2. Coenagrion puella (Linnaeus, 1758)

Fig. 2d–k, 3d–k, 4g –k, 5g –l, 7b–g, 8a–d

Reported from: throughout the Russian part of the Caucasus, but precise reports from Kabardino-Balkaria were missing while even generalised reports from Chechnya, Ingushetia and Dagestan were scarce, as summarised by Skvortsov (2010); seven localities in Adygea Republic (Shapovalov & Korotkov 2019); reported as Coenagrion ( puella) sp1 from Bil’-bil’ Kazmalyar village in Magaramkent District of Dagestan (Ilyina et al. 2022). At the Khmelevskie Lakes in Krasnodarskiy Kray reported up to 1,875 m a.s.l. (Bartenef 1930, for more detail of the locality see Kosterin 2023).

Specimens. Karachay-Cherkes Republic: 5 ♂♂ (Figs 2k, 3k, 4j) (37681170, 109341320), 4 ♀♀ (Fig. 7g), Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, shallow, sedgy and reedy pools in the Dukka River valley left side, 43.551 –43.554 N, 41.156– 41.157 E, 1695–1715 m a.s.l., 2.07.2019 ; 2 ♂♂, Zelenchuk District, 7 km SW of Nizhniy Arkhyz, Lake Taza-Kel’ (Forelevoe) at the Bol’shoy Zelenchuk River left bank, 43.633 -43.635 N, 41.390– 41.394 E, 1260–1267 m a.s.l., 3.07.2019 ; 5 ♀♀, Zelenchuk District, Dausuz village W environs, S banks of a large water reserve at the Bol’shoy Zelenchuk River, 43.793 – 43.796 N, 41.536– 41.540 E, 1012–1014 m a.s.l., 3.07.2019 ; 3 ♂♂, 4 ♀♀, Khabez District, Inzhichishkho village N margin, big ponds and the Zelenchuk River right bank, 44.211 –44.219 N, 41.714– 41.724 E, 578–582 m a.s.l., 6.07.2019 ; Karachaevsk Municipality, Teberda Town, Lake Kara-Kel’, 43.436 –43.438 N, 41.743– 41.744 E, 1340 m a.s.l., 7.07.2019 ; 2 ♀♀, Skalistyy Mountain Range, Malokarachaevsk District, 6 km SW of Tereze village, pools with cattail at the Podkumok River right bank, 43.899 –43.900 N, 42.386 E, 1144–1146 m a.s.l., 9.07.2019 , 2 ♂♂, Karachay-Cherkes Republic, Malokarachaevsk District, Uchkeken village E environs, a chain of ponds, 43.934–948 N, 42.564–570 E, 887-923 m a.s.l., 9.07.2019 . Republic of Dagestan: ♂ (165562658), Kumtorkala District, the Shura-Ozen’ River at Kumtorkala Station, next to Sarykum Barkhan, 42.9963 N, 47.2201 E, 65 m a.s.l., 31.05.2023 ; 2 ♂♂ (Figs 2i, 3i, 5k), Buynaksk District, Buynaksk S suburbs, Nizhnee Kazanishche village environs, the Orta Kopyur Reservoir, 42.782 –42.785 N, 47.135– 47.140 E, 512 m a.s.l., 2020 (no exact date), A. Tatamov leg. ; 7 ♂♂, ♀ (178696752 (Fig. 8a), 178696754, 178696758, 180156664, 180156771, 180156920, 180157200, 180157630, 180158795, 180159019, 180159369, 180159581, 180159721, 180160108), Buynaksk District, Buynaksk S environs, Verkhnee Kazanishche village environs, a swampy, reedy brook on a gentle left slope of the Akpeozen’ River valley, 42.7982 –42.7992 N, 47.1157– 47.1169 E, 487–488 m a.s.l., 29.05.2023 ; ♂, Kazbekovskiy District, 3.5 km ENE of Dylym village, Lake Gorendzhikor, 43.0607 –43.0620 N, 46.6758– 46.6780 E, 730–732 m a.s.l., 22.06.2022, E. Ilyina leg .; ♂ (177991051), Tsumadinskiy District, Lake Bezdonnoe, 42.5702 N, 46.0162 E, 2500 m a.s.l., 5.08.2023, O. Kosterin & E. Ilyina leg .; ♂, Khunzakh District, Lake Mochokh, 42.428 N, 46.524 E, 1633 m a.s.l., 8.07.2023, E. Ilyina leg .; ♀ (Fig. 7c), Sergokala District, 3 km SW of Myurego village, the Inchkheozen’ River upper valley, 42.3738 –42.3824 N, 47.6703– 47.6726 E, 617–685 m a.s.l., 19.06.2021 ; 8 ♂♂, 9 ♀♀ (167302377, 167302379 (Fig. 8d), 181376402, 186507300, 186507352, 186507391, 186507670, 186507476, 186507776. 186508006, 186508614, 186508660, 186508720, 186508895), Gergebil’ District, at Gergebil’ Water Reservoir right bank and a brook lowermost reaches including a reed swamp, 42.4496 –42.4500 N 47.0307– 47.0328 E, 799–820 m a.s.l., 24.05.2023 ; 3 ♂♂, the same place, 10.06.2023, E. Ilyina leg.; 9 ♂♂, 4 ♀♀ (178695639, 178695641, 178695642 (Fig. 8c), 178695645, 178695647, 178695648, 178695649, 178695652, 180231199, 180231269, 180231312, 180644296, 180950215, 180952129, 180952298), Tabasaran District, the Rubas River valley 2.5 km NE of Sirtych village, a swampy Dzharag-Khyan Spring, 41.8490 –41.8523 N, 48.2397– 48.2411 E, 103–105 m a.s.l., 28.05.2023 ; ♂ (186593146), Derbent District, 3.5 km NE of Belidzhi village, a rush swamp, 41.9044 N, 48.4462 E, 18 m below s.l., 25.05.2023 ; 5 ♂♂ (Figs 2g–h. 3g–h, 4i) (196674245, 196674286, 196675373, 196675450, 196675488, 196675571, 196675621), ♀, Magaramkent District, Samur Delta Forest, Bil’- Bil’ Kazmalyar village, a brook and a pond, 41.818 –41.819 N, 48.547– 48.548 E, 21–22 m a.s.l., 25.06.2021, V. Onishko & E. Ilyina leg .; 4 ♂♂, 3 ♀♀, the same place, 4.06.2022, E. Ilyina leg; 3 ♂♂, 1 ♀, the same place, 5.05.2023, E. Ilyina leg.; 4 ♂♂, 2 ♀♀ (178693668, 178693670, 181380016, 182683558, 182683220, 182680584), the same place, 27.05.2023; ♂ (178692671, 184532980), Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, environs of Samurskiy Zakaznik Stationary, a forest road, 41.8649 N, 48.5541 E, 19 m below s.l., 26.05.202316 ♂♂ (Figs 2e–f, 3e–f, 4g–h. 5 h-i), 6 ♀♀ (Fig. 7d–f) (196674331, 196674405, 196674452, 196674504, 196674584, 196674644, 196674682, 196674741, 196674839 (Fig. 8b), 196674969, 196675116, 196675169, 196675201), Rutul District, the Samur River left arm between Kufa and Kala villages, 41.569 –41.570 N, 47.358– 47.362 E, 1367–1373 m a.s.l., 26.06.2021 .

