Cicindela (s. str.) coerulea nitida Lichtenstein, 1796, 1773

Matalin, Andrey V., 2025, An intraspecific structure of Сicindela (s. str.) coerulea Pallas, 1773 (Coleoptera, Cicindelidae), Zootaxa 5583 (2), pp. 201-246 : 212-223

Cicindela (s. str.) coerulea nitida Lichtenstein, 1796

Figs 10–13 , 19–23 , 45–54 , 76–82 , 89–132

Cicindela nitida Lichtenstein, 1796: 32 (Type locality—Sibiria).

= Cicindela tricolor M.F. Adams, 1817: 278 (Type locality—Sibiria ad lacum Baikal et ad ripas fl. Lenae, ultra urbem Jakutzk), synonymized by Horn (1915: 337).

= Cicindela tricolor var. obliquefasciata Fischer von Waldheim, 1828: 21 , Tab. 1 fig. 3 (Type locality—Sibiriae), synonymized by Roeschke (in Horn & Roeschke 1891: 49).

= Cicindela tricolor var. tenuifascia Fischer von Waldheim, 1828: 22 , Tab. 1 fig. 6 (Type locality— Altai), synonymized by Roeschke (in Horn & Roeschke 1891: 49).

= Cicindela optata Fischer von Waldheim, 1828: 22 , Tab. 1 fig. 8 (Type locality—Dauria, ad litora fluvii Amur), synonymized by Motschoulski (1844: 30).

= Cicindela tricolor var. saltatoria Gistl, 1837: 38 (Type locality—Sibiria ad lacum Baikal et ad ripas fl. Lenae, ultra urbem Jakutzk, et in montosis Altaï), synonymized by Horn (1915: 337).

= Cicindela coerulea selengensis Mandl, 1936: 12 (Type locality—Selengatal, Selengafe), synonymized by Schilder (1953: 564).

Cicindela tricolor M.F. Adams, 1817 : Fischer von Waldheim 1820: 6, Tab. 1 fig. 3, Tab. 17 fig. 3; 1821: 102; 1828: 12, 21; 1829: 5; Dejean 1833: 3; 1836: 3; Dejean & Boisduval 1837: 24 Pl. 3 fig. 3; Sturm 1843: 2; Motschoulski, 1844: 30; Motschulsky, 1850: 1; Chaudoir 1865: 36; Sahlberg 1880: 5; Fleutiaux 1892: 106; Horn & Roeschke 1891: 48, Taf. V fig. 11; Tschitschérine, 1903: 94; Jakobson 1905: Tab. 1 fig. 3; Horn 1905a: 157, 158; 1905b: 27; Poppius 1906: 14; Lafer 1978: 5, figs 5, 14; 1989: 96, fig. 52.4, 97.

Cicindela hybrida tricolor M.F. Adams, 1817 : Jakobson 1905: 190; Horn 1905a: 157, 158.

Cicindela optata Fischer von Waldheim, 1828 : Motschulsky, 1850: 1.

Cicindela nitida Lichtenstein, 1796 : Lutshnik 1916: 95; Mandl 1921: 79; Shilenkov 1994: 8; Kryzhanovskij et al. 1995: 26; Berlov & Berlov 1997: 44; Sundukov 1998: 9; Matalin 2000: 676, fig. 35; Rogatnykh 2006: 30; Rogatnykh & Yakubovich 2009: 108.

Cicindela hybrida nitida Lichtenstein, 1796 : Mandl 1921: 80, 81; Winkler 1924: 4; Horn 1915: 337; 1926: 221; 1930: 402; Wu 1937: 15; Institute of Zoology Academia Sinica 1986: 417, fig. 11-1-3; Hu 1992: 374; Zhu et al. 1999: 26, 27 Tab. 4 fig. 13; Fang & Wu 2001: 69.

Cicindela grumi nitida Lichtenstein, 1796 : Schilder 1953: 564.

Cicindela nitida tricolor M.F. Adams, 1817 : Shilenkov 1987: 7.

Cicindela nitida optata Fischer von Waldheim, 1828 : Berlov & Berlov 1997: 44.

Cicindela nitida selengensis Mandl, 1936 : Kryzhanovskij et al. 1995: 27; Matalin 2000: 676, fig. 34.

Cicindela coerulea Pallas, 1773 : Rogatnykh 2009a: 9; 2009b: 69; Sundukov 2009: 89; Khobrakova 2012: 167; Putchkov 2013: 54 fig. 2.4, 62, 63, 65 fig. 4.1, 68 fig. 7.4, 73 fig. 12.5–6, 80.

Cicindela coerulea nitida Lichtenstein, 1796 : Lutshnik 1924: 21; Okninskyi 1930: 20; Mandl, 1935: 285 fig 1b, 294 figs 1–5, 296 figs 1–5; 1936: 11; 1965: 241; 1974: 137; Horn 1938: 46, Taf. 63 figs 23–25; Nakane 1960: 30, 38 Pl. I fig. d; Mandl 1970: 73 Abb. 3 fig. 6, 74; Alekseeva 1977: 5; Cassola & van Nidek 1984: 10; Werner, 1991: 18, T. 14 figs 102, 104–106, T. 15 figs 107–109; Li 1992: 5; Wiesner 1992: 117; 2020: 183; Putchkov & Shilenkov 1996: 451; Lorenz 1998: 42; 2005: 131; Puchkov & Matalin 2003: 104; Khobrakova & Sharova 2004: 144; Shook & Wiesner 2006: 10; Khobrakova & Dudko 2010: 37; Wu & Shook 2010: 78; Wu 2011: 24; Budilov 2012: 40; Sundukov 2013: 76; Khobrakova et al. 2014: 70; Koshkin et al. 2016: 312; Matalin & Dudko 2016a: 140; 2016b: 143; Tsuji et al. 2016: 3; Putchkov & Matalin 2017: 227; Matalin et al. 2019: 10; National Species List of Korea 2019: 199; Makarov et al. 2020; Putchkov 2020: 169 fig. 45.1, 173 fig. 49.4, 182 fig. 56.5, 199 fig. 61.1; Shilenkov 2021: 41.

Cicindela coerulea selengensis Mandl, 1936 : Mandl, 1935: 294 figs 6–8, 296 figs 6–10, 1936: 12; Horn 1938: 46, Taf. 63 fig. 26; Cassola & van Nidek 1984: 10; Werner, 1991: 18, T. 15 figs 112, 113; Wiesner 1992: 118; 2020: 184; Lorenz 1998: 43; 2005: 131; Puchkov & Matalin 2003: 104; Putchkov & Matalin 2017: 227.

Type material. Lectotype of Cicindela optata Fischer von Waldheim, 1828 (designated here), ♂ ( Figs 45 , 76, 77 )— C. optata Fisch., Nertshinsk [handwritten white label], Typus [typed pink label], Fischer von Waldheim [typed white label], Staatl. Museum für Tierkunde Dresden [typed white label], Lectotypus, Cicindela (s. str.) nitida f. optata F.-W., design. A. V. Matalin, 1997 [handwritten and typed red label]; LECTOTYPUS, Cicindela optata Fischer von Waldheim, 1828 , design. A. V. Matalin, 2024 [printed red label] ( MTD). Paralectotype of Cicindela optata Fischer von Waldheim, 1828 (designated here), ♂ —Staatl. Museum für Tierkunde Dresden [typed white label], Paralectotypus, Cicindela (s. str.) nitida f. optata F.-W., design. A. V. Matalin, 1997 [handwritten and typed red label], PARALECTOTYPUS, Cicindela optata Fischer von Waldheim, 1828 , design. A. V. Matalin, 2024 [printed red label] ( MTD).

