Research Article |
Corresponding author: Maxim Yu. Proshchalykin ( proshchalikin@biosoil.ru ) Academic editor: Thorleif Dörfel
© 2020 Yulia V. Astafurova, Maxim Yu. Proshchalykin, Maximilian Schwarz.
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, Schwarz M (2020) New and little-known species of the genus Sphecodes Latreille (Hymenoptera, Halictidae) from Southeast Asia. ZooKeys 937: 31-88. https://doi.org/10.3897/zookeys.937.51708
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The available information about the cleptoparasitic bees of the genus Sphecodes in Southeast Asia is summarized. Thirty-one species are currently known from this area. Four new species are described: Sphecodes discoverlifei Astafurova & Proshchalykin, sp. nov. (Laos), S. engeli Astafurova & Proshchalykin, sp. nov. (Laos, Vietnam), S. ilyadadaria Astafurova, sp. nov. (Indonesia), and S. pseudoredivivus Astafurova & Proshchalykin, sp. nov. (Laos). Nine species are newly recorded from South East Asia: S. chaprensis Blüthgen, 1927 (Laos), S. howardi Cockerell, 1922 (Malaysia, Myanmar, Thailand), S. kershawi Perkins, 1921 (Indonesia, Malaysia, Myanmar, Thailand), S. laticeps Meyer, 1920 (Thailand, Vietnam), S. montanus Smith, 1879 (Laos), S. sauteri Meyer, 1925 (Laos), S. sikkimensis Blüthgen, 1927 (Laos, Myanmar), S. simlaensis Blüthgen, 1924 (Laos), and S. turneri Cockerell, 1916 (Laos). Based on type specimens, new synonymies have been proposed for Sphecodes kershawi Perkins, 1921 = S. javanensis Blüthgen, 1927, syn. nov.; S. simlaensis Blüthgen, 1924 = S. simlaellus Blüthgen, 1927, syn. nov.; S. laticeps Meyer, 1920 = S. biroi mariae Cockerell, 1930, syn. nov. Lectotypes are designated for Sphecodes biroi Friese, 1909, S. simlaellus Blüthgen, 1927, and S. laticeps Meyer, 1920. The female of Sphecodes sauteri Meyer, 1925, and the male of S. turneri Cockerell, 1916 are described for the first time.
Anthophila, Apiformes, cleptoparasites, fauna, lectotype, taxonomy, distribution
In recent years significant progress has been made towards a better knowledge of the species of Sphecodes Latreille from central and northern Asia (
Southeast Asia is composed of eleven countries of impressive diversity in habitats and landscapes: Brunei, Myanmar (Burma), Cambodia, East Timor, Indonesia, Laos, Malaysia, the Philippines, Singapore, Thailand and Vietnam (Fig.
The first information on the genus Sphecodes from Southeast Asia was published by
In the present paper, based on a comprehensive study of specimens (including primary types) held in various collections, we report additional records of 21 species, with four species described as new and nine species recorded from Southeast Asia for the first time, resulting in a total number of 31 Sphecodes species known from this region (Table
In addition, we describe the female of Sphecodes sauteri Meyer, 1925, and the male of S. turneri Cockerell, 1916 for the first time, propose new synonymies for three specific names (S. kershawi Perkins, 1921 = S. javanensis Blüthgen, 1927, syn. nov.; S. laticeps Meyer, 1920 = S. biroi mariae Cockerell, 1930, syn. nov.; S. simlaensis Blüthgen, 1924 = S. simlaellus Blüthgen, 1927, syn. nov.), and designate lectotypes for Sphecodes biroi Friese, 1909, S. laticeps Meyer, 1920, and S. simlaellus Blüthgen, 1927 in order to clarify the status and diagnosis of type specimens.
Checklist of the Sphecodes species of Southeast Asia including distribution by countries.
