Regiscolia maculata ( Drury, 1773 ), 2025

Regiscolia maculata ( Drury, 1773) , comb. nov.

( Figs 79–85 View FIGURE 79 View FIGURE 80 View FIGURE 81 View FIGURE 82 View FIGURE 83 View FIGURE 84 View FIGURE 85 )

Sphex maculata Drury, 1773: 77 ( holotype, ♀, probably deposited at Natural History Museum , United Kingdom, not examined;

type locality “ Morea ” [= Greece, Peloponnese]). Sphex bidens Pallas, 1773: 729 ; Müller 1776: 323; Sulzer 1776: 191, tab. XXVII, fig. 3; Christ 1791: 255 (synonymised by

Saussure & Sichel 1864: 50). Sphex radula Sulzer, 1776: 192 , tab. XXVII, fig. 4 (synonymised by Saussure & Sichel 1864: 50). Scolia flavifrons Fabricius, 1775: 355 ; Fabricius 1781: 452; Fabricius 1787: 280; Cyrillo 1787: 2; Gmelin 1790: 2736; Rossi

1790: 69; Rossi 1792: 129; Fabricius 1793: 229; Fabricius 1804: 240; Klug 1805: 22; Lepeletier 1825: 393; Spinola 1839:

334; Passerini 1840: 6, 9, 12; Passerini 1841: 6, 7; Palma 1869: 34; Costa 1887: 96; Ashmead 1903: 7; Cros 1929: 242;

Bajari 1953: 124; Erlandsson 1974: 24; Vuts et al. 2012: 351–359. Scolia haemorrhoidalis Fabricius, 1787: 280 ; Gmelin 1790: 2736; Fabricius 1793: 230; Fabricius 1804: 240; Klug 1805: 24;

Germar 1817: 262; Vander Linden 1827: 288; Lepeletier 1845: 522; Eversmann 1849: 430; Burmeister 1854: 18; Mocsary

1881: 54; Costa 1887: 97; Iuga 1953: 259 (synonymised by Micha 1927: 125, and later Hamon & Osten 1994: 15). Sphex interrupta Scopoli, 1786: 60 (synonymised by Saussure & Sichel 1864: 50). Scolia hortorum Fabricius, 1787 ; Cyrillo 1787: 2; Gmelin 1790: 2736; Rossi 1790: 70; Rossi 1792: 129; Fabricius 1793: 232;

Fabricius 1804: 243; Klug 1805: 22; Vander Linden 1827: 286; Lepeletier 1845: 520; Lucas 1849: 278; Burmeister 1854:

18; Costa 1858: 6; Costa 1874: 575 (misidentification). Sphex flavifrons : Villers 1789: 239; Christ 1791: 267. Sphex hortorum : Villers 1789: 240. Vespa tricuspidata Villers, 1789: 280 (synonymised by Saussure & Sichel 1864: 50). Sphex goliath Christ, 1791: 253 (synonymised by Saussure & Sichel 1864: 51). Sphex versicolor Christ, 1791: 254 (synonymised by Saussure & Sichel 1864: 51). Sphex radula siberica Christ, 1791: 258 (synonymised by Saussure & Sichel 1864: 51). Sphex radula hungarica Christ, 1791: 258 (synonymised by Saussure & Sichel 1864: 51). Scolia quadrimaculata Fabricius, 1793: 229 ; Fabricius 1804: 240 (synonymised by Betrem & Bradley 1964: 20–21). Scolia ( Scolia) flavifrons : Guérin-Méneville 1830: 247. Scolia ( Scolia) insignis Saussure, 1858: 197 (synonymised by Bingham 1897: 77, later by Osten 2005b: 37). Scolia ( Triscolia) insignis : Saussure & Sichel 1864: 47; Cameron 1892: 101. Scolia ( Triscolia) flavifrons : Saussure & Sichel 1864: 49; Vogrin 1915: 37–38, 40; Berland 1925: 297; Betrem 1927a: xcvii;

Dusmet 1930: 16–19; Schmiedeknecht 1930: 511; Betrem 1933: 260; Betrem 1947: 414; Mingo & Compte 1963: 75. Scolia ( Triscolia) haemorrhoidalis : Saussure & Sichel 1864: 50; Cameron 1892: 102; Bingham 1897: 71–72, 77; Vogrin 1915:

36, 38, 41. Triscolia haemorrhoidalis : Gribodo 1893: 150. Scolia ( Scolia , Triscolia ) flavifrons : Dalla-Torre 1897: 158–160. Scolia ( Scolia , Triscolia ) flavifrons var. haemorrhoidalis : Dalla-Torre 1897: 160. Triscolia flavifrons : Bartlett 1912: 296. Scolia ( Triscolia) flavifrons var. haemorrhoidalis : Berland 1925: 297; Maidl 1922: 41; Dusmet 1930: 19; Schmiedeknecht 1930:

511. Scolia ( Ascoli) haemorrhoidalis : Betrem 1926: xiii. Triscolia maculata : Micha 1927: 125. Triscolia maculata maculata : Micha 1927: 126–131. Triscolia maculata bischoffi Micha, 1927: 131 . Triscolia maculata albifrons : Micha 1927: 131–132 ( lapsus calami for flavifrons in Bradley 1972: 4). Triscolia maculata barbara Micha, 1927: 132 . Syn. nov. Triscolia maculata form verticalis Micha, 1927: 132. Syn. nov. Triscolia maculata sicula Micha, 1927: 133 . Syn. nov. Triscolia maculata form excepta Micha, 1927: 133 (synonymised by Bradley 1972: 9). Scolia ( Triscolia) flavifrons haemorrhoidalis : Betrem 1927b: 291; Betrem 1933: 261; Betrem 1941: 118; Betrem 1947: 414;

Bradley 1956: 579. Scolia flavifrons haemorrhoidalis : Bradley 1928: 93. Scolia ( Triscolia) maculata : Guiglia 1928: 428; Steinberg 1962: 116–118; Friese & Muche 1967: 514. Scolia ( Triscolia) maculata flavifrons : Guiglia 1928: 429; Nadig & Nadig 1934: 36; Nadig & Nadig 1935: 4; Guiglia 1944:

146–147; Steinberg 1962: 118; Carrière 1985: 20. Scolia ( Triscolia) maculata flavifrons var. funerea Guiglia, 1928: 434 . Syn. nov. Scolia ( Triscolia) maculata flavifrons var. sicula : Guiglia 1928: 435. Scolia ( Triscolia) flavifrons var. nigra Dusmet, 1930: 19–20 . Syn. nov. Scolia ( Triscolia) flavifrons flavifrons : Betrem 1933: 260; Betrem 1941: 117; Betrem 1947: 414; Bradley 1956: 579. Scolia ( Triscolia) flavifrons flavifrons var. flavifrons : Betrem 1933: 261. Scolia ( Triscolia) flavifrons flavifrons var. verticalis : Betrem 1933: 261. Scolia ( Triscolia) flavifrons flavifrons var. sicula : Betrem 1933: 261. Triscolia maculata : Guiglia 1942: 50. Triscolia maculata flavifrons : Guiglia 1938: 5; Guiglia 1949: 32. Scolia ( Triscolia) maculata flavifrons forme barbara: Guiglia 1951: 286. Scolia ( Triscolia) maculata bischoffi : Hammer 1951: 7. Scolia flavifrons haemorrhoidalis : Atanassov 1951: 287; Erlandsson 1974: 24. Scolia flavifrons var. haemorrhoidalis : Bajari 1953: 124; Vuts et al. 2012: 352. Scolia flavifrons haemorrhoidalis : Guiglia & Bradley 1958: 93–95, 97. Scolia ( Triscolia) maculata maculata : Steinberg 1962: 118. Scolia ( Triscolia) maculata barbara : Steinberg 1962: 118. Megascolia ( Regiscolia) flavifrons flavifrons : Betrem & Bradley 1964 a: 443; Bradley & Betrem 1964: 13, 21; Nagy 1967: 222;

Bradley 1972: 9. Megascolia ( Regiscolia) flavifrons haemorrhoidalis : Betrem & Bradley 1964: 443; Bradley & Betrem 1964: 14; Nagy 1967:

223; Bradley & Betrem 1964: 14; Tkalcû 1987: 290. Megascolia ( Regiscolia) flavifrons siberica : Nagy 1967: 222. Megascolia ( Regiscolia) flavifrons form vespertina Nagy, 1967: 223. Syn. nov. Megascolia ( Regiscolia) flavifrons form vernalis Nagy, 1967: 223. Syn. nov. Megascolia ( Regiscolia) flavifrons : Bradley 1972: 4. Megascolia ( Regiscolia) flavifrons flavifrons form barbara: Bradley 1972: 5. Megascolia ( Regiscolia) flavifrons bischoffi : Bradley 1972: 6. Megascolia ( Regiscolia) flavifrons flavifrons form sicula: Bradley 1972: 15. Megascolia ( Regiscolia) flavifrons flavifrons form verticalis: Bradley 1972: 17. Megascolia ( Regiscolia) flavifrons haemorrhoidalis form insignis : Bradley 1974: 446. Scolia ( Triscolia) maculata flavifrons haemorrhoidalis : Carrière 1985: 20. Megascolia flavifrons : Piek 1986b: 45–47; Piek 1988: 37; Piek & Carrière 1990: 21–23; Carrière 1990: 16; Strumia 2019: 72,

