Thetidia, BOISDUVAL, 1840

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 : 757-761

THETIDIA BOISDUVAL, 1840

Thetidia Boisduval, 1840: 189 . Type species: Thetidia plusiaria Boisduval, 1840 , by subsequent designation by Desmarest, 1857. Spain, Algeria.

Euchloris Hübner, 1823: 283 . Type species: Phalaena smaragdaria Fabricius, 1787 , by monotypy. Austria. [Junior homonym of Euchloris Billberg, 1820 (Coleoptera) .]

Aglossochloris Prout, 1912: 17 (key), 212. Type species: Phorodesma fulminaria Lederer, 1871 , by original designation. Iran: Asterabad .

Antonechloris Raineri, 1994: 365 . Type species: Phalaena smaragdaria Fabricius, 1787 , by original designation. Austria.

Adults: Antenna of male bipectinate, filiform terminally, filiform or shortly bipectinate in female. Frons not protruding, rough-scaled. Labial palpus developed, the first and second segments rough-scaled, the second segment in female elongate and the third segment not elongate. Hind tibia of male with two pairs of spurs in both, male and female, occasionally with terminal spurs only, terminal extension absent. Frenulum absent.

Wing shape and pattern. Apex of forewing blunt, that of hindwing protruding and round; outer margin smooth or slightly wavy. Wings green with white streaks. Forewing usually with white wavy or curved antemedial line, and dentate, slightly wavy or straight postmedial line, both lines weak or broad; submarginal line usually absent, sometimes composed of short, bar-like patches; terminal line present or absent. Hindwing usually paler than that of forewing, especially on costal area; postmedial lines absent, submarginal line usually weak and close to outer margin. Discal spot usually present, round or short bar-like, white, occasionally absent. Underside: colour similar to upperside; antemedial line absent on forewing; postmedial line present on both wings.

Third sternite of male abdomen with setal patches present or not. Eighth sternite with posterior margin notched medially; eighth tergite rounded posteriorly.

Male genitalia. Uncus bifid, tapering towards apex. Socii broader than uncus, tapering, sometimes blunt terminally, usually longer than uncus and bent outwards. Gnathos with two lateral arms not joined medially. Valva simple, without processes. Vinculum quite protruding, deeply bifurcate. Aedeagus long and slender, needle-like.

Female genitalia. Sterigma developed; antrum usually present. Seventh sternite elongate longitudinally, wrinkled and sclerotized. Ductus bursae very weakly sclerotized. Corpus bursae very weak; signum absent.

Diagnosis: In the wing pattern, Thetidia is distinguished from most species of Comibaena and Protuliocnemis by the lack of brown patches on the forewing tornal angle and the hindwing apex; unlike Chlorochromodes and Argyrocosma the antemedial and postmedial lines are entire, rather than composed of small dots on the veins. In both external appearance and in the male genitalia, species of Thetidia are very close to some species of Comibaena , Comibaena hypolampes , Comibaena cenocraspis , Comibaena latilinea , Comibaena swanni , and Comibaena birectilinea , all members of Thetidia having simple valvae similar to those of the species mentioned above. However, Comibaena species have a developed frenulum, whereas the frenulum is absent in Thetidia .

Biological notes: Hausmann (2001) gave detailed biological information on four European species, and stated that the larvae are different from the genera Comibaena and Proteuchloris in that they feed on low (perennial) plants and pupate in a flimsy cocoon between leaves of the host plant or near its base.

Host plants [from 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/) are all Compositae (Asteraceae) . The larvae of several species are oligophagous on Compositae : Th. albocostaria ( Artemisia vulgaris ; Dendranthema morifolium ), Th. plusiaria ( Santolina chamaecyparissus ; Artemisia herba-alba ), Th. smaragdaris [ Achillea millefolium ; Artemisia maritima ; Artemisia albinthium ; Artemisia nutans ; Artemisia austriaca ; Tanacetum (Chrysanthemum) vulgare ; Senecio ], Th. sardinica ( Achillea millefolium ; Santolina ).

Distribution: Mainly distributed in the Palaearctic region, a few species extending to India and west Africa.

