Comibaena, HUBNER, 1823

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 : 726-745

COMIBAENA HÜBNER, 1823

Comibaena Hübner, 1823: 284 . Type species: Geometra bajularia Denis et Schiffermüller, 1775 , by subsequent designation by Moore, 1887, Austria: Vienna district.

Phorodesma Boisduval, 1840: 179 . Type species: Geometra bajularia Denis et Schiffermüller, 1775 , by subsequent designation by Fletcher, 1979.

Comobaena Agassiz, 1847: 276 . [An unjustified emendation of Comibaena Hübner. ]

Uliocnemis Warren, 1893: 355 . Type species: Phorodesma cassidara Guenée, 1858 , by original designation. India: central.

Colutoceras Warren, 1895: 88 . Type species: Colutoceras diluta Warren, 1895 , by original designation. Japan: Kiushiu.

Myrtea Gumppenberg, 1895: 477 , 478. Type species: Phalaena pustulata Hufnagel, 1767 (= Geometra bajularia Denis & Schiffermüller, 1775 ), by subsequent designation by Fletcher, 1979. [Junior homonym of Myrtea Turton, 1822 (Mollusca) .]

Probolosceles Meyrick, 1897: 73 . Type species: Comibaena quadrinotata Butler, 1889 , [Junior homonym of Probolosceles Warren, 1896c ], by original designation. Borneo: Sarawak.

Chlorochaeta Warren, 1904: 464 . Type species: Chlorochaeta longipennis Warren, 1904 , by original designation. Nigeria: Niger, Degama.

Adults: Antenna of male bipectinate, outer rami longer than inner rami, ciliate terminally; filiform and ciliate in female; a few species bipectinate in both sexes ( Comibaena argentataria , Comibaena signifera , Comibaena flavitaenia , Comibaena leucochloraria ). Frons not protruding, usually green, sometimes brown or white. Labial palpus with the third segment in female elongate. Hind tibia with two pairs of spurs in both male and female; male usually with hind tibia dilated, with hair-pencil and terminal extension.

Wing shape and pattern. Apex of forewing blunt, acute in a few species, that of hindwing round. Outer margin smooth. Wings green. Forewing with antemedial and postmedial lines distinct, wide, or very weak, the latter curving or straight, in some species accompanied by a white shadow or diffusion outside; both lines in some species composed of dots on veins (e.g. Comibaena punctaria , Comibaena cheramota , Comibaena viridifimbria , Comibaena rhodonia ), occasionally lines absent. Hindwing usually with postmedial line absent, sometimes with submarginal line. Usually with various blotches on tornal angle of forewing and apex of hindwing, white to brown, evenly colourful or pale-centred, the blotches of some species reaching M 1 on forewing and M 3 on hindwing; occasionally these blotches absent; a few species with cloud-like streaks. Both wings usually with small discal spots. Underside: usually pale green on upper half of forewing, lower half and hindwing greenish white; sometimes with green postmedial line; discal spot present on forewing, present or absent on hindwing.

Third sternite of male usually with a pair of setal patches; eighth tergite sometimes with long and narrow sclerotized tooth, or concave medially, sometimes with small spines on margin; eighth sternite membranous, protruding, or shallowly concave posteriorly.

Male genitalia. Uncus slightly to strongly sclerotized, tapering and bifid posteriorly, fused basally, or not fused throughout. Socii long and narrow, slightly to strongly sclerotized, slightly tapering, straight or curving. Gnathos undeveloped, median process absent. Valva usually with distinct costal lobe, slightly to strongly sclerotized, with distal part tapering or blunt, a median tooth sometimes present on ventral margin; sometimes valva simple, costa appearing as spinose area, or with various shapes of pointed teeth. Sacculus unmodified. Transtilla membranous. Vinculum developed, deeply or shallowly bifurcate, distance between two branches varying. Coremata present or not. Aedeagus usually needlelike, sclerotized terminally; cornutus absent; coecum penis small, often expanded at end.

