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Research Article
Review of the Epeolus julliani species group (Hymenoptera, Apidae, Epeolus Latreille, 1802), with descriptions of two new species
expand article infoYulia V. Astafurova, Maxim Yu. Proshchalykin§
‡ Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia
§ Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far East Branch of the Russian Academy of Sciences, Vladivostok, Russia
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Abstract

The nine species of the Epeolus julliani species group from the Palaearctic region are reviewed. Two new species are described and illustrated: Epeolus rasmonti Astafurova & Proshchalykin, sp. nov. (Russia, Mongolia, China) and E. kyzylkumicus Astafurova, sp. nov. (Central Asia). Epeolus julliani Pérez, 1884 and E. laticauda Bischoff, 1930 are newly recorded from Kazakhstan and E. seraxensis Radoszkowski, 1893 is newly recorded from Kazakhstan and Tajikistan. An identification key for both sexes of all members of this species group is presented.

Keywords

Anthophila, Apiformes, cleptoparasites, Palaearctic region, taxonomy

Introduction

In recent years, significant progress has been made towards a better understanding of the taxonomy of the species of Epeolus Latreille, 1802 from the Palaearctic region, in particular Europe (Bogusch and Hadrava 2018; Le Divelec 2021), the Middle East and North Africa (Bogusch 2021), Turkey (Bogusch 2018), and Central Asia and Mongolia (Astafurova and Proshchalykin 2021a, b, c, 2022). In total, about 45 species are now known from the Palaearctic. Based on specimens from several collections, the present work aims to complement these studies by providing a review of the Epeolus julliani species group.

The julliani species group includes Epeolus fasciatus Friese, 1895, E. iranicus Bogusch, 2021, E. julliani Pérez, 1884, E. laticauda Bischoff, 1930, E. seraxensis Radoszkowski, 1893, E. siculus Giordani Soika, 1944, and E. transitorius Eversmann, 1852.

Here, we add two new species to this group: Epeolus rasmonti Astafurova & Proshchalykin, sp. nov. from the East Palaearctic and E. kyzylkumicus Astafurova, sp. nov. from Central Asia.

This paper is meant as a further step towards a better documentation of the species of Epeolus and their distribution patterns in the wider Palaearctic region and adjacent areas. As previous years have shown, a greater number of undescribed species can be expected, particularly in the eastern, central and southern Palaearctic, where so far relatively little material has been available for study.

Materials and methods

The results presented in this paper are based on 257 specimens in the Epeolus julliani species group currently housed in the Zoological Institute, Russian Academy of Sciences (St. Petersburg, Russia, ZISP); Zoological Museum of the Moscow State University (Moscow, Rus­sia, ZMMU); Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far Eastern Branch of Russian Academy of Sciences (Vladivostok, Russia, FSCV); and Oberösterreichisches Landesmuseum, Biologiezentrum (Linz, Austria, OLBL).

The taxonomy and synonymy of species generally follow those of Bogusch and Hadrava (2018), except we regard E. julliani and E. transitorius as separate species, following Le Divelec (2021). Morphological terminology follows that of Michener (1944, 2007) and Engel (2001). The density of integumental punctures is described using the following formula: puncture diameter (in μm) / ratio of distance between punctures to average puncture diameter, e.g., 15–20 μm / 0.5–1.5.

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: AF – A. Fateryga, MP – M. Proshchalykin; SB – S. Belokobylskij; VG – V. Gussakovskij, VL – V. Loktionov.

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 7.7.4 Pro. All images were post-processed for contrast and brightness using Adobe Photoshop. New distributional records are noted with an asterisk (*).

The map was generated using an online tool for producing publication-quality point maps, SimpleMappr (Shorthouse 2010).

Taxonomy

Epeolus Latreille, 1802

Epeolus Latreille, 1802: 427. Type species: Apis variegata Linnaeus, 1758, monobasic.

Epeolus julliani species group

Diagnosis. Labrum with apical margin straight and without medial tooth; apically or near apical margin (as opposed to medially, as in species in the Epeolus variegatus species group, or sub-apically, as in species in the E. cruciger species group) with two sharply carinate, triangular (as seen as lateral view) teeth (tubercles). Axilla large with short apical tooth (extending well beyond midlength of scutellum but not as far back as its posterior margin).

Species included. Epeolus fasciatus Friese, 1895, E. iranicus Bogusch, 2021, E. julliani Pérez, 1884, E. kyzylkumicus Astafurova, sp. nov., E. laticauda Bischoff, 1930, E. rasmonti Astafurova & Proshchalykin, sp. nov., E. seraxensis Radoszkowski, 1893, E. siculus Giordani Soika, 1944, E. transitorius Eversmann, 1852.

Remarks. Le Divelec (2021) also included Epeolus flavociliatus Friese, 1899 to this species group, but this species clearly differs from species julliani group by small (ill-defined) labral tubercles and elongate (longer than mesoscutellum) axillae. Additionally, the placement of E. flavociliatus in this group is not supported by recent phylogenetic studies (see Onuferko et al. 2019; Lim et al. 2022). Together with E. priesneri Bogusch, 2021, E. subrufescens Saunders, 1908, E. warnckei Bogusch, 2018 and E. ruficornis Morawitz, 1875 this species rather belongs to another group.

