Stephanitis (Norba) hiurai Takeya, 1963

[Japanese name: Hiura-gunbai] Figs 2E, F, 4E, F, 7E, F, 9E, F, 11E, F, 13E, F, 15E, F, 17E, F, 19E, 21E, 23E, 25E, 28A, 29F, 31F, 36A, 36B, 41E-G

Stephanitis hiurai Takeya, 1963: 34 (in Norba group). Holotype, ♀ (Fig. 36A): Japan: Amami-Oshima, Naze [= Ryukyu Islands, Amami-Oshima Island, Naze]; OMNH (not deposited), ELKU (currently deposited).

Stephanitis hiurai takaranis Takeya, 1963: 36 (in Norba group). Holotype, ♀ (Fig. 36B): Japan: Tokara Is., Takarajima [= Ryukyu Islands, Takara Island]; OMNH (not deposited), ELKU (currently deposited). Synonymised with Stephanitis (Norba) hiurai Takeya, 1963 by Miyamoto (1964b: 524).

Stephanitis (Norba) aperta Horváth, 1912: Hayashi (2002: 137) (distribution). Misidentification.

References.

Miyamoto (1964a: 274) (distribution: part); Miyamoto (1964b: 524) (distribution: part); Miyamoto and Yasunaga (1989: 168) (checklist: Japan); Péricart and Golub (1996: 58) (catalogue: Palaearctic); Hayashi (2002: 137) (distribution); Yamada and Tomokuni (2012: 205) (monograph); Yamada and Ishikawa (2016: 433) (checklist: Japan).

Material examined.

