Comibaena signifera, SUBARGENTARIA (OBERTHUR, 1916)

Han, Hongxiang, Galsworthy, Anthony C. & Xue, Dayong, 2012, The Comibaenini of China (Geometridae: Geometrinae), with a review of the tribe, Zoological Journal of the Linnean Society 165 (4), pp. 723-772 : 751-754

COMIBAENA SIGNIFERA SUBARGENTARIA ( OBERTHÜR, 1916) ( FIGS 24 , 78 , 128 , 190 )

Phorodesma subargentaria Oberthür, 1916: 105 , pl. 387, figure 3265. Syntypes 4♂, Oriental frontier of Tibet. (ZFMK)

Comibaena signifera subargentaria: Prout, 1933: 93 .

Comibaena subargentaria: Prout, 1935: 12 , pl. 2g.

Type material examined: China: 1♂, syntype, Frontière orientale du Thibet, Chasseurs indigènes du P. Déjean, 1906 ( ZFMK) ; 1♂, syntype, Ta-Lou, Yuin- Kin, 1899, Chasseurs indigènes ( ZFMK) .

Additional material examined: China: 25♂ 4♀, southern China, April to September ( IZCAS, BMNH)

Distribution: China (Zhejiang, Fujian, Guangxi, Sichuan, Yunnan, Tibet).

COMIBAENA TAKASAGO OKANO, 1960

( FIGS 25 , 79 , 129 , 191 )

Comibaena takasago Okano, 1960: 9 , pl. 8, figure 5; text–figures 1, 2. Holotype ♂, Formosa [ Taiwan] (central): Nantow-hsien, Jen-ai-hsiang. (BMNH)

Diagnosis: The wing pattern of Comibaena takasago is unique within Comibaena in having a band-like submarginal line on the hindwing, which is brown, with white inside, and expanded on the costa and the inner margin, and is located more proximally than in other congeners.

The male genitalia of Comibaena takasago are similar to those of Comibaena flavicans and Comibaena chalybeata ( India) , and share the truncate, spinose apex of the costal lobe. The ventral distal tooth is larger and of a similar length in Comibaena takasago and Comibaena flavicans , but smaller in Comibaena chalybeata . The vinculum is deeply concave in Comibaena takasago , shallowly concave in Comibaena flavicans and very slightly so in Comibaena chalybeata . In the female genitalia, the sterigma is very large, rounded, ring-like, and expanded laterally, with the posterior margin narrowly concave at the middle.

Type material examined: China: holotype, ♂, Jen-aihsiang, Nantow-hsien, C. Formosa , v.1958, Inoue Coll. B.M. 1992-71, BMNH slides no. 16634 ( BMNH) .

Additional material examined: China: 2♂ 4♀, southern China, June ( IZCAS) .

Distribution: China (Henan, Hunan, Taiwan).

COMIBAENA FLAVICANS INOUE, 1982

( FIGS 26 , 80 , 130 )

Comibaena flavicans Inoue, 1982: 132 , figures 3c, 4c. Holotype ♂, Nepal (central): Kalbani , Kaligandaki. (BMNH)

Diagnosis: The moths of Comibaena flavicans are distinctive in having white discal markings on both wings, which are curved on the forewing. The white antemedial line of the forewing is straight, directed obliquely outwards; the postmedial line is slightly sinuous, extending slightly on veins, and bending proximally posterior to CuA 2, where it appears as a yellowish oblong patch. The tornal angle of the hindwing has a small, round, white patch, bordered with yellowish-brown, with a yellowish zigzag marking proximately. For other features see under the previous species.

Type material examined: Nepal: holotype, ♂, Kalbani, Kaligandaki, C . Nepal, 12.vii.1969, Coll. T. Miyashita, Inoue Coll. B.M. 1992-71 ( BMNH) .

Additional material examined: China: 2♀, Tibet, July ( IZCAS) .

Distribution: China (Tibet), Nepal.

