The distribution of the genus Sphecodes Latreille (Hymenoptera, Halictidae) of the Arabian Peninsula and surrounding countries with description of hitherto unknown female of S. atlanticus Warncke, 1992 and male of S. dathei Schwarz, 2010

Yulia V. Astafurova; Maxim Yu. Proshchalykin; Maximilian Schwarz

doi:10.3897/zookeys.872.35361

Research Article

The distribution of the genus Sphecodes Latreille (Hymenoptera, Halictidae) of the Arabian Peninsula and surrounding countries with description of hitherto unknown female of S. atlanticus Warncke, 1992 and male of S. dathei Schwarz, 2010

Yulia V. Astafurova^‡, Maxim Yu. Proshchalykin^§, Maximilian Schwarz^|

‡ Zoological Institute, Russian Academy of Sciences, Saint Petersburg, Russia

§ Federal Scientific Centre for East Asian Terrestrial Biodiversity, Far Eastern Branch of Russian Academy of Sciences, Vladivostok, Russia

| Unaffiliated, Ansfelden, Austria

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

Academic editor: Michael S. Engel

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 YV, Proshchalykin MYu, Schwarz M (2019) The distribution of the genus Sphecodes Latreille (Hymenoptera, Halictidae) of the Arabian Peninsula and surrounding countries with description of hitherto unknown female of S. atlanticus Warncke, 1992 and male of S. dathei Schwarz, 2010. ZooKeys 872: 13-40. https://doi.org/10.3897/zookeys.872.35361

ZooBank: urn:lsid:zoobank.org:pub:DCDEFCB1-E33D-476F-90D7-178BC08C311A

Abstract

This study summarises all available information on the bees of the genus Sphecodes in the Arabian Peninsula and surrounding countries (Israel, Jordan, and Syria). Twenty-six species are currently known from this area, while five species are newly recorded from the Arabian Peninsula: Sphecodes atlanticus Warncke, 1992 (Saudi Arabia, Yemen), S. intermedius Blüthgen, 1923 (UAE), S. nomioidis Pesenko, 1979 (UAE, Oman), S. puncticeps Thomson, 1870 (Saudi Arabia), and S. turanicus Astafurova & Proshchalykin, 2017 (Saudi Arabia). In addition, twelve species are newly recorded from Jordan, six for Syria, and four for Israel. The female of S. atlanticus Warncke, 1992 and the male of S. dathei Schwarz, 2010 are here described for the first time and a lectotype is designated for S. intermedius Blüthgen, 1923.

Keywords

Anthophila, Apiformes, cleptoparasites, fauna, lectotype, taxonomy

Introduction

The present paper is part of a series of studies dealing with the bees of the genus Sphecodes of the territory of the Palaearctic region (Warncke 1992; Bogusch and Straka 2012; Özbek et al. 2015; Astafurova and Proshchalykin 2014, 2015a, b, c, 2016a, b, 2017a, b, c, 2018; Astafurova et al. 2014, 2015, 2018a, b, c, d). The goal of this survey is to improve the knowledge on the taxonomy and distribution of Sphecodes in the Arabian Peninsula and surrounding countries (Israel, Jordan and Syria) (Fig. 1) as an essential foundation for advanced biogeographical investigations.

Figure 1.

Map of the Arabian Peninsula and surrounding lands.

For a long time, the Arabian bee fauna has been one of the lesser sampled faunas of the world. But in recent years significant progress has been made towards a better knowledge of the bees from the Arabian Peninsula, in particular regarding the family Halictidae (Dathe 2009, Engel et al. 2013). A first contemporary inventory of the Halictidae of the Arabian Peninsula was compiled by Ebmer (2008) and Dathe (2009). Later, additional species have been described and recorded by Pesenko and Pauly (2009), Schwarz (2010), Alqarni et al. (2014), Bossert (2017), and Ascher and Pickering (2019) so that there are currently 82 species from 13 genera of family Halictidae known from this area, but the Sphecodes fauna of Arabian Peninsula is particularly under-recorded.

Probably the first information on the genus Sphecodes Latreille from the Arabian Peninsula and its adjacent lands was published by Lepeletier de Saint Fargeau (Lepeletier de Saint Fargeau and Audinet-Serville 1825), who described S. olivieri from ‘Arabie’. Almost two centuries later, in his monograph on the Western Palaearctic Sphecodes, Warncke (1992) recorded several species from Israel, Syria and Lebanon (Table 1). The list of bees of the Arabian Peninsula published by Dathe (2009) included two Sphecodes species: S. olivieri and S. longuloides Blüthgen. In the recently published third volume of the “Arthropod fauna of UAE”, Schwarz (2010) described S. dathei and S. villosulus and recorded S. marginatus Hagens and S. pinguiculus Pérez from the United Arab Emirates. In total, nineteen Sphecodes species have been recorded from the Arabian Peninsula and its adjacent lands so far (Table 1). The genus Sphecodes is not yet documented from Kuwait, Bahrain, or Iraq. Clearly this cosmopolitan genus is present in these countries and it is only a matter of time before the fauna is sampled and recorded.

Table 1.

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Checklist of the Sphecodes species of the Arabian Peninsula and surrounding lands including distribution by countries.

Species		Arabian Peninsula					surrounding lands
Species		UAE	Oman	Qatar	Saudi Arabia	Yemen	Lebanon	Israel	Jordan	Syria
1	S. alternatus Smith							○●	●	○
2	S. atlanticus Warncke				●	●
3	S. barbatus Blüthgen									●
4	S. dathei Schwarz	○●			●	●
5	S. dusmeti Blüthgen	○
6	S. ephippius (Linnaeus)						○	○
7	S. gibbus (Linnaeus)							○●	●
8	S. intermedius Blüthgen	●						○●	●
9	S. longuloides Blüthgen	○
10	S. longulus Hagens							●	○●	○●
11	S. majalis Pérez								●
12	S. marginatus Hagens	○						●	●
13	S. monilicornis (Kirby)							○	○●	●
14	S. nomioidis Pesenko	●	●						○
15	S. olivieri Lepeletier	○●	●	○	●			○●	●
16	S. pellucidus Smith								●	●
17	S. pinguiculus Pérez	○	●		●			○●		●
18	S. puncticeps Thomson				●			○●	●	●
19	S. rubicundus Hagens							●
20	S. rubripes Spinola							○	●	○
21	S. ruficrus (Erichson)							○	●
22	S. rufiventris (Panzer)							○	●
23	S. tadschicus Blüthgen							●
24	S. turanicus Astafurova & Proshchalykin				●
25	S. schenckii Hagens							○	●	●
26	S. villosulus Schwarz	○●			●
Total:		9	4	1	7	2	1	16	15	9
Total:		12					20

Based on a comprehensive study of specimens in various collections, we here list 23 species of the genus Sphecodes, with five species recorded from the Arabian Peninsula for the first time. Additionally, twelve species are newly recorded from Jordan, six species newly recorded from Syria, and four species newly recorded from Israel. The female of S. atlanticus Warncke, 1992 and the male of S. dathei Schwarz, 2010 are here described for the first time and a lectotype is designated for S. intermedius Blüthgen, 1923.

Materials and methods

The results presented in this paper are based on 235 specimens collected in the Arabian Peninsula and surrounding territories and currently housed in the Natural History Museum (London, UK, NHMUK); the Zoological Institute, Russian Academy of Sciences (St. Petersburg, Russia, ZISP); Museum für Naturkunde der Humboldt Universität zu Berlin, Germany (ZISP), Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (SDEI), Oberösterreichisches Landesmuseum, Biologiezentrum, Linz, Austria (OLBL) and the private collection of Maximilian Schwarz (Ansfelden, Austria, OLBL/PCMS). The following acronyms are used for the collections where type specimens are deposited:

BLCU Utah State University, Bee Biology and Systematics Laboratory, Logan, Utah, USA;

ISZP Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland;

MNHN Muséum National d’Histoire Naturelle Paris, France;

MRSN Museo Regionale di Scienze Naturali, Torino, Italy;

MZLU Lund University, Lund, Sweden;

NHMUK Natural History Museum, London, UK;

ZMUK University of Copenhagen, Zoological Museum, Copenhagen, Denmark;

ZSN Zoologische Staatssammlung, München, Germany.

