Stephanitis (Norba) ishikawai, Souma, 2022

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) ishikawai sp. nov.

[Japanese name: Ishikawa-gunbai] Figs 3A , 5A , 8A , 10A , 12A , 14A , 16A , 18A , 19F , 21F , 23F , 25F , 28B , 30A-C , 32A-C , 41H, I

Stephanitis (Norba) aperta Horváth, 1912: Takeya (1931: 75) (distribution: part); Maa (1957: 127) (nymph); Takeya (1963: 38) (distribution: part); Miyamoto (1964a: 274) (distribution: part); Miyamoto (1964b: 524) (distribution: part); Takara and Azuma (1972: 113) (distribution); Azuma and Kinjo (1987: 34) (distribution); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Yasunaga et al. (1993: 178) (monograph: part); Hayashi (2002: 137) (distribution); Tomokuni (2006b: 67) (checklist: Taiwan); Zheng and Lin (2013: 312) (distribution); Yamada and Tomokuni (2012: 205) (monograph: part); Yamada and Ishikawa (2016: 433) (checklist: Japan). Misidentifications.

Stephanitis (Norba) exigua Horváth, 1912: Takeya (1963: 38) (distribution: part); Miyamoto (1964a: 274) (distribution: part); Miyamoto (1964b: 524) (distribution: part); Miyamoto (1964c: 105) (distribution); Takara and Azuma (1972: 113) (distribution); Azuma and Kinjo (1987: 34) (distribution); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Hayashi (2002: 137) (distribution); Tomokuni (2006b: 67) (checklist: Taiwan); Yamada and Tomokuni (2012: 204) (monograph: part); Yamada and Ishikawa (2016: 433) (checklist: Japan). Misidentifications.

Stephanitis (Norba) hiurai Takeya, 1963: Miyamoto (1964a: 272) (distribution: part); Miyamoto (1964b: 524) (distribution: part). Misidentifications.

Type series.

