Physatocheila fieberi ( Scott, 1874 )

Souma, Jun & Ishikawa, Tadashi, 2022, Taxonomic review of the tingine genera Cysteochila, Hurdchila, Physatocheila, and Xynotingis from Japan, with description of a new genus (Hemiptera: Heteroptera: Tingidae), Zootaxa 5150 (1), pp. 1-42 : 20-23

Physatocheila fieberi ( Scott, 1874)

( Figs. 8E, F , 9E , 10E , 11E , 17G, H , 18G, H )

Monanthia fieberi Scott, 1874: 291 . Syntypes: macropterous 1 ♂ 1♀, Japan; BMNH.

Physatocheila fieberi: Lethierry & Severin (1896: 21) (new combination).

Physatocheila monstrosa (non Scott, 1874): Takeya (1951: 20) (checklist: eastern Asia); Takeya (1953a: 173) (redescription); Hasegawa (1960: 45) (distribution). Misidentifications ( Drake & Maa 1954: 112).

Cysteochila fieberi: Drake & Maa (1954: 112) (new combination); Drake & Ruhoff (1965a: 168) (catalog); Miyamoto (1965: 92) (distribution); Lee (1967: 107) (distribution); Lee (1969: 198) (nymph, male genitalia); Tomokuni (1979: 137) (distribution); Tomokuni (1987: 117) (distribution); Tomokuni (1988: 652) (distribution); Miyamoto & Yasunaga (1989: 167) (checklist: Japan); Hayashi & Higashikawa (1997: 36) (distribution); Tomokuni (2006a: 60) (distribution); Miyamoto (2008: 158) (distribution); Yazaki (2009: 31) (distribution); Yamada & Tomokuni (2012: 192) (monograph); Nozaki & Nozaki (2013: 30) (distribution); Yano et al. (2013: 25) (distribution); Komatsu (2016: 99) (distribution); Yamada & Ishikawa (2016: 430) (checklist: Japan); Ito & Sasaki (2018: 18) (distribution); Okochi (2019: 1) (distribution).

Physatocheila fieberi: Péricart (1982: 363) (new combination); Péricart (1985a: 40) (distribution); Péricart & Golub (1996: 55) (checklist: Palaearctic); Kwon et al. (2001: 194) (checklist: Korea); Guilbert (2015: 538) (distribution); Guilbert & Guidoti (2018: 292) (distribution); Cho et al. (2020: 741) (checklist: Korea).

Cysteochila monstrosa (non Scott, 1874): Nakamura (2014: 357) (distribution). Misidentification.

