Type series of C. favillacea Ito, 2011 (UOPJ; Fig. 10A, B): NEPAL: Taplejung: Kharu Pokhar, 3,000 m, 17.VII.1962, T. Yasuda, holotype ♂ of C. favillacea; Ilam: Phikol, 1,460 m, 19.IV.1962, T. Yasuda paratype 2♀, of C. favillacea. CHINA: Yunnan, Mengsong, Manlvcunhanzudazhai, small hilltop, 22°07'44.0"N, 100°28'51.7"E, 1690 m, 12.VII.2011, H.Y. Han and S.W. Suk, 72♂, 42♀ (YSUW); Yunnan, Mengsong, Bengangxizhai, in forest, 22°10'34.5"N, 100°35'06.8"E, 1725 m, 11.VII.2011, H.Y. Han and S.W. Suk, 1♀ (YSUW).

Figure 3. 

The habitat of Campiglossa coei. CHINA: Yunnan, Mengsong, Manlvcunhanzudazhai, small hilltop, 22°07'44.0"N, 100°28'51.7"E, 1,690 m, 12 July 2011. Many more than 100 individuals of C. coei were collected along with at least ten other species of the subfamily Tephritinae.

This light-colored species can be diagnosed by the following characteristics. Head largely yellow-brown with grey upper occiput. Thorax with scutum entirely matte whitish grey without any outstanding dark spots or stripes; scutellum mostly matte whitish grey but ca. apical 1/3 yellow-brown. Legs entirely yellowish brown without any dark marking; fore femur with six or seven strong, brown postero-ventral setae. Wing with basal area (basal 1/3 anteriorly and basal 1/2 posteriorly) largely hyaline with only few small dark spots, especially cell br with area posterior to fork of vein Rs completely hyaline (Fig. 4A-a); male with large dark mid-anterior marking covering from mid-anterior 1/3 to posterior end of crossvein R-M; pterostigma dark brown with large round hyaline spot in both sexes (Fig. 4A-b; in the other misella group species this spot tends to be smaller or missing in male); cell r₁ posterior to pterostigma with two large hyaline spots (sometimes with tiny additional basal spot, Fig. 4A-c) in male and three large hyaline spots in female; cell r₂₊₃ without posteroapical hyaline spot. Abdomen matte whitish grey with tergites 3–5 in male and 3–6 in female each with pair of pale brown submedian spots; oviscape shiny dark brown, as long as three preceding segments.

Figure 4. 

A–F Campiglossa coei A male, lateral view B male, dorsal view C male wing D female, lateral view E female, dorsal view F female wing G–K C. misella G male, lateral view H male, lateral view I male, dorsal view J female, lateral view K female, dorsal view.

This species appears similar to C. pishanica (with only males known) but the latter species can be readily separated by the dark femora and more extensive mid-anterior wing marking with pterostigma completely dark (Figs 4A, C vs. 10E).

Figure 5. 

Genitalia of Campiglossa coei A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G spermatheca.

Body (Fig. 4A–E) predominantly matte whitish grey; setae mostly brown to dark brown but some white; setulae mostly white but some brown to dark brown; wing length 4.0–4.3 mm; thorax length 1.5–1.8 mm.

Head yellow-brown with whitish pruinosity except for dark brown ocellar triangle and grey upper occiput; head ratio 0.78–0.90, frons-head ratio 0.46–0.50, eye ratio 0.71–0.77, gena-eye ratio 0.17–0.23, antenna-head ratio 0.40–0.44, arista-antenna ratio 1.3–1.6; vertex yellow-brown; dark brown inner vertical seta approx. as long as longest diameter of eye; outer vertical seta white, 0.4× inner vertical seta; post ocellar seta white, 0.4× post ocellar seta; paravertical seta white, 0.7–0.8× post ocellar seta; ocellar seta dark brown, 3.3–4.0× ocellar triangle length; frons almost bare with frontal angle ca. 115 degree; with two dark brown frontal setae; white posterior orbital seta 0.6× dark brown anterior orbital seta; scape and pedicel yellow-brown with short brown setulae; first flagellomere 1.4–1.8× pedicel length, apically rounded, yellow-brown; arista entirely short pubescent, brown except yellow-brown basal area; face yellow-brown without distinct antennal groove; parafacial 0.4× as wide as first flagellomere; facial ridge with fine pale yellow setulae; gena with strong white genal seta and relatively long white setulae; postgena swollen with strong white postgenal seta and relatively long white setulae; postocular setae with two thick white setulae plus over ten shorter brown setulae, extended 0.5× distance from upper eye margin to lower eye margin; supracervical setae white; mouthparts geniculated with yellow-brown setulose labella; palpus with brown setulae apically and white setulae on remaining area.