Paratypes of Cicindela coerulea selengensis Mandl : 1♂ ( Fig. 113 )—Transbaic., Selenga fl. [handwritten white label], Сollect. Hauser [typed white label], C. tricolor var. [handwritten white label], v. optata , det. Ganglbauer [typed and handwritten white label], Cicindela nitida Licht. s. selengensis m., det. Ing. Mandl [typed and handwritten white label], PARATYPUS (selengensis Mandl) [typed and handwritten red label]; Cicindela (s. str.) coerulea (Pallas) selengensis Mandl 1936, Det. J. Probst [printed white label], NHMW-ZOO-COL-0002631 [printed white label]; 1♀ —Transbaic., Selenga fl. [handwritten white label], Сollect. Hauser [typed white label], v. optata , det. Ganglbauer [typed and handwritten white label], PARATYPUS (selengensis Mandl) [typed and handwritten red label], NHMW-ZOO-COL-0002632 [printed white label]; 1♀ ( Fig. 114 )— ibid, NHMW-ZOO-COL-0002633 [printed white label]; 1♀ ( Fig. 112 )—Transbaicalia, Selenga Tal., V – VI.1908 [typed white label], Сollect. Hauser [typed white label], v. optata , det. Ganglbauer [typed and handwritten white label], PARATYPUS (selengensis Mandl) [typed and handwritten red label], NHMW-ZOO-COL-0002634 [printed white label]; 1♀ —Transbaic., Selenga fl. [handwritten white label], Сollect. Hauser” [typed white label], v. optata , det. Ganglbauer [typed and handwritten white label], Cicindela nitida Licht. s. selengensis m., det. Ing. Mandl [typed and handwritten white label], PARATYPUS (selengensis Mandl) [typed and handwritten red label], NHMW-ZOO-COL-0002635 [printed white label]; 1♂ ( Figs 80, 81 )— ibid, NHMW-ZOO-COL-0002638 [printed white label]; 1♀ —Cotype [typed red label], Selenga-Tal, Transbaikalien [handwritten white label], Cicindela nitida Licht. ssp. selengensis m., det. Ing. Mandl [typed and handwritten white label], NHMW-Acquisitions-Nr.: Col-1971-6, coll. E. Grundmann [printed white label], NHMW-ZOO-COL-0002639 [printed white label] .

