Stephanitis (Stephanitis) tabidula Horvath , 1912

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) tabidula Horvath, 1912

[Japanese name: Kusu-gunbai] Figs 3C, D , 5C, D , 8C, D , 10C-E , 12C, D , 14C-E , 16C, D , 18C, D , 20B , 22B , 24B , 26C, D , 28D , 30E, F , 32E, F , 37A , 38 , 42C-G

Stephanitis (Stephanitis) tabidula Horváth, 1912: 333. Syntype(s): Japan: Kanagawa [Honshu, Kanagawa-ken (a prefecture) or Kanagawa-ku (a ward) of Yokohama-shi (a city) of Kanagawa-ken]; HNHM.

Stephanitis (Stephanitis) fasciicarina Takeya, 1931: 70: Holotype, ♂: Japan: Kyushu, Chikuzen, Akama [= Kyushu, Fukuoka-ken, Munakata-shi, Akama]; ELKU (not found, mislabeling of paratype collected at type locality?). New subjective synonym.

Stephanitis kyushuana Drake, 1948: 52: Holotype, ♂: Japan: Kyushu, Moje [= Kyushu, Fukuoka-ken, Kitakyushu-shi, Moji-ku]; USNM. Synonymised with Stephanitis (Stephanitis) fasciicarina Takeya, 1931 by Takeya (1951b: 11).

References.

Drake and Poor (1937: 403) (distribution); Drake (1948: 56) (checklist: Stephanitis ); Takeya (1951b: 12) (checklist: Japan); Drake and Maa (1953: 101) (checklist: Stephanitis ); Takara and Hidaka (1960: 188) (distribution); Takeya (1963: 42) (distribution: part); Miyamoto (1964a: 275) (checklist: Ryukyu Islands); Drake and Ruhoff (1965: 363) (catalog); Miyamoto (1965: 91) (monograph); Lee (1969: 215) (nymph, male genitalia); Miyamoto (1976: 497) (distribution); Japanese Society of Applied Entomology and Zoology (1980: 194) (pest); Jing (1981: 353) (monograph); Tomokuni (1981: 109) (distribution); Tomokuni (1985: 156) (distribution); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Yasunaga et al. (1993: 178) (monograph); Péricart and Golub (1996: 63) (catalogue: Palaearctic); Japanese Society of Applied Entomology and Zoology (2006: 250) (pest); Tomokuni and Hayashi (2006: 293) (distribution); Miyamoto (2008: 157) (monograph); Yamada and Tomokuni (2012: 208) (monograph: part); Yano et al. (2013: 25) (distribution); Maehara (2014: 60) (distribution); Yamada and Ishikawa (2016: 434) (checklist: Japan); Ahn et al. (2018: 65) (distribution); Okochi (2019: 3) (distribution); Cho et al. (2020: 742) (distribution); Souma (2021b: 31) (distribution).

Material examined.

