Cicindela (s. str.) coerulea coerulea Pallas, 1773

Matalin, Andrey V., 2025, An intraspecific structure of Сicindela (s. str.) coerulea Pallas, 1773 (Coleoptera, Cicindelidae), Zootaxa 5583 (2), pp. 201-246 : 205-212

Cicindela (s. str.) coerulea coerulea Pallas, 1773

Figs 1–9 , 14–18 , 24–44 , 65–75

Cicindela coerulea Pallas, 1773: 724 (Type locality—Irtin [= Irtysh]). = Cicindela violacea Gebler, 1817: 324 (Type locality—fl. Irtüsch), synonymized by Dejean (1833: 2). = Cicindela coerulea lucifera Mandl, 1970: 72 , Abb. 3 figs 1, 2 (Type locality—Biysk, Altai), syn. nov. Cicindela coerulea Pallas, 1773 : Pallas 1781: Tab. G fig. 12; Fischer von Waldheim 1828: 17; 1829: 5; 1842: 4; Gebler 1830:

26; 1833: 262; 1847: 268; Dejean 1833: 2; 1836: 3; Dejean & Boisduval 1837: 25 Pl. 3 fig. 4; Sturm 1843: 2; Motschoulski,

1846: 30; Motschulsky, 1850: 1; Chaudoir 1865: 36; Fleutiaux 1892: 106; Horn 1938: 46, Taf. 63 fig. 22; Rivalier 1950:

225; Schilder 1953: 564; Werner, 1991: 18, T. 14 figs 100, 101; Dubatolov et al. 1994: 4; Matalin & Zimina 2002: 230. Cicindela violacea Gebler, 1817 : Fischer von Waldheim 1820: 8, Tab. 1 fig. 4; 1821: 102. Cicindela tricolor coerulea Pallas, 1773 : Horn & Roeschke 1891: 50. Cicindela hybrida nitida Lichtenstein, 1796 : Wu 1937: 15. Cicindela hybrida coerulea Pallas, 1773 : Horn 1905a: 158; 1905b: 27; 1915: 337; 1926: 221; Winkler 1924: 4. Cicindela grumi coerulea Pallas, 1773 : Schilder 1953: 564. Cicindela coerulea tenuifascia Fischer von Waldheim, 1828 : Lorenz 1998: 43. Cicindela coerulea lucifera Mandl, 1970 : Cassola & van Nidek 1984: 10; Wiesner 1992: 118; 2020: 184; Kryzhanovskij et al.

1995: 26; Puchkov & Matalin 2003: 104; Lorenz 2005: 131; Putchkov & Matalin 2017: 227; Makarov et al. 2020. Cicindela coerulea coerulea Pallas, 1773 : Mandl, 1935: 294 figs 9, 10, 296 figs 11, 12; 1936: 9, 14; Cassola & van Nidek 1984:

10; Wiesner 1992: 117; 2020: 183; Kryzhanovskij et al. 1995: 26; Lorenz 1998: 42; 2005: 131; Puchkov & Matalin 2003:

104; Shook & Wiesner 2006: 10; Wu & Shook 2010: 77; Wu 2011: 24; Matalin & Dudko 2016a: 140; 2016b: 143; Putchkov & Matalin 2017: 227; Makarov et al. 2020; Štrunc 2020: 142.

Type material. Neotype of Cicindela coerulea Pallas, 1773 (designated here), ♂ ( Figs 1, 2 , 28 , 42 )—Irtys r. [handwritten white label], tricolor coerulea Pall. [handwritten white label], NEOTYPUS, Cicindela coerulea Pallas, 1773 , des. A. V. Matalin, 2024 [printed red label] ( ZIN ).

Paratype of Cicindela coerulea lucifera Mandl, 1970 , ♂ — Altai, Biysk, bank of Katun’ river. 16.VI.1952, Selivanovskiy [typed white label], Paratypus, Cicindela coerulea ssp. lucifera m. det. Dr. K. Mandl, 1970 [handwritten and typed red label], coll. K. Mandl [typed white label], 222, Cicindela coerulea ssp. lucifera Mandl [handwritten white label] ( NHMW).

