JHR 84: | 1-28 (202 1) or JOURNAL OF = *20ereewee openscces journal

doi: |0.3897/jhr.84.67150 RESEARCH ARTICLE () I Tymenopter a
e

https://jhr.pensoft.net The International Society of Hymenopteriss REGEARCH

New and little-known bees of the genus Epeolus
Latreille, 1802 (Hymenoptera, Apidae, Nomadinae)
from Mongolia

Yulia V. Astafurova', Maxim Yu. Proshchalykin?

| Zoological Institute, Russian Academy of Sciences, Saint Petersburg 199034, Russia 2 Federal Scientific
Center of the East Asia Terrestrial Biodiversity, Far East Branch of the Russian Academy of Sciences, Vladivostok
690022, Russia

Corresponding author: Maxim Yu. Proshchalykin (proshchalikin@biosoil.ru)

Academic editor: V. Gokhman | Received 9 April 2021 | Accepted 21 April 2021 | Published 24 October 2021
http://zoobank. org/F447C28 D-F4F4-4541-9527-99465AF7D177

Citation: Astafurova YuV, Proshchalykin MYu (2021) New and little-known bees of the genus Epeolus Latreille, 1802
(Hymenoptera, Apidae, Nomadinae) from Mongolia. In: Proshchalykin MYu, Gokhman VE (Eds) Hymenoptera
studies through space and time: A collection of papers dedicated to the 75" anniversary of Arkady S. Lelej. Journal of
Hymenoptera Research 84: 11-28. https://doi.org/10.3897/jhr.84.67150

Abstract

A review of nine species of the bee genus Epeolus Latreille, 1802 (Hymenoptera: Apidae, Nomadinae)
currently known from Mongolia is given. Two new species, F. /eleji sp. nov. and E. mongolicus sp. nov. are
described. The following five known species are newly recorded from Mongolia: E. alpinus Friese, 1893,
E. cruciger (Panzer, 1799), E. melectiformis Yasumatsu, 1938, E. nudiventris Bischoff, 1930, and E. rufi-
cornis Morawitz, 1875. A lectotype is designated for Epeolus tarsalis Morawitz, 1874.

Keywords

Anthophila, Apiformes, cleptoparasites, fauna, new species, Palaearctic, taxonomy

Introduction

Mongolia isa large, landlocked country in eastern Central Asia, covering 1,564,100 km’.
Politically, Mongolia is divided into 21 provinces named “aimags” in addition to the
capital, Ulaanbaatar (Fig. 1). This territory, as part of the Central Asian region, is an
important centre of bee diversity in the Palaearctic (Michener 1979).

Copyright Yu. V.Astafurova, M. 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.

12 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

In recent years, significant progress has been made towards a better knowledge of
the Mongolian species of some genera: Colletes Latreille, 1802 (Kuhlmann and Prosh-
chalykin 2013; Proshchalykin and Kuhlmann 2015), Aylaeus Fabricius, 1793 (Colleti-
dae) (Dathe and Proshchalykin 2016), and Sphecodes Latreille, 1804 (Halictidae) (As-
tafurova and Proshchalykin 2015; Astafurova et al. 2015). In total, 348 bee species are
currently known from Mongolia (Ascher and Pickering 2021). However, taxonomic
information about most Mongolian genera is still fragmentary. We begin here with a
reference to the genus Epeolus Latreille, 1802.

The genus Epeolus includes 109 species spread across much of the globe: they occur
throughout the Holarctic zone, from the west coast of the United States and eastwards
to Europe and as far as Japan. About 65 species are known from North and Cen-
tral America, about 35 from the Palaearctic region, of which 17 species are found in
Europe (Michener 2007; Onuferko 2018; Bogusch and Hadrava 2018). The Epeolus
fauna of Mongolia is particularly under-recorded. Only two species, Epeolus tarsalis
Morawitz, 1874 and E. variegatus (Linnaeus, 1758), have been recorded from this
country, though without precise localities (Friese 1895).

In the present paper, based on a comprehensive study of specimens deposited in
various collections, we report seven additional species, with two species described as
new and five species recorded from Mongolia for the first time, resulting in a total
number of nine Epeolus species known from this country. In addition, we designate a
lectotype for Epeolus tarsalis Morawitz, 1874 in order to clarify the status and diagnosis
of type specimens.

