Taxonomic Review of the Genus Pseudostegania Butler, 1881, with Description of Four New Species and Comments on Its Tribal

Entomological Science (2010) 13, 234–249 doi:10.1111/j.1479-8298.2010.00379.x

ORIGINAL ARTICLE

Taxonomic review of the genus Pseudostegania Butler, 1881, with description of four new species and comments on its tribal

placement in the Larentiinae (Lepidoptera: Geometridae)ens_379 234..249

Hongxiang HAN,1 Dieter STÜNING2 and Dayong XUE3 1,3Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China; and 2Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, Bonn, Germany

Abstract The genus Pseudostegania Butler is revised. In addition to the two species known, four species are described as new: P. lijiangensis sp. nov. from Yunnan Province, P. qinlingensis sp. nov. from Gansu and Shaanxi Provinces, P. zhoui sp. nov. from Sichuan Province (China) and P. burmaensis sp. nov. from Burma (Myanmar); two species are newly combined with Pseudostegania: P. distinctaria (Leech, 1897), comb. nov. and P. yargongaria (Oberthür, 1916), comb. nov. All the known species are redescribed and lectotypes are designated for P. defectata (Christoph, 1881), P. distinctaria and P. yargongaria. The generic characters, based on all species, are summarized. The tribal placement of Pseudostegania is discussed. Illustrations of moths and genitalia are presented. Key words: distribution, female genitalia, male genitalia, morphology, redescription.

INTRODUCTION their work on the tribe Asthenini and associated genera. This is because both species neither agree with the The genus Pseudostegania was described by Butler generic characters of Asthena nor belong to the tribe (1881), based on P. chrysidia Butler, 1881 from Japan, Asthenini, but they did not point out the genus where which was treated as a junior subjective synonym of these two species should be placed. In the present paper, P. defectata (Christoph, 1881) by Inoue (1977) and they are combined with the genus Pseudostegania, based Fletcher (1979), and was considered as a subspecies of P. on genitalia characters and external features. defectata by Inoue (1982). Prout (1914) hinted a pos- In recent years, new material has been collected or sible relationship of defectata with Laciniodes plurilin- discovered in collections and new taxa need to be earia Moore, 1868, but he later placed defectata into described. The purposes of this paper are to describe Asthena (Prout, 1938), just mentioning that face and four new species of Pseudostegania and to transfer palps agree better with members of Asthena. Leech another two species to this genus; to provide diagnostic (1897) described another two species, Hydrelia stra- generic characters in relation to other genera; to ensure minearia and Hydrelia distinctaria, from western China, nomenclatural stability for three species by designating and both species were transferred to Asthena by Prout lectotypes; and to redescribe all the known species. In (1914), repeated by Scoble (1999). Xue and Zhu (1999) addition, the tribal placement of Pseudostegania is dis- combined straminearia with Pseudostegania. Asthena cussed and illustrations of external features and genita- distinctaria together with Asthena yargongaria, the lia of all taxa are provided. latter described by Oberthür (1916) from Tibet, were excluded from Asthena by Xue and Scoble (2002) in MATERIALS AND METHODS Specimens studied were obtained from the Institute Correspondence: Dayong Xue, Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese of Zoology, Chinese Academy of Sciences, Beijing, Academy of Sciences, 1 Beichen West Road, Chaoyang China (IZCAS); Zoologisches Forschungsmuseum District, Beijing, 100101, China. Email: [email protected] Alexander Koenig, Bonn, Germany (ZFMK), The Received 21 August 2009; accepted 11 November 2009. Natural History Museum, London, UK (NHM),

Museum für Naturkunde der Humboldt-Universitat, black brown patches; two lines sometimes conﬂuent Berlin, Germany (MNHU) and China Agricultural or hardly differentiated; submarginal line double, the University (CAU). distal one usually diffuse or consisting of a series of Photographs of specimens were taken using various patches; subterminal or terminal lines composed of digital cameras and microscopes. Terminology of wing patches or dots between veins, sometimes indistinct. venation follows the Comstock–Needham System Hindwing: basal line present; medial line usually (Comstock 1918) and that of the genitalia is based on straight; postmedial line double, the outer one generally Pierce (1914), Klots (1970) and Nichols (1989). The more strongly waved or curved; submarginal line built digital images were enhanced and the plates compiled as a double line, or a series of patches; subterminal or using Photoshop software (Adobe Systems, San Jose, terminal lines similar to that of the forewing. Discal CA, USA). spot present on forewing and hindwing. Underside: ornamentation similar to upperside, but usually more dull, indistinct, and browner; suffusion of forewing TAXONOMY stronger on costal basal area, along postmedial and Pseudostegania Butler submarginal areas and apex. Venation (Fig. 33). Forewing with two areoles, M Pseudostegania Butler, 1881, Trans. Ent. Soc. Lond. 1 fused with lower margin of second areole at base, M 1881: 416. 2 arising from above middle of discocellulars, CuA1 not

Type species. Pseudostegania chrysidia Butler, 1881 stalked with M3. Hindwing with Sc+R1 long-stalked

(September)–(a junior synonym of Cidaria defectata with upper vein of cell, Rs long-stalked with M1, dis-

Christoph, 1881 (April) Bull. Soc. Imp. Nat. Moscou, cocellulars not angled, M2 a little closer to M1,M3 and

55 (3): 108. CuA1 separate, 3A absent. Redescription. Head and body. Antennae simple, ﬁli- Male genitalia. Tegumen large, with a ﬂattened distal form in both, male and female, upper surface of each extension, with rounded posterior margin. Extended segment scaled, often chequered pale and dark brown, part covering the reduced, knob-like uncus. On ventral lower surface shortly ciliate, yellow-brown or brown. side of tegumen there is a pair of lateral sclerites which Frons slightly protruding, smooth-scaled, whitish to may be fused basally or not. This seems to be a reduced, pale yellow; sometimes lower half black brown; upper not functional gnathus. Subscaphium basally with a margin often with a dark brown patch or a narrow slender shaft, expanded posterior part spinose. Valva band; width of central part almost equal to or slightly often narrow sub-basally, expanded and setose apically, broader than horizontal diameter of compound eye. costa sclerotized, with an oval subapical broadening, Labial palpus very short, rough-scaled, 3rd segment slightly protruding and also setose, very vague in P. nearly invisible; upper surface whitish brown to dark burmaensis and of a different shape, rather pouch-like in brown, ventral and inner surface pale brown or whitish. P. zhoui; sacculus often not distinct, sometimes with a Vertex whitish to pale yellow. Chaetosemata arranged as pointed process (P. qinlingensis, P. yargongaria); saccus a transverse band. Patagia whitish to pale yellow, mixed well-developed and broad, protruding. Transtilla a pair with black brown scales, or occasionally with a dark of small, acute sclerites connected with extraordinary brown transverse band. Tegulae white, with base black labides, the latter consisting of narrow, sclerotized shafts brown. Thorax with dorsal and ventral sides whitish. with ovally expanded, membranous and setose apices, Abdomen whitish to yellowish, sometimes with black the latter with narrow and long openings, along the brown intersegmental scales. margin of which the setae are stronger and arranged Wing shape and pattern. Forewing with costa slightly more densely, sometimes the distal parts of the setae curved, apex rounded or slightly pointed. Hindwing are broadened scale-like; the shafts are arising from the with apex and anal angle rounded, costa and inner base of the valva and are connected dorsally with the margin almost of the same length. Outer margin of transtilla by short, rod-like branches, ventrally longer both wings slightly wavy, protruding. Wings whitish branches are extending downwards to the juxta. Their to pale yellow. Forewing: yellow costal and terminal basal ends may be fused or not. Juxta a large, sclero- bands present or not; basal and antemedial lines tized, simple, pouch-like process, with posterior margin sinuous, curved, or straight and angled at posterior pointed or blunt. Anellus densely spinulose, small spines margin of costal band; medial line close to postmedial mixed with long and strong spines. Aedeagus broad, the area, similar trend as postmedial lines; postmedial lines shaft densely spinulose apically, vesica with a large, usually double, oblique and angled at posterior margin pointed cornutus and a narrow, elongate, slightly scle- of costa band, or curved, forming one to two distinct, rotized diverticulum.

