 1♂, Kiso-Hukusima, Nagano Pref., 27.iv.1976em., T. Kumata leg., Rearing code: Kumata 1520, Genitalia slide no. HS-G54, deposited in HUM (Sato, 2011: fig.II-14.3A); 1 ex, Tawamine, Konagao, Soni, Uda, Nara Pref., 18.viii.2015em., S. Kobayashi leg., 15.viii.2015(pupa); Ehime Pref.: 3♂ 4♀, Matsuyama, 23.iv.1965, H. Kuroko leg.; 1♀, Nametoko nr. Uwazima, 1.v.1981em., T. Kumata leg., K2279, HS-G57 (HUM) (Sato, 2011: fig.II-14.3B); [Hikosan, Fukuoka Pref., H. Kuroko leg.]: 1♀ 1 ex, 2.v.&22.vii.1954; 2♀ 1 ex, 1&4.v.1957; 1♀, 10.viii.1957.

Host Corylus sieboldiana: 1♂, Saiko-nishi, Fuji-Kawaguchiko, Yamanashi Pref., 16.viii.2011em., S. Kobayashi leg., 6.viii.2011(larva), SK408. 1♀, Mt. Kuroiwa, Nagano Pref., 6.iv.1987em., H. Kuroko leg., 16.x.1986(larva).

Host Ostrya japonica: 1♂, Sapporo, Hokkaido, 21.vii.1959, T. Kumata leg.

Host unknown: 5♂ 2♀ 1 ex, Mt. Wasamata, Nishihara, Kamikitayama, Nara Pref., 23&24.viii.2011, collected by light trap (L.T.), T. Hirowatari, K. Ikeuchi, Y.-S. Bae & S. Kobayashi leg., SK570.

See original description.

Adult (Fig. 2A–C). Male and female. Wing expanse 6.6–8.8 mm in Japanese specimens, 8.6, 8.8 mm in hibernating generation, and 6.6–7.7 mm in summer and autumn generations. Forewing (Fig. 2A–C) pale to dark ocherous, especially blackish brown in overwintering form (Fig. 2C). Japanese specimens of summer and autumn generations have more distinctly grayish brown scales along the costal margin and apical part of the forewing than the type series collected in July and August.

Figure 1. 

Habitats and hostplants of Coptotriche species. A–E C. minuta Diškus & Stonis, 2014, Soni, Nara Prefecture F–I C. symplocosella sp. n. A Habitat, Mitsuigatani, Konagao, 710 m B Habitat, Nagano, 600 m C Leaves of Carpinus laxiflora at Nagano D Branches of C. japonica at Kumawata, Konagao E Leaves and fruits of C. japonica at Kumawata F Type locality, Adachi Park, Kokura, Fukuoka Prefecture G Habitat, Jyozan, Mitsushima, Tsushima Is., Nagasaki Pref. H Habitat and host plants, Symplocos lucida I Symplocos lucida tree.

Figure 2. 

Adults of Tischeriidae species from Japan. A Coptotriche minuta Diškus & Stonis, 2014, male (Nara Prefecture) B Female (Nara Pref.) C Male, overwintering form (Nagano Pref.) D C. symplocosella sp. n., holotype male E Paratype female. F Tischeria kumatai sp. n., holotype male G Paratype female (Nagano Pref.) H T. relictana Ermolaev, 1986, male (Tokushima Pref.), hostplant unknown I Female (Hokkaido), hostplant: B. ermanii J Male (Nagano Pref.), hostplant: Betula grossa K Female, same hostplant L Female (Nara Pref.), hostplant unknown. Scale bar: 1 mm.

Male genitalia (Fig. 3A–C) (4 preparations examined).

Figure 3. 

Genitalia of Coptotriche minuta and C. symplocosella. A–D: C. minuta. E–H: C. symplocosella. A, E Phallus, ventral view B, F Male genitalia, ventral view C, G Right valva, inner view D, H Female genitalia, ventral view Db, Dc, Dd Same, separated two pairs of prela towards posterior part Db Lateral view Dcd ventral view De Corpus bursae and ductus spermatheca. Abbreviations: aa: apophysis anterioris; an: anellus; ap: apophysis posterioris; cb: corpus bursae; la: lateral arm of 8th tergite; pr: prela; sl: setose lobe on 9th tergite; so: socius; sp: spermatheca; tr: transtilla; un: uncus; va: valva; vi: vinculum; ve: vestibulum; 8s: 8th sternite.