Non-types collected at type locality of S. (S.) tabidula Horváth, 1912 (77 ♂♂ 69 ♀♀ 8 nymphs), JAPAN: Honshu: Kanagawa-ken, Sagamihara-shi, Chuo-ku, Tanashioda (approximate coordinates: 35°32'03.6"N, 139°20'57.7"E), 19.v.2019, leg. J. Souma (43 ♂♂ 22 ♀♀ 8 nymphs, TUA); as above but 25.v.2019 (4 ♂♂, ELKU; 22 ♂♂ 31 ♀♀, TUA); as above but 25.ii.2020, leg. J. Souma (1 ♂ 12 ♀♀, TUA); as above but 17.xi.2021 (3 ♂♂ 3 ♀♀, TUA); "Mt. Ohokusu" [= Kanagawa-ken, Yokosuka-shi, Ashina, Mt. Ogusu (approximate coordinates: 35°15'00.8"N, 139°37'40.6"E)], 7.viii.1973, leg. K. B. S. (1 ♂, KPMNH); “三崎” [= Kanagawa-ken, Miura-shi, Misaki (approximate coordinates: 35°08'31.5"N, 139°36'55.1"E)], “21-18/4/1911” [= 18-21.iv.1911], “Matsumra” (collected by Shonen Matsumura and/or deposited in Matsumura’s collection) (1 ♂ 1 ♀, ELHU) (Fig. 38 ); Kanagawa-ken, Oiso-machi, Terasaka, Nagayama, 9.vi.1991, leg. M. Enju (1 ♂, TUA); Kanagawa Pref., Ashigarashimo-gun, Manazuru-hantou, 11.xi.2000, leg. S. Nagashima (1 ♂, TUA). The type locality of Stephanitis (Stephanitis) tabidula is “Kanagawa” [= Honshu, Kanagawa-ken (a prefecture) or Kanagawa-ku (a ward) of Yokohama-shi (a city) of Kanagawa-ken] and the 146 adult individuals recorded above match the original description of the species ( Horváth 1912). The author identified S. (S.) tabidula based on the 146 adults in the present study. Paratypes of S. (S.) fasciicarina Takeya, 1931 collected at type locality (40 ♂ 45 ♀♀, ELKU; 1 ♀, KUM) (Fig. 37A ), JAPAN: Kyushu: "Chikuzen Akama" [= Fukuoka-ken, Munakata-shi, Akama (approximate coordinates: 33°48'26.1"N, 130°35'31.2"E)], 31.v.1931, leg. C. Takeya. The male holotype of S. (S.) fasciicarina is currently missing ( Souma 2021b). However, the labels shown in Fig. 37A were created after the original description (T. Mita, pers. comm. 2021). As mentioned in the section below, some of the non-types were labelled as “paratype”. Therefore, since labelling errors are suspected to have occurred for some of the type specimens, one of the 40 male paratypes collected at the type locality recorded above may correspond to the holotype. Paratypes of S. (S.) fasciicarina (59 ♂♂ 46 ♀♀, ELKU), JAPAN: Honshu: "Nagato Shimonoseki" [= Yamaguchi-ken, Shimonoseki-shi (approximate coordinates: 33°57'09.1"N, 130°55'16.8"E)], 22.viii.1930, leg. K. Yasumatsu (8 ♂♂ 5 ♀♀). Kyushu: “Fukuoka” [= Fukuoka-ken (a prefecture) or Fukuoka-shi (a city) of Fukuoka-ken (approximate coordinates: 33°35'06.7"N, 130°22'43.4"E)], 14.v.1930, leg. C. Takeya (7 ♂♂ 1 ♀♀); as above but 21.v.1930 (6 ♂♂ 7 ♀♀); as above, but 22.v.1930 (21 ♂♂ 14 ♀♀); as above, but 30.vi.1930 (1 ♂ 1 ♀); as above but 7.vii.1930 (16 ♂♂ 18 ♀♀). Although the labels of the 105 specimens were created after (T. Mita, pers. comm. 2021), their locality data match that of the original description of S. (S.) fasciicarina ( Takeya 1931). Non-types collected at type locality of S. (S.) fasciicarina (4 ♀♀, TUA), JAPAN: Kyushu: Fukuoka-ken, Munakata-shi, Akama (approximate coordinates: 33°48'26.1"N, 130°35'31.2"E), 26.ix.2021, leg. J. Souma. Non-types collected at type locality of S. (S.) kyushuana Drake, 1948 (1 ♀, TUA), JAPAN: Kyushu: Fukuoka-ken, Kitakyushu-shi, Moji-ku, Motokiyotaki (approximate coordinates: 33°56'18.9"N, 130°57'42.7"E), 16.ix.2022, leg. J. Souma. Non-types (518 ♂♂ 653 ♀♀ 60 nymphs), JAPAN: Honshu: Miyagi-ken, Oshika-gun, Onagawa-cho, Kirigasaki, 20.ix.2020, leg. H. Konno (2 ♂♂ 3 ♀♀, TUA); Miyagi-ken, Miyagi-gun, Rifu-cho, Akanuma, Tanbazawa, 22.ix.2020, leg. H. Konno (1 ♀, TUA); Ibaraki, Chôshi, 15.vi.1981, leg. M. Miyazaki (13 ♂♂ 24 ♀♀ 1 nymph, NIAES); Tochigi Pref., Uchino, Watarase-yusuichi, 