Holotype male, INDIA: Karnataka: Shimoga district, Someshwari Wildlife Sanctuary, 10 km W Agumbe, 13°28′24″N; 75°00′40″E, alt. 372 ft, early successional, wet evergreen forest, 22.ix.2005, C. R. Bartlett leg. (UDCC).

Orthopagus bartletti sp. n. can be separated from all other Orthopagus species by the very short head, hardly produced in front of eyes; the forewings with a brownish streak along nodal line connecting the pterostigmal area with the distal sublunate streak; the dorso-posterior margin of the male pygofer with a large and broad process forming two distinct angles; the robust male segment X; and the lobes of the phallobase muricate apically.

Measurements (1 male). Body length (from apex of head to tip of forewings): 11.5 mm; head length (from apex of cephalic process to base of eyes): 1.2 mm; head width (including eyes): 1.5 mm; forewing length: 9.6 mm.

Coloration (Fig. 2A–B). General colour brownish ochraceous marked with dark brown on dorsum. Head greenish ochraceous, vertex with basal corners, a pair of round patches in basal third, and apical diamond-shaped spot dark brown (Fig. 4A); frons with median area between intermediate carinae extensively dark brown anteriorly and with series of small pale fuscous spots along intermediate and lateral carinae and narrow ivory white band basally (Fig. 4C). Clypeus ivory white, with two small spots at base and apex dark brown. Compound eyes fuscous with posterior margin ochraceous; ocelli purplish-red. Pronotum brownish ochraceous, median carina, apical marginal areas of ventral lobes, and posterior lateral angles ivory white. Mesonotum brownish ochraceous (Fig. 4A). Forewings hyaline, veins ochraceous, pterostigmal area, a streak along nodal line, and a wide sublunate streak on distal third dull ochraceous; posterior (claval) margin broadly faintly brown (Fig. 3A). Hind wings hyaline, veins and an apical spot dull ochraceous. Legs pale brown; fore femora subapically and hind tibiae at base and apex (including lateral and apical spines) blackish. Abdomen dorsally and ventrally brownish ochraceous.

Structure. Head (Fig. 4A–C) very short, cephalic process practically absent. Vertex (Fig. 4A) with ratio of length at midline to width between eyes 1.4. Frons with base slightly inflated anteriad in dorsal view (Fig. 4A), with transition to vertex broadly rounded in lateral view (Fig. 4B); in ventral view, frons with ratio of length at midline to maximum width 2.2; median carina more or less obscure at base (Fig. 4C).

Male genitalia. Pygofer, in lateral view, with dorso-posterior margin forming a large, broad, biangular lobe (Fig. 4E); in ventral view (Fig. 4F) much longer than in dorsal view (Fig. 4D) with ratio of ventral to dorsal width about 3.4. Gonostyles (Fig. 4E, F) elongate, relatively narrow in basal half, with strongly sinuate dorsal margin medially. Aedeagus (Fig. 4G–I) with endosomal processes relatively short and robust, without distinct minute superficial spines, and directed laterad; phallobase with a pair of large, elongate, thumb-like ventral lobes, curved dorso-posteriad and muricate apically (Fig. 4G–H); and a pair of shorter lateral lobes, directed posteriad (Fig. 4H). Segment X, in lateral view, relatively short and robust, with ventral margin gradually widening from base to broadly truncate apex (Fig. 4E); in dorsal view broad, broadest medially, with ratio of length to maximum width 1.1 (Fig. 4D).

Female genitalia unknown.

The new species is named after Dr. Charles R. Bartlett, collector of the type specimen and curator of the insect collection at the Department of Entomology and Wildlife Ecology, University of Delaware, USA, in recognition of his kindest help and support to the first author when he visited UDCC in 2017. The species name is to be treated as a noun in genitive case.

So far only known from southwestern India (Fig. 11).

INDIA: 2 females, [no state indicated], 1934-394, T. R. Bell leg. (BMNH); West Bengal: 1 female, Calcutta [= Kolkata], 3.x.1907 (coll. Distant, BMNH); Maharashtra: 1 male, Sindhudurg district, roadside on ridge 2 km W Amboli, 15°58'04"N, 73°59'23"E, alt. 2394 ft, pasture and successional, moist deciduous forest, 28.ix.2005, C. R. Bartlett leg. (UDCC); 1 male, Pune district, 5 km E Mulshi Lake near Tamini village, 18°26'37"N, 73°25'46"E, alt. 2047 ft, deciduous forest and open areas, 2.x.2005, C. R. Bartlett (UDCC); Goa: 2 females, Sanguem district, near Bhagwan Mahaveer Sanctuary 100 m E Molem, 15°22'43"N, 74°13'52"E, alt. 342 ft, moist deciduous forest, 24–25.ix.2005, C. R. Bartlett leg. (UDCC); Kerala: 2 males, Malabar, Nadungayam [forest near Nilambur], 200 ft, 16.–22.ix.1938 (BMNH); 1 female, Parambikulam, alt. 1700–3200 ft, 16.–24.ix.1914, F. H. Gravely leg. (coll. Distant, BMNH); 2 females, Tenmalai [= Thenmala], 12.–15.v.1937 (BMNH); Tamil Nadu: 1 female, Nilgiri Hills, 11 km SE Kotagiri, Kunchappanai, 11°24ʹN, 76°56ʹE, alt. 1100 ± 100 m, 3.–15.v.2002, L. Dembický leg. (MMBC). SRI LANKA: North Western Province: 1 female, Puttalam district, Puttalam, “12.” [= ?1912] (coll. Melichar, MMBC); North Central Province: 1 male, Anuradnapura district, Wilpattu National Wildlife Park, Hunuwilagama, Wildlife Soc. Bungalow, 200 ft, 10–19.iii.1970, D. Davis & B. Rowe leg. (USNM); Central Province: 1 female, Kandy district, Talwatte, 29.xi.1995, M. Schaffer leg. (BMNH); Samaragamuwa: 1 male, Ratnapura district, Uggalkaltota, Irrigation Bungalow, alt. 350 ft, 31.i–8.ii.1970, D. Davis & B. Rowe leg. (USNM).