Additional material. RUSSIA, Altai Republic : 1♂ —Chuysky trakt, Ust-Inya, 15.VII.1907, E.G. Rodd.; 1♀ — Naryn-Gol River, 1956, Pridanieva; 1♀ — Altai, env. Ongudai, leg. A.G. Yakobson (all ZIN); 3♀♀ — ibid, 2– 3. VI.1989, S. Saluk; 1♀ —Chemal, 30.VII.[19]63; 1♂ —mouth of Chulcha River, right tributary of Chulyshman River, 29.VIII.[19]34, V. Geptner; 1♂ —mouth of Malyi Karakem River, tributary of Chulyshman River, 25. VI.1989, leg. V.N. Dremin (all ZMUM); 1♂ — Altai, Seminskiy pass, VII.1979; 2♂♂ 2♀♀ —valley of Chuya River, ZhanaAul, 1850 m, 8.VII.2022, leg. A. et E. Filippov (both MSPU); 3♂♂ 1♀ —Altaigeb., Akash, Mittellauf des Mjonka, Kuraikette, 1600–2300 m, 12.6.1992 (cJW); 2 ex. — Altai, Akash, Mjonka-Fluss, 20.05.1993, leg. A. Budarin; 9 ex. — ibid, 14.06.1991, leg. O. Legezin (both cJG); 1♀ —SE Altai, Kurai, 1.VIII.[1]964, Kerzhner; 1♂ 1♀ —Buguzun River, Chikhacheva Mnt Ridge, ~1900 m, 29.VII.[1]964, Grunin; 1♂ 1♀ —Chagan-Uzun env., bank of Chuya River, 5–6. VI.1989, V. Grichik et S. Saluk; 1♀ — ibid, A.S. Pisanenko (all ZIN); Republic of Khakassia: 2♂♂ 2♀♀ —Shira Lake, 5.VIII.1916, Khvorov; ( ZMUM); 2♂♂ — Tomsk Gouv., Fluss Abakan, Ui-Bat River, 17.VII., A. Jakobson; 1♂ 1♀ — Tomsk Gouv., Tashtyb, 16. VI., A. Jakobson (both ZIN); 1 ex. —Abakan, Krasnoe Lake, 26.07.1993, leg. A. V. Chuvilin (cFB); 2♂♂ 1♀ —Beisk District, near Tabat vill., 19.VIII.[19]63, O. Grigorjev ( ISEA); 3♂♂ 3♀♀ —2–3 km S of Abakan, 5.VII.1997, leg. V. Patrikeev; 1♂ —valley of Son River, 2. VI.1969, leg. V. Patrusheva (both MSPU); 3♀♀ —Borets, bank of Son River, 54.46 oN 90.36 oE, h~380 m, 29.06.2011, leg. K. Tomkovich (cOM); Republic of Tuva: 6♂♂ 9♀♀ —Erzin, sands on the shore of Tere-Khol, 7. VI.1962, leg. D. Berman ( ZMUM); 1 ex. — Tuva, Erzin, Erzin-River, 30.05.1990, leg. S. Gundorov (cJG); 2♀♀ —17 km NE of Khadyn Lake, 26. V.1988, leg. V. Zinchenko; 1♀ —Balgazyn, 8. VI.1948, Cherepanov (both ISEA); 1♂ — ibid, 28.VIII.1959, leg. S. V. Sharova (cISl); 1♀ —Balgazyn, 110 km SE of Kyzyl, VII.1956, Galkin; 6♂♂ —Toore-Khem, 11.08.1973, leg. B. Korotyaev; 1♂ —near Erzin, 21.07.1988, leg. S. Byzov (all ZIN); 1♂ 3♀♀ —SE shore of ToreKhol Lake, 31. V –1. VI; 1♂ —Kyzyl District, bank of Biy-Khem River, near Kara-Khaak, 20. V.1997, leg. V. Patrikeev; 2♂♂ 3♀♀ —Shagonar, 26.07.1992, A. Napolov; 2♂♂ —near Kyzyl, bank of Enisey River, 1–6.VII.1997, leg. S. Vastchenko & R. Mishustin; 6♂♂ —Khadyn Lake, 16–19. VI.1997, leg. S. Vastchenko & R. Mishustin; (all MSPU); 1♂ — ibid, 25. V.1998, leg. A. Gorodinski (cXX); 1 ex. — ibid, 20.05.1998, leg. S. Vastchenko (cFB); 1♂ 1♀ — ibid, 25. VI.1998; 2♂♂ —30 km N Balgazyn, 21.6.1994, leg. P. Smrz, Z. Kletecka (both cJW); Krasnoyarsky Krai : 1♂ —Bazaikha near of Krasnoyarsk, 14. V.[19]01, Krutovsky; 1♂ — ibid, 9. VI.[19]01; 19♂♂ 20♀♀ —Kaptyrevo; 10♂♂ — Krasnoyarsk (all ZIN); 1♂ 1♀ —O Sibiria, Berezovka, leg. Dr. J. Fleischer (cJW); 3♂ 5♀ —Sibiria, Minusinsk, Exp. Zichy, leg. Csiki ( HNHM); 1♂ —Minusinsk, Opytnoe Pole, 18. V.1928; 5♂♂ 4♀♀ — ibid, 25– 31.VII.1928 (both ISEA); 1♂ —Sibiria, Krasnoyarsk, A. Zolotarev; 3♂♂ 3♀♀ — ibid, 1899, L.A. Molchanov; 2♀♀ — ibid, VI.1902; 1♂ — ibid, 19. VI.1902; 1♂ 3♀♀ — ibid, VII.1902; 1♀ — ibid, VIII.1902; 1♂ 1♀ —Vostochnoe, 35 km S von Minusinsk, 1928, A. Zhelokhovtsev (all ZMUM); Irkutsk Oblast: 1♀ —Irkutszk, 3.VII.1967, leg. G. Rácz ( HNHM); 2 ex. — Irkutsk, 5 km NW of Cap Krestowski, 10 km S of Jelantsy, 22.06.1993, leg. R. Krause, O. Jäger; 4 ex. —Ust Orda, Kuda River, 26.06.2011, leg. W.-H. Liebig; 1 ex. —Nischnij Udinsk, Ude River valley, 28.06.2003, leg. A. Azarov; 1 ex. —Porog, Uda River valley, Privolnoye, 28.06.2003, leg. A. Azarov (all cJG); 2 ex. —Khatar-Khaday, 100 km NE of Irkutsk, 03.08.2000, leg. L. Lehmann (cAP); 1♂ —Baikal Lake, Slyudyanka dist., Kultuk vill., 12.6.2007, leg. S. Rybalkin; 1♀ —Baikal, Olkhon Peninsular, VIII.1973, Medvedev (both cJW); 1♂ — ibid, 15.08.1979, V. Shilenkov ( SFU); 1♂ —downstream of Oka River, 23.VII.1999, leg. K. Gongalsky ( MSPU); 1♂ —Malyshevka opposite Balagansk, bank of Angara River, 20.VII.[18]98, Ingenitsky; 1♂ —Shivera on Angara, 20.VIII.[18]98, Ingenitsky; 1♂ 2♀♀ —Malta (all ZIN); 2 ex. —Kodat, Mt. Togo, 2500 m, 07.07.2011 (cFB); 86 ex. —Kachug distr., bank of Lena River, h=501 m, 53.99141 N 105.72398 E, 05–15.06.2022, leg. Yu. & N. Seregin (cVS); Republic of Buryatia: 1♀ —Kyakhta ( MNHN); 1♀ — ibid, 05.07.1971; 1♂ 1♀ —near Kyakhta, Deben vill., 11.VIII.1977, leg. V. Shilenkov; 2♂♂ —Tunkinskie Goltsy, near Mondy, h~1599 m, 6–10. VI.1992; 1♂ —Mondy, valley of Irkut River, 20–30.VII.1977, leg. V. Shilenkov (all MSPU); 14 ex. — ibid, 06.2012, leg J. Gebert (cJG); 4 ex. — ibid, 06.06.2006, leg. E. Gavrischuk & O. Pekarsky; 4 ex. — ibid, 01.06.2011, leg. A. & E. Filippov (both cFB); 1♂ —Mondy, Sajan Geb.; 1♂ 2♀♀ — Burjatia, Sajany, Mondy, 1200 m, 15.7.1993; 3♂♂ 1♀ — ibid, 1.8.2002, leg. I. Legezin (all cJW); 1♂ —middle current of Oka River, Arkhabom, 19–21.VIII.1999, leg. K. Gongalsky; 4♂♂ 2♀ ♀ —between Khoronkhoi and Naushki, 50 o 25’38’’N 108 o 08’18’’E, 13–23. V.1999, leg. I. V. Melnik; 2♂♂ 1♀ —2 km E of Beloozersk, 50 o 38’30’’N 105 o 44’18’’E, 1–7.VII.1999, leg. I. V. Melnik; 1♂ 1♀ —5 km NE of Kiran, 50 o 21’43’’N 106 o 46’11’’E, 25–30. VI.1999, leg. I. V. Melnik; 2♂♂ —valley of Dzhida River, near Zakamensk, 19.06.2000, leg. P.Ya. Ustuzhanin; 2♂♂ 1♀ —Tsakir, 7.VII.2004, leg. K. Eskov; 1♂ 4♀♀ —Novaya Komushka, 51 o 47’N 107 o 38’E, 8.08.2015, S. Rudykh; 1♀ —Tunka River, tributary of Irkut River, 22. VI.1914 (all MSPU); 1♂ 2♀♀ —Ust-Kiran, 14.6.1956; 1♀ — ibid, 24.6.1951, N. Filippov; 2♂♂ — ibid; 1♂ —Tory, 17. VI.1914; 1♂ 2♀♀ —Mondy, 13–16.VII.1947, Tsvetaev (all ZMUM); 3♂♂ — ibid, 4–19. VI.1915, S. Rodionoff; 2♂♂ 1♀ — ibid, h=1600–1700 m, 7–8.06.2002, Lvovsky; 1♂ —Ust-Kiran, 20. V.[1]913; 1♂ — ibid, 30. VI.[1]913; 1♂ — ibid, 11.VII.[1]913; 1♂ 2♀♀ —Kyakhta, VII–VIII; 2♀♀ —Olkhon Island, near Ulan-Khushin, 11.VII.1973, N. Ermakova; 1♂ —Verkhneudinsk (= Ulan-Ude), 18.VII.1899, G. Suvorov (all ZIN); 1♂ 1♀ — Transbaikal, Werchne-Udinsk (= Ulan-Ude) (cJW); 3 ex. — ibid, 01.06.2014, leg. A. Filippov; 1 ex. — ibid, 12.06.2014; 1 ex. — ibid, 13.06.2015; 3 ex. —Zaigraevo, Brjanka River valley, 15.06.1997, leg. A. Petrov (all cFB); 1♂ 1♀ —Vjerchne-Ud.; 1♂ —Vjerhnje Ud., u Kulicha Bäd.; 2♀♀ —Kul. Bad. (all ZMUM); 1♂ 1♀ —Transbaikal, Werchne-Udinsk (= Ulan-Ude), leg. Dr. K. Mandl; 1♂ —Mondy, Irkut superior, 900 m, 21.VII.1973, leg. Shilenkov (both HNHM); 2♂♂ 2♀♀ —near Mondy, sandy bank of Irkut River, 09.07.1969; 1♂ —Vitim Plateau, vill. Baisa, 16.VII.1978, leg. T. Zolotarenko; 1♂ 2♀♀ — Dureny, bank of Chikoi River, 1–2.VIII.1986, leg. B. Zakharov et Yu. Chernov (all ISEA); Zabaykalsky Krai : 1♀ — Transbaic., Čita, 1910; 6♂♂ 1♀ —Transbaic., Tshita (both ZMUM); 2♂♂ — ibid ( MSPU); 1♀ — ibid, Hermann Frieb (cJW); 1♂ — ibid, coll. Jul. Isaac ( HNHM); 3♂♂ — ibid, V.1916, Yurkevich; 1♂ — ibid, VII.