Key to the species of the E. julliani species group

1 Anteromedial area of mesepisternum with deep depression (Fig. 8A, B) 2
Anteromedial area of mesepisternum normal, more or less flat 3
2 Subpleural signum positioned on a small elevated plate (Fig. 8C) and the mesepisternum lateral to anteromedian depression with a strong sharp carina (Fig. 8B) E. rasmonti sp. nov.
Subpleural signum not elevated under mesepisternum; mesepisternum lateral to anteromedian depression with weak rounded corners (Fig. 8A) E. transitorius Eversmann
3 Female 4
Male 10
4 Apical bands of metasomal terga uninterrupted 5
Apical bands of metasomal terga interrupted medially (at least on T2–T4) 6
5 Mesoscutum entirely reddish; F1 slightly longer than F2 (ca 1.5 vs 1.3–1.4 times as long as wide) E. kyzylkumicus sp. nov.
Mesoscutum entirely black or largely black; F1 slightly shorter than F2 (ca 1.2 vs 1.3–1.4 times as long as wide) E. seraxensis Radoszkowski
6 Pubescence on S2 discs dense, obscuring integument E. laticauda Bischoff
Pubescence on S2 discs sparse, sculpture of integument clearly visible 7
7 Mesoscutum and terga reddish E. iranicus Bogusch
Mesoscutum and terga black. Mesoscutum sometimes reddish along margins but never predominantly reddish 8
8 F2 1.6–1.7 times longer than F3. Integument mostly black, labrum and axilla usually black. Mesepisternum sparsely punctate E. fasciatus Friese
F2 slightly longer than F3. Integument with well developed red body coloration, labrum and axillae red. Mesepisternum densely punctate 9
9 Metasomal terga with bright yellow bands of tomentum; propodeum forms obtuse angle with mesoscutellum [known only from Sicily] E. siculus Giordani Soika
Metasomal terga with pale-yellow or whitish bands of tomentum; propodeum forms right angle with mesoscutellum [widespread] E. julliani Pérez
10 Apical bands of metasomal terga interrupted medially (at least on T2–T4) 11
Apical bands of metasomal terga uninterrupted 14
11 Metasomal terga reddish E. iranicus Bogusch
Metasomal terga black 11
12 Apical bands of T1 uninterrupted E. fasciatus Friese
Apical bands of T1 interrupted 13
13 Metasomal terga with yellowish bands of tomentum; propodeum forms obtuse angle with mesoscutellum [known only from Sicily] E. siculus Giordani Soika
Metasomal terga with whitish bands of tomentum; propodeum forms right angle with mesoscutellum [widespread] E. julliani Pérez
14 Mesoscutum and terga reddish. Pygidium narrower, 1.05–1.1 times wider than long E. kyzylkumicus sp. nov.
Mesoscutum and terga black or brownish. Pygidium wide, 1.4–1.7 times wider than long 15
15 Labral teeth positioned directly on apical margin. Hind basitarsus bordered by dense fringe of plumose setae. Pygidium wide, 1.6–1.7 times wider than long, apically distinctly bilobed. Lateral lobes of penis (best seen in dorsal view) small, triangular, extending to mid-length of penis valve (Fig. 9A, B, arrow) E. seraxensis Radoszkowski
Labral teeth usually positioned near apical margin. Hind basitarsus bordered by sparse fringe of simple setae. Pygidium narrow, 1.4 times wider than long, apically slightly bilobed or rarely straight. Lateral lobes of penis (best seen in dorsal view) large, petal shaped, elongate, extending to tip of penis valve (Fig. 9I, J, arrow) E. laticauda Bischoff

Epeolus fasciatus Friese, 1895

Epeolus fasciatus Friese, 1895: 208, ♀, ♂ (type locality: Hungary, Budapest [Pest]; Museum für Naturkunde, Berlin; Muséum National d’Histoire Naturelle, Paris; ZISP).

Material examined

Hungary, Pest [Budapest], 2.VII.1886 (1 ♂, syntype), Friese [ZISP].

Distribution

Southern and Central Europe, Turkey (Bogusch 2018).

Epeolus iranicus Bogusch, 2021

Epeolus iranicus Bogusch, 2021: 52, ♀, ♂ (type locality: Kuhre-Sefid, Bazuft, Iran; OLBL).

Material examined

None.

Distribution

Iran (Bogusch, 2021).

Epeolus julliani Pérez, 1884

Fig. 9E, F

Epeolus julliani Pérez, 1884: 318–322, ♀ (type locality: Marseille, France; Muséum National d’Histoire Naturelle, Paris).

Material examined

Austria, Wien, (1 ♀), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [ZISP]; Azerbaijan, Lenkoran, 28.VII.1930, (1 ♀), A. Shestakov [ZISP]; Georgia, Lagodehy, (1 ♀), coll. F. Morawitz [ZISP]; Iran, Shaku, Elbrus Mts., VI.1914, (1 ♀), Kirichenko [ZISP]; Tularud, 11.V.1916, (1 ♂), B. Ilyin [ZISP]; Kerman Prov., 8 km N of Bordsir, 200 m, 29°95'N, 56°58'E, 6.VI.2010, (1 ♂), Mi. Halada (OLBL); Moldova, Leovo, 25.VII.1913, (1 ♂), Chernavin [ZISP]; Kazakhstan, Kokshetau Mts., 1. VII, 3.VIII.1958, (2 ♀), V. Rudolf [ZISP]; Aktobe, Berchogur [Birshoghyr], 26.VI.1910, (7 ♀, 3 ♂), L. Bubyr [ZISP]; Russia, Dagestan Rep., 20 km W of Makhachkala, Sarykum, 23–24.V.2019, (1 ♂), MP, VL [FSCV]; Crimea, Mukhalatka, VII.1902, (1 ♀), N. Kuznetzov [ZISP]; Sevastopol, 28.VII.1916, (1 ♀), Pliginski [ZISP]; idem, 7.VII.2015, (1 ♀), V. Zhidkov [ZISP]; Tarkhankut, Atlesh, 29.VII.2008, (1 ♂), AF [ZISP]; Tarkhankut, Bolshoy Kastel, 25.VII.2015, (7 ♀), AF [ZISP]; idem, on Jurinea stoechadifolia, 8.VIII.2015, (1 ♀), V. Zhidkov [ZISP]; idem, 9.VIII.2020, (2 ♀), S. Ivanov [ZISP]; Tarkhankut, Kipchak, 16.IV.2016, (3 ♀, 1 ♂), V. Zhidkov [ZISP]; Tarkhankut, Dzhangul, 26.VII.2017, (3 ♀), AF [ZISP]; Feodossia, Karadag, 15.VI.2015, (1 ♀), AF [ZISP]; Lukull Cape, 8.VII.2015, (3 ♀), AF [ZISP]; near Sudak, 27.V.2016, (1 ♂), AF [ZISP]; Krasnodar Terr., Anapa, Bolshoy Utrish, 2.VII.2018, (1 ♀), AF [ZISP]; Orenburg Prov., Orenburg, (1 ♂), coll. F. Morawitz, Epeolus transitorius Eversm. [Morawitz det.] [ZISP]; SPAIN, Andalusia, 26.VI. (1 ♀, 1 ♂) [ZISP]; Ukraine, Akkerman[=Bilhorod-Dnistrovskyi], (1 ♂), 21.VIII.1921, Petrovich [ZISP].

Remarks

The main differences between Epeolus julliani and E. transitorius are outlined by Le Divelec 2021, who removed the former from synonymy with the latter. Here, we describe the structure of the male genitalia (Table 1).

Table 1.