Additional material. RUSSIA, Chelyabinsk Oblast: 1♂ —middle stream of Miass river , coll. I. Moiseenko (cISr) ; Novosibirsk Oblast: 1♂ — Karasuk Distr. , near Troitskoe, steppe, 28. VI.1967, leg. Zolotarenko ; 1♂ — Karasuk Distr., near Troitskoe, lake Krotovaya Lyacha , 2.VII.1979, leg. T.D. Gorelova (both ISEA) ; 8♂♂ 3♀♀ — Inskoy b., st. Meret , 21.VI.1898, 18.VIII.1898, 31.VIII.1898 ( ZIN, ZMUM, ISEA, NHMW) ; Tomsk Oblast: 2♀♀ — Tomsk ; 1♀ — Chernyi Yar , 16. VI.1908 (both ZIN) ; Altai Krai : 1♂ 1♀ — Sib. oc., step. Kulunda, 7. VI –19.VII.1922, leg. A. Reichardt ; 1♀ — Slavgorod Kluchi, Omsk reg., Slavgorod area , 29. V.1922, leg. A. Reichardt; 3♀♀ — Severnyi, Slavgorod area, Omsk reg., 24.VII.1922, leg. N.N. Pukhova ; 1♀ — Altai, env. Kolyvan , VII.1894, leg. F.F. Morawits ; 1♂ — Altai, Zmeinogorsk reg., env. Novenskoe (al ZIN) ; 1♂ — Lokot, Rubtsovsk reg., 8.VII.1920, N. Gorbunov ; 1♂ 1♀ — Mikhailovka Distr., env. Nikolaevka , 10.VII.1980, leg. Yu. Perunov ; 1♀ — Zapadno-Sibirskiy Krai , near Biysk , 28. V.1934; 2♂♂ 3♀♀ —near Biysk, Katun River , 15. VI –18.VIII.1952, Selivanovsky (all ZMUM) ; 1♂ 1♀ — ibid (cJG) ; 1♀ — Barnaul , 26. VI.1909, leg. B. Popov ( ISEA); 1♂ 2♀♀ —N Altai , 20 km S of Biysk, 10–13.VIII.1991, leg. M. Chernyakhovskiy ( MSPU) ; KAZAKHSTAN, North Kazakhstan Region: 2♂♂ 3♀♀ — Kuybyshev distr. , state farm Ruzaevskiy, plough-land, VI –VIII.1982, leg. I.B. Knor ( ISEA); Akmola Region: 1♀ — Selety River, Alchinbay mount., 15 verst from Gogolevsk , 9.VII.1900, leg. N. Petrovskiy ( ISEA) ; Pavlodar Region: 1♂ —E Kazakhstan, near Pavlodar , research station of Kaz. IZR, 5. VI.1961, leg. I. Kabak (cEK); 3♂♂ 3♀♀ — Pavlodar ; 3♂♂ — Maraldy-Romanovka, Pavlodar Area , Semipalatinsk Region, 3.VIII.1922, leg. A. Reichardt ; 2♂♂ 2♀♀ — Maraldy lake ; 1♂ — Aynakol, Pavlodar area, Akmolinsk region , 8.VIII.[1]922, leg. A. Reichardt (all ZIN) ; 1♂ —state farm Chernoyarka , steppe, 19. VI.1985, leg. Karabutova; 1♂ — ibid, meadow, 25. VI.1987, leg. Kozhukhanova ( ISEA); 12♂♂ 5♀♀ —NE Kazakhstan, right bank of Irtysh Riv., env. of Zhertumsyk (= Karla Marksa ) vill., 28 km SSE of Pavlodar City, 52°01’52’’N 77°11’54’’E, h=122 m, 23– 24.05.2021, leg. D.A. Zakharov (cDZ) ; Ulytau Oblast: 2♂♂ 4♀♀ —ur. Karaagash , 60 km NE Zhana-Arka, 3.VII.1961, leg. G. Medvedev, 19. V.1962, leg. L. Arnoldi (all ZIN); Abai Region: 1♂ — Talitsa, 1. V.