Stephanitis (Norba) aperta 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 (Norba) aperta Horvath, 1912

[Japanese name: Tabu-gunbai] Figs 2B , 4B , 6A , 7B , 9B , 11B , 13B , 15B , 17B , 19B , 21B , 23B , 25B , 27B , 29B , 31B , 34A, B , 40D-G

Stephanitis (Norba) aperta Horváth, 1912: 335. Syntype(s): Japan: Sakuna [= Honshu, Chiba-ken, Sakuna of former Toyofusa-mura in early 20th Century (current Tateyama-shi, Sakuna)]; HNHM. “Sakuna” was considered to be a misspelling of “Satsuma” ( Takeya 1931: 77).

Stephanitis (Norba) vitrea Takeya, 1931: 74. Holotype (Fig. 34A ): Japan: Yakushima Is., Onoaida-Ambo [= Ryukyu Islands, Yakushima Island, between Onoaida and Anbo]; ELKU. Synonymised with Stephanitis (Norba) exigua Horváth, 1912 by Takeya (1953: 168), with Stephanitis (Norba) aperta Horváth, 1912 by Takeya (1963: 38).

Stephanitis exigua Horváth, 1912: Takeya (1953: 168) (distribution: part). Misidentification ( Takeya 1963: 38).

References.

Takeya (1931: 75) (distribution: part); Drake (1948: 54) (checklist: Stephanitis ); Takeya (1951b: 13) (checklist: Japan); Drake and Maa (1953: 100) (checklist: Stephanitis ); Takeya (1963: 38) (distribution: part); Miyamoto (1964a: 274) (distribution: part); Drake and Ruhoff (1965: 366) (catalog); Lee (1969: 246) (male genitalia); Jing (1981: 349) (monograph); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Yasunaga et al. (1993: 178) (monograph: part); Péricart and Golub (1996: 58) (catalogue: Palaearctic); Tomokuni et al. (2000: 38) (distribution); Tomokuni (2005: 400) (distribution); Tomokuni (2006a: 350) (distribution); Yamada and Tomokuni (2012: 205) (monograph: part); Yiu and Yip (2012: 83) (distribution); Yano et al. (2013: 25) (distribution); Tomokuni (2014: 362) (distribution); Nozaki et al. (2015: 9) (distribution); Yamada and Ishikawa (2016: 433) (checklist: Japan); Maehara (2017: 146) (distribution); Okochi (2019: 2) (distribution); Souma (2021b: 31) (distribution).

Material examined.

