Review of the Dyspessa salicicola (Eversmann, 1848) species group (Lepidoptera, Cossidae: Cossinae)
Articles
DOI: 10.14258/abs.v8.e43

Review of the Dyspessa salicicola (Eversmann, 1848) species group (Lepidoptera, Cossidae: Cossinae)

Altai State University, 61 Lenina Ave., Barnaul 656049, Russia https://orcid.org/0000-0001-9512-8709
Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab. 1, St.-Petersburg, 199034, Russia https://orcid.org/0000-0003-4735-2209
Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab. 1, St.-Petersburg, 199034, Russia https://orcid.org/0000-0002-6407-5904
Altai State University, 61 Lenina Ave., Barnaul 656049, Russia https://orcid.org/0000-0001-9367-3578
Biodiversity Carpenter Moths Dyspessa fauna Central Asia Volga-Ural Region Middle East Caucasus Transcaucasia

Abstract

The article gives a revision on the species group Dyspessa salicicola (Eversmann, 1848) (Lepidoptera, Cossidae, Cossinae). The group comprises six valid taxa, namely D. salicicola salicicola, D. salicicola aschabadensis Daniel, 1953, D. saissanica Yakovlev, 2014, D. kostjuki Yakovlev, 2005, D. arabesca Yakovlev, 2005 and D. muelleri Yakovlev, sp. nov. (type locality: “Nord Jordanien, Totes Meer-Ostufer, Wadi Ibn Hammad”), united by a specific habitus (the yellow forewing with a remarkable pattern of spots). Detailed data on the distribution of all the species are provided, short diagnoses are given. The article is illustrated with images of specimens from different localities and male genitalia of all the considered taxa.

Corresponding author: Roman V. Yakovlev (yakovlev_asu@mail.ru)

Academic editor: A. Matsyura | Received 22 November 2022 | Accepted 11 December 2022 | Published 14 December 2022

http://zoobank.org/2BC10FA5-AF54-4457-AA3D-62EA59673FF8

Citation: Yakovlev RV, Shapoval NA, Shapoval GN, Naydenov AE (2022) Review of the Dyspessa salicicola (Eversmann, 1848) species group (Lepidoptera, Cossidae: Cossinae). Acta Biologica Sibirica 8: 693–712. https://doi.org/10.14258/abs.v8.e43

Keywords

Biodiversity, Carpenter Moths, Dyspessa, fauna, Central Asia, Volga-Ural Region, Middle East, Caucasus, Transcaucasia.

Introduction

Cossus salicicola Eversmann, 1848 (Lepidoptera, Cossidae, Cossinae) was described on the basis of single specimen “ich nur ein Exemplar dieses Spinners gefangen hatte… Ich kenne bis jezt nur den Mann” (Eversmann 1848: 211; Sinev et al. 2017: 410−411). The holotype and its labels are shown in plate 13, figure 14 (Sinev et al. 2017). Spuler (1910: 303) established a new combination: Dyspessa salicicola (Eversmann, 1848). Later, Silbernagel (1944) described from Makedonia (Lake Ochrid) Dyspessa salicicola f. lutescens Silbernagel, 1944, which was later synonymized with the nominative subspecies (Daniel 1962). Daniel (1953) described the subspecies Dyspessa salicicola aschabadensis on a series of specimens from Turkmenistan (“Aschabad, Suljukli, Aidere, Merw”) and Armenia (“Eriwan”). “Aschabad” was chosen as the type locality. At the further revision of Cossidae (Yakovlev 2005, 2014) based on significant differences in the structure of antennae and male genitalia, we described three species: Dyspessa kostjuki Yakovlev, 2005, D. arabesca Yakovlev, 2005 and D. saissanica Yakovlev, 2014. In this article we analyse in detail the taxonomy of the species group Dyspessa salicicola (Eversmann, 1848), united by a specific habitus (the yellow forewing with a remarkable spotty pattern), and describe a new species D. muelleri sp. nov. from Jordan.

Material and methods

Male genitalia were mounted in euparal on slides following Lafontaine and Mikkola (Lafontaine & Mikkola 1987). The imagoes were photographed using digital camera of iPhone 7. The genitalia preparations were photographed using an Olympus DP74 camera attached to an Olympus SZX16 stereomicroscope.

Abbreviation list

AHU − collection of Armin Hauenstein (Untermünkheim, Germany);

APC − collection of Alexander N. Poltavsky (Rostov-on-Don, Russia);

HMIM − Hayk Mirzayans Insect Museum (Tehran, Iran);

MGT – Simon Janashia Museum of Georgia (Tbilisi, Georgia);

MWM − Museum of Thomas Witt (Munich, Germany);

RYB − collection of Roman Yakovlev (Barnaul, Russia);

SKO − collection of Svyatoslav Knyazev (Omsk, Russia);

VSC − collection of Valeryi I. Shchurov (Krasnodar, Russia);

ZIA − Zoological Institute of Science Academy of Kazakhstan Republic (Almaty, Kazakhstan);

ZISP − Zoological Institute of the Russian Academy of Science (Saint-Petersburg, Russia);

ZMKU − Zoological Museum at Kiev State University (Kiev, Ukraine);

ZSM − Zoologische Staatssammlung Munich (Munich, Germany).

Dyspessa salicicola (Eversmann, 1848)

Figs 1−9, 27−31

Cossus salicicola Eversmann, 1848, Bull. Soc. Imp. Nat. Moscou 21 (3): 211.

Type locality: Saratovischen Gournevement, in der Gegend von Wolsk [Russia, Saratov Reg., Volsk]. Type material (holotype) in ZISP.

Cossus salicicola salicicola Eversmann, 1848

Synonymy:= Dyspessa salicicola f. lutescens Silbernagel, 1944, Z. Wien. Ent. Ges. 29: 187.

Type locality: Ochrid (Ochrid lake) [Macedonia].

Type material (holotype) in MWM.

Distribution: Bulgaria, Macedonia, Albanien, Greece, Ukraine, SW Russia, Transcaucasia, Turkey, Сentral Kazakhstan (Romanoff, 1885; Spuler, 1910; Daniel, 1962; Didmanidze, 1978; Didmanidze & Zurashvili, 1981; de Freina, 1983, 1996; de Freina & Witt, 1990; Gevorkian, 1986; Yakovlev, 2004, 2005; Didmanidze & Yakovlev, 2007; Yakovlev, 2009, 2011; Yakovlev et al., 2015, 2020; Yakovlev & Witt, 2016; Streltzov et al. 2022). Additionally, some localities in Popasnaya and Novoaydar districts of Lugansk Region (Ukraine) (https://www.inaturalist.org/observations/127394200; https://www.inaturalist.org/observations/63887630; https://www.inaturalist.org/observations/102396142) and Griechenland, Ditiki Makedonia, Siatista (western Kozani) (https://www.inaturalist.org/observations/62710006) are provided on iNaturalist.