Measurements (2 males, 8 females). Body length (from apex of head to tip of forewings): male 11.1 mm, female 12.3–14.7 mm; head length (from apex of cephalic process to base of eyes): male 1.28–1.30 mm, female 1.28–1.40 mm; head width (including eyes): male 1.40–1.45 mm, female 1.48–1.60 mm; forewing length: male 9.0–9.4 mm, female 10.2–12.2 mm.

Coloration. General coloration as in generic description (Fig. 2A). Vertex predominantly light ochraceous, with 3–4 pairs of small dark brown markings: an elongate patch on each side of midline apically, a small spot at each lateral keel medially, a roundish spot on each side of midline at basal third, and a small spot in each postero-lateral corner (Fig. 5A). Frons light ochraceous with small dark brown spots along intermediate and lateral carinae, frons base slightly infuscated (Fig. 5C). Forewing membrane pattern as in Fig. 3B. Hind wing membrane with a relatively narrow dark brown streak along the apical portion of CuA₁ vein, extending along hind wing apical margin in some specimens.

Structure. Head with cephalic process very short, not inflated (Figs 5A–C). Vertex (Fig. 5A) with ratio of length at midline to width between eyes 1.2–1.5. Transition of vertex to frons narrowly rounded in lateral view (Fig. 5B). Frons relatively broad, with ratio of length at midline to maximum width 2.3–2.7.

Male genitalia. Pygofer, in lateral view, with dorso-posterior margin produced into a relatively large but narrow, tooth-like, apically obtuse process (Fig. 5E); in ventral view (Fig. 5F) much longer than in dorsal view (Fig. 5D) with ratio of ventral to dorsal width about 2.6. Gonostyles (Fig. 5E) large and broad medially, with dorsal margin weakly sinuate medially. Aedeagus (Fig. 5G–I) relatively small and slender, endosomal processes curved laterad and slightly ventro- or dorsoanteriad, membranous, without distinct minute superficial spines; phallobase with three pairs of relatively small (not conspicuously inflated) membranous lobes: a pair of elongate lateral lobes with their apices gradually convergent and tapering posteriad (Fig. 5H–I), a pair of elongate thumb-like ventral lobes, directed ventroposteriad (Fig. 5H), and a pair of small thumb-like dorsal lobes, directed dorsolaterad (Fig. 5G). Segment X, in lateral view, elongate, basal half narrow, gradually widening to apex beyond middle (Fig. 5E); in dorsal view, widest medially, with ratio of length to maximum width 1.1 (Fig. 5D).

Female genitalia as in generic description.

India (southwestern part and West Bengal), Sri Lanka (Fig. 11).

Udugama exoleta was described from Sri Lanka as the type species of Udugama (Melichar 1903). Distant (1906) synonymized this species name with Udugama splendens described from Java, Indonesia. However, Kirkaldy (1908), probably based on comparisons of figures in Melichar (1903) and Distant (1906), commented that U. exoleta was “very different” from U. splendens in having a much longer face. Nevertheless, the synonymy of U. exoleta with U. splendens was accepted by Melichar (1912), and later included in Metcalf’s (1946) catalogue of world Dictyopharidae.

Based on our critical review of the published information and examination of Orthopagus material from Sri Lanka which agrees with the original description of U. exoleta, we propose here to resurrect Orthopagus exoletus comb. n. from the synonymy with O. splendens and to restore it as a valid species. Orthopagus exoletus can be distinguished from O. splendens by the coloration of the vertex and the structure of the male genitalia, particularly the structure of the endosomal processes of aedeagus, lobes of the phallobase and the shape of the segment X. The relative length of frons mentioned by Kirkaldy (1908) is probably not a relevant diagnostic character because the length of the head in Orthopagus species varies within a certain range.

According to Melichar (1903), U. exoleta was described based on five female specimens from “Moruwale”, deposited in the collection of the museum in Colombo, Sri Lanka. This material was not available to our study. Nevertheless, we have studied one female from Sri Lanka, “Puttalam” preserved in Melichar’s personal collection in MMBC. Even though this specimen bears original identification labels handwritten by Melichar as “Udugama” and “exoleta det. Melichar” and a dark red label “Typus” originally also attached to the specimen by Melichar, it probably cannot be considered as a syntype because it differs in its locality and deposition from the information published in the original description and probably it was also collected later than the original species description had been published. Melichar did not use type labels in the modern sense. He had rather adopted the practice of placing a ʻtypeʼ label on one or more specimens of the most taxa (even on species previously described by other authors and identified by Melichar), specimens presumably which he himself used for comparison (Young and Soós 1964, Wilson and Malenovský 2007).