A key to Mongolian Epeo/us has not been included in this paper, it is forthcoming
in a subsequent publication uniting this and the Eastern Palaearctic fauna due to their
extensive species sharing and the need for some additional work in these regions.

Materials and methods

The results presented in this paper are based on 277 specimens collected in Mongo-
lia and currently housed in the Zoological Institute, Russian Academy of Sciences
(St. Petersburg, Russia, ZISP); Oberdésterreichisches Landesmuseum, Biologiezentrum
(Linz, Austria, OLBL) and the personal collection of Maximilian Schwarz (Ansfelden,
Austria, PCMS).

The taxonomy, synonymy and distribution of species follow those of Friese (1895),
Levchenko et al. (2017) and Bogusch and Hadrava (2018). Morphological terminol-
ogy follows that of Engel (2001) and Michener (2007). The density of integumen-
tal punctures is described using the following formula: puncture diameter (in um) /
ratio of distance between punctures to average puncture diameter, e.g., 15-20 um /
0.5—1.5. Abbreviations F, T, and S are used for flagellomere, metasomal tergum and
metasomal sternum respectively. The species are listed alphabetically. The records are
first sorted alphabetically according to the aimags, and then chronologically according
to the locality. Hard brackets are used when certain data are added to specimen label

Epeolus from Mongolia i

Figure |. Administrative map of Mongolia (from Rosa et al. 2020). Aimags | Bayan-Ulgii 2 Uvs 3 Khovd
4 Zavkhan 5 Govi-Altai 6 Khuvsgul 7 Arkhangai 8 Bayankhongor 9 Bulgan 10 Orkhon I 1 Uvurkhangai
12 Umnugovi 13 Selenge 14 Darkhan-Uul 15 Tuv 16 Ulaanbaatar 17 Dundgovi 18 Khentii 19 Govi-
Siimber 20 Dornogovi 21 Dornod 22 Sukhbaatar.

information (e.g. current name of a particular locality). We have used the following
abbreviations for collectors: EN — E. Narchuk; ES — E. Sugonyaev; JH — J. Halada;
IK — I. Kerzhner; KM — M. Kozlov; MH — M. Halada; MK — M. Kadlecova; PK — P.
Kozlov; PT — P. Tymer.

Specimens were studied with an Olympus SZ51 stereomicroscope and photo-
graphs taken with a combination of a stereomicroscope (Olympus SZX10) and digital
camera (Olympus OM-D). Final images are stacked composites using Helicon Focus
6. All images were post-processed for contrast and brightness using Adobe Photoshop.

New distributional records are noted with an asterisk (*).

Taxonomy

Genus Epeolus Latreille, 1802

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

Epeolus alpinus Friese, 1893

Epeolus alpinus Friese, 1893: 34, 9, & (type locality: Goeschenen, Switzerland).
Epeolus variegatus Thomson, 1872 (nom. praeocc., nec Linnaeus 1758): 212, 2 (type
locality: unknown).

14 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

Epeolus glacialis Alfken, 1913: 36, nomen novum for E£. variegatus Thomson, 1872.

Epeolus montanus Bischoff, 1930: 9, &, J (type locality: Warnemiinde, Germany).

Epeolus pilosus Bischoff, 1930: 9-10, &, J (type locality: Rositten [=Rybachij], Kalin-
ingrad Prov., Russia).

Epeolus alpinus Bischoff, 1930 (nom. praeocc., nec Friese 1893): 9-10, & (type local-
ity: Saas, Switzerland).