Species | Southeast Asia | ||||||||
---|---|---|---|---|---|---|---|---|---|
Indonesia | Laos | Malaysia | Myanmar | Philippines | Singapore | Thailand | Vietnam | ||
No. collecting sites | 7 | 4 | 8 | 3 | 3 | – | 3 | 4 | |
No. bees examined | 11 | 16 | 11 | 3 | 6 | – | 22 | 8 | |
1 | S. bakeri Cockerell | ● | ○● | ||||||
2 | S. binghami Blüthgen | ● | ○ | ||||||
3 | S. biroi Friese | ○● | ● | ○ | ● | ||||
4 | S. bischoffi Blüthgen | ○ | ○ | ||||||
5 | S. brunneipes Friese | ○● | |||||||
6 | S. chaprensis Blüthgen | ● | |||||||
7 | S. discoverlifei Astafurova & Proshchalykin, sp. nov. | ● | |||||||
8 | S. distinctus Meyer | ● | ○ | ||||||
9 | S. duplex Blüthgen | ○● | ● | ○ | |||||
10 | S. engeli Astafurova & Proshchalykin, sp. nov. | ● | ● | ||||||
11 | S. fumipennis Smith | ● | ○ | ||||||
12 | S. howardi Cockerell | ● | ● | ● | |||||
13 | S. ilyadadaria Astafurova, sp. nov. | ● | |||||||
14 | S. insularis Smith | ○ | |||||||
15 | S. javanicus Friese | ○ | |||||||
16 | S. kershawi Perkins | ○● | ● | ● | ● | ||||
17 | S. laticeps Meyer | ○ | ● | ||||||
18 | S. luzonicus Blüthgen | ○ | |||||||
19 | S. malayensis Blüthgen | ○ | |||||||
20 | S. montanus Smith | ● | |||||||
21 | S. pseudoredivivus Astafurova & Proshchalykin, sp. nov. | ● | |||||||
22 | S. redivivus Blüthgen | ○ | |||||||
23 | S. rotundiceps Cockerell | ○ | |||||||
24 | S. samarensis Blüthgen | ● | ● | ○ | |||||
25 | S. sauteri Meyer | ● | |||||||
26 | S. sibuyanensis Cockerell | ○ | |||||||
27 | S. sikkimensis Blüthgen | ● | ● | ||||||
28 | S. simlaensis Blüthgen | ● | |||||||
29 | S. tertius Blüthgen | ○ | |||||||
30 | S. tristellus Cockerell | ○ | |||||||
31 | S. turneri Cockerell | ● | |||||||
Total: | 11 | 10 | 7 | 7 | 7 | 1 | 5 | 3 |
The results presented in this paper are based on 77 specimens collected in Southeast Asia and currently housed in the Natural History Museum (London, UK,
Morphological terminology follows that of
Specimens were studied with a Leica M205A stereomicroscope and photographs taken with a combination of stereomicroscope (Olympus SZX10) and digital camera (Olympus OM-D and Canon EOS70D). 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 (*).
Sphecodes bakeri
Cockerell, 1915: 489, ♀ (holotype: ♀, Philippines, Dapitan, Mindanao, Baker leg.;
Structurally and sculpturally this species is very similar to the male of Sphecodes samarensis Blüthgen, 1927 and the female of S. duplex Blüthgen, 1927, but from the first species it differs by weakly developed antennal tyloids (versus tyloids covering large part of ventral flagellar surface in S. samarensis) and from the second species by the ocello-ocular area (Fig.
Wings with weak yellow-brownish darkening; hind wing with angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with eight hamuli. Lateral preoccipital carina present. Female. Total body length 6–8 mm. Head strongly transverse in front view, ca. 1.3 times as wide as long (Fig.
Indonesia: 1 ♀, NE Sulawesi, 47 km WSW Kotamobagu, Dumoga-Bone N. Pk., Toraut, 211 m, VII.1985, G.R. Else (
*Indonesia, Philippines.
Sphecodes binghami
Blüthgen, 1924: 497, ♀ (holotype: ♀, Myanmar, Pegu Hill, 3.89, coll. Bingham;
The female of this species resembles Sphecodes takaensis Blüthgen, 1927 owing to a similar structure, sculpture and coloration of the body, but it differs from this species by the square F3 (as long as wide), the entirely areolate vertex and the wider pygidial plate which is as wide as metabasitarsus (in S. takaensis F3 0.7–0.8 times as long as wide; vertex with small shiny impunctate spots near ocelli; pygidial plate narrower than metabasitarsus).
Wings with brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 80°, costal margin with eight or nine hamuli. Lateral preoccipital carina present. Female. Total body length 8–9 mm. Head (Fig.
Malaysia: 1 ♀, Malaya, S. Batu Feringgi, Panang, 4.VIII.1955, H.T. Pagden (
*Malaysia, Myanmar.
