Megascolia velutina ( Saussure, 1859 )

Megascolia velutina ( Saussure, 1859)

( Figs 24 View FIGURE 24 , 28–30 View FIGURE 28 View FIGURE 29 View FIGURE 30 )

Scolia ( Scolia) velutina Saussure, 1859: 175 ( lectotype, ♂, Java, designated by Bradley 1974: 476–477; deposited in RMNH,

The Netherlands, examined). Scolia morosa Smith, 1861: 118 ; Smith 1862: 53 (synonymized by Bradley & Betrem 1964: 441 with velutina velutina ). Scolia ducalis Smith, 1861: 118 . Syn. Nov. Scolia intrudens Smith, 1862: 53 . Syn. Nov. Scolia ( Triscolia) morosa : Saussure & Sichel 1864: 39. Scolia ( Triscolia) velutina : Saussure & Sichel 1864: 41; Cameron 1892: 100. Scolia ( Triscolia) intrudens : Saussure & Sichel 1864: 42. Scolia ( Triscolia) ducalis : Saussure & Sichel 1864: 49; Cameron 1892: 101. Triscolia ducalis : Gribodo 1893: 165. Triscolia intrudens : Gribodo 1893: 166. Scolia View in CoL ( Scolia View in CoL , Triscolia ) ducalis : Dalla-Torre 1897: 155. Scolia View in CoL ( Scolia View in CoL , Triscolia ) intrudens : Dalla-Torre 1897: 166. Scolia View in CoL ( Scolia View in CoL , Triscolia ) morosa : Dalla-Torre 1897: 170. Scolia View in CoL ( Scolia View in CoL , Triscolia ) velutina : Dalla-Torre 1897: 186. Triscolia velutina : Micha 1927: 102. Triscolia velutina velutina : Micha 1927: 102. Triscolia velutina keyensis Micha 1927: 102 . Syn. Nov. Triscolia velutina morosa : Micha 1927: 103. Triscolia ducalis : Micha 1927: 141. Triscolia velutina styx Micha, 1927: 104 (synonymized by Bradley & Betrem 1964: 441 with velutina intrudens ). Triscolia halmaheira Micha, 1927: 104 (synonymized by Bradley & Betrem 1964: 441 with velutina ducalis ). Triscolia halmaheira halmaheira Micha, 1927: 104 . Scolia View in CoL ( Triscolia _ection Megascolia ) ducalis : Betrem 1928: 244. Scolia View in CoL ( Triscolia _ection Megascolia ) ducalis gribodoi Betrem, 1928: 244–245 (synonymized by Bradley & Betrem 1964: 441

with velutina intrudens ). Scolia View in CoL ( Triscolia _ection Megascolia ) velutina : Betrem 1928: 245; Betrem 1933: 256. Scolia View in CoL ( Triscolia _ection Megascolia ) velutina keyensis: Betrem 1928: 246–247; Betrem 1933: 256. Megascolia ( Megascolia) velutina velutina : Betrem & Bradley 1964: 440–441; Bradley & Betrem 1967: 314; Bradley 1974:

476–477; Osten 2005b: 45. Megascolia ( Megascolia) velutina ducalis : Betrem & Bradley 1964: 441; Bradley & Betrem 1967: 302; Osten 2005b: 45. Megascolia ( Megascolia) velutina intrudens : Betrem & Bradley 1964: 441; Bradley & Betrem 1967: 309–310; Bradley 1972:

16; Osten 2005b: 45. Megascolia ( Megascolia) velutina keyensis : Betrem & Bradley 1964: 441; Bradley 1972: 12; Osten 2005b: 45. Megascolia ( Megascolia) velutina : Gupta & Jonathan 2003: 135; Golfetti et al. 2025: 3, 10.

Type material examined. Lectotype of Scolia ( Scolia) velutina (RMNH) , ♂, [white labels] ‘K.s.r.H / Java’, ‘Museum Leiden / Triscolia / velutina / sauss. De Henr / Det: -’, ‘ Scolia / velutina / sauss / Hololectotype / det. Betrem’.

Material examined. Indonesia. Maluku province. 1♀, Ilat, Buru Island , 18.VI.2013 (local coll. leg.) [ CJBC] ; 1♀ 1♂, Ambon, Ambon island , 1.III.2018 (local coll. leg.) [ CJBC] . Central Sulawesi province. 1♀, Palolo , 11.IV.2018 (local coll. leg.) [ CJBC] ; 1♀, same place, 11.IX.2019 (local coll. leg.) [ CJBC] ; 1♂, Peleng, Tataba , X.2017 (local coll. leg.) [ CFC] ; 1♀ 1♂, same place, 12.III.2017 (local coll. leg.) [ CJBC] ; 1♂, same place, 22.XII.2018 (local coll. leg.) [ CJBC] ; 4♀, same place, 6.XII.2019 (local coll. leg.) [ CJBC] ; 1♂, same place, 21.XII.2019 (local coll. leg.) [ CJBC] ; 2♀, Kamarora , 9.IV.2018 (local coll. leg.) [ CJBC] ; 2♀, Palopo , 7.X.2017 (local coll. leg.) [ CJBC] . South Sulawesi province. 1♂, Tompobulu , 10.X.2020 (local coll. leg.) [ CJBC] ; 1♂, Maros , 12.IV.2016 (local coll. leg.) [ CJBC] .