Main differences between species of the Epeolus julliani group.

seraxensis iranicus (according to Bogusch 2021) julliani + siculus kyzylkumicus laticauda transitorius + rasmonti
Both sexes
Anteromedial area of mesepisternum normal, more or less flat with deep depression
Apical bands of metasomal terga uninterrupted interrupted medially on T2–T4 interrupted medially uninterrupted narrowly interrupted medially in female, uninterrupted in male interrupted medially
Coloration of tergal discs varies from brownish to reddish in female and from black to brownish in male reddish black reddish varies from dark brown to red- brown in female and from black to brownish in male black
Female
Length of flagellomeres (ratio L/W) F1 little shorter than F2 (ca 1.2 vs 1.3–1.4); remaining flagellomeres distinctly longer than wide F1 distinctly shorter than F2 (1.05–1.1 vs 1.35); remaining flagellomeres slightly longer than wide F1 equal or little shorter than F2 (1.2 vs 1.2–1.3); remaining flagellomeres distinctly longer than wide F1 little longer than F2 (ca 1.5 vs 1.3–1.4); remaining flagellomeres distinctly longer than wide F1 little shorter than F2 (1.2–1.3 vs 1.4–1.5); remaining flagellomeres distinctly longer than wide F1 distinctly shorter than F2 (ca 1.5 vs 1.0–1.1); remaining flagellomeres slightly longer than wide
Pubescence on S2 disc dense, obscuring integument sparse, sculpture of integument well-visible sparse, sculpture of integument well-visible moderate, almost obscuring integument dense, obscuring integument sparse, sculpture of integument well-visible
Male
Lateral lobes of penis (dorsal view) Small, triangular, extending mid-length of penis valve (Fig. 9C, D, arrow) not studied Small, petal shaped, extending mid-length of penis valve (Fig. 9E, F, arrow) Medium sized, petal shaped, not extending tip of penis valve (Fig. 9C, D, arrow) Large, petal shaped, elongate, extending tip of penis valve (Fig. 9G–J, arrow)
Pygidium very wide, 1.6–1.7 times wider than long, apically distinctly bilobed 1.3 times wider than long, apically rounded 1.25–1.35 times wider than long, apically rounded, sometimes slightly bilobed 1.05–1.1 times wider than long, apically rounded 1.4 times wider than long, apically slightly bilobed or rarely straight 1.1–1.2 times wider than long, apically rounded, sometimes slightly bilobed

The integument coloration and variability are closest to E. transitorius (see below). Unlike E. transitorius, all studied female specimens of E. julliani have a sparsely pubescent or almost glabrous clypeus (vs on that is often obscured by dense tomentum), and the lower mesepisternum is always pubescent (vs often glabrous).

Distribution

North Africa, Middle East, Europe, Caucasus, Russia (south of European part, south Ural), *Kazakhstan, Iran (Bischoff 1930; Le Divelec 2021; current data).

Epeolus kyzylkumicus Astafurova, sp. nov.

Figs 1, 2, 3, 8D, 9G, H

Material examined

Holotype : ♀, Uzbekistan, Kyzyl-kum [Kyzylkum desert], 10 km SW Arnasay [Lakes], 27.VIII.1979, Yu. Pesenko [ZISP]. Paratypes: 3 ♂, the same label as in the holotype; 1 ♀, Kazakhstan, Perowsk[=Qyzylorda], Syr-Darja Geb., 17.VII.1909, W. Nikolsky [ZISP]; 1 ♀, Tajikistan, Farap, NW Bukhara, 5.VII.1928, V. Gussakovskij [ZISP].

Diagnosis

This species is most similar to Epeolus iranicus, especially with regard to the extensive red integument coloration, but can be separated from it by the uninterrupted apical bands on the metasomal terga, dense pubescence of sterna, and longer antennae (flagellomeres distinctly longer than wide in both sexes vs slightly longer than wide in females and slightly shorter than wide in males of E. iranicus). The differences between E. kyzylkumicus sp. nov. and other species of the julliani group are outlined in Table 1.

Description

Female. Total body length 8.0 mm (Figs 1A, 2A); forewing length (without tegula) 6.0 mm.

Figure 1. 

Epeolus kyzylkumicus Astafurova, sp. nov., holotype, female (A), paratype, male (B) A, B habitus, lateral view. Scale bars: 1.0 mm.

Structure and sculpture: Head (Fig. 2B) 1.3 times as wide as long. Labrum (Fig. 2D) 1.65 (holotype) to 1.75 times (paratypes) as wide as long, angulated basally, rounded laterally and weakly concave medially, apical margin straight without medial tooth; close to apex (but not directly) with two well-visible teeth (tubercles); integument shiny, densely punctate (10–30 μm / confluent–2). Clypeus densely and finely punctate (10–15 μm / confluent–0.5), narrowly impunctate along apical margin. Frons with developed frontal keel. Frons and vertex areolate punctate (15–30 μm). Flagellomeres long, F1 1.5 times as long as wide, succeeding flagellomeres ca 1.3–1.4 times as long as wide. Mesoscutum and mesoscutellum coarsely and densely punctate (30–70 μm / confluent–0.5), small interspaces between punctures shiny and smooth. Axilla convex, apically with distinct short tooth. Mesoscutellum with deep medial longitudinal impression distinctly divided mesoscutellum on two slightly convex lobes; poste­rior margin scarcely extending over propodeum. Mesepisternum areolate-punctate (sculpture not visible under pubescence). Propodeal triangle shagreened. Metasomal terga densely and finely punctate (10–15 μm / 0.5–1), interspaces shiny and smooth; marginal zones (apical impressed area) wide, equal to length of discs. Pseudopygidial area triangular. Pygidial plate trapezoidal, apically truncate (Fig. 2C). Processes on sides of S6 normal, with short projections (Fig. 8D). Metasomal terga and sterna with punctures more or less equally dense.

Figure 2. 

Epeolus kyzylkumicus Astafurova, sp. nov., holotype, female A habitus, dorsal view B head, frontal view C S6, ventral view D labrum, frontal view E metasoma, ventral view. Scale bars: 1.0 mm (A, E); 0.5 mm (B, C); 0.3 mm (D).

Integument coloration: Body mostly reddish, but paraocular and genal areas, frons and vertex black.