Female genitalia. Apophyses anteriores and posteriores slender. Ostium situated at posterior to middle of the eighth sternite. Sterigma modified, various. Ductus bursae long and narrow (slender) or very short and broad. Corpus bursae very weak; signum absent.

Diagnosis: The wing patterns of Comibaena species are quite complex. The species in which the antemedial and postmedial lines are composed of small dots on the veins are similar to species in Chlorochromodes and some species of Argyrocosma . These species can be distinguished from the latter two genera by the male genitalia: the middle concavity of the vinculum in Comibaena is much wider than that in Chlorochromodes ; the sclerotized teeth on the costa are simpler than those of Argyrocosma . Comibaena species usually have a tornal patch on the forewing and an apical patch on the hindwing, and a few are similar to members of the genus Protuliocnemis (see diagnosis under Protuliocnemis ), but the widely bifurcate vinculum and the bifid uncus are distinctive. Some species of Comibaena (including Comibaena cenocraspis , Comibaena hypolampes , Comibaena latilinea , Comibaena swanni and Comibaena birectilinea ) are similar to members of Thetidia in wing pattern and male genitalia, but the frenulum is present in these species and absent in Thetidia .

Biological notes: As for tribe. The larvae feed on trees and shrubs. Singh (1957) and Holloway (1996)

recorded that the larvae of Comibaena cassidara feed on flowers and young pods.

Known host plants are as follows [based on Holloway (1996), Scoble (1999), Hausmann (2001), and the website of ‘HOSTS – a Database of the World’s Lepidopteran Hostplants’ (http://www.nhm.ac.uk/research-

curation/research/projects/hostplants/):

Comibaena amoenaria ( Fagaceae : Fagus crenata , Lithocarpus edulis , Quercus mongolica , Quercus serrata )

Comibaena argentataria ( Rosaceae : Rubus pungens , Rubus hirsutus )

Comibaena attenuata ( Sapindaceae : Nephelium lappaceum )

Comibaena bajularia ( Fagaceae : Quercus robur , Quercus petraea , Prunus domestica , Prunus spinosa , Fagus )

Comibaena biplaga ( Melastomataceae : Memecylon )

Comibaena cassidara (Leguminosae: Acacia nilotica , Flemingia macrophylla ; Anacardiaceae : Anacardium occidentale , Mangifera indica ; Convolvulaceae : Cuscuta ; Rutaceae : Glycosmis pentaphylla ; Rubiaceae : Ixora ; Lythraceae : Lagerstroemia indica ; Rhamnaceae : Ziziphus jujuba )

Comibaena cheramota ( Myrtaceae : Decaspermum fruticosum , Myrtus vitiensis ; Compositae : Tagetes )

Comibaena diluta ( Compositae : Farfugium japonicum ; Fagaceae : quercus glauca )

Comibaena esmeralda ( Pinaceae : Pinus patula )

Comibaena integranota (Leguminosae: Cajanus cajan ; Melastomataceae : Memecylon )

Comibaena leucospilata ( Myrtaceae : Acca ; Sterculiaceae : Hermannia ; Proteaceae : Macadamia ; Bignoniaceae : Markhamia lutea ; Markhamia platy- calyx; Anacardiaceae : Ozoroa mucronata ; Leguminosae: Acacia caffra , Acacia karroo )

Comibaena nigromacularia ( Fagaceae : Castanea crenata , Quercus serrata ; Leguminosae: Lespedeza bicolor )

Comibaena procumbaria (Leguminosae: Lespedeza bicolor , Lespedeza cyrtobotrya ; Euphorbiaceae : Mallotus japonicus ; Myricaceae : Myrica rubra ; Podocarpaceae : Podocarpus macrophyllus )

Distribution: Widely distributed in the Oriental, Palaearctic, and Australasian regions.

The Comibaena quadrinotata species group

The species Comibaena quadrinotata , Comibaena fuscidorsata , Comibaena cassidara and Comibaena attenuata fall into this group. All of them possess the tornal patch on the forewing and the apical patch on the hindwing. Both wings have small dark discal spots. The terminal extension on the male hind tibia is longer than the first tarsus ( Comibaena attenuata , Comibaena fuscidorsata , and Comibaena cassidara ) or of similar length to the first tarsus ( Comibaena quadrinotata ).