Diagnosis. Megascolia velutina ( Saussure, 1859) differs from M. procer ( Illiger, 1802) and M. scutellaris ( Gribodo, 1893) by having the scutellum and metanotum with dense and fine punctures; and the body entirely melanistic (or only the head partially marked with yellow). In contrast, in M. procer and M. scutellaris , the scutellum and metanotum have sparser and coarser punctures; and at least the head, scutellum and metanotum are marked with yellow.

Redescription. Female ( Fig. 28 View FIGURE 28 ). Length 41–47 mm.

Head ( Figs 28B, C View FIGURE 28 ). Disk of clypeus domed in the middle; densely and finely punctate, more coarsely and sparsely punctate medially. Fissura frontalis distinct, reaching the level of the lamina frontalis and ending in a pronounced pit. Frons sparsely and finely punctate, the ocellar region bearing a few more or less dense and fine punctures. Vertex almost entirely impunctate, with only a few scattered punctures.

Mesosoma ( Fig. 28D View FIGURE 28 ). Pronotum densely and finely punctate over its entire surface, except in front of the tegula with sparser and coarser punctures, revealing a more or less large impunctate surface. Pronotum with a blunt, slightly protruding tubercle in front of the tegula, sometimes barely visible. Tegula largely impunctate; basally and along scutum with fine and dense punctures; sparser behind. Scutum finely and densely punctate basally; more sparsely and coarsely punctate laterally; almost impunctate medially and apically. Scutellum finely and densely punctate all over. Metanotum densely and finely punctate; with a small impunctate area apically. Dorso-lateral area of propodeum densely and finely punctate, especially the area near the lateral area of the propodeum; anteriorly with an impunctate area (mirror); posterior part with dense and fine punctures (coalescent). Dorso-median area of propodeum densely and coarsely punctate, more sparsely punctate medially revealing an impunctate surface. Mesopleura densely and finely punctate all over. Anterior part of upper plate of metapleura densely and finely punctate on anterior two-thirds, more sparsely punctate on remainder, with a small impunctate area on posterior third. Lower plate of the metapleura mostly densely and finely punctate, with a large impunctate surface near the lateral carina of the propodeum. Apical end of hind femur bearing a row of spines. Second transverso-cubital vein arriving well after the junction between the first transverso-cubital vein and the radial area. Anterior margin of marginal cell is about 1.4 times greater than the posterior margin.

Metasoma ( Fig. 28E View FIGURE 28 ). T1 with a weak, pointed or blunt tubercle; very densely and finely punctate laterally and apically; more coarsely and less densely punctate with a small impunctate area basally. T2 impunctate basally; densely and finely punctate laterally; very sparsely punctate medially; finely and densely punctate apically. T3 almost impunctate basally, except for fine and dense punctures along T2; densely and finely punctate laterally and apically. T4 and T5 with similar punctation, T4 and T5 with similar punctation, but the central impunctate area gradually reducing. Hypopygidium with coarse and dense punctures, except apex with impunctate surface. S1 slightly domed and impunctate, except laterally with coarse punctures forming a band meeting in the middle. S2 with strong tubercle; with fine and dense punctures basally; impunctate medially; with dense and fine punctures on sides apically and with coarse and sparse punctures in the middle apically. S3 to S5 with similar punctation.

Coloration ( Figs 28 View FIGURE 28 , 29 View FIGURE 29 ). Head black ( Figs 29B, C View FIGURE 29 ), sometimes more or less yellow mainly on vertex ( Figs 28B, C View FIGURE 28 ). Antennae black. Mesosoma and metasoma entirely black. Legs black. Wings entirely dark, with blue/purple reflections. Pilosity black.

Male ( Fig. 30 View FIGURE 30 ). Length 24–35 mm.

Head ( Figs 30B, C View FIGURE 30 ). Inner margin of mandible with three distinct tooth (third apical tooth sometimes blunt and barely visible). Disk of clypeus densely punctate. Frontal spatium long, densely and finely punctate; pilosity long and dense. Antennal segment XIII about 2.2 times as long as it is wide. Median ocelli slightly depressed, larger than the posterior ocelli. Distinct fissura frontalis reaching the median ocelli and passing through the frontal pit. Vertical frontal furrow distinct, its extension reaching the upper lobe of the eye. Frons densely and finely punctate, particularly around posterior ocelli. Vertex densely and finely punctate.