Sphecodes biroi
Friese, 1909 (nec Meyer, 1925): 181, ♀, ♂ (lectotype (designated here): ♂, N. Guinea, Biro 1899 // Sattelberg, Huon-Golf. // Sphecodes biroi Fiese det., 1908, ♂ // Type // Lectotypus, Sphecodes biroi Friese, 1909, design. Astafurova et Proshchalykin, 2020 <red label>), Fig.
Sphecodes transversus
Cockerell, 1919: 556, ♀ (holotype: ♀, Philippines, Luzon, Mt Makiling, Baker leg.;
Sphecodes latifrons
Cockerell, 1919: 556, ♂ (holotype: ♂, Philippines, Luzon, Baguio, Benguet, Baker leg.;
Sphecodes abnormis
Perkins, 1921: 10–11, ♂ (holotype: ♂, “East Indies”; ?). Synonymized by
Sphecodes amboinensis
Meyer, 1925: 11, ♂ (syntypes: 4 ♂♂, Indonesia, Amboina, 1998, Biro leg.;
Sphecodes bischoffi
Meyer, 1925 (nom. praeocc., nec S. bischoffi Blüthgen, 1924): 11, ♂ (holotype: ♂, Indonesia, Java, Buitenzorg; ZMHB). Synonymized by
This species resembles Sphecodes laticeps Meyer, 1920 in having a similar structure, sculpture and coloration of the body, including the shape of the male genitalia. However, the male differs in the size of tyloids which usually cover the entire ventral flagellar surface or sometimes with a small glabrous spot on basal flagellomeres (versus antennae with well-developed medial glabrous spot on ventral surface of flagellomeres in S. laticeps); females are challenging to distinguish, but S. laticeps has T2 usually more distinctly punctate.
Wings with weak yellow-brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 80°, costal margin with seven hamuli. Lateral preoccipital carina present. Female. Total body length 6.5–7.5 mm. Head (Fig.
Malaysia: 1 ♂, Malaya, Kuala Lumpur, 17.XI.1929, H.T. Pagden (
Indonesia, *Malaysia, Philippines, *Thailand, India, Sri Lanka, New Guinea.
Records of this species in Thailand (
Sphecodes brunneipes
Friese, 1914: 14, ♀ (holotype: ♀, Indonesia, Java, Buitzotg. Schmiedek. leg., Coll. Friese; ZMHB; examined, Fig.
Unlike other species with simple mandibles in the female, this species has a preoccipital carina and a weakly curved basal vein in hind wing.
Wings with brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with seven hamuli. Lateral and dorsal preoccipital carina present. Female. Total body length 6–7 mm. Head (Fig.
Indonesia: 2 ♀♀, Lombok, near Senggigi, 18.V.2012, M. Mokrousov (
Sphecodes chaprensis
Blüthgen, 1927: 96–98, Fig.
This species resembles Sphecodes shillongensis Blüthgen, 1927 and S. simlaensis Blüthgen, 1924, sharing a similar structure and sculpture of the body, including weakly developed antennal tyloids, a densely punctate mesoscutum and scarcely punctate metasomal terga. The species differs from S. simlaensis in having dense facial pubescence obscuring integument above the antennal sockets; from S. shillongensis it differs by shorter antennae with flagellomeres (from F3 onward) ca. 1.1–1.2 times as long as wide (versus 1.3) and shape of the gonostylus with a large membranous part.
Wings hyaline; hind wing with angle between basal (M) and cubital (Cu) veins almost 90°, costal margin with six or seven hamuli. Preoccipital carina absent. Male. Total body length 4.5–5.5 mm. Head transverse, ca. 1.15 times as wide as long (Fig.
Laos: 1 ♂, Phongsaly prov., 21°44'N, 102°12'E, Ban Natsa, 9–17.V.2004, 550 m, P. Pacholatko (
*Laos, India (Bihar).
Holotype
: ♂, laos, Phongsaly prov., Phongsaly env., 21°41'N, 102°06'E, 1500 m, 1–30.VI.2003, P. Pacholatko (PCMS), Fig.
Among the oriental species lacking a preoccipital carina and with five or six hamuli in hind wing the male of the new species is recognizable by having tyloids covering the entire ventral and lateral flagellar surfaces and also in the shape of the gonostylus which has an elongate membranous part; the female is similar to Sphecodes tantalus Nurse, 1903 by combination of the strongly transverse head, the metafemur strongly enlarged in proximal half, the mesoscutum with relatively sparse punctures, the impunctate T1, the narrow pygidial plate and the reddish metasoma, but it differs by not having an elevated vertex as seen in frontal view (in S. tantalus distance from top of head to upper margin of lateral ocellus approximately a lateral ocellar diameter).