Material examined. Russia: male (holotype), Wolsk, 9 Juny, coll. Eversmann (ZISP); 1 male, 1 female, Rostov Reg., Krymskij vill., 7.vi.2010, leg. E. Khachikov (RYB); 1 male, Beta, viii.[19]28, O. Chernova (ZISP); 2 males, Kerch (ZISP); 3 males, Stavropol, 12.vi.920, Filipiev (ZISP); 2 males, Taganrog, 15.vi.1817 (ZISP); 7 males, Groznyi, 17.v.908, Rodnensky leg. (ZISP); 8 males, Mashuk Mt., 9.vi.908 (ZISP); 1 male, Gelendzhik, 9.vi.908 (ZISP); 2 males, Stavropol’, 12.vi.920, Filip’jev leg. (ZISP); 2 males, [Dagestan], Tarki, 2.vii.945, Rjabov leg. (ZISP); 1 female, Caucasus boreal, Fl. Kuma, 21.v.[18]90, Kozhannchikov (ZISP); 2 males, Neftekumsk (MWM); 2 males, Derbendt (MWM, ZISP); 1 male, 1 female, Rostov Don Reg., Manytch Gudilo lake (MWM); 2 males, Daghestan, Kapchugaj (ZMKU); 1 male, Gigant, 17.vi.2014 (APC); Rassvet, 9 males, 23–25.v.2013, 6 males, 12–18.vi.2014, 3 males, 2.vii.2014 (APC); 4 males, 1 female, Sober-Oaschkh, 19.vi.1999, 5 male, 2 females, 29.vi.2006 (VSC); 2 males, Janina Gorge, 26.vi.2000, 11 males, 1 female 27.v.2008 (VSC); 6 males, 1 female, Jakhno, 13–15.vi.2003, 11 males, 13.vii.2003, 1 male, 3.vi.2005, 3 males, 2 females, mass flying in light trap, 31.v–1.vi.2007 (VSC); 1 male, Shize, 22.vi.2004 (VSC); 1 female, Voronezhskaja, 31.vi.2006 (VSC); 3 male, Lysaja 3, 31.v.2007, 3 males, 20.vi.2007, 1 male, 5.vii.2010 (G. Schemberger) (VSC); 4 males, Akhilleon, 13.vi.2009 (VSC); more than 30 individuals for 150 minutes, Shakhan, 17.vi.2009 (VSC); 3 males, mass flying in light trap, Oblego, 18.vi.2009 (VSC); 3 males, Leninskiy Put, 25.06.2009 (VSC); 1 male, Suzdal’skaja, 26.vi.2009 (VSC); 1 male, 1 female, Gajduk, 3.vi.2010 (VSC); 2 males, Red October, 23.vi.2011 (VSC); 1 male, Sapun, 22.vii.2011 (VSC); 3 males, 1 female Lysaja 3, 26.vi.2004, 3 males, 7.07.2014 (VSC); 1 male, Il’skaja, 16.vi.2014, 1 male, 29.vi.2014 (E. Zhukov) (VSC); 3 males, Russia, Saratov Reg., Khvalynski distr., 5 km W Khvalynsk, Dacha Khrenova, 902 m, 4−10.vii.2011, leg. V.V. Anikin (RYB); 6 males, 1 female, Saratov Reg., Khvalynski distr., 5 km W Khvalynsk, Dacha Khrenova, 902 m, 1−10.vii.2017, leg. V.V. Anikin (RYB); 1 male, Volgograd region, Olkhovskyi district, Mikhaylovka, 7.vii.2018, leg. S. Legotin, M. Elistratov & S. Didenko (RYB); 8 males, Guberli (ZISP); 8 males, Russia, S. Ural, Orenburg Reg., Guberlya village, 22.vi.2003, leg. V. Zurilina (RYB); 29 males, 3 females, S. Ural, Orenburg Reg., 12.vi.1999, V. Zurilina leg. (RYB); 13 males, 2 females, S. Ural, Orenburg Reg., 12.vi.1997, V. Zurilina leg. (RYB); 18 males, 1 female, Russia, S. Ural, Orenburg Reg., near Kuvandyk, 21−23.vi.2004, V.O. Zurilina leg. (RYB); 8 males, Russia, S. Ural, Orenburg Reg., near Kuvandyk, 17.vi.2016, V.O. Zurilina leg. (RYB); 22 males, Russia, S. Ural, Orenburg Reg., near Kuvandyk, 25−29.v.2019, V.O. Zurilina leg. (RYB); 13 males, Russia, S. Ural, Orenburg Reg., near Kuvandyk, 13−18.vi.2020, V.O. Zurilina leg. (RYB); 3 males, Russia, S. Ural, Orenburg Reg., near Kuvandyk, 26−29.vi.2021, V.O. Zurilina leg. (RYB); 2 females, Bashkortostan Rep., Yantysh village, 19.vi.2007, leg. V. Zurilina (RYB). North Macedonia: 2 males, Mazedonien, Ochrid, 700 m, 23−27.vi.1936, coll. V. Bartha (MWM, ZSM); 2 males, Mazedonien, Ochrid, 700 m, 22.vi.1959, G. Thurner (MWM, ZSM); 18 males, Mazedonien, Drenovo bei Kavadar, 1−9.vi.1957, Thurner leg. (MWM, ZSM); 20 males, Stari Doiran, 150−300 m, Daniel leg (MWM); 1 male, Katlanovo, 10.vi.1979, Schaider (MWM). Bulgaria: 1 female, Kozuch, 10.vi.1981, leg. Ganev (MWM); 1 male, Mičurin-lozenec, 22.vi.1971, V. Felix legit (MWM). Greece:16 males, 3 females, Nordgriechenland, Olympos Gebiet, Litohoron, 500 m, 12−23.vi.1957, leg. Daniel (MWM); 1 female, Peloponnes, Zachlerou, Mitte vi.1959, Demelt leg. (MWM); 2 males, Peloponnes, Chelmosgeb. 