Holotype male, CHINA: Hainan: Baoting, 80 m, 21.vii.1960, S. F. Li leg. (IZCAS).

Paratypes. CHINA: Hainan: 6 males, 24 females, Baoting, 80 m, 23., 24. and 27.vii.1960, S. F. Li, X. Z. Zhang & C. Q. Li leg.; 11 males, 15 females, Tongshi, 340 m, 23., 24. and 25.vi., 31.vii., 1., 4. and 6.viii.1960, S. F. Li, X. Z. Zhang & C. Q. Li leg.; 1 male, 1 female, Yinggen, 200 m, 4.v. and 5.vii.1960, S. F. Li leg.; 1 female, Shuiman, 640 m, 25.v.1960, C. Q. Li leg.; 2 males, Wanning, 10 m, 12., 13.iv.1960, S. F. Li & C. Q. Li leg.; 3 females, Qiongzhong, 15–17.vii.1960, X. Z. Zhang & C. Q. Li leg.; 2 males, 3 females, Kwangtung, 3., 4. and 5.iv., 13. and 26.viii.1934, C. Ho leg. (all IZCAS); 1 male, Mangrin, 9.vi.1904; 1 female, Mon Boi, 29.v.1904 (both BMNH).

Orthopagus hainanensis sp. n. is similar to O. lunulifer in most characters, but can be differentiated from the latter by the ivory white to pale ochraceous vertex with a pair of dark brown spots on each side of midline in basal third (in O. lunulifer, the vertex is nearly dark brown from base to apex) and the male segment X being elongate in dorsal view, with ratio of length to width near middle 1.9–2.0 (in O. lunulifer, the male segment X is shorter and wider in dorsal view, with ratio of length to width near middle 1.2–1.3). Orthopagus hainanensis sp. n. is also similar to O. exoletus in the predominantly light vertex but it differs from the latter by a slightly longer head, the male pygofer with dorso-posterior margin produced into a broad and short lobe in lateral view (the pygofer bears a larger tooth-like process dorso-posteriorly in O. exoletus) and the aedeagus with two pairs of larger (more strongly inflated) dorsolateral membranous lobes and endosomal processes covered with minute spines (indistinct in O. exoletus).

Measurements (5 males, 10 females). Body length (from apex of head to tip of forewings): male 11.7–12.9 mm, female 13.1–14.6 mm; head length (from apex of cephalic process to base of eyes): male 1.4–1.5 mm, female 1.5–1.6 mm; head width (including eyes): male 1.6–1.7 mm, female 1.6–1.7 mm; forewing length: male 9.3–10.3 mm, female 10.6–11.8 mm.

Coloration. General coloration as in generic description (Fig. 2E–F). Head ivory white to pale ochraceous, vertex with basal corners castaneous, a pair of large fuscous patches on each side of midline at basal third, and pale fuscous apical diamond-shaped spot (Fig. 6A); frons pale with series of small pale fuscous spots along intermediate and lateral carinae, base without distinct dark spot (Fig. 6C). Forewing pattern as in Fig. 3C. Hind wing membrane with a relatively narrow dark brown streak along the apical portion of CuA₁ vein, extending along hind wing apical margin.

Structure. Cephalic process (Fig. 6A–C) relatively elongate. Vertex (Fig. 6A) with ratio of length at midline to width between eyes 1.65–1.75. Transition of vertex to frons narrowly rounded in lateral view (Fig. 6B). Frons (Fig. 6C) relatively narrow, with ratio of length at midline to maximum width 2.7–2.9.

Male genitalia. Pygofer in lateral view (Fig. 6E) with dorso-posterior margin produced into a broad and relatively short blunt lobe; in ventral view (Fig. 6F) much longer than in dorsal view (Fig. 6D) with ratio of ventral to dorsal length about 4.2. Gonostyles (Fig. 6E–F) large, broad medially, with dorsal margin weakly sinuate. Aedeagus (Fig. 6G–I) with endosomal processes covered with minute spines, extended posteriad and strongly curved dorso-anteriad; phallobase with one pair of large, strongly inflated dorsolateral lobes, their apices gradually convergent and tapering posteriad (Fig. 6G–I), and one pair of small, thumb-like ventral lobes (Fig. 6I). Segment X relatively narrow and elongate, in lateral view, narrow basally, widening beyond middle, apex subacute (Fig. 6E), in dorsal view, widest medially, with ratio of length to width 1.9–2.0 (Fig. 6D).

Female genitalia as in generic description (Fig. 7A–G).

The new species is named for its occurrence in Hainan Island, China. The specific epithet hainanensis is to be treated as a latinized adjective in nominative singular.

So far only known from Hainan Island, China.

Orthopagus lunulifer: Lectotype, male, (1) 25,18,0., Gifu, male; (2) Type, No. 3123, U.S.N.M. [red label]; (3) Cotype No. U.S.N.M. [red label] (USNM). Paralectotypes: 1 male, (1) 25,18,0., Gifu, male; (2) Type, No. 3123, U.S.N.M. [red label]; (3) Cotype No. U.S.N.M. [red label]; 1 male, 1 female, (1) 25,8,0., Gifu; (2) Type, No. 3123, U.S.N.M. [red label]; (3) Cotype No. U.S.N.M. [red label]; 1 female, (1) 20,4,27., Gifu; (2) Type, No. 3123, U.S.N.M. [red label]; (3) Cotype No. U.S.N.M. [red label]; (4) Orthopagus lunulifer Uhler [Uhler’s handwriting]; (5) 1164 [Uhler’s handwriting] (all USNM).

