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.

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.

Introduction

The present paper is part of a series of studies dealing with the bees of the genus 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 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 (Dathe 2009, Engel et al. 2013). A first contemporary inventory of the 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 known from this area, but the fauna of Arabian Peninsula is particularly under-recorded.

Probably the first information on the genus 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 from ‘Arabie’. Almost two centuries later, in his monograph on the Western Palaearctic , 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 species: and Blüthgen. In the recently published third volume of the “Arthropod fauna of UAE”, Schwarz (2010) described and and recorded Hagens and Pérez from the United Arab Emirates. In total, nineteen species have been recorded from the Arabian Peninsula and its adjacent lands so far (Table 1). The genus 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.

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

Species		Arabian Peninsula					surrounding lands
		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
		12					20

White circle – published records (Meyer 1924; Warncke 1992; Dathe 2009; Schwarz 2010; Ascher and Pickering 2019); black circle – current data. Genus are not known in Kuwait, Bahrain, and Iraq.

Based on a comprehensive study of specimens in various collections, we here list 23 species of the genus , 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 Warncke, 1992 and the male of Schwarz, 2010 are here described for the first time and a lectotype is designated for 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:

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

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

Muséum National d’Histoire Naturelle Paris, France;

Museo Regionale di Scienze Naturali, Torino, Italy;

Lund University, Lund, Sweden;

Natural History Museum, London, UK;

University of Copenhagen, Zoological Museum, Copenhagen, Denmark;

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 ( and ). 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

Smith, 1853

C46A4EBF76A657D3964CC99B537BC064

Smith, 1853: 36, ♀ (syntypes: ♀♀, Albania;

Hagens, 1882;

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.

Warncke, 1992

F3062642680F57EA9FB353FA0E5078C1

Figures 5 , 9 , 10 , 19 , 22

Figures 2–7.

Head, females, frontal view. 2 Pérez 3 Wesmael 4 Blüthgen 5 Warncke 6 Spinola 7 (Fabricius). Scale bars: 1.0 mm.

Figures 8–11.

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

Figures 16–19.

T1 (16, 17), metasoma (18, 19), females, dorsal view. 16 Pérez 17 Blüthgen 18 Wesmael 19 Warncke. Scale bars: 1.0 mm.

Figures 22–25.

Genitalia, males, dorsal view. 22 Warncke 23 Wesmael 24 Schwarz 25 Thomson. Scale bars: 0.25 mm.

Warncke, 1992: 25, Abb. 17, ♂ (holotype: ♂, Algeria: Hoggar-Geb., Guelta;

This species is similar to the Trans-Palaearctic 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, differs from by a number of characters outlined in Table 2. In addition to presence of preoccipital lateral carina, clearly differs from the species group ( Warncke, 1922, (Linnaeus, 1758), Meyer, 1922, (Panzer, 1798), Hagens, 1882, 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.

Differences between Warncke, 1992 and 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.

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.

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.

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, , 27.X.2005, J. Halada (OLBL/PCMS); 2 ♀♀, 3 ♂♂, Hawf NE Albhaydah, 200–730 m, , 14.X.2005, J. Halada (OLBL/PCMS); 2 ♀♀, 4 ♂♂, 20 km S Taizz, 1200 m, , 24.X.2005, J. Halada (OLBL/PCMS); 2 ♂♂, Jabal Bura, NEE Al Hudaydah, 200–800 m, , 30.X-1.XI.2005, J. Halada (OLBL/PCMS); 1 ♂, Wadi Aniz, SSW Sana, 1520 m, , 7.X.2005, J. Halada (OLBL/PCMS).

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

Blüthgen, 1923

517471C21052533F97CAE88F4771CD29

Figures 4 , 17

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

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

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).

*Syria; Greece, Turkey.

Remarks.

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

Schwarz, 2010

6D4AD2DBEB885FA39206B98C83009AB9

Figures 12–15 , 24

Figures 12–15.