700 m, 19−20.vi.1960, Thurner leg. (MWM). Crimea: 1 male, Bakhchisaraj distr., Sokolinoe vill., 900−1100 m, 6−11.vii.2014, leg. V. Anikin (RYB); 17 males, Feodosiya ditsr., Kurortnoe vill., 8.vi.1997, E. Rutjan (RYB); 2 males, Feodosiya ditsr., Kurortnoe vill., biostation, 44°54ˊ49.08ˊˊN 35°12ˊ6.48ˊˊE, 7−8.v.2016, leg. S. Knyazev (RYB); 1 female, Agarmysh (ZISP); 1 male, Burulcha, 17.v.902 (ZISP); 1 male, Alupka, 20.vi.18[94], Melioranski (ZISP); 1 male, Sudak (ZISP); 2 males, [Crimea], Belbek station, v.1907, N.Y. Kuznetsov (ZISP); 8 males, Staryi Krym, 14.vi.1913, S. Tschetverikov (ZISP); 1 male, Alsu, Krim, 28.vi.[19]07, W. Pliginski (ZISP); 3 males, Sebastopol, 2.v.[19]06, W. Pliginski (ZISP); 2 males, Simferopol, 6−8.vi.1920, V. Kuznetsov (ZISP). Ukraine: 1 male, Odessa (ZISP). Armenia: 2 males, Ararat Prov., Khosrov Reserve, 2000 m, 01.vii.1996, S. Andreev (RYB); 1 male, Ararat Prov., 7 km SW Surenavan vill., 1300 m, N39°48ˊ36ˊˊ; E 44°50ˊ8ˊˊ, 4.vii.2011, P. Romantsov leg. (RYB); 3 males, Ararat Prov., Khosrov Reserve, Garni, N 40°06ˊ01ˊˊ; E 44°46ˊ47ˊˊ, 1600−1800 m, 22−27.vi.2010, A.V. Krupitskiy leg. (RYB); 1 male, Vayots Dzor Prov., 12 km SW Ehegnadzor, 3 km NW Noravank Monastery, 1250 m, 39°42ˊ12ˊˊ N; 45°12ˊ26ˊˊ E, 8.vii.2011, leg. P. Romantsov leg. (RYB); 14 males, Garni, 28.vi.2000, leg. Kazaryan (RYB); 6 males, Ararat district, Dvin village, Yeranos Mts., 1600 m, 11−15.vi.2009, leg. Yu. Shevnin (RYB); 26 males, Yeghegnadzor suburbs, Aiolzdzorsky Range, 150 km SE Erevan, 2000 m, 1−27.vii.2009, leg. Yu. Shevnin (RYB); 5 males, Erivan, 15.vi.[19]28, Ryabov (ZISP); 12 males, Armenia, Inaklyu, 18.vii.1936, Ryabov (ZISP); 1 male, Leninakan [Gyumri], 7.vii.1957 (ZISP). Georgia: 1 male, South Ossetia, Tskhinval Distr., 2 km NW Grom, 42°10′6″ N / 44°11′53″ E, 930 m, 22−25.06.2021, A. Streltzov, P. Ustjuzhanin & R. Yakovlev leg. (RYB); 3 males, Kakheti Prov., Sagarejo distr., 41°29ˊ41ˊˊN; 45°17ˊ32ˊˊE, 750 m, 28.vi.2013, leg. E. Rutjan (RYB); 6 males, 2 females, Tiflis, 14.vi.1907 (MGT); 4 males, Achtala, 17.vi.1910 (MGT); 1 female, Shida Kartli, the Saguramskii reserve, 550–600 m, 7.vii.1970, E. A. Didmanidze leg. (MGT); 5 males, Kaheti, the Vashlovanskii reserve, 10.v.1973, 30.vi.1973, 31.vii.1973, 29.vi.1974, E.A. Didmanidze leg. (MGT); 5 males, 1 female, Kaheti, Shirak-Eldari Steppe, 10.v.1973, 28.vi.1973, E. A. Didmanidze leg. (MGT); 1 male, Meskhet-Dzhavakheti, Hando vill., near Ahalkalaki, 14.vii.1964, E. A. Didmanidze leg. (MGT); 2 males, Chachuny, Tiflis g., 19.v.[18]96, Mlokosevich (ZISP); 3 males, Kodzhory, Tiflis gub., 17−18.vi.916, Andrievski (ZISP); 1 male, Lagodekhi, 10.v.[18]93, Mlokosevich (ZISP); 3 males, Borzhom (ZMKU). Azerbaijan: 17 males, Helendorf [Goygol] (ZISP); 1 male, Lerik, 2.vii.1964, leg. A. Zagulyaev (ZISP); 2 males, Kusarchaj vill., 4.vi.1964, Pastukhov (ZISP); 1 male, Talysh, Astara distr., 8 km SW Gomushan vill., 1700 m, 28.vi.2003, I. Pljustch (RYB); 1 male, Talysh, 30 km SE Masalli, 800 m, 3.vi.2005, leg. V. Tikhonov (MWM). Turkey: 1 male, Ankara, Kizilcahaman, 1000 m, 28.v.71, leg. et coll. Kunz (MWM); 2 males, 1 female, Kars, vic. Kagizman, 10 km NW Kötek, 1750 m, leg. de Freina (MWM); 1 male, Agri, nördl. Cumachay, 1900-2100 m, 18.-19.vii.80, 1 cf , leg. de Freina (MWM); 1 male, Agri, 7 km E Agristepe, 2200 m, 20−22.vii.1990, leg. Gabor Ronkay (MWM). Kazakhstan: 1 male, 50 km NE Aktobe, Karabulak river, 50º53´N / 57º54´E, 370 m, 29.vi.2007, leg. P. Gorbunov (MWM); 5 males, Karaganda reg. Bektau-Ata Mts., 47.46°N / 74.79°E, 581 m, 2.vi.2015, S.V. Titov & M. Černila leg.(RYB); 6 males, Karaganda Province, Takrau River, Saryterek aul, 700 m, 48°05´N / 75°18´E, 29.v.2015, leg. P. Gorbunov (RYB); 1 male, [Akmola reg.], Kokshetau hills near Terisakkan river, 2.vii.1958, leg. Falkovich (ZISP); 1 male, Karaganda Reg., Zhana-Arka distr, 6.vi.1959, leg. Aibasov (RYB); 5 males, Karaganda Province, Zhaksy-Arganaty mountain, 49°19´N / 67°05´E, 19.vi.2010, P. Gorbunov (RYB); 1 male, Karaganda Reg., Zhana-Arka distr, Kok-Sengir Mts., 10.vi.1959, leg. Aibasov (ZIA); 1 male, Kazakhstan, Karagandy Reg., Bektau-Ata Mts., 70 km N of Balkhash, 47°26´52.20´´N / 74°41´5.92´´E, 7.vii.2018, at light, S.A. Knyazev leg. (SKO).