Thorax largely dark brown ground color with very heavy whitish pruinosity, generally appearing matte whitish grey; postpronotal lobe with single dark brown seta, yellow-brown in ground color, but appearing similar color as nearby areas due to heavy whitish pruinosity; scutum matte whitish grey with five faint brownish longitudinal bands traceable in clean specimens; two pairs of white scapular setae; acrostical setae widely separated each other, situated midway between levels of intra-alar setae and postsutural supra-alar setae; post-alar setae same level as intra-alar setae; dorsocentral setae same level as or slightly lower than transverse suture; presutural supra-alar setae approximately the same level as anterior notopleural setae; two notopleural setae dark brown with posterior seta0.5× anterior seta; scutellum mostly matte whitish grey but ca. apical 1/3 yellow-brown, slightly convex, almost bare except marginal tiny white setulae; basal scutellar setae more or less parallel, 2.3–3.5× as long as scutellum; apical scutellar setae crossed near apex, 0.9–1.3 as long as scutellum; pleura largely matte whitish grey; proepisternum with 3–5 white setulae; anepisternum matte grey with posterior 2/3 white setulose, with one strong dark brown seta and one half as long white seta ventral to it; katepisternum matte grey with a strong dark brown seta, upper area sparsely with short white setulae and lower area with long white setulae; mediotergite matte grey.

Legs entirely yellow-brown with slight grey pruinosity and brown to dark brown setae and setulae; fore coxa anteriorly with white setulae, posteriorly bare; mid coxa anteriorly with few long white setulae, posteriorly bare; hind coxa with strong white lateral seta, posteriorly largely membranous; front femur with six or seven strong brown posteroventral setae; tibiae and tarsi entirely yellow-brown; midtibial spur dark brown, 1.2–1.4 as long as tibial width.

Wing (Fig. 4A, C, D, F) hyaline with brown to dark brown pattern; area around pterostigma with sexual dimorphism (see next paragraph); cells bc, bm, bcu, alula, anal lobe almost entirely hyaline; cell c mostly hyaline with narrow brown to faint brown medial longitudinal band; pterostigma with distinct hyaline spot in both sexes (Fig. 4A-b); cell r₂₊₃ mostly without apical hyaline spot but with two large subapical spots often coalesced, one or two large hyaline spots posterior to two large r₁ spots; cell br with basal 3/5 area almost hyaline, apically dark brown with two or three hyaline spots posteriorly coalesced; cell r₄₊₅ with single large apical spot and 8–12 variably sized hyaline spots; cell dm with basal 2/3 almost hyaline, apically dark brown with 4–7 variably shaped hyaline spots; cell m with basal 3/4 almost hyaline, apically dark brown with 1–3 variably shaped hyaline spots. Wing-thorax ratio 2.4–2.5; subcostal to costa ratio 0.43–0.53; cell r₁-r₂₊₃ ratio 2.7–3.3; cell r₄₊₅-r₂₊₃ ratio 0.58–0.73. R₄₊₅ bare.