THETIDIA ALBOCOSTARIA ( BREMER, 1864)

( FIGS 38 , 92 , 142 , 198 )

Euchloris albocostaria Bremer, 1864: 76 , pl. 6, figure 22. Syntype (s), Russia: east Siberia, Amur; Ussuri , in between Noor and Ema estuaries.

Phorodesma albocostaria: Staudinger, 1871: 144 .

Thetidia albocostaria: Inoue, 1961: 75

Diagnosis: Thetidia albocostaria is distinctive within Thetidia in having large, rounded, brown ring-like, white-centred discal spots on both sides of both wings, with the discocellulars inside the discal spots brown. The brown terminal line is also unusual in Thetidia . In the male genitalia, the uncus is basally fused and the branches extend beyond the blunt socii; the lateral processes of the vinculum are nearly triangular; the length of the uncus plus tegumen is equal to the length of the vinculum. The female genitalia have a labiate antrum.

The antenna is bipectinate in both sexes. The third sternite in the male has a pair of setal patches.

Material examined: China: 37♂ 20♀, northern to southern China, April to August ( IZCAS, BMNH) ; Russia: 2♂, Far East, July ( BMNH) ; Korean Peninsula : 1♀, July ( BMNH) ; Japan: 3♂ 1♀, July ( BMNH) .

Distribution: China (Heilongjiang, Jilin, Liaoning, Inner Mongolia, Henan, Shaanxi, Gansu, Qinghai, Jiangsu, Shanghai, Anhui, Zhejiang, Hubei, Hunan), Russia, Japan, Korean Peninsula.

THETIDIA CORRESPONDENS ( ALPHÉRAKY, 1883)

( FIG. 39 )

Phorodesma fulminaria var. correspondens Alphéraky, 1883: 157 , pl. 9, figure 85. Lectotype ♂, China: Ili Valley , 2000 ft, Kuldja [Yining (Gulja)]. (ZISP)

Aglossochloris correspondens: Prout, 1912: 213 .

Thetidia correspondens: Viidalepp, 1976: 845 .

Diagnosis: The antenna is bipectinate in both sexes. The hind tibia of both sexes have only terminal spurs. The anteromedial and postmedial lines of the forewing are crenulate, and broader than in other Chinese species, but similar to those of Th. plusiaria from Spain and Algeria. The submarginal line is composed of elongate white patches. The discal spot on the forewing is white, not forming a white ring as in Th. plusiaria . The hindwing is mostly white, with the marginal area green; the submarginal line is similar to that of the forewing. The uncus of the male genitalia is widely divided and curved inwards; the valva is broad. In the female genitalia, the sterigma is rhombic, posteriorly very broad; the antrum is located at the anterior half of the sterigma, curved and striplike. (Genitalia descriptions are taken from Hausmann, 2001.)

Material examined: China: 4♂ 1♀, north-west China, end June ( ZFMK); 1♀, Alexander Gebirge, end June ( ZFMK) .

Distribution: China (Xinjiang), Russia, Central Asia, Europe (Sarmatian, southern Urals, southern Volga Plain).

THETIDIA ATYCHE ( PROUT, 1935)

( FIGS 40 , 93 , 143 , 199 )

Euchloris atyche Prout, 1935: 18 , pl. 3: c. Holotype ♂, China (west): Siao-lou. (BMNH)

Thetidia atyche: Fletcher, 1979: 204 .

Diagnosis: The transverse lines are quite slender in the holotype ( Fig. 40 ), slightly broader in the paratype.

In the male genitalia, the uncus is bifid and V-shaped, the costa of the valva is slightly curved, and the vinculum is semicircularly concave at middle, with the lateral processes tapering but blunt. The third sternite in the male possesses a pair of setal patches. The posterior margin of the eighth sternite of the male is slightly sclerotized.

In the female genitalia, the sterigma is rounded, with the anterior margin deeply concave, connected with elongate sclerotization on the seventh sternite. The antrum is funnel-shaped, with the posterior part broadened, and its posterior margin is widely and shallowly concave. The ductus bursae is very short. The corpus bursae is quite long, but weak.

Type material examined: China: holotype, ♂, Siao- Lou, Chasseurs, Indigènes du P. Déjean, 1901, BMNH slide no. 21777 ( BMNH) .

Additional material examined: China: 2♂ 1♀, southwestern China, July ( IZCAS, BMNH) .

Distribution: China (Gansu, Sichuan).

