Regiscolia azurea ( Christ, 1791 ), 2025

Regiscolia azurea ( Christ, 1791) , comb. nov.

( Figs 60–65 View FIGURE 60 View FIGURE 61 View FIGURE 62 View FIGURE 63 View FIGURE 64 View FIGURE 65 )

Sphex azurea Christ, 1791: 256 ( holotype, female, probably lost, depositories unknown; not examined; type locality unknown).

Scolia rubiginosa Fabricius, 1793: 230 ; Coquebert 1801: 55; Fabricius 1804: 241; Klug 1810: 211; Lepeletier 1845: 518–519; Burmeister 1854: 19; Smith 1855: 111; Cameron 1901: 18; Paiva 1907: 14; Vitalis de Salvaza 1919: 288 (synonymised the same year independently by Micha 1927: 117 and Betrem 1927b: 292). Syn. nov.

Scolia ( Scolia) rubiginosa : Guérin-Méneville 1830: 247.

Scolia ornata Lepeletier, 1845: 517–518 (synonymized by Burmeister 1854: 19).

Scolia ( Scolia) magnifica Saussure, 1859: 175 . Syn. nov.

Scolia ( Triscolia) magnifica : Saussure & Sichel 1864: 44; Cameron 1892: 101.

Scolia ( Triscolia) rubiginosa : Saussure & Sichel 1864: 45; Magretti 1892: 236; Cameron 1892: 101; Bingham 1897: 76-77; Mantero 1903: 33-34; Rohwer 1921: 77.

Scolia ( Triscolia) rubiginosa var. ornata : Saussure & Sichel 1864: 45–46.

Triscolia vespillo Gribodo, 1893: 168-169 . Syn. nov.

Scolia View in CoL ( Scolia View in CoL , Triscolia ) rubiginosa : Dalla-Torre 1897: 180.

Triscolia rubiginosa : Cameron 1905: 53; Leefmans 1915: 84; Leefmans 1921: 47.

Triscolia azurea : Micha 1927: 117, 153.

Triscolia azurea azurea : Micha 1927: 117–121, 153.

Triscolia azurea ornata : Micha 1927: 121, 153.

Triscolia azurea hindostana Micha 1927: 121–122 , 153.

Triscolia democratica Micha, 1927: 122 . Syn. nov.

Triscolia democratica democratica Micha, 1927: 122–123 .

Triscolia philippinensis erratica Micha, 1927: 124 (nom. nov.: azurea michaae in Betrem 1928: 233).

Scolia ( Triscolia) azurea : Betrem 1927b: 292.

Scolia ( Triscolia) philippinensis : Betrem 1927b: 292.

Scolia ( Triscolia) alecto : Betrem 1927b: 292.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea : Betrem 1928: 230; Betrem 1933: 254.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea var. intermedia Betrem, 1928: 231 ; Betrem 1933: 255 (synonymized by Guiglia & Betrem 1958: 96).

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea rubiginosa : Betrem 1928: 231–232; Betrem 1933: 255.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea rubiginosa var. magnifica : Betrem 1928: 232.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea siamensis Betrem, 1928: 232.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea cochinensis Betrem, 1928: 232. Syn. nov.

Scolia View in CoL ( Triscolia sectio Triscolia ) azurea subspecies or varietas hindostana Betrem, 1928: 232 . Scolia View in CoL ( Triscolia sectio Triscolia ) azurea var. democratica : Betrem 1928: 232–233. Scolia View in CoL ( Triscolia sectio Triscolia ) azurea michaae Betrem, 1928: 233. Scolia View in CoL ( Triscolia sectio Triscolia ) hageni Betrem, 1928: 233 . Syn. Nov. Scolia View in CoL ( Triscolia sectio Triscolia ) azurea azurea : Betrem 1933: 255. Scolia View in CoL ( Triscolia sectio Triscolia ) azurea azurea var. rubiginosa : Betrem 1933: 255. Scolia ( Triscolia) azurea hindostana : Betrem 1941: 118; Betrem 1947: 414. Scolia ( Triscolia) azurea rubiginosa : Betrem 1941: 118–119; Betrem 1947: 415. Scolia ( Triscolia) azurea michae : Betrem 1941: 118; Guiglia 1965: 323–324 ( lapsus calami for michaae). Scolia ( Triscolia) azurea azurea : Betrem 1941: 118; Betrem 1947: 415. Scolia ( Triscolia) azurea democratica : Betrem 1947: 414. Scolia ( Triscolia) azurea christiana Guiglia & Betrem, 1958: 96 (new name for the population from Myanmar and northern