Female genitalia. Ovipositor short, with lobes surfaces whitish. Vertex whitish yellow, concolorous oval, rather weak, moderately setose. Often a narrow with major part of frons. Patagia mottled with black antevaginal, sclerotized transverse band present, broad- brown scales. Tegulae whitish yellow, their bases black ening laterally, sometimes modiﬁed or absent. Antrum brown. Thorax whitish yellow dorsally and ventrally. large. Ductus bursae short and broad, usually with Abdomen colored similarly, with dorsum slightly darker a distinct, but sometimes rather weakly sclerotized yellow and venter more whitish, intersegmental areas colliculum. Corpus bursae large, usually irregularly darker. expanded laterally or subposteriorly, densely spinulose Length of forewing: ǩ13–14 mm, Ǩ14–15 mm. at the larger part of the internal surface, with longitu- Wings whitish yellow to yellow. Forewing: yellow dinal, sclerotized bands bordering this area towards costal and terminal bands indistinct, not strongly con- a smaller membranous part. A similarly membranous trasting; basal costal area densely suffused with dark diverticulum, giving rise to the ductus seminalis, present brown scales; basal line narrow, sinuous; antemedial near the ductus bursae. Also near the ductus bursae, at line similar, protruding twice, sharply angled at centre opposite side, is a shallowly concave, sclerotized area, of cell near discal spot, broadening near costa; medial opening towards a scobinate region of the membranous line weak, evenly curved outwards; postmedial area part of the bursa, close to the signum. The latter is consisting of two lines, the proximal one very vague, usually a pair of large, elongate, spinose bands or the distal one forming a sharp, outwardly directed patches, symmetrical, connected posteriorly only or over angle near M1 and a more rounded one near M3. the whole length, then with a longitudinal ridge, some- Costal half of postmedial area more or less strongly times rather small, roundish, with an anterior indenta- suffused with black brown scales. Submarginal area tion, occasionally (in P. burmaensis) absent. Spines of consisting of two lines, the proximal one continuous signum pointing inwards. from costa to hind margin, slightly wavy at costal half, Distribution. East Asia, including Myanmar, China somewhat broadening and pointing outwards near

(Jilin, Inner Mongolia, Beijing, Shaanxi, Gansu, Shanxi, M1 and M3; the distal line vague, with a strong brown

Yunnan, Sichuan, Tibet), Japan, Korea and Far East angular patch between R5 and M1, approaching the

Russia. proximal line between M3 and CuA1; terminal line often composed of small, black brown dots near Pseudostegania defectata (Christoph, 1881) margin between veins; often (especially abundant in (Fig. 1) Japanese specimens = ssp. chrysidia Butler) there is a Cidaria defectata Christoph, 1881, Bull. Soc. Imp. dark brown band connecting the postmedial and the

Nat. Moscou 55 (3): 108. Lectotype ǩ, [Russia]: submarginal area along vein M3 (Fig. 1); discal spot a Vladivostok. (MNHU) Here designated. small, round, black brown dot. Apex of forewings Larentia defectata: Staudinger, 1901, in Staudinger & often with a strong, black brown spot, internally Rebel, Cat. Lepid. Palaearct. Faunengeb. 1: 306. rounded, externally acute, sometimes reduced to a Asthena defectata: Prout, 1914, in Seitz, Macrolepid. streak or absent; fringe at apex dark brown, otherwise World 4: 273, pl. 10:h. whitish-yellow, shining. Hindwing: basal line very close Pseudostegania defectata: Inoue, 1959, Icon. Insect. to wing base, rather broad; medial line nearly straight Jap. Col. Nat. Ed.1 (Lepid.): 196, pl. 138: 26. from hind margin to discal spot, often more proximal Pseudostegania chrysidia Butler, 1881, Trans. Ent. Soc. to it; postmedial line double, distance between them Lond. 1881 (3): 417. Syntype(s), Japan: Tokyo. largest at middle, proximal line often vague or even (NHM) (synonymized with defectata by Inoue, absent; submarginal lines also double, but closer 1977, p. 272, but not stated as syn. nov.) together and parallel, distal line considerably weaker. Pseudostegania defectata chrysidia: Scoble, 1999, Terminal line appears as dots between veins. Fringe Geometrid Moths of the World, a Catalogue 2: 799. and discal spots similar to those of forewing. Under- Redescription. Antennae with ventral surface shortly side: ground color less yellowish, lines similar, but ciliate, yellow-brown, upper surface scaled, each seg- rather simple than double in hindwing, partly more ment chequered dark and pale brown at basal half diffuse, dark brown pattern near apex of forewings of antennae, terminal half with upper surface pale and costal area of submarginal lines more extended brown. Frons slightly protruding, smooth-scaled, and conﬂuent, terminal line composed of small dots whitish yellow, with a narrow black-brown band at between veins more distinct than on upperside. upper margin, and a broad, black-brown band at bottom. Male genitalia (Figs 9,17). Tegumen very large, Labial palpus very short, rough-scaled, with upper rounded, posterior margin slightly concave at middle, and lateral surfaces black brown, ventral and inner basally with a pair of large lateral, fused sclerites. Uncus

Figures 1–8 Adult of Pseudostegania. 1 P. defectata (Christoph), ǩ; 2 P. straminearia (Leech), Ǩ; 3 P. distinctaria (Leech), ǩ; 4 P. qinlingensis sp. nov. paratype, ǩ; 5 P. yargongaria (Oberthür), ǩ; 6 P. lijiangensis sp. nov., paratype, ǩ; 7 P. zhoui sp. nov., holotype, ǩ; 8 P. burmaensis sp. nov., paratype, Ǩ. Scale bars, 10 mm.