The taxonomy and distribution of species follows that of Warncke (1992), Bogusch and Straka (2012), and Astafurova and Proshchalykin (2017b). Identification keys are available in Warncke (1992), Astafurova and Proshchalykin (2017b) or Astafurova et al. (2018b), except for the two recently described new species (S. dathei and S. villosulus). A detailed synonymy can be found in Astafurova and Proshchalykin (2016b, 2017b). Morphological terminology follows that of Engel (2001) and Michener (2007). The ventral surface of some flagellomeres bear a distinctive patch of sensilla trichodea A (sensu Årgent and Svensson 1982), which we refer to as ‘tyloids’, easily observable under the microscope. Abbreviations F, T, and S are used for flagellomere, metasomal tergum and metasomal sternum respectively. The density of integumental punctures is described using the following formula: puncture diameter (in μm) / ratio of distance between punctures to average puncture diameter, e.g., 15–20 μm / 0.5–1.5. Integumental sculpture other than distinctive surface punctation is described following Harris (1979): areolate – coarse, contiguous punctures; reticulate – superficially net-like or network of raised lines; rugose – irregular, nonparallel, wrinkled raised lines (rugae); rugulose – minutely rugose; strigate – narrow, transverse or longitudinal streaks (strigae), variety of parallel lineations; tessellate – regular network of shallow grooves with flat interspaces.

Specimens were studied with a Leica M205A stereomicroscope and photographs taken with a combination of stereomicroscope (Olympus SZX10) and digital camera (Canon EOS70D). Final images are stacked composites using the program Helicon Focus 6. All images were post-processed for contrast and brightness using Adobe Photoshop.

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

Taxonomy

List of species

Sphecodes alternatus Smith, 1853

Sphecodes alternatus Smith, 1853: 36, ♀ (syntypes: ♀♀, Albania; NHMUK).

Sphecodes punctiventris Hagens, 1882; S. gracilior Morawitz, 1893; S. antigae Tournier, 1901; S. reticulatus var. algeriensis Alfken, 1914; S. alternatus lindbergi Pittioni, 1950 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 6.

Material examined

ISRAEL: 1 ♀, Rehovot s.l., 29.IV.1975, K.M. Guichard (NHMUK 013380375); JORDAN: 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/OLBL/PCMS).

Published records

Warncke 1992: 47, map (Israel, Syria); Ascher and Pickering 2019 (Israel).

Distribution

Israel, *Jordan, Syria; North Africa, South and Central Europe, Russia (east to Khakassia Republic), Turkey, Caucasus, Iran, Central Asia, Kazakhstan, NW China.

Sphecodes atlanticus Warncke, 1992

Figures 5, 9, 10, 19, 22

Sphecodes atlanticus Warncke, 1992: 25, Abb. 17, ♂ (holotype: ♂, Algeria: Hoggar-Geb., Guelta; OLBL/PCMS), examined.

Diagnosis

This species is similar to the Trans-Palaearctic Sphecodes scabricollis Wesmael, 1835 owing to the flat genal area, the developed preoccipital lateral carina, the densely punctate head and mesoscutum, the size and shape of male antennal tyloids, and in the similar gonostylar shape. However, S. atlanticus differs from S. scabricollis by a number of characters outlined in Table 2. In addition to presence of preoccipital lateral carina, S. atlanticus clearly differs from the gibbus species group (S. anatolicus Warncke, 1922, S. gibbus (Linnaeus, 1758), S. nippon Meyer, 1922, S. rufiventris (Panzer, 1798), S. schenckii Hagens, 1882, S. tadschicus Blüthgen in Popov, 1935; see Astafurova et al. 2018a) by a short distance from top of head to upper margin of lateral ocellus (2 lateral ocellar diameters as seen in dorsal view, versus those with a long vertex where this distance is at most 2.5–3.0 diameters).

Table 2.

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Differences between Sphecodes atlanticus Warncke, 1992 and S. scabricollis Wesmael, 1835.

Characters	Sphecodes atlanticus	Sphecodes scabricollis
Both sexes
Distance from top of head to upper margin of lateral ocellus as seen in frontal and dorsal views	about one lateral ocellar diameters (Fig. 5)	about two lateral ocellar diameters (Fig. 3)
Propodeal triangle/metaposnotum	equal (in female) or longer (in male) than mesoscutellum (Figs 9, 10)	distinctly shorter than mesoscutellum (Figs 8)
Metasomal terga	with coarser and denser punctures (Fig. 19)	with fine and sparser punctures, especially on T1 (Fig. 18)
Male
Mesoscutum	punctures separated by at most 1.5–2.0 puncture diameters; polished between punctures (Fig. 9)	areolate (Fig. 8)
Genitalia	gonocoxite dorsally with weak impression; gonostylar process longer (Fig. 22)	gonocoxite dorsally without impression; gonostylar process shorter (Fig. 23)
Female
Paraocular areas	with dense pubescence obscuring integument (Fig. 5)	with sparse pubescence not obscuring integument (Fig. 3)

Description of hitherto unknown female

Total body length 6.5–8.5 mm. Head (Fig. 5) black (except reddish mouthparts); transverse, 1.3 times as wide as long; vertex elevated, distance from top of head to upper margin of lateral ocellus ca. one lateral ocellar diameter as seen in frontal view and ca. two lateral ocellar diameters as seen in dorsal view; F1 and F2 transverse, 0.7–0.8 times as long as wide; F3 as long as wide; face with fine contiguous punctures (10–20 μm), clypeus with shiny interspaces between punctures separated by 0.1–0.5 of a puncture diameter; mandible with an inner tooth; paraocular areas and upper part of gena with dense adpressed, snow-white, plumose pubescence obscuring the integument.

Figures 2–7.

Head, females, frontal view. 2 Sphecodes majalis Pérez 3 S. scabricollis Wesmael 4 S. barbatus Blüthgen 5 S. atlanticus Warncke 6 S. rubripes Spinola 7 S. albilabris (Fabricius). Scale bars: 1.0 mm.

Mesosoma black; mesoscutum with coarse punctures (25–50 μm) separated by at most a puncture diameter (Fig. 10); mesoscutellum with irregular punctures separated by 0.1–4 puncture diameters; mesepisternum densely reticulate-rugose; propodeal triangle coarsely reticulate-rugose with large shiny, smooth interspaces between wrinkles (Fig. 10); lateral parts of propodeum finely and densely strigate or strigate-rugose with granulate interspaces between wrinkles; vertical part of propodeum smooth with coarse and dense punctures; legs reddish or dark brown. Hind wing costal margin with 9–10 hamuli.

Figures 8–11.

Mesosoma (8–10), dorsal view; lectotype labels (11). 8 Sphecodes scabricollis Wesmael, male 9, 10 S. atlanticus Warncke (9 – male, 10 – female) 11 S. intermedius Blüthgen, label of lectotype. Scale bars: 1.0 mm.

Metasoma (Fig. 19) with colouration varying from red on T1–T4 to entirely dark-brown; tergal discs with coarse and dense punctures (20–30 μm/ 0.5–2, sparser on anterior third of T1), marginal zone impunctate except on T1 with dense punctures (10–20 μm / 0.5–2); sterna finely tessellate with coarse setae pores; pygidial plate dull, as wide as metabasitarsus.