The male genitalia of this group bear a long costal process on the valva rather than a costal lobe as in most other species; the uncus (except in Comibaena attenuata ) has the basal part fused and the terminal part bifid; the socii are shorter than the uncus ; the eighth tergite has a developed pointed central process (except in Comibaena attenuata ). The ductus bursae and corpus bursae of the female genitalia are not distinct from each other, and the latter is quite small, folding posteriorly in the natural state (no female genitalia preparation is available for Comibaena fuscidorsata ).

COMIBAENA QUADRINOTATA BUTLER, 1889

( FIGS 1 , 55 , 108 , 176 )

Comibaena quadrinotata Butler, 1889: 22 , 105, pl. 136, figure 7. Syntype (s), India: Kangra district , Dharmsala. (BMNH)

Nemoria quadrinotata: Hampson, 1895: 503 .

Diagnosis: The wings are green, tinged with blue. The white postmedial line on the forewing is faint and forms a blunt protrusion near M 1. The hindwing lacks postmedial and submarginal lines. The forewing tornal patch is brown, with the upper margin pointed and extending beyond CuA 2 in the type. The hindwing apical patch is brown, not reaching M 1. The male genitalia are distinctive in that the valva bears a subapical spinulose tooth, and the bifid part of the uncus is quite short. The posterior margin of the eighth sternite of the male is concave at the middle and bears two blunt protrusions. In the female genitalia, the lamella postvaginalis of Comibaena quadrinotata is bell-like, with the posterior margin protruding. The lamella antevaginalis bears a pair of diverging blunt processes.

Type material examined: India: 1♂, syntype, Dharmsala, 87.59, BMNH slide no. 11824 ( BMNH) .

Additional material examined: China: 34♂ 17♀, March to October, southern China ( IZCAS); 2♂, west Celebes , May ( BMNH) .

Distribution: China (Henan, Zhejiang, Hubei, Hunan, Fujian, Taiwan, Hainan, Guangxi, Sichuan), Japan, India (north), north-east Himalayas, Vietnam, Sri Lanka, Malaysia (west), Indonesia [Celebes, Java ( ssp. pallidicincta ), Sumatra, Sulawesi].

COMIBAENA FUSCIDORSATA PROUT, 1912

( FIGS 2 , 56 , 109 , 156 )

Comibaena quadrinotata fuscidorsata Prout, 1912: 101 . Holotype ♂, India: Assam, Khasi Hills. (BMNH)

Comibaena fuscidorsata: Holloway, 1996: 243 , pl. 8, figure 245.

Diagnosis: The wing pattern of Comibaena fuscidorsata are similar to those of Comibaena quadrinotata . However the wings are tinged with a yellowish colour rather than blue, and the tornal and apical patches are larger. The brown forewing tornal patch is semicircular, but pointed in Comibaena quadrinotata , reaching or slightly extending beyond CuA 2. The hindwing apical patch is almost identical to that of Comibaena quadrinotata . In the male genitalia, the valva possesses a large, pointed, distally curved costal process, which bears a small, subapical spine; the vinculum is very long and deeply concave at the middle. The eighth sternite of the male is roundly protruded.

Type material examined: India: holotype, ♂, Khasia Hs, 94–189, BMNH slide no. 16823 ( BMNH) .

Additional material examined: China: 12♂ 4♀, southern China, April to June , September to December ( IZCAS, BMNH) ; India: 3♂, north-eastern India, February , December ( BMNH, ZFMK) .

Distribution: China (Hainan, Taiwan, Yunnan), India.

COMIBAENA CASSIDARA ( GUENÉE, 1858)

( FIGS 3 , 57 , 110 , 177 )

Phodoresma cassidara Guenée, 1858: 370 . Syntypes 1♂ 1♀, Inde centrale [central India]. (BMNH)

Geometra detenta Walker, 1861: 518 . Holotype ♀, Nepal. (BMNH)

Geometra discessa Walker, 1861: 521 . Holotype ♀, Punjab. (BMNH)

Comibaena cassidara: Holloway, 1976: 61 .