Mesosoma ( Fig. 30D View FIGURE 30 ). Pronotum finely and densely punctate all over, inter-point space almost coalescent. Tegula largely coarsely and densely punctate, the rest with micropunctures. Scutum with dense and fine punctures. Scutellum with fine and dense punctures, almost impunctate apically; inter-point space medially about equal to the diameter of a point. Metanotum finely and densely punctate. Basal part of dorso-lateral area of propodeum with an impunctate area (mirror); apical part with dense and fine punctures (coalescent). Dorso-median area of propodeum densely punctate, inter-point space clearly smaller than the diameter of a puncture.

Metasoma ( Fig. 30E View FIGURE 30 ). T1 with weak tubercle; very densely and finely punctate laterally and apically; more coarsely and less densely basally and medially.T2 very finely and densely punctate all over, inter-point space approximately equal to the diameter of a puncture medially. T3 with small triangular area almost impunctate medially, rest densely and finely punctate. T4 and T5 densely and finely punctate. S1 slightly domed; entirely impunctate, except on the sides with coarse punctures forming a band meeting in the middle. S2 with a tubercle, finely and densely punctate basally; with sparse and dense punctures medially; with dense and fine punctures on sides apically and with sparse punctures in the middle apically. S3 to S6 with similar punctation.

Coloration ( Fig. 30 View FIGURE 30 ). Entirely black, sometimes with a yellowish spot on the vertex. Wings entirely dark, with greenish or bluish-purple reflections. Body pilosity entirely black.

Genitalia. See Fig. 15C View FIGURE 15 .

Distribution. Indonesia ( Central Sulawesi, Gorontalo, Maluku, North Malaku, North Sulawesi, South Sulawesi, Southwest Papua) ( Micha 1927; Betrem 1928) ( Fig. 24 View FIGURE 24 ).

Biology. Unknown.

Remarks. Four subspecies have been described: ssp. ducalis ( Smith, 1861) from Seram, spp. intrudens ( Smith, 1861) from Sulawesi, ssp. keyensis ( Micha, 1927) from Key islands and ssp. velutina ( Saussure, 1859) from Sulawesi and Maluku ( Betrem 1928; Betrem & Bradley 1964; Osten 2005b). All these subspecies are synonymized with the nominal subspecies for the reasons given below.

M. velutina ducalis ( Smith, 1861) , syn. nov. can only be distinguished from the typical female form by the frons and vertex marked with yellow ( Fig. 28 View FIGURE 28 ), whereas the nominal subspecies is entirely black ( Betrem 1928; Betrem & Bradley 1964) ( Fig. 29 View FIGURE 29 ). Examination of a female specimen from the island of Buru, near Seram, reveals all the characteristics of ssp. velutina , with a fully melanistic head. As with ssp. intrudens (see paragraph below), frons and vertex coloration appears to be variable, and individuals intermediate between subspecies may be encountered, regardless of locality. M. velutina ducalis is therefore considered a synonym of M. velutina .

M. velutina intrudens ( Smith, 1861) , syn. nov. differs from M. velutina ducalis in females only in the yellow spot on the frons and vertex, which does not surround the ocellar triangle, unlike in ssp. ducalis , where it does ( Betrem 1928). In the nominal subspecies, the head is entirely black ( Betrem 1928; Betrem & Bradley 1964). In males, the distinction is made by the wings coloration, which shows greenish-yellow reflections on the outside of the cells and greenish-blue on the inside, whereas in the nominate subspecies, the reflections are coppery-red on the outside of the cells and greenish-yellow on the inside ( Micha 1927). In Sulawesi, female specimens are intermediate between ssp. intrudens and ssp. velutina , with the vertex very slightly marked with yellow. There is no geographical barrier, and both subspecies coexist on Sulawesi. Frons and vertex coloration appear to be intraspecific variations. It is also worth noting that female individuals intermediate between ssp. ducalis and ssp. intrudens can be found on Sulawesi, with the ocellar triangle almost entirely surrounded by yellow. Wings coloration in males does not seem to be a reliable criterion for differentiating subspecies, as it can lose its luster depending on the state of conservation and appears to vary within the same population. Thus, M. velutina intrudens is considered a synonym of M. velutina .

M. velutina keyensis ( Micha, 1927) , syn. nov. can be distinguished from the other subspecies by the bluish-purple reflections on the wings ( Micha 1927; Betrem 1928). No specimens belonging to this subspecies could be examined, but as with the other subspecies, the validity seems doubtful and a long series would have to be examined before a decision could be made. In the meantime, it is placed in synonymy with the nominal subspecies.

Genus Regiscolia Betrem & Bradley, 1964 , stat. nov.

