Corresponding author: Maxim Yu. Proshchalykin (
Academic editor: M. S Engel
The available information about the cleptoparastic bees of the genus
Astafurova YV, Proshchalykin MY, Niu Z-q, Zhu C-d (2018) New records of bees of the genus
The present paper is part of a series of works dealing with the bees of the cleptoparastic genus
The question of where the zoogeographical boundary exists between the Oriental and the Palaearctic regions in China has been discussed by many researchers working on various groups of animals (
Based on a comprehensive study of specimens in various collections, we catalogue 24 species of the genus
The results presented in this paper are based on 453 specimens collected in the Palaearctic part of China and currently housed in the
The taxonomy and distribution of species generally follow that of
Specimens were studied with a Leica M205A stereomicroscope and photographs were taken with a combination of a stereomicroscope (Olympus SZX10) and a digital camera (Canon EOS70D). Final images represent a composite of several photographs taken at different focal planes and combined using the program Helicon Focus 6. All images were post-processed for contrast and brightness using Adobe® Photoshop®.
The species are presented alphabetically and those that could not be inspected in this paper are quoted from published sources. We use the following abbreviations for collectors:
Unfortunately, we have not examined the type of
Additional species are included in this key because they are widespread in the Palaearctic and may also occur in China. These include
1 | Costal margin of hind wing with 7–14 hamuli. Base of gonocoxite dorsally without impression. Usually large species: total body length 5.0–12.0 mm |
|
– | Costal margin of hind wing with 5–6 hamuli. Base of gonocoxite dorsally with or without impression. Large or small species: total body length 3.5–11.0 mm |
|
2 | Head rounded, about as long as wide. Hind wing with basal ( |
|
– | Head transverse, wider than long. Hind wing with basal ( |
|
3 | Mesoscutum densely punctate, with confluent punctures (areolate) |
|
– | Mesoscutum sparsely punctate, medially with punctures separated by at least a puncture diameter |
|
4 | Head more transverse, 1.2 times as wide as long. Vertex long, distance from top of head to upper margin of lateral ocellus about 2.5–3.0 lateral ocellar diameters as seen in dorsal view. Tyloids on flagellomeres (at least from F4 onward) semicircular across basal 1/5–1/3 and linear across remaining flagellomeres as seen in lateral view. Mesoscutellum sparsely punctate, medially with punctures separated by more than a puncture diameter and often with impunctate areas. T1 completely red. Gonostylus larger, not narrowed apically (Fig. |
|
– | Head less transverse, 1.1 times as wide as long. Vertex shorter, distance from top of head to upper margin of lateral ocellus about two lateral ocellar diameters as seen in dorsal view. Tyloids on flagellomeres semicircular across basal 1/6–1/4, linear portion along remaining flagellomeres not developed. Mesoscutellum densely punctate, with confluent punctures. T1 black or brownish at least on basal 1/3 Gonostylus smaller, distinctly narrowed apically (Fig. |
|
5 | Vertex with a longitudinal carina. Gonostylus smaller, not overlapped apically, as in Figs |
|
– | Vertex without a longitudinal carina. Gonostylus larger, another shape, overlapped apically |
|
6 | Tyloids on flagellomeres (at least from F4 onward) are semicircular across basal 1/3–1/2. T1 with marginal zone very finely and indistinctly punctate. Body length 7.0–10.0 mm |
|
– | Tyloids on flagellomeres weakly developed, very narrow, semicircular across basal 1/7–1/5 of flagellomere. T1 with marginal zone coarsely and distinctly punctate |
|
7 | Head more transverse, 1.2 times as wide as long. Mesoscutum more coarsely punctate (30–75 μm). T2 with marginal zone impunctate. Larger: total body length 7.0–11.0 mm |
|
– | Head less transverse, 1.10–1.15 times as wide as long. Mesoscutum more finely punctate (25–40 μm). T2 with marginal zone distinctly punctate. Smaller: total body length 5.0–7.0 mm |
|