78, 80; Golfetti et al. 2025: 3, 10. Megascolia flavifrons flavifrons : Piek 1986b: 45–47; Pagliano 1987: 164; Piek & Carrière 1990: 21. Megascolia flavifrons haemorrhoidalis : Piek 1986b: 45–47; Pagliano 1987: 165; Piek 1988: 37; Piek & Carrière 1990: 21; Diniz

1991: 31; Madl 1997: 827. Megascolia flavifrons flavifrons haemorrhoidalis : Piek & Carrière 1990: 21–23. Megascolia ( Regiscolia) flavifrons haemorrhoidalis var. bischoffi : Osten 1990: 470. Megascolia ( Regiscolia) maculata maculata : Hamon & Osten 1994: 15; Hamon et al. 1995: 34; Basset 1998: 12; Osten 1999a:

424; Osten & Özbek 1999: 436; Osten 2000: 546, 551, 557; Osten 2002: 344; Osten et al. 2003: 370; Osten & Arens 2004:

312; Osten 2005a: 1455, 1461; Osten 2005b: 37; Carrière 2006: 427; Ljubomirov 2006: 532; Fallahzadeh & Saghaei 2010:

793; Makhan 2012: 1–2; Samin & Baðrýaçýk 2012: 391; Samin et al. 2014: 716; Elcin & Baðrýaçýk 2015: 50. Megascolia ( Regiscolia) maculata flavifrons : Hamon & Osten 1994: 15; Hamon et al. 1995: 36; Osten 1999a: 424; Osten 2000:

546, 551, 557; Osten 2005b: 37; Carrière 2006: 427; Carrière 2007: 247; Lohrmann & Engel 2015: 192–194; Turrisi et al.

2020: 728. Ascolia flavifrons : Argaman 1996: 188. Laskariska haemorrhoidalis : Argaman 1996: 188.

Megascolia maculata : Schmid-Egger & Burger 1998: 46; Osten 1999b: 449; Özbek & Anlas 2007: 113; Amiet 2008: 45; Japoshvili & Karaca 2010: 199; Klausnitzer et al. 2013: 97; Ceccolini & Barbagli 2014: 95, 97; Augul 2016: 125, 129; Olszewski et al. 2016: 51; Smolis et al. 2017: 161; Shorenko 2018: 126; Tischendorf & Dieterich 2020: 18–20; Verheyde et al. 2021: 3–4; Schmid-Egger & Schmidt 2021: 62; Verheyde et al. 2023: 17–20; Can & Ljubomirov 2023: 523; Golfetti et al. 2025: 3, 10.

Megascolia ( Regiscolia) maculata bischoffi : Osten 1999a: 424; Osten 2000: 546, 551, 557; Osten 2004: 503; Osten 2005b: 37.

Megascolia ( Regiscolia) maculata : Osten 2000: 546, 551; Vereecken & Carrière 2003: 71–74; Milko & Kazenas 2005: 38; Fateryga & Shorenko 2012: 17; Ruchin & Artaev 2016: 2112; Lelej & Mokrousov 2017: 13; Mokrousov & Lelej 2017: 147; Kletenkin 2019: 72; Gadallah & Brothers 2020: 178 View Cited Treatment ; Taylor 2024b: 120.

Megascolia maculata maculata : Giachino et al. 2000: 100; Vereecken & Carrière 2003: 74; Tezcan et al. 2004: 249; Bogush 2007: 168; Schedl 2007: 409; Gusenleitner et al. 2008: 58; Schedl 2010: 1031 View Cited Treatment ; Özbek & Anlas 2011: 630; Baðrýaçýk 2016: 150; Tüzün & Yalniz 2018: 161; Örgel et al. 2020: 640.

Megascolia maculata flavifrons : Vereecken & Carrière 2003: 73–76; Schedl 2006: 345; Schedl 2007: 409; Castro 2011: 387; Baldock 2014: 343; Dvořák & Boščík 2017: 380; Juillerat 2013: 173–174; Baldock et al. 2020: 16.