Specimens examined. Russia: 1ǩ (MNHU), Lectotype, here designated for reason of nomenclatural stability, “Wladiwostock”, “ex coll. Staudinger”, other labels: “Origin.”, “Cidaria defectata Chr.”, “defectata Chr.”; 1ǩ1Ǩ (MNHU), paralectotypes, [Wladiwostock], “ex coll. Staudinger”, other label: “Origin”; 1Ǩ (Zoological Institute, St. Petersburg), paralectotype, “Wladiw.[os- tok]” – “26./ 14.vi.[18]77 Chr.[istoph]” (digital images checked); 1ǩ1Ǩ (NHM), Narva. S. Ussurigebiet, 24, 28.vii.[19]21, N. Kardakoff, Joicey Bequest. Brit. Mus. 1934-120; 1ǩ (NHM), same locality and collector, 28.vii.[19]21, Rothschild Bequest B.M. 1939-1, ﬁgd. Seitz Supp. IV. 16i; 1ǩ (NHM), same locality and collector, 25.vii.[19]21, L.B.Prout Coll. B.M. 1939-643; 1ǩ (NHM), ex coll. Christoph. Vladivostok, 3.vi.[18]77, Ex. Coll. H.J. Elwes, 1920, Joicey Bequest Brit. Mus. 1934-120; 1Ǩ (NHM), Mt. Usui, near Karuisawa, 22.viii.1922, I. Sugitani, Joicey Bequest. Brit. Mus. 1934-120; 1Ǩ (NHM), Russ. Inseln, S. Ussurigebiet, 14.vii.1920, N. Kardakoff; 1ǩ (ZFMK), Wladiwostock; 1Ǩ (ZFMK), Amur, 1888; China: 1ǩ (IZCAS), Jilin: Changbaishan, 7.vii.1974; 1Ǩ (IZCAS), Shanxi: Huoxian, 28.vii.1972, coll. Wang Shuyong; 3Ǩ (IZCAS), Shaanxi: Taibai Huangbaiyuan, 10–12.vii.1980, coll. Zhang Baolin; 17ǩ4Ǩ (ZFMK), Tapaishan im Tsinling, Sued-Shensi, H. Höne, 1ǩ1Ǩ, 20.v.1936, 1700 m; 1ǩ, 16.vi.1936, 1ǩ, 28.vi.1935, 1ǩ, 30.vi.1935, all 1700 m; 2ǩ, 3.vii.1935, about 3000 m, 1ǩ, 10.vii.1936, 1700 m; 5ǩ1Ǩ, 13.vii., 23.vii., 25.vii., 26.vii., ca. 3000 m; 5ǩ2Ǩ, 5.viii., 10.viii., 12.viii.1936, 1700 m and about 3000 m; 1Ǩ (ZFMK), Mien-shan (Prov. Shensi), Obere Höhe about 2000 m, 28.vii.1937, Figure 33 Venation of P. defectata (Christoph). H. Höne; 1ǩ (CAU), Beijing: Mentougou Xiaolongmen, 30.vii.1992, coll. Yang Jikun & Liu Zhiqi; 1Ǩ (ZFMK), small, rounded, slightly sclerotized. Subscaphium Sidemi, 1889, ex coll. Staudinger; 1Ǩ (ZFMK), Mand- sclerotized, posterior half expanded and spinose. Valva schourie; 1ǩ (NHM), AMOOR, Rothschild Bequest, narrow at basal half, expanded and setose apically; B.M. 1939-1; Japan: 1ǩ1Ǩ (IZCAS, donated by K. costa sclerotized, ovally broadening and forming a Yazaki), Saiki-rindo, 1500 m, Yanagisawa–toge (Yama- curved ridge subapically. Saccus well developed and nashi Pref.), 7, 14.vii.1979, K. Yazaki leg; 1Ǩ (NHM), broad, semicircular. Labides with scale-like setae at pos- type of chrysidia, Tokyo, 80.97 (with red type label); terior margin of opening. Juxta bluntly tapering. Anellus 1ǩ (NHM), Japan, B. Bristowe, 1901-317; 1ǩ1Ǩ densely spinulose and mixed with long spines. Aedeagus (NHM), Gensan, vii.1887, Leech, Leech Coll. 1900-64; as in generic description. 1Ǩ (NHM), H. Pryer Coll. Japan, Leech Coll. 1900-64; Female genitalia (Fig. 25). Ductus bursae short and 2Ǩ (NHM), Japan, Rothschild Bequest B.M. 1939-1; broad, posterior half with slightly sclerotized collicu- 1Ǩ (NHM), Manley, 1890, Leech Coll. 1900-64; lum. Corpus bursae large, roundly expanded laterally 1ǩ (NHM), Kami kochi, 4.viii.1953, T. Haruta Lt. and medially, irregularly folded, partly sclerotized, Brit.Mus.1954-498; 2Ǩ (NHM), E.A. Cockayne, 2, about half of its internal surface densely spinulose, the 4.ix.1910, Nikko, Japan, L.B.Prout Coll. B.M.1939-643; other half membranous, both areas not separated by 3ǩ1Ǩ (ZFMK), Japan, Yokohama, 1891 (ex coll. Ober- longitudinal, sclerotized bands. Signum a pair of elon- thür, ex coll. Wehrli), 1Ǩ (ZFMK), [Japan], Chuzenyi, gate, symmetrical, spinose bands, connected posteriorly, 7.9., ex coll. Weber 13/57. spines arranged regularly. Membranous diverticulum Biological note: The larvae have been described by rather small. Sato in Sugi (1987, p. 66, 273) and two color morphs are

Figures 9–16 Male genitalia of Pseudostegania. 9 P. defectata (Christoph), MNHU; 10 P. straminearia (Leech), NHM; 11 P. distinctaria (Leech), NHM; 12 P. qinlingensis sp. nov., IZCAS; 13 P. yargongaria (Oberthür), IZCAS; 14 P. lijiangensis sp. nov., IZCAS; 14a scale-like setae on labides, of P. lijiangensis; 15 P. zhoui sp. nov., IZCAS; 16 P. burmaensis sp. nov., NHM. Scale bars, 1 mm.

Figures 17–24 Aedeagus. 17 P. defectata (Christoph); 18 P. straminearia (Leech); 19 P. distinctaria (Leech); 20 P. qinlingensis sp. nov.; 21 P. yargongaria (Oberthür); 22 P. lijiangensis sp. nov.; 23 P. zhoui sp. nov.; 24 P. burmaensis sp. nov. Scale bars, 1 mm.

figured on pl. 26, figures 15,16. It is stated that the host Pseudostegania straminearia: Xue & Zhu, 1999, Fauna plant of P. defectata is a herb Isodon kameba (Labiatae). Sinica: Insecta 15: 782, figs 958, 959, pl. XXI:9. Distribution. China (Heilongjiang, Jilin, E. Inner Mon- Redescription. Antennae darker towards apex, basal golia, Beijing, Shaanxi, Shanxi), Japan, Korea, Russia one third chequered, with upper surface of each seg- (Amur district, Ussuri district, Vladivostok region). ment pale at base and dark grayish-brown at tip. Frons slightly protruding, smooth-scaled, upper two-thirds Pseudostegania straminearia (Leech, 1897) pale yellow, lower third black brown. Labial palpus very (Fig. 2) short, rough-scaled, with upper and outer surfaces black Hydrelia straminearia Leech, 1897, Ann. Mag. Nat. brown, ventral and inner surfaces whitish. Vertex, Hist. (6) 20: 79, Holotype ǩ (NHM), China patagia, tegulae and thorax dorsally and ventrally pale (western): [Sichuan]: Wa-shan. yellow. Abdomen with dorsal and ventral sides whitish Asthena straminearia: Prout, 1914, in Seitz, Macrolepid. yellow, ventrally paler, occasionally mottled with very World 4: 273. few black brown intersegmental scales.