Holotype (♂, ELKU), "[JAPAN]: the Ryukyus, Yaeyama Isls., Yonaguni Is., Mantabaru" [=JAPAN: Ryukyu Islands (southern part): Yonaguni Island: Mantabaru (approximate coordinates: 24°27'27.6"N, 122°58'16.6"E)], 11.xi.2020, leg. J. Souma. Paratypes (141 ♂♂ 252 ♀♀), JAPAN: Ryukyu Islands (southern part): Miyako Island: Karimata, 2.ix.1958, leg. T. Hidaka (1 ♀, ELKU); Hirara, Karimata, 1.iv.1991, leg. M. Hayashi (1 ♀, TUA); as above but 21.xi.1992 (2 ♂♂ 1 ♀, ELKU); as above but 22.xi.1992 (1 ♀, TUA); Gusukube, Wipya, 2.iv.1991, leg. M. Hayashi (5 ♂♂ 9 ♀♀, TUA); Mt. Nobarudake, 2.iv.1991, leg. M. Hayashi (6 ♂♂ 12 ♀♀, TUA); as above, but 11.x.1993 (7 ♂♂ 21 ♀♀, TUA); as above, but 13.v.1995 (1 ♀, TUA); as above, but 24.vi.2008 (2 ♂♂ 1 ♀, TUA); as above but 9.xi.2018, leg. J. Souma (3 ♂♂ 2 ♀♀, TUA); Ono Mountain Forest, 18.v.1991, leg. M. Hayashi (3 ♀♀, TUA); as above but 7.xi.2018, leg. J. Souma (5 ♀♀, ELKU; 1 ♂ 2 ♀♀, TUA); Ueno-son, Mt. Nobaru-dake, 31.x.1999, leg. T. Ishikawa (3 ♂♂ 4 ♀♀, TUA); as above but leg. M. Tomokuni (2 ♂♂ 1 ♀, NSMT); Karimata, Ôno-sanrin, 1.xi.1999, leg. T. Ishikawa (1 ♂, TUA); Hirara, Higashi-sokobaru, 2.xi.1999, leg. T. Ishikawa (1 ♂ 1 ♀, TUA); Kamamamine Park, 28.iv.2002, leg. M. Hayashi (1 ♀, TUA); Ueno, Mt. Nobarudake, 27.xii.2017, leg. H. Yoshitake (2 ♂♂ 3 ♀♀, TUA); Gusukube, Yoshino, 8.xi.20118, leg. J. Souma (1 ♂ 7 ♀♀, TUA). Irabu Island: Irabu, 10.xi.2018, leg. J. Souma (1 ♂, ELKU); Makiyama, 10.xi.2018, leg. J. Souma (1 ♂ 2 ♀♀, ELKU); as above but 25.vii.2022, leg. S. Shimamoto (1 ♂ 3 ♀♀, TUA); near Makiyama-tenbôdai, 26.ix.2020, leg. H. Yoshitake (1 ♂ 1 ♀, NIAES). Kurima Island: 31.x.2007, leg. M. Hayashi (1 ♀, TUA). Ogami Island: 9.vi.2009, leg. M. Hayashi (1 ♂, TUA); 27.xii.2017, leg. H. Yoshitake (2 ♂♂ 6 ♀♀, TUA). Ishigaki Island: 5.ix.1957, leg. T. Takara (2 ♀♀, NSMT); 20.iii.1960, leg. T. Takara (1 ♂ 1 ♀, KUM); Nanbadake, 24.xi.1960, leg. K. Yasumatsu (1 ♀, KUM); Kawara, 28.x.1963, leg. Y. Hirashima (2 ♀♀, KUM); Mt. Omoto, alt. 100-526 m, 17.xi.1963, leg. G. A. Samuelson (1 ♀, KUM); Yonehara, 15.iii.1964, leg. Taira (1 ♂ 1 ♀, NIAES); Kahara-yama, 14.iii.1964, leg. Y. Miyatake (4 ♂♂ 1 ♀, KUM); as above but leg. S. Kimoto (1 ♂, KUM); as above but 18.iii.1964 (1 ♀, KUM); Yonehara, 15.iii.1964, leg. T. Shirozu (1 ♂ 3 ♀♀, KUM); as above but leg. Y. Miyatake (1 ♂ 1 ♀, KUM); as above but 12.vi.2008, leg. M. Hayashi (1 ♀, TUA); Omoto-san, 16.iii.1964, leg. Y. Miyatake (2 ♂♂ 6 ♀♀, KUM); Takada, 10.iv.1981, leg. K. Baba (1 ♀, NIAES); as above but 10.xi.1985, leg. M. Hayashi (2 ♀♀, TUA); Omoto, 15.x.1988, leg. M. Hayashi (1 ♀, TUA); as above but 2.vii.1998, leg. M. Hayashi (1 ♀, TUA); Banna-dake, 12.v.1975, leg. Y. Notsu (1 ♂, NSMT); as above but 16.xi.1984, leg. M. Tomokuni (2 ♀♀, NSMT); Mt. Bannadake, 2.iv.1976, leg. K. Murakami (1 ♂, NSMT); as above but 19.vii.2022, leg. S. Shimamoto (3 ♂♂ 6 ♀♀, TUA); Banna Park, 17.iii.1993, leg. M. Hayashi (1 ♀, TUA); Nosoko, 1.v.1998, leg. K. Takahashi (1 ♂, NSMT); Mt. Yarabu-dake, 23.iv.1999, leg. T. Ishikawa (1 ♀, TUA); as above but 6-9.v.2016, leg. N. Tsuji (2 ♀♀, ELKU); as above but 13.xi.2018, leg. J. Souma (1 ♂, TUA); Takeda-rindo, 23.v.2000, leg. H. Mizushima (1 ♂, TUA); Mt. Omoto-dake, 18.vi.1991, leg. S. Miyakawa (4 ♀♀, NSMT); as above but 22.vi.1991 (1 ♀, NSMT); as above but 12.v.1993, leg. M. Hayashi (1 ♀, TUA); as above but 24.v.2000, leg. T. Ishikawa (1 ♀, TUA); Mt. Maesedake, 20.iv.2005, leg. M. Hayashi (3 ♂♂ 3 ♀♀, TUA); as above but 21.ii.2006 (2 ♂♂ 4 ♀♀, TUA) as above but 22.iii.2006 (1 ♂, TUA); as above but 16.xi.2018, leg. J. Souma (1 ♀, TUA); as above but 6.iii.2022, leg. T. Saeki (1 ♂, TUA); as above but 20.vii.2022, leg. S. Shimamoto (5 ♀♀, TUA); Shiramizu, 29.x.2007, leg. M. Hayashi (1 ♀, TUA); Kabira, 29.x.2007, leg. M. Hayashi (1 ♀, TUA); as above but 8.x.2013, leg. R. Ito (1 ♂ 1 ♀, ELKU); Hirae, Takeda-rindô, 4.x.2013, leg. R. Ito (1 ♀, ELKU); Hirae, Omoto-dake, 7.x.2013, leg. R. Ito (1 ♂, ELKU); Nagura, 7.x.2013, leg. R. Ito (3 ♂♂ 2 ♀♀, ELKU); Yarabu, Yarabu-dake, 8.x.2013, leg. R. Ito (1 ♀, ELKU); Yarabu, 