Material examined. Dornod, 15 km W of Choibalsan, Kerulen River, 770 m,
24.VII.2007, (1 2), JH [PCMS]; Khuosgul, Terkhiyn-Tsaggan Lake, 47°11'N,
99°43'E, 2100 m, 22.VII.2005, (6 2, 2 4), JH & PT [PCMS]; Tuv, 100 km E
of Ulaanbaatar, 20 km NE of Tereltz, Tuul River, 15—21.VII.2003, (6 9, 4 4), JH
[OLBL/PCMS]; Khangaun Mts, 5 km N of Khunt, 20.VII.2005, (1 2), JH [PCMS];
75 km W Ulaanbaatar, dunes, 2.VIII.2005, (1 2), JH [PCMS]; 50 km N of Ulaan-
baatar, E of Mandal, 1180 m, 8-13.VHI.2007, (2 2), JH [PCMS]; Selenge, 90 km
N of Ulaanbaatar, Segnez River, 1450 m, 6—-8.VII.2003, (3 2, 2 3), JH [PCMS];
Ulaanbaatar, Tola River, Urga [=Ulaanbaatar], 30.VI-5.VIH.1905, (10 9, 27 3),
PK [ZISP]; Zuunmod env., 47°11'N, 106°59'E, 1630 m, 27.VII.2004, (3 2), MK
[OLBL/PCMS]; Uvurkhangai, Kholt [375 km SW Ulaanbaatar], Northern Gobi,
15-16.VII.1926, (10 2), PK [ZISP]; 12 km E of Aravaykheer, 46°22'N, 102°49'E,
1800 m, 3.VH.2004, (1 2), JH [PCMS]; Zavkhan, 40 km SW of Uliastay, dunes,
18.VII.2005, (1 3), JH [PCMS].

Distribution. *Mongolia (Dornod, Khuvsgul, Tuv, Selenge, Ulaanbaatar, Uvur-
khangai, Zavkhan); North Africa, Europe, Turkey, Iran, Russia (eastern part to Far East).

Epeolus cruciger (Panzer, 1799)

Nomada crucigera Panzer, 1799: 20, J (type locality: Austria).

Epeolus rufipes Thomson, 1870: 91, 8 (type locality: S-Sweden).

Epeolus similis Héppner, 1899: 355-356, &, & (type locality: Freisenbiittel, Germany).

Epeolus cruciger var. elegans Miller, 1921: 168, 9 (type locality: Arnswalde, Germany).

Epeolus cruciger var. rufiventris Miller, 1921: 168, 9 (type locality: Arnswalde, Ger-
many).

Epeolus marginatus Bischoff, 1930: 11, 9, J (type locality: Warnemiinde, Germany).

Material examined. Dornod, 13 km W of Dash-Balbar, Uldza River, 24.VIII.1975,
(1 2), EN [ZISP]; Khentii, 8 km N Binder, 3—5.VII.1976, (1 3), KM [ZISP]; Khovd,
20 km N Bulgan, Ulyastain-Gol River, 30.VI.1980, (1 3), IK [ZISP]; Khuvsgul,
Dzhargalant, Ider River, 19.VII.1975, (2 9, 2 3), ES [ZISP]; idem, 20.VII.1975,
(2 2), EN [ZISP]; 17 km N of Shine-Eder, 21.VII.1975, (1 4), ES [ZISP]; 25 km
SSW of Muren, 23.VII.1975, (1 2), KM [ZISP].

Distribution. *Mongolia (Dornod, Khentii, Khovd, Khuvsgul); Europe, Turkey,

Iran, Russia (eastern part to Far East).

Epeolus from Mongolia 15

Epeolus leleji sp. nov.
http://zoobank.org/CO20E9AD-B666-46A7-B076-8AB5DA1360F2
Figures 2—4

Material examined. Holotype: 4, SE Mongolia, Sukhbaatar, 100 km SSW of Ba-
ruun-Urt, 1100 m, 30.VII.2007, leg. M. Halada [OLBL/PCMS]. Paratypes: 1 9,
1 4, the same label [ZISP]; 1 2, 200 km SSE of Baruun-Urt, Moltsoy Els, 1250 m,
27.V1I.2007, leg. M. Kadlecova [OLBL/PCMS]; 5 2, Dornogovi, 28 km SE of Chatan-
Bulag, steppe, 3.VIII.2007, M. Halada leg. [OLBL/PCMS]; Umnugovi, 1 2,70 km S
of Saynshand, 1100 m, 5.VHI.2007, leg. M. Kadlecova [OLBL/PCMS].

Diagnosis. This species differs from other Palaearctic species of the genus by hav-
ing forewings with two submarginal cells (versus three cells in other species, except EF.
bischoffi Mavromoustakis, 1954) (Fig. 2A, B). The new species are clearly distinguished
from E. bischoffi by lack of long black dense hair on the whole body, position of labral
teeth closer to apical margin, the reddish female mesosoma (black in E. bischoffi) and
many other characters.