India). Syn. nov. Megascolia ( Regiscolia) azurea azurea : Betrem & Bradley 1964: 443; Gupta & Jonathan 2003: 145; Osten 2005b: 27; Pham &

Van Achterberg 2023: 383. Megascolia ( Regiscolia) azurea azurea var. rubiginosa : Betrem & Bradley 1964: 443; Osten 2005b: 27. Megascolia ( Regiscolia) azurea azurea var. democratica : Betrem & Bradley 1964: 443; Osten 2005b: 27. Megascolia ( Regiscolia) azurea christiana : Betrem & Bradley 1964: 443; Gupta & Jonathan 2003: 145–147; Osten 2005b: 27;

Kumar 2009: 105–107; Kumar & Pham 2015: 326; Kumar & Rajmohana 2017: 5; Nidup et al. 2017: 13. Megascolia ( Regiscolia) azurea christiana var. magnifica : Betrem & Bradley 1964: 444. Megascolia ( Regiscolia) azurea siamensis : Betrem & Bradley 1964: 444; Gupta & Jonathan 2003: 145; Osten 2005b: 27. Megascolia ( Regiscolia) azurea cochinensis : Betrem & Bradley 1964: 444; Gupta & Jonathan 2003: 145; Osten 2005b: 27;

Pham & Van Achterberg 2023: 383. Megascolia ( Regiscolia) azurea hindostana : Betrem & Bradley 1964: 444; Bradley 1972: 10; Gupta & Jonathan 2003: 147–148;

Osten 2005b: 27; Kumar & Rajmohana 2017: 5; Jadhav & Gaikwad 2019: 373. Megascolia ( Regiscolia) azurea rubiginosa : Bradley & Betrem 1964: 23; Baltazar 1966: 230. Megascolia ( Regiscolia) azurea azurea form democratica : Bradley 1972: 8. Megascolia ( Regiscolia) azurea azurea form rubiginosa : Bradley 1972: 8. Megascolia ( Regiscolia) azurea azurea form magnifica : Bradley 1974: 450–451. Megascolia ( Regiscolia) azurea michaae : Krombein 1978: 5, 24; Krombein 1995: 84; Gupta & Jonathan 2003: 145; Osten

2005b: 27. Guigliana azurea : Argaman 1996: 196. Megascolia ( Regiscolia) azurea : Gupta & Jonathan 2003: 145; Kumar & Rajmohana 2017: 5; Liu et al. 2021a: 103, 114; Liu et al. 2021b: 148; Taylor & Barthélémy 2021: 28; Golfetti et al. 2025: 3, 10; Barthélémy & Guenard 2025: 105. Megascolia ( Regiscolia) azurea azurea var. magnifica : Osten 2005b: 27. Megascolia cf. azurea : Khouri et al. 2022: 24–28, 37–38. Megascolia cf. hageni : Khouri et al. 2022: 24–29, 31–34, 36–38.

Type material examined. Holotype of Triscolia vespillo Gribodo, 1893 (MSNG), ♀, ‘Coll. Gribodo / Perak’, ‘Vespillo / G Grib’, ‘TYPUS’, ‘ HOLOTYPE / vespillo gr / Teste, 1929. / J.C. Bradley’ ( Fig. 66 View FIGURE 66 ).

Holotype of Scolia hageni Betrem, 1928 (RMNH), ♀, ‘Dr. B. Hagen. / Tandjong Morawa. / Serdang / (C.O. Sumatra)’, ‘ Scolia / hageni Betr / Holotype / det. Betrem’ ( Fig. 67 View FIGURE 67 ).

Allotype of Scolia hageni Betrem, 1928 (RMNH), ♂, ‘Ludeking / Sumatra’, ‘ Scolia / hageni Betr. / Allotype / det. Betrem’ ( Fig. 68 View FIGURE 68 ).