[1]912, Pisarevsky; 2♂♂ 3♀♀ — ibid, VI –VII.[1]910, I. Zadoplyaev (all ZIN); 1♂ — ibid, 10.07.1929; 2♂♂ —Chita env., 17.07.1959, N. Filippov (both MSPU); 1 ex. —Chita (Tshita), Ingoda-Tal Steppenhang, 14.06.1992, leg. Kostjuk; 1 ex. — ibid, 13.06.1994, leg. R. Krause, O. Jäger (both cJG); 1♀ —Usugli, 10.VII.1958, M. Lurie; 1♂ —Molokovka, P. V. Pisarev; 2♂♂ 1♀ —East Siberia, Sretensk; 1♂ —Sretensk District, Dunaevo env., pad Balagannaya, 17.IX.1957, G. Dlussky; 1♀ —Ust-Dunaevo, 13.IX.1957, G. Dlussky; 2♀♀ —Burgen, bank of Chitinka River, 10.VII.[1]912; 3♂♂ —Amazar, 25. V.1913; 4♂♂ 3♀♀ —near Nerchinsk, V. Podgorbinsky (all ZMUM); 1♂ —Nertschinsk; 1♂ —Bolshie Boty; 1♂ — Ingoda River, pad Argalei, 4. VI.1899, G. Suvorov; 2♂♂ —Aga [= Khyla] (all ZIN); 1♀ —Kalar District, Chara, VI.1975 ( ISEA); 2♂♂ —Drovyanaya, 2.VII.1977 (cEK; cAKr); 3♂♂ —Sokhondo Nature Reserve, cordon Agutsa, 16.06.1991, leg. V. Pekin, D. Logunov ( ISEA); 1♀ —Chita Region, near Borsya, 08.1995, leg. A. Sedov (cEK); 3♂♂ —Nertschinsk ( ZMUM); 1♀ —Transbaikal., Nerchinsk, 16.VII.1915; 1♂ —Abagatui, 17.08.1992, leg. M. Chernyakhovsky; 1♀ —Chita env., 15. VI.1983, leg. G. Nesterov; 1♀ —shore of Tsagan-Nur Lake, 4.VII.2000, leg. I. V. Melnik; 6♂♂ 3♀♀ —5 km W of Nizhny Tsasuchei, floorplan of Onon River, on sand with sparse vegetation, 14–18. VI.1995, leg. D. Fedorenko (all MSPU); 2♀♀ —Chita reg., Onon Riv., Tsasuchei vill., 3. VI.2002, leg. A. Anischenko (cJW); Republic of Yakutia : 2♂♂ 2♀♀ —near Yakutsk, 11. VI.–9.VII.1962, Zhelokhovtsev; 1♂ — ibid, 15.06.81, I. Medvedev (both ZMUM); 1♂ — ibid, 28. V.927, Nikiforov ( ZIN); 1♂ — ibid, 01. VI.1921, A. Tsvetaev (cEK); 2♂♂ 1♀ —Yakutsk, [1]901, Olenin; 1♂ — ibid, 9. VI.9127, Moskvin; 2♀♀ — ibid, 1883, Herz (all ZIN); 1 ex. — ibid, 01.07.1987, leg Z. Mracek; 2 ex. —Wladimirowka, 18.06.1989, leg. Subota (both cJG); 1♂ —Yakutsk, Botanical Garden, 24. V.1985, leg. V. V. Dubatolov; 4♂♂ — ibid, 18.VIII.1985 (both ISEA); 2♂♂ —40 verst from Yakutsk, Ytyk-Khaya, 9. VI.[19]01, Herz; 2♂♂ 2♀♀ —Ytyk-Khaya, Lena mer; 1♂ —islands on Lena River, near Ytyk-Khaya, 20–21.VII.[19]01, Herz; 1♀ —Dupsya, 8.VII.911, I. Afanasiev; 2♂♂ —Sergeleekh, 3 versty from Yakutsk, 15.VIII.911, I. Afanasiev; 1♂ —between Lena River and Verkhoyansky Mnt Ridge, [18]91, Čerski; 3♂♂ 2♀♀ —Olekminsk, 22. V.–2. VI.[19]08, Buturlin; 1♂ —near Olekminsk, 6. VI.[19]03, Kharitonov; 2♂♂ — ibid, 17. VI.[19]10 (all ZIN); 2♂♂ —right bank of Lena River, Nizhny Bestyakh, 2–17.VII.2001, S. Kurbatov; 3♀♀ — Matta River, ~ 20 km from the confluence with the Sinyaya River, 12.07.2017, leg. A. Ignatenko (both MSPU); 3 ex. —Oimyakon,Tomtor vill., leg. V.Zaritsky (cFB); Amur Oblast: 3♂♂ —30km W of Svobodnyi, near Kostyukovka, 5–9.VII.1995, leg. D. Fedorenko; 1♂ 1♀ —12 km S of Svobodnyi, Malaya Sezemka, bank of Zeya River, on heavily silted sand, 5.VII.1995, leg. D. Fedorenko; 2♂♂ —Innokentievka, 23.VII.1969, leg. Yu. Tretjakov (all MSPU); 1♀ —Kasatkino, 2.IX.1972, V. Zherikhin; 1♂ 1♀ —Arkhara, 4.IX.1972, leg. V. Zherikhin (both ZMUM); 2♂♂ 1♀ —Blagoveshchensk District, cf 15 km W of Natalino, forest, 27–31.VII.2003, leg. B. Kataev ( ZIN); 1♂ —near Blagoveshchensk, 14–17.07.1996, leg. V. Bezborodov; 3♂♂ —Zeya District, Gilyui River, 4–6.VIII.1971, L. Morozova (both ISEA); 1♂ —N of railway station Tynda, sand and pebble bank of Lena River, 18.06.1997, leg. R. Dudko, D. Lomakin ( MSPU); 1♂ 2♀♀ —Khingan Natural Reserve, cordon Erakta, 9.08.1989, leg. Yu. Tretjakov (cEK); Khabarovsky Krai : 1♀ —40 km SE of Mogdy, upstream of Amgun River, 20.07.1997, A. Brinev ( MSPU); 2♂♂ — ibid ( SFU); 1♀ —near Dzhamku, bank of Amgun River, 20–22.06.1996, leg. A.A. Kuzmin ( ISEA); Jewish Autonomous Oblast: 2♂♂ 2♀♀ —near railway station Bira, 14.VII–2.VIII.1994, leg. P. Udovichenko ( MSPU); 1♂ 1♀ —below Radde vill., Dichun River, 9.VIII.1980, leg. S. Alekseev (cSA); Primorsky Krai : 1♂ —Sidemi (= Narva River), 26. VI.1900, G. Suvorov; 1♀ —lower reaches of Tetyukhe (= Rudnaya) River, 20.VIII.[19]09, Dyukin; 3♂♂ — Vladimir Monomakh (= Monomakhovo), 3.VIII.–2.IX.[19]09, Dyukin; 3♂♂ 1♀ —Sikhote-Alin, 35 km NE of Sergeevka, 1.IX.1986, leg. B. Kataev; 3♂♂ 2♀♀ —Khanka Lake, Troitskoe, 4. V.–27. VI.[19]09, Čersky; 1♀ — Iman (= Dalnerechensk), VI.[18]99, Veselovich (all ZIN); 1♂ 2♀♀ —Khanka Lake, Novokachalinsk, 10. VI.1972, A. Ponomarenko; 1♀ —Ussuri River, Osinovka, 11. VI.[1]917, P. Elsky; 1♀ —fl. Sujfun (= Razdolnaya), Nik.- Ussurijsk, 23. V.; 1♀ —15 km SW Valentin, Glazkovka, 7.VII.1986, A. Antropov; 1♀ —Sudzukhinsky (=Lazovsky) Natural Reserve, Tyapigou (= Pribrezhnaya) River, 24.VII.[19]46, D. Panfilov; 1♂ —cove Kit, 11.VII.[19]46 (all ZMUM); 1♂ 1♀ — SU, Ussuri Region, Archipovka, 17.07.1989, leg. S. Becvar (cJW); 1♂ —Khasan District, Nature Reserve Kedrovaya Pad, 21.VIII.1964, D. Usachev ( ZMUM); 1♀ — ibid, 14.06.1968, G. Zolotarenko ( ISEA); 1♀ — ibid, 06.06.1962 ( MSPU); 1♂ —Gamova Peninsula, pass between cove Vityaz and cove Telyakovskogo, 16.VII.1979, V. V. Dubatolov; 1♂ — Gamova Peninsula, cove Astafieva, 21.VII.1993, V.K. Zinchenko 1♂ —Khasan District, Golubiny Utes, 12. VI.1968, V. Nikolaev (all ISEA); 1♀ — ibid, 22. V.2022, leg. K. Makarov & A. Matalin; 1♂ 2♀♀ —Ussurijsk District, Gornotaezhnaya Stantsiya, 20. VI.1960; 1♂ —Lazovsky Natural Reserve, cordon America, 43 o 16’16’’N 133 o 03’01’’E, 19.VII.2006, leg. K. Makarov; 1♀ —Vladivostok env., VI.1989 (all MSPU); 1♂ — Slavyanka env., VI.1993, E. Zhuravlev (cAKr); 2 ex. —Primorje, 14 km SSW of Krounovka, Krounovka-River, 11.08.1994, leg. P.-H. Schnitter, E. Grill (cJG); 1 ex. —Novonezhino, Venedivnovo, Tschernye Gory, 03.08.1990, leg A. Pütz (cAP); MONGOLIA, Uvs Aimag: 1♂ —Sandgebiet Altanels, 35 km WNW von Somon Tes, 1400 m, 3. VI.1968, leg. Dr. Z. Kaszab ( HNHM); 1♀ —Üüreg Lake, 14.08.1970, leg. B. Korotyaev ( ZIN); 1 ex. —Burgag Nuur (= Shaazgay Nuur), 03.07.2016, leg. P.-H. Schnitter (cJG); Khovd Aimag: 3 ex. —Baruun Chara, 5 km, Halbwüste, 29.07.1999, leg. A. Stubbe (cPS); Zavkhan Aimag: 1♂ 1♀ —Dzabyn-Gutu-Khudzhirt, 18–21.VII.1877, G. Potanin ( ZIN); 1♀ —Choit chunch, 26 km ONO von See Telmen nuur, 2150 m, 13.VII.1968, leg. Dr. Z. Kaszab; 1♀ —Santmargaz sum, Bajan (= Chara) nuur, 18.VIII.1963, leg. A. Bold (both HNHM); 1 ex. —W Somon, Aldarchaan, 26.07.2010, leg. A. & M. Stubbe; 3 ex. —Zawchanmandal, 09.06.2012, leg. J. Müller, A.J. Müller (both cJG); Khövsgöl Aimag: 1♀ —25 km NNE of Ulaan-Uul, 15–16.VII.[1]975, Gurieva; 1♀ —10 km SE of Muren, 23–24.VII.[1]975, Sugonyaev; 1♂ 1♀ —25 km W of Muren, 27. VI.[1]968, Arnoldi (all ZIN); 1 ex. —Khankh, 21.07.1995, leg. E. Novikov (cFB); Bulgan Aimag: 4♂♂ 1♀ —64 km W of Erdenecant, 1260 m, 47°05’N 104°05’E, 3–24. V.1990, leg. Gy. Fábián, M. Hreblay, L. Peregovits & G. Ronkay; 1♀ —11 km W von Somon, Bajannur am see Bajan nuur, 1000 m, 14. VI.1968, leg. Dr. Z. Kaszab; 5♂♂ 5♀♀ — ibid, 1000–1100 m, 24.VII.1968 (all HNHM); 1♂ 2♀♀ — ibid, 24.VIII.1968 ( NHMW); 2♀♀ —180 km SW of Ulashivet, Tarn (Lun) River, 47°19.380’N 103°41.751’E, 21.VII.2019, leg. H. Mühle (cJW); 1 ex. —Changai Mts., Orchon Brücke—Ulan Bataar lager steppe, 27.07.1999, leg. A. Stubbe (cPS); Arkhangaj Aimag: 1♀ —30 km E of Tsetserleg, 2–4.VII.