Research Article |
Corresponding author: Maxim Yu. Proshchalykin ( proshchalikin@biosoil.ru ) Academic editor: Thorleif Dörfel
© 2023 Yulia V. Astafurova, Maxim Yu. Proshchalykin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Astafurova YuV, Proshchalykin MYu (2023) The genus Epeolus Latreille, 1802 (Hymenoptera, Apidae) in Central Asia. ZooKeys 1181: 241-263. https://doi.org/10.3897/zookeys.1181.110416
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Available information about bees of the genus Epeolus in Central Asia is summarized. Twenty species are currently known from this area. Two new species are described: E. albus Astafurova & Proshchalykin, sp. nov. and E. pesenkoi Astafurova, sp. nov. Two species are newly recorded from Central Asia: E. asiaticus Astafurova & Proshchalykin, 2022 and E. nudiventris Bischoff, 1930. The hitherto unknown male of E. mikhailovi Astafurova & Proshchalykin, 2021 is described, and lectotypes are designated for E. ruficornis Morawitz, 1875 and E. vinogradovi Popov, 1952.
Anthophila, Apiformes, cleptoparasites, Palaearctic region, taxonomy
Central Asia as understood here covers the territories of Kazakhstan, Uzbekistan, Kyrgyzstan, Turkmenistan, and Tajikistan and has an area of about 4 million square kilometres, roughly half the size of Europe (Fig.
The genus Epeolus Latreille, 1802 includes more 120 species spread across much of the globe; they occur throughout the Holarctic region, from the west coast of the United States and from Japan to Europe and North Africa (
Epeolus transitorius was the first species of the genus described from Central Asia (
Epeolus species | Kazakhstan | Uzbekistan | Kyrgyzstan | Turkmenistan | Tajikistan |
---|---|---|---|---|---|
E. albus | + | + | + | ||
E. alpinus | + | ||||
E. asiaticus | + | + | |||
E. cruciger | + | + | + | + | |
E. gorodkovi | + | ||||
E. julliani | + | ||||
E. kyzylkumicus | + | + | + | ||
E. laticauda | + | + | + | + | |
E. mikhailovi | + | + | + | ||
E. mongolicus | + | ||||
E. nudiventris | + | + | + | + | + |
E. pesenkoi | + | + | + | ||
E. productulus | + | + | |||
E. rasnitsyni | + | ||||
E. ruficornis | + | + | + | + | + |
E. seraxensis | + | + | + | ||
E. tarsalis | + | ||||
E. transitorius | + | + | + | ||
E. variegatus | + | ||||
E. vinogradovi | + | ||||
Total : | 15 | 10 | 7 | 7 | 9 |
A key to Central Asian Epeolus has not been included in this paper; it is forthcoming in a subsequent publication uniting this and the entire Palaearctic fauna due to their extensive sharing of species and the need for some additional work in these regions.
The results presented in this paper are based on 354 Epeolus specimens currently housed in the Zoological Institute, Russian Academy of Sciences (St. Petersburg, Russia,
Morphological terminology follows that of
Abbreviations T and S are used for metasomal tergum and metasomal sternum, respectively.
The species are listed alphabetically. We have used the following abbreviations for collectors: MP – M. Proshchalykin; SB – S. Belokobylskij; VG – V. Gurko; VP – V. Popov; VR – V. Rudolf.
Specimens were studied with an Olympus SZ51 stereomicroscope, and photographs were taken with a combination of stereomicroscope (Olympus SZX10) and digital camera (Olympus OM-D). Final images are stacked composites generated using Helicon Focus v. 7.7.4 Pro. All images were post-processed for contrast and brightness using Adobe Photoshop. New distributional records are noted with an asterisk (*).
Epeolus Latreille, 1802: 427. Type species: Apis variegata Linnaeus, 1758, monobasic.
Trophocleptria Holmberg, 1886: 233, 275. Type species: Trophocleptria variolosa Holmberg, 1886, monobasic.
Epeolus (Diepeolus) Gribodo, 1894: 79. Type species: Epeolus giannellii Gribodo, 1894, monobasic.
Epeolus (Monoepeolus) Gribodo, 1894: 80. Type species: Apis variegata Linnaeus, monobasic.
Pyrrhomelecta Ashmead, 1899: 66. Type species: Epeolus glabratus Cresson, 1878, by original designation.
Argyroselenis Robertson, 1903: 284. Type species: Triepeolus minimus Robertson, 1902, by original designation.