Additional photographic registrations. Krasnodarskiy Kray: 1 teneral ♂ (37681097), Novorossiysk Municipality, Bol’shye Khutora village centre, a pond, 44.4782 N, 37.6023 E, 29.04.2019 ; ♂ (37520657), ♀ (37520692), Anapa Municipality, Sukko village E end, the Sukko River, 44.801 N, 37.437– 37.439 E, 7.06.2018 ; 1 ♂ (37520637), Anapa Municipality, Praskoveevka village, a pool by the Dzhanhot River, 44.459 N, 38.217 E, 7.06.2018 . Karachay-Cherkes Republic: a copula (109551771), Zelenchuk District, Dausuz village W environs, S banks of a large water reserve at the Bol’shoy Zelenchuk River, 43.7942 N, 41.5381 E, 1014 m a.s.l., 4.07.2019 . Kabardino-Balkarian Republic: 2 ♀♀ (37681248, 37681267), Mayskiy Town NE margin, an inundated quarry, 43.654 N, 44.100 E, 1.05.2019; ♂ (37681279), 2 ♀♀ (37681296, 37681331), the same place, 2.05.2019. Republic of Dagestan: ♂ (186509269), Russia, Republic of Dagestan, Gergebil’ District, a pool on the River left floodplain 1 km S of Maydanskoe village, 42.5819 N 46.9659 E, 559 m a.s.l., 24.05.2023 ; ♂, 2 ♀♀ (180153471, 180153628, 180648340), Tabasaran District, the Rubas River valley just downstream of the reservoir 2 km WSW of Tinit village, 41.8240 -41.8244 N, 48.1974– 48.1991 E, 180 m a.s.l., 28.05.2023 ; ♂ (181379714), the same place, 27.05.2023; 3 ♀♀ (182684591, 182771795, 182771980), Magaramkent District, Samur Delta Forest, a forest brook at the Bil’- Bil’ Kazmalyar village S margin, 41.8075 N, 48.5458 E, 21–22 m a.s.l., 27.05.2023 .