Material examined. Armenia. Ararat province. 1♂, Geghmahovit , 23.VI.2024 ( M. Aubert & V. Leclercq leg.) [ CJBC] ; Gegharkunik province. 1♂, Getik , 17.VI.2024 ( M. Aubert & V. Leclercq leg.) [ CJBC] . Syunik province. 1♀, Meghri , 19.VI.2024 ( M. Aubert & V. Leclercq leg.) [ CJBC] ; 1♂, Vardanidzor , 19.VI.2024 ( M. Aubert & V. Leclercq leg.) [ CJBC] . Bulgaria. Burgas region. 1♀, Ahtopol , 21.VII.1977 (local coll. leg.) [ CJBC] . Cyprus. Paphos district. 1♀ 6♂, Latchi , 29.V.2015 (local coll. leg.) [ CJBC] ; 5♀ 5♂, Konia , 5.V.2021 ( J. Demetriou leg.) [ CJBC] . Limassol district. 4♀ 5♂, Akrotíri , 14.V.2021 ( J. Demetriou leg.) [ CJBC] . France. Ardèche department. 1♀, Saint- Marcel d’Ardèche, Mas de Libian , 11.VII.2022 ( F. Cabon leg.) [ CFC]. Aude department . 3♀, Conques-sur-Orbiel , 15.VI.2006 (local coll. leg.) [ CJBC] ; 1♂, Castelnau-d’Aude, 6.VII.2011 ( J.-B. Castagnet leg.) [ CJBC] ; 1♀, Monze , 17.VII.2023 ( J.-B. Castagnet leg.) [ CJBC] ; 1♀ 1♂, same place and collector, 16.VII.2024 [ CJBC] ; 1♀, Saint-Martin-Lalande, 24.VII. 2024 ( R. Castagnet leg.) [ CJBC]. Bouches-du-Rhône department . 1♂, Marseille , VI.1991 ( R. Minetti leg.) [ CFC]. Drôme department . 2♂, Saint-Donat-sur-l’Herbasse, 1.VI.2002 (local coll. leg.) [ CJBC] ; 1♀, Donzère , 9.VIII.2013 (local coll. leg.) [ CJBC]. Haute-Corse department . 1♀ 1♂, Monte Grosso , 20.VI.2009 (local coll. leg.) [ CJBC] ; 1♀, Venaco , 4.VII.2015 ( J.-B. Castagnet leg.) [ CJBC] ; 4♀ 1♂, Bravona , 20.VI.2018 ( R. Minetti leg.) [ CFC]. Hérault department . 1♀, Roquebrun , 22.VII.2011 ( J.-B. Castagnet leg.) [ CJBC]. Landes department . 1♀, Biscarosse , 8.VII.1976 (local coll. leg.) [ CJBC] ; 1♀, Moliets-et-Maa , 12.VII.2015 ( J.-B. Castagnet leg.) [ CJBC] ; 2♀, Mimizan , 12.II.2022 ( J.-B. Castagnet leg.) [ CJBC] ; 3♀, Soustons , 12.VII.2023 ( J.-B. Castagnet leg.) [ CJBC]. Lot department . 1♀ 1♂, Gourdon , 3.VII.1973 (local coll. leg.) [ CJBC]. Var department . 1♀, Les Mayons , 1.VIII.2002 ( R. Minetti leg.) [ CFC] ; 1♀, La Cadière , 7.VIII.2012 ( R. Minetti leg.) [ CFC] ; 1♀, Hyères , 22.VII.2016 ( J.-B. Castagnet leg.) [ CJBC] ; 1♂, Rians , 12.VI.2022 ( J.-B. Castagnet leg.) [ CJBC] . Greece. Crete region. 1♂, Heraklion , 12.VI.2017 (local coll. leg.) [ CJBC] ; 2♂, Goúves , 7.VI.2013 ( A. Foucart leg.) [ CJBC] . 1♀, Palaiochora , 1.VIII.2013 (local coll. leg.) [ CJBC] ; 1♀, Chaniá , 18.VII.1998 (local coll. leg.) [ CJBC] ; 1♀, Mathiá , 20.VII.1996 (local coll. leg.) [ CJBC] ; 2♀ 2♂, Kissamos , 27.VI.2014 ( J.-B. Castagnet leg.) [ CJBC] ; 1♀ 1♂, Falassarna , 28.VI.2014 ( J.-B. Castagnet leg.) [ CJBC]. Eeastern Macedonia and Thrace region. 1♀, Loutros-Evros , 1.VII.2007 ( R. Minetti leg.) [ CFC] . Epire region. 1♀, Vrosina, 24.VII.2010 ( R. Minetti leg.) [ CFC]. Laconia region . 3♂, Neapoli, Amitsa beach, 6.V.2025 ( H. Brustel leg.) [ CFC] ; 1♀, Monemvasia , 7.V.2025 ( H. Brustel leg.) [ CFC] . North Aegean region. 1♀, Mithymna , 28.VI.2009 (local coll. leg.) [ CJBC] . Peloponnese region. 1♂, Mykines , 21.V.1995 (local coll. leg.) [ CJBC] ; 1♂, Mycenae , 7.V.2025 ( H. Brustel leg.) [ CFC] ; 2♀, Váthia , 16.VI.1996 (local coll. leg.) [ CJBC] . Thessaly region. 1♀, Meteores , 7.VI.2008 ( R. Minetti leg.) [ CFC] . Iran. Fars province. 2♂, Shiraz , 21.V.2018 (local coll. leg.) [ CJBC] . Italy. Campania region. 1♀, Salerno , 1.VI.2018 (local coll. leg.) [ CJBC] . Sardinia region. 2♀, Nuoro , 20.VI.2006 (local coll. leg.) [ CJBC] . Morocco. Rabat-Salé-Kénitra region. 1♀ 2♂, El Arba, Route de Mrirt , 22.V.2014 ( J.F. Vayssières leg.) [ CFC] ; 2♀ 1♂, Oulmes , 22.V.2015 (local coll. leg.) [ CJBC] . Russia. Rostov oblast. 1♀, Koysug , 14.VI.2019 (local coll. leg.) [ CJBC] . Krasnodar region. 1♀, Leningradskaya , 12VI.2007 (local coll. leg.) [ CJBC] . Spain. Seville province . 1♀, Seville, 19.VII.2013 ( J.-B. Castagnet leg.) [ CJBC]. Tarragona province . 1♂, Tarragona , 21.V.1992 (local coll. leg.) [ CJBC] . Türkiye. 1♀, Coavada , 10.VI.1998 ( L. Grecco leg.) [ CFC] ; 1♀, same place and collector, 11.VI.1998 [ CFC] . Ukraine. Karkiv oblast . 2♀ 2♂, Karkiv , 18.V.2014 (local coll. leg.) [ CJBC] . Odesa Oblast. 1♀, Chornomorsk , 23.VI.2024 (local coll. leg.) [ CJBC] .