8 | Vertex long, distance from top of head to upper margin of lateral ocellus about three lateral ocellar diameters as seen in dorsal view. Tyloids on flagellomeres cover at least 1/3 part of flagellomere Gonostylus with long apical process (Fig. |
|
– | Vertex shorter, distance from top of head to upper margin of lateral ocellus about two lateral ocellar diameters as seen in dorsal view. Tyloids on flagellomeres not covering more than 1/4 part of flagellomere. Gonostylus another shape at tip, as in Fig. |
|
9 | Tyloids on flagellomeres well developed, covering large part of flagellomere (as seen in lateral view, Fig. |
|
– | Tyloids on flagellomeres weakly developed, covering about 1/3 of flagellomere (as seen in lateral view, Fig. |
|
10 | T4 with marginal zone finely tessellate, without punctures (Fig. |
|
– | T4 with marginal zone distinctly punctate, smooth between punctures (rarely indistinctly tessellate) (Fig. |
|
11 | Base of gonocoxite dorsally without impression |
|
– | Base of gonocoxite dorsally with impression |
|
12 | T1 densely punctate. Gonostylus elongate (Fig. |
|
– | T1 impunctate or with a few fine punctures. Gonostylus another shape |
|
13 | Vertex coarsely and densely punctate, ocello-ocular area with confluent punctures, separated by at most a half puncture diameter |
|
– | Vertex finely and sparser punctate, ocello-ocular area with punctures, separated by at least a puncture diameter |
|
14 | Vertex with longitudinal carina (in |
|
– | Vertex without longitudinal carina |
|
15 | Vertex with well visible longitudinal carina. Felt-like areas on last flagellomeres cover at least 1/2 underside of flagellomere, F2 as long as wide (Fig. |
|
– | Vertex with weakly visible longitudinal carina. Felt-like areas on last flagellomeres cover about 1/3 underside of flagellomere, F2 slightly longer than wide (Fig. |
|
16 | Tyloids on last flagellomeres (at least from F4 onward) usually cover more than 1/2 of ventral flagellar surfaces, often up to 4/5 Membranous portion of gonostylus larger, as in |
|
– | Tyloids on last flagellomeres (at least from F4 onward) usually cover about 1/2 of ventral flagellar surfaces, rarely up to 3/4. Membranous portion of gonostylus smaller (Fig. |
|
17 | Ocello-ocular area densely punctate, with punctures separated by about one puncture diameter. Gonostylus joining apex and partly inner surface of gonocoxite (Fig. |
|
– | Ocello-ocular area sparsely punctate, with punctures separated by 1–3 puncture diameters. Gonostylus joining only apex of gonocoxite (Fig. |
|
18 | F2 shorter, 1.4–1.6 times as long as wide. Tyloids on the flagellomeres extend across 1/4–1/2 of ventral flagellar surfaces. Body length 3.5–6.0 mm |
|
– | F2 longer, 1.7–1.8 times as long as wide. Tyloids on the flagellomeres extend across 1/2–3/4 of ventral flagellar surfaces. Body length 4.0–5.0 mm |
|
19 | T1 densely punctate. Face with appressed white pubescence below and above the antennal toruli |
|
– | T1 impunctate or with sparse punctures (in |
|
20 | Tyloids variable, covering 1/2–4/5 flagellar ventral surfaces. Membranous portion of gonostylus small, triangular (Fig. |
|
– | Tyloids covering from 3/4 to entire ventral flagellar surfaces. Membranous portion of gonostylus large, close to rectangular (Figs |
|
21 | Antenna longer, F2 1.4 times as long as wide (Fig. |
|
– | Antenna shorter, F2 1.2 times as long as wide (Fig. |
|
22 | Pronotum, between dorsal and lateral surfaces, rounded, not angulate (Fig. |
|
– | Pronotum, between dorsal and lateral surfaces, with sharp angle (Fig. |
|
23 | Tyloids on flagellomeres covering less than 1/3 of ventral flagellar surfaces. Membranous portion of gonostylus larger, trapezoidal (Fig. |
|
– | Tyloids on flagellomeres (at least from F4 onward) covering about 1/2–3/4 or entire of ventral flagellar surfaces. Membranous portion of gonostylus smaller, oval or almost square (Figs |
|
24 | Clypeus with fine, simple and sparsely plumose setae, sculpturing clearly visible (Fig. |
|
– | Clypeus with densely plumose setae, partly obscuring sculpturing (Fig. |
|