Orthopagus helios: Lectotype (here designated), female, (1) Formosa, Ku Sia [handwriting, yellow label]; (2) helios Mel. [Melichar’s handwriting], det. Melichar; (3) Typus [dark red label]; (4) Transcriptio, Orthopagus helios sp.n. female [P. Lauterer’s handwriting], L. Melichar det 1912; (5) Collectio Dr. L. Melichar, Moravské museum Brno; (6) Syn-typus [red label]; (7) Invent. č. 4947/Ent., Mor. muzeum, Brno; (8) Lectotypus female, Orthopagus helios Melichar, 1912, designated by Z. S. Song & I. Malenovský, 2018 [newly added red label] (MMBC). Paralectotypes, 2 females, (1) Formosa, Ku Sia [handwriting, yellow label]; (2) Paralectotypus female, Orthopagus helios Melichar, 1912, designated by Z. S. Song & I. Malenovský, 2018 [newly added red label] (SNSD).

Orthopagus helios var. diffusus: Lectotype (here designated), male, (1) Formosa, Sauter; (2) Taihanroku, 908.; (3) v. diffusus M. [handwriting, underlined with red], det. Melichar; (4) typus [label with red frame]; (5) Hung. Nat. Hist. Museum Budapest, coll. Hemiptera [yellow label] (6) Lectotypus male, Orthopagus helios var. diffusus Melichar, 1912, designated by I. Malenovský in Song et al. 2018 [newly added red label] (HNHM). Paralectotype, male, (1) Formosa, Sauter; (2) Kosempo, 908.; (3) v. diffusus M. [handwriting, underlined with red], det. Melichar; (4) typus [label with red frame and a hindwing glued to it]; (5) Hung. Nat. Hist. Museum Budapest, coll. Hemiptera [yellow label] (6) Paralectotypus male, Orthopagus helios var. diffusus Melichar, 1912, designated by I. Malenovský in Song et al. 2018 [newly added red label] (HNHM, abdomen detached and glued to a separate label attached to the same pin).

Orthopagus elegans: Lectotype (here designated), female, (1) Formosa, Sauter; (2) Taihanroku, 908.; (3) elegantulus [Melichar’s handwriting], det. Melichar.; (4) Typus [dark red label]; (5) Collectio Dr. L. Melichar, Moravské museum Brno; (6) Orthopagus female elegans sp. n. female, L. Melichar det. 1912 [Lauterer’s handwriting], P. Lauterer’s det. 1991; (7) Syn-typus [red label]; (8) Invent. č. 4948/Ent., Mor. muzeum, Brno; (9) Lectotypus female, Orthopagus elegans Melichar, 1912, designated by Z. S. Song & I. Malenovský, 2018 [newly added red label] (MMBC). Paralectotype, female, (1) Formosa, Sauter; (2) Taihanroku, 908.; (3) elegans M. [handwriting, underlined with red], det. Melichar; (4) typus [label with red frame]; (5) Hung. Nat. Hist. Museum Budapest, coll. Hemiptera [yellow label] (6) Paralectotypus female, Orthopagus elegans Melichar, 1912, designated by I. Malenovský in Song et al. 2018 [newly added red label] (HNHM).

