Canthophorus melanopterus melanopterus ( Herrich-Schaeffer, 1835 )

Gapon, Dmitry A., 2018, Morphology of male and female terminalia and taxonomic revision of the burrower bugs genus Canthophorus (Heteroptera: Cydnidae), Annales de la Société entomologique de France (N. S.) 54 (3), pp. 183-228 : 23-28

Canthophorus melanopterus melanopterus ( Herrich-Schaeffer, 1835)

( Figures 1 C , 3 E , 4 C –4 G , 6 J –6 M , 7 C, 7 D , 8 A –8 G , 9 C , 12 B , 13 A –13 J , 14 A –14 G )

Material examined

Without locality: “ 316 ”, [date unknown] (A. Kuschakewitsch coll.), 1♀ ( ZISP); “ Fl. Bielaja ”, [date unknown] (A. Kuschakewitsch coll.), 1♀ ( ZISP); “ RussianGerman front ”, 1915 (Alexey Kiritshenko), 1♀ ( ZISP); “ Caucasus ”, [date unknown] (Barovskij), 1♀, [date unknown] (Stark), 1♀ ( ZISP); “ NW Caucasus ”, [date unknown] (Stark), 1♀ ( ZISP). Morocco: Oriental Prov., Oudjda, 1930 (coll. Sicard), 3J, 1♀ ( MNHN); Souss-Massa-Drâa, Aourir env., 30°29′34 ʺ N 9°40′31.9 ʺ W, 6,7. VI.2015 (D.A. Gapon), 1♀ ( ZISP). Algeria: “ Algérie. C[oll]. [A.F.] Morawitz [18]60.”, 1♀ ( ZISP); Sidi-bel-Abbes, [date and collector unknown], 1♀ ( ZISP); Aurés Mts, Djebel Chélia Mt. , 1800–2000 m, 22.VI.1981 (Doguet S.), 1J, 2♀ ( MNHN). Tunisia: “ Tunisia ”, 10.VI.1913 (N.L. Pastukhov), 1J, 1♀ ( ZISP); Sousse Gov., Sidi el Hani , V.1884 [collector unknown], 1J, 1♀ ( MNHN); Sfax Gov.: “ Tunisie Mérid.”, Sfax et environs, III.1922 (G. Babault), 1♀ ( MNHN); “ Hachichina ” [ Al Hushayshinah ], 13.V.1966 (coll. Hannothiaux), 1J, 1♀ ( MNHN). Libya: “ Env. de Tripoli ”, 1877 (Mouchez), 1♀ ( MNHN). Egypt: “ Aegypt.”, [date and collector unknown], 1♀ ( ZISP); “ Egypt ”, [date unknown] (coll. Eversmann), 1♀ ( ZISP); “ Egypte ”, “66–96” (Letourneux), 1J, 1♀ ( MNHN). Spain: Andalusia, Almeria, 2.V.1952 (H. Coiffait), 1♀ ( MNHN). France: Provence-Alpes-Côte d’ Azur, Nice , [date unknown] (G. V. Olsufjev), 1♀ ( ZISP). Italy: Sardinia, Asuni, 6–11.V.1900 (A. Krausse), 1J ( ZISP); Trieste Prov., Duino , 1933 (Passauro), 1J ( ZISP). Greece: “Graecia”, [date and collector unknown], 2♀ ( ZISP); Lake Kremasta env., “ de Houni à Karpenissi ”, 2. VII.1987 (A. Matocq), 1J ( AMC); Euboea Isl., Chalkis , III.1926 (Holtz), 1♀ ( ZISP); Athens, 3.III.1914 (Pastukhov), 1J, V–VI.1919 (L. Shelyuzhko), 2♀ ( ZISP), 27.III.1937 (Zhenzhurist), 1J ( ZMUM). Slovenia: “91./ Cydnus dubius Carniol [ia; Krajina ]”, 1♀ ( ZISP). Serbia: Belgrade, Topčider, 17.VII.1927, 20.IV.1928 (V. Martino), 3♀ ( ZISP). Ukraine: Khmel’ nits’ kiy Prov.: Kamenets-Podol’ skiy , 1907 (Yakubovskiy), 2♀ ( ZISP); Kamenets-Podol’ skiy env., downstream of Muksha Riv. , 14.III.1911 (Yakubovskiy), 1♀ ( ZISP); Odessa Prov.: Khadzhibeyskiy Liman, solonchak , 29.II.1920 (Alexey Kiritshenko), 1♀ ( ZISP); W coast of Khadzhibeyskiy Liman , 27.III.1921 (D.V. Znojko), 1♀ ( ZISP); Odessa, Lanzheron, 17.V.1922 (G. Tanfiljew), 1♀ ( ZISP); Luzanovka, 22.VI.1925 (Kiritshenko), 1♀ ( ZISP); Mykolaiv Prov.: “Rakovskiy poselok” [ Rakovo Settlm .], 21.VIII. (D.V. Znojko), 1♀ ( ZISP); “Nov. [yy] Worms [5 km NE Mostovoe Settlm .]”, “ Pervomayskiy okr.[ug] ”, 12.VI.1925 (A-y Kiritshenko), 1♀ ( ZISP); Kherson Prov.: Tendra Spit, VII.1898 (Birula), VII.1926 (Medvedev), 4♀ ( ZISP); “ Tavr.[icheskaya] g.[uberniya] Dnepr.[ovskiy] u.[ezd]”, Nikolaevka , 12.IX.1903 (coll. Kiritshenko), 1♀ ( ZISP); “ mon.