Diagnosis. Regiscolia maculata ( Drury, 1773) , comb. nov. differs from R. alecto (Smith, 1858) , comb. nov., R. azurea ( Christ, 1791) , comb. nov., R. capitata ( Fabricius, 1804) , comb. nov. and R. fulvifrons ( Saussure, 1855) , comb. nov. in females by having the apical part of scutum with a dense, uninterrupted band of punctures along the scutellum; the dorso-median area of propodeum short, approximately equal to the length of the metanotum; and the basal part of T2 with sparse punctures similar to those of the median part. In contrast, in R. alecto , R. azurea , R. capitata and R. fulvifrons , the apical part of scutum has sparser punctures medially or is impunctate, thus not forming a continuous band of punctures along the scutellum; the dorso-median area of propodeum is distinctly longer than the metanotum; and the basal part of T2 has finer and denser punctures forming a band, the median part being sparsely and coarsely punctate. In males, R. maculata differs by having the area around and between the posterior ocelli with very dense, tight, fine, and agglomerated punctures; and the length of the dorso-median area of propodeum nearly equal to that of the metanotum. In contrast, in R. alecto , R. azurea , R. alecto and R. fulvifrons , the area around and between the posterior ocelli has sparser and coarser punctures; and the length of the dorso-median area of propodeum is significantly greater than that of the metanotum. R. maculata is easily distinguished from R. capitata in males by having the frons and vertex distinctly more punctate. Whereas in R. capitata , the frons and vertex are almost entirely impunctate. In both sexes, the wings of R. bidens are bicolored, yellowish in the cells and dark elsewhere. Whereas in R. azurea , R. alecto , R. capitata , R. fulvifrons and R. splendida , the wings are entirely dark.

R. maculata differs from R. almoraensis ( Gupta & Jonathan, 2003) , comb. nov. and R. rubida ( Gribodo, 1893) , comb. nov. in females by having the pronotum and the basal part of scutum with very fine, tight and dense punctures. In contrast in R. almoraensis and R. rubida , the pronotum has a large impunctate area close to the tegula or mostly has coarser and sparser punctures; and the basal part of scutum has denser and coarser punctures.

