Research Article |
Corresponding author: Petr Bogusch ( bogusch.petr@gmail.com ) Academic editor: Maksim Proshchalykin
© 2021 Petr Bogusch.
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:
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. In: Proshchalykin MYu, Gokhman VE (Eds) Hymenoptera studies through space and time: A collection of papers dedicated to the 75th anniversary of Arkady S. Lelej. Journal of Hymenoptera Research 84: 45-68. https://doi.org/10.3897/jhr.84.67049
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Epeolus is one of the more species-rich genera of cuckoo bees globally. Seventeen species are known from Europe, and 23 species have been recorded from various countries in the Middle East and North Africa. Epeolus iranicus sp. nov. and E. priesneri sp. nov. are newly described in this study, and E. seraxensis Radoszkowski, 1893, stat. nov., previously a subspecies of E. transitorius Eversmann, 1852, is regarded as a valid species and a lectotype is designated. The first records of E. ibericus and E. intermedius from North Africa are given, and the distribution of E. fallax in North Africa is confirmed. This study compiles known distributional records of the species of this genus from the studied region and provides an overview of the known species richness in the studied countries. Based on available records, Turkey (14 species), Morocco (11) and Iran (9) host the highest numbers of species of Epeolus, whereas in four countries only a single species has been recorded, and the presence of this genus in seven countries has yet to be confirmed.
Distribution, Iran, Morocco, taxonomy, Turkey
Cuckoo bees form an ecological group with representatives in three families, Halictidae, Megachilidae, and Apidae, characterised by their cleptoparasitic lifestyle. Female cuckoo bees do not forage or build their own nests but instead lay eggs within the nests of host bee species (
The genus has been the focus of several taxonomic and evolutionary studies in recent years.
This study is a compilation of Epeolus records based on specimens deposited in museum collections in Europe, USA and Morocco, as well as specimens from private collections. Warncke’s notes on the distribution of bees of the genus Epeolus are also included herein. This is the first overview of bees of this genus from the Middle East and North Africa except for the comprehensive study of bees of Morocco (
Specimens of Epeolus from the following countries were examined: Algeria, Egypt, Libya, Morocco, Tunisia (= North Africa) and Bahrain, Iran, Iraq, Israel, Jordan, Kuwait, Lebanon, Oman, Palestine, Qatar, Saudi Arabia, Syria, Turkey, United Arab Emirates, and Yemen (= The Asian Part of the Middle East). The records are based on personally examined dry, pinned specimens housed in the following collections:
KUBC Kansas University Biodiversity Collection (curator Michael S. Engel);
MSIE research collection of Mohammed Shebl, Suez University, Ismailia, Egypt;
NHMB Natural History Museum, Berlin, Germany (curator Michael Ohl);
NHMC Natural History Museum, Copenhagen, Denmark (curator Lars Vilhelmsen);
Several records are adopted from unpublished notes of Klaus Warncke (cited as “K. Warncke, unpublished hand-written data”) and from published sources. The records are sorted first alphabetically according to the country, then alphabetically according to the locality, and then records from the same locality chronologically. Undated records are presented at the end of the appropriate section. The records from Turkey published by
The lists of synonymies are not presented in detail, for most species, see
Synonyms (detailed in
Northern Europe and higher altitudes in central and Southern Europe (
Iran: Elbrus, 7 km south of Chalus, 2400 m n. m., 12.VII.1977, (2 ♂), A. W. Ebmer; idem, 25.VII.1977, (1 ♀), J. Gusenleitner; idem, (1 ♀), A. W. Ebmer; Shahpasand, Tilabad, 16.VII.1977, (1 ♀), A. W. Ebmer [all P. Bogusch det.] (