Diagnosis: The reddish-brown forewing tornal patch is small and rounded, not extending beyond CuA 2. The hindwing apical patch is reddish-brown, quite narrow, and elongate. The male genitalia are almost identical to those of Comibaena fuscidorsata , but the costal process of the valva is shorter, located slightly more distally, and lacking the small tooth. The eighth abdominal segment is identical to that of Comibaena fuscidorsata . In the female genitalia, the lamella postvaginalis is similar to that of Comibaena quadrinotata , but broader. The lamella antevaginalis is band-like.

Type material examined: Phodoresma cassidara : India: 1♀, syntype, Ex. Musaeo, Ach Guenée , Ex Typicalibus Speciminibus, Ex. Oberthür Coll. Brit. Mus. 1927-3 ( BMNH); 1♂, syntype, Ex. Musaeo, Ach Guenée, Ex Typicalibus Speciminibus, Ex. Oberthür Coll., Brit. Mus. 1927-3 ( BMNH). Geometra discessa : Nepal: holotype, ♀, Punjab (with green type label). Geometra detenta : holotype, ♀ , Nepal ( BMNH) .

Additional material examined: China: 8♂ 1♀, southern China, May to September ( IZCAS, BMNH) ; India: 3♂ 1♀, northern, eastern, and western India, August ( BMNH) ; Thailand: 1♂, April ( BMNH) ; Pakistan: 1♀, northern Pakistan, May ( BMNH) ; Malaysia: 1♂ , north Borneo, August ( BMNH) ; Indonesia: 1♂, Java, May ( BMNH) .

Distribution: China (Fujian, Taiwan, Hainan, Yunnan), India, Nepal, Pakistan, Thailand, Sri Lanka, Philippines, Malaysia, Singapore, Indonesia.

COMIBAENA ATTENUATA (WARREN, 1896C)

( FIGS 4 , 58 , 111 , 178 )

Probolosceles attenuata Warren, 1896c , Novit. zool., 3: 369. Holotype ♂, Borneo : north Borneo, Mount Mulu. (BMNH)

Comibaena attenuata: Holloway, 1976: 61 .

Diagnosis: The wings are dark green, lacking any blue tinge. The transverse lines are invisible. The tornal patch on the forewing is larger than in the former three species, irregularly extending anteriorly, sometimes discontinuous and appearing as separate patches. The kidney-shaped brown apical patch of the hindwing is very large, terminating before M 3. In the male genitalia, the uncus is divided over its whole length; the valva possesses a long pointed apical tooth; the vinculum is widely concave at the middle, forming two large lobe-like processes. The eighth abdominal segment in the male is unmodified. The lamella postvaginalis of the female genitalia is a quadrate sclerite.

Type material examined: Malaysia: holotype, ♂, Mount Mulu , north Borneo, 1–4000 ft (Hose), Rothschild Bequest, B.M. 1939-1 ( BMNH) .

Additional material examined: China: 13♂ 4♀, southern China, April to July, November ; Philippines: 3♂, northern and western Philippines, February to March , July ( ZFMK, ZSM) ; Malaysia: 2♂, western Malaysia, north Borneo, May , October to November ( BMNH) ; Singapore: 2♂ 1♀ ( BMNH) ; Brunei: 1♀, February ( BMNH) ; Cambodia: 1♀ ( ZSM) ; Indonesia: 3♂ 1♀, Java, Sumatra, Sulawesi, February to March, June, December ( BMNH, ZFMK) .

Distribution: China (Hainan, Yunnan), Cambodia, Philippines, Malaysia (Sabah), Singapore, Brunei, Indonesia [Sulawesi ( ssp. prouti ), Sumatra, Java].

The Comibaena integranota species group

Only one species from China falls into this group, which also includes Comibaena integranota Hampson ( Sri Lanka, south India) and Comibaena diluta Warren ( Japan) ( Holloway, 1996). The male genitalia of this group only have a spinose costal area on the valva, and lack the separate costal lobe or process.

