Research Article |
Corresponding author: Maxim Yu. Proshchalykin ( proshchalikin@biosoil.ru ) Academic editor: Michael Ohl
© 2022 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 (2022) Review of the Epeolus julliani species group (Hymenoptera, Apidae, Epeolus Latreille, 1802), with descriptions of two new species. Journal of Hymenoptera Research 94: 191-213. https://doi.org/10.3897/jhr.94.96429
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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.
Anthophila, Apiformes, cleptoparasites, Palaearctic region, taxonomy
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 (
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.
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,
The taxonomy and synonymy of species generally follow those 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: 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 (
Epeolus Latreille, 1802: 427. Type species: Apis variegata Linnaeus, 1758, monobasic.
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.
1 | Anteromedial area of mesepisternum with deep depression (Fig. |
2 |
– | Anteromedial area of mesepisternum normal, more or less flat | 3 |
2 | Subpleural signum positioned on a small elevated plate (Fig. |
E. rasmonti sp. nov. |
– | Subpleural signum not elevated under mesepisternum; mesepisternum lateral to anteromedian depression with weak rounded corners (Fig. |
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. |
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. |
E. laticauda Bischoff |
Epeolus fasciatus
Friese, 1895: 208, ♀, ♂ (type locality: Hungary, Budapest [Pest]; Museum für Naturkunde, Berlin; Muséum National d’Histoire Naturelle, Paris;
Hungary, Pest [Budapest], 2.VII.1886 (1 ♂, syntype), Friese [
Southern and Central Europe, Turkey (
Epeolus iranicus
Bogusch, 2021: 52, ♀, ♂ (type locality: Kuhre-Sefid, Bazuft, Iran;
None.
Iran (Bogusch, 2021).
Epeolus julliani Pérez, 1884: 318–322, ♀ (type locality: Marseille, France; Muséum National d’Histoire Naturelle, Paris).
Austria, Wien, (1 ♀), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [
The main differences between Epeolus julliani and E. transitorius are outlined by
seraxensis |
iranicus (according to |
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. |
not studied | Small, petal shaped, extending mid-length of penis valve (Fig. |
Medium sized, petal shaped, not extending tip of penis valve (Fig. |
Large, petal shaped, elongate, extending tip of penis valve (Fig. |
|
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).
North Africa, Middle East, Europe, Caucasus, Russia (south of European part, south Ural), *Kazakhstan, Iran (
Holotype
: ♀, Uzbekistan, Kyzyl-kum [Kyzylkum desert], 10 km SW Arnasay [Lakes], 27.VIII.1979, Yu. Pesenko [
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
Female. Total body length 8.0 mm (Figs
Structure and sculpture: Head (Fig.
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.
Male. Structure, sculpture, coloration and pubescence are similar to those of the female (Figs
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.
Kazakhstan (Qyzylorda Prov.), Kyrgyzstan, Tajikistan.
Epeolus laticauda Bischoff, 1930: 13, ♂ (type locality: “Monda, Mongolei” [Mondy, Buryatia Republic, Russia]; Natural History Museum, Berlin).
Kazakhstan, Kulandy, Aral Sea, 13.VII.1900, (1 ♀, 1 ♂), L. Berg (ZMMU); Mergenevo, Ural River, 12.VII.1951, (1 ♀), Romadina [
Russia (Buryatia Rep.), *Kazakhstan, Tajikistan, Turkmenistan, Uzbekistan (
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.
Holotype
: ♀, Russia, Buryatia Republic, Gusinoye Lake, Baraty, 25.VII.2007, A. Lelej, M. Proshchalykin, V. Loktionov [
China, Beijing, park of the Summer Palace, 18.VIII.1954, (1 ♀), G. Bey-Bienko [
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.
Female. Total body length 7.0–9.0 mm (Figs
Structure and sculpture: Head (Fig.
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
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.
(Fig.
Epeolus transitorius var. seraxensis
Radoszkowski, 1893: 54–55, ♀, ♂ (lectotype: ♀, designated by
Azerbaijan, Araxesthal[=Nakhchivan Republic, near Ordubad], (1 ♀), Reitter leg. [
Azerbaijan, *Kazakhstan, *Tajikistan, Turkmenistan, Iran, Israel (
Records from
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: 20, ♀ (type locality: Messina, Sicily, Italy; holotype is lost).
Italia, Sicilia, (2 ♀), coll. F. Morawitz [
Italia (Sicilia) (
Epeolus transitorius
Eversmann, 1852: 102 (lectotype: ♀, designated by
Georgia, Tiflis[=Tbilisi], (1 ♀), coll. F. Morawitz, transitorius Eversm. [Morawitz det.] [
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).
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.
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.
The previous records of Epeolus transitorius from North Africa, Europe, Caucasus and Iran (
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 (
We are grateful to Maximilian Schwarz (Ansfelden, Austria), Esther Ockermueller (
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.