Description. Male (Figs 2, 3). Total body length 6.0 mm; forewing length (with-
out tegula) 3.5 mm. Structure and sculpture. Head (Fig. 3B) transverse, ca 1.35 times
as wide as long. Labrum (Fig. 3C) 1.6 times as wide as long; rounded basally and
laterally, with two small sub-apical teeth, medially (between teeth) slightly depressed,
apical margin straight; integument shiny, coarsely and densely punctate (15-30 pm /
confluent—1). Frons with developed frontal keel. Upper part of frons and vertex shiny,
smooth between punctures (15-25 um / confluent—1.5). Antennae short, Fl ca 1.0
times as long as wide, F2 and F3 0.8 times as long as wide. Mesoscutum shiny and
smooth between coarse punctures (20-40 um / confluent—1). Axilla with short acute
tooth not attaining posterior margin of mesoscutellum. Mesoscutellum with me-
dial longitudinal impression; posterior margin scarcely extending over propodeum.
Metanotum medially with small prominence, extending over propodeum (slightly vis-
ible under tomentum). Mesepisternum with confluent punctures (15—20 um). Propo-
deal triangle finely rugulose; rest vertical part of propodeum smooth. Metasomal tergal
discs shiny and smooth between tiny punctures (ca 15 um / 0.5—1.5); marginal zones
semi-transparent, smooth, with tiny and dense punctures. Pygidial plate (17) shiny,
1.2 times as long as basal width, narrowed toward apex, with shallow punctures; apical
margin slightly curved. Sterna shiny, with dense punctures. Coloration: Head black,
but mandibles yellow-red with dark apex; labrum and clypeus (apically) yellow; anten-
nae reddish with brown scape and pedicel. Mesosoma mostly black; pronotal lobe and
axilla reddish; mesoscutellum with two reddish spots on the sides of medial impres-
sion; legs reddish, spurs pale (ivory); wings hyaline, stigma light brown, veins brown.
Tergal discs dark brown, reddish brown laterally and along marginal zone; marginal
zones yellowish. Pygidial plate (T7) reddish (Fig. 3D). Visible sterna brownish with
yellow marginal zones. Pubescence: Labrum with white plumose pubescence, on api-
cal margin with thin simple setae. Face and genae with dense (obscuring integument)
whitish tomentum (sparser on frons). Vertex with relatively sparse short pubescence.

16 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

A

Figure 2. Epeolus leleji sp. nov., holotype, male A, B habitus, lateral view (A); dorsal view (B). Scale

bars: 1.0 mm.

Pronotum and metanotum with white tomentum obscuring integument. Mesoscutum
and mesoscutellum with creamy tomentum denser and lighter (whitish) peripherally.
Lateral and ventral parts of mesosoma entirely covered with white tomentum. Legs
with dense white pubescence. Tergal marginal zones with uninterrupted white tomen-
tum bands; Tl with wide basal band connected with apical (marginal) band laterally,
T2 only with lateral white tomentum spots connected with apical (marginal) band;
tergal discs with light brownish adpressed pubescence not obscuring integument.
Visible sterna with white tomentum (Fig. 3A).

Epeolus from Mongolia Ly

Figure 3. Epeolus leleji sp. nov., holotype, male A habitus, ventral view B head, frontal view C labrum,
frontal view D T4—T7, dorsal view. Scale bars: 1.0 mm (A); 0.5 mm (B=D).

Female (Fig. 4). Total body length 6 mm; forewing length (without tegula) 3.5—
4.0 mm. Structure and sculpture similar to those of the male. F1 1.6—1.8 times as long
as wide, F2 and F3 ca 1.2 times as long as wide. Mesoscutum shiny and smooth between
coarse punctures (20-30 um / confluent—0.5). T6 mostly hidden under T5, pygidial
plate truncate on apex. S5 wide, straight as seen in lateral view. Processes on sides of S6
normal, with short projections. Coloration of head and mesosoma similar to those of the
male, but antennae entirely reddish-brown and mesoscutellum entirely reddish. Mesoso-
ma reddish; pygidial plate yellow with brownish edging. Pubescence similar to that of the
male, [5 with white tomentum, pseudopygidial area short with silver-like pubescence.

18 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

A f

Figure 4. Epcolus leleji sp. nov., paratype, female A, B habitus, lateral view (A); dorsal view (B) C head,

frontal view D apex of metasoma, ventral view. Scale bars: 1.0 mm (A, B); 0.5 mm (C, D).