COMIBAENA NIGROMACULARIA ( LEECH, 1897)

( FIGS 27 , 81 , 131 , 161 , 192 )

Euchloris nigromacularia Leech, 1897: 237 . Syntypes 2♀, China (west): Chow-pin-sa; Japan: Yokohama. ( BMNH)

Uliocnemis delicatior Warren, 1897b: 391 . Holotype ♀, Japan. (BMNH)

Comibaena nigromacularia: Prout, 1912: 100 .

Phorodesma eurynomaria Oberthür, 1916: 106 , pl. 388, figure 3274. Syntype (s), China: Tse-kou. (ZFMK)

Comibaena nigromacularia delicatior: Inoue, 1961: 73 , pl. 6: 138.

Diagnosis: This and the next species, Comibaena subdelicata , resemble each other in having the forewing postmedial line angled on M 1, a tornal patch on the forewing, and a dark apical patch on the hindwing. In Comibaena nigromacularia the forewing postmedial line is pointed on M 1 but convex in Comibaena subdelicata ; the tornal patch of the former species is orange but blackish in the latter; the hindwing apical patch is tinged with reddish scales in Comibaena nigromacularia but black in Comibaena subdelicata ; and the submarginal line on the hindwing is close to the outer margin in Comibaena nigromacularia but more distant in Comibaena subdelicata .

The male genitalia have a petal-like costal lobe, with a tiny pointed process. The vinculum is short, shallowly concave at the middle, and its branches are short and broad. The aedeagus is slender, spinose, and expanded in a bulb at the middle, the posterior half of which is spinulose. The eighth sternite possesses a tiny pointed tooth at the middle of the posterior margin.

In the female genitalia, the sterigma is nearly oval. The seventh sternite is laterally sclerotized anterior to the ostium.

Type material examined: Euchloris nigromacularia : China: 1♀, syntype, Chow-Pin-Sa, 5000 ft, native coll, v. & vi.1890, Leech Coll. 1900-64, BMNH slide no. 11766 ( BMNH) . Uliocnemis delicatior : Japan: ♀, holotype of Comibaena delicatior , Japan, Rothschild Bequest, B.M.1939-1, slide no. 17270 ( BMNH) . Phorodesma eurynomaria : China: 1♂, syntype, Tsekou, P. Dubernard, 1898 ( ZFMK) .

Additional material examined: China: 83♂ 27♀, northern to southern China, April to October ( IZCAS) ; Japan: 3♂ 1♀, May to July ( BMNH, ZFMK) ; Russia: 1♂, far east Russia ( ZFMK) .

Distribution: China (Heilongjiang, Beijing, Henan, Shaanxi, Gansu, Anhui, Zhejiang, Hubei, Jiangxi, Hunan, Fujian, Taiwan, Guangxi, Sichuan, Yunnan), Russia, Japan, Korean Peninsula.

COMIBAENA SUBDELICATA INOUE, 1986

( FIGS 28 , 82 , 132 )

Comibaena subdelicata Inoue, 1986: 52 , figures 6c, d, 7c, d. Holotype ♂, Japan: Yakushima , Nagata. (BMNH)

Diagnosis: For external differences, see under previous species. The uncus of the male genitalia is fused basally, and bifid terminally with slender branches. The ventral part of the costal lobe is spinose. The vinculum is deeply concave. In the female genitalia (from Inoue, 1986: fig. 7D), the sterigma is a sclerotized ring, the ductus bursae is slender, and the corpus bursae very weak. The eighth sternite of the male is trapeziform, and its posterior margin is well sclerotized.

Type material examined: Japan: holotype, ♂, Nagata, Yakushima , Japan, 16.vi.1972, T. Watanabe, Inoue Coll. B.M. 1992-71, BMNH slide no. 16514 ( BMNH) .

Additional material examined: China: 3♂ 7♀, southern China, May to July, September ( IZCAS, ZFMK) .

Distribution: China (Zhejiang, Jiangxi, Fujian, Taiwan, Sichuan), Japan.

The Comibaena procumbaria species group

This species group contains eight species, which have simple valvae. Three species Comibaena procumbaria , Comibaena subprocumbaria , and Comibaena tenuisaria are similar in having a white apical patch on the hindwing.