Wing dimorphism. Male (Fig. 4A, C) with cell r₁ with two large hyaline spots apical to pterostigma (rarely tiny additional spot anteriorly; Fig. 4A-c); large, more or less elliptic dark brown mid-anterior marking traceable covering pterostigma, cell r₁ well beyond pterostigma, approx. basal 1/3 to 2/3 of cell r₂₊₃, and anterior areas of cells br and r₄₊₅ near crossvein R-M; vein M ratio 0.40–0.45. Female (Fig. 4D, F) – cell r₁ with three large hyaline spots apical to pterostigma; dark brown mid-anterior marking, if traceable, much smaller, or not wider than pterostigma; vein M ratio 0.62–0.76.

Male abdomen. Preabdomen slightly longer than wide, almost entirely matte pale grey; tergites 2–5 with white setulae, but tergite 5 also with 4–7 dark brown marginal setae; tergites 3–5 each with pair of pale brown submedian spots. Postabdomen (Fig. 6A–C) with proctiger short, 0.4× as long as epandrium in lateral view, microtrichosae, lower half with numerous yellow-brown setae; epandrium plus surstyli oval in caudal view; epandrium dark brown with long yellow-brown to brown setae, microtrichosae; lateral surstylar flange posteriorly serrate, with its basal width approx.1/3 as long as epandrial complex height; medial surstylus with lateral prensiseta approx.2/3 as long as medial prensiseta; preglans area of phallus strongly spinulose; glans without subapical lobe; tube-like acrophallus highly pronounced with apicodorsal opening, approx. half as long as glans; ejaculatory apodeme large, fan-shaped.

Figure 6. 

Genitalia of Campiglossa misella A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G Spermatheca.

Female abdomen. Preabdomen slightly longer than wide, almost entirely matte grey; tergites 2–6 with white setulae, and tergite 6 especially with 4–7 dark brown marginal setae; tergites 3–6 each with pair of pale brown submedial spots. Postabdomen (Fig. 6D–G) with shiny dark brown oviscape approx. as long as three preceding segments; oviscape densely with dark brown setulae but without any macrosetae, 1.8× longer than wide, cone shaped, dorsoventrally flattened; eversible membrane with taeniae approx.1/4 as long as total length of membrane; posterior 3/4 area of eversible membrane densely covered with anteriorly directed triangular spinules; spinules largest in area behind taeniae; aculeus elongated, dorsoventrally flattened, 5.5× longer than wide with apical 1/3 gradually pointed, apex with pair of tiny subapical teeth; two similar sized dark brown spermathecae, each with elliptical apical receptacle with transverse papillae and 3/5 as long narrow basal neck; spermathecal duct transparent.

Nepal, China (Yunnan).

The male wing pattern of C. coei is atypical for the genus Campiglossa (Fig. 4A, C), and that is probably why this species, based on a single male specimen, was originally classified as Tephritis by Hardy (1964). Since then, Wang (1998), under this name, recorded two males from Yunnan, China. More recently, Ito (2011) described a new species (C. favillacea syn. nov.) based on the male holotype (from the type locality of C. coei) and two female paratypes (Fig. 10A, B), but he did not mention their wing dimorphism in the description. We, fortunately, were able to collect over a hundred male and female specimens from China, showing a remarkable sexual wing dimorphism (Fig. 4A, C vs. Fig. 4D, F). Most of the specimens were collected along with at least ten other species of the subfamily Tephritinae from a small hilltop in Yunnan, China (Fig. 3; Mengsong, Manlvcunhanzudazhai, 22°07'44.0"N, 100°28'51.7"E, 1690 m, 12 July 2011). This hilltop appears to be a temporary Tephritinae hot spot due to the clearing of a small forest patch.