Portugal and Spain, North Africa, holotype from Algeria (
Algeria: Biskra, date, collector and number of specimens unknown, (K. Warncke, unpublished hand-written data). Morocco: Taroudant, Oued Souss, 24.VI.1974, (1 ♀), K. M. Guichard, G. E. Else [M. Schwarz det.] (
Holotype
from Israel, recorded also from Cyprus, Jordan, Lebanon, Syria and Turkey (
Cyprus: Akrotiri bay, 1.VII.1933, (1 ♀), 20.VII.1933, (1 ♂), 12.VII.1943, (1 ♂), 20.VII.1943, (1 ♀); Salamis, 18.VI.1957, (1 ♂); Zakaki, 23.VI.1949, (1 ♂) G. Mavromoustakis leg. [all G. Mavromoustakis det.] (KUBC). Israel: Amitai, 31.III.1952, (1 ♀), collector unknown; Jerusalem, 29.III.1940, (1 ♂), Bytinski-Salu leg. (
North Africa, Morocco, Algeria, and Tunisia (
Algeria: Bone, collector and date unknown, (1 ♂) (
Synonyms (detailed in
Widely distributed in Europe from south to north, however, it is very rare or regionally extinct in most parts of Central and Eastern Europe, while in the southwest and Western Europe it is quite widespread. It also occurs in the Middle East (Turkey and Iran), Caucasus and Siberia (
Iran: Bayne-Teheran, 6.IX.1955, (1 ♀), collector unknown; Elburz, 50 km north of Vana, 12.–13.VII.1965, (1 ♀), Giordani Soika [both P. Bogusch det.] (
Rare species occurring in the Middle East, known from Egypt, Turkey, Syria, Armenia, and Iran, male described from a specimen from Turkey (
Synonyms (detailed in
Southern and Central Europe: France, Germany, Italy, Portugal, and Spain (
Algeria: locality, date and collector unknown, (1 ♂) (
Southern and Central Europe, but recent records are known only from France and Spain (
Synonyms (detailed in
North Africa, Turkey, and Spain – Canary Islands. Specimens from the Canary Islands differ from the North African populations and represent probably a separate species (necessary to confirm by molecular studies) (
Algeria: Biskra, 27.V.1948, (1 ♂), 30.V.1948, (1 ♀), Naef, [P. Bogusch det.] (
Europe – Portugal and Spain (
Morocco: Tanger, “Mus. Drews”, (1 ♀), date and collector unknown [P. Bogusch det.] (NHMC).
Europe – France, Italy, Portugal, Spain (
Morocco: Tanger, “Mus. Drews”, (1 ♀), date and collector unknown [P. Bogusch det.] (NHMC).
Holotype
: ♀, Iran: Bazuft, Kuhre-Sefid, 1900, Escalera leg. (
Paratypes
: 2 ♂, 1 ♀ with the same labels as holotype (
This species is most similar to E. transitorius and E. seraxensis, and all three species are characterised by the presence of apical (as opposed to submedial) labral tubercles (Figs
Epeolus iranicus sp. nov. a, c–h holotype, female b, i paratype, male a, b habitus, dorsal view c head, frontal view d labrum, frontal view e mesosoma, dorsal view f mesopleuron, lateral view g metasoma, dorsal view h ventral view i pygidium, dorsal view. Scale bars: 1.0 mm (a–c, e–h); 0.1 mm (d, i).
Female. Body length: 7.7 mm (Fig.
Head. Length to width ratio = 1.3 (Fig.
Mesosoma. Pronotum reddish-brown and entirely obscured by whitish tomentum. Mesoscutum reddish with wide black medial line and whitish tomentum laterally. Punctation coarse and dense, interspaces narrower than puncture diameters but shiny. Mesoscutellum reddish, round, densely and coarsely punctate, punctures twice as large as on mesoscutum, axillar tooth (free portion of axilla) long and acute but slightly shorter than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum (Fig.
Metasoma. Metasoma entirely reddish. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1 with lateral spots of whitish tomentum at the base and near the apex, T2–T4 with apical bands of tomentum, which are broken in the middle. T2–T4 densely but finely punctate with shiny interspaces and ill-developed depressions. T5 shiny with very fine and dense punctation, whitish tomentum on sides, pseudopygidial area short, with silvery pubescence (Fig.