The terminal extension of the male hind tibia is of similar length to the first tarsus in the species Comibaena procumbaria , Comibaena subprocumbaria , Comibaena tenuisaria , and Comibaena swanni , absent in Comibaena birectilinea and Comibaena cenocraspis , and has not been examined in Comibaena hypolampes and Comibaena latilinea .

Five species ( Comibaena cenocraspis , Comibaena hypolampes , Comibaena latilinea , Comibaena swanni , and Comibaena birectilinea ) resemble members of Thetidia in wing pattern in having almost straight distinct antemedial and postmedial lines on the forewing, and a white submarginal line on the hindwing (indistinct in Comibaena latilinea and absent in Comibaena swanni ). All the male genitalia of these five species are simple, and quite similar to the pattern of Thetidia . However, species included in Thetidia usually lack the frenulum, which is present in Comibaena : as these five species also possess a frenulum, we maintained them in Comibaena .

COMIBAENA PROCUMBARIA ( PRYER, 1877)

( FIGS 29 , 83 , 133 , 162 , 193 )

Euchloris procumbaria Pryer, 1877: 232 , pl. 4: 2. Syntype (s), China: Shanghai. (BMNH)

Comibaena vaga Butler, 1881: 410 . Syntype (s), Japan: Tokyo. (BMNH)

Comibaena procumbaria: Prout, 1913a: 20 , pl. 2b.

Diagnosis: This species and Comibaena subprocumbaria lack or have only very faint transverse lines on the upperside, but bear distinct, dark green postmedial lines on the underside. The only difference between these two species lies in the white apical patch on the hindwing, which is smaller and reaches only to M 1 in Comibaena procumbaria , with veins Rs showing brown inside, whereas it is larger and reaches to M 2 in Comibaena subprocumbaria , with veins Rs and M 1 showing brown inside. The male and female genitalia could not be differentiated.

The posterior margin of the male eighth tergite is widely concave, forming two blunt processes. The eighth sternite protrudes at the middle with a slightly concave margin.

Type material examined: Euchloris procumbaria : China: 1♀, syntype, Shanghai, 80.123 ( BMNH). Comibaena vaga : Japan: 1♂, syntype , Japan, Jokei , 80.97 ( BMNH) .

Additional material examined: China: 39♂ 7♀, northern to southern China, May to August, March and October in Hong Kong ( IZCAS, BMNH, ZFMK) ; Japan: 4♂ 1♀, June to July ( BMNH, ZFMK) .

Distribution: China (Beijing, Hebei, Shanxi, Shandong, Henan, Gansu, Shanghai, Zhejiang, Hubei, Hunan, Jiangxi, Fujian, Taiwan, Guangdong, Hong Kong, Guangxi, Sichuan), Japan, Korean Peninsula.

COMIBAENA SUBPROCUMBARIA ( OBERTHÜR, 1916)

( FIGS 30 , 84 , 134 , 194 )

Phorodesma subprocumbaria Oberthür, 1916: 103 , pl. 387, figure 3259. Syntype (s), China: Siao-lou. (ZFMK)

Comibaena subprocumbaria: Prout, 1933: 93 .

Diagnosis: See the previous species.

Type material examined: China: 1♀, type, Siao-Lou, Chasseurs Indigènes, du P. Déjean, 1903 ( ZFMK) .

Additional material examined: China: 66♂ 24♀, almost all of China, March to October ( IZCAS, BMNH, ZFMK) .

Distribution: China (Beijing, Hebei, Henan, Gansu, Jiangsu, Zhejiang, Hubei, Jiangxi, Hunan, Fujian, Guangdong, Hainan, Guangxi, Sichuan, Yunnan, Tibet).

Remarks: There remains some doubt about the validity of the specific separation of this species from Comibaena procumbaria . However as we have not found intermediates, we leave them separated for the time being. Rearing may be necessary for certainty.