HUNGARY: Bdaors, Odvas hg., 18.VI.1991, B. Merz and Adams, 1♀ (YSUW). ITALY: Aosta, St. Pierre, M. Torrette, 800–850 m, 22.IV.2003, B. Merz and F. Amiet, 1♂ (YSUW). NEPAL: Taplejung: Walungchung Gola, 3,350 m, 14.VI.1962, T. Yasuda, holotype ♂ of C. roscida (UOPJ; Fig. 10C, D). SWITZERLAND: Valais 642 m, St. German/Brüke, 3.VIII.1998, B. Merz and G. Bächli, 1♀; Valais, Leuk-Rotafen, 46°18'59"N, 7°40'18"E, 640 m, 22.VII.2004, H.Y. Han and K.E. Ro, 1♂ 1♀ (YSUW); Valais, Visperterminen-Kreuz, 46°15'17"N, 7°53'52"E, 1500 m, 21.VII.2004, H.Y. Han and K.E. Ro, 2♀ (YSUW). KYRGYZSTAN: S-Issik-Kul nr. Barskaun vill., 31.VII.1995, S.V. Ovchinnikov, 1♀ (YSUW); Telash Mt. r./ N slope, Ara-Bijik rav, 2300 m, 4.VII.1998, D. Milko, 1♂ (YSUW).

Males of C. misella usually have distinct sexually dimorphic wing patterns [e.g., Fig. 4G from Kyrgyzstan is almost identical to the male syntype photograph by Hendel (1927)] but some European populations seem to show slight sexual dimorphism (e.g., Fig. 4H from Switzerland). More extensive survey is required to understand their variation, but they could still be readily diagnosed even based on our limited samples. Head largely yellowish brown with grey upper occiput. Thorax with scutum entirely ash-grey with five brownish longitudinal stripes (Fig. 4I, K); bases of acrostichal, dorsocentral, intra-alar, basal scutellar setae dark brown; scutellum ash-grey with lateral margins brown, apex yellowish brown; Legs with femora largely dark grey except for yellowish brown apices (Fig. 4G, H, J), but tibiae and tarsi yellowish brown; fore femur with six or seven dark brown posteroventral setae. Wing with basal half largely with dark spots, especially cell br posteroapical to fork of vein Rs with dark brown rectangular area (approx. twice as wide as long; Fig. 4G-a, J-a); male often with large dark mid-anterior marking covering from mid-anterior 1/3 to posterior end of crossvein R-M (Fig. 4G); pterostigma almost completely dark brown in such sexually dimorphic male (Fig. G), but with large hyaline spot in minimally dimorphic male (Fig. 4H), and female (Fig. 4J); cell r₁ apical to pterostigma with three large hyaline spots with 1^st and 3^rd spots much smaller than middle one in dimorphic male (Fig. 4G-b), but with three large similarly sized hyaline spots in female (Fig. 4J) or minimally dimorphic male (Fig. 4H); cell r₂₊₃ without posteroapical hyaline spot. Abdomen ash-grey with tergites 3–5 in male and 3–6 in female each with pair of brown submedian spots; oviscape shiny dark brown, as long as four preceding segments.

Europe, Central Asia, China (Xinjian, Shanxi, Sichuan, Xizang, Yunnan), Nepal.

This is the only species of the misella group with host feeding biology known. Interestingly, White (1988) reported that this species usually attacks the flowering spike of Artemisia vulgaris, inducing a stem gall in the first generation and developing in the capitula in the second generation in the UK. In addition to Ar. vulgaris, Korneyev and Kameneva (1993) listed Ar. santolinifoliae and Ar. dracunculus as their host plants in Central Asia (Kazakhstan).

We resurrected C. coei and C. pishanica from the synonymy of C. misella by Korneyev (2014). Our study indicates that C. coei is a valid species (Figs 1, 2). Campiglossa pishanica is somewhat similar to C. misella in having the dark femora and the large mid-anterior wing marking, but C. pishanica has the following characteristics that, we posit, are beyond the variation range of the C. misella wing pattern (Figs 4G vs. 10E): cell r₁ apical to pterostigma with two hyaline spots instead of 3, basal 3/4 of cell dm almost hyaline, and anal lobe hyaline. See also the Remarks of C. pishanica for further discussion.