Male. Body length: 7.6 mm (Fig.
Head. Length to width ratio = 1.2. Mandible light reddish, mandibular apex and preapical tooth dark brown. Labrum similar to that of female. Clypeus reddish, matt and with very fine and dense punctation, interspaces smaller than puncture diameters. Clypeus entirely covered by whitish tomentum. Frons black with basal part from clypeus to antennal tubercles reddish, with dense whitish hair around antennal socket, and with well-developed and sharp frontal keel. Vertex black, with sparse punctures, interspaces shiny and slightly greater than puncture diameters. Antenna reddish with flagellomeres dorsally partly brownish. Flagellomeres slightly shorter than wide (L/W ratio = 0.9), F3 a little longer than other flagellomeres (L/W ratio = 1.05), F2 longer (L/W ratio = 1.3).
Mesosoma. Pronotum black and entirely obscured by whitish tomentum. Mesoscutum black and with whitish tomentum laterally. Punctation coarse and dense, interspaces narrower than puncture diameters but shiny. Mesoscutellum reddish, round, densely and coarsely punctate, punctures slightly larger than on the mesoscutum, axillar tooth (free portion of axilla) long and acute but slightly shorter than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum. Propodeum brownish, very finely sculptured, matt, and laterally with whitish tomentum. Mesopleuron reddish, partly obscured with dense whitish tomentum, coarsely and densely punctate, with interspaces shiny and smaller than puncture diameters (but ill visible under tomentum). Wings brownish with dark brown venation. Legs light reddish or orange, partly covered by whitish tomentum, only tarsal claws brown, tibial spurs on middle and hind legs yellow.
Metasoma. Metasoma entirely reddish. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1 with lateral spots of whitish tomentum at the base and near the apex, T2–T4 with apical bands of tomentum, which are broken in the middle. T2–T4 densely but finely punctate with shiny interspaces and ill-developed depressions. T5 shiny with very fine and dense punctation, whitish tomentum on sides, pseudopygidial area short, with silver pubescence. T6 reddish, bearing reddish pygidial plate, with long yellowish hair (Fig.
All known specimens were collected in Iran: iranicus (-a, -um).
Host and floral associations are unknown.
Recently known only from two nearby localities in Iran and several specimens from other localities (however, not properly localised) from the same country.
North Africa – Morocco, Algeria, and Turkey. It was recorded usually in mountains (High Atlas Mts.). The holotype from “Laussitz”, which means the region on border between Germany and Poland looks very similar to the specimens from Morocco and probably was wrongly labelled (
Morocco: Oukaimeden, 8.VII.1975, (6 ♀), J. Gusenleitner leg., (2 ♀), A. W. Ebmer; 11.VII.1975, (1 ♂, 1 ♀), J. Gusenleitner [all P. Bogusch det.] (
Holotype : ♀, Egypt: Gebel Elba, date unknown, H. Priesner leg. (KUBC).
Paratypes: Chad: Tibesti, Zouar, 11.3.1953, (1 ♂), K. M. Guichard leg. (KUBC); Morocco: 20 km east of Agdz, 20.4.1995, (1 ♀), M. Halada leg. (
This species is probably closely related to E. flavociliatus, with which it shares the following morphological features: very small labral tubercles positioned near the apex, flat and shiny labrum (Fig.
Epeolus priesneri sp. nov. a, c–h holotype, female b, i paratype, male a, b habitus, dorsal view c head, frontal view d labrum, frontal view e mesosoma, dorsal view f mesopleuron, lateral view g metasoma, dorsal view h ventral view i pygidium, dorsal view. Scale bars: 1.0 mm (a–c, e–h); 0.1 mm (d, i).
Female. Body length: 7.5 mm (Fig.
Head. Length to width ratio = 1.3. Mandible yellow, mandibular apex and preapical tooth dark brown (Fig.