Figure 1.Figures 1–15. Dyspessa salicicola (Eversmann, 1848), imago males: 1. Cossus salicicola Eversmann, 1848, holotype (ZISP); 2. D. s. salicicola, Northern Macedonia (MWM); 3. D. s. salicicola, Greece (MWM); 4. D. s. salicicola, Crimea (MWM); 5. D. s. salicicola, Armenia (MWM); 6. D. s. salicicola, Armenia (MWM); 7. D. s. salicicola, Azerbaijan (MWM); 8. D. s. salicicola, Turkey, Agri Province (MWM); 9. D. s. salicicola, Turkey, Nevsehir Province (MWM); 10. D. s. aschabadensis, Paratype, Turkmenistan (MWM); 11. D. s. aschabadensis, Turkmenistan (MWM); 12. D. s. aschabadensis, Turkmenistan (MWM); 13. D. s. aschabadensis, Iran, Zanjan Province (MWM); 14. D. s. aschabadensis, Iran, Mazandaran Province (MWM); D. s. aschabadensis, Iran, Alborz Mountains (MWM).

Diagnosis. Extremely variable subspecies, distinctive in its big size (length of male forewing 11−14 mm, female 13−15 mm). Antenna bipectinate, setae 2−2.5 times longer than antenna stem diameter (in medium third). Forewing light-yellow, frame mottled, series (3–5) dark-brown spots discally. In rare cases, this series of discal spots completely reduced. Hindwing without pattern, colour from light-grey to grey. Valve apically lanceolate, with expressed crest on costal margin, proximal edge of crest sharply expressed, distal edge smooth; phallus 1/3 shorter than valve, poorly curved, gradually narrowing to apex (Figs 28−31).

Taxonomical remarks. We believe that the opinion of Daniel (1953, 1962) on the attribution of the Armenian populations to the subspecies D. s. aschabadensis Daniel, 1953 is incorrect. A large comparative material, including the serially studied male genitalia clearly showed the consubspecificity of Armenian (and other Transcaucasian and Turkish) populations with a nominative subspecies. There are minor differences in the wing pattern, the Armenian populations are characterized by a more stable spotty pattern on the forewing, than the specimens from the southern Urals, which have a very variable wing pattern (Figs 40−41).

Dyspessa salicicola aschabadensis Daniel, 1953

Figs 10−15, 32−33

Daniel, 1953, Mitt. Münch. Ent. Ges. 43: 260.

Type locality: Asia centr., Aschabad [Turkmenistan].

Type material (holotype) in ZSM.

Distribution: Turkmenistan, Iran (Daniel 1962; Yakovlev & Witt 2016; Alipanah et al. 2021).