( Figs 1B, 2B View FIGURES 1–2 , 6B View FIGURES 3–6. 3–5 , 31–92 View FIGURE 31 View FIGURE 32 View FIGURES 33–34 View FIGURES 35–36 View FIGURES 37–38 View FIGURES 39–40 View FIGURE 41 View FIGURES 42–43 View FIGURE 44 View FIGURE 45 View FIGURE 46 View FIGURE 47 View FIGURE 48 View FIGURE 49 View FIGURE 50 View FIGURE 51 View FIGURE 52 View FIGURE 53 View FIGURE 54 View FIGURE 55 View FIGURE 56 View FIGURE 57 View FIGURE 58 View FIGURE 59 View FIGURE 60 View FIGURE 61 View FIGURE 62 View FIGURE 63 View FIGURE 64 View FIGURE 65 View FIGURE 66 View FIGURE 67 View FIGURE 68 View FIGURE 69 View FIGURE 70 View FIGURE 71 View FIGURE 72 View FIGURE 73 View FIGURE 74 View FIGURE 75 View FIGURE 76 View FIGURE 77 View FIGURE 78 View FIGURE 79 View FIGURE 80 View FIGURE 81 View FIGURE 82 View FIGURE 83 View FIGURE 84 View FIGURE 85 View FIGURE 86 View FIGURE 87 View FIGURE 88 View FIGURE 89 View FIGURE 90 View FIGURE 91 View FIGURE 92 )

Scolia ( Triscolia) : Saussure 1863: 17; Saussure & Sichel 1864: 38; Rohwer 1921: 77; Betrem 1927a: xcvii; Betrem 1941:

117–118; Steinberg 1962: 115. Ascoli Agassiz, 1846: 4 (synonymized with Megascolia ( Regiscolia) by Taylor 2024b: 120). Triscolia : Gribodo 1893: 150; Micha 1927: 79. Scolia View in CoL ( Triscolia sectio Triscolia ): Betrem 1928: 228; Betrem 1933: 254–255. Megascolia ( Regiscolia) Betrem & Bradley, 1964: 441–442 ; Krombein 1978: 10, 24; Argaman 1996: 188; Osten 1999a: 424;

Osten 2000: 551; Gupta & Jonathan 2003: 138–139; Osten 2005a: 1455; Osten 2005b: 26; Kumar & Rajmohana 2017: 5;

Mokroussov & Lelej 2017: 147; Liu et al. 2021a: 103, 114; Taylor & Barthélémy 2021: 27; Pham & van Achterberg 2023: 383; Taylor 2024a: 995; Taylor 2024b: 120; Barthélémy & Guenard 2025: 105.

Ascolia Argaman, 1996: 188 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Elpaholt a Argaman, 1996: 194 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Laskariska Argaman, 1996: 188 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Zazilayza Argaman, 1996: 188 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Guigliana Argaman, 1996: 192 (synonymized with Megascolia ( Regiscolia) by Taylor & Barthélémy 2021: 28).

Paconzitva Argaman, 1996: 196 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Sugorpilfa Argaman, 1996: 196 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Xirgoniqua Argaman, 1996: 196 (synonymized with Megascolia ( Regiscolia) by Taylor 2024a: 995).

Type species. Scolia flavifrons Fabricius, 1775 [ = Regiscolia maculata ( Drury, 1773) ], by original designation.

Species included. Nine species: Regiscolia alecto (Smith, 1858) , comb. nov., R. almoraensis ( Gupta & Jonathan, 2003) , comb. nov., R. azurea ( Christ, 1791) , comb. nov., R. bidens ( Linnaeus, 1767) , comb. nov., R. capitata ( Fabricius, 1804) , comb. nov., R. fulvifrons ( Saussure, 1855) , comb. nov., R. maculata ( Drury, 1773) , comb. nov., R. rubida ( Gribodo, 1893) , comb. nov., and R. splendida ( Saussure, 1858) , comb. nov. ( Bradley & Betrem 1964; Gupta & Jonathan 2003).

Diagnosis. Regiscolia Betrem & Bradley, 1964 , stat. nov. is distinguished from the genus Megascolia Betrem, 1928 in both sexes by having a dorso-median area of propodeum flat, with a distinct transition to the postero-median area, forming a ridge; and the latero-tergal area of T3 with a groove crossing the entire sclera. In Megascolia , the dorso-median area of propodeum is sloping posteriorly, with an indistinct transition to the postero-median area; and the latero-tergal area of T3 does not have a groove. Furthermore, females of Regiscolia stat. nov. have the pronotum without tubercle in front of the tegula; the hind basitarsus with a scopa on the inner extremity; and the apex of hind basitarsus bordered by a row of spines of very unequal size. Whereas in Megascolia , the pronotum has a more or less prominent tubercle in front of the tegula; the hind basitarsus does not have a scopa; and the apex of hind basitarsus is bordered by a dense row of spines of similar size.

