Stephanitis (Stephanitis) takeyai Drake & Maa, 1955

Souma, Jun, 2022, Integrative taxonomy of the Lauraceae - feeding species of the genus Stephanitis (Hemiptera, Heteroptera, Tingidae) from Japan, Deutsche Entomologische Zeitschrift 69 (2), pp. 219-281 : 219

Stephanitis (Stephanitis) takeyai Drake & Maa, 1955

[Japanese name: Tosaka-gunbai] Figs 3E , 5E , 6B , 8E , 10F , 12E , 14F , 16E , 18E , 20C , 22C , 24C , 26E , 28E , 30G , 32G , 37B , 42H-J

Tingis globurifera Matsumura, 1905: 36 (junior primary homonym of Tingis globurifera Walker, 1873). Lectotype by subsequent designation ( Tomokuni 1994: 842), ♂ (Fig. 37B ): Japan: Gifu [= Honshu, Gifu-ken]; ELHU.

Stephanitis globurifera : Matsumura (1907: 148) (new combination).

Stephanitis takeyai Drake & Maa, 1955: 10. New name for Stephanitis globurifera (Matsumura, 1905).

References.

Horváth (1912: 330) (distribution); Drake (1923: 104) (distribution); Takeya (1930: 72) (host plant); Drake and Poor (1937: 403) (distribution); Drake (1948: 55) (checklist: Stephanitis ); Bailey (1950: 148) (invasion); Takeya (1951b: 11) (checklist: Japan); Drake and Maa (1953: 100) (checklist: Stephanitis ); Takeya (1953: 168) (distribution); Takeya (1963: 50) (distribution); Drake and Ruhoff (1965: 364) (catalog); Miyamoto (1965: 91) (monograph); Lee (1969: 226) (nymph, male genitalia); Tomokuni (1981: 109) (distribution); Japanese Society of Applied Entomology and Zoology (1980: 134) (pest); Tomokuni (1985: 156) (distribution); Ichita (1989: 33) (distribution); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Takahashi (1990a: 28) (checklist: Hyogo); Yasunaga et al. (1993: 179) (monograph); Tsukada (1994: 221) (biology); Péricart and Golub (1996: 63) (catalogue: Palaearctic); Japanese Society of Applied Entomology and Zoology (2006: 183) (pest); Miyatake (2006: 27) (host plant); Miyamoto (2008: 158) (monograph); Tsukada (2008: 349) (biology); Yamada and Tomokuni (2012: 208) (monograph); Vétek et al. (2012: 22) (distribution); Aukema et al. (2013: 72) (checklist: Palaearctic); Yano et al. (2013: 26) (distribution); Maehara (2014: 61) (distribution); Barta and Bideň (2016: 195) (distribution); Yamada and Ishikawa (2016: 434) (checklist: Japan); Ito and Sasaki (2018: 20) (checklist: Oita); Okochi (2019: 3) (distribution); Grosso-Silva et al. (2020: 371) (distribution).

Material examined.