[1]975, Narchuk; 1♀ —48 km S of Tevshruulekh, 2–3.VII.[1]975, Gurieva; 1♀ —Tevshruulekh, 13.VIII.1974 (all ZIN); 1♀ — ibid, 24.VIII.1971 ( ZMUM); 1♂ — ibid, 16.7.1975 (cJW); 1♀ —63 km O von Urdtamir, Chaalgsin chundi bei Somon Tövschruulech, 1500 m, 28. VI.1966, leg. Zanzan Tombo; 5♂♂ 2♀♀ —NO of Ecke des Sees Ogij Nuur, 1350 m, 2.VII.1964, leg. Dr. Z. Kaszab (both HNHM) 1♂ — ibid, ( NHMW); 1♂ — ibid (cJW); 2♂♂ 1♀ — Nord Mongolia, Changai, Leder ( ZIN); 1♂ 4♀♀ — ibid ( HNHM); 3♂♂ 3♀♀ — ibid ( MNHN); Selenge Aimag: 3♂♂ 5♀♀ —Ero-Gol River near Dulaankhaan, 3–4.VIII.[1]975, Gurieva; 1♂ 1♀ — ibid, 30.VIII.[1]975; 1♀ —Shaamar, summer 1984, S. Kuzmin; 2♂♂ —coupling of Suman River and Chuluut River, 29–30. VI.[1]975, Narchuk (all ZIN); 1 ex. —Eröo Somon, Eröö Gol, Hochsteppe, 26.07.1999, leg.A. Stubbe (cPS); 1♀ —Sumber Dist., 5 km S of Baruun Kharaa, 48°51.179’N 106°05.006’E, 792 m, vegetated sand plain, 7. V.2018, leg. J.A. Shetterly; 1♀ —Khongor Dist., 49°22.417’N 105°52.304’E, 668 m, vegetated low sandy dunes, leg. J.A. Shetterly (both cJW); 1 ex. —Ochron-Aue Schotterufer, Suchbaatar, 05.06.2001, leg. Stenzel; 1 ex. —Orchon-Aue, Caragana-Steppe, 06.06.2002, leg. Stenzel (cJG); 4♂♂ 2♀♀ —Transbaic., Selenga fl., Collect. Hauser, hybrida tricolor det. Ganglbauer; 1♂ —Transbaicalia, Selenga Tal, V – VI, 1, Collect. Hauser (both NHMW); 2♂♂ 2♀♀ —Mongolie sept., Selenga, J. Chaffanjon, 174-96 ( MNHN); Darkhan Uul: 1♀ —20 km NNE of Darkhan, 5.VIII.[1]975, Gurieva; 1♂ —25 km E of Darkhan, 30.VII–2. VIII.[1]975, Gurieva; 1♀ —Khara River, tributary of Selenga River, 1. VI.[18]97, Klements; 1♀ —Sharyn River, tributary of Selenga River, 31. V.[18]97, Klements (all ZIN); Ulaanbaatar: 1♂ 2♀♀ —Urga, 13.08.1913, Schultz ( HNHM); 9♂♂ —Nalaikh, 40 verst E of Urga, 1–2.VIII.[18]97, Klementz; 1♂ — ibid, 25. VI.[1]967, Zaitsev (both ZIN); Töv (= Central) Aimag: 2♂♂ —Ulan Bator, Nucht im Bogdo uul, 1800–2000 m, 27.VII.1966, leg. Dr. Z. Kaszab; 1♂ —Tel des Tola, zwischen Somon Altanbulag und Somon Tariat, 30 km ONO von Tariat, 1200 m, 25.VII.1966, leg. Dr. Z. Kaszab (both HNHM); 1 ex. —170 km W of Ulaanbataar, 16.08.2007, leg. P. Tyrner; 2 ex. —Terelz, Tuur, 21.07.2003, leg. J. Halada (both cJG); 4♂♂ 2♀♀ —45 km E of Lune, 4.IX.67, Zaitsev; 2♂♂ 4♀♀ —70 km SW of Ulan Bator, sandy steppe, 30.VII.[1]963, Arnoldi, Gurieva; Kerzhner, Zaitsev; 1♀ — ibid, 125 km SW of Ulan Bator, 30–31.VII.[1]963, Kerzhner ( ZIN); 3 ex. —SW of Altanbulag, Tola-Aue, 08.08.2009, leg. A. Stubbe (cPS); Khentii Aimag: 1♂ —20–40 km SW von Somon Zargaltchaan, S von Fluss Zencher gol, 1200 m, 23.VIII.1963, leg. Dr. Z. Kaszab ( HNHM); 5 ex. —Batshireet 3 km E of Bakh Gol, 03.08.2019, leg. P.-H. Schnitter (cPS); Govisümber Aimag: 2♀ ♀ —25–35 km NNW of Choir, 12.VII.[1]975, Gurieva ( ZIN); 1♀ —20 km SO von Čojren, 1200 m, 1.VII.1963, leg. Dr. Z. Kaszab ( HNHM); Dornogovi Aimag: 1♂ —12 km NE of Mandakh, 15. VI.[1]971, Kerzhner ( ZIN); 1♀ —Erdene-Dalay, 26. V.19[73], Sowjet-Mongol. Biol. Exped. 1973 ( ZMUM); 1♂ 2♀♀ —40 km NW von Chara-Eireg, 1150 m, 30. VI.1963, leg. Dr. Z. Kaszab ( HNHM); Dornod (= East) Aimag: 1♂ 5♀♀ —Čojbalsan, 1969, leg. Gy. Márton ( HNHM); 1♂ —50 km ESE of Choibalsan, 12.VIII.[1]976, Gurieva; 6♂♂ 1♀ — Bui Nuur (= Buir) Lake, VI.[18]99, exp. of Potanin; 1♀ —steppe W of Bui Nuur (= Buir) Lake, 25.VIII.[19]28, A. Ivanov; 1♀ —shore of Bui Nuur (= Buir) Lake, 25 km NE Bayan Nuur Lake, 17. VI.[1]971, Kozlov; 1♂ —10 km W of Dund-Bayan Lake, 27.VIII.[1]975, Gurieva; 1♂ —20 km ENE of Bayan-Uul, 8.VII.[1]976, Gurieva; 1♀ — Khökh Nuur Lake, 24–26. VI.[1]976, Gurieva; 1♂ —Derkhin-Tsagaan-Ovoo, 60 km ENE of Bayan-Burda, 21. VII.[1]971, G. Medvedev; 1♂ —Numragin-Gool River, 32 km SE of Salkhit Mnt, 24.VII.[1]971, G. Medvedev (all ZIN); 6♀♀ —Chamardavas uil, 80 km SO von Somon Chalcingol, 60 m, 13.VIII.1965, leg. Dr. Z. Kaszab ( HNHM); 1 ex. —Chalchgol E, Sambariin Tsagaan Nuur, 02.08.2013, leg. P.-H. Schnitter; 1 ex. —Hyanganï Salbar Uuls, Hingan Mts., Nömrög Gol, 02.08.2013, leg. P.-H. Schnitter (both cJG); 4 ex. — ibid; 2 ex. —Tsagaannuur, Khalkhyn Gol, 31.07.2013, leg. P.-H. Schnitter (both cPS); Bayankhongor Aimag: 1♂ —25 km W of Bömbögör, 14. VI.[1]980, G. Medvedev ( ZIN); Övörkhangai Aimag: 1♀ —Ongi River, upstream of Khalkha River, 12.VII.[1]926, Kirichenko ( ZIN); 1♂ —64 km W of Erdenecant, 1260 m, 47°05’N 104°05’E, 23–24.5.1990, leg. Gy. Fábián, M. Hreblay, L. Peregovits, G. Ronkay (cJW); 1 ex. —Harhorin Karzhorum, Changai Mts., 15.07.2001, leg. K. Gedeon (cPS); Dundgovi Aimag: S of Delgercogt, 46 o 03’01’’N 106 o 21’38’’E, 07.08.2019, leg. A. Rubenyan ( MSPU); 1♂ 1♀ — Central Mongolia, Mandal-Gobi, 6.1964, Shuvalov ( ZIN); 1 ex. —80 km S of Ulan Bataar, Mandal-Gobi, Halbwüste, 16.07.1999, leg. A. Stubbe (cPS); 2♀♀ —SW of Chuld und Delgerehangaj, Choot bulag, 1480 m, 10. VI.1967, leg. Dr. Z. Kaszab; 2♂♂ — ibid, 12.VII.1967 (both HNHM); Ömnögovi Aimag: 1♂ —60 km W von somon Bulgan, 1120 m, 21. VI.1964, leg. Dr. Z. Kaszab; 1♂ —20 km NO von Somon Bulgan, Bajan zag, 1200 m, 6–7.VII.1967, leg. Dr. Z. Kaszab (both HNHM); 1♀ —Dalansadgat, 29.6.1983, leg. Ulrich Salk Bautzen (cJW); Sükhbaatar Aimag: 3♂♂ 4♀♀ —25 km NW of Bayandelger, 19.VIII.[1]975, Gurieva; 1♂ —sands Ongon-Els, 15 km SSE of Hongor, 5– 6.VII.[1]971, Kerzhner (both ZIN); 1♂ —Ongon-Els, 10 km S von Somon Hongor, 900 m, 3–5.VIII.1965, leg. Dr. Z. Kaszab ( NHMW); 1♂ — ibid, 8.VIII.1965 ( HNHM); 1♂ —16 km W of Govi-Nuur, 30–31.VIII.1977 ( ZMUM); NORTH KOREA: 2♂♂ 1♀ —N Korea, 20–22. V.[18]97, V. Komarow ( ZIN); 1♀ — Ryanggang prov., Mupo, Tumangang, 1000 m, 29.IV.1988, leg. O. Merkl & G. Szél; 1♀ —Khamgen prov., Chongjin, 04.06.1991, leg. Ronkay & Vojnits (both HNHM); CHINA, Nei Mongol Autonomus Region: 1♂ —along Kerulen (= Herlen) River, exp. of G. Potanin ( ZIN); 1♂ — Inner Mongolia, Chifeng, Keteng, 10.IX.2006, leg. Y. Wang & M.J. Qi ( NEFU); Heilongjiang Province: 3♂♂ —Manchuria, railway station Imanpo (= Yimianpozhen), 20–21.VII.[1]914, Emelianov; 7♂♂ 7♀♀ —Manchuria, railway station Hengdaohezizhen (both ZIN); 1♂ –China, Mandzhuria, Gaolindzy st., 17.VII.1973 (cJW); 1♀ —Muleng, Xiaojinshan, 1. VI.1961; 1♂ —Mudanjiang, Ningan, 17–21.VIII.2008 (both NEFU); Liaoning Province: 1♂ 1♀ —PanJinShi, QingyanSi, 300 m, 13.VII.2007, Coll. Liu J.-Z. (cXX); Sichuan Province: 1♂ 1♀ — China, Sichuan, Wenchuan, 23.05.2012, leg. S. Murzin (cJG); Jiangxi Province: 1♂ 1♀ —Jiangxi, Ganzhou, Mount Jiulian, 24.97°N 114.73°E, 620 m, 25.IX.2016, leg. Zhenhua Liu ( SYSU); WITHOUT EXACT LOCALITY: 2♂♂ — Dauria; 1♀ —Sibiria; 1♂ —Amur, ex. coll. G. Basilewsky (all ZMUM).