Characters. Male cercus trunk short and low, very gently rounded in lateral view, without expressed apex (Fig. 3d–k), slanting down so that tubercle is as a rule hidden behind paraproct. In dorsal view (Fig. 2d–k), trunk evenly broad, without bulge, cerci separated from each other as seen behind S10 margin and moderately divaricate; tubercle very broad and rounded. Tooth disposed almost at the level of trunk apex. Process narrower triangular than in C. australocaspicum, slightly curved inside, also reaches about the middle of the height of the paraproct (Fig. 4g –k)

Male paparoct spine variably directed slanting but more behind than up in lateral view, hooked inside at tips in dorsal view (Fig. 2d–k), thick in lateral view (Fig. 3d–k), so that incision between it and prominence is weakly expressed more or less as a blunt angle. Prominence also not so well expressed, varying from bluntly rectangular to rounded. To illustrate inter- and intrapopulation variation for the paraproct shape and spine direction, eight specimens are shown in Figs 3d–k.

Male pronotum hind margin forms three waves, with smooth, broad and shallow lateral depression and a broadly rounded, somewhat raised middle lobe. The subjective perception of the pronotum shape is strongly biased by the variable presence and shape of a blue streak along its hind margin (Fig. 5g –l), which may be interrupted at the depressions (Fig. 5k) or missing on the central lobe (Fig. 5h).

Female pronotum hind lobe like in males but with ‘three waves’ more expressed as semicircular convections; again it is seen somewhat differently in the views perpendicular to the lobe surface (anteriodorsal with respect to the insect body axis, ‘flat view’), where it looks less sinuous, and to the main body axis, where it looks more wavy (Fig. 7b–h). The variable expression of its pale rim again produces an impression of variability in shape, as illustrated by three female specimens from the population of the Rutul River valley (Fig. 7d–f).

The black marking of the male S9 varies from a posterior bar (most frequently, see Fig. 8a–d) to a pair of lateroposterior spots or two dots (not shown).

The specimens from the population of the Rutul District of Dagestan have the paraproct spine somewhat narrower and longer in lateral view (Fig. 3e–f), but other Dagestanean males do not differ from those from elsewhere. Females from that population have somewhat deeper incisions on the pronotum hind margin but not so as in C. ponticum (see below). These specimens leave an expression of being of a somewhat stouter built than C. puella from elsewhere.

Female polymorphism as elsewhere in Europe (Dijkstra et al. 2020) (Fig. 8).

Distribution and habitat. A species widespread in the Caucasus, occurring in Dagestan from below the conventional sea level in the Samur Delta Forest to 2500 m above sea level at Lake Bezdonnoe on the Snegovoy Mountain Range (where just one individual was found). It is the only Coenagrion species so far recorded in Inner Dagestan. This species was found co-occurring with five other Coenagrion spp.: with C. australocaspicum in the Samur Delta Forest (but see above) and Shura-Ozen’ River (Dagestan), with C. pulchellum at Dausuz village (Karachay-Cherkes Republic) and the Shura-Ozen’ River (Dagestan), with C. ornatum at Dausuz (Karachay-Cherkes Republic), Bil’-Bil’, Sirtych and Verkhnee Kazanishche (Dagestan) villages; with C. scitulum at Lake Gorendzhikor and Belidzhi village (Dagestan).

Coenagrion puella occurs both at lentic water bodies and slow, overgrown brooks, and even seemingly prefers water with some slow current in the Caucasus.