[astery of] Korsun, niz. Dnepra” [ Korsunka Settlm .], 27.VI.1926, 24.V.1928 (Medvedev), 3♀ ( ZISP); Askaniya-Nova, 28.IV.1927, summer 1928, 7.VIII.1929 (Medvedev), 3J, 2♀, 28.VI.1929 (A. Wulf), 1♀ ( ZISP); Zaporizhia Prov.: Khortytsia I., 26.VII.1980 (T.G. Kokoreva), 1♀ ( ZISP); “ Altagir ” [Bogatyr’ Settlm.], steep bank of Molochnaya Riv. estuary , 26.VII.1980 (T.G. Kokoreva), 1♀ ( ZISP); Kharkiv Prov. [?]: Lozovaya, 23.VIII.1902 (M. Semen [ov]), 1♀ ( ZISP); Donets’ k Prov.: Velikiy Anadol forestry, under foliage , 15.VII.1905 [collector unknown], 1J, sweeping over grass, 24,28. VII.1905 [collector unknown], 2♀ ( ZISP); Mariupol, 28.IV.1933 [collector unknown], 1♀, 5.V.1933 (Zhenzhurist), 1J ( ZMUM); “mely gorn. sosn.”, Svyatogorsk, 10.VI.1941 (Arnoldi), 1♀ ( ZISP); Lugans’ k Prov.: Lugans’ k env., 28.V.1927 (Likhosherstov), 1♀ ( ZISP), 7.V.1928 (V. Talitskiy), 1♀ ( ZISP); Baronskaya [railway] station , 29. VII.1929 (V. Talitskiy), 1♀ ( ZISP); “ Novocherk.[asskiy] okr. [ug] ”, Proval’ e , 24.IV.1930 (V. Talitskiy), 3♀ ( ZISP); Autonomous Republic of Crimea: “Yuzhn. ber. Kryma [ south coast of Crimea ]”, [date unknown] (Widhalm), 1♀ ( ZISP); Crimea, 11–12.VI.1900 (Ageenko), 1♀ ( ZISP); Agarmysh Mts, [date unknown] (Kiritshenko), 2♀ ( ZISP); “ Bel’ bek” [Verkhnesadovaya] station of “Sevastopol ” [Cisdnieper] railways , 30.V.1897 (N. Kusnetzov), 31.V.1909 (W. Pliginski), 2♀ ( ZISP); Alma [Riv.], 26.V.1899 (A. Bazhenov), 12.V.1907 (Kiritshenko), 2J, 6♀ ( ZISP); Simferopol, 23.IV.1899 (A. Bazhenov), 8.IV.1907 (G. and K. Khristoforov), 2♀, 29. IX.1999 (I.S. Smirnov), 1J ( ZISP); valley of Salgir Riv. nr Simferopol , 23.IV., 1.VII.1899 (A. Bazhenov), 1J, 1♀ ( ZISP); Simferopol env. , 10.IX.1922 (Redikorzev), 1♀ ( ZISP); Simferopol, bank of Salgir Riv. , 14.VII.1924 (Kiritshenko), 2♀ ( ZISP); Kastel’ Mt. nr Alushta, 22.VI.1900 (N. Kusnetzov), 2♀ ( ZISP); “ Mukhalatka Vill.” [ Oliva Settlm .], 4.VII, 7–8.VIII.1900 (Ageenko), 2♀ ( ZISP); Kerch, VIII.1901, 5.VII.1917 (A. Kiritshenko), 2♀, 25.III.1908 (Kiritshenko), 7.VIII.1915 (Alexey Kiritshenko), 2J ( ZISP); Sebastopol, 4.VI.1901, 22,23.IV, 3,18,22,23. V.1905 (W. Pliginski), 10♀ ( ZISP); “Kastropol’” [Beregovoe Settlm.], 30.VI.1902 (N. Kusnetzov), 1♀ ( ZISP); “ Taushan-Bazar ” [ Privol’ noe Settlm .], 4.VI., 7. VII.1907 (Grigor’ ev), 4.VI.1907 (W. Pliginski), 9♀ ( ZISP); Ay-Petri Mt., 7.VII.1907 (Klementz), 1♀ ( ZISP); N slope of Ay-Petri Mt. , c. 900 m, 7.V.1999 (A.G. Koval), 1♀ ( ZISP); Eupatoria, VI– VII.1908 (Białynicki-Birula), 1♀ ( ZISP); Feodosia, 16.VI.1908 (Muralevich), 1J ( ZMUM); “ Ulusala ” [ Sinapnoe Settlm .], 15. VI.1911 (A.M. Djakonov), 1♀ ( ZISP); Voronskaya dolina [valley], 6.VIII.1913 (E. Golovleva), 1♀ ( ZISP); Chatyr-Dag Mts, 23.VI.1914 (W. Pliginski), 1♀ ( ZISP); lower plateau of ChatyrDag Mts, 28.VI.1984 (A.K. Zagulajev), 1♀ ( ZISP); Simeiz, 13,23. VI.1915, 12.VII.1926 (A. Kiritshenko), 3♀ ( ZISP); “ Limeny ” [ Goluboy Zaliv Settlm .], 27.VI.1915 (Kiritshenko), 1♀ ( ZISP); Kara-Dag Mt., NW slope of Karagach Mts , 4. VII.1915 (V. Vuchetich), 1♀ ( ZMUM), 6.V.1985 (A.K. Zagulajev), 1J ( ZISP); “ Albat ” [ Kuybyshevo Settlm .], 26. VI.1924 (Kiritshenko), 1♀ ( ZISP); Sudak, 1.IX.1925 (Lukjanovitsh), 1♀ ( ZISP); Ossoviny, 11.VI.1926 (Gordeev), 2♀ ( ZISP); “ Kikineiz ” [ Opolznevoe Settlm .], 29.VIII.1926, 4. IX.1927 (Kiritshenko), 2J ( ZISP); Staryy Krym , 7.V.1933 (B. Kusin), 1J ( ZMUM), 30.V., 21.X.1947 (K. Arnoldi), 4♀ ( ZISP); Roman-Kosh Mt., Babugan-Yayla Mts , 19.VI.1947 (K. Arnoldi), 1♀ ( ZISP); Mokryy Indol Settlm., 9 km N “ Saly ” [ Grushevka Settlm .], 15.V.1948 (K. Arnoldi), 1J ( ZISP); Gurzuf, 25. VII.1948 (K. Arnoldi), 1♀ ( ZISP); Nikita Botanical Garden, 20.IV.1989 (D.R. Kasparyan), 1J ( ZISP). Russia: Rostov Prov.: “ Kugeyskie [khutora] ” [ Kugeyskoe rural Settlm .], on ground, 7.IV.1917 [collector unknown], 2J, 2♀ ( ZISP); Man’ kovo-Berezovskaya sloboda, 28.IV.1925 ( Yu. Antova ), 1J ( ZISP); Tselina Settlm., 23.VII.1930 (Lubischew), 1♀ ( ZISP); Rostov-na-Donu, Solov’ inaya roshcha [grove] , 22.VI.2000 (S. Petrushchenko), 1♀ ( ZISP); Razdorskaya Vill. env., 7–10, 25–28.VII.2001 (A. Ponomarenko), 10.VII.2002 (E.A. Khachikov), 2J, 1♀ ( ZISP); khutor Pukhlyakovskiy, slopes, steppe meadow, 17.VI.–1.VII.2003 (D.A. Gapon), 13–25. IV.2004 (A.V. Ponomarenko), 3J, 1♀ ( ZISP); Krasnopartizanskiy Settlm., 27.VI.2003 (D.A. Gapon), 1J ( ZISP); Kalitvenskiy forestry, khutor Novonikolaevka env. , 16–29.VII.2008 (D.A. Gapon), 1♀ ( ZISP); Volgograd Prov., “ Sarepta ” [ Volgograd ], [date unknown] (Becker), 1♀ ( ZISP); Republic of Kalmykiya, Elista , 22.VI.1934 (Lukjanovitsh), 1♀ ( ZISP); Krasnodar Terr.: Ashe , 14,19. VI.1904 (Borodin), 2♀ ( ZISP); “ Sosnovka, 9 verst [9.6 km] N Ekaterinodar [ Krasnodar ]”, 1905 (N. Vorob’ ev), 1♀ ( ZISP); Krasnyy Les Settlm., 1910 (N. Bryanskiy), 1♀ ( ZISP); gorge with spring nr Gelendzhik, 1.VII.1910 (Vorob’ ev), 2♀ ( ZISP); Gelendzhik , [date unknown] (N. Vorob’ ev), 2♀ ( ZISP), VIII–IX.1934 (S. Nikulin), 1♀ ( ZMUM), 24,16. VI.1948 (Kryzhanovskij), 2♀ ( ZISP); Mar’ ino , 1912 (Sakhnovskiy), 1♀ ( ZISP); Krasnaya Polyana , “07.. ... .. V.” [2.V.1916] (Kiritshenko), 3J, 2♀ ( ZISP); Khosta, VIII.1927 (S. Nikulin), 1J, 3♀ ( ZMUM); Republic of Adygea, Maykop, Kurdzhips Riv. , 17,15. V.1933 (Shaposhnikov), 2♀ ( ZMUM); Stavropol Terr.: “ Stavr. L Fig ” [Stavropol?], [date unknown] (coll. V. Jakovlev), 1♀ ( ZISP); Stavropol, [date unknown] (Bryanskiy), 1♀ ( ZISP), 19–23.IV, 20.VI.1914 (B. Uvarov), 1♀ ( ZISP); Pyatigorsk, [date unknown] (Pr. E. Pavlov), 2♀ ( ZISP); Mineral’ nye Vody, VI–VII.1894 (N. P. Odintsov), 1♀ ( ZISP); N and S slopes of Mashuk Mt. nr Pyatigorsk, 15.IV.1908 (Skorikov), 2♀ ( ZISP); Lokhmatyy kurgan [ Lokhmataya Mt. ], W peak of Beshtau Mt., 31.V.1908 (Gerasimov), 1♀ ( ZISP); Byk Mt. nr Zheleznovodsk, 20. VI.1908 (A.P. Gerasimov), 1♀ ( ZISP); Krasnogvardeyskiy Distr., 7 km NE Ladovskaya Balka Vill ., 10.VI.1982 (B.A. Korotyaev), 1J, 2♀ ( ZISP); Republic of Dagestan: “Khodzhal-makhi” [Levashinskiy Distr., Khadzhalmakhi ], 17.VI.1944 (Rjabov), 1♀ ( ZISP); 30 km NW Rutul, 25.VII.1983 (I.M. Kerzhner), 1♀ ( ZISP). Georgia: Autonomous Republic of Abkhazia: Novyy Afon , 12.VIII.1898 (O. Birula), 1♀ ( ZISP); ravine of Bagade [nr Kodori Valley ?], 28.VII.1932 (Borchsenius), 1J ( ZISP); Bzyb Range, left bank of Bzyb Riv. , 6 km upstream of confluence with Gega Riv., 2.VII.2001 (Yu.G. Arzanov), 1♀ ( ZISP); Samtskhe-Javakheti: Sadgeri , 29.VI.1897 [collector unknown], 1♀ ( ZISP); Benara 19 km W Akhaltsikhe , 23.VI.1949 (Kiritshenko), 4♀ ( ZISP); Tbilisi, “ behind German colony of Alexanderdorf near Tiflis [ Tbilisi ]”, 23.IV.1990 (Satunin), 1♀ ( ZISP); Kakheti, Vashlovani [Nat. Park] , 13. V.1975, 25.V.1977 (O.L. Kryzhanovskij), 1J; 5♀ ( ZISP). Armenia: Lori Prov., “ Kirovakan ” [ Vanadzor ], 22.V.1958 (V. Zaitzev), 2♀ ( ZISP); Aragatsotn Prov., Burakan, 13.VII.1946 (Rikhter, Ter-Minassjan), 1♀ ( ZISP); Kotayk Prov.: “ Vokhchaberd ” [ Voghjaberd Vill .], 1938 (Rikhter, Ter-Minassjan), 2♀ ( ZISP); Dzhervezh Settlm. O Yerevan, 12. V.1938 (A. Rikhter), 1J [paratype of C. mixtus ], “ № 201–938”, 23.V.1938 [collector unknown], 1♀, 14.VII.1952 (Ter-Minassjan), 1♀, 12.IV., 13.V.1958 (V. Zaitzev), 17♀, 4, 9. V.1960 (G.M. Dlussky), 2♀ ( ZISP), 20–24, 25.VI.1960 (G. Victorov), 2J ( ZMUM); ravine of Dzhervezh, 1380 m, 9. V.1955 (Loginova), 1♀ ( ZISP); Monastery of Geghard, 6. V.1947 (Rikhter, Ter-Minassjan), 1♀ ( ZISP); Yerevan: “ № 201– 392 ”, 22.V.1938 [collector unknown], 1936 (S.A. Chernov), “zoo”, 12.V.1958 (V. Zaitzev), 3♀ ( ZISP); Ararat Prov.: Vedi Riv., upstream of “Chimankend” [ Urtsadzor Vill .], 1947 (Rikhter, Ter-Minassjan), 1♀ ( ZISP); “ Karabakhlar ” [Nerkin Karabakhlar = Urtsadzor], “ Khosrov Forest” [State Reserve ], 13.V.1955 (M. Loginova), 1♀, 31.V.1972 (E. Sugonyaev), 1♀ ( ZISP); Tigranashen env., N39°48.366 ʹ E44°58.297 ʹ, 1685 m, 19. VII.2014 (M.G. Volkovitsh), 1J ( ZISP); Vayots Dzor Prov.: “r. Arpa-chay (vost.) [Arpa Riv.] bl. [nr] Pashal. [? Pashalu = Zaritap ] mosta [bridge]”, 2–6.VII.1937 (Rjabov), 1♀ ( ZISP); Gerger [ Herher ], “Azizb.”, 15.VI.1957 (L. Zimina), 1♀ ( ZMUM); gorge of Arpa Riv. nr Vernashen Settlm., 4.VI.1958 (V. Zaitzev), 1♀ ( ZISP); Syunik Prov.: Meghri env., 5.VI.1982 (A.V. Gorochov), 2J, 2♀ ( ZISP); Meghri on Arax Riv., “ № 201– 392”, 2.V.1938 [collector unknown], 1♀ ( ZISP). Azerbaijan: Qazakh Distr., Alpoud Settlm., “Gyugazan [ Göjäzän ] Mt.”, 11. V.1960 (G.M. Dlussky), 1♀ ( ZISP); Ganja, 20.VII.1933 (Lukjanovitsh), 1J, 1♀ ( ZISP); Shusha, 12.VI.1933 (Lukjanovitsh), 1♀ ( ZISP); Lerik Distr., Govdara Vill. [17 km NNE Lerik town ], 9.VII.1909 (Kiritshenko), 2J, 1♀ ( ZISP); Lenkoran Distr., “ mont. Talysh Savdara Kreis Lenkoran ”, 9. VI.1909 [collector unknown], 3♀ ( ZISP); Nakhchivan Autonomous Republic: Babek Distr., nr [Yuxarı] Buzgov Settlm., Daralagezskiy Range , 35 km N Nakhchivan, 20. VI.1985 (M.G. Volkovitsh), 1J ( ZISP); Shahbuz Distr., “Tyurkesh” [ Türkeş ], 18–26.VII.1937 (Rjabov), 1♀ ( ZISP); Ordubad Distr.: Paraga NW of Ordubad, 25.VI., 28.VII.1933 (Znojko), 2♀ ( ZISP) [paratypes of C. mixtus ]; ChananabParaga, 28.V.1957 (Trjapitzin), 1♀ ( ZISP); 11 km E Ordubad, spring with garden and desert with slope, 29.IV.1972 (D.R. Kasparyan), 1♀ ( ZISP); valley of Araxes nr Ordubad, 27. VI.1974 (M.G. Volkovitsh), 1J ( ZISP); Nukus nr Ordubad, 3. IV.1976 (M.G. Volkovitsh), 1J ( ZISP); “Akulis” [ Ashaghy Aylis Vill .] nr Ordubad, 14.V.1976 (V.A. Rikhter), 1♀; 4 km S Bilav, old sparse Salix & “Polyurus” [Paliurus] forest along river, 1200 m, in litter & logs, 23.VI.1983 (S. Golovatch), 1J ( ZISP). Turkey: Mersin Prov., Gülek, “ Taur.[us Mts] Cilic. [ian Gates ]”, 1897 (Holtz), 1J ( ZISP); Tunceli Prov., nr Pülümür, 26–27.V.2010 (E.A. Khachikov), 1♀ ( ZISP); Van Prov., shore of Lake Van near Akdamar I. , 9.VI.2009 (D. Gapon), 1♀ ( ZISP); Bitlis Prov., Van-Bitlis Yolu nr Tosunlu N38°27.245′ E042°32.494′ 1785 m, 9.VI.2009 (D.A. Gapon), 1♀ ( ZISP); Bitlis Muş Yolu N38°33.187′ E042°05.851′ 1570 m, 10.VI.2009 (D.A. Gapon), 1J ( ZISP). Syria: “ Syria ”, Homs, 15.V.1988 (M. Garib), 1J ( ZISP). Israel: Southern Distr.: “Bet Gaddi” [ Beit HaGadi ], 4–6.V.1966 (Trjapitzin), 2♀ ( ZISP). Kazakhstan: West Kazakhstan Prov.: Dzhanibek, 28.IV.1953 (K. Arnoldi), 1♀ ( ZISP); Atyrau Prov.: “Gur’ evskiy uezd Ural’ skaya oblast’” Dossor, 25.VII.1927 (Denisova), 1♀ ( ZISP).