COMIBAENA BIPLAGA WALKER, 1861

( FIGS 5 , 59 , 112 , 179 )

Comibaena biplaga Walker, 1861: 577 . Syntype (s), Borneo : Sarawak. (OUM)

Diagnosis: This species is quite close to Comibaena attenuata in wing pattern, having a similar large kidney-shaped apical patch on the hindwing. The main difference is that the tornal patch on the forewing terminates under CuA 1, but extends beyond CuA 1 in Comibaena attenuata . The terminal extension of the male hind tibia is longer than the first tarsus. The eighth sternite of the male bears two small processes. The female genitalia also differ markedly from those of the previous four species, having an irregularly shaped sterigma and a slender ductus bursae.

Material examined: China: 9♂ 6♀, southern China, May to June, August to September, and December ( IZCAS) ; India: 2♂ 3♀, north-eastern India, February ( BMNH) ; Malaysia: 1♀, western Malaysia, September ( BMNH) ; Indonesia: 3♀, Sumatra, Java , Sarawak, February, June, October to November ( BMNH) .

Distribution: China (Hainan, Guangxi, Tibet), India, Vietnam, Malaysia ( Sarawak, Sabah), Indonesia (Sumatra, Java).

The Comibaena apicipicta species group

Nineteen species fall into this group, which contains most of the Chinese Comibaena species. All the species have a developed separated costal lobe in the male genitalia, an uncus divided over its whole length with branches of even width, and bent and hooked socii. The species Comibaena apicipicta , Comibaena striataria , Comibaena delineata , Comibaena dubernardi , Comibaena ornataria , Comibaena pictipennis , Comibaena parornataria , Comibaena bellula , Comibaena tibetensis , Comibaena tancrei , Comibaena sheni , and Comibaena signifera (altogether 12 species) bear a pointed tooth at the middle of the ventral margin of the costal lobe, which is lacking in the other species. The lateral branches of the vinculum are developed in some species, for example Comibaena apicipicta , Comibaena striataria , Comibaena delineata , Comibaena parornataria , Comibaena decora , and Comibaena auromaculata .

The female genitalia (based on the examined genitalia) have a slender ductus bursae and a very weak and small corpus bursae.

The terminal extension of the male hind tibia is absent in Comibaena apicipicta , Comibaena striataria , Comibaena dubernardi , and Comibaena tibetensis , shorter than the first tarsus in Comibaena delineata , Comibaena pictipennis , Comibaena parornataria , Comibaena bellula , Comibaena decora , Comibaena auromaculata , Comibaena sheni , Comibaena takasago , and Comibaena flavicans , of similar length to the first tarsus in Comibaena ornataria , Comibaena tancrei , Comibaena argentataria , and Comibaena signifera , and longer than the first tarsus in Comibaena nigromacularia and Comibaena subdelicata .

COMIBAENA APICIPICTA PROUT, 1912

( FIGS 6 , 60 , 113 , 157 )

Comibaena apicipicta Prout, 1912: 101 . Holotype ♂, China: Tibet, Yatung. (BMNH)

Diagnosis: Comibaena apicipicta and Comibaena striataria , are distinctive in lacking the transverse lines on both wings and the large tornal and apical patches respectively on fore- and hindwings. The wings are suffused throughout with short, white, wavy transverse lines. Small discal spots are present in the type of Comibaena apicipicta but absent in Comibaena striataria . A small pinkish apical patch is present on the hindwing of Comibaena apicipicta . The tip of the costal lobe of the valva in the male genitalia is quite narrow, appearing as a blunt spinose protrusion; the middle tooth on the ventral margin is small; the vinculum is developed, widely bifurcate, with long and tapering branches. The posterior margin of the male eighth sternite appears as a small longitudinal sclerotized ridge at the middle, forming a triangular process.

Type material examined: China: holotype, ♂, Tibet: Yatung, A.E. Hobson, 98–201 ( BMNH) .