Holotype (1 ♀, ELKU) (Fig. 36A), JAPAN: Ryukyu Islands (central part): " Amami Is. Naze " [= Amami-Oshima Island, Naze (approximate coordinates: 28°23'13.6"N, 129°29'38.2"E)], 4.v.1960, leg. T. Shibata. Holotype of Stephanitis (Norba) hiurai takaranis Takeya, 1963 (1 ♀, ELKU) (Fig. 36B), JAPAN: Ryukyu Islands (central part): " Tokara Is. Takarajima " [= Takara Island (approximate coordinates: 29°08'38.0"N, 129°12'27.8"E)], 5.vii.1960, leg. Y. Hama. Paratype of Stephanitis (Norba) hiurai takaranis Takeya, 1963 (1 ♀, ELKU), JAPAN: Ryukyu Islands (central part): " Tokara Is. Takarajima " [= Takara Island (approximate coordinates: 29°08'38.0"N 129°12'27.8"E)], 5.vii.1960, leg. Y. Hama. The original description of S. (N.) hiurai and S. (N.) hiurai takaranis (Takeya 1963) states that their type specimens are deposited in OMNH, but these holotypes and paratype are now deposited in ELKU. Non-types collected at type locality (4 ♂♂ 6 ♀♀), JAPAN: Ryukyu Islands (central part): Amami-Oshima Island: Amami-shi, Naze, Kaneku (approximate coordinates: 28°22'27.1"N, 129°29'43.6"E), 30.iv.2022, leg. J. Souma (3 ♂♂ 5 ♀♀, TUA); Amami-shi, Naze, Asani (approximate coordinates: 28°24'03.1"N, 129°29'25.3"E), 30.iv.2022, leg. J. Souma (1 ♂ 1 ♀, TUA). Non-types (50 ♂♂ 104 ♀♀ 16 nymphs), JAPAN: Ryukyu Islands (central part): Takara Island: 26.v-1.vi.1953, leg. S. Miyamoto (1 ♂ 2 ♀♀, ELKU); 27.iv.1971, leg. M. Sakai (2 ♀♀ 1 nymph, NSMT). Kikai Island: Hyakunodai, 9.iii.2019, leg. H. Kojima (1 ♂ 3 ♀♀, TUA); between Keraji and Aden, 9.iii.2019, leg. H. Kojima (9 ♀♀, TUA). Amami-Oshima Island: Hatsuno, 11.xi.1962, leg. Y. Miyatake (1 ♂, ELKU); as above but 5.x.1988, leg. M. Tomokuni (1 ♂ 3 ♀♀, NSMT); Uragami, 31.x.1966, leg. Y. Miyatake (3 ♂♂ 2 ♀♀, KUM); Santaro-toge Pass, 2.xi.1966, leg. Y. Miyatake (1 ♀, KUM); as above but 30.vi.2000, leg. Y. Nakatani (1 ♂, NIAES); Yuwan, 3.xi.1966, leg. Y. Miyatake (5 ♂♂ 10 ♀♀, KUM); Naze, 21.iv.1971, leg. M. Sakai (3 ♀♀ 1 nymph, NSMT); Naze-shi, Akazaki, 2.xi.1984, leg. M. Tomokuni (1 ♂ 5 ♀♀, NSMT); Sumiyô-son, Kawauchi, Chûô-rindô, alt. 200 m, 3.x.1988, leg. M. Tomokuni (5 ♀♀, NSMT); Mt. Yuwandake, 4.x.1988, leg. M. Tomokuni (1 ♀, NSMT); Uken-son, Mt. Yuwan-dake, 23.iii.2019, leg. Y. Hisasue (1 ♀, TUA); Shinokawa, 2.xi.2020, leg. J. Souma (3 ♂♂ 1 ♀, ELKU; 5 ♂♂ 5 ♀♀, TUA); as above but 4.xi.2020 (1 ♂ 1 ♀ 8 nymphs, TUA); Konase, 2.xi.2020, leg. J. Souma (5 ♂♂ 5 ♀♀ 1 nymph, TUA); Sokaru, 5.xi.2020, leg. J. Souma (1 ♀, TUA); Uken-son, Yuwan, 28.iv.2022, leg. J. Souma (1 ♂ 2 ♀♀, TUA); Setouchi-cho, Amurogama, 29.iv.2022, leg. J. Souma (3 ♀♀, TUA); Uken-son, Ikegachi, 30.iv.2022, leg. J. Souma (1 ♂, TUA); Amami-shi, Sumiyo-cho, Aoku, 30.iv.2022, leg. J. Souma (1 ♂ 1 ♀, TUA); Amami-shi, Sumiyo-cho, Santaro Pass, 1.v.2022, leg. J. Souma (1 ♂ 5 ♀♀ 4 nymphs, TUA); Amami-shi, Kasari-cho, Manya, 3.v.2022, leg. J. Souma (2 ♂♂ 2 ♀♀, TUA); Asani, Akazaki Park, 29.vi.2022, leg. S. Imada (1 ♀, TUA). Kakeroma Island: Shokazu, 3.xi.2020, leg. J. Souma (3 ♀♀, TUA); Kachiyuki, 3.xi.2020, leg. J. Souma (8 ♂♂ 10 ♀♀ 1 nymph, TUA). Tokunoshima Island: Asahigaoka, 11.xi.1966, leg. Y. Miyatake (2 ♀♀, KUM); Amagi-cho, Yonama, 2.x.1988, leg. M. Tomokuni (2 ♂ 1 ♀, NSMT). Okinoerabu Island: Mt. Koshiyama, 8.vii.2019, leg. Y. Tamadera (1 ♂ 3 ♀♀, TUA); Oyama Botanical Park, 23-26.vi.2022, leg. S. Imada (3 ♀♀, TUA). Okinawa Island: Yona, 19.x.1963, leg. S. Miyamoto (2 ♂♂, KUM); as above but leg. Y. Hirashima (1 ♀, KUM); as above but 25-27.v.1974, leg. M. Sato (2 ♀♀, NSMT); Izumi-Gogayama, 22.iii.1964, leg. S. Kimoto (1 ♂, KUM); as above but leg. Y. Miyatake (1 ♀, KUM); Aha, 26.xii.1973, leg. M. Hamakawa (1 ♂ 1 ♀, NSMT); Kunigami, Mt. Yonaha, 20.x.1987, leg. M. Sakai (1 ♀, NSMT); Kunigami, Ooguni-rindo, 8 km from Yona, alt. 300 m, 21.x.1987, leg. M. Tomokuni (1 ♂, NSMT); Okuni-Rindô, 16-21.iv.1997, leg. T. Ishikawa (1 ♀, TUA); Oku For. Rd., 12.ix.2005, leg. M. Hayashi (1 ♀, TUA) .

Diagnosis.