Other material examined. Cambodia. Kampong Cham province. 4♂, Lve, Kaoh Soutin , 18.II.2024 ( F. Cabon & P. Bun leg.) [ CFC] ; 1♀ 1♂, same place, 19.II.2024 ( F. Cabon & P. Bun leg.) [ CFC] . Battambang province. 1♀, Voat kor, 16.III.2024 ( F. Cabon leg.) [ CFC] . Takeo province. 5♂, Phnom Takhmao, Kirisu farms, 9.VI.2022 ( P.- O. Maquart leg.) [CFC, CJBC] . Ratanakiri province. 1♀, Veun Sai, Kaoh Piek , 15.IV.2022 ( P.-O. Maquart leg.) [ CPOM] . China. Yunnan province. 1♀, Mangyun , 12.VI.2020 (local coll. leg.) [ CJBC] ; 2♂, same place, 1.VI.2021 (local coll. leg.) [ CJBC] . India. Arunachal Pradesh state. 1♀, Dalfa hills, 12.V.2003 (local coll. leg.) [ CJBC] . Indonesia. Bali province. 1♂, mount Batur, 23.XII.2013 (local coll. leg.) [ CJBC]. Baten province . 1♀ 1♂, Tjitorek , 2.V.2017 (local coll. leg.) [ CJBC] ; 1♀ 3♂, same place, 17.II.2022 (local coll. leg.) [ CJBC] . Central Sulawesi province. 2♀ 3♂, Peleng , 10.X.2017 (local coll. leg.) [ CJBC] . East Java province. 1♀ 1♂, Mont Argopuro , 11.I.2007 (local coll. leg.) [ CJBC] ; 1♀, same place, 9.IV.2011 (local coll. leg.) [ CJBC] ; 2♀ 2♂, same place, 26.XII.2013 (local coll. leg.) [ CJBC] . Nusa tenggara province. 1♀ 2♂, Lombok Isl., Sembalun Lawang , 14.XII.2017 (local coll. leg.) [ CJBC] . West Kalimantan. 2♂, Lumar , 28.V.2019 (local coll. leg.) [ CJBC] ; 4♀, same place, 21.IV.2022 (local coll. leg.) [ CJBC] ; 1♂, Mont Bawang , 14.VII.2021 (local coll. leg.) [ CJBC] . Laos. Vientiane province. 1♀, Vientiane, 15.III.2013 (local coll. leg.) [ CJBC] . Malaysia. Perak state. 1♂, Batang Padang, Bidor , 5.VII.2021 (local coll. leg.) [ CJBC] . Sabah state. 3♀, Mount Trusmadi , 12.IV.2013 (local coll. leg.) [ CJBC] . Sri Lanka. Eastern Province. 1♂, Thiriyai , 14.III.1976 (NMNH) . Thailand. Chiang Mai province. 1♀, Chiang Mai, 23.X.2016 (local coll. leg.) [ CJBC] ; 2♂, same place, IX.2018 (local coll. leg.) [ CJBC] ; 1♀, same place, 29.IX.2019 (local coll. leg.) [ CJBC] ; 2♂, same place, 12.VII.2022 (local coll. leg.) [ CJBC] ; 2♂, same place, 20.V.2023 (local coll. leg.) [ CJBC] ; 1♀, Choeng Doi , 9.X.2016 (local coll. leg.) [ CJBC] . Vietnam. Lai Châu province. 1♂, Lai Châu, 13.IV.2023 (local coll. leg.) [ CJBC]. Lâm Đ ồng province . 3♀ 1♂, Đambri , 2.II.2017 (local coll. leg.) [ CJBC] ; 1♀ 1♂, same place, 27.VII.2017 (local coll. leg.) [ CJBC]. Qu ảng Bình province . 2♀, Lâm Th ủy, 13.III.2023 (local coll. leg.) [ CJBC] ; 1♀, same place, 12.IV.2023 (local coll. leg.) [ CJBC] ; 1♀, same place, 21.IV.2023 (local coll. leg.) [ CJBC] . Thua Thien-Huê province. 3♂, Bach Ma , 10.VIII.2016 (local coll. leg.) [ CJBC] ; 1♀ 3♂, Hue , 21.VIII.2016 (local coll. leg.) [ CJBC] . Yên Bái province. 1♂, Yên Bái, 3.III.2022 (local coll. leg.) [ CJBC] ; 1♂, same place, 7.IV.2022 (local coll. leg.) [ CJBC] ; 1♂, same place, 15.IV.2022 (local coll. leg.) [ CJBC] ; 1♀, same place, 10.VII.2022 (local coll. leg.) [ CJBC] ; 2♂, same place, 13.VII.2022 (local coll. leg.) [ CJBC] ; 2♀, same place, 11.IX.2022 (local coll. leg.) [ CJBC] ; 1♂, same place, 4.V.2023 (local coll. leg.) [ CJBC] ; 7♂, same place, 18.IV.2024 (local coll. leg.) [ CJBC] ; 2♂, same place, 26.VII.2024 (local coll. leg.) [ CJBC] .