8.x.2013, leg. R. Ito (1 ♀, ELKU); Okawa, 11.xi.2018, leg. J. Souma (1 ♂, TUA); Oganzaki, 13.xi.2018, leg. J. Souma (1 ♀, ELKU); Nosoko, 14.xi.2018, leg. J. Souma (1 ♂, ELKU); Itona, 14.xi.2018, leg. J. Souma (1 ♂, TUA); Inoda, 14.xi.2018, leg. J. Souma (1 ♀, ELKU); Hirae, 17.xi.2018, leg. J. Souma (1 ♀, TUA); Sakieda, 1.iii.2020, leg. Y. Obae (2 ♂♂ 1 ♀, ELKU). Iriomote Island: 24.iv.1951, leg. T. Takara (1 ♀, NSMT); Ôhara, 22.xi.1960, leg. K. Yasumatsu (1 ♂, ELKU); Ushikumori, 4.x.1963, leg. S. Miyamoto (1 ♂, KUM); as above but 9.iii.1964, leg. Y. Miyatake (2 ♂♂ 1 ♀, KUM); as above leg. T. Shirozu (1 ♂ 4 ♀♀, KUM); as above but 11.iii.1964, leg. Y. Miyatake (3 ♂♂ 2 ♀♀, KUM); as above but 24.iii.1973, leg. S. Azuma (1 ♂ 2 ♀♀, NSMT); Sonai, 6.x.1963, leg. S. Miyamoto (1 ♂, KUM); as above but 8.x.1963 (1 ♂ 1 ♀, KUM); Komi, 12.vii.1963, leg. Y. Miyatake (4 ♀♀, KUM); as above but 2.iv.1978, leg. K. Baba (1 ♀, NIAES); as above but 10.xi.1984, leg. M. Tomokuni (1 ♀, NSMT); as above but 11.xi.1984, leg. M. Tomokuni (1 ♂, NSMT); as above but 20.xi.1998, leg. T. Ishikawa (1 ♀, TUA); as above but 20-23.iii.2015, leg. R. Ito (1 ♀, ELKU); as above but 24.viii.2020, leg. Y. Hisasue (1 ♂, ELKU); Shirahama, 5.xi.1963, leg. H. Hasegawa (1 ♀, NIAES); Shirahama-Sonae, 5.xi.1963, leg. H. Hasegawa (1 ♀, KUM); Shirahama-Hoshidate, 8.iii.1964, leg. Y. Miyatake (2 ♂♂ 2 ♀♀, KUM); as above but leg. S. Kimoto (1 ♀, KUM); Upstream of Nakara-gawa Riv., 12.iii.1964, leg. Y. Miyatake (1 ♀, KUM); as above but leg. S. Kimoto (1 ♂ 1 ♀, KUM); as above but leg. S. Higashihirachi (1 ♂, NIAES); Ohara, 20.viii.1971, leg. S. Azuma (1 ♂, NSMT); Funaura, 28.iii.1973, leg. S. Azuma (1 ♂, NSMT); as above but 21.viii.1976, leg. K. Kyoda (1 ♀, NSMT); as above but 8.x.1977, leg. M. Taniguchi (1 ♀, NSMT); as above but 9.x.1977, leg. M. Kinjo (1 ♀, NSMT); as above but 9.x.1977, leg. M. Arasaki (1 ♀, NSMT); as above but 11.xi.1984, leg. M. Tomokuni (1 ♀, NSMT); as above but 31.iii.1996, leg. M. Hayashi (1 ♂, TUA); Hoshidate, 7.xi.1985, leg. M. Hayashi (1 ♂, TUA); Monpanare, 29.ix.1993, leg. M. Hayashi (1 ♀, TUA); Takana, Yutsun Riv., 22.xi.1997, leg. M. Tomokuni (1 ♀, NSMT); Urauchi, 19.vi.1998, leg. T. Ishikawa (1 ♀, TUA); Shirahama-rindô, 1.iii.2002, leg. T. Ishikawa (2 ♀, TUA); Mt. Sonaidake, 1.iii.2002, leg. T. Ishikawa (2 ♂♂ 1 ♀, TUA); Ôhara, Fusatoruba, 3.iii.2002, leg. T. Ishikawa (4 ♂♂ 1 ♀, TUA); Aira-gawa Rev., 7.x.2009, leg. T. Ishikawa (1 ♂, TUA); Haiminaka, 21-23.iii.2015, leg. R. Ito (1 ♀, ELKU); Sonai, 19.xi.2018, leg. J. Souma (1 ♀, ELKU); Otomi For. Rd., 20.xi.2018, leg. J. Souma (1 ♂, TUA); Unarizaki, 20.xi.2018, leg. J. Souma (1 ♂, ELKU); Ôtomi-path, 21.ii.2020, leg. R. Ito (3 ♂♂ 3 ♀♀, ELKU); Mt. Tedo-san, 14.x.2020, leg. Y. Hisasue (1 ♀, ELKU); Inaba Trail, 16.x.2020, leg. Y. Hisasue (1 ♀, ELKU). Yonaguni Island: Mt. Kubura-dake, 2.vii.1973, leg. K. Shigematsu (1 ♂, NSMT); as above but 30.iii.2020, leg. R. Ito (1 ♀, ELKU); Mt. Urabu-dake, 28.iii.1997, leg. T. Ishikawa (5 ♂♂ 1 ♀♀, TUA); as holotype (6 ♂♂ 13 ♀♀, ELKU); as holotype but 14.xi.2020 (2 ♂♂ 8 ♀♀, ELKU); Mt. Inbidake, 11.xi.2020, leg. J. Souma (1 ♂ 3 ♀♀, ELKU); as above but 13.xi.2020, leg. J. Souma (11 ♂♂ 16 ♀♀, ELKU); Higawa, 11.xi.2020, leg. J. Souma (4 ♀♀, ELKU); Iranda For. Rd., 25.iii.2021, leg. T. Saeki (1 ♂, ELKU); Promenade of Atlas Moth Museum, 26.iii.2021, leg. K. Saito (2 ♂♂ 1 ♀, ELKU). A single paratype collected in 1958 was recorded as " Stephanitis aperta ", four paratypes collected in 1960 as " Stephanitis exigua " by Takeya (1963). Of 12 paratypes collected in 1963, eight were recorded as " Stephanitis exigua " or " Stephanitis hiurai " by Miyamoto (1964a). Of 42 paratypes collected in 1964, 38 individuals were recorded as " Stephanitis exigua " by Miyamoto (1964b). Four paratypes from Iriomote Island collected by Y. Miyatake in 1963 were recorded as " Stephanitis exigua " by Miyamoto (1964c). Six paratypes from Iriomote Island collected by S. Azuma or M. Kinjo were considered to be recorded as " Stephanitis aperta " or " Stephanitis exigua " by Azuma and Kinjo (1987).