Figures 12–15.

Sphecodes dathei Schwarz, male. 12 Head, frontal view 13 antenna, frontal view 14 mesosoma, dorsal view 15 T1, dorsal view. Scale bars: 1.0 mm.

Figures 16–19.

T1 (16, 17), metasoma (18, 19), females, dorsal view. 16 Sphecodes majalis Pérez 17 S. barbatus Blüthgen 18 S. scabricollis Wesmael 19 S. atlanticus Warncke. Scale bars: 1.0 mm.

Material examined

SAUDI ARABIA: 6 ♂♂, Wadi Majarish (below Taif), 12.II.1983, K. Guichard (NHMUK 013380451, 013380453, 013380460, 013380462, 013380466, 013380459); 4 ♂♂, Fayfa, 200 m, 29.I. 1983, K. Guichard (NHMUK 013380452, 013380461, 013380463, 013380464); 1 ♂, Lodar, 800 m, 16.V.1967, K. Guichard (NHMUK 0133804446); 1 ♂, Abu Arish, 26.III.1980, K. Guichard (NHMUK 013380465); 2 ♀♀, 1 ♂, Abu Arish, Jizzan Hot Springs, 25.III.1980, K. Guichard (NHMUK 013380458, 013380441, 013380454), 1 ♂, idem, 28.I.1983 (NHMUK 013380450); 1 ♀, Wadi Maraba, 25.I.1983, 1000 m, K. Guichard (NHMUK 013380457); 1 ♀, Jeddah, Locust Research Station, 17.I.1972, A. Basha (NHMUK 013380442); YEMEN: 1 ♂, Usaifira, 1 mile N Ta’izz, 4.500 ft, 21.XII.1937, H. Scott, E. Britton (NHMUK 013380468), 1 ♂, Wadi Maytam, 12 km SE Ibb, 1600 m, 13°53'N, 44°18'E, 27.X.2005, J. Halada (OLBL/PCMS); 2 ♀♀, 3 ♂♂, Hawf NE Albhaydah, 200–730 m, 16°53'N, 53°05'E, 14.X.2005, J. Halada (OLBL/PCMS); 2 ♀♀, 4 ♂♂, 20 km S Taizz, 1200 m, 13°30'N, 43°57'E, 24.X.2005, J. Halada (OLBL/PCMS); 2 ♂♂, Jabal Bura, NEE Al Hudaydah, 200–800 m, 14°52'N, 43°24'E, 30.X-1.XI.2005, J. Halada (OLBL/PCMS); 1 ♂, Wadi Aniz, SSW Sana, 1520 m, 14°60'N, 44°09'E, 7.X.2005, J. Halada (OLBL/PCMS).

Distribution

*Saudi Arabia, *Yemen; Algeria, the Canary Islands.

Sphecodes barbatus Blüthgen, 1923

Figures 4, 17

Sphecodes barbatus Blüthgen, 1923: 497–498, ♀ (holotype: ♀, Turkey, Ak-Chehir; ZSM).

Diagnosis

Sphecodes barbatus is very similar to S. majalis. The two species are easily separable in the female, but males are difficult. The female S. barbatus differs from S. majalis by denser, distinctly plumose pubescence on paraocular areas and clypeus (Fig. 4) (sparser, weakly plumose or simple pubescence in S. majalis, Fig. 2) and by a distinctly (Fig. 17) punctate T1 (sparse and tiny punctures in S. majalis, Fig. 16).

Material examined

SYRIA: 1 ♀, Syria, 40 km NE Damaskus, 22.V.1996, H. Halada (ZISP); 2 ♂♂, Slenfe, 1200 m, 19.IV.1986, K.M. Guichard, (NHMUK 013380371, 013380372).

Distribution

*Syria; Greece, Turkey.

Remarks

Warncke (1992) interpreted Sphecodes barbatus as a subspecies of S. majalis Pérez, 1903, but later this taxon was restored as a valid species (Bogusch and Straka 2014a).

Sphecodes dathei Schwarz, 2010

Figures 12–15, 24

Sphecodes dathei Schwarz, 2010: 483–486, ♀, plates 1–12 (holotype: ♀, United Arab Emirates, Wadi Shawkah, 25°06'N, 56°02'E, 9–24.VI.2007, in water trap, A. van Harten leg.; SDEI), examined.

Diagnosis

The species is similar to Sphecodes crassus Thomson, 1870 owing to the wide female metafemur (strongly enlarged in the basal half); strongly transverse female head; sparsely punctate mesoscutum in both sexes, weakly developed male antennal tyloids (usually covering less than 1/3 of ventral flagellar surfaces). The female of Sphecodes dathei differs from S. crassus by dense, apressed, snow-white, plumose pubescence obscuring integument in paraocular areas (sparse, simple pubescence not obscuring integument in S. crassus); the male differs by densely and relatively coarsely punctate T1 (in S. crassus T1 usually with a few fine punctures, rarely with relatively coarse and dense punctures). Both species have similar gonostylar shape, but S. dathei has a narrower, trapezoidal membranous portion of the gonostylus (wider, close to oval in S. crassus, Fig. 25).

Figures 20, 21.

Habitus, females, lateral view. 20 Sphecodes albilabris (Fabricius) 21 S. rubripes Spinola. Scale bars: 1.0 mm.

Figures 22–25.

Genitalia, males, dorsal view. 22 Sphecodes atlanticus Warncke 23 S. scabri collis Wesmael 24 S. dathei Schwarz 25 S. crassus Thomson. Scale bars: 0.25 mm.

Description of hitherto unknown male

Total body length 5.0–6.5 mm. Head (Fig. 12) black (except reddish mouthparts and brownish antenna); weakly transverse, 1.1 times as wide as long; vertex not elevated; distance from top of head to upper margin of lateral ocellus ca. two lateral ocellar diameters as seen in dorsal view; antenna (Fig. 13) reaches posterior margin of mesoscutum; F1 transverse, 0.6 times as long as wide; F2 long, 1.7 times as long as wide; remaining flagellomeres 1.2–1.3 times as long as wide; tyloids weakly developed (on F2–F4 covering less than 1/6 of ventral flagellar surfaces and from F5 onward covering less than 1/3); clypeus, frons, supraclypeal and paraocular areas with fine contiguous punctures (10–20 μm); ocello-ocular area and gena with shiny interspaces, punctures separated by 0.5–1 a puncture diameter; face below and above the antennal toruli with dense adpressed snow-white plumose pubescence obscuring integument; gena with similar pubescence, but not obscuring integument.

Mesosoma (Fig. 14) black; mesoscutum and mesoscutellum with punctures (20–25 μm) separated by 0.5–4 puncture diameters; mesepisternum and hypoepimeral area densely reticulate-rugose; propodeal triangle (Fig. 14) and vertical part of propodeum coarsely reticulate-rugose with shiny, smooth interspaces between wrinkles; lateral parts of propodeum coarse reticulate- to strigate-rugose with shiny interspaces between wrinkles; legs dark brown, but tarsi and partially tibia yellow or reddish. Hind wing costal margin with 5 hamuli.

Metasomal T1–T3 red (T1 black basally, T3 – apically); tergal discs (Fig. 15) with dense punctures (10–15 μm / 0.5–1), becoming sparse along marginal zone on T1; marginal zones smooth, impunctate; sterna with numerous microscopic setae pores; gonocoxite dorsally with a deep impression; membranous portion of gonostylus small, trapezoidal (Fig. 24).