Wings hyaline, weak yellowish with light brown stigma and yellowish veins; hind wing with the angle between basal (M) and cubital (Cu) veins 90°, costal margin with five hamuli. Preoccipital carina absent. Female. Total body length 6.0–6.5 mm (Fig.
Male. Total body length 6–7 mm (Fig.
Sphecodes discoverlifei Astafurova & Proshchalykin, sp. nov., male, holotype 36 head, frontal view 37 antennae, lateral view 38 metasoma, dorsal view 39 mesosoma, dorsal view 40, 41 genitalia, dorso-lateral view (40), dorsal view (41) 42 labels. Scale bars: 1.0 mm (36–39), 0.5 mm (40, 41).
This species is dedicated to name of the website https://www.discoverlife.org (creators are J.S. Ascher and J. Pickering), in recognition of its contribution to knowledge of bee biodiversity.
Sphecodes distinctus
Meyer, 1925: 11, ♂ (holotype: ♂, Annam, Laos [Vietnam];
This species is similar to Sphecodes formosanus Cockerell, 1911 in having weakly developed tyloids, a densely punctate mesoscutum (close to areolate) and metasomal terga, but it differs in having a smaller body size (6.5–8.5 mm versus 9–10 mm), number of hamuli (seven or eight versus ten), and usually more developed tyloids (covering sometimes to basal 1/4–1/3 of distal flagellomeres compared to at most 1/5 in S. formosanus). Females of S. takaensis
Wings with weak yellow-brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins almost 80°, costal margin with seven or eight hamuli. Lateral preoccipital carina present. Male. Total body length 6.5–8.5 mm (Fig.
Thailand: 13 ♂♂, Nan, MaeCharim NP Eingang, 18°36'N, 100°58'E, 10–24.V.2012, E. & J. Holzschuh (
Meyer 1925: 11 (Vietnam);
*Thailand, Vietnam.
Annam (type locality of S. distinctus) is actually located in Vietnam, not Laos as Meyer (1925: 11) pointed out.
Sphecodes biroi
Meyer, 1925 (nom. praeocc., nec S. biroi Friese, 1909): 11, ♀ (syntypes: 2 ♀♀, “Singapure [Singapore], Biró 1898 leg.”;
Sphecodes duplex Blüthgen, 1927: 78, replacement name for S. biroi Meyer, 1925 (nec S. biroi Friese, 1909).
Structurally and sculpturally this species is extremely similar to the female of Sphecodes bakeri Cockerell, 1915, but it differs in the ocello-ocular area (Fig.
Wings with yellow-brownish darkening; hind wing with angle between basal (M) and cubital (Cu) veins almost 90°, costal margin with eight hamuli. Lateral preoccipital carina present. Female. Total body length 5–6 mm. Head strongly transverse (Fig.
Indonesia: 1 ♀, Java, Buitzorg [S. duplex Blüthgen det.] (ZMHB); Malaysia: 1 ♀, Malaya, Kuala Sleh, jungle, 15.III.1936, H.T. Pagden (
Meyer 1925: 11 (Singapore);
Indonesia, *Malaysia, Singapore.
This species is probably the unknown female of S. samarensis.
Holotype
: ♀, Laos, Hua Phan Prov., Ban Saleui, Phou Pan Mts., 20°13'30"N, 103°59'26"E, 1350–1900 m, 08.V.2012, C. Holzschuh & locals (
As with members of the Palaearctic hyalinatus species group (
Wings with weak brownish darkening, veins and stigma brown; hind wing with angle between basal (M) and cubital (Cu) veins ca. 90°, costal margin with six hamuli. Preoccipital carina absent. Female. (holotype. Fig.
Male unknown.
This species is dedicated to our colleague Dr. Michael S. Engel (University of Kansas, USA), in recognition of his significant contributions to systematic entomology.