Oxybiastes Mavromoustakis, 1954: 260. Type species: Oxybiastes bischoffi Mavromoustakis, 1954, by original destination.
Holotype
: ♂, Uzbekistan: Kashkadarya Region, Yrta-Bulak, Sundukli, 16.V.2015, M. Proshchalykin, M. Mokrousov [
Epeolus albus sp. nov. resembles E. vinogradovi Popov, 1952, E. flavociliatus Friese, 1899, E. ruficornis Morawitz, 1875, E. subrufescens Saunders, 1908, and E. warnckei Bogusch, 2018 in sharing the axillae with a pair of long, acute, curved teeth (free portion of axilla), reaching posterior margin of mesoscutellum or longer. Of them, the new species is most similar to E. subrufescens (Northern Africa, Middle East, and Turkey), with which it shares a pair of well-developed posteriorly directed processes (tubercles) between medial depression of the mesoscutellum (Fig.
Epeolus albus Astafurova & Proshchalykin, sp. nov., male, holotype (A–I), paratype (J) A–C habitus, lateral (A); dorsal (B) and ventral (C) views D head, frontal view E labrum, frontal view F pygidial plate, dorsal view G, H mesoscutellum, dorsolateral (G) and dorsal (H) views I hind basitarsus, lateral view J genitalia, dorsal view. Scale bars: 1.0 mm (A–C); 0.5 mm (D); 0.25 mm (E–J).
In E. vinogradovi axillar teeth are more elongate and strongly curved; the mesoscutellum is with a pair of long, truncate teeth (Fig.
Male (holotype). Total body length 6.0 mm; forewing length (without tegula) 4.5 mm. Structure and sculpture: head (Fig.
Integument coloration: head mostly black, but mandibles (excluding dark apex), labrum, clypeus on lower half, scape and antennae yellowish. Mesosoma mostly black; pronotal lobe, tegulae, axillar teeth, and legs (with pale spurs) yellowish; wings slightly infuscate, stigma light-brown, veins from yellow to brown. Metasomal terga black with pale and transparent marginal zones. Sterna brownish with yellow marginal zones. Pygidial plate yellow.
Pubescence: body with well-developed, snow-white tomentum of thick, plumose setae covering mostly of integument (Fig.
Female. Structure, sculpture, and pubescence are similar to those of male (Fig.
The specific name “albus” (from Latin, meaning white) is associated with the extremely well-developed white pubescence of the body in the new species.
Desert areas in Kazakhstan, Uzbekistan, and Turkmenistan.
Epeolus alpinus Friese, 1893: 34, ♀, ♂ (type locality: Goeschenen, Switzerland).
Epeolus variegatus Thomson, 1872 (nom. praeocc., nec Linnaeus, 1758): 212, ♀ (type locality: unknown).
Epeolus glacialis Alfken, 1913: 36, nomen novum for E. variegatus Thomson, 1872.
Epeolus montanus Bischoff, 1930: 9, ♀, ♂ (type locality: Warnemünde, Germany).
Epeolus pilosus Bischoff, 1930: 9–10, ♀, ♂ (type locality: Rositten [=Rybachij], Kaliningrad Prov., Russia).
Epeolus alpinus Bischoff, 1930 (nom. praeocc., nec Friese, 1893): 9–10, ♀ (type locality: Saas, Switzerland).
Kazakhstan, 1 ♂, Chagan-Obo, Saur, Semipalatinsk distr., 13.VI.1910, leg. B. Karavayev [
Kazakhstan; North Africa, Europe, Turkey, Iran, Russia (European part, Urals, Western and Eastern Siberia, Far East), Mongolia.
Epeolus asiaticus Astafurova & Proshchalykin, 2022a: 309, ♀, ♂ (type locality: Terkhin-Gol, Chulut and Khoit Rivers, Mongolia).
No records in Central Asia.
Kazakhstan, 1 ♀, Semipalatinsk, coll. F. Morawitz (
*Kazakhstan, *Kyrgyzstan; Russia (Western and Eastern Siberia), Mongolia.
Nomada crucigera Panzer, 1799: 20, ♂ (type locality: Austria).
Epeolus rufipes Thomson, 1870: 91, ♀ (type locality: S Sweden).
Epeolus similis Höppner, 1899: 355–356, ♀, ♂ (type locality: Freisenbüttel, Germany).
Epeolus cruciger var. elegans Müller, 1921: 168, ♀ (type locality: Arnswalde, Germany).
Epeolus cruciger var. rufiventris Müller, 1921: 168, ♀ (type locality: Arnswalde, Germany).