29.v.2001, leg. S. Maehara (2 ♂♂ 1 ♀, TUA); as above but 5.xi.2012 (1 ♂, TUA); Tochigi, Simotsuke City, Motoyoshida, 12.x.2013, leg. S. Maehara (2 ♀♀, TUA); as above, but 28.xi.2016 (1 ♂ 2 ♀♀); as above but 12.xi.2019 (1 ♂ 2 ♀♀, TUA); as above, but 15.x.2020 (4 ♂♂ 1 ♀, TUA); Saitama Pref., Nihongi Pass, 3.vii.1984, leg. M. Hayashi et al. (1 ♂, TUA); Saitama-ken, Iruma-gun, Moroyama-machi, Takinoiri, Shobo-goya, 28.v.2021, leg. J. Souma (2 ♂♂ 4 ♀♀, TUA); Saitama-ken, Iruma-gun, Moroyama-machi, Takinoiri, Kanikusa-hashi, 28.v.2021, leg. J. Souma (8 ♂♂ 9 ♀♀, TUA); Tokyo, 22.viii.1949, leg. R. Takahashi (1 ♂ 1 ♀, ELKU); Chiba, Tateyama, Susaki, 11.x.2001, leg. M. Tomokuni (5 ♂♂ 5 ♀♀, NSMT); Tokyo, Kiyosumi-koen, 24.vi.1972, leg. H. Hasegawa (11 ♂♂ 22 ♀♀, NIAES); Niigata-ken, Kitakanbara-gu, Seiro-machi, Mano, 14-15.vi.2013, leg. K. Nakano (1 ♂, TUA); Niigata-ken, Murakami-shi, Kashio, 13.vi.2015, leg. K. Nakano (1 ♂ 4 ♀♀, TUA); as above but 1.x.2016, leg. K. Nakano (1 ♂ 4 ♀♀, TUA); Niigata-ken, Niigata-shi, Nishi-ku, Igarashi-2no-cho, 19.vii.2015, leg. K. Nakano (1 ♀, TUA); Niigata-ken, Kariwa-gun, Kariwa-mura, Takiya, 13.vi.2021, leg. G. Mashima (1 ♂ 1 ♀, TUA); Niigata-ken, Kashiwazaki-shi, Miyagawa, 13.vi.2021, leg. G. Mashima (2 ♂♂, TUA); Ishikawa, Sekidoyama, 2.viii.1960, leg. T. Hidaka (1 ♀, ELKU); Fukui Pref., Otomi, 12.vii.1981, leg. O. Kishimoto (1 ♀, KUM); Fukui Pref., Mt. Aoba, 20.ix.1981, leg. O. Kishimoto (1 ♀, KUM); Mino, Utsumi, 21.x.1952, leg. Yasumatu (1 ♂ 1 ♀, ELKU); Izu, Kuren, 16.x.1952, leg. Yasumatsu (3 ♀♀, ELKU); Izu, Suzaki, 30.ix.1980, leg. M. Tomokuni (7 ♂♂ 7 ♀♀, NSMT); Izu, Ohno-yama, nr. Ohsawa, 1.x.1980, leg. M. Tomokuni (3 ♀♀, NSMT); Shizuoka-ken, Hamamatsu-shi, Nishi-ku, Kamigaya-cho, 16.vi.2017, leg. J. Souma (1 ♀, TUA); Aichi-ken, Kitashitara-gun, Shitara-cho, Taguchi, 17.vi.2017, leg. J. Souma (1 ♀, TUA); Mie Pref., Oosugidani, 11.viii.1970, leg. M. Tomokuni (1 ♂, NSMT); Mie Pref., Ise-shi, Shimogu, 1.xi.2000, leg. T. Ishikawa (1 ♀, TUA); Kyoto Pref., Kurama-Kibune, 12.vii.1970, leg. M. Tomokuni (1 ♀, NSMT); Osaka Pref., Higashi-Sumiyoshi, Nagai, 2.vi.1985, leg. T. Kishimoto (1 ♀, TUA); Kobe, 29.v.1941, leg. Kurosa (1 ♂, ELKU); Tajima, Sekinomiya, 8.vii.1952, leg. R. Morimoto (7 ♂♂ 6 ♀♀, ELKU); Hyogo-ken, Kobe-shi, Suma-ku, Higashisuma, 8.x.2022, leg. J. Souma (3 ♂♂ 3 ♀♀, TUA); Yamato, Nara, 14.x.1951, leg. C. Takeya (3 ♂♂ 3 ♀♀, ELKU); as above but 16.x.1951 (23 ♂♂ 30 ♀♀, ELKU; 1 ♂ 1 ♀, KUM); Prov. Kii, Wakayama, 15.x.1951, leg. S. Goto (2 ♀♀, ELKU); Wakayama, Nachi-Irokawa, 2.ix.1998, leg. leg. S. Gotoh (1 ♂, NSMT); Okayama-ken, Okayama-shi, Kita-ku, Heidan, 20.v.2022, leg. J. Souma (1 ♀, TUA); Yamaguchi-ken, Yamaguchi-shi, Atou, Shinome, 25.x.2008, leg. M. Hayashi (1 ♀, TUA); Yamaguchi-ken, Shimonoseki-shi, Hon-machi, 24.ix.2022, leg. J. Souma (6 ♂♂ 3 ♀♀, TUA). Tashiro Island: Shikishima, near Manga Island , 10.xi.2019, leg. H. Konno (1 ♂ 1 ♀, TUA). Sado Island : Kitaikari, 26.viii.2021, leg. J. Souma (2 ♂♂ 1 ♀, TUA); Kubota, 27.viii.2021, leg. J. Souma (2 ♀♀, TUA); Ishida, 27.viii.2021, leg. J. Souma (2 ♂♂ 8 ♀♀, TUA); Anyoji, 28.viii.2021, leg. J. Souma (1 ♀, TUA); Kujikawachi, 29.viii.2021, leg. J. Souma (1 ♀, TUA); Kawasaki, 29.viii.2021, leg. J. Souma (3 ♂♂ 4 ♀♀, TUA). Awa Island : Uramura, Uchiura, 3.x.2021, leg. J. Souma (9 ♂♂ 7 ♀♀, TUA). Izu Islands (northern part): Izu-Oshima Island: Kasamatsu, 4.x.2002, leg. M. Tomokuni (1 ♂♂ 2 ♀♀, NSMT). Awaji Island : Fukura, 29.ix.1972, leg. M. Tomokuni & M. Sakai (1 ♀, NSMT); Mt. Mikuma , 30.ix.1972, leg. M. Tomokuni & M. Sakai (1 ♀, NSMT). Oki Islands : Dogo Island : Saigo-cho, Mt. Daimanji , 