Material examined. Turkmenistan: 2 males (paratypes), Merw, 7.vii. (MWM); 3 males (paratypes), Ashkhabad, coll. Tancré (MWM); 2 males (paratypes), Achal-Tekke (MWM); 1 male, Kara-Kala, 14−17.v.1993 (MWM); 5 males, Kopet-Dagh Mts., 10 km S of Aidere, 27.vi.1992, 600-1000 m, 56°46E, 38°14N, leg. B. Fabian, B. Herczig, A. Podlussany and Z. Varga (MWM); 7 males, Kopet-Dagh Mts., 1000 m, Sayvana valley, cca 5 km SW of Saivana, 56°50E, 38°17N, 28.vi.1992, leg. B. Fabian, B. Herczig, A. Podlussany and Z. Varga (MWM); 1 male, Dushak, 1800 m, 23.vi.1992, leg. M. Danilevsky (MWM); 1 male, Kara-Kala distr., N slope of Syunt Mt., 12.v.1953, Kuznetsov (ZISP). Iran: Alborz Prov.: 1 male, 6 km E Âzadbar- Tâleghân Rd., 2350 m, 21.vii.1988, leg. Mirzâyâns, Badii (HMIM); Ardebil Prov.: 1 male, Ardebil, Namin, 1860 m, 20.vi.2002, leg. Hâjiesmailiân, Mofidi-Neyestânak, Deuve, Montreuil (HMIM), 1 male, Neur Lake, 2500 m, 26.viii.2010, leg. Khâni, Mofidi-Neyestânak (HMIM), 1 male, Hashtchin- Khalkhâl Rd., N 37˚26΄54.5˝, E 48˚24΄50.6˝, 837 m, 16.vii.2013, leg. Afsariân, Sarafrâzi (HMIM); East Âzarbâijân Prov.: 1 male, Kaleybar, Ghaleh Bâbak, 1500 m, 5.vii.1997, leg. Mofidi-Neyestânak, Barâri (HMIM); Gilân Prov.: 1 male, Eshkevar, Gilânchâkân, 1820 m, 27.vi.1997, leg. Barâri, Mofidi-Neyestânak (HMIM), 1 male, Rudbâr, Darreh Dasht, 900 m, 25.vi.2002, leg. Hâjiesmailiân, Serri, Mofidi-Neyestânak, Deuve, Monteuil (HMIM); Golestân Prov.: 2 males, 2 females, Golestân National Park, Dasht, 960 m, 18.vi.1977, leg. Pâzuki, Abâi (HMIM), Golestân National Park, Ghush Cheshmeh, 1000 m, 29.vi.2001, leg. Ebrâhimi, Osten, Mofidi-Neyestânak (HMIM), 2 males, Golestân National Park, Koylar, 1250 m, 23.vii.1996, leg. Ebrâhimi, V. Nazari (HMIM), 1 male, Golestân National Park, Koylar, 1350 m, 29.vii.2001, leg. Gilâsiân, Moghaddam, Ghayurfar (HMIM), 2 males 1 female, Golestân National Park, Mazârli, 350 m, 19.–20.vi.1977, leg. Pâzuki, Abâi (HMIM), 5 males, 1 female, Golestân National Park, Tang-e Gol, 620 m, 23.–25.v.1986, leg. Pâzuki (HMIM), 1 male, Golestân National Park, Tang-e Gol, 700 m, 24.–27.vii.1996, leg. Ebrâhimi, V. Nazari (HMIM), 1 female, Golestân National Park, Tang-e Râh, 930 m, 7.v.2000, leg. Badii, Ebrâhimi, Mofidi-Neyestânak (HMIM), 1 male, Khânbebin, Shir Âbâd, 80 m, 5.vii.2001, leg. Mofidi-Neyestânak, Osten, Ebrâhimi (HMIM), 2 males, Âgh Ghalâ, Chinsivly, 0 m, 2.vii.2001, leg. Ebrâhimi, Osten, Mofidi-Neyestânak (HMIM), 3 males, Gorgân, Loveh forest, 10.viii.1967, leg. Abâi, 4 males, Tuskastân, Gorgân- Shâhrud Rd., N 36˚44΄37.6˝, E 54˚34΄02.6˝, 660 m, 15.vii.2007, leg. Zahiri, Karimzâdeh (HMIM); Khuzestân Prov.: 1 male, Izeh, Susan, 600 m, 26.iv.1995, leg. Badii, Ardeh, Parchami-Arâghi (gen. prep. No. HA-2386, HMIM); Fârs Prov.: 1 male, Firuz Âbâd, Ghir, 11.iv.1975, 1160 m, leg. Borumand (HMIM); North Khorâsân Prov.: 14 males, Golestân National Park, Âlmeh, 1600 m, 26.–29.v.1986, leg. Pâzuki (HMIM), 8 males, Golestân National Park, Âlmeh, 1650 m, 17.–25.v.1988, leg. Pâzuki (HMIM), 3 males, Golestân National Park, Âlmeh, 1590 m, 6.v.1999, leg. Mofidi-Neyestânak, Barâri, Manzari (HMIM), 16 males, Golestân National Pârk, Sulgerd, 1150 m, 8.v.1999, leg. Moghaddam, Barâri, Manzari (gen. prep. No. HA-1926, HMIM), 2 males 1 female, Yâkhti Kalân, 2.vi.1986, leg. Pâzuki; Mâzandarân Prov.: 1 male, Marzan Âbâd, 400 m, 6.vii.1995, leg. Badii, Sarafrâzi, Linnavori, 1 male, Siâhbisheh, 10.vi.1974, leg. Abâi (HMIM), 1 male, Behshahr, Pasand, Pasand Res. Sta., 5.vi.1981, 6.vii.1981, no collector given (HMIM); West Âzarbâijân Prov.: 1 male, Mâku, Âvâjigh, Bâshkand, 1400 m, 16.vi.1976, leg. Pâzuki, Borumand (HMIM); Zanjân Prov.: 1 male, Elburz Mt., Gharâvol Dâgh, 2500 m, 3,4.vii.1999, leg. Hofmann, Meineke (HMIM), 2 males, Zanjân- Gilvân Rd., Gharâvol Dâgh, 2500 m, 9, 26, 27.vi.2001, leg. Hofmann, Meineke, Tremewan (HMIM), 1 male, Armaghân Khâneh, Mâri vill., N 36˚59΄52.0˝, E 48˚28΄39.0˝, 2300 m, 21.vi.2015, leg. Nâserzâdeh, Hâjiesmailiân, Montreuil (HMIM); 2 males, Iran, Zanjan, Kuh-e-Sindan (= Sandân Mt.), 25 km NE Zanjan, 2200 m, 15.vii.1997, leg. Eckweiler & Ten Hagen (MWM); 7 males, NO Persien, Khorassan (= Khorâsân), Abassabad (= Abbâs Âbâd) NP, 1200 m, 12.v.?, leg. de Freina (MWM); Mazandaran, Mt. Sah (= Shâhkuh), Azedsahr (= Âzâd Shahr which is located in Golestân Province) distr., 1000 m, 2.vi.1999, leg. Gasko (MWM); Mazandaran prov., 35 km S Amol, 700 m, 14.v.1998, leg. Fabian & Szekely (MWM); 2 males, 15 km S of Azad Shar (=Âzâd Shahr), 28–29.vi.2000, leg. K. Gasko (MWM); 2 males, Azarbaigan-e, Sarked (an erroneous spelling of Sharghi= East Azarbâijân), Dizaj, 15.vi.2000, leg. K. Gasko (MWM).1 male, road Tehran-Chalus, Marzanabad area, 2 km SE Tovir v., 900 m, 23.vi.2010, N 36˚22΄26˝; E 51˚17΄23˝, leg. K. Kolesnichenko (RYB); 2 males, Golestan, 20 km S Gorgant (= Gorgân), 900 m, 2.vi.2009, leg. K. Kolesnichenko (RYB); N. Persia, Astrabad, 26.v.1905, Filippovich (ZISP).

Diagnosis. Subspecies differs in relatively small size (length of male forewing 10−13 mm, females not examined). Forewing yellow or dark-yellow, with mottled frame and series (from 3 to 5) dark-brown spots discally, often fusing into band, spots bigger than in nominative subspecies; root, discal and postdiscal area almost always with dense sputtering of grey or brown scales. Hindwing without pattern, colour from light-grey to grey. Valve apically lanceolate, crest on costal margin more expressed than in nominative subspecies, proximal and distal margins of crest sharp, well-defined; phallus 1/3 shorter than valve, poorly curved, gradually narrowing to apex (Figs 32−33).

Taxonomical remarks. We provisionally define all the Iranian specimens as the subspecies D. s. aschabadensis (Alipanah et al. 2021). Probably, the populations from north-western portion of Iran can belong to the nominative subspecies, this suggestion needs a separate study.

Dyspessa saissanica Yakovlev, 2014

Figs 16−18, 34−35

Yakovlev, 2014: 657.

Type locality: E. Kazakhstan, Kurchum distr., Bukombaj Mts., 48º13´N / 84º43´E.

Type material. Holotype in ZISP, examined.

Distribution: E. Kazakhstan, Zaisan Valley; North-Western China (Yakovlev et al. 2016).