JAPAN: Honshu island: 1 female, Tokyo, Matsumura leg. (IZCAS); 1 male, Kamakura, ix.1913, F. Muir leg. (BPBM); 1 female, Mie prefecture, Matagari lke, 24.x.1989, C. W. O’Brien & L. B. O’Brien leg. (LBOB). CHINA: Beijing municipality: 1 female, Peiping; 9 males, 15 females, Peiping, 9., 10. and 21.vii.1938, 4., 13., 15. and 28.viii.1938, 11., 16., 19., and 24.ix.1938, T. P. Chang leg.; 1 male, 1 female, Juyongguan, 250–280 m, 3. and 6.viii.1961, S. Y. Wang & X. Z. Zhang leg.; 1 male, 1 female, Zhongguancun, 4.ix.1962, S. Y. Wang leg.; 3 males, 2 females, Shisanling, 12.ix.1962, R. Z. Xie leg. (all IZCAS); 2 males, 2 females, Changping, 8.vii.2007, Z. S. Song leg. (JSSNU); Tianjin municipality: Jixian, 4.ix.1988, K. H. Zhang leg. (IZCAS); Shandong province: 2 males, 1 female, Tsingtao [Musée Heude]; 1 male, Laoshan, 800 m [Musée Heude] (all IZCAS); Henan province: 1 male, 1 female, Henan, 8.viii.2013, D. J. Zhang (JSSNU); Anhui province: 2 males, Huang Mountain, 6.viii.1936 (IZCAS); Shangai municipality: 1 male, 1 female, 27.vii.1932, O. Piel leg. [Musée Heude] (IZCAS); Zhejiang province: 18 males, 10 females, T’ienmo Shan, 22–28.viii.1936; 2 females, Hangzhou, 24. and 25.viii.1942; 8 males, 8 females, Chusan, 7., 8., 10., 18., 20., 28. and 29.viii.1931, O. Piel leg. [Musée Heude] (all IZCAS); 1♀, Fujian province: 1 female, Jianyang, Chengguan, 90–120 m, 12.viii.1960, Y. R. Zhang leg.; 1 male, 1 female, Chongan, Xingcun, Sangang, 740 m, 12. and 20.viii.1960, Y. Zuo & C. L. Ma leg.; 1 male, 2 females, Jiangle, Longqishan, 500–700 m, 12., 13. and 19.viii.1991, S. M. Song leg. (all IZCAS); Hunan province: 1 female, Hoeng-Shan, 900 m, 1933, H. Höne leg. (MFNB); Guizhou province: 1 female, Libo, 21.viii.2000, F. M. Shi leg. (IZCAS); Sichuan province: 5 males, 3 females, Emei Mountain, Baoguosi, 550–750 m, 7., 9., 10. and 14.ix.1957, F. X. Zhu & Z. Y. Wang leg. (IZCAS); Tibet (Xizang) autonomous region: 1 female, Chayu, Xiachayu, 1900 m, 21.viii.2005, Z. S. Song leg. (IZCAS); Guangxi autonomous region: 4 males, 5 females, Guilin, Yanshan, 2., 7. and 23.viii.1952, 20., 22. and 24.vii.1953; 2 females, Pingxiang, 12. and 16.vi.1976, B. L. Zhang leg.; 5 males, 5 females, Nandan, Luofu, 350 m, 27.vii.2006, J. Liu leg. (all IZCAS); 2 males, 2 females, Guangnan, Bamei, Shiw. Taoyuan, 24°18'51"N, 105°02'08"E, 811 m, 12.viii.2012, D. Rédei leg.; 2 males, same but 24°19'08"N, 105°02'57"E, 891 m; 1 male 1 female, same but 24°19'11"N, 105°01'49"E, 834 m, 13.viii.2012 (all HNHM); Yunnan province: 1 male, 5 females, Hekou, 80 m, 5. and 7.vi.1956, K. R. Huang leg.; 3 females, Hekou, Nanxi, 200 m, 8. and 12.vi.1956, K. R. Huang leg.; 1 male, 1 female, Pingbian, 1400 m, 15.vi.1956, K. R. Huang leg. (all IZCAS); Taiwan: 1 male, Tainan, Formasa, vi.1912, H. Sauter, Orthopagus helios Mel. F. Schumacher det. [Schumacher’s handwriting] (SDEI); 2 males, 1 female, Taihanroku, vii. and 10.xi, H. Sauter leg. (MFNB); 1 male, 1 female, same data (MZPW); 5 males, 2 females, Taihanroku, 1908, Sauter leg. (HNHM); 1 male, 1 female, Hoozan, 10.vii. and 10.ix., H. Sauter leg. (MFNB); 1 female, Kotobuki, 11.vi.1935, (IZCAS); 2 females, Tainan county, ca. 350 m, 2–3 km S Kwantzuling, bamboo, shrub, 26–28.vi.1980, D. R. Davis leg. (USNM); 1 female, Chi Pen, 10.vi.1997, B. Herczig & L. Ronkay leg. (HNHM). VIETNAM: 1 female, “Indo China”, R. V. de Salvaza leg. (BMNH); 1 female, Chapa [= Sa Pa], v.–vi.1916, R. V. de Salvaza leg. (BMNH); 2 males, 1 female, Hoa Binh, vii.1939, A. de Cooma leg.; 5 males, 2 females Hoa Binh; 1 male, 1 female, Hoa Binh, Thanh-ha district, 12. and 13.vi.1966, R. Bielawski & B. Pisarski leg.; 2 males, 1 female, Hanoi, 24.vi.1959, B. Pisarski & J. Prószyński leg.; 4 males, 2 females Ninh Binh, Cuc Phuong, 5., 7. and 8.vi.1966, R. Bielawski & B. Pisarski leg.; 1 female, Nghe An district, Phu Quy, 17.vi.1959, B. Pisarski & J. Prószyński leg. (all MZPW); 1 female, Cuc Phuong, 400 m, at light, 17.x.1986, Vásárhelyi leg. (HNHM). LAOS: 1 male, Borikhane Prov., Pakkading, 31.vii.1965, native collector leg. (BPBM). INDIA: Assam: 2 males, Chabua, 10.x.1943, D. E. Hardy leg. (USNM); 1 female, Tocklai, light trap, ix.1983, 943/6, C.I.E.A. 15663 (BMNH). NEPAL: 1 male, Chitwan National Park, Island Jungle reserve, 29–30.x.1995, L. Peregovits leg. (HNHM).

Measurements (10 males, 9 females). Body length (from apex of head to tip of forewings): male 11.7–13.4 mm, female 13.0–14.9 mm; head length (from apex of cephalic process to base of eyes): male 1.33–1.50 mm, female 1.50–1.65 mm; head width (including eyes): male 1.30–1.60 mm, female 1.48–1.75 mm; forewing length: male 9.4–10.8 mm, female 10.2–11.9 mm.