Epeolus marginatus Bischoff, 1930: 11, ♀, ♂ (type locality: Warnemünde, Germany).
Kazakhstan, 1 ♀, Kyzylorda, Perovsk district, Kara-Uzyan, 15.IV.16, leg. N. Pulikovskaya [
Kazakhstan, Uzbekistan, Kyrgyzstan, Turkmenistan; Europe, Turkey, Syria, Iran, Russia (northeast to Magadan Prov.), Mongolia.
In Central Asia Epeolus cruciger is common only in northern Kazakhstan, but in the remaining Central Asian territories this species is rare and occurs mostly in the mountains. Here females have a mostly well-developed red pattern in the integument coloration and yellowish pubescence.
Epeolus gorodkovi
Astafurova in
No additional specimens examined.
Tajikistan; Afghanistan.
Epeolus julliani Pérez, 1884: 318–322, ♀ (type locality: Marseille, France).
No additional specimens examined.
Kazakhstan; North Africa, Middle East, Europe, Caucasus, Russia (south of European part, southern Urals), Iran.
Epeolus kyzylkumicus
Astafurova in
No additional specimens examined.
Kazakhstan, Uzbekistan, Tajikistan.
Epeolus laticauda Bischoff, 1930: 13, ♂ (type locality: Mondy, Buryatia Republic, Russia).
Tajikistan, 1 ♂, 15 km SW of Shaartuz, 2.VI.1982, leg. SB [
Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan; Russia (Eastern Siberia).
Epeolus mikhailovi Astafurova & Proshchalykin, 2021a: 30, ♀ (type locality: Kyrgyzstan, Toguz-bulak).
Uzbekistan, 1 ♀, Samarkand, coll. F. Morawitz [
Structure, sculpture, coloration and pubescence are similar to those of the female (see
Mountains of *Uzbekistan, Kyrgyzstan, and Tajikistan.
Epeolus mongolicus Astafurova & Proshchalykin, 2021b: 19, ♀ (type locality: 40 km SW of Uliastay, Zavkhan, Mongolia).
No additional specimens examined.
Kyrgyzstan; Mongolia, Russia (Eastern Siberia).
Epeolus nudiventris Bischoff, 1930: 14, ♀, ♂ (type locality: Mondy, Buryatia Republic, Russia).
No records in Central Asia.
Kazakhstan, 2 ♂♂, Kazalinsk, 4.VI.1900, leg. S. Berg [
*Kazakhstan, *Uzbekistan, *Kyrgyzstan, *Turkmenistan, *Tajikistan; Mongolia (Khovd), Russia (Eastern Siberia).
Holotype
: ♀, Kazakhstan: 25 km SSW of Dzhansurov [Zhansugirov], Aksu River, Zhungarskiy Alatau, 1600 m, 26.VII.1986, Yu. Pesenko [
Kyrgyzstan, 2 ♀♀, 3 ♂♂, Kungei-Alatau, 2200 m, 6–10.VII.1999, 8–9.VII. 2001, leg. Z. Klyuchko [
This species belongs to the E. variegatus species group [E. compar Alfken, 1938, E. eriwanensis Bischoff, 1930, E. intermedius Pérez, 1884, E. turcicus Bogusch, 2018, E. productulus Bischoff, 1930, E. productuloides Bogusch, 2018, and E. variegatus (Linnaeus, 1758)] in sharing the presence of labral tubercles positioned close to the middle of labrum (as opposed to apically or subapically) and the curved (as opposed to straight) S5 of the female. From other species of the group, the new species can be distinguished by the uninterrupted albeit medially narrowed apical bands of the metasomal terga and usually bright yellow tomentum on the body of the female.
Female (holotype). Total body length 7.0 mm; forewing length (without tegula) 5.5 mm. Structure and sculpture: head (Fig.
Integument coloration: head mostly black, but mandibles (excluding dark apex), labrum, clypeus on lower half, scape and F1 amber (yellow-reddish). Mesosoma mostly black; pronotal lobe, tegulae, axillae, mesoscutellum, and legs (with dark spurs) amber; wings infuscate, stigma, and veins brown. Metasomal terga mostly black, but amber on posterior half of T5; marginal zones apically yellowish and transparent. Pygidial plate amber. Sterna (including marginal zones) amber.
Pubescence: pale tomentum mostly golden-yellow (Fig.
Male. Structure, sculpture, coloration, and pubescence are similar to those of female (Fig.