Pubescence: Body with dense and mostly white tomentum (brownish only on medial part of tergal discs). Labrum with mixed thin and plumose setae. Face and genal area with dense tomentum obscuring integument, vertex with sparser and short setae. Pronotum dorsally with tomentum obscuring integument. Mesoscutum with dense tomentum peripherally and with wide paramedial strips. Metanotal integument entirely obscured by tomentum. Lower and lateral parts of thorax and propodeum laterally entirely obscured by tomentum. Legs with dense tomentum. T1 with wide basal band of tomentum connected with apical band laterally; margs of T1–T4 with uninterrupted bands of tomentum. T1–T4 discs with tomentum dense and laterally similar to that on apical margins, but somewhat sparser and brownish medially. T5 entirely obscured by tomentum. Pseudopygidial area with golden pubescence. Sterna entirely obscured by tomentum, moderate on discs and distinctly denser and longer on margins (Fig. 2E).

Male. Structure, sculpture, coloration and pubescence are similar to those of the female (Figs 1B, 3A). Total body length 6.0–7.0 mm; forewing length (without tegula) 5.5 mm. Head (Fig. 3B) ca 1.2 times as wide as long. Labrum flatter and wider than in female, 1.9 times as wide as long. F1 ca 1.3 times as long as wide, succeeding flagellomeres ca 1.2 times as long as wide. Clypeus with dense tomentum obscuring integument. Mesoscutum entirely covered by plumose setae, denser peripherally and in anterior half. Hind basitarsus boarded by sparse fringe of pale short setae (Fig. 3E). Pygidial plate (T7) shiny, coarsely and densely punctate, 1.05–1.1 times wider than long, slightly narrowed toward apex; apical margin rounded (Fig. 3D). Margins of S2 and S3 with dense uninterrupted white tomentum bands; S4 and S5 normal, with cream long setae (Fig. 3C). Genitalia as in Fig. 9C, D.

Figure 3. 

Epeolus kyzylkumicus Astafurova, sp. nov., paratype, male A habitus, dorsal view B head, frontal view C metasoma, ventral view D T7, dorsal view E hind basiratsus, dorsal view. Scale bars: 1.0 mm (A, C); 0.5 mm (B, E); 0.3 mm (D).

Etymology

The specific name “kyzylkumikus” is an adjective in the nominative singular and refers to the occurrence of this species in Kyzylkum desert of Central Asia.

Distribution

Kazakhstan (Qyzylorda Prov.), Kyrgyzstan, Tajikistan.

Epeolus laticauda Bischoff, 1930

Fig. 9I, J

Epeolus laticauda Bischoff, 1930: 13, ♂ (type locality: “Monda, Mongolei” [Mondy, Buryatia Republic, Russia]; Natu­ral History Museum, Berlin).

Material examined

Kazakhstan, Kulandy, Aral Sea, 13.VII.1900, (1 ♀, 1 ♂), L. Berg (ZMMU); Mergenevo, Ural River, 12.VII.1951, (1 ♀), Romadina [ZISP]; Tajikistan, Kulyab, 7.VIII.1933, (2 ♀), V. Popov [ZISP]; Dzhili-Kul, Vakhsh River, 12.VI.1934, (1 ♀), VG [ZISP]; Kabadian, 2.VII.1934, (1 ♀, 2 ♂), VG; Ayvadzh, 8.VIII.1934, (1 ♀), VG; Stalinabad[=Dushanbe), Botanical garden, 21.VII–8.VIII.1943, (9 ♀, 9 ♂), V. Popov [ZISP]; Kurgan-Tuybe, 14.VIII.1948, (1 ♀), V. Popov [ZISP]; Turkmenistan, Ashgabat, (1 ♀), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [ZISP]; Kara-Kala, 27.V.1953, (1 ♂), Steinberg [ZISP]; Uzbekistan, Farab, 31.V.1912, (1 ♀, 3 ♂), Golbek [ZISP]; Bukhara, 26.VI.1926, (1 ♀), V. Yakhontov [ZISP]; Khiva, 15.VI.1927, (6 ♀), VG [ZISP]; idem, 18–22.VI.1927, (1 ♀, 9 ♂), VG [ZISP]; Kattakurgan, 12.VII.1931, (1 ♀), VG [ZISP]; Dzhuma, 25–29.VI.1937, (15 ♀, 7 ♂), V. Popov [ZISP].

Distribution

Russia (Buryatia Rep.), *Kazakhstan, Tajikistan, Turkmenistan, Uzbekistan (Popov 1935, 1949, 1967; current data).

Variability

Labral tubercles in both sexes are positioned, typically, more or less close to the apical margin and rarely directly thereupon.

Females have well developed red body coloration. The labrum, clypeus, pronotal lobe, tegulae, mesepisternum, axillae, mesoscutellum, metanotum, legs, pygidial plate and sterna are always reddened. The antennae are mostly reddish, but with terminal flagellomeres usually brownish. The pronotum varies from reddish to partially black. The mesoscutum is mostly black, but red patterning sometimes also occurs laterally, or rarely it is almost entirely red-brownish. The propodeum varies from black to reddish. The metasomal terga vary from dark brown to red-brown with golden marginal zones; T5 is usually reddish.

Males are mostly black (excepting a red labrum, scape, pedicel, F1, pronotal lobe, tegulae, legs and pygidial plate). The clypeus is red (entirely or only in apical half). The mesepisternum, axillae and mesoscutellum are typically black, but sometimes can be partially or entirely red. The sterna vary from brownish to reddish. The pygidial plate apical margin is typically more or less bilobed, sometimes rather straight.

Epeolus rasmonti Astafurova & Proshchalykin, sp. nov.

Figs 4, 5, 6, 7, 8B, C, 9G, H, 10

Material examined

Holotype : ♀, Russia, Buryatia Republic, Gusinoye Lake, Baraty, 25.VII.2007, A. Lelej, M. Proshchalykin, V. Loktionov [ZISP]; Paratypes: 1 ♀, 1 ♂, the same label as in the holotype [FSCV/ZISP]; 6 ♀, 1 ♂, the same label as in the holotype, but 26.VII.2007 [FSCV/ZISP]; 1 ♂, Naushki, Selenga River, 30.VII.2007, A. Lelej, M. Proshchalykin, V. Loktionov [ZISP].