R. maculata differs from R. bidens ( Linnaeus, 1767) , comb. nov. in females by having the dorso-median area of propodeum without projections extended posteriorly; the anterior margin of marginal cell longer than posterior margin; and the T1 with a median tubercle depressed in the middle. In contrast in R. bidens , the dorso-median area of propodeum has two protruding projections prolonged towards the rear; the anterior margin of marginal cell is shorter than the posterior margin; and the T1 has a median tubercle not depressed in the middle.

R. maculata differs from R. almoraensis , R. rubida and R. bidens in males by having the dorso-median area of propodeum without protruding projections prolonged towards the rear. Whereas in R. almoraensis , R. rubida and R. bidens , the dorso-median area of propodeum has two protruding projections prolonged towards the rear.

Redescription. Female ( Fig. 79 View FIGURE 79 ). Length 29–40 mm.

Head ( Figs 79B, C View FIGURE 79 ). Mandibles long and slender. Median mandibular tooth small and prominent, sometimes blunt. Anterior margin of clypeus rounded, without prominent lateral lobes; disk of clypeus not-domed and punctate apically and laterally; impunctate medially. Fissura frontalis passing through the frontal pit and ending at the median ocelli. Frons with numerous and dense punctures. Vertex densely punctate.

Mesosoma ( Fig. 79D View FIGURE 79 ). Pronotum with fine, tight and dense punctures over its entire surface. Tegula with coarse punctures at base and along scutum; with micro-punctures often effaced on rest. Scutum with very fine, tight, coalescent and dense punctures basally; with sparse and coarse punctures laterally, except for antero-lateral corners with more dense, tight and fine punctures; largely impunctate medially; with a fine, dense, and continuous band of punctures apically. Scutellum with fine and dense punctures basally, medially and apically, with sparse punctures laterally. Metanotum with fine, tight and dense punctures, with a thin impunctate central band. Dorso-lateral area of propodeum with very dense, tight and fine punctures, without an impunctate area (mirror) anteriorly. Dorso-median area of propodeum without projections extended posteriorly, or at most with slight relief; approximately equal to the length of the metanotum; with very fine, tight dense punctures. Upper plate of metapleuron impunctate on its lower half, finely and densely punctate on the rest. Anterior margin of marginal cell longer than or equal to posterior margin. Forewing with three submarginal cells. Hind tibiae spurs of similar length.

Metasoma ( Fig. 79E View FIGURE 79 ). T1 with a tubercle depressed in the middle; finely and sparsely punctate basally revealing a small impunctate area in its center; very densely and finely punctate on laterally and apically. T2 with scattered and coarse punctures basally and medially; with very fine, tight and dense punctures apically. T3 almost entirely impunctate basally and medially; with dense, tight and fine punctures apically. T4 to T5 with similar punctation. S1 with dense and fine punctures laterally, mostly impunctate on the rest. S2 without distinct tubercle; with fine and dense punctures basally; with sparse and coarse punctures medially; with dense and fine punctures apically, with sparse punctures in the middle apically. S3 to S5 with similar punctation.

Coloration ( Figs 79–81 View FIGURE 79 View FIGURE 80 View FIGURE 81 ). Head with frons, vertex and tempora more or less marked with yellow or entirely yellow; ocellar region often black. Antennae black to brown. Mesosoma black, scutellum often marked with yellow. Metasoma black, T2 and T3 with two yellow spots sometimes fused. Legs black. Wings bicolored, yellowish in the cells, dark on the rest. Pilosity generally black, except in the colored areas in flavifrons subspecies ( Fig. 80 View FIGURE 80 ); in nominal subspecies ( Fig. 79 View FIGURE 79 ), the pilosity on the pronotum, scutum basally, and apical tergites reddish; in bischoffi subspecies ( Fig. 81 View FIGURE 81 ), only the apical tergites have reddish pilosity.

Male ( Fig. 82 View FIGURE 82 ). Length 21–32 mm.

Head ( Figs 82B, C View FIGURE 82 ). Inner margin of mandible with three distinct tooth (third apical tooth sometimes blunt and barely visible). Disk of clypeus with sparse punctures apically, dense and coarse punctures basally and laterally. Fissura frontalis visible reaching anterior ocelli. Frontal cross-furrow visible, its extension reaching the upper part of the ocular sinus. Frontal spatium with dense, tight and fine punctures. Anterior ocellus depressed, larger than posterior ocelli. Frons with dense, fine and tight punctures, impunctate in front of the anterior ocellus. Vertex with fine, tight and dense punctures, particularly around the posterior ocelli.