Length of forewing: ǩ14–16 mm, Ǩ14–17 mm. 2Ǩ (NHM), Siao-Loû, Chasseurs Indigènes du P. Wings whitish-yellow to yellow. Forewing: yellow Déjean, 1903, Ex Oberthür coll. Brit. Mus. 1927-3; costal and terminal bands not present; basal costal 1Ǩ (NHM), Tu-pa-Keo 7400 ft. 4.ix.1929, Szechwan, area slightly suffused with dark brown scales; basal Kelley-Roosevelt expedition; Rothschild bequest BM line rather straight posteriorly, sharply angled at ante- 1939-1 (other labels: Asthena straminearia Leech (in rior margin of cell; antemedial line evenly curved, Prout’s handwriting); Geometridae genitalia slide no. slightly sinuous, broadening near costa; medial line 20084); 1ǩ (IZCAS), Yunnan: Tengchong Dahaoping, weak, wavy; postmedial area consisting of two, largely 2020 m, 24–26.v.1992, coll. Xue Dayong; 1ǩ1Ǩ conﬂuent lines, forming two sharply pointed protuber- (IZCAS), Yunnan: Tengchong Heinitang, 1930 m, ances at M1 and above CuA1, the latter slightly larger 28–30.v.1992, coll. Xue Dayong; 2ǩ1Ǩ(ZFMK), than the ﬁrst. Space inside the protuberances strongly Siao-Loû, Chasseurs Indigènes du P. Déjean, 1903, ex suffused with dark brown scales; submarginal area coll. Oberthür, ex coll. Wehrli. with two weak, strongly wavy lines, separate, but Distribution. China (Sichuan, Yunnan). approaching each other near hind margin; terminal line composed of a series of tiny brown dots between Pseudostegania distinctaria (Leech, 1897) the veins, often indistinct; fringe whitish-yellow, comb. nov. (Fig. 3) shining; discal spot a small, black brown dot. Hind- Hydrelia distinctaria Leech, 1897, Ann. Mag. Nat. Hist. wing: lines slightly less distinct than those of forewing; (6) 20: 80. Lectotype ǩ, China (western): [Sichuan]: basal line thick, black brown; medial line nearly Pu-tsu-fong, Wa-Shan (NHM) Here designated. straight from hind margin to discal spot; postmedial Asthena distinctaria: Scoble, 1999, Geometrid Moths of lines double, diverging at middle, without dark brown the World, a Catalogue 1: 72. suffusion, distal line distinctly wavy; submarginal and terminal lines similar to those of forewing; discal spot Redescription. Antennae yellow on ventral side, a little smaller than that of forewing. Fringe as in upperside pale grayish-brown, on basal one third each forewing. Underside: Transverse lines similar to upper- segment chequered pale and dark grayish-brown. Frons side, partly less distinct, but dark brown suffusion slightly protruding, smooth-scaled, buffy white, with a of forewing stronger, especially on basal region near narrow black brown band at upper margin in front costa, along anterior part of submarginal line and on of antennae and a similar band, but of variable size at apex; a weak, brown terminal line present on forew- lower margin. Labial palpus very short, rough-scaled, ing, dots between veins on hindwing more distinct. with upper and lateral surfaces black brown, ventral and Male genitalia (Figs 10,18). Similar to defectata, but inner surfaces whitish. Vertex whitish yellow. Patagia tegumen much shorter and narrower at apex; valva with whitish. Tegulae whitish yellow, its bases black brown. apex not expanded as distinctly as in defectata; oval Thorax with dorsal and ventral sides whitish yellow, broadening of costa less developed and truncate api- abdomen of the same color, dorsum with sparse black cally; saccus larger and more strongly protruding; juxta brown scales on intersegmental borders. slightly broader posteriorly and spinulose area of anellus Length of forewing: ǩ 14–16 mm. Wings white with smaller than in defectata. Aedeagus as in the generic a weak creamy tinge, forewing with a broad yellow description, diverticulum not everted in Figure 18. costal band, gradually broadening towards apex, reach-

Female genitalia (Fig. 26). Ductus bursae short and ing vein M1 posteriorly. Forewing: transverse lines out- broad, colliculum strongly sclerotized, but situated in a wardly curved, partly oblique, acutely angled towards transverse position. Corpus bursae large, expanded lat- costa at the border of the yellow costal band; basal line erally, internal surface consisting of two sclerotized more or less evenly curved outwards; antemedial line layers, both almost completely spinulose, just leaving rather straight below yellow costal band, sometimes the area around the signum membranous. Signum a pair forming a small angle on anal fold, broadened near of elongate, symmetrical, spinose bands, connected lon- costa; costal band between basal and antemedial lines gitudinally or separated shortly at anterior end, spines with irregular black suffusion; medial line slightly arranged regularly. Membranous diverticulum rather excurved posteriorly, forming a right angle at the border strong. of the costal band; postmedial area consisting of two Specimens examined. China: 1ǩ (NHM), Holotype, lines, the proximal one almost parallel to medial line, labeled “Wa-Shan, 6000 ft, A.E. Pratt coll. vi.1889, but rather vague or absent near hind margin; the distal

Leech Coll. 1900-64, Figured in Seitz. Vol. IV.Pl.” line approaching the proximal one between CuA1 and

(other labels: Hydrelia straminearia sp. n. type ǩ; anal fold, and between the veins M1 and M3, broadened

Geometridae genitalia slide no. 20083; red type label); and roundly protruding on M3, with a distinct tooth at

Figures 25–32 Female genitalia of Pseudostegania. 25 P. defectata (Christoph), MNHU; 26 P. straminearia (Leech), ZFMK; 27 P. distinctaria (Leech), ZFMK; 28 P. qinlingensis sp. nov. paratype, IZCAS; 29 P. yargongaria (Oberthür), IZCAS; 30 P. lijiangensis sp. nov., paratype, ZFMK; 31 P. zhoui sp. nov., paratype, IZCAS; 32 P. burmaensis sp. nov., paratype, NHM. Scale bars, 1 mm. the border of the yellow band; costal part broad, inter- costal band; terminal line composed of a series of small rupted in the middle. Submarginal line narrow, accom- dots, much larger towards apex; discal spot a small panied by an indistinct diffuse line outside, conﬂuent black brown dot. Hindwing: basal line black brown; with the submarginal line in the middle of the forewing medial line nearly straight from hind margin to discal or not, forming two black patches, a larger one at the spot, rarely more basally; postmedial line double, distal border and a smaller one in the middle of the yellow one more distinct and curved more strongly outwards

at middle than the inner one; submarginal line evenly Labial palpus with most outer surface dark brown, part curved outwards, distinct and continuous, accompany- of ventral surface whitish. Vertex whitish. Patagia dark ing line faint or absent; terminal line rarely composed brown, with a few white scales anteriorly. Tegulae white, of a series of evenly round dots, in most specimens faint with a black-brown base. Thorax white on dorsal and or absent; discal spot a small black brown dot, much ventral sides. Abdomen dorsally and ventrally white, a smaller than that of forewing. Fringe whitish. Under- dark line present on posterior margin of the ﬁrst segment, side: yellow costal band almost invisible; lines similar to small dark dots in the middle of the posterior margin of upperside, but less distinct, except costal part of post- the next two or three tergites. medial and submarginal lines and basal part of discoidal Length of forewing: ǩǨ14–15 mm. Wings white, cell which are broadened and strongly suffused with with a slight creamy tinge, forewing with broad pale dark brown scales. yellow costal band, its posterior border reaching vein