Material examined

SAUDI ARABIA: 1 ♀, Wadi Majarish, 800 m, 12.II.1983, K. Guichard (NHMUK 013380455); UNITED ARAB EMIRATES: 1 ♀, Hatta, 24.IV.1992 (NHMUK 013380414); 1 ♂, idem, 19–20.V.1988 (NHMUK 013380431); 7 ♂♂, idem, 14.IV.1990, I. Hammer (NHMUK 013380428, 013380429, 013380430, 013380432, 013380433, 013380434, 013380430); YEMEN: 1 ♀, Lawdar, NE Aden, 1140 m, 13°53'N, 45°48'E, 28.X.2005, J. Halada (OLBL/PCMS).

Published records

Schwarz 2010: 483 (United Arab Emirates).

Distribution

United Arab Emirates, *Saudi Arabia, *Yemen.

Sphecodes gibbus (Linnaeus, 1758)

Sphex gibba Linnaeus, 1758: 571, ♀ (syntypes: ♀♀, Sweden; ZMUK).

Apis glabra Füessly, 1775; Andrena ferruginea Olivier, 1789; Apis gibbosa Christ, 1791; Melitta sphecoides Kirby, 1802; M. picea Kirby, 1802; Andrena austriacaFabricius, 1804; Dichora analis Illiger, 1806; Sphecodes apicatus Smith, 1853; S. nigripennis Morawitz, 1876; S. sutor Nurse, 1903; S. gibbus var. rufispinosus Meyer, 1920; S. g. var. turkestanicus Meyer, 1920; S. castilianus Blüthgen, 1924; S. pergibbus Blüthgen, 1938; S. lustrans Cockerell, 1931; S. angarensis Cockerell, 1937 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 17.

Material examined

JORDAN: 1 ♀, Jordan Valey, Dayr Alla, 27.IV.1996, M. Halada (OLBL/PCMS); 1 ♀, N. Shuna env., 20–22.IV.1996; 1 ♀, idem, 29–30.IV.1996, M. Halada (OLBL/PCMS); SYRIA: 1 ♂, 20 km NE Latakia, 25.V.1996, M. Halada (OLBL/PCMS); ISRAEL: 1 ♀, Rehovot s.l., 29.IV.1975, K.M. Guichard (NHMUK 1975-248, 013380378); 1 ♀, Ein Gedi, 200 m, 11.III.1975, K.M. Guichard (NHMUK 1975-154, 013380379); 1 ♂, Jericho (Wadi Quilt), 250 m, 13–22.V.1975, K.M. Guichard (NHMUK 1975-248, 013380373); 1 ♀, Jericho, 200 m, 6–27.III.1975, K.M. Guichard (NHMUK 1975-154, 013380374).

Published records

Warncke 1992: 30 (Israel).

Distribution

Israel, *Jordan; North Africa, Europe (north to 63°), Russia (east to Yakutia), Turkey, Iran, Pakistan, Central Asia, Kazakhstan, Mongolia, NW China, India.

Sphecodes intermedius Blüthgen, 1923

Figure 11

Sphecodes intermedius Blüthgen, 1923: 500 (lectotype (designated here): ♂, Type <red label> // Caucas Portz // Sph. intermedius ♂, Type., P. Blüthgen det. // <blue circle> // Lectotypus, Sphecodes intermedius Blüthgen, 1923, ♂, des. Astafurova & Proshchalykin, 2018; paralectotype: ♀, Type <red label> Sph. intermedius ♀, Type, P. Blüthgen det. // Paralectotypus, Sphecodes intermedius Blüthgen, 1923, ♀, des. Astafurova & Proshchalykin, 2018; ISZP, examined, Fig. 11).

Sphecodes lactipennis Meyer, 1925 (Synonym).

Diagnosis

See Astafurova et al. 2018a: 20.

Material examined

UNITED ARAB EMIRATES: 1 ♂, Hatta (Hotel), 28. IV.1989, (NHMUK 013380370); 1 ♂, idem, 23.VIII.1991 (NHMUK 013380409); 1 ♂, idem, 14.IV.1990, I. L. Hamer [D. Baker det., 1992 as S. punctatissimus Meyer] (NHMUK 013380361); ISRAEL: 1 ♀, Jerusalim, 16.VII.1930, S. Bodenheimer [det. Blüthgen] (MNHB); 1 ♀, Tiberias, 200 m, 22.III.1975, K.M. Guichard (NHMUK 013380410); 1 ♀, Jericho (Hisham Palace), 200 m, 8.III.1975, K.M. Guichard (NHMUK 1975-248, 013380408); JORDAN: 1 ♀, N. Shuna env., 20–22.IV.1996; 1 ♀, idem, 29–30.IV.1996, M. Halada (OÖLM)

Published records

Ascher and Pickering 2019 (Israel)

Distribution

*United Arab Emirates, Israel, *Jordan; North Africa, South Europe (east to Ukraine), Russia (south of the European part, Urals), Caucasus, Turkey, Kazakhstan, Central Asia, Pakistan, China (Gansu).

Remarks

Sphecodes intermedius Blüthgen, 1923 was described from specimens of both sexes collected in “Caucas” [Caucasus] (Fig. 11). There are two specimens (female and male) in ISZP from this locality, which correspond to the original description of P. Blüthgen. One of these specimens (male) is designated here as a lectotype of S. intermedius to avoid any confusion about the status of the type specimens and to properly diagnose this species.

Sphecodes longulus Hagens, 1882

Sphecodes longulus Hagens, 1882: 226, Fig. 25, ♂ (syntypes: ♂♂, Germany; ? Dominican monastery, Venlo, Nederland).

Sphecodes longulus var. eupidus Hagens, 1882; S. nitidulus Hagens, 1882; S. subfasciatus Blüthgen, 1934; S. amakusensis Yasumatsu & Hirashima, 1951; S. sabulosus Tsuneki, 1983; S. crassicornis Tsuneki, 1983; S. tsunekii Haneda, 1994 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 21.

Material examined

JORDAN: 1 ♀, 30 km N Tafila, 2.V.1996, M. Halada (OLBL/PCMS); 1 ♂, 20 km SW Madaba, 26.V.2007, 400 m, Z. Kejval (OLBL/PCMS); 1 ♂, Ajlun, 35 km W Jarash, 850 m, Z. Kejval (OLBL/PCMS); 1 ♀, 20 km S North Shuna Tall al Arbatin, 19.IV.1996, M. Halada (OLBL/PCMS); ISRAEL: 1 ♂, Dafna, 27.V.1991, K. Warncke (OLBL/PCMS); 1 ♂, North Galeleya, Nature Reserve ”Khule”, 23.V.1968, V. Trjapitzin (ZISP); 1 ♀, 5 km W Jericho, Wadi Qelet, St. Georg Mon., 6.V.1996, O. Niehuis (OLBL/PCMS); SYRIA: 1 ♂, Damask, 20–21.V.1980, M. Halada (OLBL/PCMS).

Published records

Warncke 1992: 17 (Syria); Ascher and Pickering 2019 (Jordan).

Distribution

*Israel, Jordan, Syria; Europe (north to Finland, Sweden, Denmark, England), Russia (east to Far East), Turkey, Iran, Central Asia, Kazakhstan, China, Japan.

Sphecodes majalis Pérez, 1903

Figures 2, 16

Sphecodes majalis Pérez, 1903: 219, ♀, ♂, (syntypes: ♀, ♂, France, Spain; MNHN).

Sphecodes gracilior Pérez, 1903; S. opacifrons Pérez, 1903; S. problematicus Schulz, 1906 (Synonyms).

Diagnosis

Refer to the diagnosis S. barbarus, above.

Material examined

JORDAN: 35 ♀ ♀, 5 ♂♂, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS); 2 ♀, 10 km N Jarash, 20.IV.2002, M. Snizek (OLBL/PCMS); 1 ♀, Ajlun S of Anjara, 27.IV.2002, M. Snizek (OLBL/PCMS).

Distribution

*Jordan; North Africa, South Europe, Russia (south of the European part), Turkey, Iran.