Additional material

China, Beijing, park of the Summer Palace, 18.VIII.1954, (1 ♀), G. Bey-Bienko [ZISP]; Mongolia, Khutag, Selenga River, 25.VII.1975, (1 ♂), E. Narchuk [ZISP]; 200 km SEE of Baruun-Urt, Moltsoy Els, 1250 m, 27.VII.2007, (3 ♀, 14 ♂), M. Kadlecova, M. Halada, P. Tymer [OLBL]; 100 km NE of Ondorkhaan, Kerulen River, 970 m, 22.VII.2007, (3 ♂), M. Kadlecova [OLBL]; 100 km W Choibalsan, 820 m, 23.VII.2007, (1 ♂), M. Halada [OLBL]; 15 km W Choibalsan, Kerulen River, 770 m, 24.VII.2007, (1 ♂), M. Halada [OLBL].

Diagnosis

This species is most similar to Epeolus transitorius, with which it uniquely shares a deep depression in the anteromedian area of the mesepisternum, but can be separated from it by the position of the subpleural signum on a small elevated plate (Fig. 8C) and the presence of a strong, sharp carina on the mesepisternum lateral to the anteromedian depression (Fig. 8B) (vs weak rounded corners, Fig. 8A). The differences between Epeolus rasmonti sp. nov., E. transitorius and other species of the julliani group are outlined in Table 1.

Description

Female. Total body length 7.0–9.0 mm (Figs 4, 5A); forewing length (without tegula) 6.0–7.0mm.

Figure 4. 

Epeolus rasmonti Astafurova & Proshchalykin, sp. nov., holotype, female, habitus, lateral view. Scale bar: 1.0 mm.

Structure and sculpture: Head (Fig. 5B) 1.3 times as wide as long. Labrum (Fig. 5C) 1.55 times as wide as long, more or less rounded basally and laterally, weakly concave medially, apical margin straight without medial tooth; apically with two well-visible carina-shaped teeth (tubercles); integument shiny, areolate punctate (10–30 μm). Clypeus densely and finely punctate (10–15 μm / confluent–0.5), narrowly impunctate along apical margin. Frons with developed frontal keel. Frons and vertex coarsely and areolate punctate (30–40 μm). F1 long, ca 1.5 times as long as wide, F2 distinctly shorter than F1 (1.0–1.1 times as long as wide), succeeding flagellomeres slightly longer than wide (ca 1.2 times). Mesoscutum and mesoscutellum coarsely and mostly areolate punctate (30–70 μm), a few small interspaces between punctures shiny and smooth. Axilla slightly convex laterally, apically with distinct short tooth. Mesoscutellum with shallow medial longitudinal impression; poste­rior margin scarcely extending over propodeum. Mesepisternum areolate punctate on upper half and with dull, delicately shagreened interspaces on lower part; anteromedian area with deep depression, laterally from anteromedian depression with strong sharp carina (Fig. 8B). Propodeal triangle shagreened. Metasomal terga densely and finely punctate (10–15 μm / confluent–1), interspaces shiny; marginal zones wide, equal to length of discs. Pseudopygidial area narrow, linear. Pygidial plate trapezoidal, apically truncate. Processes on sides of S6 normal, with short projections. Metasomal terga and sterna with punctures more or less equally dense (Fig. 5D).

Figure 5. 

Epeolus rasmonti Astafurova & Proshchalykin, sp. nov., holotype, female A habitus, dorsal view B head, frontal view C labrum, frontal view D metasoma, ventral view. Scale bars: 1.0 mm (A, D); 0.5 mm (B); 0.3 mm (C).

Integument coloration: Head mostly black, but mandibles (excluding darker apex), labrum, clypeus along apical margin, scape and F1 reddish (amber). Mesosoma mostly black; pronotal lobe, tegulae, axillae, mesoscutellum, metanotum medially and legs (including spurs) reddish (amber); wings with brownish darkening, stigma and veins brown. Mesepisternum partially reddish (usually in middle part, lower scobal area). Metasomal terga black; marginal zones brownish to dark-golden apically. Py­gidial plate reddish with brownish edging. Sterna brownish to reddish; margins golden.

Pubescence: Body with relatively sparse pubescence; tomentum white, except black on tergal discs. Labrum with thin sparse setae. Paraocular area with dense tomentum obscuring integument; remaining part of face and vertex with sparse thin pubescence. Upper half of frons with relatively long erect simple setae mixed with adpressed sparse plumose pubescence. Genal area with relatively dense plumose setae, but not obscuring integument. Pronotum dorsally with tomentum obscuring integument, but medially setae sparse and short. Mesoscutum with dense tomentum peripherally and with narrow paramedial strips. Mesepisternum with sparse, short plumose pubescence or glabrous on lower part and with relatively dense and longer plumose pubescence on upper half. Metanotal integument obscured by tomentum except medially. Legs with sparse pubescence. T1 with basal band of tomentum interrupted medially and connected with apical band laterally; margins of T1–T4 with widely interrupted bands of tomentum. Black tomentum on T1–T4 discs and in interrupted area sparser than pale tomentum. T5 obscured by white tomentum laterally and black medially. Pseudopygidial area with white pubescence. Discs of metasomal sterna with short brownish plumose setae, sparse on S2 and entirely obscured by tomentum on S3–S5; margins with dense and pale tomentum interrupted medially.

Male. Structure, sculpture, coloration and are similar to those of the female (Figs 6, 7), but pubescence more developed. Total body length 5.5–8.0 mm; forewing length (without tegula) 6.0–6.5 mm. F1 1.3–1.4 times as long as wide, succeeding flagellomeres ca 1.1 times as long as wide. Labrum with dense, plumose setae medially. Face obscured by plumose tomentum, but on upper frons sparser and mixed with long setae (Fig. 7B). Mesoscutum with paramedial strips wider than in female. Axillae and mesoscutellum black or reddish. Mesepisternum mostly areolate-punctate, with shiny, smooth interspaces on lower part; entirely obscured by tomentum. Hind basitarsus boarded by dense fringe of long golden setae (Fig. 7C). Metasomal terga with apical bands narrower interrupted than in female. Pygidial plate (T7) reddish, shiny, coarsely and densely punctate, ca 1.2 times wider than long, narrowed toward apex; apical margin rounded, slightly bilobed or almost straight (Fig. 7D). Margins of S2 and S3 with dense uninterrupted white tomentum bands; S4 and S5 normal, with gold long setae. Genitalia as on Fig. 9G, H.

Figure 6. 

Epeolus rasmonti Astafurova & Proshchalykin, sp. nov., paratype, male A, B habitus, lateral view (A) ventral view (B). Scale bars: 1.0 mm.

Figure 7. 