Diagnosis. Regiscolia azurea ( Christ, 1791) , comb. nov. differs from R. almoraensis ( Gupta & Jonathan, 2003) , comb. nov., R. bidens ( Linnaeus, 1767) , comb. nov., R. maculata ( Drury, 1773) , comb. nov., and R. rubida ( Gribodo, 1893) , comb. nov. in females by having the apical part of scutum with sparse punctures medially or without punctures, thus not forming a continuous band of punctures along the scutellum; the dorso-median area of propodeum 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 contrast, in R. almoraensis , R. bidens , R. maculata , and R. rubida , the apical part of scutum has a denser and uninterrupted band of punctures along the scutellum; the dorso-median area of propodeum is shorter, approximately equal to the length of the metanotum; and the basal part of T2 has sparser punctures similar to those of the median part. In males, R. azurea differs by having the area around and between the posterior ocelli with sparse and coarse punctures; and the length of the dorso-median area of propodeum significantly greater than that of the metanotum. In contrast, in R. almoraensis , R. bidens , R. maculata , and R. rubida , the area around and between the posterior ocelli has denser, tighter, finer, and agglomerated punctures; and the length of the dorso-median area of propodeum is nearly equal to that of the metanotum. In both sexes, the wings of R. azurea are entirely dark, whereas in R. almoraensis , R. bidens , R. maculata , and R. rubida they are bicolored, yellowish in the cells and dark elsewhere (in R. almoraensis the wings are almost entirely black).

R. azurea is easily distinguished from R. capitata ( Fabricius, 1804) , comb. nov. in both sexes by having the head with coarse and dense punctures; and the mesosoma entirely melanistic. In contrast, in R. capitata the head is almost entirely impunctate, with finer and sparser punctures; and the scutellum, metanotum, dorso-median area of propodeum are marked with yellow.

R. azurea is distinguished from R. alecto (Smith, 1858) , comb. nov. in female by having the pronotum with coarse and sparse punctures on the posterior half, and a large impunctate area in front of the pronotal excavation; the transition between the dorso-median and postero-median areas of propodeum forming a protruding ridge; and the basal part of T2 with dense and fine punctures forming a band interrupted medially. In contrast, in R. azurea the pronotum has denser, tighter, and finer punctures over its entire surface; the transition between the dorso-median and postero-median areas of propodeum forms an angle without a protrusion; and the basal part of T2 has denser and finer punctures forming a continuous band across its entire width. In males, R. azurea differs by having the basal part of T1 with an impunctate area at the level of the tubercle; and the metasoma with reddish apical pilosity. Whereas in R. alecto , the T1 has denser and finer punctures across its entire surface; and the metasomal pilosity is entirely black.

R. azurea is distinguished from R. fulvifrons ( Saussure, 1855) , comb. nov. in females, by having the apical margin of clypeus without distinct lateral lobes; and the disk of clypeus flat, impunctate, and weakly striated. In contrast, in R. fulvifrons , the apical margin of clypeus has two prominent lateral lobes; and the disk of clypeus is domed and fully reticulated. In males, R. azurea differs by having the T3 with a large impunctate central area. Whereas in R. fulvifrons , the T3 has a small impunctate area in the middle. In both sexes, R. azurea differs by having the metasoma with red pilosity apically. Whereas in R. fulvifrons , the metasoma has black pilosity, yellow in the colored areas.

R. azurea is distinguished from R. splendida ( Saussure, 1858) , comb. nov., by having the mesosoma entirely melanistic and the metasoma with red pilosity apically. Whereas in R. splendida , the mesosoma is generally marked with orange on pronotum, scutum and scutellum (except in ssp. floresensis ) and the metasoma has black pilosity, yellow in the colored areas.

Redescription. Female ( Fig. 60 View FIGURE 60 ). Length 28–40 mm.

Head ( Figs 60B, C View FIGURE 60 ). Mandibles short and thickened at the apex. Median mandibular tooth small and prominent, sometimes blunt. Anterior margin of clypeus rounded, without prominent lateral lobes; disk of clypeus not-domed and reticulated apically. Fissura frontalis strong ending at the frontal pit. Frons with numerous and sparse punctures. Vertex almost impunctate, except behind posterior ocelli with some sparse punctures.