[19]08; 3♂♂ 3♀♀ — Semipalatinsk (both ZIN) ; 1♀ — Semipalatinsk , Sibirien ( NHMW) ; 2♂♂ 1♀ — Kazakhstan, near Semipalatinsk , 15–22. V.1991, leg. S. Khvylya ( MSPU); 1♂ — Kazakhstan, 60 km NE of Semipalatinsk , 3.VII.1958 (cEK) ; 1♂ —E Kazakhstan, Semipalatinsk Area, Novopokrovka , 15.05.1972, leg. A.S. Badenko (cBK) ; 1♂ 1♀ — Semipalatinskiy leskhoz, Kashtakskoe lesnichestvo, 10.VIII.1955 ( ZMUM) ; 1♂ — Dzhertasskiy (= Djartas), Semipalatinsk oblast , 16. V.1971, leg. Merenkova ( ISEA); East Kazakhstan Region: 2♂♂ 6♀♀ — Ust-Kamenogorsk, 10. VI.30, VI.1908 ( ZIN, cPU); 1♀ —E Kazakhstan, Ust-Kamenogorsk City, bank of Irtysh Riv. , 14.06.1979, leg. V. Ryzhikh (cBK) ; 1♂ — Korosteli, Kaz. №1, Rubtsovsk Reg. , 1.VII.1923, Isaev ; 1♂ 1♀ — Berezovka, Bukhtarma river , Ust-Kamenogors Reg .; 2♀♀ — Kaznakovo , 20. V.1930, F. Zaitsev; 2♂♂ 4♀♀ — Altai mer., suso Zaysan, 1991 ; 1♂ 1♀ —Irtysch; 2♀♀ — Belaya river , S Altai, 6.VIII.1930, leg. Lyukyanovich (all ZIN) ; 5♂♂ 3♀♀ — East Kazakhstan reg., Kyzylkum sands, 19 km S Kulynzhon, h=223 m, 48.72187°N, 083.28732°E, 09.05.2022, leg. S. V. Kolov ( IZK) ; 3♂♂ 3♀♀ — Zaysan , 15 km SW of Kaznakovka, sands, 20.05.80, leg. Nikolaev; ibid, 25–31. V.89, M. Danilevsky ( ZIN, MSPU, ISEA, cJG) ; 4♂♂ 2♀♀ —N Zaysan, 300 m, V.1989 (cJW); 1♂ 1♀ — ibid ( HMHN) ; 3♂♂ 2♀♀ — N Zaysan Lake, env. of Kaznakovka vill., h=398 m, 21–31. V.1989, leg. M. Danilevsky ( ZIN, MSPU); 1♂ — ibid, (cBK) ; 339 ex. — ibid, 29.04.1997, leg. A.A. Klimenko (cAK) ; 8♂♂ 3♀♀ — ibid, 3.05.1997, leg. S.A. Toropov (cST) ; 1♂ — ibid, 9.05.2000, leg. S. Nekrashevich (cDF) ; 10♂♂ 7♀♀ — E Kazakhstan, N Zaysan Lake , 20.IX.1990, 27.V.1996, 25.V.1997, 25.V.2000, leg. A. Klimenko (cPU, MSPU) ; 1♂ —E Kazakhstan, N Zaysan, N part of Kuludzhanskie sands, 8–10. V.1999, leg. A. Korolyuk, I. Smelyanskiy; 1♀ —cape Krasnoyarsk, Irtysh River , 22. VI.1925, leg. V.I. Vereshchagin (both ISEA) ; 1♂ 1♀ — Aigyrkum desert, 5 km W of Buran vill., 26.05.2003, leg. Zhdanko (cSS); WITHOUT EXACT LOCALITY : 1♂ — Altai , foothills, VII.1979, ex. koll. G.A. (cISl) ; 2♂♂ —Altai; 1♂ 2♀♀ —Camp. Kirgiss. or.; 2♂♂ 3♀♀ —fl. Irtysh (all ZMUM) ; 1♀ — Irtysh ( ZIN) ; 1♂ — Siberie ( MNHN) .