Male genitalia (Figs 11,19): Similar to defectata, M1 and gradually broadening towards apex. Forewing: but tegumen much broader; valva narrower sub-basally, transverse lines except submarginal line outwardly with apex expanded more strongly; basal shaft of sub- oblique and angled inwards at the border of the costal scaphium much stronger, aedeagus without distinct yellow band, interrupted or continuous inside the band; differences compared to defectata. basal line indistinct; antemedial line sharply angled on Female genitalia (Fig. 27): Ductus bursae short and the upper margin of discoidal cell, broadened near costa, broad, with elongate, weakly sclerotized colliculum. slightly dentate on other parts; costal region between Corpus bursae large, two-thirds of internal surface basal and antemedial lines with some additional dark spinulose, remaining part membranous. Spinulose area patches; medial line curved outwards, close to postme- bordered on both sides by sclerotized, longitudinal dial area; postmedial line consisting of two lines, the bands, the surface of which is also, but less, spinulose. proximal line almost parallel to medial line at middle, The concave, sclerotized area near the opening of the but diverging near costa and hind margin; the distal ductus bursae large, the membranous diverticulum line strongly curved outwards, between CuA1 and inner very large, elongate. Signum large, oval, densely spinose, margin, the proximal one there straight, but very faint with a longitudinal ridge. or absent, near the area of M2 both lines conﬂuent. Specimens examined. China: 1ǩ (NHM), Lectotype, Submarginal line a series of small, black brown patches, here designated for reason of nomenclatural stability, tapering towards inner margin; terminal line composed Pu-tsu-Fang, 9820 ft. Native coll. vi. & vii.1890, Leech of a series of small dots between the veins, much larger Coll. 1900-64 (other labels: Hydrelia distinctaria sp. n. towards apex; discal spot a small black brown dot. type ǩ; Geometridae genitalia slide no. 20087; red type Hindwing: basal line a short, black brown streak; label); 1ǩ (NHM), paralectotype, Pu-tsu-Fang, 9820 ft. medial line nearly straight, slightly angled in discal cell; Native coll. vi. & vii.1890, Leech Coll. 1900-64; 1ǩ postmedial line double, the distal one more curved out- (NHM), paralectotype, Wa-shan, 6000 ft, A.E. Prout wards at middle than the proximal one; submarginal coll. vi.1889, Leech Coll. 1900-64; 1ǩ (ZFMK), Chas- line curved outwards, discontinuous; terminal line com- seurs Indigènes des Missionaires de Ta-tsien-Loû, 1906, posed of a series of round dots equal in size; discal spot ex coll. Oberthür, ex coll. Wehrli; 20ǩ6Ǩ (Coll. M. a small black brown dot, smaller than that of forewing. Sommerer, Munich), China, Sichuan, Daxue Shan, Fringe whitish, black brown on veins’ end and apex of Gongga Shan, Moxi, 14–19.vii.1999, 2850 m, 29°41′N, forewing. Underside: yellow band almost invisible; lines 101°51′E, leg. Siniaev & Plutenko. similar to upperside, but less distinct, pale brown suffu- Distribution. China (Sichuan). sion present along postmedial, submarginal and mar- Taxonomic note. Xue and Scoble (2002) excluded ginal area as well as on discoidal cell. distinctaria from Asthena, but did not state the genus to Male genitalia (Figs 12,20). Tegumen large, rounded, which it should belong; here it is newly combined based posterior margin slightly concave at middle, basally with on similar external features and the unique, apomorphic a pair of large, lateral, fused sclerites. Uncus a sclerotized genitalia characters with Pseudostegania. projection. Subscaphium sclerotized, shaft broad, expanded apically and spinose. Valva narrow, moder- Pseudostegania qinlingensis Xue & ately expanded and setose apically; costa sclerotized Han sp. nov. (Fig. 4) at base, oval terminal broadening present; sacculus with Description. Antennae with lower surface yellow- a long and pointed process, sclerotized. Saccus well- brown, upper surface pale brown at base, then chequered developed and broad. Labides with some broad scale-like dark and pale brown, terminal two-thirds brown. Frons setae at posterior margin of opening. Juxta blunt. Anellus whitish, with a narrow black band at upper margin. undeveloped. Aedeagus as in the generic description.

Female genitalia (Fig. 28). Antevaginal, transverse Redescription. Antennae with lower surface brown, band indistinct. Antrum enlarged, strongly sclerotized. upper surface white at base, then chequered white and Ductus bursae short, with a broad colliculum. Corpus pale grayish-brown till one-third, terminal two-thirds bursae rather small, irregularly folded and wrinkled, grayish-brown. Frons slightly protruding, smooth- with a squarish, protruding part posteriorly on left side. scaled, pale yellow, with a dark brown round patch About half of its internal surface densely spinulose; between antennae. Labial palpus very short, rough- membranous diverticulum very large; signum a small scaled, white; vertex white. Patagia dark brown; tegulae spinose patch, incurved anteriorly. white, with a black brown base. Thorax white dorsally Diagnosis. On wing pattern, qinlingensis is closest to and ventrally, abdomen dorsally white, with contrasting distinctaria, the difference mainly lie in the submar- gray-brown bands along posterior margins of tergites, ginal lines of both, forewing and hindwing: submar- underside dull white with transverse bands less distinct. ginal line of qinlingensis is composed of a series of Length of forewing: Ǩǩ15–16 mm. Wings chalk- small patches, while that of distinctaria is completely white, forewing with broad yellow costal and terminal linear. In the male genitalia, qinlingensis is close to bands, both getting gradually broader towards apex, yargongaria, both bearing a pointed saccular process. the costal band reaching vein M1 posteriorly. Forewing: The process of qinlingensis, however, is much larger transverse lines except those of submarginal and termi- and located more posteriorly. The shaft of sub- nal region straight and outwardly oblique, sharply scaphium is broader in qinlingensis, and the juxta is angled inwards at the border of the costal yellow band, less acute. Female genitalia of qinlingensis have the these costal parts may be continuous or interrupted in signum rather similar to that of yargongaria and lijian- the middle; medial line together with two lines of post- gensis, but they are unmistakable because of the pecu- medial area similarly strong, straight lines, close to each liar, enlarged and sclerotized antrum (see Fig. 28). The other near costal band and distinctly diverging near hind colliculum is rather short and broad. margin; two submarginal lines gently curved outwards, Holotype: ǩ (IZCAS), labeled: “Gansu: Wenxian almost parallel to margin, but abruptly bent to the Qiujiaba, 2350 m, 22.vii.1999, Coll. Yao Jian”. opposite direction near costa; proximal line smooth and Paratypes. China: 1ǩ (IZCAS), Gansu: Wenxian continuous, distal line close to the first, but somewhat Fanba, 800 m, 26.vi.1998, Coll. Zhang Xuezhong; 1Ǩ more diffuse. Subterminal line composed of a series of (IZCAS), ibidem, Coll. Yao Jian; 2Ǩ (IZCAS), Gansu: round dots between the veins, gradually increasing in Zhouqu Shatanlinchang, 2350 m, 5.vii.1998, Coll. size towards the much larger, oval, subapical patch; Wang Hongjian; 3Ǩ (IZCAS), same locality, 4.vii.1998, discal spot a small black brown dot. Hindwing: basal Coll. Yao Jian; 1Ǩ (IZCAS), same locality and date, line black brown; medial line nearly straight, connected Coll. Zhang Xuezhong; 3Ǩ (IZCAS), Shaanxi: Liuba with the discal spot or running just proximal or distal to Miaotaizi, 1350 m, 21.vii.1998, Coll. Yao Jian. 1ǩ it; two parallel, postmedial lines, the outer one bent (IZCAS, ex. ZFMK), Tapaishan im Tsinling, Sued- inwards near costa; submarginal line double, curved Shensi (China), 26.vi.1935, E (= 3000 m), H. Höne; outwards almost parallel to termen, proximal one 1ǩ1Ǩ (NHM, ex. IZCAS), Gansu: Zhouqu Shatanlin- continuous, accompanying distal one faint and diffuse, chang, 2400 m, 14.vii.1999, Coll. Yao Jian; 7ǩ3Ǩ sometimes incomplete; subterminal line composed of a (ZFMK), Tapaishan im Tsinling, Sued-Shensi (China), series of evenly round dots; discal spot a very small H. Höne, 1ǩ, 19.vi.1935, L; 1ǩ, 20.vi.1935, L; black brown dot, smaller than that of forewing. Fringe 1ǩ1Ǩ, 22.vi.1935, L; 2ǩ, 23.vi.1935, F; 1ǩ1Ǩ, whitish, with black brown scales on veins’ end of both 24.vi.1935, F; 1Ǩ, 25.vi.1935, F; 1ǩ, 2.vii.1935, F. L (= wings and at apex of forewing. Underside: upperside 1700 m) and F (= 3000 m) have been used by H. Höne yellow bands represented by dark brown bands on in the years 1934 and 1935 to indicate the elevation of underside, a yellow patch present in costal band post- the collecting site on the labels. medially. Lines similar to those of upperside, but more Distribution. China (Shaanxi, Gansu). diffuse and partly confluent on forewing, discoidal cell Etymology. The species is named after the type locality. with dark suffusion; on hindwing, medial and first postmedial lines visible but indistinct, the second postmedial line distinct, bent outwards at middle, submarginal line Pseudostegania yargongaria (Oberthür, 1916) indistinct and sometimes discontinuous, terminal line as comb. nov. (Fig. 5) on upperside, but dots larger. Asthena yargongaria Oberthür, 1916, Études Lépid. Male genitalia (Figs 13,21). Tegumen very large, Comp. 12: 167, pl. 399. Lectotype ǩ, [China]: Yargong rounded, posterior margin slightly concave at middle, Zambala, P. Soulié. 1903. (ZFMK) Here designated. basally with a pair of large, lateral, fused sclerites. Uncus