Mesosoma ( Fig. 60D View FIGURE 60 ). Pronotum finely and densely punctate basally, more coarsely and sparsely punctate on the rest. Tegula with coarse punctures at base and along scutum; with micro-punctures often effaced on rest. Scutum coarsely and sparsely punctate basally; more sparsely laterally, except for lateral corners with very dense and fine punctures; largely impunctate medially and apically. Scutellum with fine and dense punctures along the scutum; sparser and coarser medially and apically, with distinctly impunctate surfaces. Metanotum finely and densely punctate with a small impunctate central band. Dorso-lateral area of propodeum densely and finely punctate, basally with an impunctate area (mirror). Dorso-median area of propodeum clearly longer than the length of the metanotum; punctures fine and dense over its entire surface. 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. 60E View FIGURE 60 ). T1 with a tubercle, either depressed in the middle or not; coarsely and sparsely punctate at base revealing a small impunctate area medially; very densely and finely punctate laterally and apically. T2 densely and finely punctate basally, punctation interrupted in the middle; very sparsely punctate medially; very finely and densely punctate apically. T3 almost entirely impunctate basally and medially; densely and finely punctate apically. T4 to T5 with similar punctation, but the central impunctate area gradually reduced. S1 densely and finely punctate medially and apically. S2 with weak tubercle; with fine and dense punctures at base; with sparse and coarse punctures in center; with dense and fine punctures on sides apically, sparsely punctate in the middle apically. S3 to S5 with similar punctures.

Coloration ( Figs 60 View FIGURE 60 , 61 View FIGURE 61 ). Frons, vertex and tempora orange-reddish (generally ssp. hindostana and michaae) or yellowish orange (generally ssp. azurea ). Mesosoma black with black pilosity, sometimes with red spots on pronotum. In the subspecies hindostana ( Fig. 61 View FIGURE 61 ), metasoma black with a pair of large, rounded reddish spots on T3, and most of T4 and T5 also exhibit reddish coloration. Metasomal pilosity black, except for reddish pilosity on T3–T6 (occasionally T2 with a few reddish setae, or the pygidium with black pilosity). In subspecies michaae, as in hindostana, but clypeus partially or entirely orange-reddish. In subspecies azurea ( Fig. 60 View FIGURE 60 ), metasoma black, T4 and T5 often marked with small orange-reddish spots. Metasomal pilosity black, except for reddish pilosity on T4–T6 (sometimes also apically on T3). Some specimens from Malaysia exhibit entirely black metasomal pilosity or almost so, in contrast to the typically reddish pilosity observed in azurea . (see remarks).

Male ( Fig. 62 View FIGURE 62 ). Length 22–33 mm.

Head ( Figs 62B, C View FIGURE 62 ). Inner margin of mandible with three distinct tooth (third apical tooth sometimes blunt and barely visible). Disk of clypeus with few coarse punctures medially, denser and finer punctures laterally. Fissura frontalis indistinct, not reaching the median ocelli. Frontal cross-furrow barely visible, its extension reaching the upper lobe of the eye. Frontal spatium with dense, tight and fine punctures. Anterior ocelli shallowly depressed, larger than posterior ocelli. Frons with dense and coarse punctures, except in front of anterior ocellus with an impunctate area. Vertex behind the ocelli with sparse and fine punctures, dense punctures near the occipital carina.

Mesosoma ( Fig. 62D View FIGURE 62 ). Pronotum with dense and fine punctures over its entire surface, inter-points spaces almost coalescent. Tegula with few coarse punctures anteriorly, with shallow and dense micropunctures on the rest. Scutum with fine and dense punctures, denser and finer basally, coarser laterally. Scutellum with fine and dense punctures basally, sparser and coarser medially and mostly impunctate apically. Metanotum with fine and dense punctures laterally, impunctate line in the middle. Dorso-lateral area of propodeum with an impunctate area (mirror); with fine and dense punctures. Dorso-median area of propodeum with coarse and dense punctures over its entire surface, inter-point spaces smaller than the diameter of a punctures. Anterior margin of marginal cell longer than posterior margin. Forewing with three submarginal cells.

Metasoma ( Fig. 62E View FIGURE 62 ). T1 with a strong tubercle; very densely and finely punctate laterally and apically; more coarsely and sparsely basally and medially. T2 with fine and dense punctures; more denser and finer punctures basally and apically, at these levels pilosity forming a large band of punctures. T3 impunctate medially, with very dense and fine punctures apically, less dense and sparse basally. T4-T6 similarly punctate, but the central impunctate area smaller. S1 raised in the middle, with dense punctures medially, with few punctures apically. S2 with dense and fine punctures anteriorly, sparser laterally, mostly impunctate medially and apically. S3 with very sparse punctures medially, more denser laterally. S4-S6 similarly punctate.

