Megascolia procer ( Illiger, 1802 )

Megascolia procer ( Illiger, 1802)

( Figs 19–21 View FIGURE 19 View FIGURE 20 View FIGURE 21 , 23 View FIGURE 23 , 24 View FIGURE 24 )

Scolia procer Illiger, 1802: 196 ( holotype, ♀, Sumatra, deposited in NMB, Germany, not examined); Fabricius 1804: 238;

Lepeltier 1825: 393; Burmeister 1854: 19; Tullgren 1904: 465; Richards 1919: 128. Scolia ( Scolia) procer : Guérin-Méneville 1830: 247. Scolia procera : Lepeletier 1845: 519; Smith 1855: 111; Smith 1858a: 89; Cameron 1901: 18. Scolia ( Triscolia) procer : Saussure & Sichel 1864: 43; Smith 1871: 343; Cameron 1892: 100; Bingham 1897: 75; Bingham

1906: 37; Rohwer 1921: 77. Scolia ( Triscolia) procera : Magretti 1892: 236; Mantero 1903: 33. Triscolia procer : Gribodo 1893: 152. Triscolia procer forma scapulata Gribodo, 1893: 163 (synonymised with procer procer by Micha 1927: 104 and later again by

Betrem & Bradley 1964: 441). Triscolia procer var. sumatrensis Gribodo, 1893: 164 . Syn. nov. Triscolia procer var. bimaculata Gribodo, 1893: 164 . Scolia ( Scolia , Triscolia ) procera : Dalla-Torre 1897: 174. Scolia ( Scolia , Triscolia ) procera var. bimaculata : Dalla-Torre 1897: 174. Scolia ( Scolia , Triscolia ) procera var. scapulata : Dalla-Torre 1897: 174. Scolia ( Scolia , Triscolia ) procera var. sumatrensis : Dalla-Torre 1897: 174. Scolia ( Triscolia) procera forma scapulata Mantero, 1903: 33. Scolia ( Triscolia) procera var. nigriventris Mantero, 1903: 33 . Triscolia procera : Cameron 1905: 52. Scolia ( Triscolia) procer var. bimaculata : Rohwer 1921: 77. Triscolia procer : Micha 1927: 104. T riscolia procer procer : Micha 1927: 104–108. Triscolia procer form luteifrons Micha, 1927: 108 (synonymised with procer procer by Betrem & Bradley 1964: 441). Scolia ( Triscolia sectio Megascolia ) procer : Betrem 1928: 240. Scolia ( Triscolia sectio Megascolia ) procer var. sarawakensis Betrem, 1928: 241 . Syn. nov. Scolia ( Triscolia sectio Megascolia ) procer var. bimaculata : Betrem 1928: 242. Scolia ( Triscolia sectio Megascolia ) procer var. nigriventris : Betrem 1928: 242. Scolia ( Triscolia sectio Megascolia ) procer procer : Betrem 1933: 255. Scolia ( Triscolia sectio Megascolia ) procer luteifrons: Betrem 1933: 256. Megascolia ( Megascolia) procer procer : Betrem & Bradley 1964: 438, 441; Betrem in Bradley 1972: 12; Bradley 1973a: 286;

Gupta & Jonathan 2003: 135; Kumar & Rajmohana 2017: 5; Ming & Soh 2020: 110. Megascolia ( Megascolia) procer procer var. sumatrensis : Betrem & Bradley 1964: 441. Megascolia ( Megascolia) procer procer var. sarawakensis : Betrem & Bradley 1964: 441; Osten 2005b: 41. Megascolia ( Megascolia) procer javanensis Betrem & Bradley, 1964: 439 , 441; Bradley & Betrem 1966: 81; Gupta & Jonathan

2003: 135; Osten 2005b: 41; Ming & Soh 2020: 110. Syn. nov. Megascolia ( Megascolia) procer nigriventris : Betrem & Bradley 1964: 439, 441; Bradley 1973a: 285; Gupta & Jonathan 2003:

135; Osten 2005b: 41. Megascolia ( Megascolia) procer bimaculata : Betrem & Bradley 1964: 440, 441; Petersen 1970: 64–65; Bradley 1973a: 278;

Gupta & Jonathan 2003: 135; Osten 2005b: 28, 41. Megascolia ( Megascolia) procer procer form sumatrensis: Bradley 1973a: 287; Osten 2005b: 41. Megascolia procer : Argaman 1996: 199. Megascolia ( Megascolia) procer : Osten 2001: 439; Gupta & Jonathan 2003: 135; Kumar & Rajmohana 2017: 5; Ming & Soh

2020: 110; Golfetti et al. 2025: 3, 10. Megascolia ( Megascolia) procer procer form luteifrons: Osten 2005b: 41. Megascolia cf. procer : Khouri et al. 2022: 24–29, 31–34, 36–38.

Type material examined. Holotype of Scolia ( Triscolia) procera var. nigriventris (MSNG) , ♀, ‘‘Is. Nias / Lelemboli / Agosto 1886 / Modigliani’’, ‘‘ HOLOTYPE / nigriventris Man. / Teste, 1929 / J.C. Bradley’’( Fig. 25 View FIGURE 25 ).