Holotype ♂: KOREA: Gyeongsangbuk-do, Bonghwa-gun, Myeongho-myeon, Mt. Cheongnyangsan, 36°46'43.6"N, 128°55'0.8"E, 600 m, 30.VI.2007, H.Y. Han et al. (NIBR). Paratypes: RUSSIA: Primorsky-Krai: between Chernyatino and Pokrovk, 43°57'32.7"N, 131°32'24.1"E, 55 m, 26.VI.2008, H.Y. Han and H.S. Lee, 3♂ 3♀; Khasansky-District, Kedrovaya Pad, 43°05'09.4"N, 131°35'06.0"E, 22 m, 23.VI.2008, H.Y. Han and H.S. Lee, 1♂; Khasansky-District, Barabash, 43°10'46.9"N, 131°28'20.0"E, 61 m, 22.VI.2008, H.Y. Han and H.S. Lee, 1♀; Ussuriysk, 43°47'05.4"N, 132°01'37.8"E, 19 m, 26.VI.2008, H.Y. Han and H.S. Lee, 1♀. All paratypes in YSUW.

The specific epithet is derived from the closely related species melaena prefixed with para.

This new species can be diagnosed by the following characteristics. Head largely yellowish brown with grey upper occiput. Thorax with scutum entirely ash-grey with five faint brownish longitudinal stripes (Fig. 7B, E); bases of acrostichal, dorsocentral, intra-alar, basal scutellar setae dark brown; scutellum ash-grey with apex yellowish brown; Legs with femora largely dark grey except for yellowish brown apices (Fig. 4G, H, J), but tibiae and tarsi yellowish brown; fore femur with six or seven dark brown posteroventral setae. Wing with basal area (basal 1/3 anteriorly and basal 1/2 posteriorly) largely hyaline with only few small dark spots, especially cell br with area posterior to fork of vein Rs completely hyaline (Fig. 7C-a); male with large dark mid-anterior marking covering from pterostigma to posterior end of crossvein R-M; male pterostigma almost completely dark brown, at most with tiny hyaline spot (Fig. 7C-b); female pterostigma with large round hyaline spot (Fig. 7D-a); cell r₁ posterior to pterostigma with three large hyaline spots in both sexes; cell r₂₊₃ without posteroapical hyaline spot (Fig. 7C-c, D-a). Abdomen ash-grey with tergites 3–5 in male and 3–6 in female each with pair of brown submedian spots; oviscape shiny dark brown, as long as three preceding segments.

Figure 7. 

A–E Campiglossa paramelaena sp. nov. A male, lateral view B male, dorsal view C male wing D female, lateral view E female, dorsal view F female, wing G–K C. melaena G male, lateral view H male, dorsal view I male, wing J holotype male, wing K female, wing J, K Reproduced from Korneyev and Ovchinnikova (2004) with permission from Valery Korneyev.

Campiglossa paramelaena sp. nov., appears similar to C. misella but the former species can be readily separated by the almost hyaline basal area of the wing, and the area posterior to the fork of vein Rs in particular is completely hyaline while the latter species has a distinctly dark spot on that area (Fig. 7C-a vs. Fig. 4G-a, J-a).

Body (Fig. 7A–F) predominantly ash-grey; setae mostly dark brown but some white; setulae mostly white but some dark brown; wing length 3.0–3.8 mm; thorax length 1.2–1.5 mm.