Stephanitis (Norba) hiurai 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 7E, F, 9E, F, 11E, F, 13E, F, 15E, F, 17E, F, 19E, 21E, 23E); calli light brown; body in male 2.0-2.1 times (in female 1.8-2.0 times) as long as maximum width across hemelytra (Figs 2E, F, 4E, F); rostrum not reaching metasternum; pronotum unicarinate (Fig. 25E); 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, with posterior margin not extending to middle of pronotal disc; median carina of pronotum with 2 rows of areolae at highest part; pronotal disc lustrous; paranotum less erect, narrowed posteriorly, 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. 28A); apices of hemelytra close to each other in rest; costal area with 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 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 in male not carinate (in female carinate); pygophore elevated at centre of venter, posterior margin slightly emarginate in middle part (Fig. 29F); and paramere stout, weakly curved inwards at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 31F).

Intraspecific variation.

According to the original description (Takeya 1963), Stephanitis (Norba) hiurai is divided into two subspecies, S. (N.) h. hiurai Takeya, 1963 and S. (N.) h. takaranis . However, previous studies by Miyamoto (1964a, 1964b) used only a few specimens which included misidentified S. (N.) ishikawai sp. nov., described below, and synonymised S. (N.) h. takaranis with the nominotypical subspecies. Nevertheless, subsequent authors ( Péricart and Golub 1996; Yamada and Tomokuni 2012; Yamada and Ishikawa 2016) continued to treat S. (N.) h. takaranis as a valid subspecies without providing any justification. Many specimens of S. (N.) hiurai from various localities were observed in the present study and the differences in colouration and shape of the hood cited in the original description of the two subspecies (Takeya 1963) are herein considered to be intraspecific variations depending on individual and size, respectively. Therefore, the synonymy of S. (N.) h. takaranis with S. (N.) hiurai, proposed by Miyamoto (1964a, 1964b) is supported.

Remarks.

The partial COI gene pairwise sequence distances between Stephanitis (Norba) hiurai and S. (N.) aperta are only 0.006632-0.009310 (Suppl. material 3) and both species resemble each other in general habitus. However, the former is easily distinguished from the latter by the following characters: hood wider than vertex at widest part (as wide as vertex in S. (N.) aperta), incompletely covering eye (not covering in S. (N.) aperta) (Figs 7B, E, F, 9B, E, F, 25B, E); pronotal disc lustrous (opaque in S. (N.) aperta); and R+M (radiomedial) vein in female carinate (not carinate in S. (N.) aperta) (Fig. 17B, E, F).

Distribution.

Japan (Ryukyu Islands (central part): Takara Island, Kikai Island, Amami-Oshima Island, Kakeroma Island, Tokunoshima Island, Okinoerabu Island, Okinawa Island) (Fig. 46) (Takeya 1963; Miyamoto 1964a, 1964b, 1964c; Hayashi 2002; Yamada and Tomokuni 2012; Yamada and Ishikawa 2016; present study). The previous record from Ishigaki Island, the southern part of the Ryukyu Islands (Miyamoto 1964a, 1964b), corresponds to Stephanitis (Norba) ishikawai sp. nov., described below. Hundreds of specimens from the southern part of the Ryukyu Islands possessing an unicarinate pronotum were examined, but all of them belonged to S. (N.) ishikawai sp. nov. Therefore, S. (N.) hiurai is probably not distributed in the southern part of the Ryukyu Islands. Stephanitis (Norba) hiurai inhabits the laurilignosa in a subtropical climate of the central part of the Ryukyu Islands, which is in the Oriental Region.

Host plants.

Machilus thunbergii, “Tabunoki” ( Lauraceae) (Fig. 43H) (present study). Stephanitis (Norba) hiurai feeds only on this lauraceous tree and is monophagous. This lace bug was also collected from Symplocos myrtacea Siebold et Zucc., “Hainoki” ( Symplocaceae) without any data on its development (Yamada and Tomokuni 2012).

Biology.

Stephanitis (Norba) hiurai feeds on the abaxial surface of leaves of Machilus thunbergii (present study. Adults were collected in almost all seasons (Takeya 1963; Miyamoto 1964a, 1964b; present study). Nymphs were collected in April and November (present study).