References. RUSSIA, Altai Republic : 7 km W of Edigan, right bank of Katun River; Ulus-Cherga; bank of Karakem River, 2 km of mouth; near Chibit; Kuray; Kuray Steppe, floodplain terrace of Kurayka River; Kuray Steppe, NW Kuray; 6–8 km NE of Kokorya, Kyzylshin River, h~1900–2000 м; valley of Kyzyl-Aryk River (all Matalin & Dudko 2016a); Chemal; Chulyshman Upland, upstream of Chulyshman River; bank of Chuya River, near Chagan-Uzun; Chikhacheva Mnt. Ridge, Buguzun River, h~ 1900 m (all Matalin & Dudko 2016b); Krasnoyarsky Krai : Jenisseisk ( Mandl 1921; Wu 1937); near Krasnoyarsk, Mokhovaya, Sobakina, Narmut, Karaulnaya and Bazaikha Rivers (all Okninskyi 1930); Irkutsk Oblast: Katschug ( Poppius 1906); Irkutsk ( Mandl 1921); Malyshevka opposite Balagansk, bank of Angara River; Shivera, bank of Angara River (both Lafer 1978); Tazheran steppe, salineland on the bank of Holbo-Nur Lake ( Shilenkov 2021); Republic of Buryatia: Vjerhnje Oudinsk (= Ulan-Ude); Kiakhta (both Motchoulski 1844); Botyi (Jakobson 1907); Werchne-Udinsk (= Ulan-Ude); Troizkosawsk (= Kyakhta) (both Mandl 1921; 1931); Kalenovo (Imelhenova 1980); valley of Dzhida River (Shilenkov & Anichtchenko 1998); Ulan-Ude (Khobrakova 2003); Tamgata ( Khobrakova & Dudko 2010); Alla River; Angir-Nuur; Bagdarin; Baisa; Barguzin; Beloozersk; Bilutai; Bolshoe Akubycrit Lake; Verkhnyaya Ivolga; Guzhiry; Gunda; Gusinoe Lake; Gusinoozersk; Dalan-Turu; Dzhida; Dzhirga; Dureny; Deben; Zhakhna; Zakamensk; Zun-Murino; Indola; Kabansk; Karasinoe Lake; Karaftit; Kindikan; Kokorino; Kudara-Somon; Kuitun; Kumyn; Kyren; Maisky; Malyi Amalat; Moigoty; Monahovo; Mongolzhon; Mondy; Mostovoi; Murochi; Muya; Nizhnee Beloe Lake; Novaya Komushka; Novaya Kurba; Nukhu-Daban; Orlik; Orkhabom; Perilly; Savva; Sentsa River; Sentsyn-Tala; Seya River; Taezhnyi; Taksimo; Tankhoi; Tapkhar; Temnik (Atsula); Tissa River; Tunka; Turka; Uoyan; Ust’-Kiran; Kyakhta; Ust’- Kyakhta; Ust’-Muya; Ukherik; Khaigas; Khara-Khuzhir; Kharagun; Kharibyaty; Khoito-Gol; Khoyur-Gorkhon River; Khulugaisha; Tscistokhalo Lake; Khulugaisha; Khutel; Shasnur; Shishkovka (all Khobrakova et al. 2014); Kudara, Kentei-Geb. ( Mandl 1921); Zabaykalsky Krai : Tschita ( Mandl 1921); Ingoda River, pad Argalei; mouth of Aga River (both Lafer 978); Khilok (Khobrakova et al. 2014); Republic of Yakutia : Ytyk-haja ( Poppius 1906); Jakutsk ( Mandl 1921); Chochur-Muran, bank of Lena River ( Lafer 1978); Amur Oblast: Ulunga River; Natalino, bank of Zeya River; Saskal, mouth of Bereya River; Ekimchan, Selemdzha River (all Lafer 1978); 2.5 km E of Blagoveshchensk, floodplane of Zeya River ( Rogatnykh 2009a); Jewish Autonomous Oblast: Bastak Nature Reserve ( Rogatnykh 2006); Bastak Nature Reserve, kordon Zaimka Ivakina ( Budilov 2012); Khabarovsky Krai : Bureinsky Natural Reserve, near kordon Strela, bank of Bureya River, 51°38’ N 134°15’ E; mouth of Seregekta River, 51°36’ N 134°08’ E (both Koshkin et al. 2016); Primorsky Krai : Nikolsk-Ussurijsk; Pokrofka; SutschanGebiet (= Partizansky District) ( Mandl 1921); Khasan; Gornoe, Ananievka River; Ussury Reserve; Lesnoy Kordon; Benevka River (tributary of Kievka River); Chernigovka (all Lafer 1978); Lazovsky Nature Reserve: Vtoroi Log; Nogeevskaya; America; Petrova; Staraya Kamenka; Pasechnaya; Kievka; Benevskoe; Lazo (all Sundukov 2009); MONGOLIA, Selenge Aimag: Shaamar (Ulykpan 2008); Khövsgöl Aimag: Bayanzürkh ( Lafer 1978); Arkhangaj Aimag: Changai-Geb. ( Mandl 1921); NO Ecke des Sees Ogij nur ( Mandl 1965); Zavkhan Aimag: Santmargaz sum, Bajan (Chara) nur; Songino (both Mandl 1965); Gobi-Altai Aimag: Zavchan-Niederung, ca 20 km N von Zargalan, 47º20’N 95º48’E ( Mandl 1974); CHINA, Nei Mongol Autonomus Region: Baotou, 40.747706 N 109.791142 E, 11.06.2017 (https://www.inaturalist.org/observations/196973519); Heilongjiang Province: Suifen (Süd-UssuriGebiet) ( Mandl 1921); Liaoning Province: 41.198646 N 123.267497 E, 12.04.2020 (https://www.inaturalist.org/ observations/41983527).