Coloration ( Figs 62–64 View FIGURE 62 View FIGURE 63 View FIGURE 64 ). Head black, sometimes with orange or reddish spot on ocular sinus and behind eyes. Mesosoma black. In the subspecies hindostana ( Fig. 63 View FIGURE 63 ) and azurea ( Fig. 62 View FIGURE 62 ), metasoma black with a pair of large, rounded reddish spots on T3 (sometimes absent in ssp. azurea ), and most of T4-T6 and often T7 also exhibit reddish coloration. Metasomal pilosity black, except for reddish pilosity on T3–T7 (occasionally T2 shows a few reddish setae apically in ssp. hindostana , or sometimes black pilosity on T 3 in ssp. azurea ). In michaae ( Fig. 64 View FIGURE 64 ), as in hindostana, but the head and clypeus mostly orange-reddish.

Genitalia. See Fig. 31C View FIGURE 31 .

Distribution. Bangladesh, Bhutan, Cambodia, China ( Anhui, Fujian, Guangdong, Hainan, Hong Kong, Guangxi, Guizhou, Hunan, Yunnan), India (Andaman Islands, Arunachal Pradesh, Assam, Chhattisgarh, Himachal Pradesh, Karnataka, Kerala, Maharashtra, Manipur, Meghalaya, Mizoram, Nagaland, Odisha, Sikkim, Tamil Nadu, Tripura, Uttarakhand, West Bengal), Indonesia ( Bali, Bangka Belitung Islands, Baten, Central Java, Central Sulawesi, East Java, Lampung, Jakarta, West Java, West Kalimantan, West Nusa Tenggara, West Sumatra), Laos, Malaysia ( Perak, Sabah, Selangor), Myanmar, Nepal, Singapore, Sri Lanka, Thailand, Vietnam ( Paiva 1907; Vitalis de Salvaza 1919; Micha 1927; Betrem 1928, 1941, 1947; Krombein 1978; Gupta & Jonathan 2003; Kumar, 2009; Kumar & Rajmohana 2017; Nidup et al. 2017; Jadhav & Gaikwad 2019; Liu et al. 2021a; Taylor & Barthélémy 2021) ( Fig. 65 View FIGURE 65 ).

Biology. The only known hosts are Oryctes rhinoceros ( Linnaeus, 1758) and Xylotrupes gideon ( Linnaeus, 1767) ( Leefmans 1915, 1921; Betrem 1928). In Cambodia, groups of males were observed flying around bamboo clumps in semi-urban areas (FC pers. obs.). A female was observed walking and searching on the ground in a papaya field (FC pers. obs.).

Remarks. To date, six subspecies have been recognized ( Gupta & Jonathan 2003; Osten 2005b): ssp. azurea ( Christ, 1791) from Borneo to Java, Sumatra, Malakka (city), Southern China, ssp. christiana ( Guiglia & Betrem, 1958) from Myanmar, Bangladesh and northern India, ssp. cochinensis ( Betrem, 1928) from Vietnam, ssp. hindostana ( Micha, 1927) from southern India, michaae ( Betrem, 1928) from Sri Lanka and ssp. siamensis ( Betrem, 1928) from Thailand. Betrem & Bradley (1964) listed only five, but the ssp. michaae Betrem, 1928 from Sri Lanka seems to have been overlooked. These subspecies differ in the absence or presence of maculation on certain parts of the body, and in the extent of reddish pilosity on the metasoma. Only the subspecies ssp. azurea, ssp. hindostana and ssp. michaae are considered valid here.

R. azurea cochinensis is known only from two females in southern Vietnam (Cap St. Jacques, now Vũng Tàu). Betrem (1928) gave only a brief description: “ like subspecies azurea , but abdomen entirely black except for two spots on T3 and T4. Abdomen entirely black, with only a few red setae behind the spots ”. R. azurea hindostana differs from R. azurea christiana in females only by the presence of black setae on the pygidium, whereas they are reddish in ssp. christiana , and in males by subtle variations in coloration on the head and pilosity ( Gupta & Jonathan 2003; Nidup et al. 2017). All the female specimens examined in this study from Cambodia, China, India, Thailand and Vietnam belong to either ssp. christiana or ssp. hindostana , and many females show a mixture of reddish and black setae on the pygidium. It is therefore difficult to know to which subspecies these females should be attributed. The presence of ssp. azurea in China ( Gupta & Jonathan 2003) seems doubtful and should be attributed to ssp. christiana ( Betrem 1941, 1947; Guiglia & Betrem 1958). Examination of a large sample of R. azurea specimens from throughout its range revealed three distinct groups corresponding to well-defined subspecies. The first subspecies, R. azurea michaae ( Fig. 64 View FIGURE 64 ), is strictly localized to Sri Lanka; the second, R. azurea azurea ( Figs 60 View FIGURE 60 , 62 View FIGURE 62 ), is distributed in insular and peninsular Southeast Asia, mainly in Indonesia, Peninsular Malaysia and East Malaysia; and the third, R. azurea hindostana ( Figs 61 View FIGURE 61 , 63 View FIGURE 63 ), occupies continental Asia. R. azurea cochinensis ( Betrem, 1928) , syn. nov. and R. azurea christiana ( Guiglia & Betrem, 1958) , syn. nov. are therefore considered junior subjective synonyms of R. azurea hindostana ( Micha, 1927) .