COMIBAENA TENUISARIA (GRAESER, 1889)

( FIGS 31 , 85 , 135 , 163 , 195 )

Phorodesma tenuisaria Graeser, 1889: 385 . Holotype ♂, Russia: Amurlandes , Vladivostok.

Euchloris tenuisaria: Staudinger, 1901: 262 .

Comibaena tenuisaria: Prout, 1912: 99 .

Diagnosis: Unlike the two previous species, Comibaena tenuisaria has slender antemedial and postmedial lines on the forewing; the apical patch on the hindwing differs from the previous two species in that it terminates at M 1, contains within it a perfect reddish-brown ‘8’ marking, and its proximal margin is wavy and bordered by a white submarginal line.

In the male genitalia, the uncus of Comibaena tenuisaria is longer than that of Comibaena subprocumbaria and Comibaena procumbaria ; the socii are slightly curved, but not as strongly as in Comibaena procumbaria . The costa of the valva of Comibaena tenuisaria is slightly concave, in contrast to the slight convexity in Comibaena subprocumbaria and Comibaena procumbaria , and the branches of the vinculum are narrower than in those two species.

The posterior margin of the male eighth segment is shallowly concave in the tergite and flat in the sternite.

The female genitalia resemble those of members of Thetidia in having a small, bulb-like sterigma and distinct antrum, which is situated at the middle of the sterigma, with the posterior margin slightly concave.

Material examined: 2♂ 12♀, southern China , May to July ( IZCAS, ZFMK); North Korea : 1♂ ( ZFMK); Russia : 5♂, Far East, July to August ( BMNH, ZFMK) .

Distribution: China (Shanxi, Henan, Shaanxi, Gansu, Jiangsu, Anhui, Fujian), Russia, Korean Peninsula.

COMIBAENA CENOCRASPIS PROUT, 1926

FIGS 32 , 86 , 136 , 164 , 196 )

Comibaena cenocraspis Prout, 1926: 133 . Syntypes 7♂, Upper Burma, Hparé. (BMNH)

Diagnosis: This and the next three species have similar patterns, and are best considered together. Comibaena cenocraspis and Comibaena hypolampes lack discal spots on the underside, which are present in Comibaena latilinea and Comibaena swanni ; only Comibaena latilinea has a distinct brown terminal line; the fringes are brown in Comibaena swanni but whitish in the other three species; the first two species have white submarginal lines.

These four species have only small differences in the male genitalia: for example, the costa in Comibaena cenocraspis and Comibaena hypolampes is slightly swollen, whereas it bears a tiny subapical sclerite in Comibaena latilinea and is distinctly expanded apically in Comibaena swanni . The vinculum is shallowly bifurcate in Comibaena swanni but deeply so in the other three species. The branches of the vinculum in Comibaena hypolampes are considerably narrower than in Comibaena cenocraspis and Comibaena latilinea , and strongly hooked. The uncus is much shorter than the socii in Comibaena latilinea and Comibaena swanni .

The female genitalia bear a rounded sterigma, and more than half of the posterior margin of the antrum is concave.

Type material examined: Myanmar: 3♂, syntypes, Hparê , north-east Burma, ix.1923, A.E. Swann XII, L.B. Prout Coll. B.M. 1939-643 ( BMNH) ; 1♂, syntype, same data, BMNH slide no. 21779 ( BMNH) ; 1♂, syntype, same data, Joicey Bequest, Brit. Mus. 1934- 120 ( BMNH) ; 1♂, syntype, same data ( ZFMK) .

Additional material examined: China: 8♂ 2♀, southern China, April , July to August ( IZCAS, BMNH) .

Distribution: China (Sichuan, Yunnan), India, Myanmar.

COMIBAENA HYPOLAMPES PROUT, 1918

( FIGS 33 , 87 , 137 )

Comibaena hypolampes Prout, 1918: 19 . Syntype (s) ♂, China: Tibet, Vrianatong. (BMNH)

Diagnosis: As for Comibaena cenocraspis . The antemedial and postmedial lines are broader than in Comibaena cenocraspis . The eighth sternite of the male is slightly concave at the middle of the posterior margin.