Diagnosis

Membrane brownish, opaque ( Figure 1 C ); only in one examined specimen from South Anatolia membrane slightly smoky, almost white and transparent. Yellowish rib on lateral margins of pronotum and corium broader than base of antennal segment II. Vesica with long membranous lateral branch except North African and Syrian specimens having short brunch. Spicule basal plate on dorsolateral lobes of conjunctiva with narrow distal denticle ( Figure 7 C, 7 D ). Posterior pouches of gynatrial sac approximately as long as wide ( Figures 8 B , 12 B ), sometimes slightly shortened, but rather long posteriorly (beyond dorsal inner margin). Sclerites inside pouches usually symmetrical, cone- or cup-shaped, somewhat varying in size ( Figures 8 B , 12 B , 13 A ); in specimens from North Africa and Southern Europe, sometimes in specimens from south of Eastern Europe and Caucasus, they differentiated into anterior and posterior parts of different shapes (see below), often sharply asymmetrical or represented only by anterior lamellate part lying parallel and close to anterior inner wall of pouch ( Figures 13 B – 13 J , 14 A –14 G ). Tubular part of spermathecal capsule 2–3 times as long as transverse diameter of spherical part ( Figure 9 C ).

Measurements

Body 5.10–8.35 in length, 3.10–4.90 in width; pronotum 2.65–4.35 in width; head 1.29–1.81 in width; ocular index 2.69–3.68; length of antennal segments I–V: 0.27–0.41: 0.45–0.66: 0.52–0.87: 0.78–1.06: 0.84–1.05.

Geographic variability

The nominotypical subspecies evinces polymorphism in the structure of the genitalia of males and especially females, namely, in relative length of lateral branch of vesica, shape of spicula at dorsolateral lobes of conjunctiva, and shape and size of sclerites in posterior pouches of gynatrial sac. The distribution of forms described below is shown in Figure 16 .

All males examined can be divided into two forms according to the relative length of the lateral branch of vesica.

In males from North Africa [Morocco, Oudjda, 1930, coll. Sicard, 3 males. Algeria, Aurés, Djebel Chelia, 22. VI.1981, Doguet S., 1 male. Tunisia: Sidi el Hani, V.1884, 1 male; Al Hushayshinah, coll. Hannothiaux, 1 male. Egypt: “Egypte”, “66–96”, Letourneux, 1 male] and male from Syria, lateral branch of vesica distinctly shortened ( Figure 4 G ). Lateral branch in remaining males distinctly long, 2.6–3.4 times as long as vesica [the vesici in the specimens from Italy were not examined since their aedeagi was prepared and severely damaged by the previous researcher, but these specimens have the usual, not narrowed spicules at dorsolateral lobes (vide infra), and relying on this it can be assumed that the Italian specimens also had the long lateral branch of vesica]. In males from North Africa, spicule at dorsolateral lobes of conjunctiva uniformly narrow or very slightly widened subapically, with almost straight outer margin before apex ( Figure 7 D ); in all males, except for three specimens from Syria, Egypt and Morocco, spicule less strongly curved than in other specimens examined. In specimens from remaining part of natural habitat of the species, spicule rather strongly widened subapically and curved, with distinctly convex outer margin before apex.

The females examined can be divided into six forms by the shape of sclerites in the posterior pouches of the gynatrial sac.

Form 1. In specimens from Ukraine, Russia, Transcaucasia, Turkey and Israel, sclerites of conical or cotyloid shape, not differentiated into parts, with broad margin faced anteriad in dorsal ( Figure 8 B ) or postero-inner ( Figure 12 B ) view. Usually these sclerites large, occupying almost entire cavity of pouch, but sometimes decreased in varying degree (e.g. as in Figure 13 A ) without obvious geographical regularity. Two specimens from Kazakhstan, [Dzhanibek, 28.IV.1953, K. Arnoldi] and Russia [Stavropol, Bryanskiy] possessing unusual short, widely cotyloid sclerites.

Form 2. Both sclerites or only right sclerite divided into two parts. Anterior part subconical or cotyloid; posterior part flask-shaped, conical, cotyloid or of different shape and connected or closely adjacent to anterior part.

In female presumably from Slovenia [“91./ Cydnus dubius Carniol ”], sclerites symmetrical ( Figure 13 B ); anterior part of each sclerite shaped as broad and rather long cone; posterior part resembling a flask with narrow neck directed anteriad and rather small dilated part rounded in dorsal view and having anterior inner wall elongated into triangular process directed ventrally in postero-inner view.

In female from France [Nice, G.V. Olsufjev], pouches rather strongly elongated ( Figure 13 C ). Anterior parts of sclerites cotyloid, wide and short; posterior part within right pouch flask-shaped, large, approximately as long as wide, with triangular projection along ventral inner wall of pouch and with neck connected with anterior part of sclerite; within left pouch, posterior part of sclerite absent.

In specimen from Dagestan [30 km NW Rutul, 25. VII.1983, I.M. Kerzhner] ( Figure 13 D ), anterior parts of sclerites large and conical; posterior part of right sclerite small, conical, directed posteriad by its apex and connected with anterior part of sclerite by narrow short ligament; posterior part of left sclerite shaped as plate lying along inner margin of anterior part of sclerite, connected with anterior inner angle of latter, thickened at posterior end connected with convex wall of anterior part of sclerite by short ligament and bearing process directed ventromediad.

In specimen from Stavropol Territory of Russia [N and S slopes of Mashuk Mt., Pyatigorsk, 15.IV.1908, Skorikov], sclerites similar to those of previous specimen, with difference that anterior parts of sclerites somewhat narrowed posteriorly; posterior part of right sclerite with directed dorsad lamellar projection on outer margin from dorsal side; left sclerite without posterior part.

In specimen from Ukraine, Crimea, [Alma Riv., 26. V.1899, A. Bazhenov] ( Figure 13 E ), sclerites similar to those of two previous specimens: anterior parts of sclerites subconical; posterior part of right sclerite large, broadly conical, with apex directed posteriad, anteriorly connected by weakly sclerotised ligament with anterior part of sclerite; left pouch without posterior part of sclerite, with small weakly sclerotised patch on inner wall near pouch apex. Another specimen from Ukraine [Mariupol, 28.IV.1933, collector unknown] with anterior parts of sclerites as in previous specimen and with symmetrical, large, broadly conical posterior parts adjacent to anterior parts.