Mesosoma. Pronotum reddish-brown and entirely obscured by yellowish tomentum. Mesoscutum black with yellowish tomentum, with dense punctation, interspaces narrower than puncture diameters. Mesoscutellum reddish, round, densely and coarsely punctate, punctures slightly larger than on the mesoscutum, axillar tooth (free portion of axilla) long and acute, slightly longer than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum (Fig.
Metasoma. Metasoma entirely reddish. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1–T4 with entire bands of tomentum. T2–T4 densely but finely punctate with shiny interspaces and ill-developed depressions. T5 shiny with very fine and dense punctation, whitish tomentum on sides, pseudopygidial area short, with silver pubescence (Fig.
Male. Body length: 7 mm (Fig.
Head. Length to width ratio = 1.27. Mandible yellow, mandibular apex and preapical tooth dark brown. Labrum similar to that of female. Clypeus yellow, shiny and with very fine and dense punctation in the middle, interspaces smaller or similar as puncture diameter. Lateral parts of clypeus with punctation of similar size but very sparse with large interspaces. Frons black, with dense whitish hair around antennal socket, and with well-developed frontal keel. Vertex with fine and sparse punctures, interspaces shiny and greater than puncture diameters. Antenna reddish, only scape, pedicel and F1 completely yellow, last four flagellomeres darker, brownish. Flagellomeres slightly shorter than wide (L/W ratio = 0.85), F2 longer than other flagellomeres (L/W ratio = 1.4).
Mesosoma. Pronotum black and entirely obscured by yellowish tomentum. Mesoscutum black with yellowish tomentum, with dense punctation, interspaces narrower than puncture diameters. Mesoscutellum reddish, round, densely and coarsely punctate, punctures slightly larger than on the mesoscutum, axillar tooth (free portion of axilla) long and acute, slightly longer than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum. Propodeum black, very finely sculptured and shiny in the middle, and laterally with yellowish tomentum. Mesopleuron black with reddish macula in the middle, entirely obscured with dense yellowish tomentum, coarsely and densely punctate, with interspaces shiny and similar in size to puncture diameters. Wings brownish with dark brown venation. Legs light reddish or orange, only coxae black, tibial spurs on middle and hind legs yellow.
Metasoma. Metasoma reddish except base of T1. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1–T6 with entire bands of tomentum. T2–T4 densely but finely punctate with shiny interspaces and ill-developed depressions. T7 (pygidium) reddish with large punctures (Fig.
The species is named in dedication to Hermann Priesner (1891–1974), Austrian entomologist, specialist on Heteroptera and Thysanoptera, and the person who collected the holotype. The epithet priesneri is masculine and declined in the genitive case.
Host and floral associations are unknown.
Only four specimens were found in collections, two from Morocco, one from Egypt and Chad (Tibesti Mts.). The species probably occurs in North Africa and in Sahara, distribution south of the Sahara and in the nearest countries in Asia and South Europe is likely but yet unconfirmed.
Turkey (
Europe – Bulgaria, France, Greece, Italy, Portugal, Russia (Volgograd), Spain, Switzerland, the Middle East (
Turkey: Nemrut, 7.–8.VI.1992, (1 ♀), M. Hradský; Karadut env., 1.6.2001, (1 ♂, 1 ♀), K. Deneš [all P. Bogusch det.] (
Synonyms (detailed in
Southern and Central Europe, where the species is rare. Recorded also in the Middle East (
Turkey: Kazan-Kizilcahaman, 30.VI.1976, (1 ♂), Desmier de Chenal; idem, 21.–22.VI.1965, (1 ♂), Holzschuh [M. Schwarz det.]; Sultan Daglari, Yalvaz env., 5.VII.1993, (1 ♀), K. Deneš [all P. Bogusch det.] (
Epeolus transitorius var. seraxensis
Radoszkowski, 1893: 54–55 (syntypes: ♀♀, ♂♂, Serax, Turkmenistan, NHMB,
Syntypes : Turkmenistan, Serax, date unknown, (1 ♂, 1 ♀), Radoszkowski’s collection (NHMB), examined. The female is designated here as a lectotype.