Regiscolia stat. nov. is distinguished from the genus Gigascolia Castagnet & Cabon , gen. nov. in both sexes by having the T1 with a prominent tubercle instead of a vertical groove. In females, Regiscolia stat. nov. is distinguished from the genus Gigascolia Castagnet & Cabon , gen. nov. by having the inner margin of mandible with a central tooth; the head rounded; the lamina frontalis strong and prominent; the callosity of pronotum without carina; the coxae III elongated; and the T1 campaniform, gradually widening. In Gigascolia gen. nov., the inner margin of mandible has a sharp apical tooth; the head is sub-quadrangular; the lamina frontalis is weaker, not very prominent; the callosity of pronotum has a vertical carina; the coxae III is rounded; and the T1 is pyriform, narrowing at the basal quarter. In males, Regiscolia stat. nov. is distinguished from the genus Gigascolia Castagnet & Cabon , gen. nov. by having the inner margin of mandible with three distinct teeth; and the mesosoma long, about 1.2 to 1.3 times as long as wide. In Gigascolia gen. nov., the inner margin of mandible has two distinct teeth; and the mesosoma is larger, almost as long as wide.

Redescription. Female. Mandibles with a single central tooth on the inner margin. Clypeus about twice as long as wide. Anterior margin of clypeus wide, truncated or subtruncated in median part; disk of clypeus not domed, impunctate or with rugosities; bordered by a fringe of dense setae. Lamina frontalis strong, elevated and densely punctate at the level of the frontal spatium. Frontal area forming an elongated isosceles triangle. Frons and vertex more or less densely and coarsely punctate. Rounded head, longer than wide. Mesosoma longer than wide. Callosity of pronotum without vertical carina. Pronotum without tubercle in front of tegula. Scutum wider than long. Dorso-median area of propodeum wider than long; flat, transition to postero-median area forming pronounced ridge; at the same level as the dorso-lateral areas of propodeum. Coxae III elongated, without prominent tubercle on inner surface. Hind metatarsus with scopa at inner end; apex of hind metatarsus bordered by a row of spines of very unequal size. Forewing with three submarginal cells (except for one species: R. fulvifrons ). Second transverso-cubital vein arrives at or before the junction between the first transverso-cubital vein and the radial area. T1 with a transverse tubercle, sometimes depressed in the center, but without a vertical transverse groove; campaniform, gradually widening. Apical margin of basal tergites with tight and fine punctures; long and dense pilosity forming a band. Lateral tergal area of T3 with a groove crossing the tergite from one end to the other, this part bearing dense micro-punctures. S1 short, raised in the middle and densely punctate over most of its surface.

Male. Inner margin of mandible with three distinct tooth (third apical tooth sometimes blunt and barely visible). Antennal articles short, segment III 1.4 to 1.8 times longer than wide. Frontal area forming an elongated isosceles triangle. Mesosoma long, clearly longer than wide. Scutum 1.4 to 1.5 times wider than long. Dorso-median area of propodeum flat, wider than long, transition to postero-median surface forming a pronounced ridge. Dorso-median and dorso-lateral areas densely and finely punctate, sometimes with a small impunctate surface. T1 with a strong tubercle, without vertical transverse groove. Latero-tergal area of T3 with furrow crossing tergite from one end to the other, with dense micro-punctures. S1 raised in the middle and densely punctate over most of its surface.

Genitalia ( Fig. 31 View FIGURE 31 ). Gonocoxites forming two sclerotized half-rings that touch in the middle on the ventral side and at the base on the dorsal side. Ventral part of each gonocoxite subrectangular, obliquely arranged; dorsal part larger, slightly swollen. Side of each gonocoxite extended by an elongated gonostyle, fairly wide, with subparallel margin, widening slightly towards the apex, subtruncate at the apex. Gonostylus with numerous setae; setae on outer surface fine, fairly long and dense; setae on inner surface shorter, thicker, spiniform and less numerous, in addition to fine, longer, scattered setae.

Distal part of volsella elongated (about twice as high as wide), progressively narrowed, pointed at apex, like a knife blade. Basal part with rounded basal lobe at proximal edge. Basal and distal parts of volsella separated by distinct, fairly deep notch. Ventral side of basal part of volsella has fairly long, spaced setae. Distal part has a few spaced, mostly short setae, a few longer setae on outer edge. Brown sensory conules, 25 to 50 μm long, present at different levels of the volsella: a dozen closely spaced conules are located at the end of the lateral lobe of the basal part; numerous conules arranged in several irregular rows along the lateral edge of the volsella, in its middle part; two groups of two or three conules in a more distal position.

Aedeagus almost reaching the gonostyles in length. Sclerified penile valves joined by membrane, which forms a rounded cap at the apex of the aedeagus. Ventral edge of the distal part of the penile valve provided with about fifteen sclerified teeth, arranged in an almost straight line, progressively increasing from the apex to the base; basal teeth spaced further apart, irregularly shaped.