25 | Antenna short, middle flagellomeres as long as or slightly longer than wide. Tyloids on flagellomeres covering entire of ventral flagellar surfaces. Membranous portion of gonostylus longer, reach penis valve (Fig. |
|
– | Antenna long, flagellomeres (from F3 onward) 1.2–1.3 times as long as wide. Tyloids on flagellomeres (at least from F4 onward) covering about 1/2–3/4 of ventral flagellar surfaces. Membranous portion of gonostylus shorter, not reach penis valve (Fig. |
|
26 | F2 short, 0.9–1.0 times as long as F3. Tyloids on flagellomeres (at least from F4 onward) usually cover entire ventral flagellar surfaces. Gonostylus with trapezoidal membranous portion (Fig. |
|
– | F2 longer, 1.1–1.2 as long as F3. Tyloids on flagellomeres shorter, covering at most 4/5 the ventral flagellar surfaces (in |
|
27 | Tyloids on flagellomeres covering more than 3/4 flagellar ventral surfaces. Gonostylus with large, trapezoidal membranous portion (Fig. |
|
– | Felt-like areas on flagellomeres cover less 1/3 underside of flagellomere. Gonostylus with oval membranous portion or without one |
|
28 | Head less transverse, 1.05 times as wide as long. Mesoscutum sparsely punctate, medially with punctures mostly separated by 1–3 puncture diameters. T1–T3 usually red, rarely terga entirely black. Gonostylus with oval membranous portion (Fig. |
|
– | Head more transverse, at least 1.15 times as wide as long. Mesoscutum very densely punctate, with confluent punctures (areolate). Terga usually wholly black, rare T1 dark red. Gonostylus without membranous portion (Fig. |
|
1 | Hind wing with basal ( |
|
– | Hind wing with basal ( |
|
2 | Vertex less elevated (distance from top of head to upper margin of lateral ocellus less two lateral ocellar diameters as seen in frontal view), with longitudinal sharp carina (Fig. |
|
– | Vertex more elevated (distance from top of head to upper margin of lateral ocellus more two lateral ocellar diameters as seen in frontal view), acarinate, but sometimes with weak (indistinct) longitudinal ridge |
|
3 | Face and gena with sparse, semi-erect, gray pubescence not obscuring integument. T1 with finer punctures (3–10 μm). Body length 6.0–8.0 mm |
|
– | Face and gena with dense, appressed, snow-white pubescence obscuring integument. T1 with coarser punctures (10–30 μm) |
|
4 | Mesoscutum coarsely punctate (25–75 μm). T2 with marginal zone impunctate. Larger: body length 8.0–11.0 mm |
|
– | Mesoscutum relatively finely punctate (25–40 μm). T2 with marginal zone distinctly punctate. Smaller: body length 6.5–8.5 mm |
|
5 | Gena flat. Preoccipital lateral carina developed (Fig. |
|
– | Gena swollen. Preoccipital carina not developed |
|
6 | Vertex shorter, distance from top of head to upper margin of lateral ocellus about 2 lateral ocellar diameters as seen in dorsal view (Fig. |
|
– | Vertex longer, distance from top of head to upper margin of lateral ocellus equal to 2.5–3.0 lateral ocellar diameters as seen in dorsal view (Fig. |
|
7 | T4 with marginal zone impunctate, finely tessellate (Fig. |
|
– | T4 with marginal zone distinctly punctate, smooth between punctures (rarely indistinctly tessellate) (Fig. |
|
8 | Mesoscutum densely punctate, with punctures separated by less than a puncture diameter (Fig. |
|
– | Mesoscutum sparsely punctate, medially with punctures separated by at least 2 puncture diameters (Fig. |
|
9 | Head rounded-rectangular on upper margin, square-shaped as seen in frontal view (Fig. |
|
– | Head uniformly rounded on upper margin, oval as seen in frontal view (Fig. |
|
Females of this vicarious species are very difficult to distinguish morphologically; however, |
||
10 | Mandible simple (without an inner tooth) |
|
– | Mandible bidentate |
|
11 | Head narrower, at most 1.15 times as wide as long (Fig. |
|
– | Head broader, 1.2–1.3 times as wide as long |
|
12 | Face, gena and mesepisternum with gray, sparse, semi-erect pubescence, not obscuring integument. Metasoma coarsely punctate (10–15 μm). Pygidial plate as wide as metabasitarsus. Body length 5.0–8.0 mm |
|
– | Face, gena and mesepisternum with dense, snow-white, appressed, pubescence obscuring integument (Fig. |
|