Other material. Iran: Anaesthal, date unknown, (1 ♂), Ritter; Ghezir (Gherir), 4.X.1975, (1 ♀), collector unknown; 30 km west of Karaj, 19.VII.1975, (1 ♂, 3 ♀), 15 km south of Karaj, 21.VII.1975, (1 ♀), P. F. Torchio [all P. Bogusch det.] (
This species is probably closely related to E. transitorius and E. iranicus, and all three species are characterised by the presence of apical (as opposed to submedial) labral tubercles (Fig.
Epeolus seraxensis stat. nov. a, c–h lectotype, female b, i syntype, male a, b habitus, dorsal view c head, frontal view d labrum, frontal view e mesosoma, dorsal view f mesopleuron, lateral view g metasoma, dorsal view h ventral views i pygidium, dorsal view. Scale bars: 1.0 mm (a–c, e–h); 0.1 mm (d, i).
Female. Body length: 9 mm (Fig.
Head. Length to width ratio = 1.3. Mandible light reddish, mandibular apex and preapical tooth dark brown (Fig.
Mesosoma. Pronotum reddish-brown and entirely obscured by whitish tomentum. Mesoscutum black, reddish only laterally, with whitish tomentum laterally and white paramedian bands reaching from the base somewhere behind the middle part of mesoscutum. Punctation coarse and dense, interspaces narrower than puncture diameters but shiny. Mesoscutellum reddish, round, densely and coarsely punctate, punctures twice as larger as on mesoscutum, axillar tooth (free portion of axilla) long and acute but slightly shorter than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum (Fig.
Metasoma. Metasoma entirely reddish. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1 with whitish tomentum at the base and an unbroken apical band, T2–T4 with unbroken apical bands of tomentum. T2–T4 densely but finely punctate with shiny interspaces and well-developed depressions. T5 shiny with very fine and dense punctation, whitish tomentum on sides, pseudopygidial area short, with silver pubescence (Fig.
Male. Body length: 10 mm (Fig.
Head. Length to width ratio = 1.25. Mandible light reddish, mandibular apex and preapical tooth dark brown. Labrum similar to that of female. Clypeus reddish, matt and with very fine and dense punctation, interspaces smaller than puncture diameters. Clypeus entirely covered by whitish tomentum. Frons black with basal part from clypeus to antennal tubercles reddish, with dense whitish hair around antennal socket, and with well-developed and sharp frontal keel. Vertex black, with dense punctures, interspaces shiny and smaller than puncture diameters. Antenna reddish with flagellomeres dorsally partly brownish. Flagellomeres slightly shorter than wide (L/W ratio = 0.95), F2 longer than other flagellomeres (L/W ratio = 1.4).
Mesosoma. Pronotum reddish-brown with black line in the middle, and entirely obscured by whitish tomentum. Mesoscutum black, with whitish tomentum laterally and ill-developed white paramedian bands reaching from the base to the middle part of mesoscutum. Punctation coarse and dense, interspaces narrower than puncture diameters but shiny. Mesoscutellum reddish, round, densely and coarsely punctate, punctures twice as large as on mesoscutum, axillar tooth (free portion of axilla) long and acute but slightly shorter than mesoscutellum. Mesoscutellum with posterior margin extending over propodeum. Propodeum black, very finely sculptured, matt, dorsally and laterally with whitish tomentum. Mesopleuron black, entirely obscured with dense whitish tomentum, coarsely and densely punctate, with interspaces shiny and smaller than puncture diameters (but ill visible under the tomentum). Wings brownish with dark brown venation. Legs light reddish or orange, on dorsal side entirely covered by whitish tomentum, only tarsal claws brown, tibial spurs on middle and hind legs yellow.
Metasoma. Metasoma dark brown. T1 finely and sparsely punctate, interspaces shiny and larger than puncture diameters. T1 with whitish tomentum at the base and an unbroken apical band, T2 with C-shaped pattern of whitish tomentum on each side, apically connecting into an unbroken apical band. T3–T6 with unbroken apical bands of tomentum. T2–T5 densely but finely punctate with shiny interspaces and well-developed depressions. T7 reddish, bearing reddish wide and apically bilobed pygidial plate, with long brown hair on basal part (Fig.