Epeolus rasmonti Astafurova & Proshchalykin, sp. nov., paratype, male A habitus, dorsal view B head, frontal view C hind basiratsus, dorsal view D T7, dorsal view. Scale bars: 1.0 mm (A); 0.5 mm (B–D).

Etymology

The specific epithet is a patronym honoring our colleague and friend Prof. Pierre Rasmont (University of Mons, Belgium) on the occasion of his 65th birthday and in recognition of his many contri­butions to the study of bee diversity.

Distribution

(Fig. 10). Russia (Buryatia Rep.), Mongolia (Bulgan, Dornod, Khentii, Sukhbaatar), China (Beijing).

Epeolus seraxensis Radoszkowski, 1893

Fig. 9A, B

Epeolus transitorius var. seraxensis Radoszkowski, 1893: 54–55, ♀, ♂ (lectotype: ♀, designated by Bogusch 2021: 59, Serax, Turkmenistan; Natu­ral History Museum, Berlin).

Material examined

Azerbaijan, Araxesthal[=Nakhchivan Republic, near Ordubad], (1 ♀), Reitter leg. [ZISP]; Kazakhstan, Tartugay, 3–15.VI.1929, (3 ♀, 2 ♂), A. Shestakov [ZISP]; Charyn valley, W Chundza, 650 m, 43°37'N, 79°21'E, 29–31.V.2001, (2 ♀), M. Hauser [OLBL]; 30 km ENE Shelek, 43°41.24'N, 78°38.50'E, 500 m, on Tamarix sp., 2.VIII.2002, (1 ♂), M. Kuhlmann [OLBL]; 3 km NE Boradysu, 30 km E Shelek, 19.VI.2003, (1 ♀), V. Kazenas [ZISP]; Charyn valley, 12 km W Chundzha, 12.VI.2004, (1 ♀, 1 ♂), V. Kazenas [ZISP]; Tajikistan, Dushanbe, 1931, (1 ♀, 1 ♂), N. Fursov [ZISP]; Turkmenistan, Kopetdag, 12 km SW Kyzyl-Arvat, 24.V.1953, (1 ♂), Odintzova [ZISP].

Distribution

Azerbaijan, *Kazakhstan, *Tajikistan, Turkmenistan, Iran, Israel (Bogusch 2021; current records).

Remarks

Records from Bogusch (2021: 59) from Iran (“Anaesthal”) belong to specimens collected in Araxesthal, Azerbaijan, near Ordubad (see, Reitter 1890, 1905).

Variability

Females have well developed red body coloration, including the labrum, clypeus, pronotal lobe, tegulae, mesepisternum, axillae, mesoscutellum, metanotum, legs, pygidial plate and sterna. The antennae are brownish with the scape, pedicel and F1 entirely reddish. The mesoscutum is mostly black, sometimes with small red patterning peripherally and a pair red stripes posteriorly. The propodeum varies from entirely reddish to entirely black. The tergal discs vary from brownish to reddish. The males are mostly black, except the red labrum, scape, pedicel, F1, pronotal lobe, tegulae, legs and pygidial plate. The axillae and mesoscutellum are black or red. Tergal discs vary from black to brownish, and sterna from brownish to reddish.

Epeolus siculus Giordani Soika, 1944

Epeolus siculus Giordani Soika, 1944: 20, ♀ (type locality: Messina, Sicily, Italy; holotype is lost).

Material examined

Italia, Sicilia, (2 ♀), coll. F. Morawitz [ZISP]; Sicilia, 35 km SW Ragusa, 18–22.VI.2002, (1 ♂), J. Halada (OLBL).

Distribution

Italia (Sicilia) (Giordani Soika 1944).

Epeolus transitorius Eversmann, 1852

Figs 8A, 10

Epeolus transitorius Eversmann, 1852: 102 (lectotype: ♀, designated by Le Divelec 2021: 16, Indersk District, Atyrau Province, Kazakhstan; ZISP).

Material examined

Georgia, Tiflis[=Tbilisi], (1 ♀), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [ZISP]; Kazakhstan, Zaysan, IX.1908, (1 ♂), coll. Gussakovskij [ZISP]; Fort Aleksandrovsk[=Fort-Shevchenko], 5.VII.1909, (1 ♂), Nasonov [ZISP]; Berchogur[=Birshoghyr], 26.VI.1910, (1 ♂), L. Bubyr [ZISP]; Chernoretsk, 11.VII.1925, (2 ♂), coll. Gussakovskij [ZISP]; near Chelkar Lake, Bolshiye Barsuki desert, 20.VI–10.VII., (1 ♀) [ZMMU]; Yanvartsevo, 19.VI–17.VIII.1950, (1 ♀, 5 ♂), V. Rudolf [ZISP]; 10 km N Zharkol Lake, 9.VII.1950, (2 ♀), V. Rudolf [ZISP]; Tengiz Lake, 3.VII.1957, (2 ♂), V. Rudolf [ZISP]; Kokshetau, on Allium glabrosum, 11.VII.1958, (1 ♀), V. Rudolf [ZISP]; 50 km E Balkhash Lake, 26–27.VI.1992, (1 ♀, 2 ♂), J. Halada [OLBL]; 5 km W Shardara, 250 m, 41°16'N, 67°53'E, 23–24.V.2016, (1 ♀, 1 ♂), J. Halada [OLBL]; Russia, Astarakhan Prov., 35 km NNE of Astrakhan, 26.VII.2015, (1 ♀, 6 ♂), MP, VL, S. Belokobylskij, M. Mokrousov [ZISP]; Altai Terr., Novovoznesenka, 11.VII.1922, (2 ♂), A. Reygardt [ZISP]; Crimea, 8 km N of Dmitrovka, 5.VI.2018, (1 ♀), V. Savchuk [ZISP]; Mirny, Donuslav Lake, 17.VII.2017, (1 ♀), AF [ZISP]; Dagestan Rep., 20 km W of Makhachkala, Sarykum, 20–29.V.2019, (2 ♀, 1 ♂), MP, VL [FSCV]; Kalmykia Rep., 22 km E of Yashkul, 16–18.VII.2015, (1 ♀, 2 ♂) MP, VL, S. Belokobylskij, M. Mokrousov [FSCV/ZISP]; 17 km SWW of Artezian, Kuma River, 18–24.VII.2015, (2 ♀, 9 ♂), MP, VL, S. Belokobylskij, M. Mokrousov [FSCV/ZISP]; Orenburg Prov., Mayachnaya, 4.VIII.1993, (1 ♀), L. Zimin [ZMMU]; Orenburg, 9.VIII–3.IX.1926, (6 ♀, 1 ♂), P.A. Vorontzovskiy [ZISP]; Volgograd Prov., Sarepta, (2 ♀, 2 ♂), coll. F. Morawitz [ZISP]; Tinguta, 11.VIII.1952, (2 ♂), G. Viktorov [ZMMU]; Tajikistan, Yagnob River, (1 ♂), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [ZISP]; Stalinabad[=Dushanbe], 18.VII.1936, (2 ♀), VG [ZISP]; idem, 15.VII.1943, (2 ♀, 1 ♂), V. Popov [ZISP]; Ukraine, Yareski, 21.VII.1922, (2 ♀), coll. V. Gussakovskij [ZISP]; Kharkov, (1 ♂), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [ZISP]; Uzbekistan, Kurgan-Tyube[=Kurgantepa], Fergana, 27.V., 17.VII.1938, (1 ♀, 1 ♂), V. Popov [ZISP].