References. RUSSIA, Tyumen Oblast: Tobolsk ( Mandl 1921); Omsk Oblast: N of vill. Malyi Atamas, bank of Irtysh River, 54.014266 N 74.941458 E, 30.V.2021,S.Knyazev(https://www.inaturalist.org/observations/81663853); Tomsk Oblast: Tomsk ( Mandl 1921); Altai Krai : Kolywan ( Mandl 1921; Wu 1937); near Mikhaylovskoe; Laptev Log; near Uglovskoe (all Matalin & Dudko 2016a); KAZAKHSTAN, Abai Region: Semipalatinsk ( Mandl 1921); Beskaragay District, 51.042429 N 79.465608 E, 18.V.2022, A. Murtazina (https://www.inaturalist.org/ observations/117848591); East Kazakhstan Region: Ust-Bukon; 20 km S Kaznakovka, Bukonskie Peski (both Matalin & Dudko 2016a); Novopokrovka ( Matalin & Dudko 2016b).

Diagnosis. The nominotypical subspecies is easily recognized among other ones by entirely lacking ( Figs 1–5 , 24–26, 28, 29, 31–35, 38, 39 ) or strongly reduced ( Figs 27, 30, 36, 37, 40, 41 ) white elytral pattern (HI = 0.0–6.0, mean = 0.4). From other subspecies, except C. coerulea nitida , the nominotypical subspecies is well distinguished by the long internal sack sharply rounded towards to the ventral side of aedeagus tube ( Figs 65–70 vs. Figs 167–170 , 190–195 , 233–240 ).

Redescription. TL = 12.5–15.0 mm (mean = 14.06 mm, n = 41) in males, 13.7–16.0 mm (mean = 14.85 mm, n = 33) in females. Specimens as a rule unicoloured blue, blue-violet or black ( Figs 1, 3, 4 ); sometimes bicolored with blue-violet head and pronotum and blue-green or green elytra, or with blue-green head and pronotum and cupreous or cupreous-green elytra with golden tinge in some specimens ( Fig. 5 ).

Head often black, violet with bluish tinge or blue with light greenish lustre, rarely blue with bright golden-green sides of clypeus, central portion of frons, vertex and occiput, as well as supraorbital plates. Labrum with 4–14 (mean = 8, n = 78) submarginal setae, LW/LL = 2.0–2.5 (mean = 2.2, n = 41) in males, 1.92–2.2 (mean = 2.07, n = 33) in females ( Figs 6–9 ). Mandibles relatively stout, ML/MW = 5.43–7.78 (mean 6.41, n = 36) in males and 6.0–8.5 (mean 7.3, n = 24) in females. Scape besides 3–5 apical setae with 1–3, rarely with 4–5 additional setae or glabrous.

Pronotum transverse, PW/PL = 1.28–1.46 (mean 1.35, n = 41) in males and 1.33–1.50 (mean = 1.41, n = 33) in females, black, violet or blue, in some specimens bluish-green with golden tinge on disk and bright blue anterior and posterior grooves and midline ( Figs 14–18 ).

Elytra with smooth punctured sculpture except slightly granular humeral area, EL/EW = 1.45–1.63 (mean 1.55, n = 41) in males and in females 1.40–1.58 (mean 1.50, n = 33), as a rule without white pattern ( Figs 1–5 , 24–26, 28, 29, 31–35, 38, 39 ); sometimes with basal portion of humeral lunula and apical portion of apical lunula ( Fig. 30 ), with narrow apical lunule separated into two small dots ( Fig. 37 ), with apical portion of humeral lunule and subsutural portion of medial band ( Fig. 41 ); or with humeral and apical lunula divided into two small spots and narrow subsutural portion of medial band ( Figs 27, 36 ).

Aedeagus relatively long ( Figs 42–44 ), EL/AL= 1.7–1.86 (mean 1.78, n = 19) with short, slightly curved basal portion and sharp, medium-sized apical hook. Internal sac long, sharply rounded towards the ventral side so В distinctly or virtually contact with the aedeagus tube, mt large with sharply curved apex, VA and DA the largest, DLL relatively large ( Figs 65–70 ).

Distribution. RUSSIA: Chelyabinsk, Tyumen, Omsk, Novosibirsk and Tomsk Oblasts, Altai Krai ; KAZAKHSTAN: North Kazakhstan, Akmola, Pavlodar, Ulytau, Abai and East Kazakhstan Regions ( Fig. 246 ).