Turkmenistan (type location), in the Middle East currently reported from Iran and Israel.
Synonyms (detailed in
Known from the Middle East and North Africa: Algeria, Egypt, Iran, Jordan, Libya, Sudan, Syria, Turkey, and UAE, (
Algeria: Biskra, 24.IV.1897, (1 ♀), collector unknown [P. Bogusch det.] (
Synonyms (detailed in
South and Central Europe, the Middle East and North Africa (known from Cyprus, Egypt, Jordan, Morocco, Syria, and Turkey), Russia (to the Urals) (
Cyprus: Akrotiri bay, 12.VII.1943, (1 ♂, 3 ♀), 18.VII.1965, (1 ♀), 19.VII.1933, (1 ♀), 10.VII.1943, (1 ♂); Enkomi, 20.VI.1947, (1 ♂, 1 ♀), collector unknown [all P. Bogusch det.] (
Turkey (
Synonyms (detailed in
Widely distributed across Europe (
Iran: Elburs, 25.VII.1977, (2 ♂), J. Gusenleitner [P. Bogusch det.] (
Turkey (
In total, 23 species of the genus Epeolus are known to occur in the studied region. With Triepeolus tristis (Smith, 1854) known from Turkey, 24 species of the tribe Epeolini occur in the Middle East and North Africa. Of these species, several are widely distributed in the whole or in most of the region (e.g. E. subrufescens and E. variegatus) while others have more restricted distributions. For example, several species occur in North Africa only (E. collaris, E. priesneri sp. nov.) and several are known from both sides of the Mediterranean (Southwestern Europe and North Africa) (E. aureovestitus, E. fallax, E. ibericus, E. intermedius). Other species occur only in Asia (E. bischoffi, E. eriwanensis, E. iranicus sp. nov.), including three species known from only one country (Turkey: E. productuloides, E. turcicus, E. warnckei). Several species widely distributed in Europe also occur in the Middle East – E. cruciger, E. schummeli, E. transitorius and E. variegatus. Regarding the number of records, E. flavociliatus is probably the most common and numerous species (recorded from seven countries) in the region, especially in North Africa. Another species, E. subrufescens, was recorded in nine countries, but represented by far fewer specimens. Most of the records presented in this study support the previously known distribution records of the species. Records of E. ibericus and E. intermedius from Morocco are the first finds of both species outside of Europe.
The numbers of species known from the studied countries vary from 0 to 14. The highest numbers of species were recorded in Turkey (14), Morocco (11) and Iran (9), while only one species was recorded in Lebanon, Libya, Oman, and Palestine. No species of Epeolus were recorded from Bahrain, Iraq, Kuwait, Qatar, Saudi Arabia and Yemen. The diversity of Epeolus in every country is the result of multiple factors. First, larger countries and countries with heterogeneous landscape (as Turkey, Morocco or Cyprus) are expected to host more species than countries, where the majority of the landscape is covered by deserts (Saudi Arabia), or small countries (Bahrain, Kuwait). Second, the political situation in some countries makes impossible or exceedingly difficult to collect insects legally and safely, resulting in marked differences in the reported numbers of species between some neighbouring countries. Studied countries with the numbers of known species of Epeolus are shown in Fig.
After three species described from Turkey (E. productuloides, E. turcicus and E. warnckei by
Further studies on Epeolus of the Middle East and North Africa are needed to bring additional information on the distribution of the species and on the changes in their distribution. Several specimens in collections of
I would like to thank to all owners and curators of collections, from which I could study material of Epeolus and to Yulia Astafurova (Saint Petersburg, Russia) and Thomas Onuferko (Toronto, Canada) for their helpful comments. Special thanks belong to Marc Taylor (London, UK) for help with specimens from