Distribution. Mainly Palaearctic, Indomalayan, and Australasian (only the islands north of Australia and east of Papua): Afghanistan, Albania, Algeria, Armenia, Austria, Belgium, Bhutan, Bosnia and Herzegovina, Bulgaria, Cambodia, China, Croatia, Cyprus, Czechia, Egypt, France, Georgia, Greece, Hong-Kong, Hungary, India, Indonesia, Iraq, Iran, Israel, Italy, Kazakhstan, Kirgistan, Kuwait, Laos, Libya, Macedonia, Malta, Malaysia, Morocco, Myanmar, The Netherlands, Uzbekistan, Pakistan, The Philippines, Portugal, Romania, Russia, Serbia and Montenegro, Slovakia, Slovenia, Spain, Switzerland, Syria, Tajikistan, Thailand, Tunisia, Turkey, Turkmenistan, Ukraine, Vietnam ( Betrem 1928; Guiglia 1940; Betrem 1941; Betrem 1947; Steinberg 1962; Baltazar 1966; Nagy 1967; Pagliano 1987; Osten 2000; Gupta & Jonathan 2003; Osten 2005a; Bogusch 2007; Özbek & Anlaş 2011; Juillerat 2013; Samin et al. 2014; Olszewski et al. 2016; Nidup et al. 2017; Al-Azab 2020; Baldock et al. 2020; Verheyde et al. 2021; Demetriou et al. 2021; Liu et al. 2021a; Taylor & Barthélémy 2021; Verheyde et al. 2023; Pham & van Achterberg 2023) (fig. 32).

Remarks. The taxon philippinensis Rohwer, 1921 , usually regarded as a valid species, is here treated as a senior synonym of alecto Smith, 1858 . Osten (2005b) includes one additional species in the subgenus Regiscolia , Scolia ( T.) hageni Betrem, 1928 . Nevertheless, this species is placed here in synonymy with Regiscolia azurea ( Christ, 1791) (see remarks of R. azurea ). Argaman (1996) proposed splitting the subgenus Regiscolia sensu Betrem & Bradley (1964) into nine genera ( Ascolia , Elpaholta , Guigliana , Laskariska , Paconzitva , Regiscolia , Sugorpilfa , Xirgoniqua , Zazilayza ), practically one genus for each known species. This subdivision was revised and invalidated by Taylor (2024a).

Key to species of Regiscolia Betrem & Bradley, 1964 , stat. nov.

Females

(undescribed in R. splendida )

1. Apical part of the scutum with a dense, uninterrupted band of punctures along the scutellum ( Fig. 33A View FIGURES 33–34 ). Dorso-median area of propodeum short, approximately equal to the length of the metanotum. Basal part of T2 with sparse punctures, similar to the punctures of the median part ( Fig. 34A View FIGURES 33–34 ). Wings bicolored, yellowish in the cells, dark on the rest (except in almoraensis , wings almost entirely black).................................................................................. 2

- Apical part of the scutum with sparse punctures in the middle or impunctate, not forming a continuous band of punctures along the scutellum ( Fig. 33B View FIGURES 33–34 ). Dorso-median area of propodeum clearly longer than the length of the metanotum. Basal part of T2 with fine, tight and dense punctures forming a band; punctation of median part sparse and coarse ( Fig. 34B View FIGURES 33–34 ). Wings entirely dark................................................................................................ 5

2. Pronotum close to tegula with a large impunctate area or mostly with coarse and sparse punctures ( Figs 35A, B View FIGURES 35–36 ). Basal part of scutum with dense and coarse punctures ( Figs 35A, B View FIGURES 35–36 ). Dorso-median area of propodeum with two protruding projections prolonged towards the rear ( Figs 36A, B View FIGURES 35–36 ).................................................................. 3

- Pronotum mostly with very fine, tight and dense punctures ( Figs 35C, D View FIGURES 35–36 ). Basal part of scutum with very dense, tight and fine punctures ( Figs 35C, D View FIGURES 35–36 ). Dorso-median area of propodeum with two protruding projections prolonged towards the rear or without projections extended posteriorly ( Figs 36C, D View FIGURES 35–36 )....................................................... 4

3. Frons with fine and sparse punctures. Pronotum close to tegula with a large impunctate area ( Fig. 35A View FIGURES 35–36 ). Anterior margin of marginal cell longer than posterior margin. Wings bicolored, yellowish in the cells, dark on the rest. Pilosity entirely ferruginous. Most of the integument reddish-brown. T2 and T3 marked with yellow............................ R. rubida (Gribodo)

- Frons with coarser and denser punctures. Pronotum with coarse and sparse punctures, without distinct impunctate area ( Fig. 35B View FIGURES 35–36 ). Anterior margin of marginal cell equal than posterior margin. Wing almost entirely black, yellowish area restricted to the costal margin near the marginal cell. Pilosity entirely black. Integument predominantly black. T3 marked with orange-reddish, sometimes T4 weakly...................................................... R. almoraensis (Gupta & Jonathan) .