Female genitalia (Fig. 3D) (3 preparations examined). Similar to C. japoniella, but different in having a long ductus spermathecae with convolutions; anterior part with minute spines (Fig. 3De).

Russia: the Russian Far East (Primorskiy Territory) (Stonis et al., 2014); Japan: Honshu (Nagano, Yamanashi and Nara Prefectures), Shikoku (Ehime Prefecture), Kyushu (Fukuoka Prefecture).

Carpinus cordata Blume, C. japonica Blume, C. laxiflora (Siebold & Zucc.) Blume, C. tschonoskii Maxim., Corylus sieboldiana Blume and Ostrya japonica Sarg. (Betulaceae).

(Figs 7–8). The larvae were observed from June to October and hibernated in the final larval stage in Nara Prefecture. The first to second larval instars form a short linear mine towards the leaf edge (Fig. 8F, H). Later instar larvae fold the leaf edge down, forming a blotch mine, then widening it into the surrounding area (Fig. 8C, D); there are usually one to two mines per leaf (Figs 7, 8A, B, G). Frass is ejected through holes in the mine (Fig. 8H). Late and final instar larvae are about 4.0–5.5 mm long and pale yellowish green in coloration (Fig. 8E, I). The folded mines in the leaf edge are 10–20 mm in length, and the late blotch mines are 2–6 mm in width and 7–15 mm (Fig. 7B, C, E) or 20–46 mm (Fig. 7A, D) in length and ocherous in coloration. Pupation takes place within the mine.

Two pairs of prelae were observed to expand caudally and form a hump-shape in the female genitalia of some specimens (Fig. 3Db–d).

The folded, leaf-edge mines of this species resemble at first sight those of foreign congeneric species feeding on Fagaceae and Rosaceae, e.g. Coptotriche citrinipennella (Clemens, 1859), C. gaunacella (Duponchel, 1843), C. crataegifoliae (Braun, 1972) and C. agrimoniella (Braun, 1972). However, larvae of C. citrinipennella form more tightly folded and narrow mines (Braun 1972), while other species form more expanded mines. Fully expanded mines of C. minuta are easily distinguishable from those of the other Japanese Coptotriche species as shown in Fig. 13, although all of them are irregular blotch mines lined with a few folds. The mine of C. minuta is most similar to that of C. angusticollella, but the fold of C. minuta is obviously smaller than that of C. angusticollella. A mine of C. minuta may look like a pupal shelter of Roeslerstammia pronubella ([Denis & Schiffermüller], 1775), Roeslerstammiidae, which utilizes the same hostplant, Carpinus laxiflora (Hirowatari et al. 2012, figs 7, 8).

47(11♂ 9♀ 27 exs)

Holotype ♂, JAPAN: Kyushu: Adachi Park, Kokura, Fukuoka Pref., 9.iv.2012em., host Symplocos lucida, 20.iii.2012(larva), S. Kobayashi leg. Paratypes 10♂ 9♀, Mikata, Mitsushima, Tsushima, Nagasaki Pref., 25.iv.–6.v.2012em., S. Kobayashi leg., host S. lucida, 27.iii.2012 (larva), SK402–405.

20 exs, same data as paratypes.

Pupae. 7 exs, Mikata, Mitsushima, Tsushima, Nagasaki Pref., 27.iv.2012, S. Kobayashi leg., host S. lucida, 27.iii.2012(larva).

The color of the scaling is very similar to that of many other Coptotriche species; the new species differs from other members of the genus in the combination of the rather long uncus (Fig. 3F), and the gently curved slender valva (Fig. 3G) in the male genitalia, and the very small corpus bursae (Fig. 3H) in the female genitalia.

