Integrative taxonomy of the Lauraceae - feeding species of the genus Stephanitis (Hemiptera, Heteroptera, Tingidae) from Japan Author Souma, Jun https://orcid.org/0000-0002-2238-5015 Entomological Laboratory, Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka, Japan & Research Fellowship for Young Scientists (DC 1), Japan Society for the Promotion of Science, Tokyo, Japan kodokusignal@gmail.com text Deutsche Entomologische Zeitschrift 2022 2022-12-15 69 2 219 281 http://dx.doi.org/10.3897/dez.69.89864 journal article http://dx.doi.org/10.3897/dez.69.89864 1860-1324-2-219 BFE2BE4759E9450A807079B702A38625 9EE9E317E8195E3A94756A0DBBB1C35B Stephanitis (Stephanitis) tomokunii sp. nov. [Japanese name: Tomokuni-gunbai] Figs 3F , 5F , 8F , 10G , 12F , 14G , 16F , 18F , 20D , 22D , 24D , 26F , 28F , 30H , 32H , 39 , 42K, L Stephanitis (Stephanitis) tabidula Horvath , 1912: Takeya (1963 : 42) (distribution: part); Tomokuni and Ishikawa (2002 : 170) (distribution); Yamada and Tomokuni (2012 : 208) (monograph: part); Yamada and Ishikawa (2016 : 434) (checklist: Japan). Misidentifications. Type series. Holotype (♂, ELKU), "[JAPAN]: Izu Isls., Hachijo, Is., Mitsune" [=JAPAN: Izu Islands (southern part): Hachijo Island: Mitsune (approximate coordinates: 33°07'16.3"N , 139°48'21.6"E )], 17.v.2021, leg. J. Souma. Paratypes (66 ♂♂ 134 ♀♀), JAPAN: Izu Islands: (southern part): Miyake Island: Ako, 15.v.1999, leg. T. Kishimoto (1 ♂ 1 ♀); Izu, 12.v.2018, leg. T. Ishikawa (8 ♂♂ 12 ♀♀, TUA); Tosa For. Rd., 12.v.2018, leg. J. Souma (2 ♂♂ 8 ♀♀, TUA); as above but leg. T. Saeki (1 ♂ 1 ♀, TUA); Sannomiya For. Rd., 12.v.2018, leg. J. Souma (20 ♂♂ 18 ♀♀, TUA); Nanto For. Rd., 13.v.2018, leg. J. Souma (1 ♂ 2 ♀♀, TUA). Hachijo Island: "八丈" [= Hachijo Island], Arakawa, "20/V/909" [= 20.v.1909] (1 ♂ 2 ♀♀, ELHU) (Fig. 39 ); Minemura, 7.viii.1948, leg. Fujiyama (1 ♀, ELKU); Kaminato, 24.v.1949, leg. I. Fujiyama (1 ♀, NIAES); Mitsune Beach, 28.v.1949, leg. T. Aoki (1 ♀, NIAES); Noboryo Pass, 11.viii.1949, leg. I. Fujiyama (1 ♀, NIAES); as above but alt. 350 m, 5.vii.2001, leg. M. Tomokuni (3 ♂♂ 3 ♀♀, NSMT); 19.vii.1957, leg S. Hisamatsu (1 ♀, NSMT); as holotype but 16.v.2021 (4 ♀♀, ELKU); Mt. Miharayama, alt. 200-560 m, 2.vii.2001, leg. M. Tomokuni (1 ♂ 2 ♀♀, NSMT); Mt. Hachijo-fuji, alt. 250-530 m, 4.vii.2001, leg. M. Tomokuni (3 ♀♀, NSMT); as above but alt. 560-850 m, 3.vii.2001 (3 ♀♀, NSMT); as holotype (4 ♂♂ 5 ♀♀, ELKU); as holotype but 18.v.2021 (10 ♂♂ 30 ♀♀, ELKU); as holotype but 19.v.2021, leg. J. Souma (1 ♂ 1 ♀, ELKU; 5 ♂♂ 16 ♀♀, TUA); as holotype but 20.v.2021 (6 ♂♂ 14 ♂♂, TUA); Mistune, Mihara For. Rd., 17.v.2021, leg. J. Souma (2 ♂♂ 7 ♀♀, ELKU); Sueyoshi, 17.v.2021, leg. J. Souma (1 ♂ 2 ♀♀, ELKU). Three paratypes collected in 1909 are deposited in Matsumura's collection. A single paratype collected in 1948 and 12 paratypes collected in 3-5.vii.2001, were recorded as " Stephanitis tabidula " in previous studies ( Takeya 1963 ; Tomokuni and Ishikawa 2002 ; Yamada and Tomokuni 2012 ). Figure 39. Paratypes of Stephanitis (Stephanitis) tomokunii sp. nov. deposited in Matsumura's collection of ELHU and their 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 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 ). Diagnosis. Stephanitis (Stephanitis) tomokunii 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 8F , 10G , 12F , 14G , 16F , 18F , 20D , 22D , 24D ); calli dark brown; body in male 2.3 times (in female 2.1 times) as long as maximum width across hemelytra (Figs 3F , 5F ); rostrum not reaching metasternum; pronotum tricarinate (Fig. 26F ); 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, posterior margin not extending to middle of pronotal disc; median carina of pronotum with 1-2 rows of areolae at highest part; pronotal disc; paranotum less erect, narrowed posteriorly, 3 rows of areolae at widest part, anterolateral angle slightly protruding anteriad, outer margin gently curved inwards at posterolateral angle, maximum height shorter than height of eye (Fig. 28F ); apices of hemelytra close to each other in rest; costal area with 3 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, posterior margin slightly emarginate in middle part (Fig. 30H ); and paramere stout, weakly curved inward at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 32H ). 