4. Fissura frontalis inconspicuous, not reaching anterior ocelli. Dorso-median area of propodeum with two protruding projections prolonged towards the rear ( Fig. 36C View FIGURES 35–36 ). Anterior margin of marginal cell shorter than posterior margin ( Fig. 37A View FIGURES 37–38 ). T1 with a median tubercle not depressed in the middle ( Fig. 38A View FIGURES 37–38 ). Antennae orange-red, except scape and pedicel.................................................................................................... R. bidens (Linnaeus)

- Fissura frontalis reaching anterior ocelli. Dorso-median area of propodeum without projections extended posteriorly, or at most with slight relief ( Fig. 36D View FIGURES 35–36 ). Anterior margin of marginal cell longer than posterior margin ( Fig. 37B View FIGURES 37–38 ). T1 with a median tubercle depressed in the middle ( Fig. 38B View FIGURES 37–38 ). Antennae entirely black, sometimes brownish........... R. maculata (Drury)

5. Frons and vertex with fine, shallow and very scattered punctures ( Fig. 39A View FIGURES 39–40 ). Ocello-ocular distance at least 3 times longer than the distance between the posterior ocelli. Basal part of scutum at notauli level with a large rounded area with very fine, dense and colescent punctures, forming a tuft of long setae ( Fig. 40A View FIGURES 39–40 ). Scutellum, metanotum, dorso-median area of propodeum, T1 generally marked with yellow. Pilosity of pronotum, metanotum, propodeum and metasoma forming tufts of very dense, long setae.............................................................................. R. capitata (Fabricius)

- Frons and vertex with coarse and deep punctures, frons punctation always denser ( Fig. 39B View FIGURES 39–40 ). Ocello-ocular distance 1.9 to 2.2 times longer than the distance between the posterior ocelli. Basal part of scutum at notauli more or less coarsely and densely punctate, with normal pilosity ( Fig. 40B View FIGURES 39–40 ). Scutellum, metanotum, dorso-median area of propodeum and T1 entirely black. Mesosoma View in CoL and metasoma pilosity less dense and long........................................................ 6

6. Apical margin of clypeus with two prominent lateral lobes; disk of clypeus domed and fully reticulated. Vertex with denser punctation ( Fig. 41A View FIGURE 41 ). Apical row of punctures on T3, of similar height along most of its length, the middle slightly narrowed. Metasoma marked yellow-orange on T3, often T4 and T5, sometimes T2, black pilosity except in the colored areas. Usually two submarginal cells, sometimes three.................................................. R. fulvifrons (Saussure)

- Apical margin of clypeus more rounded, without distinct lateral lobes; disk of clypeus flat, entirely impunctate, weakly striated only at apex. Vertex with few scattered punctation ( Fig. 41B View FIGURE 41 ). Apical row of punctures in T3, very indented in the middle, revealing an impunctate triangular surface. Metasoma entirely black with black or red pilosity if T3 and/or T4 marked with orange, pilosity reddish on apical tergites. Three submarginal cells.............................................. 7

7. Pronotum with dense, tight and fine punctures over its entire surface, at most a small area with sparse punctures in front of the excavation of pronotum ( Fig. 42A View FIGURES 42–43 ). Transition area between the dorso-median and postero-median areas of propodeum forming an angle without protrusion. T1 with a median tubercle not depressed ( Fig. 43A View FIGURES 42–43 ). T2 basally with dense and fine punctures forming a continuous band across its entire width ( Fig. 43A View FIGURES 42–43 ). Metasoma black. Pilosity black (except in ssp. regnatrix with reddish pilosity on T2 apically and pygidium).................................................. R. alecto (Smith)

- Pronotum with coarse and sparse punctures in the posterior half, revealing a large impunctate surface in front of the excavation of pronotum ( Fig. 42B View FIGURES 42–43 ). Transition area between the dorso-median and postero-median areas of propodeum generally forming a protruding ridge. T1 with a median tubercle often depressed in the middle ( Fig. 43B View FIGURES 42–43 ). T2 basally usually with dense and fine punctures forming a band, punctation interrupted in the middle ( Fig. 43B View FIGURES 42–43 ). Metasoma black, often marked with orange-red on T4, T5. Pilosity reddish on the last tergites, exceptionally with entirely melanistic pilosity............. R. azurea (Christ)

Males

1. Frons and vertex almost entirely impunctate, with fine, scattered and shallow punctures ( Fig. 44A View FIGURE 44 ). Frontal lamina short, the extension of the frontal cross-furrow delimiting frontal spatium and frons ending in the lower part of the ocular sinus. Distance between the posterior ocelli, 1.7 times shorter than the ocello-ocular distance. Scutellum, metanotum, dorso-median area of propodeum, T1, and T3 marked with yellow. Mesosoma View in CoL and metasoma pilosity forming very dense and long tufts.......................................................................................... R. capitata (Fabricius)

- Frons and vertex more densely punctate, with more coarse, dense and deep punctures ( Fig. 44B View FIGURE 44 ). Frontal lamina long, the extension of the frontal cross-furrow delimiting frontal spatium and frons ending in the middle of the ocular sinus or in its upper part. Distance between the posterior ocelli, equal to or greater than the ocello-ocular distance. Metanotum, dorso-medial area of propodeum, and T1 generally entirely black. Pilosity of the mesosoma and metasoma long and dense................ 2