12.ix.1984, leg. M. Tomokuni (3 ♀♀, NSMT). Shikoku: Tokushima-ken, Tokushima-shi, Bizan-cho, Mosukegahara, 2.x.2022, leg. J. Souma (1 ♂ 1 ♀, TUA); "Sanuki Wada-mura" [= Kagawa-ken, Kannonji-shi, Toyohama-cho, Wada], 10.vi.1930, leg. M. Hanada (2 ♂ 3 ♀♀, ELKU; 1 ♀, KUM); Tosa, Yamakita-mura, 10.xi.1951, leg. K. Yasumatsu (3 ♂♂ 2 ♀, ELKU); Ehime P., Matsuyama C., 25.x.1972, leg. H. Hasegawa (11 ♂♂ 7 ♀♀ 4 nymphs, NIAES). Kyushu: as paratype collected at type locality (1 nymph, ELKU); Fukuoka, Hirao, 22.x.1931, leg. Fujino & Hashimoto (1 ♀, ELKU); as above but 16.x.1932, leg. T. Shrozu (1 ♂ 1 ♀, ELKU); as above but 3.vi.1951, leg. Matsuda (1 ♀, ELKU); as above but 21.vi.1957, leg. Y. Miyatake (1 ♀, KUM); Chikuzen, Inunakiyama, 25.x.1931, leg. K. Yasumatsu (1 ♀, KUM); as above, but 15.vi.1969, leg. S. Miyamoto (1 ♀, KUM); Chikuzen, Tsutsugatake, 6.xii.1931, leg. C. Takeya (1 ♂ 14 ♀♀, ELKU); Tikuzen, Kyûdai-Kasuya-Ensyûrin, 20-22.vi.1944, leg. S. Ito (1 ♀, ELKU); Buzen, Hikosan, 12.vii.1948, leg. S. Miyamoto (1 ♀, KUM); as above, but 5.viii.1951, leg. C. Takeya (5 ♂♂ 9 ♀♀, ELKU); as above, but 6.viii.1951, leg. C. Takeya (8 ♂♂ 15 ♀♀, ELKU); as above but 15.vii.1958, leg. Y. Miyatake (2 ♂♂ 1 ♀, KUM); as above, but 6.vi.1959, leg. Y. Miyatake (1 ♂, KUM); as above but 7.vi.1959, leg. Y. Miyatake (1 ♀, KUM); as above, but 14.vi.1959 (3 ♀♀, KUM); as above, but 14.vi.1959, leg. K. Yasumatsu (1 ♀, ELKU); Chikugo, Korasan, 25.vii.1951, leg. C. Takeya (3 ♂♂ 3 ♀♀, ELKU); as above, but 7.viii.1951, leg. S. Miyamoto (1 ♂ 1 ♀, ELKU); as above, but 1.ix.1951, leg. C. Takeya (1 ♂ 3 ♀♀, ELKU); as above but 26.v.1953 (4 ♂♂ 5 ♀♀, ELKU); Fukuoka, Fukuoka, Korasan, 25.vii.1951, leg. S. Miyamoto (1 ♂ 3 ♀♀, KUM); as above but 7.viii.1951 (2 ♀♀, KUM); as above, but 11.vi.1961 (3 ♀, KUM); Chikuzen, Fukuoka, 27.vii.1951, leg. C. Takeya (6 ♂♂ 7 ♀♀, ELKU); Chikuzen, Narutake-yama, 19.viii.1951, leg. Y. Hirashima (1 ♀, ELKU); as above but 8.vi.1961, leg. C. Takeya (1 ♀, ELKU); Fukuoka, Mt. Kumado , 23.viii.1952, leg. S. Miyamoto (4 ♂♂ 3 ♀♀, ELKU; 9 ♂♂ 6 ♀♀ 5 nymphs, KUM); Aburayama, 28.viii.1952, leg. C. Takeya (1 ♀, ELKU); as above, but 5.v.1961, leg. S. Miyamoto (5 ♂♂ 9 ♀♀ 17 nymphs, KUM); as above, but 11.v.1961, leg. S. Miyamoto (16 ♂♂ 11 ♀♀, KUM); as above, but 4.vi.1961, leg. S. Miyamoto (1 ♀, KUM); Fukuoka, Magarifuchi, 23.ix.1952, leg. C. Takeya (6 ♂♂ 4 ♀♀, ELKU); as above, but 16.vii.1961, leg. S. Miyamoto (3 ♂♂ 4 ♀♀, KUM); Chikuzen, Mt. Kosho , 14.vi.1953, leg. C. Takeya (1 ♀, ELKU); Chikugo, Himeharu, 3.vi.1954, leg. Gyotoku (3 ♂♂ 3 ♀♀, ELKU); Fukuoka, Sasayama, ix.1954, leg. S. Miyamoto (1 ♂ 2 ♀♀, KUM); Fukuoka, Gokoku Shrine, 5.ix.1959, leg. S. Miyamoto (39 ♂♂ 36 ♀♀, KUM); as above, but 24.vii.1961 (9 ♂♂ 8 ♀♀, KUM); Fukuoka, Kashii, 6.ix.1959, leg. S. Miyamoto (1 ♀, KUM); as above, but 2.xi.1961 (1 ♂, KUM); Fukuoka, Nogochi, 16.vi.1961, leg. S. Miyamoto (13 ♂♂ 9 ♀♀ 3 nymphs, KUM); Fukuoka, Tachibanayama, 23.vii.1961, leg. S. Miyamoto (2 ♂♂ 7 ♀♀, KUM); Fukuoka, 24.vii.1961, leg. S. Miyamoto (2 ♂♂ 6 ♀♀, KUM); Fukuoka, Atagoyama, 4.viii.1961, leg. S. Miyamoto (1 ♂ 1 ♀, KUM); Fukuoka, Hakozaki, Kyushu University, 12.viii.1961, leg. S. Miyamoto (134 ♂♂ 124 ♀♀ 14 nymphs, KUM); Fukuoka, Kanetake, 13.ix.1969, leg. S. Miyamoto (5 ♂♂ 12 nymphs, KUM); Fukuoka, Kamado Shrine, 27.ix.1977, leg. S. Miyamoto (2 ♂♂, KUM); as above, but 5.xi.1977 (1 ♀, KUM); Fukuoka, Tenmangu, 6.x.1977, leg. S. Miyamoto (1 ♀, KUM); Fukuoka-ken, Munakata-shi, Yamada, 28.viii.2017, leg. J. Souma (1 ♀, TUA); Fukuoka-ken, Fukuoka-shi, Nishi-ku, Motooka, Kyushu University, 23.v.2020, leg. J. Souma (2 ♂♂ 1 ♀, ELKU); as above, but 21.ix.2020 (3 ♂♂ 1 ♀, ELKU); Fukuoka-ken, Fukuoka-shi, Nishi-ku, Kusaba, 24.v.2020, leg. J. Souma (1 ♂, ELKU); Fukuoka-ken, Itoshima-shi, Shimanokita, 24.v.2020, leg. J. Souma (14 ♂♂ 28 ♀♀, ELKU); Fukuoka-ken, Fukuoka-shi, Sawara-ku, Itaya, Mt. Sefuri , 30.v.2020, leg. S. Yagi (1 ♀, ELKU); Fukuoka-ken, Dazaifu-shi, Kitadani, Mt. Homan , 4.vi.2020, leg. J. Souma (7 ♂♂ 11 ♀♀, ELKU; 1 ♂ 4 ♀♀, TUA); Fukuoka-ken, Kasuya-gun, Sasaguri-machi, Tsubakuro, 5.vi.2020, leg. J. Souma (2 ♂♂ 1 ♀, TUA); Fukuoka-ken, Itoshima-shi, Nijofukui, Nijo Forest Park, 20.vi.2020, leg. S. Chuman (4 ♂♂ 2 ♀♀, TUA); Fukuoka-ken, Tagawa-gun, Soeda-machi, Mt. Hikosan , 21.vi.2020, leg. J. Souma (3 ♀♀, TUA); as above but 20.vi.2020, leg. S. Yagi (1 ♀, ELKU); Fukuoka-ken, Itoshima-shi, Tomari, 29.v.2022, leg. J. Souma (1 ♀, ELKU); as above, but 10.vii.2022, leg. J. Souma (1 ♂, TUA); Saga Pref., Fujitsu-gun, Tara-cho, Oura, Ushioro, alt. 100 m, 12.vii.2021, leg. M. Nishida (1 ♂, ELKU); Hizen , Kunimiyama , 16.vi.1950, leg. T. Shrozu (1 ♂ 2 ♀♀, ELKU); Higo , Yatsushiro , 8.x.1953, leg. C. Takeya (1 ♀, ELKU); Kumamoto , Yuyama , Mt. Ichifusa , 5.viii.1985, leg. M. Miyazaki (3 ♀♀, NIAES); as above, but 6.viii.1985 (2 ♂♂ 4 ♀♀, NIAES); Bungo , Tsukumi , 29.vii.1951, leg. R. Matsuda (2 ♂♂ 1 ♀, ELKU); Bungo , Hachiya , 11.x.1951, leg. S. Nakao (1 ♀, KUM); Ôita Pref., Saiki-shi , Ume , Minamitabaru , Takadoya-jinja , 26.v.2019, leg. R. Ito (1 ♂, TUA); Oita Pref., Saeki-shi , Ume , Kitagawa dam, 25-26.v.2019, leg. N. Tsuji & S. Imada (1 ♀, TUA); Hyuga , Takaoka , 25.viii.1952, leg. A. Ema (6 ♂♂ 10 ♀♀, ELKU); Hyuga , Mt. Kirishima , 12.ix.1952, leg. Nakao & Ogata (1 ♂, ELKU); Miyazaki Pref., Hinokage-chô, Mt. Tansuke-dake , 1.vi.2019, leg. R. Ito (1 ♀, TUA); Miyazaki-ken , Higashimorokata-gun , Aya-cho , Takeno , Omoridake For. Rd. , 8.x.2021, leg. T. Saeki (1 ♂, ELKU); Kagoshima Pref., Mianmiôsumi-chô, Sata , Hetsuka , 30.v.2020, leg. R. Ito (1 ♀, TUA). Nokonoshima Island : 27.vi.1987, leg. S. Miyamoto (1 ♀, KUM). Tsushima Island : Izuhara-Sasutôge, 7.vi.1941, leg. T. Shirôzu (1 ♀, ELKU); Izuhara , 31.x.1962, leg. S. Miyamoto (1 ♀, ELKU; 1 ♂, KUM); Mt. Ariake , 12.vii.1968, leg. S. Miyamoto & A. Nakanishi (1 ♀, KUM); Observatory , 28.viii.1988, leg. S.+ K. M. (1 ♀, KUM); Kuda , 29.viii.1988, leg. S. + K. M. (1 ♂ 1 ♀, KUM); Kamiagata-cho , Sago , 6.ix.2017, leg. J. Souma (5 ♂♂ 4 ♀♀, TUA); as above but 8.ix.2017 (3 ♀♀, TUA); Izuhara-cho , Tsutsu , Mount Tatera , 7.ix.2017, leg. J. Souma (1 ♂, TUA); Kamiagata-machi , near Mt. Konoki-yama , 8.ix.2017, leg. T. Ishikawa (11 ♂♂ 8 ♀♀ 3 nymphs, TUA); Mitsushima-machi , Kechi , 8.ix.2017, leg. J. Souma (12 ♂♂ 17 ♀♀, TUA). Iki Island : Gônoura-chô, Katabaruhure , Takenotsuji , 16-18.vii.2016, leg. R. Ito (1 ♂, TUA). Amakusa Islands : Shimoshima Island : “Kakuyama” [= Miyajidake-machi, Mt. Kakuyama], 18.ix.1931, leg. H. Hori (1 ♂ 1 ♀, ELKU). Goto Islands : Fukue Island : Miiraku-Kahara , 4.viii.1933, leg. T. Shirozu (2 ♀, KUM). Ryukyu Islands (northern part): Yakushima Island : Kosugidani , 23.viii.1952, leg. C. Takeya & Y. Hirashima (2 ♂♂ 5 ♀♀, ELKU); Koseda , 21.ix.2013, leg. N. Tsuji (1 ♀, ELKU). Eight specimens from "Sanuki Wada-mura" and “Kakuyama” were labelled as “paratype” of S. (S.) fasciicarina . However , their labels were created subsequently (T. Mita, pers. comm. 2021) and the locality data of these eight individuals do not match those of the original description of S. (S.) fasciicarina ( Takeya 1931). Therefore , the eight specimens do not seem to have paratype status. Four specimens from “Tachibanayama” were identified as " S. (N.) aperta possessing lateral carina" in a previous study ( Takeya 1963) .