Specimens examined. Non-types (70 ♂♂ 70 ♀♀ 1 nymph), JAPAN: Honshu: Yamagata-ken , Asahi-machi , Shirakura , 3817'30" N 14000 '60"E, 500 m alt., 23.viii.2017, leg. Y. Fukuda (1 ♀, TUA) ; Gunma Pref., Mt. Akagi-san , 3.vi.1997, leg. T. Ishikawa (1 ♀, TUA) ; Gunma-ken, Niiharu-mura, Amemi-rindo , 4.v.1998, leg. S. Arai (1 ♂, TUA) ; Gunma-ken, Tone-gun, Katashina-mura , Hanasaka , 15.vi.2020, leg. T. Saeki (3 ♂♂ 2 ♀♀, TUA) ; Saitama Pref., Higashi-matsuyama City, Sakagawa-rind , 21.v.1998, leg. S. Arai (1 ♂, TUA) ; Saitama Pref., Naguri-mura, Mt. Warabi-san , 21.v.1998, leg. S. Arai (1 ♀, TUA) ; Saitama-ken, Chichibu-shi, Ôtaki , Saikaigaid , 9.vi.2018, leg. K. Okuda (2 ♂♂ 2 ♀♀, TUA) ; as above but 14.vii.2018 (1 ♀, TUA) ; Tokyo-to, Hachioji-shi, Uratakao-machi , Kogesawa-rindo , 3538'37.5" N 13914 '29.4"E, 20.vi.2017, leg. J. Souma (1 ♂ 2 ♀♀, TUA) ; as above but 27.ix.2017 (1 ♂ 1 ♀, TUA) ; Kanagawa-ken, Odawara-shi, Hayakawa , Sarusawa-rindo , 20.ix.2017, leg. J. Souma (4 ♀♀, TUA) ; Kanagawa-ken, Ashigarakami-gun, Yamakita-machi , Kurokura , 3525'54.9" N 13904 '30.1"E, 6.ix.2018, leg. J. Souma (1 ♂ 3 ♀♀, TUA) ; Nagano-ken, Matsumoto-shi, Azumi , 31.v.2017, leg. K. Okuda (1 ♀, TUA) ; Nagano-Pref., Minamisaku-gun, Minamimaki-mura , 13.ix.2021, leg. K. Okuda et al. (1 ♂ 3 ♀♀, TUA) ; Yamanashi-ken, Kosuge-mura, Shiraitonotaki , 3545'15" N 13853 '58"E, 850 m alt., 8.ix.2018, leg. Y. Fukuda (1 ♂, TUA) ; Shizuokaken, Kakegawa-shi, Kamihijikata , 18.vi.2017, leg. J. Souma (1 ♂, TUA) ; Shizuoka-ken, Shizuoka-shi, Aoi-ku , Hatori , 19.vi.2017, leg. J. Souma (1 ♂ 2 ♀♀, TUA) ; Shizuoka-ken, Shizuoka-shi, Suruga-ku , Maruko , Sakasagawa , 19.vi.2017, leg. J. Souma (2 ♂♂ 2 ♀♀, TUA) ; Aichi-ken, Kitashitara-gun, Shitara-cho , Taguchi , 17.vi.2017, leg. J. Souma (4 ♂♂ 4 ♀♀, TUA) ; Shiga Pref., Taga-cho, Kawachi , 3.x.1998, leg. T. Ishikawa (2 ♂♂ 2 ♀♀, TUA) ; Yamashiro, Kyoto – Kifune , 7.ix.1952, leg. E. Nakanishi (1 ♂, KUM) . Sado Island: Niigata-ken, Sado-shi , Shimoaikawa , Osado Sky Line , 6.vi.2020, leg. T. Saeki (1 ♂, TUA) . Kyushu: Fukuoka, Mt. Hikosan , 22–26.viii.1961, leg. S. Miyamoto (1 ♂ 1 ♀, KUM) ; Ôita Pref., Yuhu-shi, Yuhuin-ch, Kawakami , 10.viii.2016, leg. R. Ito (1 ♀, TUA) . Okinoshima Island: Chikuzen , 25–28.vii.1958, leg. Hirashima, Murakami, Y. Miyatake (1 ♂, KUM) . Ryukyu Islands: Yaeyama Islands: Ishigakijima , Nanbadake , 24.xi.1960, leg. K. Yasumatsu (1 ♀, ELKU) ; Ishigaki Is. , Yonehara, 11.iii.1973, leg. H. Hasegawa (2 ♂♂, NIAES) ; Yaeyama Isls., Ishigaki Is., Tomino , 17.x.1990, leg. M. Hayashi (1 ♂ 2 ♀♀, TUA) ; as above but 17.xi.1992 (9 ♂♂ 10 ♀♀, TUA) ; as above but 20.iv.2005 (1 ♂, TUA) ; as above but 14.xi.2018, leg. J. Souma (7 ♂♂ 2 ♀♀ 1 nymph, TUA) ; Yaeyama Isls., Ishigaki Is., Kainan , 3.iv.1996, leg. M. Hayashi (1 ♂ 1 ♀, TUA) ; Yaeyama Isls., Ishigaki Is., Chiteibaru , 27.vi.1997, leg. M. Hayashi (1 ♂, TUA) ; Yaeyama Isls., Ishigaki Is., Hirakubo , 19.iv.2005, leg. M. Hayashi (1 ♀, TUA) ; Yaeyama Isls., Ishigaki Is., Tomino–Yonehara , 22.vi.1997, leg. M. Hayashi (1 ♂, TUA) ; Yaeyama Isls., Ishigaki Is., Takada , 21.ii.2006, leg. M. Hayashi (1 ♀, TUA) ; Yaeyama Isls., Ishigaki Is., Yonehara , 12.vi.2008, leg. M. Hayashi (2 ♂♂ 1 ♀, TUA) ; Yaeyama Isls., Ishigaki Is., Hoshino , 1.x.1993, leg. M. Hayashi (1 ♂, TUA) ; Yaeyama Isls., Ishigaki Is., Okawa , 2421'42.2" N 12409 '52.5"E, 11.xi.2018, leg. J. Souma (8 ♂♂ 5 ♀♀, TUA) ; as above but 16.xi.2018 (1 ♂, TUA) ; Yaeyama Isls., Ishigaki Is., Mt. Nosokodake , 2428'58.1" N 12414 '53.7"E, 14.xi.2018, leg. J. Souma (7 ♂♂ 11 ♀♀, TUA) ; Yaeyama Isls., Ishigaki Is., Itona , 2427'49.4" N 12413 '36.3"E, 14.xi.2018, leg. J. Souma (1 ♀, TUA) ; Okinawa-ken, Ishigaki-jima Is., Yonehara , 18.x.2020, leg. S. Imada (1 ♂, ELKU) ; Iriomote I., Komi , 10.xi.1984, leg. M. Tomokuni (1 ♂, NSMT) . TAIWAN: Yilan County, Yuanshan Township, Huxi Village , 16.iii.1999, leg. T. Mizusawa (1 ♂ 3 ♀♀, TUA) . A single male collected from Kyushu in 1961 has been recorded as Cysteochila fieberi by Miyamoto (1965). A single specimen from the Ryukyu Islands deposited in ELKU is labeled “ Cysteochila fieberi ishigakiensis Takeya, 1962 ” (manuscript name). A single nymph from the Ryukyu Islands is in a poor condition and is therefore not described in the present article.