Sphecodes marginatus Hagens, 1882

Sphecodes marginatus Hagens, 1882: 223, Fig. 18, ♂ (syntypes: 2 ♂, Germany: Cleve; ? Dominican monastery, Venlo, Nederland).

Sphecodes atratus Hagens, 1882; S. nigritulus Hagens, 1882; S. biskrensis Pérez, 1903 (Synonyms).

Diagnosis

This species belongs to the miniatus species group (S. creticus Warncke, 1992, S. haladai Warncke, 1992, S. larochei Warncke, 1992, S. marginatus Hagens, 1882, S. miniatus Hagens, 1882, S. nomioidis Pesenko, 1979, S. schawrzi Astafurova & Proshchalykin, 2014, and S. sandykachis Astafurova & Proshchalykin, 2018), with the same length and transverse F1–F3 in females. Among species of this group S. marginatus is most close to S. miniatus and S. nomioidis as they have a similar sculpture and structure of the body. Hence females of the three species are challenging to distinguish, but the male differs from the other two species by smaller triangular gonostylus. Differences between these three species are outlined by Bogusch and Straka (2012) and between females of this species group by Astafurova et al. (2018c).

Material examined

ISRAEL: 1 ♀, 1 ♂, Jerusalim, 18.VI.1930; 1 ♂, idem, 10.VI.1931, S. Bodenheimer [det. Blüthgen] (MNHB); JORDAN: 1 ♀, W Jordan Valey, Mubalath, 27.IV.1996, M.Halada (OLBL/PCMS); 1 ♀, n. Shuna, 20–22.IV.1996, M. Halada(OLBL/PCMS); 2 ♂♂, NW of Ailun, 850 m, 20.V.2007, Z. Kejval (ZISP); 1 ♀, Jericho (Wadi Quilt), 250 m, 6.III.1975, K.M. Guichard (NHMUK 1975-154, 013380467).

Published records

Schwarz 2010: 486 (United Arab Emirates); Ascher and Pickering 2019 (United Arab Emirates).

Distribution

United Arab Emirates, *Israel, *Jordan; North Africa, Europe (north to Germany and Denmark, east to Belarus).

Sphecodes monilicornis (Kirby, 1802)

Melitta monilicornis Kirby, 1802: 47, ♂ (syntypes: ♂♂, England, NHMUK).

Sphecodes maculatus Lepeletier de Saint Fargeau, 1841; S. subquadratus Smith, 1845; S. ruficrus Dalla Torre, 1896; S. hanuman Nurse, 1903; S. monilicornis var. nigerrima Blüthgen, 1927; S. caucasicus Meyer, 1920; S. cephalotes Meyer, 1920; S. smyrnensis Meyer, 1920; S. monilicornis quadratus Meyer, 1920; S. monilicornis berberus Warncke, 1992; S. quadratus cephalotiformis Pittoni, 1950 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 24.

Material examined

SYRIA: 1 ♀, 50 km W Homs, 12.V.1996, M. Halada (OLBL/PCMS); 1 ♀, 60 km S Damask, Khabab, 14.V.1996, M. Halada (OLBL/PCMS); 1 ♀, 20 km S North Shuna Tall al Arbatin, 19.IV.1996, M. Halada (OLBL/PCMS); 1 ♀, 10 km W Jarasch, 1.V.1996, M. Halada (OLBL/PCMS); 2 ♀♀, Jisr ash Shunhur, 26.V.1996, M. Halada (OLBL/PCMS); 4 ♂♂, 20 km NE Latakia, 25.V.1996, M. Halada (OLBL/PCMS); 1 ♂, 30 km W Damask, 19.VI.2000, M. Halada (OLBL/PCMS); JORDAN: 1 ♀, Jarash env., 1.V.1996, M. Halada (OLBL/PCMS); 1 ♀, 10 km N Jarash, 1.V.1996, M. Halada (OLBL/PCMS); 1 ♂, 16 km WN Aijun, 600 m, 21.V.2077, Z. Kejval (OLBL/PCMS); 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS).

Published records

Ascher and Pickering 2019 (Jordan).

Distribution

Jordan, *Syria; North Africa, Europe (north to 64°), Russia (east to Far East), Caucasus, Turkey, Iran, Pakistan, Central Asia, Kazakhstan, Mongolia, China.

Sphecodes nomioidis Pesenko, 1979

Sphecodes nomioidis Pesenko, 1979: 860, ♀ (holotype: ♀, Ukraine: Donetsk Province, Yenaktsevo, 10.VIII.1978, V. Radchenko leg.; ZISP).

Diagnosis

Refer to diagnosis for S. marginatus, above.

Material examined

UNITED ARAB EMIRATES: 4 ♂♂, Hatta, 14.IV.1990, I. Hamer (NHMUK 013380411, 013380415, 013380416, 013380417); OMAN: 1 ♂, Rostaq, 350 m, 21–31.III.1976, K. Guichard (NHMUK 013380443).

Published records

Bogusch and Straka 2012: 14 (Jordan).

Distribution

*United Arab Emirates, *Oman, Jordan; South and Central Europe (west to Austria), Ukraine, Russia (SW of the European part), Turkey.

Sphecodes olivieri Lepeletier de Saint Fargeau, 1825

Sphecodes olivieri Lepeletier de Saint Fargeau in Lepeletier de Saint Fargeau and Audinet-Serville, 1825: 448, ♂ (syntypes: ♂♂, ‘Arabie’).

Sphecodes collaris Spinola, 1843; S. hispanicus var. abyssinicus Sichel, 1865; S. ruficornis Sichel, 1865; S. punctulatus Sichel, 1865; S. subpunctulatus Sichel, 1865; S. rufithorax Morawitz, 1876; S. verticalis Hagens, 1882; S. desertus Nurse, 1903; S. chionospilus Cockerell, 1911; S. chionospilus var. sanguinatus Cockerell, 1911; S. tenuis Meyer, 1920; S. olivieri var. niveatus Meyer, 1925 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 25.

Material examined

UNITED ARAB EMIRATES, 1 ♂, Digdaga, 8.VIII.1984, J.N. Brown [D. Baker det, 92] (NHMUK 013380368); 1 ♂, Hatta (Hotel), 21.VIII.1987, I.L. Hamer [D. Baker det, 92] (NHMUK 013380366); 1 ♂, Soweihan Rd, 12.IV.1988, I.L. Hamer [D. Baker det, 92] (NHMUK 013380367); 1 ♂, Jebal Ali, 15.II.1991, I.L. Hamer [D. Baker det, 92] (NHMUK 013380361); SAUDI ARABIA, 1 ♀, 1♂, Jeddah, 15.II.1972, K.M. Guichard (NHMUK 013380399, 013380398); 1 ♀, 3 ♂♂, Riyadh area, 16–21.IV.1980, K.M. Guichard (NHMUK 013380456, 013380395, 013380396, 013380397); 1 ♂, Jeddah, 13.IV.1980 (NHMUK 013380393); 2 ♀♀, idem, 15.IV.1980, K.M. Guichard (NHMUK 013380392, 013380394); JORDAN: 1 ♀, 20 km W At Tafila, 1.VI.2007, Z. Kejval (OLBL/PCMS); OMAN, 1 ♀, Wadi Qurvat, Ag. Stn. 500 m, 5.III.1976, K. Guichard (NHMUK 013380383); 2 ♀♀, Tinaf, 650 m, 7.III.1976, K. Guichard (NHMUK 013380381, 013380380); 1 ♀, Rostaq, 350 m, 21–31.III.1976, K. Guichard (NHMUK 013380382);

ISRAEL, 1 ♂, Ein Bokek Zohar, 350 m, 25.V.1975, K.M. Guichard (NHMUK 1975-248, 013380385); ISRAEL: 1 ♀, Jericho (Wadi Kelt), 200 m, 6.III.1975, K.M. Guichard (NHMUK 1975-248, 013380387).