Material examined. Kazakhstan. Holotype, male, E. Kazakhstan, Kurchum distr., Bukombaj Mts., 550 m, 48º13´N / 84º43´E, 9−10.vi.2012, leg. R. & A. Yakovlev (ZISP); paratype, male, Zaisan (SW part of Altai), coll. A. Meinhardt (ZISP). Not type material: 16 males, E. Kazakhstan, Altai Mts., Kurchum distr., 26 km SE of Barak-Batyr vill., unnamed mountain massif, gravelly slopes, 610 m, 48º29´14´´'N / 84º07´02´´E, 20.vi.2014, Volynkin A.V. & Titov S.V. lg. (RYB); 14 males, 1 female, North-East Kazakhstan, South Altai., Kyzyltas Mts., 6 km W of Kyzyltas vill., 620 m, 48°16´N / 85°26´E, 6.vi.2016, leg. P. Gorbunov (RYB); 1 male, Eastern Kazakhstan, Saur Mts, 20 km SEE Zaisan, 47°22´N / 85°09´E, 1225‒1250 m, 20.vi.2018, R. Yakovlev, V. Doroshkin, V. Rudoj & A. Naydenov leg. (RYB); 1 female, Eastern Kazakhstan, Saur Mts, Kenderlik river Valley (left bank), 2 km W Kenderlik village, 24.vi.2018. 47°28´N / 85°12´E, 740 m, R. Yakovlev, V. Doroshkin, V. Rudoj & A. Naydenov leg. (RYB). China: 1 male, Chamil-hami, Tancre (ZSM).

Diagnosis. Length of forewing 13−15 mm, light-coloured, forewing creamy, with thin grey frame (in D. salicicola, frame mottled), with poorly noticeable series of dark spots discally (often reduced), hindwing light-grey. Valve more narrow than that of D. salicicola, crest on costal margin of valve less expressed, phallus 1/4 shorter than valve.

Length of forewing in females 14−15 mm. Forewing light-yellow, almost without pattern; hindwing dark-grey.

Dyspessa kostjuki Yakovlev, 2005

Figs 19−21, 36

Eversmannia 3/4: 22.

Type locality: Ukraine, “Proval’skaya Stepp” Naturschutzgebiete.

Type material (holotype) in MWM.

Distribution: SE Ukraina, Crimea and SW Russia (Yakovlev 2005, 2007, 2011; Yakovlev et al. 2015; Suchkov 2019).

Material examined. Ukraine: Holotype (male), Ukraine, “Proval’skaya step’” Naturschutgebiete, licht, 11.vii.1997, I. Kostjuk leg. (MWM); 19 males (paratypes), same data (MWM); 1 male (paratype), Ukraine, Nikolaevskaja-Gebiet, Pervomajskij-Bezirk, Kuziptschino, 20.vi.1992, leg. A. Bidzilja & Kljutschko (MWM); 1 male, Donetsk Reg., Khomutovskaya Steppe, 15.vi.1975, leg. A. Korovin (RYB); 1 male, 1 female, Donetsk Reg., Amvrosievka, 15.vi.1975, leg. A. Korovin (RYB). Crimea: 2 males (paratypes), Ukraine, Krim, Karadag, 4.vii.1995, leg. Z.F. Kljutschko; 10 males (paratypes), Ukraine, Crimea, Karadagh, 16.-24.vi.1993, leg. Budashkin (MWM); 15 males (paratypes), same data, 30.v.−20.vi.1994, leg. Budashkin (MWM); 4 males (paratypes), Ukraine, Krym, Karadagh, 24.vi.1974, Z. Klyuchko (AHU); 1 male (paratype), Krym, Karadagh, 6.vii.1994, leg. Budashkin (AHU); 2 males (paratypes), Krym, Feodosia distr., Kurortnoe vill., 8.vi.1997, E. Rutjan leg. (AHU); 9 males, Simferopol distr., Krasnolesie vill., balka Tavelchuk, 26.v.2013, leg. V. Savchuk (RYB); 10 males, Bakhchisaraj distr., Nauchnyi vill., Sel-Bukhra Mt., 11.vi.2013, leg. V. Savchuk (RYB); 2 males, Feodosia distr., Kurortnoe vill., 13.vi.1997, E. Rutjan leg. (RYB); 1 male, Tarkhankut, Dzhangul, 15.vi.1996, A. Belov leg. (RYB). Russia: 9 males (paratypes), Russland, Rostov-Don, 40 km W Nedwigovka, 12.vi.1993, ex coll. Schintlmeister (MWM); 2 males, Dagestan Rep., Rutul district, Rutul vill., Lalaan riv., 1400 m, 31.v.2021, leg. V. Zurilina (RYB); 1 male, Dagestan Rep., Urban district of Makhachkala, Talgi vill., 8.vi.2021, leg. V. Zurilina (RYB); 1 male, 8 km S Makhachkala (MWM); 1 male, Daghestan, Makhachkala city, Karaman-2 distr., 25.vi.2009 (APC); 1 male, Daghestan, Makhachkala city, Karaman-2 distr., 21.vi.2011 (APC); 1 male, Bessergenevskaja, 3.vi.2009 (APC); 1 ex., Volochaevsky, 7.vii.2009 (APC); Gigant, 1 male, 7.vi.2011, 2 males, 4.vi.2012 (APC); 2 males, Kamensk, 9.vi.2008 (APC); Konygin, 1 males, 16.vi.1993, 3 males, 2–18.vi.2007, 1 male, 5.vi.2010 (APC); 2 males, Kuibyshevo, 20.v.2009 (APC); 4 males, Nesvetay, 9.vi.2008 (APC); 4 males, Nizhnesazonov, 22.vi.2011 (APC); 7 males, Olympia-camp, 6–8.vi.2011 (APC); Rasswet, 1 male, 29.v.2013, 6 males, 9–30.vi.2013 (APC); 1 male, Ternovoy, 10.vi.2012 (APC).

Diagnosis. Length of forewing in males 9−12 mm. Setae 1.5 times longer than antenna stem diameter (in its medium part), gap between setae processes wider than the process thickness. Forewing light-yellow or ochre, with mottled frame and fringe, postdiscally on wing – series of small, sometimes poorly expressed 2−4 dark round spots. About one third of specimens have completely reduced spots (particularly, pattern reduced in specimens from eastern Ukraine). Hindwing grey, fringe light-yellow. Valve extended in medium third, with trapezoidal process on costal margin. Phallus thin, poorly curved, twice shorter than valve.

Length of forewing in female 9−11 mm, pattern resembles that of male.

Dyspessa arabeska Yakovlev, 2005

Figs 22−24, 37

Eversmannia 3/4: 23.