Adult (Fig. 2D–E). Wing expanse 8.7 mm; forewing length 4.0 mm in holotype, 7.3–9.2 mm in paratypes (8.0 mm on average for 21 specimens). Head: palpi cream white to ocherous; frons smooth, blackish brown; vertex tuft blackish brown centrally, frontal and lateral tufts brown; collar brown to grayish black apically, comprised of slender lamellar scales; antenna minimally 2/3 length of forewing, brown to golden. Thorax: anterior part black, posterior part grayish black. Forewing brown to ocherous with blackish scales densely covering apex, and tipping termen and tornus; termen with brown scales. Cilia and hindwing blackish gray. Legs pale ocherous. Abdomen: black; anal tuft grayish ocherous.

Male genitalia (Fig. 3E–G) (2 preparations examined). Uncus with oblong claw-shaped lateral lobes. Socii membranous. Tegumen broad and rather long. Valva slender, gently curved inwards (Fig. 3G). Transtilla present. Anellus membranous, indistinct. Vinculum with rather short triangular ventral plate, with rounded anterior part. Phallus tulip-shaped, slender with broad ended apical part (Fig. 3E).

Female genitalia (Fig. 3H) (2 preparations examined). Similar to C. japoniella and C. bifurcula, except corpus bursae very small, slender, with two narrow signa with minute spines, and a short slender ductus spermathecae.

Pupa. (Fig. 11) (3 preparations examined). Brown to dark brown, 4.4–5.1 mm in length. Vertex (Fig. 11A–B) smooth, with a pair of short setae laterally (Fig. 11C). Dorsum A2–A7 with a pair of long setae, and a concentration of very small spines (Fig. 11E). Dorsum A8–A10 (Fig. 11G, H) with a pair of long dorsal spines and a pair of long lateral spines; A10 (Fig. 11F–H) furcated with a pair of short acute processes from caudal apex, rolled on the dorsal side.

Japan: Kyushu (Fukuoka and Nagasaki (Tsushima Is.) Prefectures).

Symplocos lucida (Thunb.) Siebold & Zucc. (Symplocaceae).

The specific epithet, symplocosella, refers to the genus of the hostplant, Symplocos.

(Figs 9–10). Because many young larvae were observed in leaf mines in March, the species seems to overwinter in the larval stage. The larvae mine leaves of an evergreen tree, Symplocos lucida, forming an elongate full-depth blotch mine beginning with a slender, linear shape (Fig. 9A, E), and gradually expanding as they feed and grow (Fig. 9B, F, G); about ~3 cm in length, white to dark yellow; the older mines turn brown in coloration (Fig. 9C). There were usually 1–3 mines per leaf (Fig. 9G). The larva ejects frass through circular holes (Fig. 10). From shed larval head capsules in the mine, we estimate that the species has six larval instars (Figs 9I, 10). The semifinal and final instar larvae are 6.0–7.0 mm long and pale green in coloration. Head capsule widths are 1st instar: 0.21 mm, 2nd: 0.25 mm, 3rd: 0.30 mm, 4th: 0.40 mm. The mature larva lines the mine with silk, so that the upper surface of the mine shows a few folds (Fig. 9C); a pupal cocoon is situated at the end of the mine.

The pupal characters of the new species are similar to those of other Coptotriche species, but the new species has rather short caudal processes.

6 (3♂ 3♀)

Holotype ♂, JAPAN: Hokkaido: Teine, 14.v.1959, host Tilia japonica, T. Kumata leg. (genitalia slide no. OPU-SK486). Paratypes Host. Tilia japonica: 1♂ 1♀, same locality and data of holotype, SK485; 1♀, Mt. Maruyama, Sapporo, Hokkaido, 2.v.2007em., H. Sato leg., 7.ix.2006(larva); 1♀, Oshirakawa, Azumi, Nagano Pref., 31.v.1992em., N. Hirano leg., 7.ix.1991, SK567. Host Tilia sp.: 1♂, Mt. Maruyama, Sapporo, Hokkaido, 24.iii.2007em., H. Sato leg., 7.ix.2006(larva).