Holotype of Stephanitis (Norba) vitrea Takeya, 1931 (1 ♀, ELKU) (Fig. 34A ), JAPAN: Ryukyu Islands (northern part): "Yakushima Onoaida-Ambô” [= Yakushima Island, between Onoaida and Anbo (approximate coordinates: 30°16'08.3"N, 130°36'44.2"E)], 3.viii.1929, leg. H. Hori. The labels shown in Fig. 34A were created after (T. Mita, pers. comm. 2021), but the condition of the specimen matched the photograph of the original description ( Takeya 1931). Therefore , the female individual seems to correspond to the holotype of S. (N.) vitrea . Suspected syntype of S. (Stephanitis) propinqua Horváth, 1912 (1 ♀, ELHU) (Fig. 34B ), JAPAN: Kyushu : “カゴシマ” [= Kagoshima-ken (approximate coordinates: 31°35'59.5"N, 130°32'59.6"E)], “7/10” [= 10.vii]. This specimen was labelled as " Stephanitis gratiosa Horv." (apparently an unpublished name) and “カゴシマグンバイ” (unknown Japanese name). According to the original description ( Horváth 1912), the syntype (s) of S. (S.) propinqua was (were) deposited in ELHU and this female individual is the only specimen in Matsumura’s collection of ELHU matching the label data of syntype (s) of S. (S.) propinqua . However , the morphological characteristics of the specimen match the original description of S. (N.) aperta ( Horváth 1912), but not that of S. (S.) propinqua . Conversely , the general habitus of S. (S.) propinqua recorded from Korea by previous studies ( Takeya 1932, 1963) differs from that of all the East Asian species of Stephanitis (present study). Therefore , if there is no syntype of S. (S.) propinqua in other institutions, then S. (S.) propinqua should be synonymised with S. (N.) aperta and a new name should be given to " S. (S.) propinqua " distributed in Korea. Non-types (534 ♂♂ 777 ♀♀ 16 nymphs), JAPAN: Honshu: Ibaraki-ken, Higashiibaraki-gun, Oarai-machi, Isohama-cho, 12.ix.2020, leg. J. Souma (7 ♂♂ 13 ♀♀ 3 nymphs, TUA); Tochigi Pref., Uchino, Watarase-yusuichi, 26.xi.2016, leg. S. Maehara (4 ♂♂ 2 ♀♀, TUA); as above but 2.x.2020 (4 ♂♂ 3 ♀♀, TUA); Chiba-ken, Tateyama-shi, Ohto (approximate coordinates: 34°58'19.8"N, 139°52'33.9"E), 22.v.2021, leg. J. Souma (1 ♀, TUA); Chiba-ken, Tateyama-shi, Shimosanagura, 22.v.2021, leg. J. Souma (1 ♂ 2 ♀♀, TUA); Chiba-ken, Tateyama-shi, Tateyama, 22.v.2021, leg. J. Souma (2 ♂♂ 6 ♀♀, TUA); as above but 23.v.2021 (4 ♀♀, ELKU; 2 ♂♂ 2 ♀♀ 1 nymph, TUA); Chiba-ken, Tateyama-shi, Sunozaki, 23.v.2021, leg. J. Souma (11 ♂♂ 32 ♀♀, TUA); Chiba-ken, Tateyama-shi, Sunozaki Shrine, 23.v.2021, leg. J. Souma (1 ♂ 2 ♀♀, TUA); Chiba-ken, Tateyama-shi, Masaki, 23.v.2021, leg. J. Souma (7 ♂♂ 14 ♀♀, TUA); Chiba-ken, Tateyama-shi, Higashinagata, 24.v.2021, leg. J. Souma (1 ♂ 1 ♀, TUA); Chiba-ken, Tateyama-shi, Yamogi (approximate coordinates: 34°58'08.7"N, 139°53'34.4"E), 