Mesosoma ( Fig. 82D View FIGURE 82 ). Pronotum with dense, tight and fine punctures over its entire surface, inter-points spaces almost coalescent. Tegula with dense and coarse punctures anteriorly and along the scutum, largely impunctate near the wing. Scutum with very fine, coalescent and dense punctures in its anterior third; with dense and fine punctures on the rest. Scutellum with fine and dense punctures basally; sparser medially; mostly impunctate apically. Metanotum with fine and dense punctures laterally, impunctate line in the middle. Dorso-lateral area of propodeum without impunctate area (mirror) anteriorly; with fine, dense and tight punctures. Dorso-median area of propodeum without protruding projections; with dense, fine and coalescent punctures over its entire surface. Anterior margin of the marginal cell longer than posterior margin. Forewing with three submarginal cells.

Metasoma ( Fig. 82E View FIGURE 82 ). T1 with a strong tubercle; with dense, fine and tight punctures over its entire surface. T2 with fine, tight and dense punctures, slightly sparse medially. T3-T6 similarly punctate. S1 raised in the middle, with dense punctures medially. S2 without tubercle; with dense and fine punctures basally and laterally, sparser medially and apically. S3 with sparse punctures medially, more denser laterally. S4-S6 similarly punctate.

Coloration ( Figs 82–84 View FIGURE 82 View FIGURE 83 View FIGURE 84 ). Head black, ocular sinus often marked with yellow-orange, sometimes tempora. Antennae black. Mesosoma black. Metasoma black, except T2-T3 marked with two yellow spots, sometimes fused; occasionally T4 marked with yellow (mainly in bischoffi subspecies). Pilosity generally black, except in the colored areas in flavifrons subspecies ( Fig. 83 View FIGURE 83 ); in nominal subspecies ( Fig. 82 View FIGURE 82 ), the pilosity on the pronotum, scutum, and apical tergites reddish; in bischoffi subspecies ( Fig. 84 View FIGURE 84 ), only the apical tergites have reddish pilosity. Wings bicolored, yellowish in the cells, dark on the rest.

Genitalia. See Fig. 31G View FIGURE 31 .

Distribution. Albania, Algeria, Armenia, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Cyprus, Czechia, Egypt, France ( Corsica include) Georgia, Germany, Greece ( Creta, Cyclades, Dodecanese Islands include), Hungary, Iraq, Iran, Israel, Italy ( Sicily include), Kazakhstan ( Atyrau), Kuwait, Macedonia, Malta, Morocco, Netherlands, Portugal, Romania, Russia ( Adygea, Bashkortostan, Chechnya, Chelyabinsk, Dagestan, Ingushetia, Kabardino-Balkaria, Kalmykia, Karachay-Cherkessia, Krasnodar, Nizhny Novgorod, North Ossetia-Alania, Penza, Rostov, Samara, Saratov, Stavropol, Ulyanovsk, Vladimir), Serbia and Montenegro, Slovakia, Slovenia, Spain (Mallorca and Balearic islands include), Switzerland, Syria, Tunisia, Türkiye, Turkmenistan, Ukraine ( Guiglia 1938; Betrem 1941; Steinberg 1962; Friese & Muche 1967; Nagy 1967; Muche 1975; Pagliano 1987; Hamon et al. 1995; Osten 2000; Osten 2002; Osten 2005a; Schedl 2006; Ljubomirov 2006; Bogusch 2007; Fallahzadeh & Saghaei 2010; Özbek & Anlaş 2011; Makhan 2012; Juillerat 2013; Samin et al. 2014; Lohrmann & Engel 2015; Olszewski et al. 2016; Ruchin & Artaev 2016; Lelej & Mokrousov 2017; Mokrousov & Lelej 2017; Dvořák & Boščík 2017; Tüzün & Yalniz 2018; Gadallah & Brothers 2020; Baldock et al. 2020; Örgel et al. 2020; Tischendorf & Dieterich 2020; Verheyde et al. 2021; Demetriou et al. 2021; Verheyde et al. 2023) ( Fig. 85 View FIGURE 85 ). Tischendorf & Dieterich (2020) report an observation in Germany ( Hesse) and discuss the possible acclimatization and establishment in the country.

Biology. The biology of Regiscolia maculata has been extensively studied ( Passerini 1840, 1841; Lepeletier 1845; Fabre 1891; Schumacher 1917; Berland 1925; Cros 1929; Carrière 1990, 2006; Vereecken & Carrière 2003). The species is known to parasitize the following species: Anoxia villosa (Fabricius, 1781) , Lucanus cervus ( Linnaeus, 1758) , Oryctes nasicornis ( Linnaeus, 1758) and Polyphylla fullo ( Linnaeus, 1758) ( Vereecken & Carrière 2003) .