Etymology. The new species is named in honor of Prof. Arkady Lelej (Vladivostok,
Russia), an outstanding hymenopterist and our friend.
Distribution. Mongolia (Dornogovi, Sukhbaatar, Umnugovi).

Epeolus from Mongolia 19

Remarks. It is noteworthy that two other cleptoparasitic genera, Nomada Scopoli,
1770 (Apidae) and Sphecodes (Halictidae), also have a small group of species with two
submarginal cells (Proshchalykin and Lelej 2010; Astafurova et al. 2020).

Epeolus melectiformis Yasumatsu, 1938

Epeolus melectiformis Yasumatsu, 1938: 224, 2, 3 (type locality: Ookawa-mura, Tosa,
Shikoku, Japan).

Material examined. Arkhangai, Chuluut Gol River, 47°48'N, 100°19'E, 1940 m,
23.VII.2005, (1 2, 1 3), PT [OLBL/PCMS]; 100 km NE of Tsetserleg, Ogui Lake,
29.VII.2005, (2 3), JH [OLBL/PCMS]; Bulgan, 170 km W of Ulaanbaatar, dunes,
1070 m, 16.VIII.2007, (1 2), JH [OLBL/PCMS]; Dornod, 33 km SE Khalk-Gol,
Khalkin-Gol River, 31.VII.1976, (5 9), KM [ZISP]; 100 km W of Choilbalsan,
820 m, 23.VII.2007, (1 2, 2 4), MH [OLBL/PCMS]; 15 km W of Choibalsan, Ker-
ulen River, 770 m, 24.VII.2007, (17 4), JH [OLBL/PCMS]; 50 km SW Choibalsan,
960 m, 25.VII.2007, (2 3), JH [OLBL/PCMS]; Khentii, 100 km NE of Ondorkhaan,
Kerulen River, 970 m, 22.VII.2007, (22 4), MK [OLBL/PCMS]; Khovd, 75 km WSW
Salkhit, 24.VII.1971, (1 2), IK [ZISP]; Modon-Obr Mts, 30 km ENE of Tsagan-Ula,
25.VII.1971, (2 4), KM [ZISP]; Khuvsgul, 20 km SE of Toson-Tsengel, 25.VII.1975,
(1 2), KM [ZISP]; Sedenge, Ero-Gol River near Dulan-Khan, 4.VIII.1975, (1 2), KM
[ZISP]; Sukhbaatar, 200 km SSE of Baruun-Urt, Moltsoy Els, 1250 m, 27.VII.2007,
(14 3), MH [PCMS]; 210 km SSE of Baruun-Urt, 29.VII.2007, (2 9, 2 4), JH
[OLBL/PCMS]; 100 km SSW of Baruun-Urt, 1100 m, 30.VII.2007, (10 9,8 4), MH
[PCMS]; idem, (1 9), PT [OLBL/PCMS]; 7uv, Khangaun Mts, 5 km N of Khunt,
20.VII.2005, (2 4), JH [PCMS]; 75 km W of Ulaanbaatar, dunes, 2.VIII.2005, (1
3), JH [OLBL/PCMS]; Ulaanbaatar, Tola River, Urga [=Ulaanbaatar], 23.VII.1905,
(1 9, 1), PK [ZISP].

Distribution. *Mongolia (Arkhangai, Bulgan, Dornod, Khentii, Khovd, Khuvs-
gul, Selenge, Sukhbaatar, Tuv, Ulaanbaatar); Russia (Buryatia, Far East), Japan (Hok-
kaido, Honshu, Shikoku, Kyushu, Ryukyu).

Epeolus mongolicus sp. nov.
http://zoobank.org/O2AAC04F-7 1 F7-46BE-9 DEF-8272BC1 D7AB9
Figures 5, 6

Material examined. Holotype: 2°, W Mongolia, Zavkhan, 40 km SW of Uliastay,
dunes, 18.VII.2005, JH (OLBL/PCMS). Paratypes: 11 9, with the same labels
(OLBL/PCMS, 2 2 — ZISP); 3 2, Bulgan, 170 km W of Ulaanbaatar, dunes, 1070 m,
16.VII.2007, MK (OLBL/PCMS); 4 9, 160 km W Ulanbaatar, dunes, 1220 m,
16.VIII.2007, JH (OLBL/PCMS).