Coloration. General coloration as in generic description (Figs 1A, 2G–H). Vertex dark brown with five light ochraceous streaks: along median carina in anterior third (in some specimens, the whole median carina is light), along each lateral carina subapically and along each lateral carina basally, the latter streaks being sickle-shaped and curved to median carina at the base (Fig. 8A). Frons light ochraceous with small dark brown spots along intermediate and lateral carinae, frons base slightly infuscated (Fig. 8C). Forewing membrane pattern as in Fig. 3D. Hind wing membrane with a dark brown streak along the apical portion of CuA₁ vein, extending along hind wing apical margin.

Structure. Head with cephalic process moderately long, not inflated (Figs 8A–C). Vertex (Fig. 8A) with ratio of length at midline to width between eyes 1.6–2.0. Transition of vertex to frons relatively sharp, almost angular in lateral view (Fig. 8B). Frons relatively narrow, with ratio of length at midline to maximum width 2.6–3.0.

Male genitalia. Pygofer, in lateral view (Fig. 8E), with dorso-posterior margin produced into a short and broad lobe; in ventral view (Fig. 8F) much longer than in dorsal view (Fig. 8D) with ratio of ventral to dorsal length about 4.0. Gonostyles (Fig. 8E–F) large, broad medially, with dorsal margin weakly sinuate. Aedeagus (Fig. 8G–I) with endosomal processes extended posteriad and strongly curved dorso-anteriad, relatively short, their apices not reaching the base of phallobase; phallobase with one pair of large, strongly inflated dorsolateral lobes (Fig. 8H), their apices blunt, and one pair of small, thumb-like ventral lobes, directed posteriad (Fig. 8I). Segment X, in lateral view, narrow basally, widening to apex beyond middle, apex subacute (Fig. 8E); in dorsal view, relatively short and broad in dorsal view, widest medially, with ratio of length to maximum width 1.2–1.3 (Fig. 8D).

Female genitalia as in generic description.

Widely distributed in tropical, subtropical and temperate eastern Asia (Japan, China, Korea, Vietnam, Laos, northeastern India and Nepal).

Adult O. lunulifer was reported as a minor pest of leaves of Morus alba L. (Pu and Mao 2012) and Camellia oleifera Abel in southeastern China (Zhao et al. 2013). Matsumura (1910) listed ”Anagnia splendens“ among pests of sugarcane in Taiwan; this record perhaps also refers to O. lunulifer.

Erroneously according to Metcalf (1946), Matsumura (1905a, b) considered O. lunulifer to be a junior synonym of Anagnia splendens (Germar). As he probably studied material from Japan, the description and illustration of “Anagnia splendens” in Matsumura (1905a) probably refer to O. lunulifer (i.e., the only Orthopagus species currently confirmed from Japan). The same is probable for the records and a figure of “Anagnia splendens” from Okinawa and Taiwan published in Matsumura (1905b and 1910, respectively).

Melichar (1912) differentiated O. elegans, O. helios and O. lunulifer based on slight differences in the transparency of the forewing membrane, extent of the dark brown apical band on the forewing and the shape of frons. Based on a study of material from Taiwan, Schumacher (1915) suggested that Orthopagus helios and O. elegans described by Melichar (1912) belong to the same species. However, his synonymisation of O. helios var. diffusus Melichar and O. elegans Melichar under O. helios Melichar was not widely accepted (Metcalf 1946). We have examined the corresponding type specimens and additional specimens from the same series collected by H. Sauter in Taiwan and currently deposited in HNHM, MFNB, MMBC, SDEI, and SNSD, and confirm here Schumacher’s conclusion. Simultaneously, we suggest that O. helios should be treated as a junior synonym of O. lunulifer because we consider the differences among these taxa listed by Melichar (1912) to represent intraspecific variation. We designate here the lectotypes for O. elegans, O. helios and O. helios var. diffusus to stabilize the nomenclature according to Article 74 of ICZN (1999).

Liang (1996) designated the lectotype for O. lunulifer, and provided a left lateral view of male genitalia for this species. Detailed illustrations of the male and female genitalia (but no detailed description) were also provided for O. lunulifer by Song et al. (2016d, 2018).

The single male specimen examined from Nepal (Chitwan National Park) is identical in external characters to specimens of O. lunulifer from Japan, China, Taiwan and Vietnam. However, it differs in the shape of the lobe on the dorso-posterior margin of the pygofer which is smaller (shorter and simply angular) than in the rest of O. lunulifer males studied. The phallobase of this specimen could not be sufficiently compared as its membranous lobes failed to inflate during the preparation. More specimens and data are needed to confirm the identification.

Lectotype male (here designated), (1) Los Banos, P.I. Baker; (2) 1311; (3) philippinus [Melichar’s handwriting] det. Melichar; (4) Collectio Dr. L. Melichar, Moravské museum Brno; (5) Orthopagus male philippinus sp.n., L. Melichar, 1914 [Lauterer’s handwriting], P. Lauterer det. 1991; (6) Syn-typus [red label]; (7) Invent. č. 4954/Ent., Mor. muzeum, Brno; (8) Lectotypus male, Orthopagus philippinus Melichar, 1914, designated by Z. S. Song & I. Malenovský, 2018 [newly added red label] (MMBC).

Paralectotypes, 4 females, same locality labels as holotype but Inv. nos 4949–4951, 4953;1 female, Mt. Makiling, Luzon, Baker (MMBC, Inv. no. 4952, all paralectotypes bearing the following label: Paralectotypus female, Orthopagus philippinus Melichar, 1914, designated by Z. S. Song & I. Malenovský, 2018).