In specimen from Armenia [ravine of Dzhervezh, 9. V.1955, Loginova] ( Figure 13 F ), anterior parts of sclerites cotyloid, short and broad; posterior part of right sclerite shaped as spacious conical sclerotised area occupying entire pouch space in apical third and widely adjacent to anterior part of sclerite; posterior part of left sclerite transverse, with widely rounded posterior margin, closely adjacent to anterior part of sclerite in inner part, with short process anterior to narrowed inner end. Sclerite in left pouch of another specimen from Armenia [Geghard, 6. V.1947, Rikhter, Ter-Minassjan] similar to right sclerite of previous specimen, with large conical posterior part closely adjacent to cotyloid anterior part; posterior part of right sclerite absent.

In one more specimen from Armenia [Dzhrvezh O of Yerevan, “No. 201–938”, 23.V.1938, collector unknown] ( Figure 13 G ), anterior parts of sclerites large, subconical; posterior part of right sclerite large, hemispherical, closely adjacent to anterior part of sclerite, with acute-angular process directed anteroventrad; posterior part of left sclerite in the form of rather narrow, hook-shaped process connected with anterior part of sclerite and directed posteroventrad.

Form 3. Anterior parts of sclerites in dorsal view shaped as very thin, weakly sclerotised plates perpendicular to sclerite longitudinal axis and lying parallel to anterior inner wall of pouch; posterior parts of sclerites flask-shaped, slightly larger and more elongated than in specimen of previous two forms.

In female from Morocco [nr Aourir, 6,7. VI.2015, D.A. Gapon] ( Figure 13 H ), sclerites symmetrical, anterior lamellar parts small, posterior parts of sclerites without processes. In specimen from Egypt [“Egypt”, coll. Eversmann] ( Figure 13 I ), anterior lamellar parts of sclerites larger, necks of flask-shaped posterior parts wider than in specimen from Morocco; neck of left sclerite just behind anterior lamellar part with process directed towards inner corner of pouch, widened part of left sclerite slightly smaller than the same of right sclerite. In female from Algeria [“Algérie”, 1860, Coll. A.F. Morawitz] ( Figure 13 J ), both sclerites slightly sclerotised, with rather large lamellar anterior parts and asymmetrical posterior parts; right sclerite flask-shaped narrower posteriorly than in previous specimens, with very narrow neck and without process; left sclerite shaped as narrow straight band strongly narrowing anteriorly.

Form 4. Anterior parts of sclerites as in form 3; posterior parts strongly reduced or absent.

In female from Tunisia [Sfax and environs, III.1922, G. Babault] ( Figure 14 A ), anterior parts of sclerites in the form of well visible large plates; posterior part of right sclerite shaped as short narrow band strongly narrowing anteriorly and connecting with anterior part of sclerite; posterior part of left sclerite small, transversely rounded and separated from anterior part. In another specimen from Tunisia [Sidi el Hani, V.1884] ( Figure 14 B ), small area of very weak and poorly visible sclerotisation lying behind well visible, rather large anterior lamellar part of left sclerite; right sclerite without posterior part. Pouches in both females from Tunisia shortened. In female from Libya [“Env. De Tripoli”, 1877, Mouchez] ( Figure 14 C ), anterior parts of sclerites represented by rather large plates, well visible in dorsal view; posterior parts of sclerites absent.

Form 5 combines characters of the forms 1, 3 and 4. Sclerites significantly asymmetrical, with large, more or less wide posterior parts and thin, more or less developed plates corresponding to anterior parts of sclerites, closely adjoining to wide anterior margins of posterior parts and lying parallel to anterior inner wall of pouch.

In female from Algeria [Sidi-bel-Abbes] ( Figure 14 D ), posterior part of left sclerite large, transverse, with rounded posterior margin; posterior part of right sclerite diminished, subtriangular; anterior platelike parts of sclerites almost not developed. Female from Tunisia [10.VI.1913, N.L. Pastukhov] ( Figure 14 E ) with posterior part of left sclerite of the same shape as in previous specimen, but smaller; posterior part of right sclerite as long as wide, greatly diminished, with rounded posterior margin; anterior platelike parts of sclerites quite distinct, especially in right pouch. In another female from Tunisia [Al Hushayshinah, coll. Hannothiaux] ( Figure 14 F ), posterior parts of sclerites elongated, rather weakly sclerotised, and asymmetrical, wide in left pouch and narrow in right one; anterior platelike parts of sclerites distinct.

Form 6. In one female from Morocco [Oudjda, 1930, coll. Sicard], two from Algeria [Aurés, Djebel Chelia, 22. VI.1981, Doguet S.], two from Egypt [“Egypte”, “66–96”, Letourneux; and “Aegypt.”] ( Figure 14 G ), and one female from Andalusia [Almeria, 2.V.1952, H. Coiffait], sclerite within each pouch shaped as rather small, almost transparent plate with poorly appreciable dorsal rib looking as very thin strip in dorsal view; in view from anterior inner wall of pouch, contours and weak sclerotisation of plate barely appreciable.

For the time being, it is not possible to ascertain the taxonomic significance of the listed characters and the taxonomic status of the described forms due to a scarcity of the examined material from Southern and Western Europe.

Notes on synonymy

Canthophorus melanopterus niger ( Vidal, 1950) was erected as a variety Sehirus dubius var. nigra distinguished by completely black body dorsum, some points of which in the transverse depression of the pronotum sometimes with blue or purplish luster, and by the rather deep (“assez accentuée”) transverse depression of the pronotum. The described by two specimens collected at Mount Bou Iblan in the Middle Atlas at an altitude of 2500–2800 m. Vidal left his type specimens in Rabat, and for a long time the types of this author were not available for examination. They remain such now and probably should be considered lost. Wagner (1956) distinguished this variety by the colour and the deep transverse depression on pronotum and noted that he found only a few specimens with the black dorsum among more than 1000 specimens examined by him. Subsequently, this raised to the subspecific status by Stichel (1961).