13 | Pygidial plate at least 1.2 times wider than metabasitarsus, usually dull. Mesoscutum densely punctate, punctures usually separated by less than two puncture diameters. Total body length 7.0–11.0 mm |
|
– | Pygidial plate equal to or narrower than metabasitarsus, shiny. Mesoscutum usually sparsely punctate, disc medially with punctures separated by more than two puncture diameters. Total body length 4.0–9.0 mm |
|
14 | Vertex with longitudinal carina (Figs |
|
– | Vertex without longitudinal carina |
|
15 | Vertex with obvious longitudinal carina (Fig. |
|
– | Vertex with weakly visible longitudinal carina (Fig. |
|
16 | Pygidial plate 1.6–1.7 times as wide as metabasitarsus. Gena wider, 0.8 times as wide as eye in lateral view. Mesoscutum densely punctate, with punctures separated by at most a puncture diameter (Fig. |
|
– | Pygidial plate 1.2–1.5 times as wide as metabasitarsus. Gena narrower, 0.7 times as wide as eye in lateral view. Mesoscutum sparsely punctate, medially with punctures usually separated by 1–2 puncture diameters (Fig. |
|
17 | Head more transverse, 1.30–1.35 times as wide as long; vertex, behind ocelli, not elevated in frontal view. Setae on scape distinctly longer than width of scape. Pygidial plate 1.3–1.5 times as wide as metabasitarsus |
|
– | Head less transverse, 1.20–1.25 times wider than long; vertex, behind ocelli, weakly elevated. Setae on scape shorter than width of scape. Pygidial plate 1.2–1.4 times as wide as metabasitarsus |
|
Females of these species are very difficult to distinguish morphologically; however, |
||
18 | Clypeus densely punctate, punctures separated by less than one puncture diameter. Pronotum, between dorsal and lateral surfaces, rounded, not angulate (Fig. |
|
– | Clypeus sparsely punctate, punctures separated by at least one puncture diameter. Pronotum, between dorsal and lateral surfaces, with sharp angle (Fig. |
|
19 | Hind femur narrow, regularly pointed toward distal end, its length more than 3.5 times its maximum width. Body length 6.0–7.5 mm |
|
– | Hind femur widened in proximal half, its length at most 3 times its maximum width |
|
20 | Vertex, behind ocelli, weakly elevated in frontal view (Fig. |
|
– | Vertex not elevated in frontal view (Fig. |
|
21 | Thorax ventrally with sculpture finer that on sides (Fig. |
|
– | Thorax ventrally with sculpture as coarse as that on sides (Fig. |
|
22 | Vertex longer, distance from top of head to upper margin of lateral ocellus equal to about 3–3.5 times lateral ocellar diameters as seen in dorsal view. Upper half of gena with appressed, dense pubescence obscuring integument |
|
– | Vertex shorter, distance from top of head to upper margin of lateral ocellus equal to about two lateral ocellar diameters as seen in dorsal view. Gena with erect, sparse pubescence |
|
23 | Mesoscutum and mesoscutellum very sparsely punctate, with tiny punctures separated by 1–7 diameters (Fig. |
|
– | Mesoscutum and mesoscutellum more densely punctate, with coarse punctures separated by 1–3 puncture diameters (Fig. |
|
24 | F3 transverse, 0.6–0.7 times as long as wide, as long as F1. Pygidial plate 0.9–1.0 as wide as metabasitarsus |
|
– | F3 square, as long as wide, longer than F1 Pygidial plate 0.6–0.8 as wide as metabasitarsus |
|
25 | Paraocular area with dense, strongly plumose setae below the antennal toruli (Fig. |
|
– | Face with sparse, simple and weakly plumose setae (Fig. |
|
26 | Head more transverse, 1.25 times as wide as long. Labrum trapezoidal, 0.7 times as long as wide. Hind femur strongly enlarged on proximal half, maximum width 0.4 times its length. Body length 5.0–8.0 mm |
|
– | Head less transverse, 1.1 times as wide as long. Labrum semicircular, 0.5 times as long as width. Hind femur weakly enlarged on proximal half, maximum width 0.35 times its length. Body length 4.5–6.5 mm |
|
(Females of this vicarious species are very difficult to distinguish morphologically; however,
(Females of these species are very difficult to distinguish morphologically; however,
CHINA:
*China (Liaoning, Inner Mongolia, Hebei, Beijing, Shanxi, Gansu), Central Asia, Russia, Europe (north to Finland and Sweden), Turkey, Syria, Caucasus, North Africa, Israel, India.