Other material examined. China. Yunnan province. 1♀, Mangyun , 4.X.2020 (local coll. leg.) [ CJBC] . Indonesia. Kalimantan Barat province. 2♀ 2♂, Loemar , 15.IX.2016 (local coll. leg.) [ CJBC] ; 2♂, same place, 6.VI.2019 (local coll. leg.) [ CJBC] . West Kalimantan province. 4♂, Mount Bawang , VII.2022 (local coll. leg.) [ CJBC] . Laos. Oudoxmay province . 1♀, Namkat Yorla Pa , 3.V.2019 ( S. Collard leg.) [ NPYPBM] . Malaysia. Perak state. 1♀ 2♂, Perak, 2.XI.2005 (local coll. leg.) [ CJBC] ; 2♂, same place, 21.IV.2013 (local coll. leg.) [ CJBC] . Selangor state. 1♀, Kuala Selangor, 30.I.2013 (local coll. leg.) [ CJBC]. The Philippines. Palawan province. 1♀, Roxas , 30.XI.2020 (local coll. leg.) [ CJBC] ; 1♀, same place, XII.2022 (local coll. leg.) [ CJBC] ; 1♂ 1♀, same place, 7.XI.2023 (local coll. leg.) [ CFC] ; 1♂, same place, 22.IX.2023 (local coll. leg.) [ CJBC] ; 2♀, same place, 4.II.2021 (local coll. leg.) [ CJBC] ; 3♀ 1♂, same place, 13.X.2023 (local coll. leg.) [ CJBC] ; 4♀, same place, 1.XI.2023 (local coll. leg.) [ CJBC] ; 1♀, same place, VIII.2022 (local coll. leg.) [ CJBC] ; 1♀, same place, 25.XII.2023 (local coll. leg.) [ CJBC] ; 2♀ 1♂, same place, 25.I.2024 (local coll. leg.) [ CJBC] ; 1♀ 1♂, same place, 8.II.2024 (local coll. leg.) [ CJBC] ; 2♀, same place, 25.IV.2024 (local coll. leg.) [ CJBC] . Singapore. North region . 1♂, Sembawang , 29.I.2021 (local coll. leg.) [ CJBC] ; 1♀, same place, 31.VII.2022 (local coll. leg.) [ CJBC] . Thailand. Nakhon Si Thammarat province. 1♂, Khao Phra , 13.IX.2017 (local coll. leg.) [ CJBC] . Vietnam. Ha Giang province. 1♀, Ha Giang, VII.2023 (local coll. leg.) [ CFC] . Lam Dong province. 1♂, Dambri , III.2023 (local coll. leg.) [ CJBC]. Th ừa Thiên Hu ế province . 1♀ 1♂, Bạch Mã mountain , 4.VIII.2016 (local coll. leg.) [ CJBC] ; 1♀, same place, 12.III.2017 (local coll. leg.) [ CJBC] .

Diagnosis. Megascolia procer ( Illiger, 1802) differs from M. velutina ( Saussure, 1859) and M. scutellaris ( Gribodo, 1893) in both sexes by having the pronotum largely impunctate medially and apically; and the T1 with a broad impunctate basal area. In contrast, in M. scutellaris and M. velutina , the pronotum is more densely punctate throughout, without distinct impunctate area (sometimes with a small scattered area apically); and the T1 is largely punctate, basally with sparser and coarser punctures. In addition, M. procer is more largely marked with yellow (pronotum, scutellum, metanotum, and generally on T1 and T3). Whereas M. velutina and M. scutellaris are entirely black, sometimes with yellow markings mainly on the scutellum and metanotum.

Redescription. Female ( Fig. 19 View FIGURE 19 ). Length 39–57 mm.

Head ( Figs 19B, C View FIGURE 19 ). Disk of clypeus domed in the middle; coarsely and sparsely punctate medially, more finely and densely punctate on the rest. Fissura frontalis distinct, reaching the level of the lamina frontalis and ending in a pronounced pit. Frons almost impunctate, with only the ocellar triangle bearing a few punctures. Vertex almost entirely impunctate, with only a few scattered punctures.

Mesosoma ( Fig. 19D View FIGURE 19 ). Pronotum finely and densely punctate basally, almost impunctate on the rest. Pronotum with a distinct tubercle, often blunt but protruding in front of the tegula. Tegula largely impunctate; with fine and dense punctures basally; along scutum with scattered and coarse punctures. Scutum finely and densely punctate basally; more sparsely punctate laterally; largely impunctate medially and apically. Scutellum finely and densely punctate basally, much more sparsely punctate apically. Metanotum finely and sparsely punctate; almost impunctate apically. Dorso-lateral area of propodeum densely and finely punctate, especially the region near the lateral area of the propodeum, anteriorly with an impunctate surface (mirror). Dorso-median area of propodeum densely and finely punctate; medially more sparsely punctate revealing an impunctate surface. Mesopleura densely and finely punctate all over. Anterior part of upper plate of metapleura densely and finely punctate on anterior half, more sparsely punctate on posterior half. Lower plate of metapleura densely and finely punctate over almost its entire surface, except for a small 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 rib and the radial area. Anterior margin of marginal cell almost twice as long as posterior margin.