Published records

Lepeletier de Saint Fargeau 1825: 448 (‘Arabie’); Warncke 1992: 46, map (Israel); Dathe 2009: 385 (United Arab Emirates); Schwarz 2010: 486 (United Arab Emirates); Ascher and Pickering 2019 (United Arab Emirates, Qatar).

Distribution

United Arab Emirates, *Oman, Qatar, *Saudi Arabia, Israel, *Jordan; North Africa, South Europe, Russia (South of European part), Turkey, Caucasus, Iran, Pakistan, Central Asia, Kazakhstan, NW China.

Sphecodes pellucidus Smith, 1845

Sphecodes pellucidus Smith, 1845: 1014, ♀, ♂ (syntypes: ♀♀, ♂♂, England; NHMUK).

Sphecodes pilifrons Thomson, 1870; S. brevicornis Hagens, 1874; S. volatilis Smith, 1879; S. pellucidus var. algirus Alfken, 1914; S. pellucidus var. hybridus Blüthgen, 1924; S. pellucidus var. niveipennis Meyer, 1925 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 27.

Material examined

SYRIA: 1 ♀, 30 km N Dara, Nawa, 18.V.1996, M. Halada (OLBL/PCMS); JORDAN: 1 ♀, Jordan valley, S. Shuna, 17.IV.1996, M. Halada (OLBL/PCMS); 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS).

Distribution

*Jordan, *Syria; North Africa, Europe (north to 66°), Russia (east to Far East), Turkey, Iran, Central Asia, Kazakhstan, Mongolia, China.

Sphecodes pinguiculus Pérez, 1903

Sphecodes pinguiculus Pérez, 1903: CCXX, ♀ (syntypes: ♀♀, Spain: Catalonia; MNHN).

Sphecodes sareptensis Meyer, 1922; S. excellens Meyer, 1922; S. punctatissimus Meyer, 1922; S. hungaricus Blüthgen, 1923; S. coelebs Blüthgen, 1923; S. consobrinus Blüthgen, 1923; S. persicus Blüthgen, 1924; S. capverdensis Pauly & La Roche, 2002 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 30.

Material examined

SYRIA: 1 ♂, 80 km E Palmira, 450 m, 22.IV.1992, K. Warncke (OÖLM); SAUDI ARABIA, 1 ♀, Hofut, 145 m, 21–6.IV.1980, K. Guichard (NHMUK 013380359); 1 ♀, Hatta, 10.IV.1983, I.L. Hamer (NHMUK 013380389); 1 ♀, idem, 6.VI.1986, I.L. Hamer (NHMUK 013380390); 2 ♀♀, Khor-Fakkan, 20.III.1987, I.L. Hamer (NHMUK 013380364, 013380377); 1 ♀, Soweihan Rd, 12.IV.1988, I.L. Hamer (NHMUK 013380391); OMAN, 1 ♀, Wadi Qurvat, Ag. Stn. 500 m, 5.III.1976, K. Guichard (NHMUK 013380407); 2 ♀♀, Rostaq, 350 m, 21–31.III.1976, K. Guichard (NHMUK 013380405, 013380406);

ISRAEL: 1 ♂, Tel-Aviv, 22.IV.1966, Bytinski-Salz (OLBL/PCMS); 3 ♀♀, Jericho (Hisham Palace), 200 m, 8.III.1975, K.M. Guichard (NHMUK 1975-248, 013380408).

Published records

Schwarz 2010: 486 (United Arab Emirates, Israel); Ascher and Pickering 2019 (Israel, United Arab Emirates).

Distribution

United Arab Emirates, *Oman, *Saudi Arabia, Israel, *Syria; Cape Verde Islands, North Africa, South Europe, Russia (east to Buryatia), Turkey, Iran, Central Asia, Kazakhstan, Mongolia, North China.

Sphecodes puncticeps Thomson, 1870

Sphecodes puncticeps Thomson, 1870: 99, ♀, ♂ (syntypes: ♀♀, ♂♂, Sweden; MZLU).

Sphecodes bituberculatus Pérez, 1903; S. opacifrons Pérez, 1903; S. puncticeps var. cretanus Strand, 1921 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 31.

Material examined

SAUDI ARABIA: 1 ♂, Riyadh area, 16–21.IV.1980, K.M. Guichard (NHMUK 013380403); JORDAN: 1 ♀, Jordan valley, S. Shuna, 17.IV.1996, M. Halada (OLBL/PCMS); 2 ♀, 3 ♂♂, Jordan valley, Dayr Alla, 27.IV.1996, M. Halada (OLBL/PCMS); 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS); 1 ♀, 10 km SE Suwayda Kafr, 19.V.1996, M. Halada (OLBL/PCMS); SYRIA: 1 ♀, Latakia s.l., 17.VI.1986, K. Guichard (NHMUK 013380360); 2 ♀♀, 10 km SE Suwayda Kafr, 19.V.1996, M. Halada (OLBL/PCMS); ISRAEL: 2 ♂♂, Jerusalem, 21.IX.1922, P.A. Buxton (NHMUK 013380401, 013380400); 1 ♂, Rehovot s.l., 29.IV.1975, K.M. Guichard (NHMUK 1975-248, 013380404); 3 ♂♂, Jericho (Wadi Quilt), 250 m, 13–22.V.1975, K.M. Guichard (NHMUK 1975-248, 013380365, 013380376, 013380402).

Published records

Warncke 1992: 19 (Israel); Ascher and Pickering 2019 (Israel).

Distribution

*Saudi Arabia, Israel, *Jordan, *Syria; North Africa, Europe (north to Finland and Sweden), Russia (east to Far East), Turkey, Iran, Central Asia, Kazakhstan, Mongolia.

Sphecodes rubicundus Hagens, 1875

Sphecodes rubicundus Hagens, 1875: 318 (syntypes: ♂♂, ♀♀, Germany; ? Dominican monastery, Venlo, Nederland).

Sphecodes rubicundus altisilesiacus Torka, 1927 (Synonyms).

Diagnosis

The female of this species as well as S. ruficrus is most close to S. pellucidus and S. ephippius owing to a densely punctate head and mesosoma, relative wide pygidial plate and impunctate T1, but differs by having a distinctly elevated vertex with the distance between vertex and upper margin of lateral ocellus at least a lateral ocellar diameter as seen in frontal view (versus 0.2–0.5). S. rubicundus differs from S. ruficrus by white pubescence of head and mesosoma (with brown setae in S. ruficrus) and a less curved basal (M) vein in hind wing. The male most closely resembles S. pesenkoi Astafurova & Proshchalykin, 2018 and S. ruficrus (Erichson, 1835) owing to a similar gonostylar shape (elongate, spoon-shaped). The male of S. rubicundus differs from S. pesenkoi by an areolate mesoscutum (versus punctures separated by 1–3 puncture diameters) and coarsely and densely punctate T1 (a few fine punctures in S. pesenkoi).

According to the phylogenetic analysis (Habermannová et al. 2013) Sphecodes rubicundus, S. ruficrus, S. pellucidus, and S. ephippius belong to the same clade. Relationship between these species also is well supported by morphological characters.

Material examined

ISRAEL, 2 ♂♂, Jerusalem, 800 m, 20.III.1975, K.M. Guichard (NHMUK 1975-154, 013380388, 013380386); 1 ♀, Tiberias, 200 m, 22.III.1975, K.M. Guichard (NHMUK 1975-154, 013380384); 1 ♀, Jerusalem, 20.III.1993, D. Ahal (OLBL/PCMS).