Sphecodes fumipennis
Smith, 1853: 36, ♀ (holotype: ♀, N. India, coll. J.S. Baly;
The male of this species resembles Sphecodes assamensis Blüthgen, 1927 in having a similar size of antennal tyloids and shape of the gonostylus, but it differs from this species by shining interspaces on vertex and mesoscutum (dull in S. assamensis), and the number of hamuli (ten or twelve versus eight). The female differs from other oriental species by combination of the following characters; lack of a preoccipital carina, large body length (9.5–12 mm), eleven or twelve hamuli, mesoscutum mostly punctate-areolate and vertex strongly elevated. With these characteristics the female is similar to the palaearctic S. albilabris (Fabricius, 1793), but it differs in a sparsely punctate T1 disc with punctures separated by 2–6 puncture diameters (versus 0.5–2 in S. albilabris).
Wings with strong brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with eleven or twelve hamuli. Female. Total body length 9.5–12 mm. Head (Fig.
Laos: 2 ♀♀, 1 ♂, Louang Phrabang pr., Ban Song Cha, 1200 m, V. 1999, V. Kuban (
Sphecodes howardi
Cockerell, 1922: 12, ♀ (holotype: ♀, Canton [= Guangdong], China, C.W. Howard Collection, Type No 24885USNM;
This species is most close to Sphecodes kershawi and also resembles S. formosanus Cockerell, 1911, S. takaensis Blüthgen, 1927, and S. binghami owing to similar structure, sculpture, and coloration of the body, including a densely punctate disc and marginal zone of T1 (differences between females of these species are outlined in Table
Differences between females of Sphecodes howardi, S. formosanus, S. takaensis, S. binghami, and S. kershawi.
Characters | Sphecodes species | ||||
---|---|---|---|---|---|
howardi | formosanus | takaensis | binghami | kershawi | |
F3 | About as long as wide | About as long as wide | 0.7–0.8 times as long as wide | About as long as wide | About as long as wide |
Mesoscutum | With coarse and confluent punctures, but medially with interspaces 0.5–1.0 puncture diameter | Densely punctate with punctures separated at most a puncture diameter | With coarse and confluent punctures, but medially with interspaces 0.5–1.0 puncture diameter | With coarse and confluent punctures, but medially with interspaces 0.5–1.0 puncture diameter | Mostly areolate-punctate |
T4 marginal zone | Tessellate | Smooth | Smooth | Smooth | Smooth or unclearly tessellate |
Pygidial plate | Narrower than metabasitarsus | As wide as metabasitarsus | Narrower than metabasitarsus | As wide as metabasitarsus | Narrower than metabasitarsus |
Number of hamuli | 7–8 | 9–10 | 8 | 8–9 | 6–7 |
Distribution | Malaysia, Myanmar, China (Guangdong) | China (Taiwan) | China (Taiwan) | Malaysia, Myanmar | Indonesia, Malaysia, Myanmar, Thailand, China (Macao) |
Wings with brownish darkening; hind wing with angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with seven or eight hamuli. Lateral preoccipital carina present. Female. Total body length 8.5–9.5 mm. Head transverse, ca. 1.25 times as wide as long; vertex elevated with distance from top of head to upper margin of lateral ocellus approximately a lateral ocellar diameter as seen in frontal view; labrum semi-oval, 0.4 times as long as basal width; face and vertex areolate-punctate; paraocular (below and above the antennal sockets), supraclypeal areas and gena with adpressed white pubescence obscuring integument. Mesoscutum and mesoscutellum mostly with areolate punctures (40–75 μm), but medially with a few shining interspaces of approximately a puncture diameter (Fig.
Malaysia: 1 ♀, Titi Serong Perak, 29.III.1930, H.T. Pagden (
*Malaysia, *Myanmar, China (Guangdong).
Holotype
: ♂, indonesia, West Java, Gunung Halimun N.P., Tea Plantation, Citalahab, 850 m, 6.77607N, 106.85401E, 20.VIII.2004, P. Hartmann (
The new species most closely resembles Sphecodes insularis Smith, 1858, from which it differs by having an areolate punctate mesoscutum (with interspaces between punctures up to a puncture diameter in S. insularis).
Wings with weak yellow-brownish darkening, veins and stigma brown; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with seven or eight hamuli. Lateral preoccipital carina well developed. Male. (holotype, Fig.
Female unknown.
The species is named after the author’s daughter Darya Gayday and her husband Ilya Gayday, who recently married. It is to be treated as a noun.
Sphecodes kershawi
Perkins, 1921: 9, ♂ (syntypes: ♂♂, China, Macao, J.C. Kershaw leg.;
Sphecodes javanensis Blüthgen, 1927: 69–70, ♂ (holotype: ♂, Indonesia, Java, Buitenzorg, VIII. 1920; NHMW, examined). Syn. nov.