20 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

A

Figure 5. Epeolus mongolicus sp. nov., holotype (A, B) and paratype (C), female A-C habitus, lateral
view (A); dorsal view (B, C). Scale bars: 1.0 mm.

Diagnosis. This species is clearly distinguished from other Palaearctic species by
red metasoma with spectacular bright copper-reddish (Fig. 5A, B) or gold-yellowish
(Fig. 5C) tomentum entirely covering terga and well-developed on head and meso-
soma. Unlike other Palaearctic species, the new species does not possess contrast col-

Epeolus from Mongolia 21

oration of pubescence, forming light spots or bands. ‘The species is structurally closest
to E. alpinus and E. cruciger, since it also belongs to E. cruciger species group, and
especially resembles F. a/pinus in having long setae on vertex and labrum with almost
straight apical margin (slightly curved).

Description. Female. Total body length 6.5—-8.0 mm; forewing length (without
tegula) 5.5—7.0 mm. Structure and sculpture: Head (Fig. 6A) transverse, ca 1.3 times
as wide as long. Labrum (Fig. 6B) 1.6 times as wide as long, rounded basally and later-
ally, apical margin slightly curved with small distinct medial tooth; sub-apically with
two well-visible teeth, medially (between teeth) slightly depressed; integument shiny,
coarsely and densely punctate (15-30 um / confluent—1). Clypeus dull, densely and
finely punctate (10-15 um / confluent—0.5), widely shiny and impunctate along api-
cal margin. Frons with developed frontal keel. Upper half of frons densely punctate
(15-25 pm / confluent—0.5), shiny and smooth between punctures. Flagellomeres
ca 1.5 times as long as wide. Mesoscutum and mesoscutellum coarsely and densely
areolate-punctate (25-40 um /confluent—0.5). Axilla pointed apically, but without
distinct tooth (Fig. 6C). Mesoscutellum with medial longitudinal impression; poste-
rior margin scarcely extending over propodeum. Mesepisternum areolate-punctate.
Propodeal triangle shagreened; rest vertical part of propodeum shiny and smoother
(finely tessellate to smooth). Metasomal terga densely and finely punctate (10-15 um
/ 1-2), interspaces smooth and dull; marginal zones transparent under tomentum.
Pseudopygidial area (Fig. GE) short, triangular. Pygidial plate trapezoidal, truncate on
apex. Processes on sides of S6 normal, with short projections. Sterna densely punc-
tured like terga (Fig. 6F). S5 wide, straight as seen in lateral view (Fig. 6D). Colora-
tion: Head mostly black, but mandibles yellow-red with dark apex; labrum entirely
yellow-red; clypeus entirely yellow-red or black on upper half; antennae yellow-red
on basal segments and ventrally. Mesosoma mostly black; mesepisternum entirely
black or partially red (on upper half and ventrally and laterally; pronotal lobe, axilla,
mesoscutellum, metanotum (medially) and legs (including spurs) yellow-red; wings
with brownish darkening, stigma and veins brown. Metasoma yellow-reddish. Py-
gidial plate red with brownish edging. Pubescence: Body without contrast coloration
of pubescence, only with tomentum of approximately the same color: bright cop-
per-reddish (Fig. 5A, B) or gold-yellowish (Fig. 5C). Labrum with thin yellow setae
denser and longer around sub-apical teeth. Face and genae with dense tomentum
obscuring integument (sparser on upper half of frons). Upper half of frons with long
thin setae. Vertex with short thick setae, dense but not obscuring integument. Mesos-
cutum entirely covered with tomentum (tomentum can be strongly shabby). Lateral
and ventral parts of mesosoma, metanotum entirely covered with dense tomentum.
Legs with sparse yellow setae. Metasomal terga entirely covered with dense tomentum
obscuring integument. Pseudopygidial area with silver-like pubescence. Sterna with
golden short and relatively dense setae (not entirely obscuring integument, but denser
on S4 and S5).

Male. Unknown.

Etymology. ‘The specific epithet is named after the country of origin.

Distribution. Mongolia (Bulgan, Zavkhan).

22 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11—28 (2021)

y pogel
amt

Figure 6. Epeolus mongolicus sp. nov., holotype, female A head, frontal view B labrum, frontal view
C mesosoma dorso-lateral view D, E apex of metasoma, lateral view (D); ventral view (E) F metasoma,

ventral view. Scale bars: 0.5 mm.