Head yellow-brown with whitish pruinosity except for dark grey ocellar triangle and upper occiput; head ratio 0.85–0.92, frons-head ratio 0.47–0.53, eye ratio 0.75–0.83, gena to eye ratio 0.17–0.22, antenna-head ratio 0.41–0.46, arista-antenna ratio 1.3–1.7; vertex yellow-brown; dark brown inner vertical seta approximately as long as longest diameter of eye; outer vertical seta white, 0.4× inner vertical seta; post ocellar seta white, 0.3–0.4× post ocellar seta; paravertical seta white, 0.7–0.9× post ocellar seta; ocellar seta dark brown, 3.0–3.5× ocellar triangle length; frons almost bare with frontal angle 110–115 degree; with two dark brown frontal setae; white posterior orbital seta 0.6–0.8× dark brown anterior orbital seta; scape and pedicel yellow-brown with short dark brown setulae; first flagellomere 1.5–2.1× pedicel length, apically rounded, yellow-brown but with greyish tinge in some individuals; arista entirely short pubescent, dark brown except yellow-brown basal area; face yellow-brown without distinct antennal groove; parafacial 0.4–0.5× as wide as first flagellomere; facial ridge with fine pale yellow setulae; gena with strong white genal seta and relatively long white setulae; postgena swollen with strong white postgenal seta and relatively long white setulae; postocular setae with two thick white setulae plus ten or more shorter dark brown setulae, extended 0.6× distance from upper eye margin to lower eye margin; supracervical setae white; mouthparts geniculated with labella yellow-brown setulose; palpus with brown setulae apically, white setulae on remaining area.

Thorax largely dark brown in ground color with heavy whitish grey pruinosity, generally appearing ash-grey; postpronotal lobe with single dark brown seta, yellow-brown in ground color, therefore, appearing paler than nearby areas; scutum ash-grey with five faint brownish longitudinal bands traceable in clean specimens; two pairs of white scapular setae; acrostical setae widely separated, situated midway between levels of intra-alar setae and postsutural supra-alar setae; post-alar setae same level as intra-alar setae; dorsocentral setae approximately same level as transverse suture; presutural supra-alar setae slightly above level of anterior notopleural setae; two notopleural setae dark brown with posterior seta 0.5× anterior seta; bases of acrostichal, dorsocentral, intra-alar, basal scutellar setae dark brown; scutellum mostly ash-grey but ca. apical 1/5 yellow-brown, slightly convex, almost bare except marginal tiny white setulae; basal scutellar setae more or less parallel, 3.1–3.6× (in males) and 2.4–3.0× (in females) as long as scutellum; apical scutellar setae crossed near apex, 1.1–1.4× (in males) and 0.9–1.1× (in females) as long as scutellum; pleura largely ash-grey; proepisternum with 3–5 white setulae; anepisternum ash-grey with posterior 2/3 white setulose, with single strong dark brown seta and one seta half as long and white ventral to it; katepisternum ash-grey with a strong seta, upper area sparsely covered with short white setulae and lower area with long white setulae; mediotergite ash-grey. Legs yellow-brown ground color with ash-grey pattern and brown to dark brown setae and setulae; fore coxa yellow-brown with posterobasal 1/3 grey, anteriorly with white setulae, posteriorly bare; midcoxa yellow-brown, anteriorly with few long white setulae, posteriorly bare; hind coxa greyish yellow-brown, with white lateral seta, posteriorly largely membranous; femora largely ash-grey except yellow-brown apices; tibiae and tarsi entirely yellow-brown; midtibial spur dark brown, 1.0–1.3× as long as wide.

Wing (Fig. 5A, C, D, F) hyaline with brown to dark brown pattern; area around pterostigma with distinct sexual dimorphism (see next paragraph); cells bc, bm, bcu, alula, anal lobe almost entirely hyaline; cell c mostly hyaline with narrow brown to faint brown medial longitudinal band; cell r₁ with basal 1/4 hyaline, apical 3/4 dark brown with three large hyaline spots apical to pterostigma; cell r₂₊₃ without apical hyaline spot but with two large subapical spots often coalesced, two large hyaline spots posterior to three r₁ spots, two or three tiny spots apical to them; cell br with basal 2/3 almost hyaline, apically dark brown with 1–3 hyaline spot; cell r₄₊₅ with single apical spot and 8–12 variably shaped hyaline spots; cell dm with basal 2/5 almost hyaline, apically dark brown with 4–7 variably shaped hyaline spots; cell m with 5–7 hyaline spots; cell cu₂ with six or seven large hyaline spots coalesced each other resulting in largely hyaline background with few small brown spots. Wing-thorax ratio 2.4–2.6, subcosta-costa ratio 0.53–0.64, cell r₁-r₂₊₃ ratio 2.2–2.7, cell r₄₊₅-r₂₊₃ ratio 0.54–0.67. R₄₊₅ bare.