Diagnosis.

Stephanitis (Stephanitis) tabidula is recognised amongst other species of Stephanitis by a combination of the following characters: head, calli, pronotal disc, marking on hemelytra and ventral surface in various shades of brown (Figs 8C, D , 10C-E , 12C, D , 14C-E , 16C, D , 18C, D , 20B , 22B , 24B ); body in male 2.1 times (in female 2.0 times) as long as maximum width across hemelytra (Figs 3C, D , 5C, D ); rostrum not reaching metasternum; pronotum unicarinate or tricarinate (Fig. 26C, D ); hood pale, shorter than median carina of pronotum, as wide as or wider than vertex at widest part, not or incompletely covering eye, as high as median carina of pronotum at highest part, with posterior margin not extending to middle of pronotal disc; median carina of pronotum with 2 rows of areolae at highest part; pronotal disc opaque; paranotum more erect, slightly narrowed posteriorly, with 3 rows of areolae at widest part, anterolateral angle protruding anteriad, outer margin gently curved inwards at posterolateral angle, maximum height longer than height of eye (Fig. 28D ); apices of hemelytra close to each other in rest; costal area with 3-5 rows of areolae at widest part; subcostal area in male with 2 rows (in female with 2-3 rows) of areolae at widest part; discoidal area with 3-4 rows of areolae at widest part; sutural area with 3-4 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. 30E, F ); and paramere stout, strongly curved inwards at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 32E, F ).