Figure 8. 

Epeolus transitorius Eversmann, 1852, female (A), E. rasmonti Astafurova & Proshchalykin, sp. nov., paratype, female (B, C), E. kyzylkumicus Astafurova, sp. nov., female (D) A, B lower mesepisternum, ventral view C subpleural signum (arrow), ventral view D S6, lateral view. Scale bars: 0.5 mm (A, B); 0.3 mm (C, D).

Variability

Females typically have well developed red body coloration. The labrum, pronotal lobe, tegulae, metanotum (medially), legs, and sterna are always red. The clypeus is usually red, but rarely can be mostly black or dark brownish (except with a red apical margin). The antennae are typically brownish with a red scape, pedicel and F1. The pronotum varies from black to partially red. The mesoscutum is mostly black, but red patterning sometimes also occurs laterally. Coloration of the mesepisternum varies from entirely red to partially black. The axillae and mesoscutellum are usually red, but rarely black peripherally. The terga are black or brownish, but T5 apically is typically red; sometimes red patterning occurs also on the anterior half of T1 and along margins. Pubescences of female is moderate. The face (except upper half of frons) is often obscured by dense tomentum, but 45% of specimens examined have the clypeus with sparse pubescence or almost glabrous. The mesepisternum is densely tomentose on the upper half and with sparse pubescence or glabrous below the scobal suture.

Figure 9. 

Male, genitalia A, C, E, G, I genital capsule, dorsal view B, D, F, H, J penis, dorsal view A, B Epeolus seraxensis Radoszkowski, 1893 C, D Epeolus kyzylkumicus Astafurova, sp. nov. E, F E. julliani Pérez, 1884 G, H E. rasmonti Astafurova & Proshchalykin, sp. nov. I, J E. laticauda Bischoff, 1930. Scale bars: 0.5 mm.

The males do not show significant variability. The body is mostly black, but the labrum, scape, pedicel, pronotal lobe, tegulae, legs and pygidial plate are red. The clypeus is black or with red patterning apically. The axillae and mesoscutellum are usually black, often with red axillar teeth, sometimes partially red on mesoscutellum. The sterna are black or brownish to red along margins; marginal zones are golden. The pygidial plate is apically rounded, sometimes more or less straight or slightly bilobed.

Distribution

(Fig. 10). Greece, Georgia, Ukraine, Russia (south of European part, south Ural, SW Siberia), Kazakhstan, Uzbekistan, Tajikistan (Le Divelec 2021; current data).

Figure 10. 

Distribution of Epeolus transitorius Eversmann (red dots) and E. rasmonti Astafurova & Proshchalykin, sp. nov. (blue dots).

Remarks

The previous records of Epeolus transitorius from North Africa, Europe, Caucasus and Iran (Friese 1893, 1895; Bischoff 1930; Popov 1967; Bogusch and Hadrava 2018; Bogusch 2021) are mostly wrong (see Le Divelec 2021). Records from Morawitz (1875: 144, from Tajikistan [=Levchenko et al. 2017: 317, as Uzbekistan]) belongs to E. michailovi Astafurova & Proshchalykin, 2021; records from Turkmenistan (Levchenko et al. 2017: 317) refer to E. seraxensis Radoszkowski, 1893.

Discussion

The Epeolus julliani species group is distributed only in the southern Palaearctic, ranging from northern Africa to China. Eight of the nine species occur in the Western Palaearctic, and only Epeolus rasmonti sp. nov. is known only from the Eastern Palaearctic. There are no species with a Trans-Palaearctic range. Epeolus julliani is the most widespread species in this group, occurring from North Africa and Southern Europe east to the Middle East and Central Asia and north to the Urals. Epeolus transitorius occurs from Eastern Europe and Central Asia to Western Siberia. Epeolus seraxensis is distributed from the Middle East and Caucasus to Central Asia. Epeolus fasciatus Friese, 1895 occurs from Southern Europe to Turkey. The remaining species are endemics—Epeolus siculus – Sicily; E. laticauda Bischoff, 1930 and E. kyzylkumicus sp. nov. – Central Asia; E. iranicus – Iran.

Species of the group do not demonstrate as much intraspecific variation as has been observed within those in the E. cruciger species group (Astafurova and Proshchalykin 2022). The main combination of features to distinguish species in the julliani group is coloration and pubescence of the body, length of flagellomeres, and shape/width of the male pygidial plate (Table 1). The shape of the lateral lobes of the penis is an additional feature that can reliably distinguish species in this group (Fig. 9).

Acknowledgements

We are grateful to Maximilian Schwarz (Ansfelden, Austria), Esther Ockermueller (OLBL) and Alexander Antropov (ZMMU) for pro­viding Epeolus specimens and Michael Orr (Beijing, China) for checking the English grammar, as well as to the subject editor (Michael Ohl, Berlin, Germany) and two reviewers (Thomas Onuferko, Ottawa, Canada and Petr Bogusch, Hradec Králové, Czech Re­public) for their valuable comments, which helped to improve the quality of this paper.

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.