Additional material examined.

Non-types (13 ♂♂ 8 ♀♀ 13 nymphs), JAPAN: Ryukyu Islands: (southern part): Iriomote Island: Shirahama-Hoshidate , 8.iii.1964, leg. Y. Miyatake (1 nymph); Ôhara, Fusatoruba , 3.iii.2002, leg. T. Ishikawa (4 nymphs, TUA). Yonaguni Island : as holotype (1 nymph) . TAIWAN: Taipei City: Taihoku [= Taipei], 16.iv.1917, leg. M. Maki (2 ♂♂, ELKU); as above but 15.i.1927, leg. R. Takahashi (1 ♀, ELKU); as above but 2.iv.1930, leg. C. Takeya (1 ♂ 1 ♀, ELKU); as above, but 10.ii.1932, leg. R. Takahashi (4 ♂♂ 2 ♀♀, ELKU; 1 ♂, KUM); Da'an District , 5.x.2021, leg. Y.-J. Tsai (3 ♂♂ 1 ♀ 7 nymphs, NMNS); as above but 22.xi.2021 (2 ♂♂ 3 ♀♀, NMNS). Twelve specimens collected from Taiwan in the early 20th century were recorded as " Stephanitis aperta " or " Stephanitis exigua " by Takeya (1931, 1963). The Taiwanese specimens are most similar to Stephanitis (Norba) ishikawai sp. nov. in morphological characteristics and were provisionally identified as pertaining to the new species in the present study. All 13 nymphs recorded above are in poor condition and are thus not described here .