Intraspecific variation.

Stephanitis (Stephanitis) fasciicarina (including its junior synonym S. (S.) kyushuana ) was previously distinguished from S. (S.) tabidula by the following characters ( Takeya 1931; Drake 1948): body larger (smaller in S. (S.) tabidula ); antennal segment III less than 2.0 times as long as antennal segment IV (2.0 times in S. (S.) tabidula ); anterior margin of hood extending beyond clypeus (slightly extending in S. (S.) tabidula ); markings on hemelytron indistinct (distinct in S. (S.) tabidula ); and costal area with 4 rows of areolae at widest part (3 rows in S. (S.) tabidula ). The author’s examination of 1,496 adults from various localities in Japan, including the type localities of S. (S.) tabidula and S. (S.) fasciicarina , together with the photographs of the holotype of S. (S.) kyushuana ( United States National Museum of Natural History 2021) revealed that no other significant differences in the external morphology could be found between the two forms and these characters have considerable intraspecific variability, with transitional individuals between the extreme forms (Figs 3C, D , 5C, D , 8C, D , 10C-E , 12C, D , 14C-E , 16C, D , 18C, D , 20B , 22B , 24B , 26C, D , 30E, F , 32E, F , 32D, E , 37A , 38 , 42C-F ). In addition, the partial COI gene pairwise sequence distances between the two forms from the vicinity of the type localities of S. (S.) tabidula and S. (S.) fasciicarina (Suppl. material 2) were only 0.001321-0.003973 (Suppl. material 3). Furthermore, the Bayesian tree (Fig. 1 ), based on the partial COI gene, shows that the two forms are monophyletic with a high posterior probability. In conclusion, S. (S.) tabidula and S. (S.) fasciicarina are currently impossible to distinguish, based on morphological and molecular data either at the species or subspecies level. Therefore, the following new subjective synonymy is proposed: Stephanitis (Stephanitis) tabidula Horváth, 1912 = Stephanitis (Stephanitis) fasciicarina Takeya, 1931 syn. nov.