Type material examined: China: 1♂, syntype (maybe holotype), Vrianatong , Tibet, Joicey Coll., B.M. 1929- 122 ( BMNH) .

Additional material examined: China: 35♂ 5♀, northern Yunnan, July to August ( IZCAS, ZFMK) .

Distribution: China (Yunnan, Tibet).

COMIBAENA LATILINEA PROUT, 1912

( FIGS 34 , 88 , 138 , 165 )

Comibaena latilinea Prout, 1912: 101 . Holotype ♂, China (west): Pu-tsu-Fong. (BMNH)

Phorodesma theodoraria Oberthür, 1916: 114 , pl. 388, figure 3277. Syntypes 2♂, China: Szechwan [Sichuan], Tachien-lu [Kangding] region. (ZFMK)

Diagnosis: As for Comibaena cenocraspis . The male eighth sternite is identical to that of Comibaena hypolampes .

Material examined: China: 1♂, holotype, Pu-tsu- Fang, 9820 ft, native coll. vi.-vii.1890, Leech Coll. 1900-64 ( BMNH) ; 1♂, syntype of Phorodesma theodoraria : Ta-tsien-Lou, Chasseurs du P. Déjean, 1904 ( ZFMK) .

Distribution: China (Sichuan).

Figures

Figures 1–26. Comibaenini moths. 1, Comibaena quadrinotata; 2, Comibaena fuscidorsata; 3, Comibaena cassidara; 4, Comibaena attenuata; 5, Comibaena biplaga; 6, Comibaena apicipicta; 7, Comibaena striataria; 8, Comibaena delineata; 9, Comibaena dubernardi; 10, Comibaena ornataria; 11–15, Comibaena pictipennis; 11, from Darjeeling; 12, syntype of Comibaena pictipennis superornataria; 13, from Tengchong, Yunnan; 14, from Baoshan, Yunnan; 15, from Nanchuan, Sichuan; 16, Comibaena parornataria sp. nov., holotype; 17, Comibaena bellula sp. nov., holotype; 18, Comibaena decora sp. nov., holotype; 19, Comibaena auromaculata sp. nov., holotype; 20, Comibaena tibetensis sp. nov., holotype; 21, Comibaena tancrei; 22, Comibaena sheni sp. nov., holotype; 23, Comibaena argentataria; 24, Comibaena signifera subargentaria; 25, Comibaena takasago; 26, Comibaena flavicans. Scale bar = 13 mm.

Figures 67–78. Male genitalia. 67–69, Comibaena pictipennis. 67, from Dahaoping, Tengchong, Yunnan; 68, from Nyingchi, Tibet; 69, from Qingchengshan, Sichuan; 70, Comibaena parornataria; 71, Comibaena bellula; 72, Comibaena decora; 73, Comibaena auromaculata; 74, Comibaena tibetensis; 75, Comibaena tancrei; 76, Comibaena sheni; 77, Comibaena argentataria; 78, Comibaena signifera subargentaria. Scale bars = 1 mm.

Figures 108–131. Aedeagus. 108, Comibaena quadrinotata; 109, Comibaena fuscidorsata; 110, Comibaena cassidara; 111, Comibaena attenuata; 112, Comibaena biplaga; 113, Comibaena apicipicta; 114, Comibaena striataria; 115, Comibaena delineata; 116, Comibaena dubernardi; 117, Comibaena ornataria; 118–119, Comibaena pictipennis; 118, from Tengchong, Yunnan; 119, from Nyingchi; 120, Comibaena parornataria; 121, Comibaena bellula; 122, Comibaena decora; 123, Comibaena auromaculata; 124, Comibaena tibetensis; 125, Comibaena tancrei; 126, Comibaena sheni; 127, Comibaena argentataria; 128, Comibaena signifera subargentaria; 129, Comibaena takasago; 130, Comibaena flavicans; 131, Comibaena nigromacularia. Scale bars = 1 mm.