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

Schwarz, 2010: 483–486, ♀, plates 1–12 (holotype: ♀, United Arab Emirates, Wadi Shawkah,

The species is similar to 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 differs from by dense, apressed, snow-white, plumose pubescence obscuring integument in paraocular areas (sparse, simple pubescence not obscuring integument in ); the male differs by densely and relatively coarsely punctate T1 (in T1 usually with a few fine punctures, rarely with relatively coarse and dense punctures). Both species have similar gonostylar shape, but has a narrower, trapezoidal membranous portion of the gonostylus (wider, close to oval in , Fig. 25).

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).

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, , 28.X.2005, J. Halada (OLBL/PCMS).

Schwarz 2010: 483 (United Arab Emirates).

United Arab Emirates, *Saudi Arabia, *Yemen.

(Linnaeus, 1758)

5F913968CCE556978526570A1C4BB74D

Linnaeus, 1758: 571, ♀ (syntypes: ♀♀, Sweden;

Füessly, 1775;

See Astafurova et al. 2018a: 17.

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).

Warncke 1992: 30 (Israel).

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

Blüthgen, 1923

7CC2FBA91AFE532A829B9A3365B2C600

Figure 11

Blüthgen, 1923: 500 (lectotype (

Meyer, 1925 (Synonym).

See Astafurova et al. 2018a: 20.

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 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)

Ascher and Pickering 2019 (Israel)

*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).

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 to avoid any confusion about the status of the type specimens and to properly diagnose this species.

Hagens, 1882

43EBA5F314925AF7A2E67FBE8AB2EAED

Hagens, 1882: 226, Fig.

Hagens, 1882;

See Astafurova et al. 2018a: 21.

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).

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

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

Pérez, 1903

8A5ED1678AF25BA98265B2151EB7EBBC

Figures 2 , 16

Pérez, 1903: 219, ♀, ♂, (syntypes: ♀, ♂, France, Spain;

Pérez, 1903;

Refer to the diagnosis , above.

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).

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

Hagens, 1882

DCA18E56EF965AF78B75623ACCAAF279

Hagens, 1882: 223, Fig.

Hagens, 1882;

This species belongs to the miniatus species group ( Warncke, 1992, Warncke, 1992, Warncke, 1992, Hagens, 1882, Hagens, 1882, Pesenko, 1979, Astafurova & Proshchalykin, 2014, and Astafurova & Proshchalykin, 2018), with the same length and transverse F1–F3 in females. Among species of this group is most close to and 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).

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).

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

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

(Kirby, 1802)

4AC06390842B516CAC0E5EECE61A4859

Kirby, 1802: 47, ♂ (syntypes: ♂♂, England,

Lepeletier de Saint Fargeau, 1841;

See Astafurova et al. 2018a: 24.

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).

Ascher and Pickering 2019 (Jordan).

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

Pesenko, 1979

5BE05D899B75554B805681C443A35A68

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

Refer to diagnosis for , above.

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).

Bogusch and Straka 2012: 14 (Jordan).

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

Lepeletier de Saint Fargeau, 1825

8379E7A2BCD151EE998204A763B18992

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

Spinola, 1843;

See Astafurova et al. 2018a: 25.

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).

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).

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.

Smith, 1845

80F345845F915F009BB3FD1A6E8B1BBD

Smith, 1845: 1014, ♀, ♂ (syntypes: ♀♀, ♂♂, England;

Thomson, 1870;

See Astafurova et al. 2018a: 27.

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).

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

Pérez, 1903

1BA02AED9E255C609B3F70042BC8AE7F

Pérez, 1903: CCXX, ♀ (syntypes: ♀♀, Spain: Catalonia;

Meyer, 1922;

See Astafurova et al. 2018a: 30.

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).

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

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.

Thomson, 1870

E61D565668005FD398FBE21910268C45

Thomson, 1870: 99, ♀, ♂ (syntypes: ♀♀, ♂♂, Sweden;

Pérez, 1903;

See Astafurova et al. 2018a: 31.