The color of the scaling of this new species has little or no diagnostic value. However, the female genitalia exhibit good diagnostic characters, especially the thickened plate-like vestibulum (antrum) (Fig. 4I). Among Tischeria species having a similarly unusual antrum (e.g. T. ptarmica Meyrick (see van Nieukerken 2010) and T. zestica Meyrick (see Puplesis and Diškus 2003), the new species is most similar to T. unca Diškus & Stonis from the Russian Far East (feeding on Quercus), but is recognizable by the slender posterior plate of the antrum and the lack of spines in the corpus bursae in the female genitalia, and the long spiral shaped juxta (Fig. 4E–F) and the valva with a very slender basal half (Fig. 4A) in the male genitalia. The South African species, T. antilope Puplesis, Diškus & Mey (female unknown) also has a similarly shaped juxta and valva, but differs from the new species by the narrow ventral plate of the vinculum, the longer valva and the lack of a pair of short lateral processes on the juxta (Puplesis and Diškus 2003, figs 586, 589). A Far Eastern Russian species, T. puplesisi Kozlov (female unknown), differs from the new species by the broader valva and short, stout juxta (Kozlov 1986, fig. 2).

Figure 4. 

Genitalia of Tischeria kumatai. A Left valva, inner view B Male genitalia with phallus, juxta and left valva removed, ventral view C Left lateral lobe of uncus, lateral view D Juxta, ventral view E Phallus and juxta, lateral view F Same, ventral view G Valvae, phallus, juxta and vinculum, ventral view H Female genitalia, distal part, ventral view I Antrum, ventral view J Corpus bursae, ventral view. Abbreviations: ju: juxta; ph: phallus; so: socius; un: uncus; va: valva; vi: vinculum.

Adult (Fig. 2F–G). Wing expanse 7.3 mm; forewing length 3.5 mm in holotype, 7.4 and 8.4 mm (3.3 and 3.9 mm) in paratypes. Head: palpi cream white; face smooth, cream white; vertex tuft white mixed with slender pale ocherous lamellar scales; antenna slightly longer than or equal to half of forewing, basally pale ocherous, apically ocherous to brown. Thorax pale ocherous. Forewing pale ocherous with scattered brown scales on the costal half towards the apical area. Cilia and hindwing blackish gray. Legs white to pale ocherous. Abdomen: brown; anal tuft grayish ocherous.

Male genitalia (Fig. 4A–G) (1 preparation examined). Uncus with very slender lateral lobes (Fig. 4C). Socii membranous. Tegumen narrow, marginally reinforced with a pair of slightly inwardly curved arms (Fig. 4B). Valva long, basally very slender, apically rounded, distally covered with fine setae and with a long slender dorsal process (Fig. 4A). Ventral plate of vinculum long and narrow, rounded anteriorly. Juxta very long, comprising two pairs of spiral curved processes (Fig. 4D–F); one pair connecting to the middle of the phallus, equal to the length of valva, apically forming a spiral shape (Fig. 4E–F); the other pair more slender, half the length of the valva, basally slightly curved laterally (Fig. 4D). Phallus slender, distinctly broadened at basal end, forming a pale slender membranous structure from the middle to apex.

Female genitalia (Fig. 4H–I) (2 preparations examined). Antrum (Fig. 4I) strongly sclerotized, plate-like, slightly broadened medially, with a pair of pale spatulate plates at 1/3 of antrum and a pair of very slender long processes. Ductus spermathecae membranous and slender, with 3–4 coils. Corpus bursae small and smooth (Fig. 4H).

Japan: Hokkaido, Honshu (Nagano Prefecture).

Tilia japonica (Miq.) Simonk. (Malvaceae).

The specific epithet, kumatai, is dedicated to Dr Tosio Kumata, who is one of the great Lepidoptera taxonomists and collected the holotype and some of the paratypes.

The larvae form dark gray blotch mines on the leaf edge which are very similar to the folded mines of Coptotriche minuta on Carpinus japonica. Tischeria kumatai is common in Nagano Prefecture.

Russia: the Russian Far East (Sakhalin).

6(3♂ 3♀)

Host Betula ermanii: 1♂ 1♀, Sapporo, Hokkaido, 1.v.1959, T. Kumata leg., HK387♂, SK487♀.