Diagnosis. Cicindela coerulea nitida is recognized by complete and isolated white elytral spots (HI = 8.0–16.0, mean = 9.8) with distinctly sinuate medial band ended by round apical extension (typical pattern Figs 89–106 ). Moreover, from other subspecies except nominotypical one C. coerulea nitida is clearly distinguished by long and sharply rounded internal sack so В distinctly or virtually contact with the ventral side of aedeagus tube ( Figs 117–132 vs. Figs 167–170 , 190–195 , 233–240 ).

Redescription. TL = 12.0– 15.2 mm (mean = 13.76 mm, n = 93) in males, 12.4–15.4 mm (mean = 14.31 mm, n = 67) in females. Colouration of top side variable from unicoloured green, blue, or violet ( Figs 76, 78–80 ) to bicolored with green or bluish-green head and pronotum and cupreous elytra ( Fig. 82 ).

Head green or bluish-green, rarely bronze-green with golden or golden-cupreous reflection on genae, frons, vertex, occiput and ocular plates as well as with blue or greenish-blue shallow interocular cavities; in some specimens genae bright blue or head completely blue, violet or black. Labrum with 4–14 (mean = 8, n = 160) submarginal setae ( Figs 10–13 ); LW/LL = 1.83–2.44 mm (mean = 2.14 mm, n = 93) in males, 1.77–2.3 mm (mean = 2.05 mm, n = 67) in females. Mandibles relatively stout, ML/MW = 5.71–7.4 (mean = 6.45, n = 20) in males and 7.2–7.75 (mean = 7.48, n = 20) in females. Scape besides 3–5 apical setae with 1–5 additional setae, rarely glabrous.

Pronotum green or bluish-green with bright blue anterior and posterior grooves as well as midline and golden or cupreous-golden reflection especially on anterior and posterior lobes; in some specimens completely blue or violet, rarely black; PW/PL = 1.27–1.46 (mean = 1.37, n = 93) in males and 1.28–1.57 (mean = 1.41, n = 67) in females ( Figs 19–23 ). Thoracic segments bright green or blue-green with golden or golden-cupreous reflection or violet with blue tinge on mesepimeron, rarely entirely black.

Elytra generally cupreous with golden or golden-green reflection especially along suture, not infrequently green, blue or violet, rarely black; EL/EW = 1.45–1.71 (mean = 1.55, n = 93) in males and 1.41–1.57 (mean = 1.50, n = 67) in females; white elytral pattern typically complete, narrow or relatively wide, humeral and apical lunulae always isolated from marginal band, middle band distinctly curved with relatively long transverse portion, oblique or virtually vertical branch ended by rounded apical extension ( Figs 89–103, 105 ); in some specimens humeral lunula divided into two separate spots ( Figs 104, 106 ); sometimes middle band coupling via marginal band with apical lunule ( Figs 112, 113 ) or with both humeral and apical lunulae ( Figs 114–116 ); rarely middle band short and lacking rounded apical portion ( Figs 110, 111 ) or white elytral pattern completely undeveloped ( Figs 107–109 ).

Abdominal sternites green with golden or golden-cupreous tinge, or blue-violet with greenish-gold reflection.

Aedeagus relatively long ( Figs 45–54 ), EL/AL = 1.64–2.0 (mean = 1.79, n = 30), apex clearly curved with sharp, medium-sized hook. Internal sac long and sharply rounded towards to ventral side so В distinctly or virtually contact with the aedeagus tube, mt large with smoothly curved apex, VA and DA the largest, DLL relatively large ( Figs 117–132 ).

Distribution. RUSSIA: Republics of Altai, Khakassia, Tuva, Buryatia and Yakutia ; Krasnoyarsky, Zabaykalsky, Khabarovsky and Primorsky Krai ; Irkutsk, Amur and Jewish Autonomous Oblasts; MONGOLIA: Uvs, Khovd, Zavkhan, Khövsgöl, Bulgan, Arkhangaj, Selenge, Töv (= Central), Khentii, Govisümber, Dornogovi, Dornod (= East), Bayankhongor, Gobi-Altai, Övörkhangai, Dundgovi, Ömnögovi and Sükhbaatar Aimags, Darkhan Uul, Ulaanbaatar; NORTH KOREA: Rason City, North Hamgyong and Ryanggang Provinces; CHINA: Nei Mongol Autonomus Region; Heilongjiang, Jilin, Liaoning, Sichuan and Jiangxi Provinces. The eastern part of the Altai Piedmont Plain is the transition zone between C. coerulea nitida and the nominotypical subspecies; the valley of the Lakes between Gurvansaikhan and Khangai Mountains is the transition zone between C. coerulea nitida and C. coerulea kaszabi while the extreme south of Manchurian Plain is the transition zone between C. coerulea nitida and C. coerulea shantungensis ( Fig. 246 ).

Larva. See Putchkov & Shilenkov (1996) and Putchkov (2013; 2020).

Ecology. This is one of the most common Siberian and Mongolian tiger beetle inhabiting dry steppes, semideserts and open sands both near and far from water.

Development. The life cycle is biennial with spring–early summer reproduction. In the spring beetles both of new and previous generations copulate, and, after some time, females lay eggs. The larvae develop in soil burrows, and the third-instar larvae overwinter. Teneral beetles emerge in late summer–early autumn of the next year, feed and then overwinter. In spring of third year overwintered immature beetles maturate and start reproducing (Institute of Zoology Academia Sinica 1986).

Notes. In 1936 Mandl described the green specimens of Cicindela coerulea from the Selenga River valley as C. coerulea selengensis . Mainly green colour of the beetles, marginal band connected with apical lunula (HI = 15–16, mean = 15.3) and poorly developed rounded apical extension on medial band ( Figs 80 , 112–114 ) were the main criteria for this. Mandl noted that he had only one blue specimen, and he saw one dark brown specimen from the same locality ( Mandl 1936: 12). However, in the collection of Mandl (NHMW) five males and two females with the identical locality labels as in the type specimens were found (see chapter Addition material). Among them, one specimen has blue, one specimen has cupreous-green, and five specimens have cupreous elytra. Moreover, the marginal band in these beetles is short or entirely reduced, and all spots are separated from each other. Thus, by the white elytral pattern (HI = 8–10, mean = 9.6) these specimens are indistinguishable from typical C. coerulea nitida (HI = 8–14, mean = 9.4). Considering the high variability of the colouration as well as of the white elytral pattern in C. coerulea nitida , and also the location of the Selenga River valley virtually in the centre of the distribution area of the aforementioned subspecies, C. coerulea selengensis should be considered only as a pattern-coloured morph of C. coerulea nitida . Thus, I accept the opinion of Schilder (1953: 564), who considered Cicindela coerulea selengensis Mandl, 1936 as a synonym of Cicindela coerulea nitida Lichtenstein, 1796 . The absence of significant differences in basic proportions of body between C. coerulea nitida and C. coerulea selengensis ( Figs 241–245 ) reinforces this conclusion.

In the recent Checklist of the tiger beetles of the World ( Wiesner 2020), the names cyanescens Poppius, 1905; viridis Poppius, 1908; nigra Mandl, 1921; cobaltina Lutshnik, 1924; poppiusi Lutshnik, 1924; starodubtzevi Lutshnik, 1924 and starovi Lutshnik, 1924 were listed among the synonyms of C. coerulea nitida . However, according to the Articles 12.1 and 45.6.2 of ICZN (1999) the name is infrasubspecific if its author used one of the terms “ aberration ” and “ morph ” (or any their abbreviations), as well as according to the Articles 12.1 and 45.5 of ICZN (1999) the name is infrasubspecific if it was published as a fourth name as an addition to a trinomen. Thus, the names Cicindela tricolor ab. cyanescens Poppius, 1905; Cicindela tricolor ab. viridis Poppius, 1908; Cicindela hybrida nitida ab. nigra Mandl, 1921; Cicindela coerulea nitida ab. cobaltina Lutshnik, 1924; Cicindela coerulea nitida ab. poppiusi Lutshnik, 1924; Cicindela coerulea nitida ab. starodubtzevi Lutshnik, 1924; Cicindela coerulea nitida ab. starovi Lutshnik, 1924 are unavailable and should be excluded from the species group.