Type locality: Asia min., Turcia, 50 km östl. Istanbul.

Type material (holotype) in MWM.

Distribution: Western Turkey (Istanbul prov.).

Material examined. Male (Holotype) Asia min., Turcia, 50 km östl. Istambul, 6.vii.65, 50 m/ü.M., leg. M. u W. Glaser (MWM). Paratypes: 8 males, same data, GenPr MWM: 9823, 9822, 9812 (MWM); 4 males, Asia min., Turcia, Gebze (Izmit), 2 u. 3.vi.1969, leg. F. Hahn (MWM).

Diagnosis. Length of forewing in males 8–10 mm. Setae equal to antenna stem diameter (in medium third). Forewing light-yellow, with mottled yellow frame and fringe, on wing postdiscally – series of elongated brown spots forming longitudinal stripes, often fused between each other and making bands, brown stroke in discal cell, similar stroke underlining discal cell from below. In a few specimens, thin brown stroke along costal margin of fore wing. Hindwing grey, fringe light-yellow. Valve extended in middle, crest on costal margin poorly expressed, semicircular. Phallus 1/3 shorter than valve, poorly curved.

Figure 2.Figures 16–27. Dyspessa, imago specimens: 16. D. saissanica, male, Holotype (ZISP); 17. D. saissanica, male, NE Kazakhstan (RYB); 18. D. saissanica, female, NE Kazakhstan (RYB); 19. D. kostjuki, male, Holotype (MWM); 20. D. kostjuki, male, Ukraine (RYB); 21. D. kostjuki, female, Ukraine (RYB); 22. D. arabesca, male, Holotype (MWM); 23. D. arabesca, male, Paratype (MWM); 24. D. arabesca, male, Paratype (MWM); 25. D. muelleri, male, Holotype (MWM); 26. D. muelleri, male, Paratype (MWM); 27. D. salicicola salicicola, female, Greece (MWM).

Dyspessa muelleri Yakovlev, sp. nov .

http://zoobank.org/3844667C-9EA8-40EA-9F94-ED8644CD47A0

Figs 25−26, 38−39

Material: Holotype (male), “Nord Jordanien, Totes Meer-Ostufer, Wadi Ibn Hammad, minus 360 m, 30.v.1999, leg. G. Müller” (GenitalPräparat Heterocera MWM: 28.232).

Paratype: 6 males, “Nord Jordanien, ca. 20 km W Irbid, Wadi et Taiyba, Jordan graben, 100 m, 18.vi.1999, leg. G. Müller” (GenitalPräparat Heterocera MWM: 28.231).

Diagnosis. The new species is similar to the small species D. kostjuki and D. arabesca. The poorly expressed pattern of the forewing is similar to that of D. kostjuki, but the new species reliably differs in the male genital structure: the valve is not extended in the medium third and the bump on the costal margin of the valve is poorly expressed.

Description. Length of forewing in male 9−12 mm. Antenna bipectinate, setae 1.5 times longer than antenna stem diameter (in medium third). Forewing ochre-yellow, with mottled frame and series of poorly expressed blurred brown spots (2−4 in number) postdiscally, fringe yellow. Hindwing grey, without pattern.

Male genitalia. Uncus triangular, with poorly sclerotized apex; gnathos arms thick, of medium length; gnathos poorly structured; valve lanceolate, poorly curved, costal margin (on border between medium and distal thirds) with small bumpy process, sclerotized more than the rest of valve surface; transtilla process short, poorly curved, basally very thick, apically curved as a hook; juxta small, saddle-like; saccus semicircular, quite robust; phallus 1/3 shorter than valve, almost straight, poorly narrowing to apex, vesica aperture in dorso-apical position, in length about 1/3 of phallus, vesica without cornuti.

Female unknown.

Distribution. Northern Jordan.

Etymology. The new species is named after the well-known German entomologist, researcher in tropical medicine and fauna of Lepidoptera of Eurasia and Africa, Dr. Günter Müller (Munich).

Discussion

Members of the Dyspessa salicicola species group are widespread in western Palaearctic, the eastern limit of its distribution range lies in eastern Kazakhstan and north-western China (D. saissanica). D. salicicola is the most widely distributed species of the group, however the question about the boundaries of its subspecies’ ranges remains open. It seems promising to conduct a molecular genetic study of this species group representatives to confirm the status of the taxa indicated here, as well as to clarify the subspecies affiliation of D. salicicola populations from the Transcaucasus and Turkey.

Figure 3.Figures 28–33. Dyspessa salicicola (Eversmann, 1848), male genitalia: 28. D. s. salicicola, Russia, Southern Ural, Orenburg Reg., Kuvandyk (RYB); 29. D. s. salicicola, Russia, Southern Ural, Orenburg Reg., Kuvandyk (MWM); 30. D. s. salicicola, Armenia, Garni (MWM); 31. D. s. salicicola, Turkey, Agri Province, Cumaçay (MWM); 32. D. s. aschabadensis, Turkmenistan, Kopet-Dagh, Kara-Kala (MWM); 33. D. s. aschabadensis, Iran, Alborz Mts., Chalush Region (MWM).

Figure 4.Figures 34–39. Dyspessa, male genitalia: 34. D. saissanica, Holotype (ZISP); 35. D. saissanica, NE Kazakhstan, Southern Altai (RYB); 36. D. kostjuki, Paratype, Crimea (MWM); 37. D. arabesca, Paratype (MWM); 38. D. muelleri, Holotype (MWM); 39. D. muelleri, Paratype (MWM).

Figure 5.Figures 40–41. Series of males of Dyspessa salicicola (RYB): 40. S. Ural, Orenburg Region, near Kuvandyk; 41. Armenia, Yeghegnadzor suburbs, Aiolzdzorsky Range, 150 km SE Erevan.

Figure 6.Figure 42. Map showing distribution of Dyspessa salicicola species group.

Acknowledgements

The authors are indebted to Mr. Thomas Witt† and Mr. Harald Sulak (Museum Witt, Munich), Sergei Sinev (Sankt-Petersburg), Helen Alipanah (Tehran), Axel Hausmann and Günter Müller (Munich), Eteri Didmanidze (Tbilisi), Igor Kostjuk and Igor Pljushtch (Kiev), Armin Hauenstein (Untermünkheim), Svyatoslav Knyazev (Omsk), and Valeryi I. Shchurov (Krasnodar) for their kind assistance provided during the study of the deposited material, and to Anna Ustjuzhanina (Tomsk) for language improvements.