a small sclerotized projection. Subscaphium sclerotized, quered dark and pale brown till one-third, terminal with slender base, expanded apically and spinose. Valva two-thirds brown. Frons slightly protruding, smooth- narrow sub-basally, expanded and setose apically; scaled, pale yellow; with a dark brown round spot costa sclerotized at base, the terminal broadening rather between antennae. Labial palpus very short, rough- weakly so; sacculus with a short and pointed process, scaled, terminal segment hardly visible, with upper sclerotized. Saccus well developed and broad. Labides surface and lateral-upper half black brown; tip, lateral- with setae on posterior margin of opening much broader lower half and ventral surface pale brown (almost than other setae. Juxta tapering, nearly triangular. uniform pale brown in female). Vertex whitish. Patagia Anellus densely spinulose. Aedeagus as in the generic dark brown, with a few white scales anteriorly. Tegulae description, diverticulum quite shorter than in other white, with a black brown base. Thorax white on dorsal species. and ventral sides. Abdomen white, with contrasting Female genitalia (Fig. 29). Ductus bursae short and dark gray bands along posterior margins of tergites, broad, with distinctly sclerotized colliculum. Corpus underside dull white or pale brown, without transverse bursae large, expanded and irregularly folded and bands. wrinkled laterally; two sclerotized bands extending Length of forewing: ǩǨ15 mm. Wings cream-white, anteriorly (not visible in Fig. 29); internal surface almost forewing with broad yellow costal and terminal bands, completely spinulose. Signum a small rounded spinulose costal band gradually broadening towards apex, its pos- patch, deeply incurved anteriorly. Membranous diver- terior border reaching vein M1; terminal band tapering ticulum large. towards anal angle, often not reaching it. Forewing: Specimens examined. China: 1ǩ (ZFMK), Lectotype, transverse lines black brown, all except basal and sub- here designated for reason of nomenclatural stability, marginal lines outwardly oblique and angled inwards at Yargong Zambala, P. Soulié, 1903, ex coll. Ch. Ober- the border of the costal yellow band, but interrupted thür, ex coll. Wehrli; 1ǩ (ZFMK), paralectotype, inside the band; basal line ovally curved; antemedial Yarégong, P. Soulié, 1902, ex coll. Ch. Oberthür, ex coll. line straight, broadened near the angle and near costa; Wehrli; 1Ǩ (NHM), paralectotype, Yarégong, P. Soulié, the angle often touching the discal spot; costal region 1902, Ex Oberthür Coll. Brit. Mus. 1927-3 (other between basal and antemedial lines with some addi- labels: Asthena yargongaria Obthr.; Geometridae geni- tional dark patches; medial line narrow, parallel to the talia slide no. 20088; red type label); 1Ǩ (NHM), para- proximal line of postmedial area at middle, but dis- lectotype, Thibet Yargong-Zambala, P.Soulié 1903, tinctly diverging near costa and hind margin; postmedial Ex Oberthür coll. Brit. Mus. 1927-3; 4ǩ4Ǩ (ZFMK), area consisting of two narrow, similar lines, the distal

A-tun-tse, Nord-Yünnan, H. Höne, 1Ǩ, mittlere Höhe, line with a strong, tooth-like protuberance at vein M1, about 4000 m, 5.vii.1936; 4ǩ3Ǩ, obere Höhe, about but only slightly excurved near M3 and slightly or not

4500 m, 29.vi., 8-12.vii.1936; 2ǩ1Ǩ (ZFMK), bent inwards between vein CuA2 and vein A; distinctly Li-kiang, Provinz Nord-Yuennan, H. Höne, 1ǩ, broadening near costa, often conﬂuent with the costal 2800 m, 2.iv.1935; 1ǩ, 30.vi.1935; 1Ǩ, about 4000 m, part of the proximal line; submarginal two lines gently 25.vii.1935; 1ǩ1Ǩ (IZCAS, ex. ZFMK), A tun tse curved almost parallel to margin, but abruptly bent to (Nord Yünnan), Obere Höhe (about 4500 m), the opposite direction near costa; proximal line narrow, 10.vii.1936, 17.vii.1936, H. Höne; 1ǩ (IZCAS), continuous, distal line consisting of a series of small Yunnan: Lijiang Yulongshan, 10.vii.1982, coll. Song black brown dots, more or less continuous from costa Shimei; 13ǩ2Ǩ (IZCAS), Yunnan: Lijiang Ganheba, towards inner margin, decreasing in size and often not 3296 m, 23.vi.2009, coll. Han Hongxiang, Yangchao & reaching it; subterminal line composed of a series of Qifeng. round dots, getting much larger towards the oval apical Distribution. China (Yunnan, Tibet). patch; discal spot a small black brown patch, sometimes Taxonomic note. Xue and Scoble (2002) excluded connected with medial line. Hindwing: basal line a small yargongaria from Asthena, but did not state the genus to black brown patch; medial line nearly straight, broader which it should belong, here it is newly combined with than that of forewing, often connected with the discal Pseudostegania based on similar external features and spot, but also often running just anterior or posterior to genitalia characters with Pseudostegania. it; postmedial line double, parallel, the interior one slightly weaker; two submarginal lines curved outwards, Pseudostegania lijiangensis Xue & Stüning the distal one often indistinct or diffuse; subterminal sp. nov. (Fig. 6) line composed of a series of evenly round dots which Description. Antennae with lower surface yellow- are variable in size and may also be completely absent; brown, upper surface with base pale brown, then che- discal spot a black brown dot. Fringe whitish, with