I did not find any significant differences in the colour and relief of the pronotum between examined North African specimens of C. melanopterus from the collections of MNHN and ZISP (including specimens collected at an altitude of 1800–2000 m in Algeria) and specimens from other parts of the species natural habitat. The darkest specimens were two from Egypt, but they have the metallic luster of the entire body dorsum. And in general, the darkest specimens of C. melanopterus , which I saw and which mostly correspond to the diagnosis of Vidal’ s variety, originate from the Crimea [Mokryy Indol Settlm., 15.V.1948 (Arnoldi), 1J; Staryy Krym, 30.V.,1947 (Arnoldi), 1♀; Simferopol, 14.VII.1924 (Kiritshenko), 1♀. These specimens were labeled in the ZISP collection as], and the depth of the transverse depression on the pronotum is slightly variable irrelatively of the distribution of individuals. In view of these reasons, I consider the name Canthophorus melanopterus niger pertaining to the rare colour (and perhaps, highmountainous) place this name in synonymy with Canthophorus melanopterus melanopterus .

Distribution

North Africa, Southern Europe, France, Austria, Eastern Europe (in Russia, only the south of the European part), Transcaucasia, Turkey, Syria, Iraq, Iran, Israel, Jordan and European part of Kazakhstan ( Lis 2006). The record of the species for North-East China, as mentioned by Lis (2006) with a question mark, probably should be recognised as erroneous.

Figures

Figure 1. Hemelytral membranes in Canthophorus spp. A, C. dubius dubius (Scopoli, 1763), France: Roquefort-les-Pins; B, C. dubius sanigarum n. ssp., Russia: Krasnaya Polyana, paratype; C, C. melanopterus melanopterus (Herrich-Schaeffer, 1835), Ukraine: Ascania Nova; D, C. melanopterus contrarius (Wagner, 1956), Cyprus: Château de Buffavento; E, C. impressus impressus (Horváth, 1880), Russia: Khodzhal-Makhi; F, C. impressus hajastanicus n. ssp., Georgia: gorge of Paravanichet Riv., paratype. Abdomens removed to show membrane colour in transparency.

Figure 3. Right parameres in Canthophorus and Adomerus spp. A, B, C. dubius dubius (Scopoli, 1763), France: Seine-et-Marne; C, D, C. dubius sanigarum n. ssp., Russia: Krasnaya Polyana, paratype; E, C. melanopterus (Herrich-Schaeffer, 1835), Russia: Krasnaya Polyana; F, C. impressus impressus (Horváth, 1880), Ukraine: Demir-kapu; G, C. impressus hajastanicus n. ssp., Armenia: Gekhmakhan Mt., paratype; H, C. wagneri Asanova, 1964, Russia: Makhachkala; I, C. mixtus mixtus Asanova, 1964, Armenia: Kipchakh; J, C. mixtus mixtus Asanova, 1964, Armenia: Gyumri; K, C. mixtus hissaricus Asanova, 1964 n. stat., Tajikistan: urochishche Ruydasht, paratype; L, C. niveimarginatus Scott, 1874, Russia: Kyzyl; M, C. coeruleus (Reuter, 1902), Kyrgyzstan: urochishche Ak-terek; N, O, A. aeneus (Walker, 1867) n. comb.; P, Q, A. maculipes (Mulsant & Rey, 1852) n. comb. Outer-ventral view (A, C, E, F–M), inner view (N, P), and dorsal view (B, D, O, Q). Scale bar: 0.25 mm.

Figure 4. Aedeagi and vesici in Canthophorus spp. A, B, C. dubius dubius (Scopoli, 1763), Russia: Krasnaya Polyana, paratype; C–G, C. melanopterus melanopterus (Herrich-Schaeffer, 1835), Russia: Pukhlyakovskiy (C, D), Ukraine: Opolznevoe (E), Greece: Lake Kremasta (F), Tunisia: Sidi el Hani (G); H, C. melanopterus contrarius (Wagner, 1956). Aedeagus in lateral view (A) and in dorsal view (B, C); vesica in dorsoposterior view (D) and in lateral view (E–H). a.l – apical lobes of conjunctiva; dl.l – dorsolateral lobes of conjunctiva; l.b – lateral branch of vesica; sp – spicule of dorsolateral lobes; v.l – ventral lobe of conjunctiva; vs – vesica. Scale bares: 0.25 mm.

Figure 6. Ventral lobes of conjunctiva in Canthophorus spp. A–C, C. dubius dubius (Scopoli, 1763), France: Forêt des Morières; D–I, C. dubius sanigarum n. ssp., Russia: Krasnaya Polyana, paratype (D–F), Russia: Tuapse (G–I); J–M, C. melanopterus (Herrich- Schaeffer, 1835), Ukraine: Yalta (J, K), Armenia: Tigranashen (L, M); N, O, C. impressus (Horváth, 1880), Ukraine: Velikiy Anadol; P, Q, C. wagneri wagneri Asanova, 1964, Russia: Akhty; R, S, C. wagneri hyrcanicus n. ssp., Turkmenistan: gorge Khozly; T, U, C. mixtus Asanova, 1964, Kazakhstan: Dzhanibek; V, W, C. niveimarginatus Scott, 1874, Russia: Kyzyl; X, Y, C. coeruleus (Reuter, 1902), Kyrgyzstan: urochishche Ak-terek. Ventral view (A, D, G, J, L, N, P, R, T, V, X), lateral view (B, E, H, K, M, O, Q, S, U, W, Y), and anterior view (C, F, I). Scale bar: 0.25 mm.

Figure 7. Spicules of conjunctival dorsolateral lobes in Canthophorus spp.A, C. dubius dubius (Scopoli, 1763), France:Forêt des Morières; B, C. dubius sanigarum n. ssp., Russia: Krasnaya Polyana, paratype; C, D, C. melanopterus melanopterus (Herrich-Schaeffer, 1835), Ukraine: Opolznevoe (C), Tunisia, Sidi el Hani (D); E, C. melanopterus contrarius (Wagner, 1956); F, C.melanopterus mariae n. ssp., Tajikistan:Kondara Gorge, paratype; G, C. impressus impressus (Horváth, 1880), Ukraine:Demir-kapu; H, C. impressus hajastanicus n. ssp., Armenia:Gekhmakhan Mt., paratype; I, C. wagneri wagneri Asanova, 1964, Russia: Il’ Mt. SE Vladikavkaz; J, C. wagneri hyrcanicus n. ssp., Turkmenistan: gorge Khozly, paratype; K–Q, C. mixtus mixtus Asanova, 1964, Kazakhstan: Chagan-Aba (K), Kazakhstan: Yanvartsevo (L); Armenia: Kipchakh (M), Kazakhstan:Khanskaya stavka(N),Tajikistan:KhozretishiRange[Gurjeva] (O,P), Kazakhstan:Trans-Ili Alatau,gorge Levyy Talgar(Q);R,S, C. mixtus hissaricus Asanova, 1964 n. stat., Tajikistan: urochishche Ruydasht, paratype (R), Tajikistan: Khozretishi Range, 75 km NE Muminabad (S); T, U, C. niveimarginatus Scott, 1874, Russia: Kyzyl (T), Mongolia: Stariy Tsakhir (U); V, C. coeruleus (Reuter, 1902), Kyrgyzstan: urochishche Ak-terek. Scale bar: 0.25 mm.