Epeolus from Mongolia 2S

Epeolus nudiventris Bischoff, 1930
Epeolus nudiventris Bischoff, 1930: 14, 9, 3 (type locality: Mondy, Buryatia, Russia).

Material examined. Khovd, 50 km SSW of Uench, Utyn-Mod, 27.V1.1980, (1 9),
IK [ZISP].
Distribution. *Mongolia (Khovd); Russia (Buryatia).

Epeolus ruficornis Morawitz, 1875

Epeolus ruficornis Morawitz, 1875: 144, 9, J (type locality: Varzaminor near Aykul
Lake, Tajikistan).

Material examined. Dornogovi, 5,65 km SE of Chatan-Bulag, 1020 m, 2.VIII.2007,
(49,13), MK & JH [OLBL/PCMS]; Khovd, 12 km SW of Altai, Bodonchin-
Gol River, 22. VII.1970, (1 @), V. Zaytzev [ZISP]; Govi-Altai, 60 km SE of Bugat,
Khaychi-Bulak, 19.VII.1970, (1 @), IK [ZISP]; Uvurkhangai, 159 km of SW Ara-
vaykheer, 45°11'N, 101°26'E, 1250 m, 5.VII.2004, (1 @), JH [OLBL/PCMS].

Distribution. *Mongolia (Dornogovi, Khovd, Uvurkhangai); Azerbaijan, Tajik-
istan, Turkmenistan, China (Gansu) (Morawitz 1890, 1894).

Epeolus tarsalis Morawitz, 1874

Epeolus tarsalis Morawitz, 1874: 182-183, 9, 3 (lectotype (designated here): 3’, Der-
bent [Dagestan Republic, Russia] // x. ®. Mopasuma [Collection of F Morawitz]
/| Epeolus tarsalis Mor. [handwritten by EF Morawitz] // Lectotypus Epeolus tarsalis
Mor., 4, design. Astafurova & Proshchalykin, 2021 <red label>, ZISP).

Epeolus praeustus Pérez, 1884: 324-326, & (type locality: Pyrenees).

Epeolus rozenburgensis Van Lith, 1949: 105-112, & (type locality: the Netherlands).

Epeolus himukanus Hirashima, 1955: 40-41, J (type locality: Kyushu, Japan).

Epeolus tarsalis ssp. tirolensis Van Lith, 1956: 99, 2 (type locality: Tirol, Austria).

Material examined. Dornod, 55 km NNE of Khavirga, 21.VIH.1975, (1 ¢), KM
[ZISP]; 13 km W of Dash-Balbar, Uldzy River, 24.VIII.1975, (2 2, 1 4), EN [ZISP];
Selenge, Ero-Gol River near Dulan-Khan, 4.VIII.1975, (1 3), KM [ZISP]; 13 km E of
Bayan-Gol, 7.VIII.1975, (1 @), EN [ZISP]; Sukhbaatar, 50 km SSW of Barun-Urta,
19. VIII.1975, (1 4), KM [ZISP]; Juv, 75 km W Ulaanbaatar, dunes, 2.VIII.2005,
(1 3), JH [OLBL/PCMS]; Ulaanbaatar, “Tzorgol-Khayrkhan” [SW Ulaanbaatar],

24 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 84: 11-28 (2021)

23.VII.1909, (1 4), PK [ZISP]; Uvs, Tarialan, 13. VIII.1965, (1 3), Dlabola (OLBL);
Zavkhan, Songino, 28—29.V1I.1965, (1 3), Dlabola (OLBL).

Distribution. Mongolia (*Dornod, *Selenge, *Sukhbaatar, *Tuv, *Ulaanbaatar,
*Uvs, *Zavkhan); Europe, Caucasus, Russia (eastern part to Far East), Korea, Japan
(Honshu, Kyushu).

Remarks. ‘This species was previously reported from North Mongolia (Friese 1895;
Pittioni 1947) without an exact locality.

Epeolus variegatus (Linnaeus, 1758)

Apis variegata Linnaeus, 1758: 577, &, J (type locality: Sweden).

Apis murcaria Christ, 1791: 188-189, 9, J (type locality: Germany).

Apis festiva Christ, 1791: 190-191, 9, J (type locality: Germany).

Epeolus pictus Nylander, 1848: 174-175, 9, J (type locality: Siberia, Russia).
Epeolus productus Thomson, 1870: 91, 9, J (type locality: Sweden).