A subspecies known as siamensis Betrem, 1928 is found locally in Southeast Asia (part of Thailand, Cambodia), and concerns female specimens with the pronotum more or less marked with red ( Betrem, 1928). As this taxon is restricted to females and a limited geographical area, it is preferable to consider the ssp. siamensis as a simple local form rather than as a distinct taxonomic entity.

Examination of the types male and female of Scolia ( Triscolia) hageni Betrem, 1928 , syn. nov. ( Figs 67 View FIGURE 67 , 68 View FIGURE 68 ) reveals that this species is a synonym of R. azurea . The female has an abnormally black head ( Figs 67B, C View FIGURE 67 ), but displays all the criteria of R. azurea . A similar variation in the yellow coloration of the head in females is observed in R. maculata , with specimens ranging from entirely yellow to almost entirely black.

In Malaysia, particularly in the vicinity of Malacca (city) and on Borneo island, some female specimens of R. azurea exhibit an entirely black pilosity, or occasionally only the pygidium or T5 is partially tinged with reddish. These fully melanistic specimens, in which only the head has yellow-orange markings, display a general habitus very similar to R. alecto and can easily be misidentified (see key to Regiscolia species). Betrem (1928, 1941) mentioned the locality of Malacca for two melanistic female specimens, which he associated though with some doubt, with Scolia ( Triscolia) alecto , hesitating between that and S. ( T.) azurea . In doing so, he synonymized Triscolia vespillo , which shows the same habitus and also comes from Malacca, with S. ( Triscolia) alecto (as Micha 1927 had before him). Nevertheless, examination of the holotype female of Triscolia vespillo reveals a problematic specimen with features intermediate between azurea and alecto ( Fig. 66 View FIGURE 66 ). However, the pronotum shows coarse and sparse punctures in its posterior half, leaving a large impunctate area characteristic of azurea . To date, only melanistic females have been found, and the male of alecto has never been reported from Malacca, pending the examination of a larger number of melanistic specimens in Malaysia, we therefore consider, contrary to Betrem (1928), that Triscolia vespillo should be treated as a junior synonym of azurea , not alecto . The two specimens mentioned by Betrem (1928) more plausibly refer to melanistic individuals of R. azurea .

The status of Scolia rubiginosa Fabricius, 1793 has changed considerably over time and remains uncertain today. Dalla-Torre (1897) first linked this taxon with some doubt to Sphex azurea Christ, 1791 , then later actually considered it to be a subspecies of Scolia ( Triscolia) azurea ( Micha 1927; Betrem 1927b). The type specimen of S. rubiginosa is a male that cannot now be found ( Guiglia & Betrem 1958; Bradley & Betrem 1964). Consequently, the drawing by Coquebert (1801) was designated by Bradley & Betrem (1964) as the lectotype. Fabricius (1793) was very vague as to the origin of the type specimen: “ in India Orientali ” referred at that time more or less to South and Southeast Asia (from India to Indonesia). Based on their interpretation of Fabricius’ description, Micha (1927) and Betrem (1928, 1941) applied it mainly to specimens from continental Asia. Guiglia & Betrem (1958) later synonymized this name with the nominal subspecies on the basis of a reinterpretation of Fabricius’ description concerning the coloration of the pilosity fringe on the T3 (a criterion used to distinguish male subspecies according to Betrem (1928)). They thus erected a new subspecies, christiana Guiglia & Betrem, 1958, for specimens formerly known as S. rubiginosa (mainly from continental Asia). Taylor & Barthélémy (2021) rightly expressed doubts about the interpretation of Fabricius’ description of S. rubiginosa by the various authors, especially as the precise type locality of this specimen is unknown. They declined to comment on the subspecies present in Hong Kong, pending clarification of the status of R. azurea subspecies. In fact, the pilosity coloration criteria proposed by Betrem (1928) to determine male subspecies of R. azurea are not reliable, as pilosity coloration is variable, particularly in males within the same population. Furthermore, as the type specimen of S. rubiginosa has disappeared and the type locality is not known with certainty, interpretation on the basis of a drawing is also prohibited. S colia rubiginosa Fabricius, 1793 , syn. nov. is therefore retained here as a synonym of R. azurea .