CHINA:
*China (Gansu, Xinjiang), Central Asia, Europe, Russia (south of European part and east to Khakassia Republic), Turkey, Iran, North Africa.
CHINA:
*China (Inner Mongolia, Shanxi), Central Asia, Mongolia, Russia, Europe (north to 64°), Caucasus, Turkey, Iran, Japan, North Africa.
CHINA:
China (*Heilongjiang, *Jilin, *Liaoning, *Inner Mongolia, *Hebei, Tianjin, Beijing, *Shanxi, *Shandong, *Shaanxi, *Ningxia, *Xinjiang), Europe (north to Sweden), Korea, Russia, Caucasus, Turkey, Central Asia, Mongolia.
CHINA:
*China (Xinjiang), Russia (east to Irkutsk Prov.), Mongolia, Central Asia, Caucasus, Turkey, Europe (north to 62°).
CHINA:
*China (Beijing, Shanxi), Central Asia, Russia, Europe (north to 66°), Caucasus, Turkey, Japan.
CHINA:
*China (Heilongjiang, Shanxi), Central Asia, Europe (north to 66°), Russia (east to Far East), Turkey, Near East, Mongolia, Japan.
CHINA:
Meyer, 1920: Yarkand (Xinjiang), as
China (Xinjiang), Central Asia, Russia (east to Yakutia), Pakistan, Mongolia, Europe (north to 63°), Turkey, Israel, North Africa, India.
CHINA:
China (Jilin, Hebei, *Shanxi, *Shaanxi, Anhui, Jiangsu, Shanghai, Shanghai, Zhejiang, Sichuan, Yunnan, Xizang, Guandong).
The female of this species is challenging to distinguish from West-Palaearctic
CHINA:
*China (Gansu), Central Asia, Europe, Russia (European part, Ural), North Africa, Caucasus, Turkey.
CHINA:
*China (Inner Mongolia, Shanxi, Ningxia), Mongolia (Dornod Aimag, Khentii Aimag).
CHINA:
*China (Hebei), Russia (Far East), Japan.
CHINA:
China (*Inner Mongolia, *Hebei, *Shaanxi, Gansu, *Xinjiang), Central Asia, Russia, Europe (north to Finland, Sweden, Denmark, England), Turkey, Syria, Japan.
No material examined.
China (Liaoning).
Known only from the holotype.
CHINA:
*China (Heilongjiang), Central Asia, Mongolia, Russia, North Pakistan, Europe (north to 64°), Caucasus, Turkey, North Africa.
CHINA:
China (*Heilongjiang, *Inner Mongolia, *Hebei, *Tianjin, *Beijing, *Shaanxi, Gansu), Russia (East Siberia, Far East), Mongolia, Japan.
CHINA:
*China (Xinjiang), Tajikistan.
CHINA:
*China (Gansu, Xinjiang), Central Asia, South Europe, Russia (south of European part), Caucasus, Turkey, Iran, Pakistan, India, Israel, United Arab Emirates, Qatar, North Africa.