Material examined. Bayankhongor, 56 km NW of Bayankhongor, 46°33'N, 100°12'E,
2200 m, 12.VII.2004, (1 2), MK [OLBL/PCMS]; 2 km S of Bayankhongor, 46°12'N,
100°43'E, 12.VII.2004, 1880 m, 10.VII.2004, (1 2), JH [OLBL/PCMS]; Selenge,
90 km N of Ulaanbaatar, Segnez River, 1450 m, 6-8.VII.2003, (1 2), JH [OLBL/
PCMS]; Tuv, 50 km N of Ulaanbaatar, E of Mandal, 1180 m, 8—13.VIII.2007, (24 9),
JH [OLBL/PCMS]; Ulaanbaatar, Tola River, Urga [=Ulaanbaatar], 14—16.VII.1905,
(1 2), PK [ZISP].

Distribution. Mongolia (*Bayankhongor, *Selenge, *Tuv, *Ulaanbaatar), North
Africa, Europe, Russia (east to Yakutia), Turkey, Georgia, Central Asia, Iran, Pakistan.

Remark. Friese (1895: 208) determined some specimens from North Mongolia
(without an exact locality) as Epeolus pictus Nylander, 1848, but noted that these speci-
mens differed from FE. variegatus. However, according to Morawitz (1865: 50), Epeolus
pictus is a synonym of £. variegatus, and this is supported by Bogusch and Hadrava
(2018) who re-examined type specimens.

Discussion

The genus Epeolus in Mongolia is poorly known, and this study is the first special
review of this taxon in the country. In total, nine species of Epeolus are now known
from Mongolia (seven of these are newly recorded from the studied region, including
two species that are new to science). For comparison, 17 species are known from Eu-
rope (Bogusch and Hadrava 2018), eleven from Turkey (Bogusch 2018), twelve from
Russia (Levchenko et al. 2017), and only two from China (Morawitz 1890; Niu et al.
2018). Both Epeolus alpinus and E. melectiformis seem to be relatively common and
widespread in Mongolia. Four species, Epeolus alpinus, E. variegatus, E. cruciger, and

Epeolus from Mongolia 25

E. tarsalis are widespread in the Palaearctic, unlike E. melectiformis, which is an eastern
Palaearctic species distributed only from Japan to Khovd (Mongolia). Epeolus ruficornis
and E. nudiventris are xerophilic Central Asian species, and Mongolia is the eastern-
most region of their distribution ranges. The two new species are desert Mongolian
endemics distributed in South-Eastern Mongolia (E. /eleji sp. nov.) and North-Eastern
Mongolia (E. mongolicus sp. nov.).

Unlike other Epeolini, all Epeo/us species are so far known as cleptoparasites of spe-
cies of Colletes (Colletidae). The Mongolian fauna includes 38 Colletes species, of which
more than a half are found only in Mongolia or adjacent territories of China, Kazakh-
stan and Siberia (Russia) (Proshchalykin 2017). Although Colletes are found in all
aimags of Mongolia, the fauna of the steppe and desert zones is the richest one. Most
Colletes species prefer open landscapes with dry and well-warmed soils suitable for
nesting. In addition, some Colletes are oligolectic, visiting a limited taxonomic range of
flowering plants. It is obvious that the biodiversity of cleptoparasites is associated with
their hosts. In this regard, Mongolia, like the rest of Central Asian territories, is very
promising for the study of the genus Epeolus. It is quite certain that new records for
regional faunas and species that are new to science will be found during further studies.

Acknowledgements

We are grateful to Fritz Gusenleitner (OLBL) for his help during our visit to Austria,
Maximilian Schwarz (Ansfelden, Austria) and Esther Ockermueller (OLBL) for pro-
viding Epeolus specimens and Michael Orr (Beijing, China) for checking the English
grammar, as well as to the guest editor (Vladimir Gokhman, Moscow, Russia) and two
reviewers (Max Kasparek, Heidelberg, Germany and Petr Bogusch, Hradec Kralové,
Czech Republic) for their valuable comments, which helped to improve the quality of
this paper.

This investigation was supported by the Russian Foundation for Basic Research
(grant numbers 19-04—00027 and 20-54—44014) and the state research project
AAAA-A19-—119020690101-6.

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