Diagnosis.

Stephanitis (Norba) ishikawai sp. nov. 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 8A , 10A , 12A , 14A , 16A , 18A , 19F , 21F , 23F ); calli light brown; body in male 2.1 times (in female 2.0 times) as long as maximum width across hemelytra (Figs 3A , 5A ); rostrum reaching metasternum; pronotum unicarinate (Fig. 25F ); hood pale, shorter than median carina of pronotum, wider than vertex at widest part, incompletely covering eye, as high as median carina of pronotum at highest part, posterior margin extending to middle of pronotal disc; median carina of pronotum with 2 rows of areolae at highest part; pronotal disc opaque; paranotum less 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. 28B ); 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, posterior margin slightly emarginate in middle part (Fig. 30A-C ); and paramere stout, weakly curved inwards at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 32A-C ).

Description.

Male. Head, pronotal disc, marking on hemelytra and ventral surface in various shades of brown; calli light brown; eye dark red; areolae of pronotum and hemelytron transparent; hood pale; pronotal disc opaque; pubescence on body yellowish (Figs 3A , 8A , 12A , 16A , 19F , 21F ).

Body 2.1 times as long as maximum width across hemelytra (Fig. 3A ). Head (Figs 8A , 12A , 19F , 25F ) glabrous; a pair of frontal spines close to each other at apices, not reaching apex of clypeus; median spine as long as frontal spines, reaching bases of frontal spines; pair of occipital spines longer than median spine, reaching middle part of eyes; antenniferous tubercles obtuse, slightly curved inwards; clypeus smooth. Compound eye round in dorsal view. Antenna densely covered with pubescence; segment I cylindrical; segment II cylindrical, shortest among antennal segments; segment III longest amongst antennal segments; segment IV cylindrical, longer than segment I. Bucculae closed at anterior ends, with 3 rows of areolae throughout length. Rostrum reaching metasternum.