Wing dimorphism. Male (Fig. 7A, C) with pterostigma entirely dark brown or at most with tiny hyaline spot; large, more or less elliptic dark brown mid-anterior marking traceable covering pterostigma, cell r₁ adjacent to pterostigma, basal 1/4 to 3/5 of cell r₂₊₃, and anterior areas of cells br and r₄₊₅ near crossvein r-m; vein M ratio 0.29–0.43. Female (Fig. 5D, E) with pterostigma dark brown with distinct round hyaline spot; large mid-anterior marking not traceable; such marking interrupted by distinct round hyaline spot on pterostigma and 2–4 small round spots on cell br posterior to it; vein M ratio 0.41–0.53.

Male abdomen. Preabdomen slightly longer than wide, almost entirely ash-grey; tergites 2–5 with white setulae, but tergite 5 also with 5–7 dark brown marginal setae; tergites 3–5 each with pair of brown submedian spots. Postabdomen (Fig. 8A–C) with proctiger short, 0.4× as long as epandrium in lateral view, microtrichosae, lower half with numerous yellow-brown setae; epandrium plus surstyli oval in caudal view; epandrium dark brown with long yellow-brown to brown setae, microtrichosae; lateral surstylar flange posteriorly serrate, with its basal width approx.1/3 as long as epandrial complex height; medial surstylus with lateral prensiseta approx.2/3 as long as medial prensiseta; preglans area of phallus strongly spinulose; glans without subapical lobe; tube-like acrophallus highly pronounced with apicodorsal opening, approx. half as long as glans; ejaculatory apodeme large, fan-shaped.

Figure 8. 

Genitalia of Campiglossa paramelaena sp. nov. A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus D female postabdomen with aculeus and eversible membrane pulled out, ventral view E magnified view of aculeus tip F magnified view of oviscape and eversible membrane G spermatheca.

Female abdomen. Preabdomen slightly longer than wide, almost entirely ash-grey; tergites 2–6 with white setulae, tergite 6 especially with dark brown marginal setae; tergites 3–6 each with pair of brown submedial spots. Postabdomen (Fig. 8D–G) with shiny dark brown oviscape approx. as long as three preceding tergites; oviscape densely covered by dark brown setulae but without any macrosetae, 1.3× longer than wide, cone shaped, dorsoventrally flattened; eversible membrane with taeniae approx. 1/3 as long as total length of membrane; posterior 2/3 area of eversible membrane densely covered with anteriorly directed triangular spinules; spinules largest in area behind taeniae; aculeus elongated, dorsoventrally flattened, approx. 4× longer than wide with apical 2/5 gradually pointed, apex with pair of tiny subapical teeth; two similar sized dark brown spermathecae, each with pear-shaped apical receptacle with transverse wrinkles and half as long narrow basal neck; spermathecal duct transparent.

Korea, the Russian Far East.

Individuals of C. paramelaena sp. nov., have DNA barcodes (Figs 1, 2) indistinguishable from those of C. melaena, which is a distinctly darker species with a more extensive wing pattern (Fig. 7G–I). Superficially, C. paramelaena sp. nov., more closely resembles C. misella (see Diagnosis), while the average barcode distance between these two species is 1.9 % (range 1.7–2.1 %). We postulate that C. paramelaena sp. nov., is not a light-colored seasonal form of C. melaena, because both species are from the same collecting lot in the Russian Far East (see Type material). Moreover, this species not only has a lighter body coloration but also has a much sparser wing pattern on the anal area then in C. melaena. In addition, the male surstylar flange of C. melaena is relatively larger (the base of the flange is approx. half as long as the height of the epandrial complex in the lateral view) than that of C. paramelaena sp. nov. (the base of the flange is distinctly shorter than half the height of the epandrial complex) (Fig. 9A vs. Fig. 8A).

Figure 9. 