THETIDIA CHLOROPHYLLARIA ( HEDEMANN, 1879)

( FIGS 41–42 , 94–95 , 144 , 166 , 200 )

Phorodesma chlorophyllaria Hedemann, 1879: 510 , pl. 3, figure 7. Syntype (s), Russia: Amur, Askold Island .

Phorodesma jankowskiaria Oberthür, 1879: 8 . Syntype (s), Russia: Askold .

Nemoria chlorophyllaria: Gumppenberg, 1895: 490 .

Euchloris chlorophyllaria: Staudinger, 1901: 263 .

Thetidia chlorophyllaria: Inoue, 1961: 75 .

Euchloris pekingensis Chu, 1981: 116 , pl. 30, figure 791, Holotype ♀, China: Beijing. (IZCAS) syn. nov.

Thetidia pekingensis: Han & Xue, 2011 , Fauna Sinica: Insecta 54: 163, pl. 15. 8.

Diagnosis: The antemedial line of the forewing is approximately vertical to the inner margin and roundly curved inwards onto the costa. In the male genitalia, the length of the uncus plus tegumen is equal to the length of the vinculum; the vinculum is deeply concave at middle; the gnathos is very broad compared to other Chinese representatives of the genus. In the female genitalia, the sterigma is roundly protruding; the antrum is rounded, with two tiny lateral teeth on the posterior margin and two blunt protrusions on the anterior margin; the ductus bursae is quite long, with the central part swollen. The corpus bursae is quite weak.

The third sternite in the male possesses a pair of weak setal patches. The posterior margin of the eighth tergite is slightly concave, and that of the sternite is flat.

Type material examined: Euchloris pekingensis : China: holotype, ♀, Beijing: Baihuashan, 9.vii.1973, coll. Han Yinheng ( IZCAS) .

Additional material examined: China: 37♂ 55♀, northern to south-western China, May to August ( IZCAS, ZFMK, BMNH) ; Russia: 2♂ 5♀, Far East, Siberia, July ( BMNH) .

Distribution: China (Heilongjiang, Inner Mongolia, Beijing, Hebei, Shanxi, Shandong, Shaanxi, Gansu, Qinghai, Sichuan), Russia, Japan.

Remarks: The single specimen of Th. pekingensis differs externally from other species of Thetidia in that the antemedial and postmedial lines are broadened near the anal fold and connected to each other, appearing as two crossed lines. However, as the female genitalia are identical to those of Th. chlorophyllaria , and no further specimens have turned up in the type locality over many years, we think it likely that this is an aberration of Th. chlorophyllaria , and consequently synonymize it with the latter.

THETIDIA KANSUENSIS ( DJAKONOV, 1936)

FIGS 43–44 , 96–97 , 145 , 167 )

Euchloris kansuensis Djakonov, 1936: 3 , pl. 1a: 8. Syntypes 2♂, China: Kansu (south), Bandshuka. (NHRS)

Thetidia kansuensis: Scoble, 1999: 936 .

Diagnosis: In wing pattern, Th. kansuensis is similar to Th. chlorophyllaria , but the forewing postmedial line is much closer to the apex, and the hindwing is much paler, in the type material. The male genitalia of the two species also differ in the size of the vinculum, which is less than the length of the uncus plus tegumen in Th. kansuensis but of equal length in Th. chlorophyllaria .

Type material examined: Euchloris kansuensis : 1♂, syntype, Kina: S. Kansu, Sven Hedins Exp. Ctr. Asien, Dr. Hummel (NHRS) .

Additional material examined: China: 6♂, Tibet, July to August ( IZCAS) .

Distribution: China (Gansu, Tibet).

Remarks: The material from Tibet has some external differences from the material from Gansu in that the postmedial line on the forewing is closer to the apex, and the hindwing is much darker. However, they have almost identical male genitalia, and we consider them to be one species.

THETIDIA SMARAGDARIA ( FABRICIUS, 1787)

FIGS 45 , 98 , 146 , 201 )

Phalaena smaragdaria Fabricius, 1787: 192 . Syntype (s), Austria.

Geometra smaragdaria: Villers, 1789: 499 .

Euchloris smaragdaria: Hübner, 1826: 283 .

Hemithea smaragdaria: Duponchel, 1829: 251 , pl. 152: 5.