black brown scales on veins’ end and apex, the latter 29.vi.1934, 3200–3500 m; 1ǩ, 14.vi., 1ǩ, 18.vi., often weaker on hindwing and absent when the subter- 1ǩ, 20.vi.1935, 2400–2800 m; 1Ǩ, 18.vi., 1Ǩ, minal dots are missing. Underside: Lines similar to 22.vi.1935, 4000 m; 1ǩ, 3.vii., 2ǩ, 5.vii.1935, upperside, less distinct, with strong brown suffusion 4000 m, 1ǩ, 8.vii.1934, 2400–2800 m, 1ǩ, 9.vii.1935, especially along postmedial, submarginal and marginal 2400–2800 m, 1ǩ, 10.vii.1934, 2400–2800 m; 1ǩ, area as well as on discal cell. 10.vii.1934, 3200–3500 m; 1ǩ, 11.vii.1934, 2400– Male genitalia (Figs 14,14a,22). Tegumen as broad as 2800 m; 2ǩ, 11.vii.1934, 3200–3500 m; 3ǩ, in distinctaria, with posterior margin rounded, uncus 12.vii.1934, 3200–3500 m; 2ǩ, 14.vii.1934, 2400– larger, sclerites of reduced gnathos not fused in the 2800 m; 1ǩ, 14.vii.1934, 3200–3500 m; 3ǩ, middle. Shaft of subscaphium rather broad, slightly 14.vii.1935, 3200–3500 m; 1ǩ, 15.vii.1934, 2400– spinulose distally, expanded apical part densely spinose. 2800 m; 3ǩ, 15.vii.1935, 3200–3500 m; 1ǩ, Valva sub-basally not distinctly narrow, apex only mod- 16.vii.1934, 3200–3500 m; 1ǩ1Ǩ, 16.vii.1934, erately expanded, subapical costal broadening distinct. 2400–2800 m; 6ǩ, 16.vii.1935, 3200–3500 m; 1Ǩ, Sacculus without process; saccus moderately protrud- 17.vii.1934, 4000 m; 1ǩ, 18.vii.1934, 2400–2800 m; ing, angled near base. Labides with shaft rather short, 1ǩ, 19.vii.1934, 3200–3500 m; 1ǩ,1Ǩ, 19.vii.1934, margin of opening bearing a series of broad, scale-like 3200–3500 m; 1ǩ, 20.vii.1934, 4000 m; 1ǩ, setae, but the latter may be lost easily during prepara- 20.vii.1935, 3200–3500 m; 4ǩ5Ǩ, 21.vii.1934, tion. Juxta blunt. Aedeagus as in the generic description 4000 m; 1ǩ, 21.vii.1935, 3200–3500 m; 1ǩ, Female genitalia (Fig. 30). Ductus bursae short and 21.vii.1935, 4000 m; 1Ǩ, 22.vii.1934, 2400–2800 m; broad, anterior half with distinctly sclerotized collicu- 2ǩ, 22.vii.1935, 4000 m; 1ǩ, 23.vii.1934, 2400– lum. Corpus bursae large, folded and wrinkled; two 2800 m; 1ǩ, 23.vii.1934, 3200–3500 m; 2ǩ, elongate sclerotized bands present, bordering the spinu- 24.vii.1934, 3200–3500 m; 2ǩ, 24.vii.1935, 3200– lose internal surface of the bursa. Signum small, a 3500 m; 1ǩ, 24.vii.1935, 2400–2800 m; 1ǩ1Ǩ, 25. spinose, rounded, concave patch, shortly incurved ante- vii.1934, 3200–3500 m; 1Ǩ, 25.vii.1935, 4000 m; riorly (not visible in lateral view). The concave, sclero- 1ǩ, 26.vii.1935, 2400–2800 m; 1ǩ, 26.vii.1936, tized area near the opening of the ductus bursae and the 3200–3500 m; 1ǩ, 29.vii.1935, 3200–3500 m; 1ǩ, membranous diverticulum large. 2.viii.1935, 2400–2800 m; 1ǩ, 4.viii.1934, 2400– Diagnosis. Most similar to yargongaria, but with the 2800 m; 1Ǩ, 7.viii.1935, 4000 m; 1ǩ, 9.viii.1934, ground color yellowish-white (in yargongaria chalk- 2400–2800 m; 1ǩ, 13.viii.1935, 2400–2800 m; 1ǩ, white). Wing pattern is similar in both species but lines 14.viii.1934, 2400–2800 m; 1ǩ, 19.viii.1934, 2400– are narrower in lijiangensis, the medial line is slightly 2800 m; 1ǩ, 21.viii.1935, 2400–2800 m; 3ǩ2Ǩ curved and the distal of the two postmedial lines are (IZCAS, ex. ZFMK), 1ǩ, 30.vi.1934, 3200–3500 m; curved outwards at or just posterior to vein M3 (in 1Ǩ, 12.vii.1934; 1ǩ, 16.vii.1935, 3200–3500 m; yargongaria all three lines are almost straight in their 1ǩ1Ǩ, 21.vii.1934, 4000 m. central parts). In the male genitalia the two species can be Distribution. China (Yunnan). distinguished by the saccular process, which is lacking in Etymology. The species is named after the type locality. lijiangensis. Male genitalia otherwise are similar to those of distinctaria, straminearia and defectata, but Pseudostegania zhoui Xue & Han sp. nov. the labides bear a series of very broad scale-like setae at (Fig. 7) the distal opening in lijiangensis, smaller or lacking in the Description. Antennae with lower surface yellow- three other species. For comparison of other slight differ- brown, upper surface pale brown. Frons slightly pro- ences see Figures 9–11 and 14. Female genitalia are truding, smooth-scaled, upper half pale brown, lower similar in yargongaria and lijiangensis, but lijiangensis half black brown, with a small dark brown dot between has a shorter, more strongly sclerotized colliculum. antennae. Labial palpus with upper and lateral surfaces Holotype ǩ (ZFMK), labeled: “Li-kiang. (China), X. black brown, ventral surface whitish. Vertex whitish, Provinz Nord-Yuennan, 1.7.1934, H. Höne.” (yellow, sparsely mixed with dark brown scales. Tegulae pale printed label, with collection no. 413034 on under- yellow, with base black brown. Thorax and abdomen side; X = elevation-symbol used by H. Höne for with dorsal sides pale yellow, and ventral sides whitish. 2400–2800 m). Length of forewing: ǩ12.5–14.5 mm; Ǩ14–15 mm. Paratypes. China: 76ǩ18Ǩ (ZFMK), Li-kiang Wings yellow. Forewing: yellow costal and terminal (China), Provinz Nord-Yuennan, H. Höne, 1ǩ, bands not present; wing base diffused with black brown 27.v.1934; 5ǩ, 13.v. (2), 15.v., 18.v., 19.v.1935, 3200– scales; basal and antemedial lines irregularly sinuous 3500 m; 1ǩ2Ǩ, 2.vi., 1ǩ, 6.vi., 1ǩ, 9.vi.1935, 1Ǩ, or curved outwards; medial line sinuous, incurved at

middle; postmedial line distinctly excurved, forming two Sichuan: Tianquan Labahe, 1300 m, 29.vii.2004, coll. protrusions on vein M1 and below M3; a large black Ren Yingdang; 1ǩ1Ǩ, Sichuan: Pingwu Laohegou brown patch present between medial and postmedial Caiyuchang, 1630 m, 14.viii.1999, coll. Zhou Xin. lines, partly extended to costa and terminating near Distribution. China (Sichuan).