2. Area around and between the posterior ocelli with very dense, tight, fine and agglomerated punctures ( Fig. 45A View FIGURE 45 ). Length of the dorso-median area of propodeum almost equal to the length of the metanotum. Bicolored wings, yellowish in the cells, dark on the rest (except in almoraensis , wings almost entirely black)................................................... 3

- Area around and between the posterior ocelli with more sparse and coarse punctures (in R. fulvifrons , punctation between the posterior ocelli often more fine and dense, but not agglomerated) ( Fig. 45B View FIGURE 45 ). Length of the dorso-median area of propodeum significantly greater than the length of the metanotum. Wings entirely dark....................................... 5

3. Upper margin of the marginal cell equal or almost equal to the lower margin ( Fig. 46A View FIGURE 46 ). Yellow antennae (except for the scape and pedicel). T3 marked with two yellow spots.............................................. R. bidens (Linnaeus)

- Upper margin of the marginal cell significantly longer than the lower margin ( Fig. 46B View FIGURE 46 ). Black or orange-reddish antennae. T2 and T3 marked with two yellow spots, sometimes fused or T3 with two orange-reddish spot.......................... 4

4. Basal third of the scutum with dense, fine, and tight punctures. Dorso-median area of propodeum with two protruding projections prolonged towards the rear.............................................................................. 5

- Basal third of the scutum with extremely dense, fine, and coalescent punctures. Dorso-median area of propodeum without protruding projections prolonged towards the rear, or at most with a slight relief................... R. maculata (Drury)

5. Orange-reddish antennae; flagellomere 1 with micropunctures similar to that of the following flagellomeres. Wings bicolored, yellowish in the cells, dark on the rest. Pilosity entirely ferruginous. Most of the integument reddish-brown. T2 and T3 marked with yellow........................................................................... R. rubida (Gribodo)

- Black antennae; flagellomere 1 without micropunctation, unlike the following flagellomeres. Wing almost entirely black, yellowish area restricted to the costal margin near the marginal cell. Pilosity entirely black. Integument predominantly black. T3 marked with orange-reddish, sometimes T4 weakly............................ R. almoraensis (Gupta & Jonathan) .

6. Tegula with coarse and dense punctures over two-thirds of its surface, the rest with micropunctures. Scutum with dense and fine punctures, interspaces between punctures approximately equal to the diameter of a point; without any distinct impunctate area medially ( Fig. 47A View FIGURE 47 ). T3 with dense punctures across its entire surface, small area in the middle with sparse punctures. Metasoma marked with yellow-orange on T3, sometimes on T4, T5, and T6. Black pilosity, yellow on the colored areas. Usually two submarginal cells, sometimes three............................................ R. fulvifrons (Saussure)

- Tegula with coarse and sparse punctures on its basal third, the rest micropunctures more or less visible. Scutum with coarser and sparser punctures, in the middle interspaces between punctures significantly larger than the diameter of a puncture; with a more or less large impunctate area medially ( Fig. 47B View FIGURE 47 ). T3 with dense punctures, with a large impunctate central area. Metasoma entirely black or marked with yellow-orange on T3 or reddish on the apical tergites. Black pilosity, yellow on the colored surfaces, or reddish on the apical tergites. Three submarginal cells.............................................. 7

7. T1 with dense and fine punctures across its entire surface; area around the tubercle with interspaces between punctures equal to or smaller than the diameter of a puncture; on the tubercle punctation sometimes more spaced ( Fig. 48A View FIGURE 48 ). Frons, vertex, and tempora entirely yellow-orange, sometimes also clypeus. Metasoma entirely black or marked with yellow-orange on T3. Pilosity of the metasoma entirely black, or black pilosity with yellow on the colored surfaces......................... 8

- T1 with dense and fine punctures laterally and apically; the area around the tubercle more or less impunctate, with coarse punctures and the interspaces between punctures significantly greater than the diameter of a puncture; on the tubercle punctation very sparse ( Fig. 48B View FIGURE 48 ). Head black, sometimes marked with orange-reddish in the ocular sinus or with a fine band behind frontal spatium (except in michaae ssp. with head mostly reddish). Metasoma black, with T4, T5, T6, and T7 generally marked with orange-reddish, sometimes T3. Pilosity of metasoma black, with reddish pilosity on the apical tergites R. azurea (Christ)

8. Mesosoma View in CoL and metasoma black. The Philippines, Malaysia, Indonesia (Borneo, Sulawesi, Makian islands)..................................................................................................... R. alecto (Smith)

- Mesosoma View in CoL black or marked with yellow-orange on the pronotum, scutum, and scutellum. Metasoma black, marked with yellow-orange on T3. Indonesia ( Flores, Wetar and Rote islands).................................... R. splendida (Saussure)