Lectotype (1 ♂, ELHU) (Fig. 37B ), JAPAN: Honshu: Gifu [= Honshu, Gifu-ken (approximate coordinates: 35°26'05.5"N, 136°46'16.6"E)]. Non-types (157 ♂♂ 169 ♀♀ 2 nymphs), JAPAN: Honshu: Saitama-ken, Iruma-gun, Moroyama-machi, Takinoiri, Kanikusa-hashi, 28.v.2021, leg. J. Souma (3 ♀♀, TUA); Tokyo-to, Hachioji-shi, Uratakao-machi, Kogesawa-rindo, 6.ix.2016, leg. J. Souma (11 ♂♂ 7 ♀♀, TUA); as above but 12.v.2017 (1 ♂ 1 ♀, TUA); as above 22.v.2017, leg. Y. Kato (1 ♀, TUA); Kanagawa-ken, Sagamihara-shi, Minami-ku, Asamizodai, 4.vi.2017, leg. J. Souma (2 ♂♂ 1 ♀, TUA); Kanagawa-ken, Sagamihara-shi, Minami-ku, Shimomizo, 10.ix.2020, leg. J. Souma (1 ♀, TUA); Kanagawa-ken, Atsugi-shi, Funako, 12.v.2016, leg. J. Souma (1 ♂, TUA); as above but 7.iii.2017 (1 ♀, TUA); as above, but 5.v.2017 (23 ♂♂ 18 ♀♀, TUA); as above but 6.v.2017 (20 ♂♂ 9 ♀♀, TUA); as above, but 7.v.2017, leg. H. Shigetoh (1 ♂, TUA); as above but 11.v.2017 (43 ♂♂ 52 ♀♀, TUA); as above, but 31.v.2017 (4 ♀♀, ELKU); as above but 5.vi.2017 (1 ♀, TUA); Kanagawa-ken, Atsugi-shi, Nanasawa, 6.vi.2017, leg. J. Souma (8 ♂♂ 5 ♀♀, TUA); Kanagawa-ken, Isehara-shi, Oyama, 31.v.2017, leg. Y. Yamada (1 ♂, TUA); Yamanashi-ken, Hokuto-shi, Takane-cho, Kiyosato, 5.ix.2022, leg. J. Souma (1 ♂, TUA); Nagano-ken, Matsumoto-shi, Azumi, 7.ix.2022, leg. J. Souma (2 ♂♂ 1 ♀, TUA); Hiroshima Pref., Kitahiroshima, Nakaso, 23.vii.2022, leg. Y. Uehara (1 ♂, TUA); as above but leg. H. Hashimoto (1 ♂, TUA). Sado Island: Chigusa, 28.viii.2021, leg. J. Souma (1 ♂ 2 ♀♀, TUA); Kamiyokoyama, 28.viii.2021, leg. J. Souma (3 ♂♂ 5 ♀♀, TUA); Kujikawachi, 29.viii.2021, leg. J. Souma (2 ♀♀, TUA); Noura, 29.viii.2021, leg. J. Souma (3 ♀♀, TUA). Shikoku: Kagawa-ken, Takamatsu-shi, Nishiueta-cho, Fujio, 28.iv.2020, leg. Y. Waki (1 ♂, TUA); Kagawa-ken, Takamatsu-shi, Nishiueta-cho, Donguri Land, 23.v.2021, leg. Y. Waki (1 ♀, TUA); Kochi-ken, Kochi-shi, Shigekura, 2.vii.2020, leg. J. Souma (2 ♂♂ 1 ♀ 2 nymphs, ELKU). Kyushu: Fukuoka-ken, Kasuya-gun, Sasaguri-machi, Tsubakuro, 5.vi.2020, leg. N. Tsuji (5 ♂♂ 9 ♀♀, ELKU); Fukuoka Pref., Tagawa, Soeda, Mt. Hiko-san , Takanosubaru, 28.v.2022, leg. Y. Uehara (1 ♂ 1 ♀, TUA); Oita-ken, Kusu-gun, Kokonoe-machi, 12.vii.2017, leg. S. Imada (1 ♀, ELKU); Kumamoto Pref., Aso, Minamiaso, Kain, 12.vi.2022, leg. H. Hashimoto (1 ♀, TUA); Kumamoto-ken, Kuma-gun, Mizukami-mura, Mt. Ichifusa-yama , 27.vii.2022, leg. S. Inoue (1 ♀, TUA); Oita-ken, Hita-shi, Maetsue-machi, Ono, 21.viii.2022, leg. J. Souma (1 ♂ 1 ♀, TUA); Oita-ken, Hita-shi, Maetsue-machi, Yugi, 21.viii.2022, leg. J. Souma (1 ♂, TUA). Ryukyu Islands (northern part): Yakushima Island : Kosugidani, 23.viii.1952, leg. C. Takeya & Y. Hirashima (2 ♂♂ 3 ♀♀, ELKU); as above, but 29.vii.1963, leg. T. Okada (1 ♀, KUM); Hananoego, 24.viii.1952, leg. C. Takeya & Y. Hirashima (17 ♂♂ 19 ♀♀, ELKU; 1 ♀, KUM); as above, but 26.x.1979, leg. S. Makihara (4 ♂♂ 7 ♀♀, NSMT); Onoaida, 20.viii.2021 (3 ♂♂ 5 ♀♀, ELKU).

Diagnosis.