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 ). Description. Male. Head, pronotal disc, marking on hemelytra and ventral surface in various shades of brown; calli dark brown; eye dark red; areolae of pronotum and hemelytron transparent; hood pale; pronotal disc opaque; pubescence on body yellowish (Figs 3F , 8F , 12F , 16F , 20D , 22D ). Body 2.3 times as long as maximum width across hemelytra (Fig. 3F ). Head (Figs 8F , 12F , 20D , 26F ) glabrous; 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 amongst antennal segments; segment III longest amongst antennal segments; segment IV cylindrical, longer than segment I. Bucculae closed to each other at anterior ends, with 2 rows of areolae throughout its length. Rostrum not reaching metasternum. Pronotum (Figs 8F , 12F , 26F , 28F ) 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, as wide as vertex at widest part, not covering eye, as high as median carina of pronotum at highest part, posterior margin not extending to middle of pronotal disc, 4 rows of areolae at highest part, dorsal margin slightly arched. Median carina straight, extending to apex of posterior process, 1-2 rows of areolae at highest part, dorsal margin arched. Calli smooth. Paranotum less erect, narrowed posteriorly, 3 rows of areolae at widest part, with anterolateral angle slightly protruding anteriad, outer margin gently curved inwards at posterolateral angle, maximum height shorter than height of eye. Posterior process triangular, obtuse at apex. 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 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 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. Hemelytron (Fig. 16F ) 2.5 times as long as its maximum width, extending beyond apex of abdomen, glabrous; 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 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. 12F ) smooth in anterior part, coarsely punctate in posterior part. Ostiolar peritreme oblong. Sternal laminae (Fig. 20D ) 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. 3F ) smooth, densely covered with pubescence; femora thickest at middle. Figure 46. Collection sites of Stephanitis (Norba) hayashii sp. nov. and S. (N.) hiurai used in the present study. Red-filled areas = localities; circles = small isolated islands. 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. 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. Abdomen oblong in dorsal and ventral views. Pygophore (Figs 22D , 30H ) compressed dorsoventrally, semicircular in ventral view, elevated at centre of venter, posterior margin slightly emarginate in middle part, covered with pubescence. Paramere (Fig. 32H ) 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 3.1-3.4 mm; maximum width across hemelytra 1.4-1.5 mm; length of antennal segments I to IV 0.2 mm, 0.1 mm, 1.2-1.3 mm and 0.6 mm, respectively; pronotal length 1.2-1.4 mm; pronotal width across paranota 0.8-0.9 mm; hemelytral length 2.4-2.6 mm; maximum width of hemelytron 1.0-1.1 mm. Female. General habitus very similar to that of male (Figs 5F , 10G , 14G , 18F , 24D ) except for the following characters: body 2.1 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; maximum width across subcostal area wider than in male, with 2-3 rows of areolae at widest part; and apical part of abdomen pentagonal in ventral view. Measurements (n = 20). Body length with hemelytra 3.3-3.6 mm; maximum width across hemelytra 1.5-1.7 mm; length of antennal segments I to IV 0.2 mm, 0.1 mm, 1.0-1.1 mm and 0.6 mm, respectively; pronotal length 1.3-1.5 mm; pronotal width across paranota 0.9-1.0 mm; hemelytral length 2.5-2.7 mm; maximum width of hemelytron 1.0-1.2 mm. Remarks. The partial COI gene pairwise sequence distances between Stephanitis (Stephanitis) tomokunii sp. nov. and S. (S.) tabidula are only 0.002645-0.007978 (Suppl. material 3) and both species are very similar in general habitus. In fact, S. (S.) tomokunii sp. nov. was misidentified as S. (S.) tabidula in previous studies ( Takeya 1963 ; Tomokuni and Ishikawa 2002 ; Yamada and Tomokuni 2012 ). However, the former is easily distinguished from the latter by the following characters: body in male 2.3 times (in female 2.1 times) as long as maximum width across hemelytra (in male 2.1 times and in female 2.0 times in S. (S.) tabidula ) (Figs 3C, D, F , 5C, D, F ); paranotum less erect (more erect in S. (S.) tabidula ), narrowed posteriorly (slightly narrowed in S. (S.) tabidula ), with anterolateral angle slightly protruding anteriad (protruding in S. (S.) tabidula ), with maximum height shorter than height of eye (longer in S. (S.) tabidula ) (Figs 8C, D, F , 10C-E, G , 12C, D, F , 14C-E, G , 26C, D, F , 28 , D, F); and apex of paramere weakly curved inwards (strongly curved in S. (S.) tabidula ) (Fig. 32E, F, H ). Distribution. Japan (Izu Islands (southern part): Miyake Island, Hachijo Island) (Fig. 48 ) ( Takeya 1963 ; Tomokuni and Ishikawa 2002 ; Yamada and Tomokuni 2012 ; present study). Stephanitis (Stephanitis) tomokunii sp. nov. inhabits the laurilignosa in a temperate climate of the southern part of the Izu Islands, which is in the Palaearctic Region. Etymology. The new species is named in honour of Masaaki Tomokuni, a Japanese heteropterist who collected some of the paratype specimens. Host plants. Machilus thunbergii , "Tabunoki" (Fig. 44F ) (present study). Stephanitis (Stephanitis) tomokunii sp. nov. feeds only on this lauraceous tree and is monophagous. Biology. Stephanitis (Stephanitis) tomokunii sp. nov. feeds on the abaxial surface of leaves of Machilus thunbergii (present study). Adults were collected in May, July and August ( Takeya 1963 ; Tomokuni and Ishikawa 2002 ; present study); the nymph and overwintering stage are unknown.