Figure 8. Female terminalia in Canthophorus spp. A–G, C. melanopterus (Herrich-Schaeffer, 1835), Ukraine: Alma Riv. (A), Azerbaijan: Ganja (B), Ukraine: Kamenets-Podol’skiy (C), Armenia: Dzhervezh (D), Ukraine: Khadzhibeyskiy liman (E), Morocco: Aourir (F); Egypt: “Aegipt.” (G); H, C. dubius dubius (Scopoli, 1763), France: Forêt des Morières; I, C. dubius sanigarum n. ssp., Russia: Krasnaya Polyana, paratype; J, K, C. impressus impressus (Horváth, 1880), Ukraine: “yayla Besh-tekme” (J), Russia: Beshtau Mt. (K); L, C. impressus hajastanicus n. ssp., Armenia: Gekhmakhan Mt., paratype; M, N, C. wagneri wagneri Asanova, 1964, Russia: Anapa (M), Armenia: Sevan Lake (N); O–S, C. mixtus mixtus Asanova, 1964, Ukraine: Kremenets Mt. nr Izyum (O), Armenia: Spitak (P), Ukraine: Aleshki, paratype (Q), Ukraine: Nyrkovo – Vyemka, paratype (R), Azerbaijan: Kars (S); T, U, C. mixtus hissaricus Asanova, 1964; n. stat., Tajikistan: urochishche Ruydasht [paratypes]; V, W, C. niveimarginatus Scott, 1874; Russia: Kosh-Agach (V), Russia: Kamen’-Rybolov (W); X, Y, C. coeruleus (Reuter, 1902), Kyrgyzstan: urochishche Ak-terek (X), Kyrgyzstan: Berk-Suu Riv (Y). External terminalia (genital plates) and vestibulum of gynatrium, ventral view (A); gynatrial sac, dorsal view (B); shape of right first ramus (C–Y). a.g – anterior part of gynatrial sac; d.d – distal part of spermathecal duct; f.r – first rami; g.c – gynatrial cone; gn – gonangulum; gp.I – gonapophyses I; gp.II – gonapophyses II; gx.II – gonocoxites II; m.d – median part of spermathecal duct; p.d – proximal part of spermathecal duct; p.g – posterior part of gynatrial sac; p.m – posterior margin of gynatrial sac; p.p – posterior pouches of gynatrial sac; r.s – ring sclerites; tr – triangulum. Scale bars: 0.25 mm.

Figure 9. Female terminalia in Canthophorus spp. A, B, C. dubius (Scopoli, 1763), Russia: Krasnaya Polyana (A), Italy: Lecco (B); C, C. melanopterus melanopterus (Herrich- Schaeffer, 1835), Azerbaijan: Ganja; D, C. melanopterus mariae n. ssp., Tajikistan: Kondara Gorge, paratype. Gynatrial sac, dorsal view (B), capsule of spermatheca (C, D). Scale bars: 0.25 mm.

Figure 12. Posterior pouches of gynatrial sac in Canthophorus spp., posterio-inner view. A, C. dubius (Scopoli, 1763), Italy: Lecco; B, C. melanopterus (Herrich-Schaeffer, 1835), Ukraine: Alma Riv.; C, C. impressus impressus (Horváth, 1880), Ukraine: “yayla Besh-tekme”; D, C. impressus hajastanicus n. ssp., Armenia: Gekhmakhan Mt., paratype; E, C. wagneri Asanova, 1964, Russia: Anapa (some diagnostic characters is indicated by arrows); F, C. mixtus Asanova, 1964, Ukraine: Aleshki, paratype (pouch is shown straightened). Scale bar: 0.25 mm.

Figure 13. Posterior part of gynatrial sac in Canthophorus melanopterus melanopterus (Herrich-Schaeffer, 1835), dorsal view. A, Ukraine: Kamenets-Podol’skiy; B, Slovenia, Krajina; C, France: Nice; D, Russia: 30 km NW Rutul; E, Ukraine: Alma Riv.; F, Armenia: ravine of Dzhervezh; G, Armenia: Dzhrvezh O Yerevan; H, Morocco: Aourir; I, “Egypt” [coll. Eversmann]; J, Algérie [coll. A.F Morawitz]. Scale bar: 0.25 mm.

Figure 14. Posterior part of gynatrial sac in Canthophorus melanopterus (Herrich-Schaeffer, 1835), dorsal view. A, Tunisia: Sfax; B, Tunisia: Sidi el Hani; C, Libya: Tripoli; D, Algeria: Sidi-bel-Abbes; E, Tunisia [N.L. Pastukhov]; F, Tunisia: Al Hushayshinah; G, Egypt: “Aegypt.”; H, C. melanopterus contrarius (Wagner, 1956); I, C. melanopterus mariae n. ssp. [holotype]. Scale bar: 0.25 mm.

Figure 16. Geographical variability of male and female terminalia in Canthophorus melanopterus (Herrich-Schaeffer, 1835). 1–6 in legend correspond to numbers of forms 1–6 by female terminalia in C. m. melanopterus; 7, females of C. m. mariae n. ssp.; 8, males with long lateral branch of vesica; 9, males with short lateral branch of vesica.