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).

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

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

Hagens, 1875

68AD66DC2D4A583CBEFB7835049C7E1E

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

Torka, 1927 (Synonyms).

The female of this species as well as is most close to and 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). differs from by white pubescence of head and mesosoma (with brown setae in ) and a less curved basal (M) vein in hind wing. The male most closely resembles Astafurova & Proshchalykin, 2018 and (Erichson, 1835) owing to a similar gonostylar shape (elongate, spoon-shaped). The male of differs from by an areolate mesoscutum (versus punctures separated by 1–3 puncture diameters) and coarsely and densely punctate T1 (a few fine punctures in ).

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

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).

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

Spinola, 1839

AED78E1E647A59999CE12B44195BAF69

Figures 6 , 21

Figures 20, 21.

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

Spinola, 1839: 512, ♀ (syntypes: ♀♀, Cyprus;

Lepeletier, 1841;

The female of differs from 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 , Fig. 7). Both sexes also differ by mainly red legs, except brown coxae and trochanters, Fig. 21 (at most reddish tarsi and tibia in , Fig. 20). These two species also differ in phenology (males of were recorded in the early spring while males of were found in the summer) and have different hosts (Bogusch and Straka 2012, Cross 2017). 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 The past records of from Israel and Syria refer to We examined material of from Morocco and Tunisia, but we do not have any specimens of from the Arabian Peninsula or surrounding lands.

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

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

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

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

(Erichson, 1835)

4DD92252598B52318ED7B28E7E56E63D

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

Wesmael, 1836;

Refer to diagnosis for , above.

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

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

Israel, *Jordan; North Africa, southwestern Europe.

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 sensu Warncke (1992).

(Panzer, 1798)

EA4EA9F64BA55BF3A1FC3AF064E687E7

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

Schenck, 1853;

See Astafurova et al. 2018a: 34.

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

Ascher and Pickering 2019 (Israel).

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

Hagens, 1882

30EA22E5954652D3BF5B8627DB0398EC

Hagens, 1882: 217, ♂ (holotype: ♂, no locality, Rudow leg. [see

Pérez, 1903;

See Astafurova and Proshchalykin 2017b: 274.

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).

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

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

Blüthgen, 1935

C6671BF631855F19A9D51ED3E1E89547

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

See Astafurova et al. 2018a: 39.

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

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

Astafurova & Proshchalykin, 2017

7D2BE7510D0857F28E38D5D19D9C4E52

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

See Astafurova et al. 2018a: 41.

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

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

Schwarz, 2010

D12FCFAA24895241B1E7149122E79AA7

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

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 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 ). The male of recalls 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).

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).

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

United Arab Emirates, *Oman, *Saudi Arabia.

Discussion

In total, 26 species of 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.

List of 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 distribution of species are given according to Özbek et al. 2015 (Turkey), Astafurova et al. 2018d (Iran), Warncke 1992 (North Africa), Astafurova and Proshchalykin 2017b and Astafurova et al. 2018a, c (Central Asia).

The 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 , , , , , , , and are widespread from north to south of the Palaearctic region and occur in biomes ranging from forest to desert. However, two of these ( and ) are recorded from the Arabian Peninsula and the remainder all are found only in Mediterranean areas.

, Hagens, , and are steppe species, distributed in Europe, Turkey and the Caucasus to Iran. Of them, only is recorded from the Arabian Peninsula.

, , , and are widespread from steppe to desert in the Western Palaearctic. Of these only is not recorded from the Arabian Peninsula.

, , and are possibly purely Mediterranean species not reaching the Arabian Peninsula. In contrast, turns out to be Sahara-Arabian. and are Mediterranean-Arabian species.

and are Irano-Turanian species reaching the Arabian Peninsula.

Finally, two species, and 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 (, , , , , and ) are common to both and these are all widespread in the Western Palaearctic.