Additional material examined: China: 12♂ 1♀, southern China, July to September ( IZCAS) .

Distribution: China (Fujian, Yunnan, Tibet).

COMIBAENA STRIATARIA ( LEECH, 1897)

( FIGS 7 , 61 , 114 , 180 )

Euchloris striataria Leech, 1897: 239 . Holotype ♀, China (western): Che-tou. (BMNH)

Comibaena striataria: Prout, 1913a: 20 , pl. 2c.

Diagnosis: The small pointed process on the male eighth sternite is more slender than in Comibaena apicipicta . Compared to Comibaena apicipicta , the uncus is much longer, projecting beyond the socii; the tip of the costal lobe is blunt and decorated with a row of small spines, as opposed to rather narrow; the pointed tooth at the middle of the ventral margin is much bigger; the vinculum is more widely bifurcate, and the two branches are shorter. The female genitalia are distinctive in the broad, quadrate, ring-like sterigma; the ductus bursae is broader than in other species covered in the present work.

Type material examined: China: holotype, ♀, Che-tou , 11070 ft, native coll (collector), vii.-viii.1890 ( BMNH) .

Additional material examined: China: 55♂ 1♀, south-western China, March to September ( IZCAS, ZFMK) .

Distribution: China (Shaanxi, Sichuan, Yunnan).

COMIBAENA DELINEATA ( WARREN, 1893)

( FIGS 8 , 62 , 115 , 181 )

Uliocnemis delineata Warren, 1893: 356 , pl. 31, figure 14. Syntypes ♂ ♀, India: Sikkim. (BMNH) Geometra delineata: Hampson, 1895: 497 . Comibaena delineata: Prout, 1913a: 20 .

Diagnosis: The wing pattern of Comibaena delineata has a superficial resemblance to that of Comibaena flavicans : both have a narrow yellowish posterior and enlarged part of the postmedial line on the forewing and a small white tornal patch bordered with purple on the hindwing. However, they differ markedly in the following features: the transverse lines are yellowish-white in Comibaena delineata but white in Comibaena flavicans ; the discal spots on both wings are small, blackish-brown dots in Comibaena delineata but elongate white stripes in Comibaena flavicans ; the submarginal line of the hindwing is absent in Comibaena flavicans , but developed in Comibaena delineata , pale yellow, and positioned far from the outer margin, forming a small tooth near M 3 and a larger pointed one below CuA 2. In the male genitalia, Comibaena delineata shares the characteristic socii with Comibaena ornataria and Comibaena bellula , with its most basal part very broad, and the terminal part narrow and hooked, forming a blunt process; but the basal part of the socii in Comibaena delineata is much broader than in the other two species; a long spinose terminal process is present on the costal lobe; the vinculum is developed, and so deeply concave so as to be semicircular at middle in one specimen from Yadong, Tibet (slide no. 21770 in BMNH), but only shallowly concave at middle, and with short branches in one IZCAS specimen ( Fig. 62 ). The female genitalia have an inverse butterfly-shaped sterigma.

Type material examined: India: 2♂, syntypes, Sikkim, O. Möller, Collectio H.J. Elwes, Rothschild Bequest, B.M. 1939-1 ( BMNH) ; 1♀, syntype, Sikkim, Knyvett, Collectio H.J. Elwes, Rothschild Bequest, B.M. 1939-1 ( BMNH) .

Additional material examined: China: 11♂ 5♀, Tibet, July to August ( IZCAS, BMNH) ; Bhutan: 1♂ ( BMNH) ; India: 1♂, Sikkim ( ZFMK) .

Distribution: China (Tibet), India (Sikkim), Bhutan.

COMIBAENA DUBERNARDI ( OBERTHÜR, 1916)

( FIGS 9 , 63 , 116 , 182 )

Phorodesma dubernardi Oberthür, 1916: 114 , pl. 388, fig. 3276. Syntypes ♂, China: Tse-kou. (ZFMK)

Comibaena rectilineata Sterneck, 1927: 13 . Holotype ♀, China (west).