Host Betula grossa, N. Hirano leg.: 1♂, Oshirakawa, Azumi, Matsumoto, Nagano Pref., 25.iv.1990em., 23.x.1989(larva), SK565; 1♀, same locality, 20.v.2004em., 11.x.2003(larva), SK566.

Host unknown: 1♀, Mt. Wasamata, Nishihara, Kamikitayama, Nara Pref., 23.viii.2011(L.T.), T. Hirowatari, K. Ikeuchi, Y.-S. Bae & S. Kobayashi leg., SK569. 1♂, Kumosa-yama, Kamiyama, Tokushima Pref., 22.viii.2010(L.T.), K. Yamada, T. Hirowatari, K. Ikeuchi, S. Kobayashi and K. Akita leg., SK463, deposited in TKPM.

Tischeria relictana resembles Coptotriche species associated with Rosaceae in that the wings and thorax are covered with gray scales. However, this species can be regarded as a member of Tischeria by the presence of a developed juxta in the male genitalia. Although having divided valvae as well as some other congeneric species (e.g., T. zestica Meyrick and T. martinkrugeri Puplesis & Diškus), T. relictana clearly differs from the others in the double juxta with anteriorly semicircular sclerotized diaphragma (Fig. 5A). A Far Eastern Russian species, T. sichotensis Ermolaev, has female genitalia similar in shape to T. relictana, but the former is separated from the latter by the presence of two acute lateral lobes of the antrum and the short spine-like pectinations in the caudal part of the corpus bursae (Stonis et al. 2014, figs 42, 43).

Figure 5. 

Genitalia of Tischeria relictana. A Male genitalia, ventral view, with right valva removed B Right valva, inner view C Female genitalia, ventral view (hostplants: Betula grossa) D Antrum, spermatheca and corpus bursae (hostplants unknown). Abbreviations: an: antrum.

Adult (Fig. 2H–L). Male and female. Wing expanse 6.1–7.1 mm; forewing length 3.0–3.2 mm in Japanese specimens. Head: palpi blackish brown; face smooth, blackish brown; vertex tuft blackish brown; antenna blackish brown, equal to half of forewing in length. Thorax black. Forewing blackish brown to black. Cilia and hindwing blackish gray. Legs blackish brown. Abdomen blackish brown; anal tuft grayish ocherous.

Male genitalia (Fig. 5A, B) (2 preparations examined). Uncus with long and very slender lateral lobes. Socii membranous. Tegumen strongly sclerotized marginally with a pair of slightly inwardly curved frames. Valva (Fig. 5B) broad, covered distally with fine setae, and having a long, slender, dorsal process. Ventral plate of vinculum narrow, triangular. Juxta short, comprising two pairs of processes, one pair connecting to the middle of the phallus, half the length of the valva, narrow medially, broadened apically; the other pair needle-shaped, 1/4 length of valva, slightly broadened basally. Transtilla absent. Diaphragma anteriorly sclerotized, a semicircular plate, folding round the phallus and contacting the needle-shaped part of the juxta ventrally. Phallus (Fig. 5A) slender, distinctly broadened at basal end, forming a pale slender membranous structure from the middle to apex.

Female genitalia (Fig. 5C, D) (3 preparations examined). Similar to T. sparmanniae and T. zestica, but differs in having short apophyses anteriores and posteriores, a slender ductus bursae and the corpus bursae without spines.

Russia: the Russian Far East (Sakhalin) (Ermolaev 1986); Japan: Hokkaido, Honshu (Nagano and Nara Prefectures), Shikoku (Tokushima Prefecture).

Betula ermanii Cham., B. grossa Siebold & Zucc. (Betulaceae).

The detailed biology of this species is unknown. The larvae mine leaves of Betula spp., according to label data of adult specimens.

We collected a female adult of this species at a light trap on Mt. Wasamata, Nara Prefecture, where we also collected tischeriid mines on Betula grossa (Fig. 12). The larvae formed an ocherous to dark gray oblong mine, similar to that of Coptotriche minuta, on the leaf edge or along the leaf vein. Unfortunately, adults did not emerge. We also collected a male adult of this species in a light trap in a deciduous broadleaf forest where Betula grossa grows, on Mt. Kumoso, Tokushima Prefecture. C. Doorenweerd (pers. comm. E.J. van Nieukerken) collected a pupa in Hokkaido from a folded leafmine at the leaf edge on Betula of which the DNA barcode groups with other species of Tischeria and not Coptotriche. Judging from these data, the mines on Betula can be considered to have been made by T. relictana.