Stephanitis (Stephanitis) takeyai is recognised amongst other species of Stephanitis by a combination of the following characters: head, calli, pronotal disc, marking on hemelytra and ventral surface black (Figs 8E , 10F , 12E , 14F , 16E , 18E , 20C , 22C , 24C ); body in male 2.1 times (in female 1.9 times) as long as maximum width across hemelytra (Figs 3E , 5E , 6B ); rostrum reaching metasternum; pronotum tricarinate (Fig. 26E ); hood dark, longer than median carina of pronotum, wider than maximum width of head across eyes, completely covering eye, higher than median carina of pronotum at highest part, with posterior margin extending near posterior margin of pronotal disc; median carina of pronotum with 2 rows of areolae at highest part; pronotal disc opaque; paranotum more erect, not narrowed posteriorly, 3 rows of areolae at widest part, anterolateral angle slightly protruding anteriad, outer margin gently curved inwards at posterolateral angle, maximum height longer than height of eye (Fig. 28E ); apices of hemelytra close to each other in rest; costal area with 3-4 rows of areolae at widest part; subcostal area in male with 3 rows (in female with 4 rows) of areolae at widest part; discoidal area with 3-4 rows of areolae at widest part; sutural area with 3 rows of areolae at widest part; hypocostal lamina with a single row of areolae throughout its length; R+M (radiomedial) vein carinate; pygophore elevated at centre of venter, with posterior margin slightly emarginate in middle part (Fig. 30G ); and paramere stout, weakly curved inwards at apex, with outer margin cuspidate in middle part, inner margin slightly curved inwards in basal part (Fig. 32G ).

Remarks.

Amongst the Japanese species of Stephanitis , S. (Stephanitis) takeyai is similar to S. (S.) svensoni Drake, 1948, which feeds on Illicium anisatum L. ( Schisandraceae ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012), in general habitus, but the former is easily distinguished from the latter by the following characters: hood longer than median carina of pronotum (shorter in S. (S.) svensoni ), wider than maximum width of head across eyes (narrower in S. (S.) svensoni ), completely covering eye (incompletely covering in S. (S.) svensoni ) (Figs 8E , 10F , 26E ); and subcostal area of hemelytron in male with 3 rows (in female 4 rows) of areolae at widest part (in male 4 rows and in female 5 rows in S. (S.) svensoni ) (Figs 16E , 18E ).

Teratological form.

Segmental oligomery of the antenna was confirmed in S. (S.) takeyai and one examined specimen lacked the left antennal segment IV (Fig. 6B ), as reported in many tingids ( Štusák and Stehlík 1978; Souma 2020b, 2020d, 2020e).

Distribution.

Japan (Honshu; Sado Island; Oki Islands: Dogo Island; Shikoku; Kyushu; Ryukyu Islands (northern part): Yakushima Island) (Fig. 48 ); Austria; Belgium; Canada; Czech Republic; France; Great Britain; Germany; Hungary; Italy; Netherlands; Poland; Portugal; Slovakia; Switzerland; U.S.A. ( Takeya 1963; Yamada and Tomokuni 2012; Vétek et al. 2012; Aukema et al. 2013; Barta and Bideň 2016; Yamada and Ishikawa 2016; Grosso-Silva et al. 2020; present study). Stephanitis (Stephanitis) takeyai is native to Japan, but it has invaded many countries in Europe and North America ( Yamada and Tomokuni 2012). The previous record from India ( Drake and Ruhoff 1965) is an error resulting from the confusion between the type locality of Tingis globulifera Walker, 1873 [= Cochlochila (Physodictyon) bullita ( Stål, 1873)] described from India and T. globulifera Matsumura, 1905 [= S. (S.) takeyai Drake & Maa, 1955] described from Japan (cf. Drake and Maa 1955). The previous records from Amami-Oshima Island, the central part of the Ryukyu Islands ( Takeya 1963), are misidentifications of S. (S.) pyrioides and a nymph pertaining to another genus. In Japan, S. (S.) takeyai inhabits the deciduous broad-leaved forest in a temperate climate of Japan proper (Honshu, Shikoku and Kyushu) and its surrounding islands, which is in the Palaearctic Region.

Host plants.