Comibaena dubernardi: Prout, 1933: 92 .

Diagnosis: The postmedial line on the forewing is more proximal compared to other species, straight, and sometimes slightly bent inwards below CuA 2. The hindwing is distinctive in having a large, yellowishpurple tornal patch below CuA 2, and in the absence of any transverse lines. The male genitalia are different from the other species in having a triangular dorsal process on the costal lobe as well as the central ventral tooth; in addition, the tip of the costal lobe is truncate, decorated with irregular spines, and bears a pointed apical tooth; the vinculum is very short, and only slightly concave at the middle, not forming branches. The sterigma of the female genitalia is labiate.

Type material examined: China: 1♂, syntype, Thibet, Tsekou, R.P. Dubernard ( ZFMK) ; 2♂, syntypes, Tsekou, P. Dubernard, 1898 ( ZFMK) .

Additional material examined: China: 20♂ 2♀, southwestern China, June to August ( IZCAS, ZFMK) .

Distribution: China (Yunnan).

COMIBAENA ORNATARIA ( LEECH, 1897)

( FIGS 10 , 64 , 117 , 158 , 183 )

Euchloris ornataria Leech, 1897: 238 . Syntypes 6♂, China (western): Pu-tsu-fong. (BMNH & ZFMK)

Comibaena ornataria: Prout, 1912: 100 .

Diagnosis: Comibaena ornataria and the following species, Comibaena pictipennis , are similar in having a convex postmedial line on the forewing, a forewing subtornal patch, and a hindwing apical patch. However, the postmedial line of Comibaena ornataria is closer to the outer margin than in Comibaena pictipennis and the subtornal patch is narrower in Comibaena ornataria but forms a rounded patch in Comibaena pictipennis . The most distinctive difference is in the hindwing: the reddish-brown apical patch is much narrower and smaller in Comibaena ornataria , but quite large in Comibaena pictipennis ; the submarginal line is closer to the outer margin and only bent inwards near the tornal angle in Comibaena ornataria , but in Comibaena pictipennis it forms a large tooth near M 2 and CuA 2 and has two U-shaped convexities; the hindwing tornal patch is small in Comibaena ornataria , but very large in Comibaena pictipennis .

The posterior central area of the male eighth sternite is sclerotized, with a small ridge. The male genitalia are similar to those of Comibaena delineata , but the terminal process of the costal lobe is less narrow and pointed than in Comibaena delineata ; the vinculum is short, shallowly and widely concave at middle, almost flat, with two quite small branches; the tooth on the ventral margin of the costal lobe is sometimes very small, as in much of the Tibetan material. In one dissected syntype, the valva is asymmetric ( Fig. 64 ), the right costal terminal process being expanded, and less pointed than the left one. The sterigma of the female genitalia is hat-like, with the posterior margin concave at the middle, and the anterior margin expanded.

Type material examined: China: 4♂, syntypes, Pu-tsu-Fang , 9820 ft, native coll, vi.-vii.1890, Leech Coll. 1900-64 ( BMNH) ; 1♂, ibidem ( ZFMK) .

Additional material examined: China: 68♂ 11♀, south-western China, June to August ( IZCAS, BMNH, ZFMK) .

Distribution: China (Sichuan, Yunnan).

COMIBAENA PICTIPENNIS BUTLER, 1880

( FIGS 11–15 , 65–69 , 118–119 , 184 )

Comibaena pictipennis Butler, 1880: 215 . Syntype (s), India: Darjeeling. (BMNH)

Geometra pictipennis: Hampson, 1895: 496 .

Comibaena pictipennis: Prout, 1933: 92 , pl. 11b.

Phorodesma superornataria Oberthür, 1916: 104 , pl. 387, figures 3262, 3263. Syntypes 9♂ ♀, [ China], Siao-Lou ; Tien-tsuen. (ZFMK) syn. nov.

Diagnosis: As for previous species. The male genitalia are very similar to those of Comibaena ornataria in the shapes of the uncus , the socii, and the valvae, but differ from that species in that the ventral tooth on the costal lobe is longer, and the vinculum is much more deeply concave and not truncate as in Comibaena ornataria . The sterigma in the female genitalia is quadrate. The ductus bursae is broader, but more slender than that of Comibaena striataria .