Russia: the Russian Far East (Primorskiy Territory).

29(18♂ 11♀)

Host Quercus acutissima: 2♂ 3♀, Komaga-take-SA, Komagane, Nagano Pref., 12–15.viii.2010em., S. Kobayashi leg., 1.viii.2010(larva), SK554, 594; 9♂ 8♀, Tsubata-cho, Ishikawa Pref., ix.1983em., I. Togashi leg., viii.1983, HK705, 706; 1♂, Awara-cho, Fukui Pref., 25.viii.1987em., HK948; 1♂, Kazura, Soni, Uda, Nara Pref., 31.vii.2013em., S. Kobayashi leg., 13.vii.2013(larva), SK555; 2♂, Iwawaki-san, Osaka Pref., 23.v.1980em., H. Kuroko leg., 3.xi.1979.

Host Quercus crispula: 2♂, Minodo, Nagano Pref., 13.v.1980em., H. Kuroko leg., SK595.

Host unknown: 1♂, Ohdaigahara, Kamikitayama, Yoshino, Nara Pref., 20.vii.2009(L.T.), T. Hirowatari, K. Ikeuchi, S. Kobayashi, K. Akita, A. Inotsuka & T. Yoshida leg., SK214.

See Stonis et al. (2014).

Male genitalia (Fig. 6). (9 preparations examined). See Sato (1993, 4A–E) and Stonis et al. (2014, fig. 26–29, 31, 33, 35).

Figure 6. 

Male genitalia of Japanese specimens of Tischeria decidua siorkionla. A, D, G, J, K Right valva, outer view B, E, H Juxta, ventral view C, F, I Phallus, ventral view L, M Preparation by Dr H. Kuroko A–F, J, L Host Quercus acutissima G–I Host Q. crispula K Host unknown A–C Osaka Pref., genitalia slide no. SK596 D–F Nagano Pref., SK594 G–I Nagano Pref., SK595 J Nara Pref., SK555 K Nara Pref., SK214 L Ishikawa Pref., HK705 M Same data, HK706. Scale bar 100 μm.

Female genitalia See Sato (1993, fig. 4F).

Russia: the Russian Far East (Primorskiy Territory); Japan: Hokkaido, Honshu, Kyushu (Tsushima Is.).

Quercus acutissima Carruth., Q. crispula Blume, Q. dentata Thunb., Q. serrata Thunb., and Q. variabilis Blume, Fagaceae in Japan (Sato 1993). Q. mongolica Fisch. ex Ledeb. and Q. serrata in Russia (Kozlov 1986; Stonis et al. 2014).

(Fig. 13–3). See Sato (1993) and Stonis et al. (2014).

In Japan, this species had been treated as ‘T. decidua Wocke’ until the East Asiatic subspecies T. decidua siorkionla was described by Kozlov (1986). Stonis et al. (2014) reported that Japanese representatives of T. decidua belonged to the subspecies siorkionla Kozlov. In the Japanese specimens we studied, the apex of the valva is broader (present study: 50–65 μm; Stonis et al. (2014): 65 μm), but other characters were considered to lie within the range of individual variation, e.g., some specimens have a rather prominent median bulge and sinuous inner margin of the valva (Fig. 6A, D, G, L; Sato 1993, 4D), i.e. more similar to that of the nominotypical European subspecies; others have a rather longer but less prominent median bulge and nearly straight inner margin of the valva (Fig. 6J, K, M; Sato 1993, fig. 4A, E), i.e. more similar to that of T. d. siorkionla. The chitinized basal part of the phallus tends to be less developed and the transverse bar is shorter in Japanese material than in the nominotypical European subspecies (Fig. 6C, F, I), as shown by Stonis et al. (2014). In conclusion, we treat the Japanese representatives as T. decidua siorkionla following Stonis et al. (2014) on the basis of the broader apex of the valva and the less developed basal part of the phallus (Fig. 6).