24.v.2021, leg. J. Souma (2 ♂♂ 4 ♀♀, TUA); Chiba-ken, Minamiboso-shi, Chikura-cho, Minamiasai, 24.v.2021, leg. J. Souma (1 ♀, TUA); Chiba-ken, Kamogawa-shi, Yomogi, 25.v.2021, leg. J. Souma (1 ♀, TUA); Tokyo Met., Imperial Palace, 5.vi.1996, leg. A. Saito (7 ♂♂ 7 ♀♀, NSMT); as above but 16.x.1996, leg. M. Tomokuni (8 ♂♂ 15 ♀♀, NSMT); 19.xi.1997, leg. M. Tomokuni (9 ♂♂ 6 ♀♀, NSMT); as above but 25.v.1998 (2 ♀♀, NSMT); as above but 28.v.1998, leg. M. Tomokuni (4 ♂♂ 3 ♀♀, NSMT); as above but 16.vii.2009, leg. M. Tomokuni (1 ♀, NSMT); as above but 28.v.2012, leg. M. Tomokuni (1 ♀, NSMT); as above but 23.vii.2012, leg. M. Tomokuni (1 ♂ 2 ♀♀, NSMT); as above but 10.ix.2012, leg. M. Tomokuni (1 ♀, NSMT); Tokyo-to, Minato-ku, Shiba-koen, 8.vii.2022, leg. J. Souma (4 ♂♂ 1 ♀, TUA); Kanagawa-ken, Sagamihara-shi, Minami-ku, Shimomizo, 15.xi.2021, leg. J. Souma (9 ♂♂ 8 ♀♀ 3 nymphs, TUA); Kanagawa-ken, Sagamihara-shi, Chuo-ku, Tanashioda, 19.v.2019, leg. J. Souma (2 ♀♀, TUA); Kanagawa-ken, Sagamihara-shi, Chuo-ku, Tana, 19.v.2019, leg. J. Souma (22 ♂♂ 16 ♀♀, TUA); as above but 17.xi.2021 (9 ♂♂ 11 ♀♀, TUA); Kanagawa-ken, Sagamihara-shi, Midori-ku, Oshima, 17.xi.2021, leg. J. Souma (1 ♂ 3 ♀♀ 1 nymph, TUA); Kanagawa-ken, Zama-shi, Iriyanishi, 1.i.2022, leg. J. Souma (1 ♂ 14 ♀♀, TUA); Kanagawa-ken, Ebina-shi, Kamiimaizumi, 1.i.2022, leg. J. Souma (1 ♂ 1 ♀, TUA); Kanagawa-ken, Atsugi-shi, Funako, 29.v.2017, leg. J. Souma (37 ♂♂ 41 ♀♀, TUA); as above but 30.v.2017 (32 ♂♂ 30 ♀♀, TUA); as above but 31.v.2017 (4 ♀♀, ELKU; 52 ♂♂ 55 ♀♀, TUA); as above but 2.vi.2017 (6 ♂♂ 16 ♀♀, TUA); as above but 5.vi.2017 (30 ♂♂ 17 ♀♀, TUA); Kanagawa-ken, Atsugi-shi, Nanasawa, 6.vi.2017, leg. J. Souma (1 ♀, TUA); Kanagawa-ken, Aiko-gun, Aikawa-machi, Nakatsu, 26.v.2021, leg. J. Souma (3 ♂♂ 11 ♀♀, TUA); as above but 15.xi.2021 (3 ♀♀, TUA); Kanagawa-ken, Yokohama-shi, Isogo-ku, Negishihachiman Shrine, 31.v.1999, leg. M. Takakuwa (1 ♀, KPMNH); Kanagawa-ken, Yokohama-shi, Kanazawa-ku, Noukendaimori, 15.vi.2017, leg. J. Souma (3 ♂♂, TUA); Kanagawa-ken, Yokosuka-shi, Kamoi, 27.vi.2017, leg. J. Souma (4 ♀♀, TUA); Kanagawa-ken, Chigasaki-shi, Yanagishima, 1.vi.2019, leg. J. Souma (10 ♂♂ 7 ♀♀, TUA); Kanagawa-ken, Hadano-shi, Tsurumaki, Mt. Azuma , 19.xi.2021, leg. J. Souma (2 ♂♂ 5 ♀♀, TUA); Kanagawa-ken, Hadano-shi, Soya, Mt. Kobo , 19.xi.2021, leg. J. Souma (4 ♀♀, TUA); Kanagawa-ken, Ashigarashimo-gun, Manazuru-machi, Manazuru, 23.v.2021, leg. J. Souma (11 ♂♂ 30 ♀♀, TUA); Niigata-ken, Niigata-shi, Nishi-ku, Igarashi-2no-cho, 19.vii.2015, leg. K. Nakano (2 ♀♀, TUA); as above but 16.vi.2021, leg. G. Mashima (12 ♂♂ 14 ♀♀, TUA); Niigata-ken, Niigata-shi, Chuo-ku, Sekiya, Nishikaigan Park, 22.x.2016, leg. K. Nakano (4 ♂♂ 10 ♀♀, TUA); Niigata-ken, Niigata-shi, Chuo-ku, Seigoro, Toyanogata