CuA1 posteriorly, medial and postmedial lines approach- Etymology. The species is named after the collector. ing each other near CuA2, then diverging again towards inner margin; submarginal line curved outwards; termi- Pseudostegania burmaensis Xue & Han nal line composed of tiny dots between veins, indistinct sp. nov. (Fig. 8) for bad condition of specimens; discal spot a round Description. Antennae pale yellow below, with black dot, close to proximal margin of the postmedial basal half of segments of basal one third silvery on patch. Hindwing: basal line a small black brown streak; the upper side; frons yellow, with basal half dark medial line nearly straight, broader than that of forew- brown; labial palpus yellow, rough-scaled on both, ing; postmedial line double, sinuous, area between the inner and outer surfaces; vertex and tegulae yellow, the lines slightly broadened at middle; submarginal line latter in some specimens with a brown spot at centre parallel to outer margin; terminal line and discal spot of base. Thorax whitish on dorsal side. Abdomen similar to those of forewing. Fringe pale yellow. Under- dorsally pale yellow brown, mixed with a few brown side: lines similar to upperside, but more indistinct; the scales at middle. large central black brown patch on forewing almost Length of forewing: ǩǨ15 mm. Wings pale yellow. identical to that of upperside. Lines diffuse. Forewing with a large black brown patch Male genitalia (Figs 15,23). Tegumen bluntly taper- present at base; basal and antemedial lines indistinct; ing, uncus quite small, subscaphium densely spinulose, medial and postmedial lines forming a wide black becoming gradually broader towards apex. Valva not brown band, protruding outwards between costa and narrowed sub-basally and not extended apically, costa M2, and between M3 and CuA2; submarginal line a sclerotized, forming a small pouch-like structure sub- narrow black brown band, with inner margin irregularly apically. Saccular process absent. Saccus strongly wavy; terminal line absent; discal spot an oval black dot. protruding. Labides with scale-like setae at posterior Hindwing: basal line a small black brown streak; medial margin of distal opening. Juxta almost triangular, line absent, postmedial line forming a wide band, curved anellus appearing as a spinulose patch. Aedeagus as in outwards; submarginal band similar to that of forewing; the generic description, obviously without differential discal spot much smaller than that of forewing. Fringe characters. pale yellow. Underside: lines similar to upperside, but Female genitalia (Fig. 31). Ductus bursae short and more diffuse. broad, colliculum developed, sclerotized. Corpus bursae Male genitalia (Figs 16,24). Tegumen rounded, large, posteriorly bearing a sclerotized, pouch-like pro- posterior margin not concave. Uncus a small projection. trusion; three-fourths of internal surface densely spinu- Subscaphium densely spinulose. Valva simple, narrow lose, no sclerotized bands present. Signum a pair of sub-basally and slightly expanded apically, costa with a elongate, symmetrical, spinose patches, connected pos- very vague oval subapical broadening. Labides with teriorly, spines arranged regularly. apex rounded. Saccus semicircular. Juxta blunt. Anellus Diagnosis. Externally, the much larger patch on forew- appearing as densely spinose patch. Aedeagus with ing of zhoui distinguishes it from the rather similar cornutus on vesica very long, narrow and pointed, diver- straminearia and defectata. In male genitalia, the valva ticulum also quite long and narrow. of zhoui bears, besides other differences, a bag-like Female genitalia (Fig. 32). Antevaginal, transverse structure subapically, which is distinctive in Pseudoste- band indistinct. Ductus bursae not separated from gania. Female genitalia show a rather close relation to corpus bursae, colliculum absent. Corpus bursae large, straminearia and defectata by the shape of the signum, elongate, concave on right side, with a band-like, scle- but in zhoui it is shorter and broader and not so deeply rotized area in the middle; the largest part of the internal divided anteriorly. Moreover, the small round, pouch- surface densely, but ﬁnely spinulose. Signum absent. like protrusion near the ductus bursae is distinctive. Diagnosis. Pseudostegania burmaensis is distinctive in Holotype ǩ (IZCAS, ex. Nankai University), labeled: wing pattern: the transverse lines are represented by “Sichuan: Mabian Yonghong, 1200 m, 22.vii.2004, diffuse bands. In the male genitalia, burmaensis is dis- coll. Ren Yingdang”. tinctive in the almost absent subapical modiﬁed struc- Paratype. China: 1Ǩ (IZCAS, ex. Nankai University), ture on costa and the cornutus being quite long and thin. same locality and collector as holotype, 1500 m, In the female genitalia, burmaensis is the only species 23.vii.2004; 1ǩ (IZCAS, ex. Nankai University), without a signum in Pseudostegania.

Holotype ǩ (NHM), labeled: “BURMA: Mishmi resolved. Species of Laciniodes have, besides the labides, Hills. Dingliang, 2450 ft, 11.iii.1935, M. Steele, valves, subscaphium and juxta also very similar, but Brit. Mus. 1935-312, Geometridae genitalia slide no. they have, on the other hand, a well-developed, distally 19709”. pointed uncus, the tegumen is not expanded posteriorly Paratypes: 1Ǩ (NHM), BURMA: Mishmi Hills. and the aedeagus has a bundle of spines instead of a Dingliang, 2450 ft, 14.iii.1935, M. Steele, Geometridae large, single cornutus. A further remarkable feature is genitalia slide no. 19710); 1ǩ1Ǩ (NHM), BURMA: that both genera have a gnathus: in Laciniodes well Mishmi Hills. Dingliang, 2450 ft, 11.iii.1935, M. Steele. developed and functional, in Pseudostegania reduced to Distribution. Myanmar. sclerites which are attached to the ventral side of the Etymology. The species is named after the earlier name tegumen and which are not functional. In Asthenini and of the type locality. other related tribes the gnathus is completely absent. In the female genitalia, the signum in Laciniodes is simple, Diagnosis and tribal placement of patch- or band-like, not a compact, often divided plate Pseudostegania with internal spines as in Pseudostegania. In both Externally, species of Pseudostegania are reminiscent genera, but also in the Eupitheciini, the shafts of the of several species presently included within genera of labides are arising from the base of the valve and are the tribe Asthenini. As a consequence, earlier authors connected dorsally with the transtilla by short, rod- described some of them as members of Asthena (yargon- like branches, ventrally longer branches are extending garia Oberthür) or Hydrelia (distinctaria Leech, stra- downwards to the juxta. In Eupitheciini, however, the minearia Leech). The latter two have been transferred to ventral branches of the labides are connected with a Asthena subsequently. central constriction of the juxta, the latter thus having Xue and Zhu (1999) combined straminearia an hourglass-like shape. In Pseudostegania (and Lacin- with Pseudostegania. Xue and Scoble (2002) excluded iodes) the juxta is more or less triangular. yargongaria, distinctaria and straminearia from Asthena Considering all these facts, neither Laciniodes nor (and also from the tribe Asthenini), because of the large Pseudostegania really fit into Asthenini, Eupitheciini or differences in male and female genitalia, and they did the other tribes mentioned above; they probably deserve not decide where to place them correctly. a tribe of their own. But further, more comprehensive Differences excluding species of Pseudostegania from studies are necessary to decide this, especially since even Asthenini have been the distinctive unusual labides in the distinction between the Asthenini and Eupitheciini male genitalia, which in all the members of Asthenini are is considered to be only weakly supported (Holloway not expanded distally and are not extended basally into 1997, p. 120). prominent ventral arms. In the female genitalia, the corpus bursae bears a signum (or signa) composed of ACKNOWLEDGMENTS radiating denticles or spines in Asthenini, in Pseudoste- gania the signum is a sclerotized plate of different shape, We express sincere thanks to the staff of the Natural densely spined internally. History Museum, London, UK, for giving us the oppor- Asthenini are rather close to Eupitheciini. Xue and tunity to study the collections under their care. We espe- Scoble (2002) transferred three genera from the latter cially thank Sir Anthony Galsworthy, London, for his tribe to the Asthenini. However, species of the tribe kind help on checking head characters of straminearia, Eupitheciini, and also those of the Perizomini and the distinctaria and burmaensis and label data in the collec- Operophterini, have distally expanded labides which are tion of NHM; he also gave useful comments on an united distally; the Perizomini, Operophterini and a part earlier draft of the manuscript. Dr Wolfram Mey, of Eupitheciini show this unusual character (e.g. Xue & Museum für Naturkunde der Humboldt-Universität Zhu 1999, p. 886, fig. 1117). Compared to Pseudoste- zu Berlin, Germany, kindly gave us the opportunity to gania, the labides are at first glance similar, but in detail study the collections and loaned two syntypes of defec- they are different, often elongate (Holloway 1997, p. tata. Dr V. G. Mironov, Systematics of Insects Labora- 119). Labides of Pseudostegania are unique in having a tory, Zoological Institute of the Russian Academy of deep groove across the distal, expanded part, its margins Sciences, provided us with digital images of another being densely covered with specialized, apically scale- syntype of defectata and its labels and gave us useful like spines. The shape of this distal part is similar, but information about its historical background. Thanks without the groove, in the genus Laciniodes. Also of also to Katsumi Yazaki, Tokyo, for his generous gift of the latter genus, which is similar to Pseudostegania specimens of Pseudostegania defectata chrysidia and Dr in a number of characters, the tribal placement is not Li Houhun, for giving specimens of Pseudostegania

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