Notes. Cicindela coerulea was described by Pallas (1773) based on collections from his trip to Western Siberia in 1771. In the original description he indicated that the beetles are bright blue, but their colouration is variable from violet-black to completely black. Additionally, Pallas indicated that this species is abundant in different sandy habitats south of Irtin (= Irtysh River). Pallas’ Coleoptera collection is housed partly in ZIN and partly in MFNB. According to Horn and his co-authors ( Horn et al. 1990) the portion of the collection of Pallas housed in St.- Petersburg was lost due to a fire. However, the only fire in the Kunstkamera occurred on the night of December 4–5, 1747 ( Hartanovich & Hartanovich 2014), and Pallas’s collection had not yet been deposited there by that time. There were no further fires that led to the destruction of collections neither in the Kunstkamera or in the Zoological Museum. Nevertheless, I could not locate a single specimen of C. coerulea from the collection of Pallas either in ZIN or in MFNB. At the same time in 1781, the illustration of C. coerulea was published ( Pallas 1781: Tab. G fig. 12). According to the Article 73.1.4 of ICZN (1999) the designation of an illustration of a single specimen as a holotype is to be treated as designation of the specimen illustrated; the fact that the specimen no longer exists or cannot be traced does not invalidate the designation. However, Pallas had a series of these specimens and mentioned Article of ICZN that cannot be applied. I believe the types of C. coerulea are lost and, according to the Article 75.3 of ICZN (1999). a neotype must be designated. Because “Irtin (= Irtysh River)” was indicated by Pallas (1773: 724) as the type area, a male specimen from the collection of ZIN labelled as “Irtys r.” ( Figs 1, 2 , 28 , 42 ) is designated here as the neotype of Cicindela coerulea Pallas, 1773 (see chapter Type material).

The elytral colouration in the nominotypical subspecies is highly variable from black through violet, blue or blue green to greenish-cupreous and bright reddish-cupreous ( Figs 24–41 ). Reddish-cupreous beetles (three males) from the environs of Biysk ( Russia, Altai Krai ) were described by Mandl (1972) as Cicindela coerulea lucifera . However, similarly coloured specimens have been observed in the other parts of the range of the nominotypical subspecies, for example in Zhana-Arka, Kazakhstan ( Figs 5 , 38, 39 ). Furthermore, in the same habitat both reddish-cupreous and bluish-green ( Figs 30, 35–37, 40, 41 ) as well as reddish-cupreous and violet-black ( Figs 5 , 26, 38, 39 ) specimens are observed. The shape of the aedeagus and the internal sack in these specimens are identical ( Figs 65–70 ). Moreover, the main body proportions in different coloured specimens do not differ significantly, except the differences between males and females ( Figs 71–75 ). Thus, a new synonymy is established: Cicindela coerulea coerulea Pallas, 1773 = Cicindela coerulea lucifera Mandl, 1970 , syn. nov.

During the current study, a male of C. coerulea coerulea with the label “Issyk-Kul” was found in the collection of ZIN. Considering the great gap between the above-mentioned locality and the main distribution area of the nominotypical subspecies, old age of this specimen as well as the lack of information on the collector, date and exact locality, I consider this record to be mislabelled.

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 1–5. Cicindela coerulea coerulea, habitus, dorsal view: 1, 2—Irtysh River; 3—Zaysan Lake, Kaznakovka; 4— Meret; 5—Zhana-Arka; 1, 2—neotype; 2—labels; 1–4—males; 5—female.

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 24–41. Cicindela coerulea coerulea, left elytron: 24—Chernoyarka; 25—Maraldy-Romanovka; 26, 38, 39—Zhana- Arka; 27—Meret; 28—Irtysh; 29—Chernyi Yar; 30, 35–37, 40, 41—Biysk; 31—Lokot; 32—Kuludzhanskie sands; 33—Zaysan Lake, Kaznakovka; 34—Ryzaevskiy; 28—neotype; 24, 25, 28, 31–33, 36, 37, 39, 41—males; 26, 27, 29, 30, 34, 35, 38, 40— 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 65–70. Cicindela coerulea coerulea, internal sack: 65, 68—Zhana-Arka; 66, 69—Chernoyarka; 67, 70—Zaysan Lake, Kaznakovka; 65–67—right lateral view; 68–70—left lateral view; without scales. The abbreviations of humps and sclerites see in the text.

FIGURES 71–75. Morphometric features of Cicindela coerulea coerulea: 71, 72—relation of the proportions in different coloured specimens (71—relation of the proportions EL/EW and PW/PL; 72—relation of the proportions LW/LL and PW/ PL; ellipses—90% confidence intervals); 73–75—differences of the body proportions (73—EL/EW; 74—PW/PL; 75—LW/ LL; small circles in the rectangles—medians; rectangles—25–75 percentiles; whiskers—min–max; solid circles—males; open circles—females; different letters indicate significant differences).

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.