Hipparchus smaragdarius Curtis, 1830: 300 .

Phorodesma smaragdaria: Boisduval, 1840: 179 .

Thetidia smaragdaria: Viidalepp, 1976: 845 .

Diagnosis: Thetidia smaragdaria is one of the three Chinese species that have discal spots on the forewing. The other two species are Th. albocostaria and Th. correspondens . However, the discal spots of Th. smaragdaria , which are white dots, differ from those of Th. albocostaria , which are large and brown ringed, and from the elongate white patches of Th. correspondens . The male genitalia are similar to those of the congeners, except that the uncus is widely bifid; the lateral branches of the vinculum vary from narrow to blunt; and the length of the vinculum is distinctly shorter than the length of the uncus plus tegumen. In the female genitalia, the sterigma is almost quadrangulate; the antrum is cup-like, with the posterior margin concave; the ductus bursae is quite long; and the corpus bursae is long and weak.

The antenna is bipectinate in both sexes. The third sternite in the male possesses weak setal patches. The posterior margin of the male eighth sternite is slightly concave at the middle.

Material examined: China: 17♂ 8♀, northern and southern China, May to August ( IZCAS, ZFMK) ; Europe : 1♂ 1♀, June, August ( BMNH) ; Asia Minor: 1♂ 1♀ ( BMNH) .

Distribution: China (Inner Mongolia, Xinjiang, Jiangsu), Russia, Mongolia, Japan, central and western Asia, India, Europe. Hausmann (2001) gave detailed distribution information on several subspecies.

Remarks: The materials examined in the collection of IZCAS most closely resemble Thetidia smaragdaria anomica (Prout) in wing pattern, but the male genitalia differ from BMNH slide Geom 21781 (from Issyk-kul, the type locality) in the lateral branches of the vinculum, which are narrow and blunt in BMNH slide Geom 21781 but hooked in the materials from Inner Mongolia. Four subspecies of Th. smaragdaria have been described in addition to the nominate subspecies. Viidalepp (1996) and Hausmann (2001) treated Thetidia volgaria as one subspecies of Th. smaragdaria ; Hausmann (2001) also noted that the status of Th. smaragdaria volgaria is not clear. In our opinion the subspecies is not distinct from nominate Th. smaragdaria .

Species not occurring in China

Thetidia albosagittata ( Ebert, 1965)

Aglossochloris albosagittata Ebert, 1965: 5 , pl. 1, figure 4; pl. 2, figure 4. ( Afghanistan)

Thetidia bilineata Hausmann, 1991

Thetidia bilineata Hausmann, 1991: 117 , pl. 1, figure 8; pl. 10, figure 72. ( Jordan)

Thetidia crucigerata ( Christoph, 1887)

Phorodesma crucigerata Christoph, 1887: 93 , pl. 4, figure 11. ( Turkmenistan)

Thetidia euryrithra ( Prout, 1935)

Aglossochloris euryrithra Prout, 1935: 18 , pl. 3: c. ( Jordan)

Thetidia fulminaria ( Lederer, 1870) [specimen(s)]

Phorodesma fulminaria Lederer, 1870: 17 , pl. 2, figure 4. ( Iran)

Thetidia hammeri ( Ebert, 1965)

Aglossochloris hammeri Ebert, 1965: 5 , pl. 1, figure 5; pl. 2, figure 5. ( Afghanistan)

Thetidia hazara ( Ebert, 1965)

Aglossochloris hazara Ebert, 1965: 4 , pl. 1, figure 3; pl. 2, figure 3. ( Afghanistan)

Thetidia mabillei ( Thierry-Mieg, 1893)

Phorodesma mabillei Thierry-Mieg, 1893: 162 . ( Russia)

Thetidia pallidmarginata ( Pajni & Walia, 1984)

Euchloris pallidmarginata Pajni & Walia, 1984: 176 , figures 1–6. ( India)

Thetidia persica Hausmann, 1996 [male and female genitalia photographs, Hausmann, (2001: fig. 24c)]

Thetidia persica Hausmann, 1996: 27 , figures 12, 114–116, 147. ( Iran)

Thetidia plusiaria Boisduval, 1840 [specimen(s); male and female genitalia]

Thetidia plusiaria Boisduval, 1840: 189 . ( Spain; Algeria)

Thetidia radiata Walker, 1863 .