Figures 188–199. Female genitalia. 188, Comibaena sheni; 189, Comibaena argentataria; 190, Comibaena signifera subargentaria; 191, Comibaena takasago; 192, Comibaena nigromacularia; 193, Comibaena procumbaria; 194, Comibaena subprocumbaria; 195, Comibaena tenuisaria; 196, Comibaena cenocraspis; 197, Comibaena birectilinea; 198, Thetidia albocostaria; 199, Thetidia atyche. Scale bars = 1 mm.

Figures 79–90. Male genitalia. 79, Comibaena takasago; 80, Comibaena flavicans; 81, Comibaena nigromacularia; 82, Comibaena subdelicata; 83, Comibaena procumbaria; 84, Comibaena subprocumbaria; 85, Comibaena tenuisaria; 86, Comibaena cenocraspis; 87, Comibaena hypolampes; 88, Comibaena latilinea; 89, Comibaena swanni; 90, Comibaena birectilinea. Scale bars = 1 mm.

Figures 27–54. Comibaenini moths. 27, Comibaena nigromacularia; 28, Comibaena subdelicata; 29, Comibaena procumbaria; 30, Comibaena subprocumbaria; 31, Comibaena tenuisaria; 32, Comibaena cenocraspis; 33, Comibaena hypolampes; 34, Comibaena latilinea; 35, Comibaena swanni; 36, Comibaena birectilinea sp. nov., holotype; 37, Microbaena pulchra minor; 38, Thetidia albocostaria; 39, Thetidia correspondens; 40, Thetidia atyche; 41, Thetidia chlorophyllaria, 42, holotype of Thetidia pekingensis; 43–44, Thetidia kansuensis, 43, from Gansu, 44, from Tibet; 45, Thetidia smaragdaria; 46, Proteuchloris neriaria; 47, Linguisaccus subhyalina; 48, Linguisaccus minor sp. nov., paratype; 49, Chlorochromodes rhodocraspeda sp. nov., holotype; 50, Argyrocosma inductaria; 51, Protuliocnemis castalaria; 52, Protuliocnemis falcipennis; 53, Protuliocnemis candida sp. nov., paratype; 54, Protuliocnemis dissimilis sp. nov., paratype. Scale bar = 13 mm.

Figures 156–175. The male eighth segment. In Figures 156–174, the tergite is on the left and the sternite is on the right. 156, Comibaena fuscidorsata; 157, Comibaena apicipicta; 158, Comibaena ornataria; 159, Comibaena auromaculata; 160, Comibaena tancrei; 161, Comibaena nigromacularia; 162, Comibaena procumbaria; 163, Comibaena tenuisaria; 164, Comibaena cenocraspis; 165, Comibaena latilinea; 166, Thetidia chlorophyllaria; 167, Thetidia kansuensis; 168–170, Linguisaccus subhyalina; 171, Linguisaccus minor; 172, Chlorochromodes rhodocraspeda; 173, Protuliocnemis candida; 174, Protuliocnemis dissimilis; 175, tympanal organ of Pro. dissimilis. Scale bar = 1 mm.

Figures 132–155. Aedeagus. 132, Comibaena subdelicata; 133, Comibaena procumbaria; 134, Comibaena subprocumbaria; 135, Comibaena tenuisaria; 136, Comibaena cenocraspis; 137, Comibaena hypolampes; 138, Comibaena latilinea; 139, Comibaena swanni; 140, Comibaena birectilinea; 141, Microbaena pulchra; 142, Thetidia albocostaria; 143, Thetidia atyche; 144, Thetidia chlorophyllaria; 145, Thetidia kansuensis; 146, Thetidia smaragdaria; 147, Proteuchloris neriaria; 148, Linguisaccus subhyalina; 149, Linguisaccus minor; 150, Chlorochromodes rhodocraspeda; 151, Argyrocosma inductaria; 152, Protuliocnemis castalaria; 153, Protuliocnemis falcipennis; 154, Protuliocnemis candida; 155, Protuliocnemis dissimilis. Scale bars = 1 mm.