Type material examined: Comibaena pictipennis pictipennis : India: 1♀, syntype, Darjiling, 79.57 ( BMNH) ; 1♀, syntype?, Darjeeling , Moore Coll. 94–106 ( BMNH) . Comibaena pictipennis superornataria : China: 1♂, syntype, Siao-Loû , Chasseurs Indigènes du P. Déjean, 1903 ( ZFMK) ; 1♀, syntype, Siao-Loû , Chasseurs Indigènes du P. Déjean, 1903 ( ZFMK) .

Additional material examined: China: 33♂ 10♀, southern China, May to June , August to October ( IZCAS, BMNH, ZFMK) ; India: 8♂ 1♀, north India, April to July ( BMNH) ; Bhutan: 2♂ ( BMNH) .

Distribution: China (Hunan, Taiwan, Sichuan, Yunnan, Tibet), India, Nepal, Bhutan, Kashmir district.

Remarks: A long series of material shows that the submarginal line varies in position from very close to the outer margin to much further from it, especially in the material from Yunnan ( Figs 13, 14 ) and Sichuan ( Fig. 15 ). This line in Comibaena pictipennis superornataria is also further from the outer margin ( Fig. 12 ), and the forewing tornal patch is a little more proximally placed than in most Indian specimens, but it falls well within the range of variation of Comibaena pictipennis , and no distinguishing features were found to separate Comibaena pictipennis superornataria consistently from material from elsewhere.

The male genitalia of Comibaena pictipennis from the type locality have broad arms to the vinculum ( Fig. 65 ), whereas most specimens from the northeastern Himalayas, which are externally indistinguishable, have narrower arms, although in a variety of shapes ( Figs 66–69 ). Even materials from the same locality, for example, Dahaoping, Tengchong, and Yunnan ( Fig. 13 ), which again have an identical wing pattern, differ slightly in the shape of the vinculum ( Figs 66 , 67 ).

The material from Baoshan, Yunnan ( Fig. 14 ), Nyingchi, Tibet, and Sichuan ( Fig. 15 ) are smaller compared to material from elsewhere, and externally similar to Comibaena nepalensis Yazaki , but the male genitalia are not identical to those of the latter. For example, the male genitalia of the material from Sichuan ( Fig. 69 ) have very narrow arms to the vinculum, which are not identical to those of Comibaena nepalensis , and the costal lobe is less strongly expanded than in Comibaena nepalensis .

Examination of this long series has not fully clarified the situation, and the large range of variation may even bring into question the separate specific status of Comibaena nepalensis , which is also externally very similar to Comibaena pictipennis . The absence so far of females to go with any of the smaller males is a little suspicious, and may suggest a single species with a high degree of variability in the male genitalia. By contrast, a complex of species may well be involved, and the series from Sichuan could be distinct. If females for these smaller Comibaena pictipennis are found in due course, they may be helpful in clarifying the species limits.

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 55–66. Male genitalia. 55, Comibaena quadrinotata; 56, Comibaena fuscidorsata; 57, Comibaena cassidara; 58, Comibaena attenuata; 59, Comibaena biplaga; 60, Comibaena apicipicta; 61, Comibaena striataria; 62, Comibaena delineata; 63, Comibaena dubernardi; 64, Comibaena ornataria; 65–66, Comibaena pictipennis, 65, from Darjeeling; 66, from Dahaoping, Tengchong, Yunnan. 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 176–187. Female genitalia. 176, Comibaena quadrinotata; 177, Comibaena cassidara; 178, Comibaena attenuata; 179, Comibaena biplaga; 180, Comibaena striataria; 181, Comibaena delineata; 182, Comibaena dubernardi; 183, Comibaena ornataria; 184, Comibaena pictipennis; 185, Comibaena parornataria; 186, Comibaena bellula; 187, Comibaena tancrei. Scale bars = 1 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 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.