The male of this species is closest to Sphecodes sibuyanensis Cockerell, 1925 owing to similar short antennae with tyloids covering the entire ventral flagellar surface, a densely punctate metasoma (including marginal zone on T1) and in the shape of the gonostylus, with an elongate membranous part. From S. sibuyanensis it differs by a mostly punctate-areolate mesoscutum (versus a lot of mesoscutellar puncturesseparated by 0.5–1 puncture diameter). The female is closest to S. howardi and is also similar to S. formosanus, S. takaensis, and S. binghami owing to a similar structure, sculpture and coloration of the body, including the densely punctate disc and marginal zone of T1 (differences between females of these species are outlined in Table
Wings with strong brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with six, seven or eight hamuli. Lateral preoccipital carina present. Female. Total body length 7.5–8 mm. Head transverse (Fig.
Indonesia: 1 ♂, Ceylon, Calutara, O.S. Wickwar, 1912-189 (
Indonesia, *Malaysia, *Myanmar, *Thailand, China (Macao).
Sphecodes laticeps
Meyer, 1920: 121, ♀, ♂ (lectotype (designated here): ♂, Formosa, Takao, H. Sauter S.G., 8.12.09 // Sphec. laticeps Meyer det. n. spec., !Type // Lectotypus, Sphecodes laticeps Meyer, 1920, design. Astafurova et al. 2020 <red label>; ZMHB); Paralectotypes: 2 ♀♀, Formosa, Taihorinsho, Sauter S.V., VIII. // Sphec. laticeps Meyer det. n. spec., Type, ZMHB,
Sphecodes candidius
Meyer, 1925: 10, ♀ (holotype: ♀, Taiwan, “Lake Candidius 25./9/-10./10/ 1907”;
Sphecodes biroi mariae
Cockerell, 1930: 162, ♂ (holotype: ♂; Thailand, “Siam, Nam, Jan. 8, 1928 (Cockerel)”;
This species resembles Sphecodes biroi Friese, 1909 and S. samarensis Blüthgen, 1927 owing to a similar structure, sculpture and coloration of the body, including the shape of the male gonostylus. S. laticeps differs from S. samarensis by the shining and more elevated vertex with distance from top of head to upper margin of lateralocellus ca. one and a half or two lateral ocellar diameters as seen in frontal view (versus dull, areolate vertex with distance from top of head to upper margin of lateral ocellus half or one ocellar diameter. The female of S. samarensis is unknown, but these features are suitable for both sexes). The male of S. laticeps differs from S. biroi in having less developed tyloids and a glabrous spot on the ventral surfaces of flagellomeres (versus tyloids usually covering entire ventral flagellar surface or sometimes with small non-setae spot on basal flagellomeres). The females of S. laticeps and S. biroi are difficult to distinguish, but S. laticeps has T2 usually more distinctly punctate.
Wings with brownish darkening; hind wing with angle between basal (M) and cubital (Cu) veins almost 90°, costal margin with seven hamuli. Lateral preoccipital carina present. Female. Total body length 7–7.5 mm. Head strongly transverse (Fig.
Vietnam: 4 ♂♂, Gia Prov., Lai-Contum, Tran Lap, 20 km N Buon Luoi, 22–25.XI.1988, Sharkov (
Sphecodes montanus
Smith, 1879: 27, ♀, ♂ (syntype: ♀, Northern India, Masuri [Uttaranchal: Mussoorie], 7000 ft, B.M. Type HYM.17a549;
This species is closest to Sphecodes kozlovi Astafurova & Proshchalykin, 2015 and S. simillimus Smith, 1873, both displaying a similar form to the male genitalia with a large membranous section of the gonostylus (Fig.
Wings hyaline to weak brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 90°, costal margin with five or six hamuli. Vertex with longitudinal carina; lateral preoccipital carina weakly developed (Fig.
Laos: 1 ♂, Prov. Hua Phan, Phou Pan, Umg. Ort Ban Saleui, 20°13'N, 103°59'E, 1350–1900 m, 10–14.V.2012, C. Holzschuh & locals (
Holotype
: ♀, Laos, Louang Prabang prov., 20°33'N, 102°14'E, Ban Songcha, 1200 m, 24.IV–16.V.1999 (
This species is sculpturally closest to Sphecodes malayensis Blüthgen, 1927, S. redivivus Blüthgen, 1927 and S. sauteri Meyer, 1925 (refer to diagnosis of S. sauteri below) and possibly is the unknown female of S. redivivus owing to a similar sculpture of the hypoepimeral area.