PHILIPPINES: Luzon island: 1 male, Mt. Makiling, Baker leg. (USNM); 1 male, Los Baños, 11.xii.1913, D. T. Fullaway leg. (BPBM); 1 male, Los Baños, i.1913, P. Ledyard leg (LBOB); 2 males, Mt. Montalban, Rizal, Wa-wa Dam, 150–200 m, 6. and 17.iii.1965, H. M. Torrevillas leg. (BPBM); 1 male, 4 females, Manila, G. Boettcher leg. (MMBC); 1 male, 2 females, “B. M. 1925-491”, E. M. Ledyard leg.; 1 female, “Acc. No. 6625, Lot, Bu. of Sci., P. I.,1908-228”, C. S. Banks leg.; 1 male, “Acc. No. 5364, Lot, Govt. Lab. Coll., 1908-228”, C. S. Banks leg. (all BMNH).

Measurements (4 males, 9 females). Body length (from apex of head to tip of forewings): male 10.6–11.4 mm, female 12.9–13.9 mm; head length (from apex of cephalic process to base of eyes): male 1.25–1.33 mm, female 1.35–1.45 mm; head width (including eyes): male 1.30–1.43 mm, female 1.50–1.60 mm; forewing length: male 8.5–9.1 mm, female 10.3–11.4 mm.

Coloration. General coloration as in generic description (Figs 2I–J). Vertex predominantly dark brown, lateral margins with a pale ochraceous S-shaped streak in posterior three quarters (Fig. 9A). Frons light ochraceous with small dark brown spots along intermediate and lateral carinae and a large roundish dark brown spot at base (Fig. 9C). Forewing membrane pattern as in Fig. 3E. Hind wing membrane with a narrow dark brown infuscation along the apical portion of CuA₁ vein, not extending along hind wing apical margin.

Structure. Head with cephalic process moderately elongate (Figs 9A–B). Vertex (Fig. 9A) with ratio of length at midline to width between eyes 1.6–1.8. Transition of vertex to frons blunt, broadly rounded in lateral view, cephalic process thus appearing inflated and bulbous apically (Fig. 9B). Frons relatively narrow, with ratio of length at midline to maximum width 2.7–3.1.

Male genitalia. Pygofer, in lateral view, with dorso-posterior margin produced into a relatively large but narrow, tooth-like, apically obtuse process (Fig. 9E); in ventral view (Fig. 9F) much longer than in dorsal view (Fig. 9D) with ratio of ventral to dorsal width about 2.5. Gonostyles (Fig. 9G) large, broad medially, with dorsal margin weakly sinuate. Aedeagus (Fig. 9H–J) with endosomal processes covered with minute superficial spines, extended posteriad and strongly curved laterad and dorso-anteriad but not reaching the base of phallobase; phallobase with one pair of large, strongly inflated dorsolateral lobes, their apex convergent and tapering posteriad (Fig. 9I), and one pair of smaller, thumb-like ventral lobes, directed ventro-posteriad (Fig. 9I–J). Segment X, in lateral view, relatively narrow basally, widening to apex beyond middle, apex blunt (Fig. 9E); in dorsal view, relatively large and broad, widest in apical third, ratio of length to maximum width 1.0–1.1 (Fig. 9D).

Female genitalia as in generic description.

So far only known from the Luzon island, Philippines.

Orthopagus philippinus can be easily recognized from other species of the genus by the shape of the head and the presence of a relatively large roundish dark spot at base of frons which is present in all specimens studied.

Melichar (1914) indicated that he described this species based on five (male and female) specimens from “Luzon, Los Baños, Mt. Maquiling (C. F. Baker)”. Six specimens (1 male, 5 females) preserved in Melichar personal collection in MMBC and labelled as either from Los Baños or Mt. Makiling and collected by Baker are considered here as the original syntypes. The male specimen is designated here as the lectotype for O. philippinus to stabilize the nomenclature according to Article 74 of ICZN (1999).

CHINA: Hainan island: 1 male, Hainan, 5.v.1934, C. Ho leg.; 2 females, Shuiman, 640 m, 25.v.1960, C. Q. Li leg.; 2 males, 5 females, Tongshi, 340 m, 23. and 25.vi.1960, C. Q. Li leg.; 1 female, Yinggen, 200 m, 6.vii.1960, S. F. Li leg.; Yunnan province: 1 male, 1 female, Hekou, 80 m, 7.vi.1956, K. R. Huang leg.; 3 females, Hekou, Xioananxi, 200 m, 8.vi.1956, K. R. Huang leg.; 1 male, Gaoligong Mountain, 1000 m, 20.viii.1958, C. L. Li leg.; 1 male, Malipo, 20.vii.1958; 1 female, Xishuangbanna, Menga, 1050 m, 18.ix.1957, F. J. Pu leg.; 1 male, 1 female, Xishuangbanna, Menglun, 600 m, 9.ix.1993, H. L. Xu & L. L. Yang leg.; 1 male, Xishuangbanna, Menglun, 10.ix.1993, X. Y. Cheng leg. (all IZCAS). VIETNAM: 1 female, Lao Kay, 31.v.1960, at light, K. Galewski leg. (MZPW); 1 male, 1 female, Ninh Binh, Cuo-phuong, 5., 7, and 8.vi.1966, R. Bielawski & B. Pisarski leg. (MZPW); 2 males, 22 km S of Nha Trang, 20–26.xi.1960, C. M. Yoshimoto leg. (BPBM); 1 male, DaiLanh, N of Nha Trang, 30.xi.–5.xii.1960, C. M. Yoshimoto leg. (BPBM). THAILAND: Trang province: 1 male, Khaophappha Khaochang, 200–400 m, 3.i.1964, G. A. Samuelson leg. (BPBM). INDIA: Assam: 1 female, Mazbat near Mangaldai, 11–15.x.1910 (coll. Distant, BMNH). MALAYSIA: Penang: 1 male, Island of Penang, Baker leg. (USNM). INDONESIA: Java: 1 male, Java (West), Djasinga, 5.i.1966, J. Stusak (BPBM); 1 female, Samarang, iv.1909, E. Jacobson; 1 female, same but vi.1909 (both coll. Melichar, MMBC); 1 female, Wied. in CW. Java, GW [handwriting] (coll. Zool. Mus. Leipzig Übernahme 1971, SNSD); 1 male, Java [handwriting] (MFNB).