Pronotum (Figs 8A , 12A , 25F , 28B ) unicarinate, 1.3 times as long as maximum width across paranota, sparsely covered with pubescence. Pronotal disc coarsely punctate. Hood shorter than median carina of pronotum, wider than vertex at widest part, incompletely covering eye, as high as median carina of pronotum at highest part, posterior margin extending to middle of pronotal disc, 4 rows of areolae at highest part, dorsal margin arched. Median carina straight, extending to apex of posterior process, 2 rows of areolae at highest part, dorsal margin arched. Calli smooth. Paranotum less 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. Posterior process triangular, obtuse at apex.

Hemelytron (Fig. 16A ) 2.4 times as long as its maximum width, extending beyond apex of abdomen, sparsely covered with pubescence; maximum width across hemelytra 1.6 times as much as maximum width across paranota; apices close to each other in rest; costal area with 3-4 rows of areolae at widest part; subcostal area with 2 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; C (costal), R+M (radiomedial) and Cu (cubital) veins carinate.

Thoracic pleura (Fig. 12A ) smooth in anterior part, coarsely punctate in posterior part. Ostiolar peritreme oblong. Sternal laminae (Fig. 19F ) lower than bucculae; pro- and mesosternal laminae open in both anterior and posterior ends; metasternal laminae as high as mesosternal laminae, open at anterior ends, fused each other at posterior ends. Legs (Fig. 3A ) smooth, densely covered with pubescence; femora thickest at middle.

Abdomen oblong in dorsal and ventral views. Pygophore (Figs 21F , 30A-C ) compressed dorsoventrally, semicircular in ventral view, elevated at centre of venter, with posterior margin slightly emarginate in middle part, covered with pubescence. Paramere (Fig. 32A-C ) stout, expanded in middle part, weakly curved inwards at apex, outer margin not sinuate in middle part, inner margin nearly straight in basal part, covered with pubescence in middle part of outer and inner margins.

Measurements (n = 20). Body length with hemelytra 2.9-3.2 mm; maximum width across hemelytra 1.4-1.6 mm; length of antennal segments I to IV 0.2 mm, 0.1 mm, 1.2 mm and 0.7-0.8 mm, respectively; pronotal length 1.2-1.3 mm; pronotal width across paranota 0.8-1.0 mm; hemelytral length 2.2-2.5 mm; maximum width of hemelytron 0.9-1.1 mm.

Female. General habitus very similar to that of male (Figs 4D , 9D , 13D , 17D , 23D ), except for the following characters: body 2.0 times as long as maximum width across hemelytra; antennal segment III shorter than in male; pronotum 1.4 times as long as maximum width across paranota; hood wider than in male; hemelytron 2.3 times as long as maximum width; maximum width across hemelytra 1.7 times as much as maximum width across paranota; subcostal area wider than in male, with 3 rows of areolae at widest part; and apical part of abdomen pentagonal in ventral view.

Measurements (n = 20). Body length with hemelytra 3.0-3.4 mm; maximum width across hemelytra 1.5-1.7 mm; length of antennal segments I to IV 0.2 mm, 0.1 mm, 1.0 mm and 0.7-0.8 mm, respectively; pronotal length 1.2-1.4 mm; pronotal width across paranota 0.9-1.0 mm; hemelytral length 2.3-2.5 mm; maximum width of hemelytron 1.0-1.1 mm.

Remarks.

