Tingis (Tropidocheila) matsumurai Takeya, 1962

(Figs. 1A, 2 A–C, 3A, B, 4A, 5A, B, 6A–C)

Tingis (Tropidocheila) matsumurai Takeya, 1962: 60 . Holotype: macropterous ♀, Japan: Honshu, Kii, Mt. Koya; ELKU (http:// konchudb.agr.agr.kyushu-u.ac.jp/elkutype/exec/refile.cgi?&lang=jp&no=0112&tax= Tingis %20( Tropidocheila)%20matsu murai%20Takeya).

References. Ishihara et al. (1974: 73) (distribution); Miyamoto & Yasunaga (1989: 168) (checklist: Japan); Péricart & Golub (1996: 68) (checklist: Palaearctic); Maehara (2010: 133) (distribution); Yamada & Tomokuni (2012: 210) (monograph); Yano et al. (2013: 26) (distribution); Maehara (2014: 62) (distribution); Komatsu (2016: 101) (distribution); Yamada & Ishikawa (2016: 435) (checklist: Japan); Ahn et al. (2018: 70) (monograph); Hayashi & Kadowaki (2018: 299) (distribution); Ito & Sasaki (2018: 21) (distribution); Ikeda et al. (2019: 60) (distribution); Nishida (2019: 191) (distribution); Okochi (2019: 3) (distribution); Cho et al. (2020: 743) (checklist: Korea).

Specimens examined. Non-types (macropterous 36 ♂♂ 28 ♀♀ 1 terminalia missing), JAPAN: Honshu: Tochigiken, Utsunomiya-shi, Tsuruta-machi, Tsuruta Swamp, 29.iv.2018, leg. T. Saeki (11 ♂♂ 2 ♀♀ 1 terminalia missing, TUA) ; Tochigi-ken, Tochigi-shi, Ohira-machi, Shimominagawa, 24.vi.2020. leg. S. Maehara (4 ♂♂ 1 ♀, TUA) ; Tôkyô-to, Ôme-shi, Mitakesan, Mt. Mitake-yama, N 3547.039' E13909.008', Alt. 870 m, 14.v.2016, leg. S. Shimamoto (1 ♂, TUA) ; Kanagawa-ken, Sagamihara-shi, Minami-ku, Asamizodai, 3532'02.5" N 13924 '43.3"E, 16.xii.2017, leg. J. Souma (1 ♂, TUA) ; Kanagawa-ken, Yamato-shi, Kamisohyagi, 3528'39.7" N 13926 '40.3"E, 3.ix.2018, leg. J. Souma (1 ♂, TUA) ; Kanagawa-ken, Atsugi-shi, Funako, Atsugi Camp. of TUA, N 35.431171 E 139.347602, 23.iv.2014, leg. S. Shimura (1 ♂, TUA) ; Kanagawa-ken, Atsugi-shi, Funako, 3525'58.3" N 13920 '48.6"E, 25.v.2017, leg. J. Souma (1 ♂, TUA) ; Kanagawa-ken, Atsugi-shi, Ono, Komachi-ryokuchi, 7.iv.2006, leg. T. Ishizaki (1 ♂, TUA) ; Kanagawa-ken, Atsugi-shi, Iiyama, Shiroyama, 3528'17.9" N 13917 '54.8"E, 15.iv.2017, leg. H. Kidokoro (1 ♂, TUA) ; Kanagawa-ken, Isehara-shi, Mt. Oyama, 3526'26.9" N 13913 '50.0"E, 27.iii.2018, leg. J. Souma (1 ♂, TUA) ; Kanagawa-ken, Hadano-shi, Minamiyana, 3522'30.5" N 13915 '01.1"E, 26.iv.2018, leg. T. Saeki (1 ♂, TUA) ; Kanagawa-ken, Hadano-shi, Minoge, Harutake-rindô, 4.v.2009, leg. T. Ishikawa (2 ♂♂ 7 ♀♀, TUA) ; Kanagawaken, Hadano-shi, Tochikubo, Near Mt. Zukkô Yama, N 3521.730' E13911.211', Alt. 230 m, 27.ii.2017, leg. S. Shimamoto (2 ♀♀, TUA) ; Kanagawa-ken, Ashigarakami-gun, Yamakita-machi, Yaga, 30.iv.2018, leg. J. Souma (3 ♀♀, TUA) ; Kanagawa-ken, Ashigarakami-gun, Yamakita-machi, Nakagawa, 35°26'45.5"N 139°02'52.4"E, 30.vi.2018, leg. J. Souma (1 ♂, TUA) ; Kanagawa-ken, Odawara-shi, Kamisoga, 3519'12.9" N 13911 '20.8"E, 20.iii.2019, leg. J. Souma (1 ♂ 3 ♀♀, TUA) . Kesennuma-Oshima Island: Miyagi-ken, Kesennuma-shi, Oshima, 1920. v.1994, leg. Asia C. (1 ♀, TUA) . Sado Island: Niigata-ken, Sado-shi, Kanaishinbo, Osado Sky Line, 5.vi.2020, leg. T. Saeki (1 ♂ 1 ♀, TUA) ; Niigata-ken, Sado-shi, Shimoaikawa, Osado Sky Line, 6.vi.2020, leg. T. Saeki (1 ♂, TUA) . Shikoku: Ehime, Shigenobu-chô, Kanbayashi, 9.vii.2003, leg. T. Kurihara (1 ♀, TUA) . Kyushu: Fukuoka Pref., Fukuoka-shi, Nishi-ku, Motooka, Kyushu University, Ito Camus, 25.iv.2020, leg. N. Tsuji (1 ♂, ELKU) ; as above but 27.iv.2020 (1 ♀, ELKU); as above but 15.vi.2020 (4 ♂♂ 4 ♀♀, ELKU); as above but 3335'50.3" N 13012 '52.4"E, 17.vi.2020, leg. J. Souma (1 ♀, ELKU); Fukuoka-ken, Tagawa-gun, Soeda-machi, Ochiai, Fukakurakyo, 19.iv.2019, leg. Y. Tamadera (1 ♀, ELKU) ; Saga Pref., Fuji-ch, Hakuzan Dam, 6.vi.2015, leg. N. Tsuji (1 ♂, ELKU) (Fig. 7).