Female (holotype, Fig.
Mesosoma black with legs yellow-brownish to light yellow; wings hyaline, with weak yellowish darkening and light brown stigma and veins; stigma wide, 0.4 times as long as wide; hind wing with angle between basal (M) and cubital (Cu) veins ca. 90°, costal margin with five hamuli; mesoscutum and mesoscutellum (Fig.
Metasomal T1 almost impunctate with a few minute setae pores (Fig.
Sphecodes pseudoredivivus Astafurova & Proshchalykin, sp. nov., female, holotype 104 head, frontal view 105 vertex, dorsal view 106 scutum, dorsal view 107 propodeum, dorsal view 108 hypoepimeral area and mesepisternum, lateral view 109 T1–T3, dorsal view. Scale bars: 1.0 mm (104, 109), 0.5 mm (105–108).
Male unknown.
The species name highlights the morphological similarity with S. redivivus Blüthgen.
Only known from the type locality in Laos.
Sphecodes samarensis
Blüthgen, 1927: 73, Fig.
This species is closest to Sphecodes bakeri (refer to diagnosis of S. bakeri, above). The male is also similar to S. biroi Friese, 1909 and S. laticeps Meyer, 1920 owing to a similar structure, sculpture and coloration of the body, including the shape of the male genitalia. The species differs from S. laticeps by the areolate and less elevated vertex with distance from top of head to upper margin of lateral ocellus half or one of a lateral ocellar diameter as seen in frontal view (versus shining vertex with interspaces between punctures and distance from top of head to upper margin of lateral ocellus one and a half or two lateral ocellar diameters. The female of S. samarensis is unknown, but these features would work in both sexes). From S. biroi the species differs in the shape of tyloids with a glabrous medial spot on ventral surface of flagellomeres (versus tyloids usually covering entire ventral flagellar surface or sometimes with a small non-setae spot on basal flagellomeres). The unknown female is probably closest to S. duplex and S. bakeri.
Wings with weak yellow-brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins almost 80°, costal margin with seven hamuli. Lateral preoccipital carina present. Male. Total body length 5–6.5 mm. Head (Fig.
Indonesia: 1 ♂, Sumatra, 500 m, Sirggalang Annai Valley n.r., 14.X.2005, S. Jakl (
*Indonesia, *Malaysia, Philippines.
Sphecodes sauteri
Meyer, 1925: 10, ♂ (holotype: ♂, “Formosa [Taiwan], Mt. Hoozan, 1910, III, Sauter [leg.]”;
This species is sculpturally closest to Sphecodes malayensis Blüthgen, 1927, S. pseudoredivivus sp. nov. and S. redivivus Blüthgen, 1927 including a scarcely punctate metasomal terga and smoothed hypoepimeral area (differences between males of these species are outlined in Table
Differences between males of Sphecodes sauteri, S. malayensis, and S. redivivus.
Characters | Sphecodes species | ||
---|---|---|---|
sauteri | malayensis | redivivus | |
Head | 1.2 times as wide as long | 1.25 times as wide as long | 1.2 times as wide as long |
Tyloids | Semi-oval across 1/3 basal flagellar surfaces | Semi-oval across 1/2 basal flagellar surfaces | Semi-oval across 1/4 basal flagellar surfaces |
F3 | 1.2 as long as wide | Square | Square |
Hypoepimeral area | Smooth with tiny and sparse punctures | Smooth with a few microscopical punctures | Smooth to finely rugulose with dense punctures |
Number of hamuli | 6-7 | 5 | 5 |
Antennae coloration | Red | Brown | Brown |
Wings with weak yellow-brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 90°, costal margin with six or seven hamuli. Preoccipital carina absent. Female (new). Total body length 6 mm. Head (Fig.
Laos: 1 ♀, Phongsaly prov., Phongsaly env., 21°41'N, 102°06'E, 1500 m, 28.V.-20.VI.2003, V. Kuban (
*Laos, China (Taiwan).
Sphecodes sikkimensis
Blüthgen, 1927: 54, Fig.
This species differs noticeably from other described Oriental species with a lateral preoccipital carina by a combination of large total body length (12–15 mm) and the greatest number (12–15) of hamuli (other large oriental species are usually at most 12 mm in length and have hind wings with at most eleven or twelve hamuli).