This study was supported by the grant of the Russian Science Foundation No. 22-24-00265 “Cossoid Moths (Lepidoptera, Cossoidea) of tribus Endagriini Duponchel, 1844 of the World: systematic, phylogeny and evolution of the group”, https://rscf.ru/project/22-24-00265/

References

Alipanah H, Yakovlev R, Falsafi H, Witt Th, Saldaitis A (2021) Cossidae (Lepidoptera) of Iran: a review with description of two new species. Zootaxa 5062, 100 pp. https://doi.org/10.11646/zootaxa.5062.1.1

Eversmann E (1848) Beschreibung einiger neuen Falter Russlands. Bulletin de la Société Impériale des Naturnalistes de Moscou 21(3): 205−232.

Daniel F (1953) Neue Heteroceren. Mitteilungen der Münchner Entomologischen Gesellschaft 43: 256–261.

Daniel F (1962) Monographie der palaearktischen Cossidae. VI. Genus Dyspessa Hbn.Erster Teil. Mitteilungen der Münchner Entomologischen Gesellschaft 52: 1–38.

Didmanidze EA (1978) Lepidoptera of arid landscapes of Georgia (Heterocera). Tbilisi, 318 p. (in Russian).

Didmanidze EA, Yakovlev RV (2007) Cossidae (Lepidoptera) of Georgia. Entomofauna 28(1): 1–16.

Didmanidze EA, Zurashvili TM (1981) Materials on study of Macrolepidopteraof Vashlovanskii Reserve. Zapovedniki Gruzii 5: 76–118 (in Russian).

Freina JJ de (1983) 4. Beitrag zur systematischen Erfassung der Bombyces- und Sphinges-Fauna Kleinasiens. Neue Kenntnisse über Artenspektrum, Systematik und Nomeklatur sowie Beschreibungen neuer Taxa. Mitteilungen der Münchner Entomologischen Gesellschaft 72: 57–127.

Freina JJ de (1996) Cossidae. In: Karsholt O, Razowski J (Eds.) The Lepidoptera of Europe. A distributional Checklist. Apollo Books, Soro, 129–130.

Freina JJ de, Witt TJ (1990) Die Bombyces und Sphinges der Westpalaearktis (Insecta, Lepidoptera). Bd. 2. Cossoidea, Hepialoidea, Pyraloidea, Zygaenoidea. Edition Forschung & Wissenschaft, München, 134 p.

Kemal M, Koçak AÖ (2009) Revised Checklist of the Lepidoptera of Turkey. PRIAMUS Serial Publication of the Centre for Entomological Studies Ankara, Supplement 17: 1−253.

Lafontaine JD, Mikkola K (1987) Lock–and–key system in the inner genitalia of Noctuidae(Lepidoptera) as taxonomic character. Entomologiske Meddelelser 55: 161–167.

Romanoff NM (1885) Les Lepidopteres de la Transcaucasie. Deuxième partie. Mémoires sur lesLépidoptères par N. M. Romanoff 2: 1–118.

Sinev SY, Anikin VV, Zolotuhin VV (2017) Volga-Ural Bombyces described by E. Eversmann. Proceedings of the Museum Witt Munich 7: 409−411.

Silbernagel A (1944) Die Schmetterlinge der Ochrid-Gegend in Macedonien. Zeitschrift der Wiener Entomologischen Gesellschaft 29: 184−187.

Spuler A (1910) Die Schmetterlinge Europas, 2. E. Schweizerbart, Stuttgart, 523 pp.

Streltzov AN, Ustjuzhanin PYa, Morozov PS, Naydenov AE, Spitsyn VM, Yakovlev RV (2022) Lepidoptera of South Ossetia (Northern Transcaucasia). Part II. Cossidae, Limacodidae, Erebidae (Lymantriinae, Arctiinae, Syntominae, Notodontinae), Lasiocampidae, Lemoniidae, Saturniidae, Sphingidae, Drepanidae and Cimeliidae. Acta Biologica Sibirica 8: 647−654.

Suchkov SI (2019) Additional data to the Lepidopterofauna of Reserve “Kamennye Mogily”. Biodiversity of the steppe zone of Ukraine: study, preservation, playback. Drukarskii dvir, Slovyansk: 157−158 (in Ukrainian).

Yakovlev RV (2004) Carpenter-Moths (Lepidoptera, Cossidae) of Siberia. Euroasian entomological journal 3(2): 155–163 (in Russian).

Yakovlev RV 2005 New data on distribution and systematic of Cossidae (Lepidoptera) of Europe and adjacent territories. Eversmannia 3−4: 18−27 (in Russian).

Yakovlev RV (2007) Carpenter moths (Lepidoptera: Cossidae) of Russia. Eversmannia 9: 11−33 (in Russian).

Yakovlev RV (2011) Catalogue of the Family Cossidae of the Old World. Neue Entomologische Nachrichten, 66, 1–129.

Yakovlev RV (2014) Carpenter moths (Cossidae) of Altai mountain country. Zoologicheskii zhurnal 93(5): 651−661 (in Russian).

Yakovlev RV, Poltavsky AN, Ilyina EV, Shchurov VI, Witt Th (2015) Cossidae (Lepidoptera) of the Russian Caucasus with the description of a new species. Zootaxa 4044(2): 270–288. http://dx.doi.org/10.11646/zootaxa.4044.2.5

Yakovlev RV, Saldaitis A, Pekarsky O (2016) A new species of Dyspessa Hübner, (Lepidoptera, Cossidae) from Western China, with catalogue of Chinese species of the genus. Zootaxa, 4107(1): 085–088. http://doi.org/10.11646/zootaxa.4107.1.5

Yakovlev RV, Shapoval NA, Ivonin VV, Knyazev SA, Kuftina GN, Masharskiy AE (2020) A new species of Carpenter Moths (Lepidoptera, Cossidae) from Tarbagatai (NE Kazakhstan) and Altai (SW Siberia, Russia) Mountains. Zootaxa 4896(1): 085–095. https://doi.org/10.11646/zootaxa.4896.1.3

Yakovlev RV, Witt TJ (2016) Carpenter-moths of Turkmenistan (Lepidoptera: Cossidae). Biological Bulletin of Bogdan Chmelnitskiy Melitopol State Pedagogical University 6(3): 164–173.