Female. Total body length 5.5–7.5 mm. Coloration of clypeus varies from entirely amber to dark brown/black along upper margin. The supraclypeal area is sometimes reddish on lower frontal keel. The metasomal terga are usually black (except amber on posterior half of T5) but may be reddish or brownish laterally along the marginal zones and on the posterior part of T1. The pubescence of the mesoscutum is usually as described in the holotype but sometimes has dense tomentum obscuring much of the anterior part instead of the usual distinct paramedian strips. Two specimens from the type locality have narrowly interrupted apical bands of tomentum. Male. Total body length 5.0–7.0 mm. The labrum varies from black to reddish. The basal band of tomentum on T1 is usually uninterrupted and is often almost merged with the apical band (except a small central area), rarely interrupted medially. Coloration of pale tomentum is usually less bright than in the female, from yellow to whitish.
The specific epithet is dedicated to Yuri Andreevich Pesenko (1944–2007), a renowned melittologist and, during his lifetime, one of the leading experts on the systematics of halictid bees.
Mountains of Kazakhstan, Uzbekistan, and Kyrgyzstan.
Epeolus productulus Bischoff, 1930: 4, ♂, ♀ (type locality: Sarepta, Volgograd Prov., Russia).
Schwartz et al. 1999: 486 (Kazakhstan, Uzbekistan).
Kazakhstan, 3 ♀♀, Ber Tschogur, Mugodjar Mts, 3.VI.1910, coll. L. Wolmann [
Kazakhstan, Uzbekistan; Central and South Europe, Turkey, Syria, Ukraine, Russia (south of European part, Orenburg Prov.).
Epeolus rasnitsyni Astafurova & Proshchalykin, 2021c: 11, ♂ (type locality: the mouth of the Shakhdara River, Tajikistan).
No additional specimens examined.
Tajikistan (Gorno-Badakhshan Autonomous Region).
Epeolus ruficornis
Morawitz, 1875: 144, ♀, ♂ (type locality: Ayni, Tajikistan). Lectotype (designated here): ♀, 11.VI.[1870] // Варзаминоръ [Varzaminor (= Ayni), Tajikistan // Epeolus ruficornis Mor. [handwritten by F. Morawitz], [illustrated on Fig.
Kazakhstan, 3 ♀♀, 3 ♂♂, Bayrakum near Dzhulek, 10.VI.1907, leg. L. Wolmann [
The species is closest to Epeolus warnckei Bogusch, 2018, but differs by having poorly developed labral teeth (Fig.
*Kazakhstan, *Uzbekistan, Kyrgyzstan, Turkmenistan, Tajikistan, Mongolia (Dornogovi, Khovd, Uvurkhangai), Azerbaijan, China (Xinjiang, Gansu).
Epeolus transitorius var. seraxensis Radoszkowski, 1893: 54–55, ♀, ♂ (type locality: Serax, Turkmenistan).
Epeolus seraxensis:
Kazakhstan, 3 ♀♀, Charyn River, 10 km NW of Chundzha, 22–23.VII.1988, leg. M. Volkovish [
Kazakhstan, Turkmenistan, Tajikistan, Azerbaijan, Iran, Israel.
Epeolus tarsalis Morawitz, 1873: 182–183, ♀, ♂ (type locality: Dagestan, Russia).
Epeolus praeustus Pérez, 1884: 324–326, ♀ (type locality: Pyrenees).
?Epeolus sibiricus Radoszkowski, 1887: 295, ♀, ♂ (type locality: Vladivostok, Russia).
Epeolus rozenburgensis van Lith, 1949: 105–112, ♀ (type locality: the Netherlands).
Epeolus himukanus Hirashima, 1955: 40–41, ♂ (type locality: Kyushu, Japan).
Epeolus tarsalis ssp. tirolensis van Lith, 1956: 99, ♀ (type locality: Tirol, Austria).
No additional specimens examined.
Kazakhstan, Europe, Russia (European part, Eastern Siberia, Far East), Mongolia, Korea, Japan.
Epeolus transitorius
Eversmann, 1852: 102, ♀ (type locality: Indersk District, Atyrau Province, Kazakhstan;
No additional specimens examined.
Kazakhstan, Uzbekistan, Tajikistan; Greece, Georgia, Ukraine, Russia (southern European part, southern Ural, Western Siberia).
Apis variegata Linnaeus, 1758: 577, ♂ (type locality: Sweden).
Apis notata Christ, 1791: 188–189, ♀, ♂ (type locality: Germany).