Aesculus turbinata Blume, “Tochinoki” ( Sapindaceae ) ( Takeya 1963); Cerasus jamasakura (Siebold ex Koidz.) H.Ohba, “Yamazakura” ( Rosaceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Cinnamomum camphora , “Kusunoki” ( Lauraceae ) ( Takeya 1930, 1963; Yamada and Tomokuni 2012; Okochi 2019); Diospyros kaki Thunb., “Kaki” ( Ebenaceae ) ( Takeya 1963; Yasunaga et al. 1993); Elliottia paniculata (Siebold et Zucc.) Hook.f., “Hotsutsuji” ( Ericaceae ) ( Takeya 1963); Hydrangea hydrangeoides , “Iwagarami” (Siebold et Zucc.) B.Schulz ( Hydrangeaceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Illicium anisatum , “Shikimi” ( Schisandraceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Lindera obtusiloba , “Dankobai” ( Lauraceae ) ( Takeya 1963; Yamada and Tomokuni 2012; present study); L. praecox (Siebold et Zucc.) Blume, “Aburachan” ( Takeya 1963); L. triloba , “Shiromoji” ( Takeya 1963); Litsea cubeba (Lour.) Pers., “Aomoji” ( Lauraceae ) ( Takeya 1930, 1963; Yamada and Tomokuni 2012); Lit. umbellata Thunb., “Kuromoji” (Fig. 44E ) ( Takeya 1930, 1963; Yamada and Tomokuni 2012; Okochi 2019; present study); Lit. sericea (Siebold et Zucc.) Blume, “Kekuromoji” ( Takeya 1963; Yamada and Tomokuni 2012); Lyonia ovalifolia (Wall.) Drude, “Nejiki” ( Ericaceae ) ( Takeya 1963; Yamada and Tomokuni 2012; Maehara 2014; Okochi 2019); Machilus thunbergii , “Tabunoki” ( Lauraceae ) (present study); Pieris japonica (Thunb.) D.Don ex G.Don, “Asebi” ( Ericaceae ) ( Takeya 1930, 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; Maehara 2014); Platanus spp., “Suzukakenoki” or “Puratanasu” ( Platanaceae ) ( Miyatake 2006); Pourthiaea villosa (Thunb.) Decne. “Ushikoroshi” ( Rosaceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Rhododendron japonoheptamerum Kitam. var. hondoense (Nakai) Kitam., “Honshakunage” ( Okochi 2019); Salix sp., “Yanagi” ( Salicaceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Styrax japonicus Siebold & Zucc, “Egonoki” ( Styracaceae ) ( Takeya 1930, 1963). Stephanitis (Stephanitis) takeyai feeds on an extremely wide range of angiosperms and is euryphagous, unlike many tingids ( Schuh and Weirauch 2020). However, some records of this species, collected from Hydrangeaceae , Rosaceae , Salicaceae , Sapindaceae , Schisandraceae and Styracaceae ( Takeya 1930, 1963; Yamada and Tomokuni 2012), do not include data on the insect’s development. As this lace bug is the most common Japanese lace bug ( Yamada and Tomokuni 2012), some of the previous host records may be of plant families from which S. (S.) takeyai was accidentally collected. This lace bug sometimes occurs on plantings of C. camphora , D. kaki and P. japonica in its distribution range (present study). This tingid species has been known to be a pest of D. kaki and P. japonica (Japanese Society of Applied Entomology and Zoology 1980, 2006) and, occasionally, it can also cause harm on C. camphora .

Biology.

Stephanitis (Stephanitis) takeyai feeds on the abaxial surface of leaves of the various host plants in Japan (present study). In Japan, this lace bug is trivoltine ( Tsukada 1994, 2008); adults were collected in almost all seasons ( Takeya 1953; Tomokuni 1981, 1985; Ichita 1989; Yasunaga et al. 1993; Tsukada 1994, 2008; Yamada and Tomokuni 2012; Yano et al. 2013; Maehara 2014; Ito and Sasaki 2018; present study); nymphs were collected from April to October ( Tsukada 1994, 2008; Maehara 2014; present study); the overwintering stage is the egg, but third-generation adults are found until March of the following year ( Tsukada 1994; present study).

Figures

Figure 3. Male habitus of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov. from Yonaguni Island, southern part of Ryukyu Islands; B. S. (N.) mendica from Honshu; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai from Shikoku; F. S. (S.) tomokunii sp. nov. from Miyake Island, southern part of Izu Islands. Scale bar: 1.0 mm.

Figure 5. Female habitus of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov. from Miyako Island, southern part of Ryukyu Islands; B. S. (N.) mendica from Honshu; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai from Kyushu; F. S. (S.) tomokunii sp. nov. from Miyake Island, southern part of Izu Islands. Scale bar: 1.0 mm.

Figure 6. Teratological forms of two Stephanitis species from Japan, dorsal view: A. S. (Norba) aperta from Izu-Oshima Island, northern part of Izu Islands; B. S. (Stephanitis) takeyai from Kyushu. Arrows indicate antennae with segmental oligomery and malformed paranotum. Scale bar: 1.0 mm.

Figure 8. Male pronota of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica lacking lateral carina; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov .. Scale bars: 0.2 mm.