Thetidia radiata Walker, 1863: 1567 . ( India)

Thetidia recta ( Brandt, 1941)

Aglossochloris recta Brandt, 1941: 865 , pl. 28, figure 14. ( Iran)

Thetidia sardinica ( Schawerda, 1934) [specimen(s); male and female genitalia photographs, Hausmann (2001: fig. 25)]

Euchloris sardinica Schawerda, 1934: 463 , fig. 5. (Sardinia)

Thetidia serraria ( Staudinger, 1892) [specimen(s)]

Phorodesma smaragdaria var.? serraria Staudinger, 1892: 141 . (central Asia)

Thetidia silvia Hausmann, 1991

Thetidia silvia Hausmann, 1991: 117 , pl. 1, figures 10, 11; pl. 10, figure 71. ( Jordan)

Thetidia smaragdularia ( Staudinger, 1892)

Phorodesma smaragdularia Staudinger, 1892: 142 . (Central Asia)

Euchloris ? viridifrons Warren, 1897b: 389. ( Afghanistan)

Thetidia undulilinea ( Warren, 1905) [specimen(s)]

Eucrostes undulilinea Warren, 1905: 384 . ( Sierra Leone)

PROTEUCHLORIS HAUSMANN, 1996

( FIGS 46 , 99 , 147 , 202 )

Proteuchloris Hausmann, 1996: 24 . Type species: Geometra neriaria Herrich-Schäffer, 1852 .

Adults: Antenna of male with basal four-fifths bipectinate with long rami; dentate with very short forked branches in female. Frons flat. Labial palpus long, with second segment bushy-scaled, third segment naked. Hind tibia of male with two pairs of spurs and terminal extension. Frenulum absent.

Wing shape and pattern. Apex of forewing blunt, that of hindwing round. Wings green. Forewing with antemedial line slightly wavy; postmedial line angled at upper half, sinuous near inner margin; a white ringed brown tornal patch present. Hindwing with narrow, white-ringed brown apical patch and a small whitish anal patch. Both wings with small discal spots. Underside: pale green, lower area of forewing and hindwing greenish white; both wings with indistinct postmedial lines; discal spot present on both wings.

Posterior and anterior edge of sternite 8 strongly sclerotized, notched medially on posterior edge; tergite 8 rounded posteriorly.

Male genitalia. Uncus bifid, distinctly tapering. Socii curved. Gnathos weak. Valva simple. Vinculum deeply bifurcate. Coremata absent. Aedeagus long, slender, straight.

Female genitalia. Antrum round to subrhombic, laterally slightly furrowed. Signum absent.

Diagnosis: The absence of the frenulum and the simple valva are identical to those of Thetidia , whereas the terminal extension of the hind tibia is absent in Thetidia but present in Proteuchloris .

Biological notes: Hausmann (2001) illustrated the larva of Proteuchloris neriaria and stated that this species feeds on Quercus .

Distribution: East-Mediterranean, Crete, Turkey, Armenia, Cyprus, Levant.

Species not occurring in China

Proteuchloris neriaria (Herrich-Schäffer, 1852) [specimens, photographs, male and female genitalia] Geometra neriaria Herrich-Schäffer, 1852: 62 .

( Greece)

Figures

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 91–102. Male genitalia. 91, Microbaena pulchra; 92, Thetidia albocostaria; 93, Thetidia atyche; 94–95, Thetidia chlorophyllaria; 96–97, Thetidia kansuensis; 98, Thetidia smaragdaria; 99, Proteuchloris neriaria; 100, Linguisaccus subhyalina; 101, Linguisaccus minor; 102, Chlorochromodes rhodocraspeda. Scale bars = 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.

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 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 200–210. Female genitalia. 200, Thetidia chlorophyllaria; 201, Thetidia smaragdaria; 202, Proteuchloris neriaria; 203, Linguisaccus subhyalina; 204, Linguisaccus minor; 205, Chlorochromodes rhodocraspeda; 206, Argyrocosma inductaria; 207, Protuliocnemis castalaria; 208, Protuliocnemis falcipennis; 209, Protuliocnemis candida; 210, Protuliocnemis dissimilis. Scale bars = 1 mm.