Amongst the Japanese species of Stephanitis , S. (Norba) ishikawai sp. nov. is most similar to S. (N.) hayashii sp. nov. in general habitus, but the former is easily distinguished from the latter by the following characters: length of antennal segment IV 0.7-0.8 mm (0.6 mm in S. (N.) hayashii sp. nov.); paranotum less erect (more erect in S. (N.) hayashii sp. nov.) (Figs 7D , 8A , 9D , 10A , 11D , 12A , 13D , 14A ); and paramere stout (slender in S. (N.) hayashii sp. nov.), with inner margin nearly straight in basal part (slightly curved inward in S. (N.) hayashii sp. nov.) (Figs 31C, D , 32A-C ). Moreover, the partial COI gene pairwise sequence distances between both species are 0.042503-0.048201 (Suppl. material 3). Stephanitis (Norba) ishikawai sp. nov. was misidentified as S. (N.) aperta , S. (N.) exigua and S. (N.) hiurai in previous studies ( Takeya 1931, 1963; Maa 1957; Miyamoto 1964a, 1964b; Takara and Azuma 1972; Yasunaga et al. 1993; Yamada and Tomokuni 2012; Zheng and Lin 2013), as these four species are very similar to each other. However, S. (N.) ishikawai sp. nov. is easily distinguished from the three species by the following characters: rostrum reaching metasternum (not reaching in S. (N.) aperta , S. (N.) exigua and S. (N.) hiurai ) (Fig. 19B, C, E, F ); and hood slightly higher than median carina of pronotum at highest part (as high as S. (N.) aperta , S. (N.) exigua and S. (N.) hiurai ), with posterior margin extending to middle of pronotal disc (not extending in S. (N.) aperta , S. (N.) exigua and S. (N.) hiurai ) (Figs 7B, C, E, F , 8A , 9B, C, E, F , 10A , 11B, C, E, F , 12A , 13B, C, E, F , 14A , 25B, C, E, F ).

Distribution.

Japan (Ryukyu Islands (southern part): Miyako Island, Irabu Island, Kurima Island, Ogami Island, Ishigaki Island, Iriomote Island, Yonaguni Island); Taiwan (northern part) (Fig. 47 ) ( Takeya 1931, 1963; Maa 1957; Miyamoto 1964a, 1964b, 1964c; Takara and Azuma 1972; Azuma and Kinjo 1987; Yasunaga et al. 1993; Hayashi 2002; Yamada and Tomokuni 2012; Zheng and Lin 2013; Yamada and Ishikawa 2016; present study). Judging from the photographs, living individuals identified as " Stephanitis (Norba) aperta " or " S. (N.) exigua " in previous studies (Yasunaga et al. 193; Yamada and Tomokuni 2012; Zheng and Lin 2013) corresponded to the new species. Stephanitis (Norba) ishikawai sp. nov. inhabits the laurilignosa in a subtropical climate of the southern part of the Ryukyu Islands and northern Taiwan, which is in the Oriental Region.

Etymology.

The new species is named in honour of Tadashi Ishikawa, a Japanese heteropterist who collected part of paratypes and taught the author how to describe new species.

Host plants.

Cinnamomum camphora , “Kusunoki” ( Lauraceae ) (present study); Litsea japonica , “Hamabiwa” ( Lauraceae ) (Fig. 43I ) (present study); Machilus thunbergii , “Tabunoki” ( Lauraceae ) (present study). Stephanitis (Norba) ishikawai sp. nov. feeds only on lauraceous trees and is oligophagous.

Biology.

Stephanitis (Norba) ishikawai sp. nov. feeds on the abaxial surface of leaves of the three host plants (present study). Adults were collected in almost all seasons ( Miyamoto 1964a, 1964b, 1964c; Takara and Azuma 1972; Yasunaga et al. 1993; Yamada and Tomokuni 2012; present study); nymphs were collected in March and November (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 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 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 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 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 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 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 41. Living individuals of three species of Stephanitis from Japan: A-D. S. (Norba) hayashii sp. nov. from the central part of Ryukyu Islands, male (A), female (B) and fifth (C) and fourth (D) instar nymphs from Kakeroma Island; E-G. S. (N.) hiurai from Amami-Oshima Island, central part of Ryukyu Islands, male (E) and female (F) and fifth instar nymph (G); H, I. S. (N.) ishikawai sp. nov. from Yonaguni Island, southern part of Ryukyu Islands, female (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 47. Collection sites of Stephanitis (Norba) ishikawai sp. nov. and S. (N.) mendica used in the present study. Red-filled areas = localities; circles = small isolated islands.