Diagnostic characters. Recognized among other species of Tropidocheila by a combination of the following characters: general color black (Figs. 1A, 6A, B); pubescence on dorsum shorter than diameter of compound eye; head with five spines (Fig. 4A); a pair of frontal spines reaching beyond tip of clypeus; median spine reaching bases of frontal spines; a pair of occipital spines reaching middle of compound eyes; rostrum reaching middle of metasternum (Fig. 2A); anterior margin of hood curved outward throughout its length; lateral carinae of pronotum distinct on pronotal disc; paranotum ridge-shaped, subvertical, with a single row of areolae in anterior half, without areolae in posterior half (Fig. 2B); costal area of hemelytron with 2 rows of areolae throughout its length, narrower than subcostal area at widest part of each (Fig. 2C); subcostal area with 3 rows of areolae at widest part; discoidal area with 9 rows of areolae at widest part; and sutural area with 11–12 rows of areolae at widest part.

Description of genitalia. Pygophore (Figs. 3A, 5B) compressed dorsoventrally, hexagonal in ventral view, elevated at center of venter, concave at anterior margin of dorsum. Paramere (Fig. 5A) expanded in middle part, curved inward in apical part; outer and inner margins covered with pubescence in middle part. Female terminalia (Fig. 3B) pentagonal in ventral view, covered with pubescence; ovipositor with ovivalvula at base.

Distribution. Japan (Honshu, Kesennuma-Oshima Island, Sado Island, Dogo Island, Nakanoshima Island, Shikoku, Kyushu, Fukue Island); Korea.

In Japan, Tingis (Tropidocheila) matsumurai inhabits deciduous forests of temperate climate.

Host plant. In Japan, Cornus controversa Helms. ex Prain and C. macrophylla (Cornaceae) have been confirmed as host plants of Tingis (Tropidocheila) matsumurai (Maehara 2010, 2014; Yamada & Tomokuni 2012; Komatsu 2016; Ito & Sasaki 2018). In Korea, C. controversa has been confirmed as a host plant of this species (Ahn et al. 2018).

Biology. Although many tingids generally feed on the abaxial surface of leaves of their respective host plants (Schuh & Weirauch 2020), several individuals of Tingis (Tropidocheila) matsumurai were collected from flowers and fruits of the host tree Cornus controversa and C. macrophylla in Japan (Maehara 2010, 2014) (Fig. 6C), suggesting that this lace bug readily feed on the flowers and fruits.

In Japan, adults were collected in almost all seasons; nymphs were collected in June; the overwintering stage is adult (Takeya 1962; Ishihara et al. 1974; Maehara 2010, 2014; Yamada & Tomokuni 2012; Yano et al. 2013; Komatsu 2016; Hayashi & Kadowaki 2018; Ito & Sasaki 2018; Ikeda et al. 2019; Nishida 2019; Okochi 2019; present study).