Figure 10. Female pronota of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica possessing lateral carina; C-E. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (E) possessing lateral carina and from Honshu (D) lacking lateral carina; F. S. (S.) takeyai; G. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 12. Male pronota of five Stephanitis species from Japan, dorsolateral view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica lacking lateral carina; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 14. Female pronota of five Stephanitis species from Japan, dorsolateral view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica possessing lateral carina; C-E. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (E) possessing lateral carina and from Honshu (D) lacking lateral carina; F. S. (S.) takeyai; G. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 16. Male hemelytra of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 18. Female hemelytra of five Stephanitis species from Japan, dorsal view: A. S. (Norba) ishikawai sp. nov.; B. S. (N.) mendica; C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 20. Rostra of four Stephanitis species from Japan, dorsal view: A. S. (Norba) mendica; B. S. (Stephanitis) tabidula; C. S. (S.) takeyai; D. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 22. Male terminalia of four Stephanitis species from Japan, dorsal view: A. S. (Norba) mendica; B. S. (Stephanitis) tabidula; C. S. (S.) takeyai; D. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 24. Female terminalia of four Stephanitis species from Japan, dorsal view: A. S. (Norba) mendica; B. S. (Stephanitis) tabidula; C. S. (S.) takeyai; D. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 26. Line drawings of the pronota of four Stephanitis species from Japan, dorsal view: A, B. S. (Norba) mendica lacking lateral carina (A) and possessing lateral carina (B); C, D. S. (Stephanitis) tabidula from Honshu (C) and Kyushu (D); E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 28. Line drawings of the pronota of six Stephanitis species from Japan, lateral view: A. S. (Norba) hiurai; B. S. (N.) ishikawai sp. nov.; C. S. (N.) mendica lacking lateral carina; D. S. (Stephanitis) tabidula lacking lateral carina from Honshu; E. S. (S.) takeyai; F. S. (S.) tomokunii sp. nov. Scale bars: 0.2 mm.

Figure 30. Line drawings of the pygophore of five Stephanitis species from Japan, dorsal view: A-C. S. (Norba) ishikawai sp. nov. from Miyako (A) and Yonaguni (B) Islands, central part of Ryukyu Islands and Taiwan (C); D. S. (N.) mendica; E, F. S. (Stephanitis) tabidula from Honshu (E) and Tsushima Islands (F); G. S. (S.) takeyai; H. S. (S.) tomokunii sp. nov. Scale bars: 0.1 mm.

Figure 32. Line drawings of the paramere of five Stephanitis species from Japan, dorsal view: A-C. S. (Norba) ishikawai sp. nov. from Miyako (A) and Yonaguni (B) Islands, central part of Ryukyu Islands and Taiwan (C); D. S. (N.) mendica; E, F. S. (Stephanitis) tabidula from Honshu (E) and Tsushima Islands (F); G. S. (S.) takeyai; H. S. (S.) tomokunii sp. nov. Scale bars: 0.1 mm. Arrows indicate important morphological differences.

Figure 37. Two specimens of Stephanitis species from Japan. A. Paratype of S. (Stephanitis) fasciicarina syn. nov. [= S. (S.) tabidula] collected at type locality deposited in ELKU and its labels; B. Lectotype of Tingis globulifera [= S. (S.) takeyai] deposited in Matsumura's collection of ELHU and its labels.

Figure 42. Living individuals of four species of Stephanitis from Japan: A, B. S. (Norba) mendica from Honshu, male (A) and female (B); C-G. S. (Stephanitis) tabidula, male (C) and female (D) from Honshu and male (E), female (F) and fifth instar nymph (G) from Kyushu; H-J. S. (S.) takeyai, male (H) and fifth instar nymph (J) from Shikoku and female from Honshu (I); K, L. S. (S.) tomokunii sp. nov. from Hachijo Island, southern part of Izu Islands, male (K) and female (L).

Figure 44. Lauraceous host plants of four species of Stephanitis from Japan: A. Cinnamomum yabunikkei from Kyushu, damaged by S. (Norba) mendica; Machilus thunbergii (B) from Honshu, Neolitsea sericea (C) from Sado Island and C. camphora (D) from Kyushu, all damaged by S. (Stephanitis) tabidula; E. Lindera umbellata from Sado Island, damaged by S. (S.) takeyai; F. M. thunbergii from Hachijo Island, damaged by S. (S.) tomokunii sp. nov.

Figure 48. Collection sites of Stephanitis (Stephanitis) tabidula, S. (S.) takeyai and S. (S.) tomokunii sp. nov. used in the present study. Red-filled areas = localities; circles = small isolated islands.