Park, 30.x.2016, leg. K. Nakano (1 ♂ 4 ♀♀, TUA); Niigata-ken, Niigata-shi, Higashi-ku, Matsuzono, 21.vii.2019, leg. K. Nakano (2 ♂♂ 1 ♀, TUA); Niigata-ken, Niigata-shi, Nishi-ku, Akatsuka, Sakata, 9.x.2019, leg. K. Nakano (1 ♂ 3 ♀♀, TUA); Niigata-ken, Niigata-shi, Kita-ku, Nigorikawa, 7.vi.2020, leg. K. Nakano (5 ♀♀, TUA); Niigata-ken, Niigata-shi, Chuo-ku, Nishifunami-cho, 12.vi.2021, leg. G. Mashima (12 ♂♂ 17 ♀♀, TUA); Niigata-ken, Niigata-shi, Chuo-ku, Bandaijima, 30.viii.2021, leg. J. Souma (18 ♂♂ 10 ♀♀, TUA); Shizuoka-ken, Shimoda-shi, Suzaki, Tsumekizaki, 21.vii.2020, leg. J. Souma (1 ♂, TUA); Shizuoka-ken, Hamamatsu-shi, Nishi-ku, Kamigaya-cho, 16.vi.2017, leg. J. Souma (15 ♂♂ 33 ♀♀, TUA); Shizuoka-ken, Numazu-shi, Kamikanuki, Higashihongo-cho, 27.xii.2021, leg. J. Souma (1 ♀, TUA); Yamaguchi-ken, Shimonoseki-shi, Mimosusogawa-cho, 16.ix.2022, leg. J. Souma (1 ♂ 3 ♀♀, ELKU). Izu Islands (northern part): Izu-Oshima Island: Okada, Minatono-mieru-oka, 4.vi.2019, leg. Y. Tamadera (19 ♂♂ 37 ♀♀, TUA). Jogashima Island : 4.vi.2019, leg. J. Souma (11 ♂♂ 9 ♀♀, TUA). Ebisu Island : 21.vii.2020, leg. J. Souma (3 ♂♂, TUA). Shikoku: Ehime Pref., Kashima, 2.vi.1971, leg. M. Tomokuni (1 ♂ 1 ♀, NSMT); Tosa, Nakagawa, 19.viii.1953, leg. G. Yamamoto (1 ♂, ELKU); Kochi Pref., Cape Ashizuri, 7.vi.1971, leg. M. Tomokuni (1 ♀, NSMT); Kochi-ken, Kochi-shi, Hitsuzan-cho, 30.vi.2020, leg. J. Souma (1 ♀, TUA). Okinoshima Island (Kochi Prefecture): 11.viii.1951, leg. T. Esaki (1 ♀, ELKU; 1 ♀, KUM). Kyushu: Prov. Buzen, Kokura, 20.xi.1951, leg. A. Yamasaki (1 ♂, KUM); Fukuoka, Tachibanayama, 23.vii.1961, leg. S. Miyamoto (6 ♂♂ 8 ♀♀, KUM); as above but 8.ix.1961 (1 ♀, KUM); Fukuoka-ken, Fukuoka-shi, Nishi-ku, Motooka, Kyushu University, 23.v.2020, leg. J. Souma (1 ♀, ELKU); Fukuoka-ken, Itoshima-shi, Tomari, 16.v.2022, leg. J. Souma (1 ♀, ELKU); as above but 10.vii.2022 (1 ♀, ELKU); Kumamoto-Pref., Kumamoto-City, Kuwamizuhonmachi, 4.i.2021, leg. K. Goto (2 ♀♀, ELKU); Ôita Pref., Saiki-shi, Yonouzu, Tsurumisaki, 2.vii.2017, leg. R. Ito (1 ♀, ELKU); Ôita Pref., Saiki-shi, Kamiura, Niinameura, 19.vii.2020, leg. R. Ito (1 ♂ 3 ♀♀, ELKU); Miyazaki Pref., Nichinan-shi, Miyaura, 12.v.2018, leg. R. Ito (1 ♂ 1 ♀, ELKU); Miyazaki Pref., Hyûga-shi, Okuragahama, 1.vi.2019, leg. R. Ito (2 ♀♀, ELKU); Kagoshima, 21.v.1953, leg. S. Miyamoto (1 ♂ 1 ♀, KUM); Oosumi, Izashiki~Ootomari, 25.v.1953, leg. Yoshida (1 ♂, ELKU); Osumi, Sata, 29.v.1953, leg. Yoshida (1 ♂ 1 ♀, ELKU); Osumi, Sata Cape, 30.v.1953, leg. I. Hiura (4 ♀♀, KUM); Kagoshima-ken, Kagoshima-shi, Shiroyama-cho, 4.vii.2017, leg. J. Souma (1 ♂, ELKU); Kagoshima, Minamiosumi-T., Sugiyama-dani Valley , 31.vii.2017, leg. N. Tsuji (1 ♀, ELKU); Kagoshima Pref., Minamiôsumi-chô, Sata, Hetsuka, 30.v.2020, leg. R. Ito (9 ♂♂ 6 ♀♀, ELKU). Okinoshima Island (Fukuoka Prefecture): 25-28.vii.1958, leg. Hirashima, Murakami & Y. Miyatake (35 ♂♂ 70 ♀♀, ELKU; 1 ♀, KUM). Tsushima Island : Izuhara-machi, Kitazato, Kamisaka, 27.vii.2022, leg. Y. Uehara (1 ♀, ELKU). Amakusa Islands : Shimoshima Island : Tomioka, 12.ix.1931, leg. Hori & Chô (2 ♂♂ 4 ♀♀, ELKU; 1 ♀, KUM). Goto Islands : Fukue Island : 1. ix.1962, leg. S. Miyamoto (1 ♂ 1 ♀, KUM); as above but leg. S. Miyamoto & Kawarabata (1 ♀, ELKU); Inuyamaze, 2.ix.1962, leg. S. Miyamoto (2 ♂♂ 1 ♀, KUM); as above but (1 ♂ 1 ♀, KUM); Arakawa, 3.ix.1962, leg. S. Miyamoto (2 ♀♀, KUM). Sakura Island : Kurokami-cho, 27.vii.2021, leg. Y. Obae (18 ♂♂ 13 ♀♀, TUA). Koshiki Islands : Kamikoshiki Island : Nakano, Mt. Tomeki , 2.v.2019, leg. N. Kaneko (1 ♂ 4 ♀♀ 1 nymph, TUA). Shimokoshiki Island : Teuchi, 27-29.viii.1960, leg. K. Morimoto (1 ♂ 2 ♀♀, ELKU); 25.v.1975, leg. Y. Watanabe (1 ♂ 1 ♀, TUA). Ryukyu Islands (northern part): Tanegashima Island : Nishinoomote, 31.viii.1952, leg. C. Takeya & Y. Hirashima (2 ♂♂, ELKU; 2 ♂♂ 2 nymphs, KUM); Nakatane-cho, Masuda, 10.vii.2021, leg. T. Saeki (1 ♀, TUA). Yakushima Island : Onoaida , 26.viii.1952, leg. C. Takeya & Y. Hirashima (3 ♂♂ 1 ♀, ELKU); as above but 27.viii.1952 (8 ♂♂ 11 ♀♀ 1 nymph, ELKU); Miyanoura, 28.viii.1952, leg. C. Takeya & Y. Hirashima (1 ♂ 4 ♀♀, ELKU); as above but 18.v.2022, leg. J. Souma (1 ♀, TUA); Shiratani-unsuikyô, alt. 300-600 m, 14.vii.2017, leg. R. Ito (1 ♂, ELKU); Kurio, Koyojigawa For. Rd. , 7.vii.2021, leg. T. Saeki (1 ♂, ELKU); Kurio, 15.viii.2021, leg. J. Souma (3 ♂♂, ELKU); Koseda, 17.viii.2021, leg. J. Souma (1 ♂, ELKU); as above but 19.v.2022, leg. J. Souma (10 ♂♂ 14 ♀♀, TUA); Funayuki, 17.viii.2021, leg. J. Souma (1 ♀, ELKU); Hirauchi, 19.viii.2021, leg. J. Souma (1 ♂ 4 ♀♀, ELKU); Anbo , 20.viii.2021, leg. J. Souma (9 ♂♂ 14 ♀♀, ELKU); Tabugawa, 18.v.2022, leg. J. Souma (1 ♂ 3 ♀♀, TUA). Nakanoshima Island : 3-13.vi.1953, leg. S. Miyamoto (1 ♀, ELKU); Okizaki, 5.vii.2017, leg. J. Souma (2 ♀♀, TUA); as above but 6.vii.2017 (1 ♀, TUA); Kusuki, 7.vii.2017, leg. J. Souma (4 ♂♂ 7 ♀♀, TUA). Taira Island : Shûraku, 8-10.x.2016, leg. H. Yoshitake (3 ♀♀, TUA); Higashinohama, 8-10.x.2016, leg. H. Yoshitake (1 ♀, TUA). Akuseki Island : 24.iv.1971, leg. M. Sakai (2 ♂♂ 3 ♀♀ 4 nymphs, NSMT); 17.vi.2016, leg. H. Yoshitake (1 ♀, NIAES); Yudomari, 8.vii.2017, leg. J. Souma (1 ♂ 3 ♀♀, TUA). Seven specimens collected from “Ohto” and “Yamogi” adjacent to the type locality “Sakuna” well match the original description of Stephanitis (Norba) aperta ( Horváth 1912). In the present study, the author identified S. (N.) aperta based on these seven individuals . Syntype (s) of S. (N.) aperta exist(s) in the collection of HNHM (D. Rédei, pers. comm. 2021).