Distribution

*Israel; Europe (north to 56°), Russia (south of the European part), Turkey, Caucasus, Iran.

Sphecodes rubripes Spinola, 1839

Figures 6, 21

Sphecodes rubripes Spinola, 1839: 512, ♀ (syntypes: ♀♀, Cyprus; MRSN).

Sphecodes africanus Lepeletier, 1841; S. rufipennis Cockerell, 1931, S. atrescens Cockerell, 1931 (Synonyms).

Diagnosis

The female of S. rubripes differs from S. albilabris by the pubescence of paraocular area (Fig. 6) with brown erect setae not obscuring integument (versus white plumose appressed pubescence obscuring integument usually with admixture of brownish erect setae in S. albilabris, Fig. 7). Both sexes also differ by mainly red legs, except brown coxae and trochanters, Fig. 21 (at most reddish tarsi and tibia in S. albilabris, Fig. 20). These two species also differ in phenology (males of S. rubripes were recorded in the early spring while males of S. albilabris were found in the summer) and have different hosts (Bogusch and Straka 2012, Cross 2017). S. albilabris is widespread in the Palaearctic from the Atlantic Ocean to Russian Far East; however, the distribution of the species in the Mediterranean Region is unclear due to confusion with S. rubripes. The past records of S. albilabris from Israel and Syria refer to S. rubripes. We examined material of S. albilabris from Morocco and Tunisia, but we do not have any specimens of S. albilabris from the Arabian Peninsula or surrounding lands.

Material examined

JORDAN: 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS).

Published records

Meyer 1924: 3 (Syria, as S. fuscipennis rubripes); Warncke 1992: 31 (Israel, as S. albilabris rubripes); Ascher and Pickering 2019 (Israel, as S. albilabris (Fabricius)).

Distribution

*Jordan, Israel, Syria; North Africa, South-Western Europe, Cyprus.

Remarks

Mayer (1924) and later Warncke (1992) interpreted Sphecodes rubripes as a subspecies of S. albilabris (Fabricius, 1793), but this taxon was restored as a valid species (Bogusch and Straka 2014b).

Sphecodes ruficrus (Erichson, 1835)

Dichroa ruficrus Erichson, 1835: 101, ♀, (syntypes: ♀♀, Spain; ZMHB).

Sphecodes hispanicus Wesmael, 1836; S. rufipes Smith, 1853; S. gibbus var. tunetanus Gribodo, 1894; S. atrohirtus Pérez, 1903 (Synonyms).

Diagnosis

Refer to diagnosis for S. rubicundus, above.

Material examined

JORDAN: 1 ♀, W Jordan Valley, env. of S. Shuna, 17.IV.1996, M. Halada (OLBL/PCMS).

Published records

Warncke 1992: 21 (Israel); Ascher and Pickering 2019 (Israel).

Distribution

Israel, *Jordan; North Africa, southwestern Europe.

Remarks

Russia is mistakenly listed as within the distribution by Bogusch and Straka (2012) as well as Turkey and Armenia by Özbek et al. (2015) due to confusion with S. ruficrus rubicundus sensu Warncke (1992).

Sphecodes rufiventris (Panzer, 1798)

Tiphia rufiventris Panzer, 1798: 4, ♀ (syntypes: ♀♀, Germany; ZMHB).

Sphecodes subovalis Schenck, 1853; S. brevis Hagens, 1875; S. singularis Meyer, 1920; S. combinatus Blüthgen, 1927; S. subovalis austrinus Erlandsson, 1979; S. rufiventris hethiticus Warncke, 1992 (Synonyms).

Diagnosis

See Astafurova et al. 2018a: 34.

Material examined

JORDAN: 1 ♂, W Jordan Valey, Mubalath, 27.IV.1996, M. Halada (OLBL/PCMS).

Published records

Ascher and Pickering 2019 (Israel).

Distribution

Israel, *Jordan; North Africa, Europe, (north to 57°), Russia (east to Khakassia Republic), Turkey, Iran, Central Asia, Kazakhstan.

Sphecodes schenckii Hagens, 1882

Sphecodes schenckii Hagens, 1882: 217, ♂ (holotype: ♂, no locality, Rudow leg. [see Blüthgen 1923: 444]; MNHB).

Sphecodes sulcicollis Pérez, 1903; S. caspicus Meyer, 1920 (Synonyms).

Diagnosis

See Astafurova and Proshchalykin 2017b: 274.

Material examined

JORDAN: 1 ♂, NW Ajlun, 850 m, 20.V.2007, Z. Kejval (OLBL/PCMS) ; 1 ♀, 10 km N Petra, 3.V.1996, M. Halada (OLBL/PCMS); SYRIA: 3 ♂♂, 20 km NE Latakia, 25.V.1996, M. Halada (OLBL/PCMS).

Published records

Warncke 1992: 27 (Israel, as Sphecodes schenckii caspicus Meyer); Ascher and Pickering 2019 (Israel).

Distribution

Israel, *Jordan, *Syria; Europe (north to Germany), Russia (European part), Turkey, Caucasus, ? Iran.

Sphecodes tadschicus Blüthgen, 1935

Sphecodes tadschicus Blüthgen in Popov, 1935: 366, ♂, ♀ (holotype: ♂, near Kulab [Tajikistan], 25.VII.1935, V. Popov leg.; ZISP).

Diagnosis

See Astafurova et al. 2018a: 39.

Material examined

ISRAEL: 1 ♀, 8 ♂♂, Jerusalem, 10–25.VIII.1960, Bytinski (MNHB).

Distribution

*Israel; Turkey, Iran, Central Asia, Kazakhstan.

Sphecodes turanicus Astafurova & Proshchalykin, 2017

Sphecodes turanicus Astafurova & Proshchalykin, 2017b: 274, ♂, ♀ (holotype: ♀, Turkmenistan, Chardzhou, 16.IV.1988, Dialentov leg.; ZISP).

Diagnosis

See Astafurova et al. 2018a: 41.

Material examined

SAUDI ARABIA: 1 ♂, Riyadh, El Ha’ir, 16–21.IV.1980, K.M. Guichard (NHMUK 013380449).

Distribution

*Saudi Arabia; Central Asia, Kazakhstan, China (Gansu).

Sphecodes villosulus Schwarz, 2010

Sphecodes villosulus Schwarz, 2010: 486–491, ♀, ♂ (holotype: ♀, United Arab Emirates, Dubai, Nakhalai, 28–30.IV.1984, in Malaise trap, E. Sugden leg.; BLCU).

Diagnosis

This species differs from other small Palaearctic species with 5–6 hamuli in the hind wing by having a unique combination of simple mandibles and the male gonocoxite dorsally with an impression. The female is closest to S. armeniacus owing to dense appressed snow-white pubescence obscuring the integument on face, a transverse head and sparsely punctate mesoscutum, but differs from this species by sparser and finer punctate ocello-ocular area (3–5 μm / 2–3 versus 5–10 μm / 1–2) and strongly transverse F3 (almost square in S. armeniacus). The male of S. villosulus recalls S. miniatus in the rectangular gonostylar shape, but clearly differs from this species by the less developed tyloids on the flagellomeres extending to approximately a half of ventral flagellar surfaces (versus those across 4/5).

Material examined

UNITED ARAB EMIRATES: 1 ♀, 1♂, Abu Dhabi, 30.I.1987, I. Hamer (NHMUK 013380419, 013380423); 1 ♀, 2♂♂, Abu Dhabi, 31.III.1987 (NHMUK 013380418, 013380421, 013380420); 1 ♀, idem, 10.IV.1987, I. Hamer (NHMUK 013380422); 1 ♀, Hatta, 20.XII.1990 (NHMUK 013380424); 2 ♂♂, idem, 23.VIII.1991 (NHMUK 013380426, 013380427); 1 ♀, idem, 5.III.1993, I. Hamer (NHMUK 013380425); 3 ♀♀, North Ras, Al Khaimah, 17.II.2018 (M. Mokrousov) (ZISP); SAUDI ARABIA: 1 ♀, Riyadh, El Ha’ir, 19.III.1980, K.M. Guichard (NHMUK 013380448); OMAN: 1 ♀, Rostaq, 350 m, 21–31.III.1976, K.M.Guichard (NHMUK 013380444).