Wings with strong brownish darkening and metallic violet lustre; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 80°, costal margin with 12–15 hamuli. Female. Total body length 12–15 mm. Head transverse, 1.25 times as wide as long; vertex elevated with the distance from top of head to upper margin of a lateral ocellus ca. two lateral ocellar diameters as seen in frontal view; labrum short, semi-oval, 0.45 times as long as basal width; face and ocello-ocular area areolate-punctate; paraocular areas and gena with sparse pubescence not obscuring integument. Mesoscutum and mesoscutellum areolate-punctate (50–100 μm). Propodeal triangle (metapostnotum) coarsely reticulate-rugose; mesepisternum reticulate-rugose. Metasoma red, coarsely and densely punctate (ca. 25 μm), sparser on T1. Marginal zone impunctate, except T1 with fine punctures (ca. 10 μm); pygidial plate dull, 1.2 times as wide as metabasitarsus. Male unknown.
Laos: 1 ♀, Prov. Hua Phan, Phou Pan, Umg. Ort Ban Saleui, 20°13'N, 103°59'E, 1350–1900 m, 28.IV.2012, C. Holzschuh (
*Laos, *Myanmar, NE India, China (Guangdong).
Sphecodes simlaensis
Blüthgen, 1924: 514–515, ♀ (syntypes: 2 ♀♀, India, Simla, VIII. and IX.[18]98, Nurse leg.; were not found in
Sphecodes simlaellus
Blüthgen, 1927: 46–48, Fig.
This species is close to the Palaearctic Sphecodes geoffrellus (Kirby 1802) owing to a similar structure, sculpture, coloration of the body and shape of the male gonostylus. Females of S. simlaensis and S. geoffrellus are difficult to distinguish morphologically, but the male of S. simlaensis is easy discerned by the weakly developed tyloids, covering at most 1/4 of the basal ventral surfaces of the flagellomeres, Fig.
Wings with weak yellowish or brownish darkening; hind wing with basal vein strongly curved with angle between basal (M) and cubital (Cu) veins ca. 80°, costal margin with five hamuli. Preoccipital carina absent. Female. Total bodylength 5–5.5 mm. Head (Fig.
Laos: 1 ♂, Phongsaly pr., Phogsaly env., 1500 m, 21°41'N, 102°06'E, VII.2003, Pacholatko (
Sphecodes turneri
Cockerell, 1916: 430, ♀ (holotype: ♀, India, Assam, Shilong, 5.03., B. Turner, 1905-125. 17a.561;
This species differs from other species of the genus by having forewings with two submarginal cells (Fig.
Wings with brownish darkening; hind wing with the angle between basal (M) and cubital (Cu) veins ca. 70°, costal margin with eight or nine hamuli. Female. Total body length 8–9 mm. Head transverse (Fig.
Laos: 1 ♀, 1 ♂, Prov. Hua Phan, Phou Pan, Umg. Ort Ban Saleui, 20°13'N, 103°59'E, 1350–1900 m, 15.IV.2012, C. Holzschuh & locals (
*Laos, India (Meghalaya).
It is noteworthy that the other cleptoparasitic genus Nomada Scopoli (Apidae) also has a small group of species with two submarginal cells (
The most important figures, on which the study is based, are shown in Table
Most species recorded in Southeast Asia have montane distributions and are found up to 1900 m. The range of S. biroi, which is distributed from New Guinea to India, is the widest among Southeast Asian species. Sphecodes simlaensis, S. montanus, S. sikkimensis, and S. fumipennis are also probably widespread in the mountainous areas of the Oriental Region.
Morphologically, a large proportion of Southeast Asian species have close affinities to some of the Palaearctic species or belong to one of the Palaearctic species groups. Sphecodes engeli belongs to the hyalinatus species group (for the composition of Palaearctic species groups see
It is quite certain that new species will be found in further studies, and through synonymy and the association of sexes described as separate species, numerous changes in the species spectrum can be expected in the future.
We are grateful to Fritz Gusenleitner (OÖLM) for help during the Russian co-authors’ visit to Austria, Andrew Grace (Hastings, UK) for checking the English grammar, and David Notton (
This investigation for Yulia Astafurova was supported by the Russian Funds for Basic Research (grant number 19–04–00027) and the state research project AAAA–A17–117030310210–3 and the Presidium RAS program no.41 “Biodiversity of natural systems and biological sources of Russia”.