CHINA:
*China (Gansu, Xinjiang), Central Asia.
CHINA:
China (*Gansu, Xinjiang, Sichuan), Central Asia, Russia, Mongolia, Europe (north to 66°), Caucasus, Turkey, North Africa.
CHINA:
Ascher and Pickering, 2018 (Zhejiang, Jiangsu).
China (*Hebei, *Beijing, *Shaanxi, Jiangsu, Shanghai, Zhejiang, *Sichuan), Russia (Far East).
CHINA:
*China (Inner Mongolia, Gansu), Central Asia, Mongolia, Russia, South Europe, Caucasus, Turkey, Iran, Israel, United Arab Emirates, North Africa, Cape Verde Islands.
CHINA:
*China (Heilongjiang, Liaoning, Beijing, Shaanxi, Qinghai, Zhejiang), Kazakhstan (East Kazakhstan), Russia, Europe (north to S England and Latvia), Caucasus, Turkey, Iran, South Korea, Japan, India.
CHINA:
*China (Gansu), Central Asia.
Genitalia, males, dorsal view.
Genitalia, males, dorsal view.
Head, females, frontal view.
Diagnostic characters of
Diagnostic characters of
Diagnostic characters of
Scutum, females, dorsal view.
Antennae, males.
In total, 33 species of
Checklist of the
Species | Province | Published data | Type of areal | |
---|---|---|---|---|
1 | Gansu, Liaoning, Inner Mongolia, Shanxi | first record | P | |
2 | Xinjiang, Gansu | first record | P | |
3 | China (exactly locality is unknown) |
|
? | |
4 | Inner Mongolia, Shanxi | first record | P | |
5 | Xinjiang, Inner Mongolia, Ningxia, Liaoning, Hebei, Shandong, Shanxi, Shanxi, Heilongjiang, Jilin, Beijing, Tianjin | P | ||
6 | Xinjiang | first record | P | |
7 | Shanxi, Beijing | first record | P | |
8 | Taiwan |
|
O | |
9 | Guandong | Meyer 1927 | O | |
10 | Xinjiang | P | ||
11 | Shanxi, Inner Mongolia | first record | P | |
12 | Sichuan, Shanghai; Hebei, Shaanxi, Shanxi, Jilin, Jiangsu, Anhui, Zhejang, Xizang, Guandong, Yunnan | PO | ||
13 | S. |
Guandong | O | |
14 | Gansu | first record | P | |
15 | Guandong | Meyer 1927 | O | |
16 | Inner Mongolia, Ningxia, Shanxi | first record | P | |
17 | Hebei | first record | P | |
18 | Taiwan |
|
O | |
19 | Gansu, Shanxi, Hebei, Inner Mongolia | P | ||
20 | Liaoning |
|
P | |
21 | Heilongjiang | first record | P | |
22 | Gansu, Inner Mongolia, Shaanxi, Heilongjiang, Beijing, Gansu | P | ||
23 | Xinjiang | first record | P | |
24 | Xinjiang, Gansu | first record | P | |
25 | Sichuan, Shanghai, Shanxi, Hebei, Beijing , Zhejiang, Jiangsu | PO | ||
26 | Gansu, Inner Mongolia | first record | P | |
27 | Xinjiang, Gansu | first record | P | |
28 | Xinjiang, Sichuan, Gansu | P | ||
29 | Taiwan |
|
O | |
30 | Qinghai, Zhejiang, Shaanxi, Heilongjiang, Beijing | first record | P | |
31 | Taiwan |
|
O | |
32 | Guandong |
|
O | |
33 | Gansu | first record | P |
P – Palaearctic species; O – Oriental species; PO – Palaearctic and Oriental species
The majority of the Palaearctic Chinese
The other eight Palaearctic species have smaller distributional ranges.
None of the above Palaearctic species are recorded below 30°N. However, two species,
We are grateful to Maximilian Schwarz and Fritz Gusenleitner (OÖLM) for help during our visit to Austria. We thank Hege Vårdal (