Diagnosis.

Stephanitis (Norba) aperta is recognised amongst other species of Stephanitis by a combination of the following characters: head, pronotal disc, marking on hemelytra and ventral surface in various shades of brown (Figs 7B , 9B , 11B , 13B , 15B , 17B , 19B , 21B , 23B ); calli dark brown; body in male 2.1 times (in female 2.0 times) as long as maximum width across hemelytra (Figs 2B , 4B , 6A ); rostrum not reaching metasternum; pronotum unicarinate (Fig. 25B ); hood pale, shorter than median carina of pronotum, as wide as vertex at widest part, not 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 less erect, narrowed posteriorly, with 3 rows of areolae at widest part, with anterolateral angle protruding anteriad, with outer margin gently curved inwards at posterolateral angle, maximum height shorter than height of eye (Fig. 27B ); 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 2 rows (in female with 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. 29B ); and paramere stout, weakly curved inwards at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 31B ).

Remarks.

Amongst the Japanese species of Stephanitis , S. (Norba) aperta is similar to S. (N.) exigua in general habitus, but it is easily distinguished by the following characters: calli dark brown (light brown in S. (N.) exigua ) (Figs 7B, C , 9B, C , 11B, C , 13B, C ); pronotal disc opaque (lustrous in S. (N.) exigua ); paranotum with 3 rows of areolae at widest part (2 rows in S. (N.) exigua ), with anterolateral angle protruding anteriad (slightly protruding in S. (N.) exigua ); and subcostal area of hemelytron in female with 3 rows of areolae at widest part (2 rows in S. (N.) exigua ) (Figs 15B, C , 17B, C ). The place name “Sakuna” was considered to be a misspelling of “Satsuma” [= Kyushu, Kagoshima-ken, former Satsuma-gun in the early 20th century (current Satsumasendai-shi and Satsuma-cho)] by Takeya (1931). However, the former is the name of an actual place in Honshu. The present author confirms the occurrence of S. (N.) aperta in “Ohto” and “Yamogi”, adjacent to “Sakuna” (see material examined). Therefore, “Sakuna” seems to correspond to the type locality of S. (N.) aperta .

Teratological form.

The segmental oligomery of the antenna was confirmed in Stephanitis (Norba) aperta , and one examined specimen lacks the left antennal segment IV (Fig. 6A ), as reported in many tingids ( Štusák and Stehlík 1978; Souma 2020b, 2020d, 2020e). Additionally, the right paranotum of this teratological individual is shorter than that of its normal left side, with two rows of areolae at the widest part (three rows on the left side).

Distribution.

Japan (Honshu; Izu Islands (northern part): Izu-Oshima Island; Jogashima Island; Ebisu Island; Shikoku; Okinoshima Island (Kochi Prefecture); Kyushu; Okinoshima Island (Fukuoka Prefecture); Tsushima Island; Amakusa Islands: Shimoshima Island; Goto Islands: Fukue Island; Sakura Island; Koshiki Islands: Kamikoshiki Island, Shimokoshiki Island; Ryukyu Islands (northern part): Tanegashima Island, Yakushima Island, Nakanoshima Island, Taira Island, Akuseki Island) (Fig. 45 ) ( Horváth 1912; Takeya 1931, 1963; Yamada and Tomokuni 2012; Yamada and Ishikawa 2016; present study). Previous records from China in the 20th century ( Drake and Ruhoff 1965; Jing 1981) do not list the examined specimens and appear to be erroneous. Judging from the photographs, a recent record from Hong Kong ( Yiu and Yip 2012) corresponds to another species, as the pronotum has lateral carina. Therefore, the presence of S. (N.) aperta in China remains unconfirmed. The previous record from the central part of the Ryukyu Islands ( Miyamoto 1964b; Azuma and Kinjo 1987; Hayashi 2002) corresponds to Stephanitis (Norba) exigua , S. (N.) hayashii sp. nov. or S. (N.) hiurai . The previous records from the southern part of the Ryukyu Islands and northern Taiwan ( Takeya 1963; Miyamoto 1964a; Azuma and Kinjo 1987; Hayashi 2002) correspond to S. (N.) ishikawai sp. nov., described below. Hundreds of specimens from the central and southern parts of the Ryukyu Islands possessing the unicarinate pronotum were examined, but all of them belong to S. (N.) exigua , S. (N.) hayashii sp. nov., S. (N.) hiurai or S. (N.) ishikawai sp. nov. Therefore, S. (N.) aperta is probably not distributed in the central and southern parts of Ryukyu Islands. Stephanitis (Norba) aperta inhabits the laurilignosa in a temperate climate of Japan proper (Honshu, Shikoku and Kyushu) and its surrounding islands and the northern part of the Izu and Ryukyu Islands, which is in the Palaearctic Region.