Remarks. Three subspecies of Regiscolia maculata are recognized ( Osten 2000, 2005b): ssp. flavifrons ( Fabricius, 1775) from Italy to Portugal ( Figs 80 View FIGURE 80 , 83 View FIGURE 83 ), ssp. bischoffi ( Micha, 1927) from Cyprus ( Figs 81 View FIGURE 81 , 84 View FIGURE 84 ) and ssp. maculata ( Drury, 1773) from Southwest France and eastern Mediterranean (except Cyprus) ( Figs 79 View FIGURE 79 , 82 View FIGURE 82 ). These subspecies are geographically distinct despite some areas of overlap ( Osten 2000) and are based mainly on the pilosity coloration ( Osten 2000). Piek (1986) and Piek & Carrière (1990) reported the presence of atypical specimens of ssp. flavifrons in southeastern France, showing partially reddish pilosity on the metasoma, similar to that observed in the nominal subspecies.According to them, the coexistence of both forms within the flavifrons population suggests there is no clear geographical separation, thus calling into question the validity of these subspecies. They propose using the term “variation” to distinguish between the two forms ( flavifrons and maculata ). They also observed a possible correlation between low winter temperatures and the appearance of reddish-haired specimens in the southeastern French population ( Piek & Carrière 1990). However, although this hypothesis is intriguing, it should be approached with caution, as specimens from the eastern Mediterranean also show reddish pilosity on the abdomen despite not experiencing harsh winters.

According to Schmid-Egger & Schmidt (2021), DNA barcoding of specimens representing the three subspecies ( maculata , flavifrons , and bischoffi) seems not reveal any genetic sub-clustering. DNA barcoding therefore does not support their subspecies status, and considering their partially overlapping distribution, the current subspecies most likely represent geographically based colour variation. Further studies are needed to clarify the validity of these subspecies.

Numerous subspecies, forms, and varieties have been described: Triscolia maculata barbara Micha 1927 , Triscolia maculata form verticalis Micha 1927, Triscolia maculata sicula Micha 1927 , Scolia ( Triscolia) maculata flavifrons var. funerea Guiglia 1928 , Scolia ( Triscolia) flavifrons var. nigra Dusmet 1930 , Megascolia ( Regiscolia) flavifrons form vespertina Nagy 1967, and Megascolia ( Regiscolia) flavifrons form vernalis Nagy 1967. These taxa are distinguished solely by a few characters related to the coloration of the body and pilosity, such as the extent of black pigmentation on the head in Triscolia maculata barbara Micha 1927 and Triscolia maculata sicula Micha 1927 . However, these coloration characteristics are variable within populations of R. maculata . Consequently, the taxa Triscolia maculata barbara Micha 1927 , syn. nov., Triscolia maculata form verticalis Micha 1927, syn. nov., Triscolia maculata sicula Micha 1927 , syn. nov., Scolia ( Triscolia) maculata flavifrons var. funerea Guiglia 1928 , syn. nov., Scolia ( Triscolia) flavifrons var. nigra Dusmet 1930 , syn. nov., Megascolia ( Regiscolia) flavifrons form vespertina Nagy 1967, syn. nov., and Megascolia ( Regiscolia) flavifrons form vernalis Nagy 1967, syn. nov., are synonymized with R. maculata .

The three valid subspecies can be distinguished using the following key.

Key to subspecies of Regiscolia maculata

Females and males

1. Metasoma with entirely black pilosity ( Figs 80 View FIGURE 80 , 83 View FIGURE 83 ). Mainly North Africa and western Mediterranean....... ssp. flavifrons (Fabricius)

- Metasoma with reddish pilosity on apical tergites ( Figs 79 View FIGURE 79 , 81 View FIGURE 81 , 82 View FIGURE 82 , 84 View FIGURE 84 ). Mainly eastern Mediterranean, plus southwest France................................................................................................... 2

2. Mesosoma with entirely black pilosity ( Figs 81 View FIGURE 81 , 84 View FIGURE 84 ). T4 often marked with yellow (in male). Cyprus.... ssp. bischoffi (Micha)

- Mesosoma with reddish pilosity on pronotum, scutum partially or entirely, sometimes on the scutellum, metanotum and propodeum ( Figs 79 View FIGURE 79 , 82 View FIGURE 82 ). T4 generally black. Eastern Mediterranean (except Cyprus) and southwest France.................................................................................................. ssp. maculata (Drury)