Chinese literature sources mention that C. coerulea nitida is recorded in total in 17 provinces (Institute of Zoology Academia Sinica 1986; Hu 1992; Zhu et al. 1999). It should be noted, that these records refer to at least three subspecies: C. coerulea nitida , C. coerulea kaszab i, and C. coerulea shantungensis . I was unable to examine the specimens from Jilin, Henan, Shaanxi, Jiangsu, Anhui, Zhejiang, Fujian, Guangdong and Yunnan Provinces. However, according to the presently known distribution of different subspecies, I provisionally consider the specimens from Jilin Province to be C. coerulea nitida and the specimens from Henan and Shaanxi Provinces to be C. coerulea shantungensis . The localities in Sichuan and especially in Jiangxi provinces are placed far away from the main distribution area of the species. Due to the scant material (male and female in each of them) I cannot reliably establish the subspecific status of these beetles, but preliminarily consider them as C. coerulea nitida . The records from other Chinese provinces mentioned above are unconfirmed and the status of the specimens from them is still unclear.

Figures

FIGURE 246. Map of the distribution of subspecies of C. coerulea (blue open circles—C. coerulea coerulea; red solid circles—C. coerulea nitida; light blue solid circles—C. coerulea kaszabi; red open circles—C. coerulea shantungensis; green solid circles—C. coerulea przewalskii; lilac solid circles—C. coerulea grumi; dark blue shading—transition zone between C. coerulea coerulea and C. coerulea nitida; light blue shading—transition zone between C. coerulea nitida and C. coerulea kaszabi; red shading—transition zone between C. coerulea nitida and C. coerulea shantungensis. Maps source—https:// mapswire.com/maps/asia-physical-maps/.

FIGURES 6–13. Cicindela coerulea ssp., labrum: 6–9—C. coerulea coerulea (6— Kuludzhanskie sands; 7—Biysk; 8— Zhana-Arka; 9—Semipalatinsk); 10–13—C. coerulea nitida (10—Sergeevka; 11—Buir-Nuur; 12—Kurai; 13—Burgen); 6, 7, 10, 11—males; 8, 9, 12, 13—females.

FIGURES 14–23. Cicindela coerulea ssp., pronotum: 14–18—C. coerulea coerulea (14—Chernoyarka; 15—Kashtakskoe lesnichestvo; 16—Miass River; 17—Ryzaevskiy; 18—Zhana-Arka); 19–23—C. coerulea nitida (19—Kerulen; 20— Verkhneudinsk (= Ulan-Ude); 21—Darkhan; 22—Balgazyn; 23—Chemal); 16, 19–21—males; 14, 15, 17, 18, 22, 23— females.

FIGURES 42–64. Cicindela coerulea ssp, aedeagus, left lateral view: 42–44—C. coerulea coerulea (42—Irtysh; 43— Kuludzhanskie sands; 44— Chernoyarka); 45–54—C. coerulea nitida (45—Nertshinsk; 46—Tere-Khol Lake; 47—Minusinsk; 48—Buir-Nuur; 49—Arkhara; 50—Yakutsk; 51—Sergeevka; 52—Nalaikh; 53—Jiulian Mt. (image by Keyi Wang); 54— Mandal Cobi); 55, 56—C. coerulea shantungensis (55—Peking; 56—Pin-Faun-Cha sands); 57–61—C. coerulea kaszabi (57, 58—60 km W v. Somon Bulgan; 59—60 km WNW of Bayan-Dalay; 60, 61—Barkull); 62—C. coerulea przewalskii, Amdo; 63—C. coerulea grumi, р. БадО-хО и ХЫ-хО-дО; 64—C. coerulea kozlovi, ДабасуН ГОби; 58—internal sack (without scale); 42—neotype; 45, 62—lectotypes; 57, 58—holotype; 63, 64—paralectotypes.

FIGURES 76–88. Cicindela coerulea spp., habitus, dorsal view: 76–82—C. coerulea nitida (76, 77—Nertshinsk (images by Jörg Gebert); 78—Khodyn Lake; 79—Kiran; 80, 81—Selenga fl. (images by Matthias Seidel and Helena Shaverdo); 82— Chita); 83–88—C. coerulea kaszabi (83, 84—60 km W v. Somon Bulgan; 85— Bayan Dalay; 86—Gulin-Tal; 87—Beger-Nur; 88—Tsagan-Gol River); 76, 77—lectotype of C. optata; 80, 81—paratype of C. coerulea selengensis; 83, 84—holotype; 77, 81, 84—labels; 78, 80, 83, 86, 88—males; 76, 79, 82, 85, 87—females.

FIGURES 89–106. Cicindela coerulea nitida, left elytron, typical pattern: 89—Hanka Lake; 90—Wenchuan (image by Keyi Wang); 91, 94, 100, 104, 106—Minusinsk; 92—Olekminsk; 93—Muren; 95—Mandakh; 96, 101—Luna; 97—Changai; 98— Jiulian Mt.; 99—central Yakutia; 102, 103—Mondy; 105—Nalaikh; 89, 91, 94–96, 100–103, 105, 106—males; 90, 92, 93, 97–99, 104—females.

FIGURES 107–116. Cicindela coerulea nitida, left elytron, atypical pattern: 107—Altai, Seminsky Mnt. pass; 108—Chita; 109, 110—Ongudai; 111—Amur; 112–114—Transbaikalia, Selenga fl. (images by Matthias Seidel and Helena Shaverdo); 115, 116—Mandal Gobi; 112–114—paratypes of C. coerulea selengensis; 107, 108, 111, 113, 16—males; 109, 110, 112, 114, 115— females.

FIGURES 117–124. Cicindela coerulea nitida, internal sack, right lateral view: 117—Khodyn Lake; 118—Abakan; 119— Mondy; 120—Olekminsk; 121—Dzabyn-Gutu-Khudzhirt; 122—Natalino; 123—Blagoveshchensk; 124—Innokentievka; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 125–132. Cicindela coerulea nitida, internal sack, left lateral view: 125—Khodyn Lake; 126—Abakan; 127— Mondy; 128—Olekminsk; 129—Dzabyn-Gutu-Khudzhirt; 130—Natalino; 131—Blagoveshchensk; 132—Innokentievka; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 167–170. Cicindela coerulea kaszabi, internal sack: 167, 169—Gulin-Tal; 168, 170—Barkull; 167, 168—right lateral view; 169, 170—left lateral view; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 190–195. Cicindela coerulea shantungensis, internal sack: 190, 193—canyon of Moudjik (= Si-He) River; 191, 194— Yangmadianzi (images by Xinxin Xiong); 192, 195—K’ing-iang-fou; 190–192—right lateral view; 193–195—left lateral view; 190, 193—partly inflated; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 233–240. Cicindela coerulea ssp., internal sack: 233, 237—C. coerulea przewalskii, Amdo; 234–236, 238–240—C. coerulea grumi (234, 238—р. БадО-хО и ХЫ-хО-дО; 235, 239—Qinghai, Wulan; 236, 240—C. coerulea kozlovi, ДабасуН ГОби); 223–236—right lateral view; 237–240—left lateral view; 233, 234, 236–238, 240—lectotypes; 236, 240—partly inflated; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 241–245. Morphometric features of Cicindela coerulea ssp: 241, 242—relation of the proportions in different subspecies (241—relation of the proportions PW/PL andEL/EW; 242—relation of the proportions LW/LLand PW/PL; ellipses— 90% confidence intervals; solid light blue circles—C. coerulea coerulea; solid dark blue circles—C. coerulea nitida; solid green circles—C. coerulea selengensis; solid lilac circles—C. coerulea kaszabi; solid red circles—C. coerulea shantungensis; open magenta circles—C. coerulea przewalskii; open yellow circles—C. coerulea grumi; open orange circles—C. coerulea kozlovi); 243–245—differences of the body proportions (243—EL/EW; 244—PW/PL; 245—LW/LL; small solid circles in the rectangles—medians; rectangles—25–75 percentiles; whiskers—min–max; open circles—outliers; asterisks—extremes; different letters indicate significant differences).