Host plants.

Cinnamomum camphora (L.) J.Presl, “Kusunoki” ( Lauraceae ) (Fig. 43D ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; present study); Neolitsea sericea (Blume) Koidz., “Shirodamo” (Fig. 43C ) (present study); Machilus thunbergii Siebold et Zucc., “Tabunoki” ( Lauraceae ) (Fig. 43B ) ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012; Okochi 2019; present study). Stephanitis (Norba) aperta feeds only on lauraceous trees and is oligophagous. This lace bug was also collected from Symplocos glauca (Thunb.) Koidz., “Mimizubai” ( Symplocaceae ), without any data on its development ( Takeya 1963; Yasunaga et al. 1993; Yamada and Tomokuni 2012). This lace bug sometimes occurs on plantings of C. camphora and M. thunbergii in its distribution range (present study), suggesting that it can become a pest of both lauraceous trees.

Biology.

Stephanitis (Norba) aperta feeds on the abaxial surface of leaves of the three known host plants (present study). Adults were collected in almost all seasons ( Takeya 1931, 1953; Yasunaga et al. 1993; Yano et al. 2013; present study); nymphs were collected in April, May, August, September and November (present study); the overwintering stage is represented by the adult (present study).

Figures

Figure 2. Male habitus of five Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua from Honshu; B. S. (Norba) aperta from Honshu; C. S. (N.) exigua from Minamidaito Island, Daito Islands; D. S. (N.) hayashii sp. nov. from Senaga Island, central part of Ryukyu Islands; E, F. S. (N.) hiurai from Kakeroma (E) and Takara (F) islands, central part of Ryukyu Islands. Scale bar: 1.0 mm.

Figure 4. Female habitus of five Stephanitis species from Japan, dorsal view: A. S. (Stephanitis) ambigua from Honshu; B. S. (Norba) aperta from Honshu; C. S. (N.) exigua from Okinawa Island, central part of Ryukyu Islands; D. S. (N.) hayashii sp. nov. from Aguni Island, central part of Ryukyu Islands; E, F. S. (N.) hiurai from Okinawa (E) and Takara (F) Islands, central part of Ryukyu 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 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 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 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 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 15. Male hemelytra 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 17. Female hemelytra 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 19. Rostra 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. Scale bars: 0.2 mm.

Figure 21. Male terminalia 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. Scale bars: 0.2 mm.

Figure 23. Female terminalia 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. 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 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 29. Line drawings of the pygophore 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.

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 34. Three specimens of Stephanitis species from Japan. A. Holotype of S. (Norba) vitrea [= S. (N.) aperta] deposited in ELKU and its labels; B. Suspected syntype of S. (Stephanitis) propinqua [= S. (N.) aperta] deposited in Matsumura's collection of ELHU and its labels; C. Syntype of S. (N.) exigua [= S. (S.) pyrioides] deposited in Matsumura's collection of ELHU and its labels.

Figure 40. Living individuals of three species of Stephanitis from Japan: A-C. S. (Stephanitis) ambigua from Honshu, male (A) and fifth (B) and fourth (C) instar nymphs; D-G. S. (Norba) aperta from Honshu, male (D), female (E) and fifth (F) and fourth (G) instar nymphs; H, I. S. (N.) exigua from Okinawa Island, central part of Ryukyu Islands, male (H) and fifth instar nymph (I).

Figure 43. Lauraceous host plants of six species of Stephanitis from Japan: A. Lindera glauca from Honshu, damaged by S. (Stephanitis) ambigua; Machilus thunbergii (B), Neolitsea sericea (C) and Cinnamomum camphora (D) from Honshu, all damaged by S. (Norba) aperta; E. M. thunbergii from Okinawa Island, damaged by S. (N.) exigua; Litsea japonica (F) from Kakeroma Island and C. yabunikkei (G) from Tokashiki Island, both damaged by S. (N.) hayashii sp. nov.; H. M. thunbergii from Amami-Oshima Island, damaged by S. (N.) hiurai; I. Lit. japonica from Miyako Island, damaged by S. (N.) ishikawai sp. nov.

Figure 45. Collection sites of Stephanitis (Stephanitis) ambigua, S. (Norba) aperta and S. (N.) exigua used in the present study. Red-filled areas = localities; circles = small isolated islands.