Published records

Schwarz 2010: 486 (United Arab Emirates); Ascher and Pickering 2019 (United Arab Emirates).

Distribution

United Arab Emirates, *Oman, *Saudi Arabia.

Discussion

In total, 26 species of Sphecodes are recorded from the Arabian Peninsula and surrounding lands (Israel, Jordan and Syria) (Table 1). This is a comparable number to the Iranian fauna, but distinctly less in comparison with the adjacent fauna of Turkey, North Africa and Central Asia (Table 3).

Table 3.

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List of Sphecodes species recorded in Arabian Peninsula and surrounding lands (AP+SL), Turkey, Iran, North Africa (Morocco, Algeria, Libya, Tunisia, Egypt) and Central Asia (Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, Tajikistan).

	Sphecodes species	AP+SL	Turkey	Iran	North Africa	Central Asia
1	S. albilabris (Fabricius, 1793)	–	+	+	+	+
2	S. alternatus Smith, 1853	+	+	+	+	+
3	S. anatolicus Warncke, 1992	–	+	+	–	+
4	S. armeniacus Warncke, 1992	–	+	–	+	+
5	S. atlanticus Warncke, 1992	+	–	–	+	–
6	S. atlassa Warncke, 1992	–	–	–	+	–
7	S. barbatus Blüthgen, 1923	+	+	–	–	–
8	S. crassanus Warncke, 1992	–	+	–	–	+
9	S. crassus Thomson, 1870	–	+	+	+	+
10	S. cristatus Hagens, 1882	–	+	–	–	+
11	S. croaticus Meyer, 1922	–	+	+	+	+
12	S. dathei Schwarz, 2010	+	–	–	–	–
13	S. dusmeti Blüthgen, 1924	+	+	–	+	–
14	S. ebmeri Astafurova & Proshchalykin, 2018	–	–	+	–	–
15	S. ephippius (Linné, 1767)	+	+	+	–	+
16	S. ferruginatus Hagens, 1882	–	+	–	–	+
17	S. geoffrellus (Kirby, 1802)	–	+	–	+	+
19	S. gibbus (Linnaeus, 1758)	+	+	+	+	+
20	S. hakkariensis Warncke, 1992	–	+	–	–	+
21	S. haladai Warncke, 1992	–	–	+	+	+
22	S. hyalinatus Hagens, 1882	–	–	–	–	+
23	S. hirtellus Blüthgen, 1923	–	–	–	+	–
24	S. intermedius Blüthgen, 1923	+	+	–	+	+
25	S. longulus Hagens, 1882	+	+	+	–	+
27	S. longuloides Blüthgen, 1923	+	–	–	+	–
28	S. majalis Pérez, 1903	+	+	+	+	–
29	S. marginatus Hagens, 1882	+	–	–	+	–
30	S. monilicornis (Kirby, 1802)	+	+	+	+	+
31	S. niger Hagens, 1874	–	+	–	–	–
32	S. nomioidis Pesenko, 1979	+	+	–	–	–
33	S. nurekensis Warncke, 1992	–	–	–	–	+
34	S. olivieri Lepeletier de Saint Fargeau, 1825	+	+	+	+	+
35	S. pectoralis Morawitz, 1876	–	–	+	–	+
36	S. pellucidus Smith, 1845	+	+	+	+	+
37	S. pesenkoi Astafurova & Proshchalykin, 2018	–	–	–	–	+
38	S. pinguiculus Pérez, 1903	+	+	+	+	+
39	S. pseudofasciatus Blüthgen, 1925	–	+	–	+	–
40	S. puncticeps Thomson, 1870	+	+	+	+	+
41	S. reticulatus Thomson, 1870	–	+	+	–	+
42	S. rubicundus Hagens, 1875	+	+	+	–	–
43	S. rubripes Spinola, 1839	+	–	–	+	–
44	S. ruficrus (Erichson, 1835)	+	–	–	+	–
45	S. rufiventris (Panzer, 1798)	+	+	+	+	+
46	S. sandykachis Astafurova & Proshchalykin, 2018	–	–	–	–	+
47	S. saxicolus Warncke, 1992	–	–	+	–	+
48	S. scabricollis Wesmael, 1835	–	+	+	–	+
49	S. schenckii Hagens, 1882	+	+	+	–	–
50	S. schwarzi Astafurova & Proshchalykin, 2015	–	–	–	–	+
51	S. spinulosus Hagens, 1875	–	+	+	+	+
52	S. tadschicus Blüthgen, 1935	+	+	+	–	+
53	S. trjapitzini Astafurova & Proshchalykin, 2018	–	–	–	–	+
54	S. turanicus Astafurova & Proshchalykin, 2017	+	–	–	–	+
55	S. zangherii Noskiewicz, 1931	–	+	–	–	–
56	S. villosulus Schwarz, 2010	+	–	–	–	–
Total:		26	34	25	26	35

The Sphecodes fauna of the Arabian Peninsula and surrounding lands is a complex of Mediterranean, Sahara-Gobian, endemic, and species widespread in the Palaearctic region. Eight species, namely S. alternatus, S. ephippius, S. gibbus, S. longulus, S. monilicornis, S. marginatus, S. pellucidus, and S. puncticeps are widespread from north to south of the Palaearctic region and occur in biomes ranging from forest to desert. However, two of these (S. marginatus and S. puncticeps) are recorded from the Arabian Peninsula and the remainder all are found only in Mediterranean areas.

Sphecodes majalis, S. schenckii Hagens, S. rubicundus, and S. nomioidis are steppe species, distributed in Europe, Turkey and the Caucasus to Iran. Of them, only S. nomioidis is recorded from the Arabian Peninsula.

Sphecodes olivieri, S. intermedius, S. rufiventris, and S. pinguiculus are widespread from steppe to desert in the Western Palaearctic. Of these only S. rufiventris is not recorded from the Arabian Peninsula.

Sphecodes barbatus, S. rubripes, and S. ruficrus are possibly purely Mediterranean species not reaching the Arabian Peninsula. In contrast, S. atlanticus turns out to be Sahara-Arabian. Sphecodes dusmeti and S. longuloides are Mediterranean-Arabian species.

Sphecodes tadschicus and S. turanicus are Irano-Turanian species reaching the Arabian Peninsula.

Finally, two species, S. dathei and S. villosulus are endemic to the Arabian Peninsula.

Although the Arabian fauna of the genus is not fully studied it is now clear that the Arabian fauna differs from that of the Mediterranean; of 26 recorded species only six (S. olivieri, S. intermedius, S. marginatus, S. nomioidis, S. pinguiculus, and S. puncticeps) are common to both and these are all widespread in the Western Palaearctic.

Acknowledgments

We are grateful to Fritz Gusenleitner (OLBL) for help during our visit to Austria, and Andrew Grace (Hastings, UK) for checking the English. We thank Hege Vårdal (NHRS), Lukasz Przybylowicz (ISZP), Michael Ohl, and Viola Richter (ZMHB) for providing Sphecodes type specimens. We also thank Michael Engel, Petr Bogusch, and anonymous reviewer for their comments and suggestions to streamline and improve the earlier drafts of the manuscript.

This investigation was supported by the Russian Funds for Basic Research (grant numbers 17-04-00259 and 19-04-00027) and the state research project (№ АААА-А19-119020690101-6).

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