References

  • Astafurova YuV, Proshchalykin MYu (2021a) Review of the Epeolus tarsalis species group (Hymenoptera: Apidae, Epeolus Latreille, 1802), with description of a new species. Zootaxa 5006(1): 26–36. https://doi.org/10.11646/zootaxa.5006.1.6
  • Astafurova YuV, Proshchalykin MYu (2021b) New and little-known bees of the genus Epeolus Latreille, 1802 (Hymenoptera, Apidae, Nomadinae) from Mongolia. Journal of Hymenoptera Research 84: 11–28. https://doi.org/10.3897/jhr.84.67150
  • Astafurova YuV, Proshchalykin MYu (2021c) A new species of the genus Epeolus Latreille, 1802 (Hymenoptera, Apoidea: Apidae) from the Pamirs, with a checklist of Central Asian species. Far Eastern Entomologist 437: 10–15. https://doi.org/10.25221/fee.437.3
  • Astafurova YuV, Proshchalykin MYu (2022) Review of the Epeolus cruciger species group (Hymenoptera: Apidae, Epeolus Latreille, 1802) of Asia, with the description of two new species. Journal of Hymenoptera Research 92: 305–328. https://doi.org/10.3897/jhr.92.90098
  • Bogusch P (2018) Three new species and new records of cuckoo bees of the genus Epeolus in Turkey (Hymenoptera: Apidae: Nomadinae). Acta Entomologica Musei Nationalis Pragae 58: 127–135. https://doi.org/10.2478/aemnp-2018-0010
  • Bogusch P (2021) The cuckoo bees of the genus Epeolus Latreille, 1802 (Hymenoptera, Apidae) from the Middle East and North Africa with descriptions of two new species. Journal of Hymenoptera Research 84: 45–68. https://doi.org/10.3897/jhr.84.67049
  • Bogusch P, Hadrava J (2018) The bees of the genera Epeolus Latreille, 1802 and Triepeolus Robertson, 1901 (Hymenoptera: Apidae: Nomadinae: Epeolini) of Europe: taxonomy, identification key, distribution, and ecology. Zootaxa 4437(1): 1–60. https://doi.org/10.11646/zootaxa.4437.1.1
  • Eversmann E (1852) Fauna hymenopterologica volgo-uralensis (Continuatio). Bulletin de la Société impériale des Naturalistes de Moscou 25(2/3): 3–137.
  • Friese H (1893) Die Bienenfdauna von Deutschland und Ungarn. R. Friedlander & Sohn, Berlin, 79 pp.
  • Friese H (1895) Die Bienen Europa‘s (Apidae europaeae) nach ihren Gattungen, Arten und Varietäten auf vergleichend morphologisch-biologischer Grundlage. Theil I: Schmarotzerbienen R. Friedländer, Berlin, 218 pp. https://doi.org/10.5962/bhl.title.160173
  • Giordani Soika A (1944) Contributo alla conoscenza dei Vespidi solitari e sociali della regione etiopica. Atti del Regio Istituto Veneto di Scienze 103: 149–178.
  • Latreille PA (1802) Histoire naturelle, generale et particuliere, des Crustaces et des Insectes. Historie naturelle des Crustaces et des Insectes 3: 1–467. https://doi.org/10.5962/bhl.title.15764
  • Le Divelec R (2021) The West Palaearctic Epeolini Linsley & Michener, 1939 housed in the Muséum national d’Histoire naturelle (Paris) with some taxonomic notes (Hymenoptera: Apidae: Nomadinae). Annales de la Société entomologique de France (N.S. ) 57(4): 313–345. https://doi.org/10.1080/00379271.2021.1942206
  • Lim K, Lee S, Orr M, Lee S (2022) Harrison’s rule corroborated for the body size of cleptoparasitic cuckoo bees (Hymenoptera: Apidae: Nomadinae) and their hosts. Scientific Reports 12: 10984. https://doi.org/10.1038/s41598-022-14938-9
  • Levchenko TV, Byvaltsev AM, Proshchalykin MYu (2017) Family Apidae. In: Lelej AS, Proshchalykin MYu, Loktionov VM (Eds) Annotated Catalogue of the Hymenoptera of Russia. Volume I. Symphyta and Apocrita: Aculeata. Proceedings of the Zoological Institute RAS, Supplement 6: 263–276.
  • Michener CD (1944) Comparative external morphology, phylogeny, and a classification of the bees. Bulletin of the American Museum of Natural History 82: 151–326.
  • Morawitz F (1875) A travel to Turkestan by the member-founder of the society A. P. Fedtschen­ko, accomplished from the Imperial Society of Naturalists, Anthropologists, and Ethnog­raphists on a Commission from the General-Governor of Turkestan K. P. von Kaufmann (Issue 9). Vol. II. Zoogeographical Investigations. Pt. V. (Division 7). Bees (Mellifera). Pt. I [Apidae genuinae]. Izvestiya Imperatorskogo Obshchestva Lyubiteley Estestvoznaniya, Anthropologii i Ethnografii 21(3): 1–160. [in Russian]
  • Onuferko TM, Bogusch P, Ferrari R, Packer L (2019) Phylogeny and biogeography of the cleptoparasitic bee genus Epeolus (Hymenoptera: Apidae) and cophylogenetic analysis with its host bee genus Colletes (Hymenoptera: Colletidae). Molecular Phylogenetics and Evolution 141: e106603. https://doi.org/10.1016/j.ympev.2019.106603
  • Pérez J (1884) Contribution a la faune des Apiaires de France. Actes de la Société linnéenne de Bordeaux 37: 257–378.
  • Popov VB (1935) Contributions to the bee fauna of Tajikistan (Hymenoptera, Apoidea). Trudy Tajikskoi Basy Akademii Nauk SSSR 5: 351–408. [in Russian]
  • Popov VB (1949) Notes on the bee fauna of Tajikistan (Hymenoptera, Apoidea). Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 8: 688–699. [in Russian]
  • Popov VB (1967) The bees (Hymenoptera, Apoidea) of Middle Asia and their associations with angiosperm plants. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 38: 11–329. [in Russian]
  • Radoszkowski O (1893) Faune hyménoptèrologique transcaspienne. (Suite et Fin). Horae Societatis Entomologicae Rossicae 27(1/2): 38–81.
  • Reitter E (1890) Eine neue mit Athous verwandte Elateriden. Gattung aus russisch Armenien. Entomologische-Nachrichten 16: 247–248.
  • Reitter E (1905) Neun neue Coleopteren aus der palaearktischen Fauna. Wiener Entomologische Zeitung 24(5–6): 201–206.
  • Shorthouse DP (2010) SimpleMappr, an online tool to produce publication-quality point maps. https://www.simplemappr.net/ [accessed 12 October 2022]
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