Measurements (3 males, 11 females). Body length (from apex of head to tip of forewings): male 10.3–11.8 mm, female 12.7–13.6 mm; head length (from apex of cephalic process to base of eyes): male 1.28–1.36 mm, female 1.35–1.48 mm; head width (including eyes): male 1.30–1.49 mm, female 1.50–1.59 mm; forewing length: male 8.5–9.1 mm, female 10.4–11.3 mm.

Coloration. General coloration as in generic description (Figs 1B, 2K–L). Vertex predominantly dark brown, lateral margins with a pale greenish or ochraceous S-shaped streak in posterior three quarters, median carina anteriorly light in some specimens (Figs 1B, 10A). Frons light ochraceous with small dark brown spots along intermediate and lateral carinae and indistinct infuscation at base (Fig. 10C). Forewing membrane pattern as in Fig. 3F. Hind wing membrane with a narrow dark brown infuscation along the apical portion of CuA₁ vein, extending into a narrow infuscation along hind wing apical margin.

Structure. Head with cephalic process relatively short (Fig. 10A–B). Vertex (Fig. 10A) with ratio of length at midline to width between eyes 1.5–1.6. Transition of vertex to frons narrowly rounded in lateral view, cephalic process not inflated apically (Fig. 10B). Frons relatively narrow, with ratio of length at midline to maximum width 2.8–3.1.

Male genitalia. Pygofer, in lateral view, similar to O. philippinus, with dorso-posterior margin with a large and relatively narrow, tooth-like, apically obtuse process (Fig. 10E); pygofer with ratio of ventral to dorsal width about 2.6 (Fig. 10D, F). Gonostyles (Fig. 10E) large, broad medially, with dorsal margin weakly sinuate. Aedeagus (Fig. 10G–I) with endosomal processes very long, extended posteriad and strongly curved dorso-anteriad, reaching the base of phallobase; phallobase with one pair of relatively small, inflated dorsolateral lobes, their apex elongate, thumb-like, directed posteriad (Fig. 10H), and one pair of relatively large ventral lobes, convergent and tapering posteriad (Fig. 10H–I). Segment X, in lateral view, narrow basally, widening to apex beyond middle (Fig. 10E); in dorsal view, elongate, widest at apical third, ratio of length to maximum width 1.5–1.6 (Fig. 10D).

Female genitalia as in generic description.

Confirmed records are from Indonesia (Java), West Malaysia, Vietnam, Thailand, southern China (Hainan, Yunnnan) and north-eastern India (Assam). Based on the description and illustrations in Yang and Yeh (1994), O. splendens probably also occurs in Taiwan (see also Tsaur 2005). Published records from the Philippines (Luzon; Stål 1861, Distant 1906) probably refer to O. philippinus, while the records from Sri Lanka (Melichar 1903, 1912; Distant 1906) concern O. exoletus and the ones from Japan (Matsumura 1905a, b, 1910) refer to O. lunulifer. Records from western India (Stål 1861), Myanmar (Distant 1906), Singapore, and Indonesia: Sumatra (Bierman 1908) still need to be checked.

Flata splendens was described by Germar (1830) from Java, and was designated as the type species of Anagnia by Stål (1861). Udugama flavocarinata Bierman, 1907 from Java was considered as a junior synonym of O. splendens by Melichar (1912). Orthopagus splendens var. tibialis Kirkaldy was differentiated from the typical form by having “the fore and middle tibiae distinctly bi- or tri-angulate with brownish” (Kirkaldy and Muir 1913). However, this pattern is typical for all Orthopagus species.

The synonymy of Dictyophara indiana Walker, 1851 with Orthopagus splendens proposed by Stål (1861) is considered doubtful (see above).

Orthopagus splendens has been frequently confused in literature and museum collections with other species of the genus. It can be differentiated from them by the combination of the following characters: a relatively small body size, a moderately short head, an extensive dark pattern on vertex, the pygofer with dorso-posterior margin bearing a relatively large but narrow, simply tooth-like process, the endosomal processes of the aedeagus relatively long and robust, covered with minute superficial spines and phallotheca bearing two pairs of membranous lobes.