Diagnosis. Recognized among other species of Physatocheila by a combination of the following characters: general color brown ( Figs. 8E, F , 9E , 10E , 11E ); median spine on head extending beyond bases of frontal spines; rostrum reaching posterior margin of metasternum; pubescence on pronotum less than 0.5 times as long as diameter of compound eye; lateral carinae of pronotum close to each other in anterior part, concealed by paranotum in anterior part; paranotum completely covering pronotal disc in anterior part, not touching each other, not concealing median carina of pronotum, widened posteriorly, bulged upward in posterior part, forming a cyst; cyst of paranota concave in posterior part; outer margin of paranotum gently curved outward in its entire length; costal area of hemelytron wider than subcostal area at widest part, less than 0.5 times as wide as discoidal area at widest part, with 2 rows of areolae in its entire length; and subcostal area subvertical, with 2 rows of areolae in its entire length.

Description of genitalia. Pygophore ( Figs. 10E , 17G, H ) compressed dorsoventrally, hexagonal in ventral view, strongly concave at anterior margin of dorsum, elevated at center of venter, smooth on surface, irregularly punctate in middle part of dorsum. Paramere ( Fig. 18G, H ) expanded in middle part, curved inward in apical part; outer and inner margins covered with pubescence in middle part. Female terminalia ( Fig. 11E ) pentagonal in ventral view, covered with pubescence.

Remarks. Physatocheila fieberi was transferred from Cysteochila to Physatocheila by Péricart (1982) and was treated as a member of Cysteochila by Japanese researchers until quite recently (Tomokuni 1987, 2006a; Miyamoto 2008; Yamada & Tomokuni 2012; Yamada & Ishikawa 2016). This species certainly belongs to Physatocheila because of the presence of generic characteristics, as described in the Diagnosis section.

Physatocheila fieberi closely resembles P. chiniana comb. nov. in the shape of the hemelytron but can be distinguished from it by the following characteristics: rostrum reaching posterior margin of metasternum ( Fig. 9E ); paranota not touching each other, touching median carina of pronotum ( Fig. 8E, F ); cyst of paranotum concave in posterior part.

In Japan, P. fieberi has two infraspecific variants despite the lack of distinct morphological differences in pygophores ( Fig. 17G, H ) and parameres ( Fig. 18G, H ). One of these variants, which matches the photographs (https:// data.nhm.ac.uk/media/c0138d47-da95-4d37-9f4f-f942ed55a173; https://data.nhm.ac.uk/media/a457f7de-7d63- 469f-a270-1c98a4aa45de) and illustration ( Péricart 1985a) of the syntypes, is present in a population distributed in Japan proper, and the other variant, which is newly recognized in the present article, is distributed in a population on the Ryukyu Islands. Individuals in the first population ( Fig. 8E ) differ from those in the second one ( Fig. 8F ) in having a larger and broader body. The infraspecific variants of P. fieberi in eastern Asia need to be clarified first.

Distribution. Japan (Hokkaido, Honshu, Sado Island, Shikoku, Kyushu, Okinoshima Island, Tsushima Island, Ryukyu Islands: Ishigaki Island, Iriomote Island), Bhutan, China, India, Korea, Taiwan, Thailand, Vietnam.

Physatocheila fieberi is recorded from Sado Island and the Ryukyu Islands for the first time.