Apis pulchella Christ, 1791: 194–195, ♀, ♂ (type locality: Germany).
Apis muscaria Christ, 1791: 195–196, ♀, ♂ (type locality: Germany).
Apis festiva Christ, 1791: 190–191, ♀, ♂ (type locality: Germany).
Epeolus pictus Nylander, 1848: 174–175, ♀, ♂ (type locality: Siberia, Russia).
Epeolus productus Thomson, 1870: 91, ♀, ♂ (type locality: Sweden).
Kazakhstan, 1 ♀, Akmola, 6 km NE of Imkty-kol’ Lake, 21.VI.1957, leg. VR [
Kazakhstan, North Africa, Europe, Russia (to Eastern Siberia), Turkey, Georgia, Kazakhstan, Iran, Pakistan, Mongolia.
In Central Asia Epeolus variegatus is only known from northern Kazakhstan, where it is a common species. Reports of this species from other Central Asian countries are not confirmed by our examined material. In Russia, according to currently available data, the species is distributed east to the province of Irkutsk. A record from Yakutia by
Epeolus vinogradovi
Popov, 1952: 108, ♀, ♂ (type locality: Dzhebel, Turkmenistan). Lectotype (designated here): ♀, ст. Джебел, Туркмен. [Dzhebel, Turkmenistan], 12.VI.[1]934, В. Попов [V. Popov leg.] // к. Ф. Моравица // Epeolus vinogradovi sp.n. Holotype! ♀, Popov det. 1935 [illustrated on Fig.
Turkmenistan, ♀ (paralectotype), Молла-кара, бл. Джебела, Туркм. [Molla-Kara, Dzhebel, Turkmenistan], 1.VII. [1]934, В. Попов [leg. V. Popov] // Epeolus vinogradovi sp.n. Paratype ♀, Popov det. 1935; 1 ♂ (paralectotype), Mulla Kara // Epeolus spinosus F. Morawitz [by Morawitz] // Epeolus vinogradovi sp.n. ♂, Popov det. 1935; 1 ♂ (paralectotype), Репетек [Repetek], 3.VI.27, Н. Умнов [leg. N. Umnov] // Epeolus vinogradovi sp.n. Paratype ♂, Popov det. 1935; 1 ♂, Uzun-Ada, 23.V.1896, leg. Varentzov; 2 ♂♂, Kara-Bogaz, 40 km N of Kyzyl-Arvat, 3.VII, 18.VI.1953, leg. Kryzhanovskiy; 1 ♀, Repetek, 8–13.VI.1976, leg. V. Kaplin.
Turkmenistan.
In total, 20 species are recorded from Central Asia, or, in other words, almost half of the Palaearctic fauna of the genus. For comparison, 18 species are known from Europe, 16 from the Middle East, 12 from North Africa, nine from Mongolia, 12 from Siberia and the Russian Far East, five from Japan, and only four from China.
The core of the Epeolus fauna of Central Asia is formed by seven endemic species (E. albus, E. gorodkovi, E. kyzylkumicus, E. mikhailovi, E. pesenkoi, E. rasnitsyni, and E. vinogradovi) and four species distributed from Central Asia to Mongolia and southern Siberia (E. asiaticus, E. laticauda, E. mongolicus, and E. nudiventris). Of them, Epeolus laticauda is a fairly common species in desert biotopes. Epeolus gorodkovi, E. mikhailovi, and E. rasnitsyni occur mostly in the mountains.
Nine species are more widely distributed. Epeolus julliani, E. productulus, and E. transitorius are widespread in the Palaearctic. Epeolus ruficornis and E. seraxensis are distributed in arid territories of the Central Palaearctic, from the Middle East and Caucasus to Central Asia and Mongolia. Only four species recorded in Central Asia are widespread in the whole Palaearctic region (E. alpinus, E. cruciger, E. tarsalis, and E. variegatus), but in Central Asia they are rare (E. alpinus, E. cruciger, and E. tarsalis) or distributed only in northern part (E. variegatus).
We are grateful to Maximilian Schwarz (Ansfelden, Austria), Esther Ockermueller (
The authors have declared that no competing interests exist.
No ethical statement was reported.
This investigation was supported by the Russian Funds for Basic Research (grant number 20-54-44014) and the state research projects 122031100272-3 and 121031000151-3.
All authors have contributed equally.
Yulia V. Astafurova https://orcid.org/0000-0003-0557-7792
Maxim Yu. Proshchalykin https://orcid.org/0000-0001-7870-8226
All of the data that support the findings of this study are available in the main text.