Remarks.

Stephanitis (Stephanitis) tabidula and S. (Norba) aperta are distributed in the same regions and feed on the same host plants ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; present study), but the former can be readily distinguished by the tricarinate pronotum (unicarinate in S. (N.) aperta ) (Figs 7B , 8C, D , 9B , 10C, E , 25B , 26C, D ). However, individuals of S. (S.) tabidula rarely have an unicarinate pronotum (Fig. 10D ). Such specimens, although they strongly resemble S. (N.) aperta in general habitus, can still be easily differentiated from the latter species by the following characteristics: paranotum more erect (less erect in S. (N.) aperta ), slightly narrowed posteriorly (narrowed in S. (N.) aperta ), with maximum height longer than height of eye (shorter in S. (N.) aperta ) (Figs 11B , 13B , 14D , 27B , 28D ); and apex of paramere strongly curved inwards (slightly curved in S. (N.) aperta ) (Figs 31B , 32E ).

Teratological form.

As mentioned in the above section, Stephanitis (Stephanitis) tabidula rarely has a unicarinate pronotum (Figs 10D , 14D ).

Distribution.

Japan (Honshu; Izu Islands (northern part): Izu-Oshima Island; Tashiro Island; Sado Island; Awa Island; Awaji Island; Oki Islands: Dogo Island; Shikoku; Kyushu; Nokonoshima Island; Tsushima Island; Iki Island; Amakusa Islands: Shimoshima Island; Goto Islands: Fukue Island; Ryukyu Islands (northern part): Yakushima Island) (Fig. 48 ); southern Korea ( Horváth 1912; Takeya 1931, 1963; Takara and Hidaka 1960; Miyamoto 1976; Tomokuni 1985; Yasunaga et al. 1993; Tomokuni and Hayashi 2006; Yamada and Tomokuni 2012; Yamada and Ishikawa 2016; Ahn et al. 2018; present study). Judging from the description and illustration ( Jing 1981), the Chinese population differs from the Japanese population in the structure of the pronotum and seems to correspond to another species. According to the present author’s examination, the specimens previously recorded from Okinawa Island (the central part of the Ryukyu Islands) ( Takara and Hidaka 1960) correspond to Stephanitis (Norba) mendica . Judging from the photographs and specimens, the previous records from the southern part of the Izu Islands ( Takeya 1963; Tomokuni and Ishikawa 2002; Yamada and Tomokuni 2012) correspond to S. (S.) tomokunii sp. nov., described below. In Japan, S. (S.) tabidula inhabits the laurilignosa in a temperate climate of Japan proper (Honshu, Shikoku and Kyushu) and its surrounding islands and the northern part of Izu and Ryukyu Islands, which is located in the Palaearctic Region.

Host plants.

Cinnamomum camphora , “Kusunoki” ( Lauraceae ) (Fig. 44D ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; present study); C. yabunikkei , “Yabunikkei” ( Yasunaga et al. 1993; present study); Laurus nobilis L., “Gekkeiju” ( Lauraceae ) ( Takeya 1963; Yamada and Tomokuni 2012); Neolitsea sericea , “Shirodamo” (Fig. 44C ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; Okochi 2019; present study); Machilus japonica Siebold et Zucc. ex Blume, “Aogashi” or “Hosobatabu” ( Lauraceae ) ( Takeya 1963; Yamada and Tomokuni 2012; Okochi 2019; present study); M. thunbergii , “Tabunoki” (Fig. 44B ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; Okochi 2019; present study). Stephanitis (Stephanitis) tabidula feeds only on lauraceous trees and is oligophagous. This lace bug sometimes occurs on plantings of C. camphora and M. thunbergii in its distribution range (present study). This tingid species has been known to be a pest of C. camphora (Japanese Society of Applied Entomology and Zoology 1980, 2006) and occasionally also causes harm on M. thunbergii .

Biology.

Stephanitis (Stephanitis) tabidula feeds on the abaxial surface of leaves of the five host plants in Japan (present study). In Japan, adults were collected in almost all seasons ( Takeya 1931; Miyamoto 1976; Tomokuni 1981, 1985; Yasunaga et al. 1993; Tomokuni and Hayashi 2006; Yano et al. 2013; Okochi 2019; present study); nymphs were collected from May to July and in September and October (present study); the overwintering stage is the adult (present study).

Figures

Figure 1. Bayesian tree using the cytochrome c oxidase subunit I (COI) gene. Numbers associated with branches indicate posterior probabilities. Each sample ID is followed by a species name and collection locality.

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 7. Male pronota of five Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov.; E, F. S. (N.) hiurai from Amami-Oshima (E) and Kikai (F) Islands, central part of Ryukyu Islands. Scale bars: 0.2 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 9. Female pronota of five Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov.; E, F. S. (N.) hiurai from Amami-Oshima (E) and Kikai (F) Islands, central part of Ryukyu Islands. 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 11. Male pronota of five Stephanitis species from Japan, dorsolateral view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov.; E, F. S. (N.) hiurai from Amami-Oshima (E) and Kikai (F) Islands, central part of Ryukyu Islands. 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 13. Female pronota of five Stephanitis species from Japan, dorsolateral view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov.; E, F. S. (N.) hiurai from Amami-Oshima (E) and Kikai (F) Islands, central part of Ryukyu Islands. 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 25. Line drawings of the pronota of six Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov.; E. S. (N.) hiurai; F. S. (N.) ishikawai sp. nov. Abbreviations: ca, calli; co, collar; ho, hood; lc, lateral carina; mc, median carina; pa, paranotum; pd, pronotal disc; pp, posterior process. 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 27. Line drawings of the pronota of four Stephanitis species from Japan, lateral view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D. S. (N.) hayashii sp. nov. Abbreviations: bu, buccula; ca, costal area; da, discoidal area; hl, hypocostal lamina; ho, hood; lc, lateral carina; mc, median carina; pa, paranotum. 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 31. Line drawings of the paramere of five Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua; B. S. (Norba) aperta; C. S. (N.) exigua; D, E. S. (N.) hayashii sp. nov. from Kakeroma (D) and Aguni (E) Islands, central part of Ryukyu Islands; F. S. (N.) hiurai. Scale bars: 0.1 mm. Arrows indicate important morphological differences.

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 38. Non-types of Stephanitis (Stephanitis) tabidula collected at type locality deposited in Matsumura's collection of ELHU and their 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.