Host plant. In Japan, Korea, and Taiwan, P. fieberi occurs on the following Boehmeria species (Urticaceae): B. gracilis C.H.Wright [= B. tricuspis (Hance) Makino var. unicuspis Makino ] ( Kwon et al. 2001; Okochi 2019); B. japonica (L.f.) Miq. [= B. longispica Steud. var. dura (Satake) Satake ] ( Miyamoto 1965, 2008; Kwon et al. 2001; Yamada & Tomokuni 2012); B. nivea (L.) Gaudich. var. nivea (present study); B. nivea (L.) Gaudich. var. concolor Makino [= B. nipononivea Koidz. var. concolor (Makino) Ohwi ] ( Kwon et al. 2001); B. platanifolia (Maxim.) Franch. et Sav. ex C.H.Wright ( Kwon et al. 2001) ; B. silvestrii (Pamp.) W.T.Wang ( Hasegawa 1960; Okochi 2019); B. spicata (Thunb.) Thunb. (Takeya 1953a; Hasegawa 1960; Miyamoto 1965, 2008; Yamada & Tomokuni 2012; Komatsu 2016); and Boehmeria sp. (Tomokuni 2006a) .

Biology. A number of individuals of Physatocheila fieberi were collected from the leaves of the host plants B. spicata and B. nivea var. nivea in Japan proper and the Ryukyu Islands, respectively (present study). Therefore, this species appears to be folivorous, similar to many tingids ( Schuh & Weirauch 2020).

In Japan proper, adults were observed in almost all seasons (Takeya 1953a; Hasegawa 1960; Tomokuni 1979, 1987, 1988; Hayashi & Higashikawa 1997; Yazaki 2009; Yamada & Tomokuni 2012; Nozaki & Nozaki 2013; Yano et al. 2013; Maehara 2014; Komatsu 2016; Ito & Sasaki 2018; Okochi 2019; present study), but it is unknown when nymphs can be collected. In Japan proper, adults were collected in February ( Hasegawa 1960), suggesting that the adult stage is the overwintering stage in P. fieberi , similar to that in many other tingids in temperate regions ( Schuh & Weirauch 2020). In the Ryukyu Islands, adults were observed from February to April, and in June, October, and November, and nymphs were collected in November (present study).

Figures

FIGURE 8. Nine Physatocheila species from Japan. Dried specimens, dorsal view: P. chiniana comb. nov. (A); P. consueta comb. nov. (B); P. distinguenda (C); P. euthenia comb. nov. (D); P. fieberi from Japan proper (E); P. fieberi from the Ryukyu Islands (F); P. miyatakei (G); P. nigrintegerrima (H); P. orientis (I); P. veteris (J). Scale bar: 1.0 mm.

FIGURE 9. Rostra and sternal laminae of nine Physatocheila species from Japan, ventral view: P. chiniana comb. nov. (A); P. consueta comb. nov. (B); P. distinguenda (C); P. euthenia comb. nov. (D); P. fieberi (E); P. miyatakei (F); P. nigrintegerrima (G); P. orientis (H); P. veteris (I). Scale bars: 0.2 mm.

FIGURE 10. Male terminalia of nine Physatocheila species from Japan, ventral view: P. chiniana comb. nov. (A); P. consueta comb. nov. (B); P. distinguenda (C); P. euthenia comb. nov. (D); P. fieberi (E); P. miyatakei (F); P. nigrintegerrima (G); P. orientis (H); P. veteris (I). Scale bars: 0.2 mm.

FIGURE 11. Female terminalia of nine Physatocheila species from Japan, ventral view: P. chiniana comb. nov. (A); P. consueta comb. nov. (B); P. distinguenda (C); P. euthenia comb. nov. (D); P. fieberi (E); P. miyatakei (F); P. nigrintegerrima (G); P. orientis (H); P. veteris (I). Scale bars: 0.2 mm.

FIGURE 17. Pygophores of eleven species of Cysteochila and its close relatives from Japan, dorsal view: C. abundantis (A); Limnostatua lewisi comb. nov. (B); Physatocheila chiniana comb. nov. (C); P. consueta comb. nov. (D); P. distinguenda (E); P. euthenia comb. nov. (F); P. fieberi from Japan proper (G); P. fieberi from the Ryukyu Islands (H); P. miyatakei (I); P. nigrintegerrima (J); P. orientis (K); P. veteris (L). Scale bars: 0.1 mm.

FIGURE 18. Parameres of eleven species of Cysteochila and its close relatives from Japan, dorsal view: C. abundantis (A); Limnostatua lewisi comb. nov. (B); Physatocheila chiniana comb. nov. (C); P. consueta comb. nov. (D); P. distinguenda (E); P. euthenia comb. nov. (F); P. fieberi from Japan proper (G); P. fieberi from the Ryukyu Islands (H); P. miyatakei (I); P. nigrintegerrima (J); P. orientis (K); P. veteris (L). Scale bars: 0.1 mm.