Male genitalia of Campiglossa melaena A epandrial complex, lateral view B epandrial complex, caudal view C glans and preglans of distiphallus.

. Russia: Primorsky-Krai: Khasansky-District, Barabash, 43°10'46.9"N, 131°28'20.0"E, 61m, 22.VI.2008, H.Y. Han and H.S. Lee, 3♂ (YSUW); Nadezhdinsky-District, Vol’no-Nadezhdinskoye, grassland near restaurant, 43°22'31.6"N, 132°01'43.1"E, 61m, 22.VI.2008, H.Y. Han and H.S. Lee, 3♂ (YSUW).

This is the darkest species of the misella group, showing the least wing dimorphism (Fig. 7G, I, J vs. K). Head largely brown with dark grey upper occiput. Thorax with dark grey scutum with five brownish longitudinal stripes (Fig. 7H); scutellum dark grey; Legs with coxae and femora largely dark grey but tibiae and tarsi brown; fore femur with 5–7 dark brown posteroventral setae. Wing almost entirely brown to dark brown with numerous hyaline spots; male with large dark mid-anterior marking covering from pterostigma to posterior end of crossvein R-M; male pterostigma almost completely dark brown, at most with tiny hyaline spot (Fig. 7I-a); female pterostigma with larger hyaline spot (Fig. 7K-a); cell r₁ posterior to pterostigma with three large hyaline spots in both sexes; cell r₂₊₃ with posteroapical hyaline spot (Fig. 7I-b). Abdomen almost entirely dark grey.

North east China, the Russian Far East.

Hering’s (1941) original description and wing drawing of the holotype from north east China fall clearly within the variation range of the specimens we obtained from the Russian Far East. Unfortunately, we were not able to collect any female specimens, but Korneyev and Ovchinnikova’s (2004) illustrations (Fig. 7J, K) show a similar sexual dimorphism of the wing pattern as in the other misella group species. Individuals of C. melaena have DNA barcodes (Figs 1, 2) indistinguishable from those of C. paramelaena sp. nov. (see the Remarks of the latter species for further discussion).

This is an interesting species showing the characteristics of both C. coei and C. misella. The only known C. pishanica male wing pattern is very similar to that of C. coei (Fig. 10E vs. Fig. 4A, C), but Fig. 10E shows the following differences: pterostigma almost completely dark with very tiny hyaline spot (Fig. 10E-a; C. coei male consistently has a much larger spot, Fig. 4A-a), and fork of vein Rs and area posterior to it with dark spot. Except for the much lighter basal wing area, C. pishanica body appears very similar to that of C. misella, which also has dark femora and a scutum with five stripes.

Figure 10. 

A Holotype male of Campiglossa favillacea Ito, 2011 (new synonym of C. coei, UOPJ) B from left, two paratype females and holotype of C. favillacea [UOPJ] C holotype female of C. roscida Ito, 2011 (new synonym of C. misella) D ditto E holotype or paratype male wing of C. pishanica F holotype male wing of C. propria E, F reproduced from Wang (1998) with permission from Xing-Jian Wang.

Only three males (the type series) known from China (Xinjian).

We are not sure if this species actually belongs to the misella group, because the only known male (holotype) does not show close similarity to any known member of the group except for its large mid-anterior dark wing marking (Fig. 10F). This male also shows an unusual enlargement of cell r₁ resulting in a distinctly more rounded anterior wing margin than other species (Fig. 10F-a vs. Fig. 10E-a). In addition to this peculiar enlarged cell r₁, C. propria male can also be diagnosed based on the following characteristics: scutum ash-grey with five brownish longitudinal stripes; legs entirely yellowish; cell r₁ apical to pterostigma with three tiny hyaline spots plus a large subapical hyaline spot; cell r₂₊₃ basal to crossvein R-M dark without any spot, apical to R-M with six hyaline spots including posteroapical spot; abdominal tergite 3–5 each with pair of large brown submedian spots.

Only known from the type locality (Gansu, China).