Scolia ( Scolia) magnifica Saussure, 1859 has at times been considered as a variety or form of several subspecies of R. azurea ( Betrem 1928; Betrem & Bradley 1964; Osten 2005b). According to Betrem (1928), this taxon can only be distinguished by having the T1 and T2 with black setae mixed with red setae, whereas in other taxa (subspecies, varieties, and forms), they are covered with entirely black pilosity. The examination of numerous azurea specimens in this study reveals that pilosity coloration is sometimes variable in both males and females. A few specimens show an atypical pilosity on T1 and T2, with black mixed with reddish setae, thus presenting the same variation described by Betrem (1928) for the taxon magnifica . These specimens fall within the extreme range of pilosity coloration observed in azurea and do not warrant a distinct taxonomic status. Consequently, Scolia ( Scolia) magnifica Saussure, 1859 , syn. nov. is here placed in synonymy with R. azurea .

Triscolia democratica Micha, 1927 (with unknown syntype locality) was at times treated as a mere variety, form, or subspecies of R. azurea ( Betrem 1928, 1947; Betrem & Bradley 1964; Bradley 1972; Osten 2005). According to Betrem (1928), it differs from other azurea taxa only by having the T4 and most sternites with entirely black pilosity, whereas in other azurea taxa, the T4 has at least some red pilosity on the sides. Later, Betrem (1947) listed the subspecies of the taxon azurea and, without justification, attributed the locality “ Sri Lanka ” to the taxon democratica . In the same publication, the taxon michaae Betrem, 1928, which he had described a few years earlier and which is normally endemic to Sri Lanka, is absent. We can only speculate about this: either Betrem synonymized the taxon democratica with the taxon michaae without explanation, or it is a lapsus calam i for michae. However, neither Micha’s (1927) description nor Betrem’s (1928) description of T. democratica matches that of that of the specimen from Sri Lanka (head largely marked with reddish yellow, including the clypeus in both sexes). This association therefore appears to be an error. For the same reasons as for Scolia ( Scolia) magnifica Saussure, 1859 , we here consider Triscolia democratica Micha, 1927 , syn. nov. to be a junior synonym of R. azurea .

A key to the three subspecies considered valid here is proposed. Some rare specimens show an atypical T1 punctation or metasoma coloration, particularly in areas where the ranges of ssp. azurea and ssp. hindostana overlap.

Key to subspecies of Regiscolia azurea

Females

1. Head except the apex of clypeus reddish yellow. Sri Lanka................................... ssp. michaae (Betrem)

- Frons and vertex reddish yellow or reddish, clypeus black ( Figs 60 View FIGURE 60 , 61 View FIGURE 61 ). From India to Indonesia, excluding Sri Lanka..... 2

2. T1 with the depression behind the tubercle generally shallow and more elongated, more or less distinct; usually with a large impunctate area behind the tubercle, sparse and coarse punctures ( Fig. 60 View FIGURE 60 ). Head yellowish. T3 often entirely black. Insular and peninsular Southeast Asia................................................................ spp. azurea (Christ)

- T1 with the depression behind the tubercle generally deep, more or less rounded; usually with a small impunctate area behind the tubercle, sparse but denser and finer punctures ( Fig. 61 View FIGURE 61 ). Head reddish-yellow. T3 with two large orange spots, sometimes fused (rarely missing). Continental Asia................................................ spp. hindostana (Micha)

Males

1. Head and clypeus mostly reddish-yellow ( Fig. 64 View FIGURE 64 ). Sri Lanka................................. ssp. michaae (Betrem)

- Head and clypeus mostly black ( Figs 62 View FIGURE 62 , 63 View FIGURE 63 ). From India to Indonesia, excluding Sri Lanka.......................... 2

2. T1 generally with a large impunctate area behind the tubercle, with coarse and sparse punctures. Disk of T 2 in the middle usually with very sparse punctures, clearly more spaced than punctures on the sides ( Fig. 62 View FIGURE 62 ). Insular and peninsular Southeast Asia................................................................................. spp. azurea (Christ)

- T1 generally with a small impunctate area behind the tubercle, sparse but denser and finer punctures. Disk of T 2 in the middle usually with slightly sparser punctures than punctures on the sides ( Fig. 63 View FIGURE 63 ). Continental Asia..... spp. hindostana (Micha)