Metasoma ( Fig. 19E View FIGURE 19 ). T1 with strong, bifid tubercle, largely impunctate; impunctate basally; with dense and fine punctures laterally and apically. T2 impunctate basally; densely and finely punctate laterally; very sparsely punctate medially; finely and densely punctate apically. T3 impunctate basally, except for fine and dense punctures along T2; densely and finely punctate laterally and apically. 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 coarse and sparse punctures in the middle apically. S3 to S5 with similar punctation.

Coloration ( Figs 19 View FIGURE 19 , 20 View FIGURE 20 , 25 View FIGURE 25 ). In the nominotypical subspecies ( Fig. 19 View FIGURE 19 ), head black, marked with yellow on frons and vertex with only the ocellar region black. Antennae black. Mesosoma black, marked with yellow on pronotum, scutellum and metanotum. Metasoma black, yellow markings on T1 and two yellow spots on T3. Legs black. Wings entirely dark, with blue/greenish highlights. Pilosity black; yellow on the colored areas. In the subspecies bimaculata ( Fig. 20 View FIGURE 20 ), head with the same coloration as the nominotypical subspecies, mesosoma black, sometimes faintly marked with yellow on the scutellum, metasoma black, with two small yellow spots on T3. In the subspecies nigriventris ( Fig. 25 View FIGURE 25 ), head with the same coloration as the nominotypical subspecies, mesosoma and metasoma entirely black.

Male ( Fig. 21 View FIGURE 21 ). Length 27–40 mm.

Head ( Figs 21B, C View FIGURE 21 ). Inner margin of mandible with three distinct tooth (third apical tooth sometimes blunt and barely visible). Disk of clypeus densely punctate, largely impunctate medially. Antennal segment XIII about 2.4 times longer than wide. Distinct fissura frontalis reaching the median ocelli and passing through the frons fossa. Cross-furrow distinct, its extension ending in the ocular sinus. Frontal spatium short, densely and finely punctate; pilosity fine and dense. Median ocelli slightly depressed, larger than posterior ocelli. Frons sparsely and finely punctate, except for an impunctate band in front of the median ocelli. Vertex densely and finely punctate. Vertex sparsely and coarsely punctate.

Mesosoma ( Fig. 21D View FIGURE 21 ). Pronotum finely and sparsely punctate, inter-point space sometimes very large, revealing an impunctate area. Tegula with few coarse punctures in its apical part and along the scutum, the rest with fine and dense punctures. Scutum with dense, fine punctures, very sparse medially, revealing two impunctate bands. Scutellum with fine and dense punctures, more sparser medially; inter-point space medially often well over twice the diameter of a point. Metanotum finely and sparsely punctate, with large impunctate area medially. Basal part of dorso-lateral area of propodeum with impunctate area (mirror); apical part finely and densely punctate, more finely punctate under the mirror. Dorso-median area of propodeum densely punctate, inter-point space approximately equal to the diameter of a puncture.

Metasoma ( Fig. 21E View FIGURE 21 ). T1 with a strong tubercle; very densely and finely punctate laterally and apically; largely impunctate basally and medially. T2 very finely and densely punctate all over. 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 ( Figs 21–23 View FIGURE 21 View FIGURE 22 View FIGURE 23 ). In the nominotypical subspecies ( Figs 21 View FIGURE 21 , 23 View FIGURE 23 ), head almost entirely yellow, only the mandibles in part, ocellar region, frontal spatium, tempora behind the eyes and occiput black. Antennae black, last antennal segments often marked with orange-brown underneath. Mesosoma black, with yellow markings on pronotum, scutellum and metanotum. Metasoma black, with a yellow spot on T1 and two yellow spots on the basal part of T3 (sometimes absent). Legs black. Wings entirely dark, with blue/purple reflections. Pilosity entirely black; yellow on the colored areas. In the subspecies bimaculata ( Fig. 22 View FIGURE 22 ), clypeus largely black, mesosoma black, sometimes with faint yellow markings on the pronotum and/or scutellum, and T1 entirely black. The male of subspecies nigriventris is unknown.

Genitalia. See Fig. 15A View FIGURE 15 .

Distribution. China ( Yunnan; new record), India ( Arunachal Pradesh), Indonesia ( Bengkulu, East Kalimantan, East Java, Jakarta, Jambi, Lampung, North Sumatra, South Sumatra, West Java, West Kalimantan, West Sumatra), Laos (Oudoxmay; new record), Malaysia ( Pahang, Perak, Phatthalung, Sabah, Sarawak, Selangor), Myanmar, The Philippines ( Palawan), Singapore, Thailand, Vietnam ( Micha 1927; Betrem 1928; Betrem & Bradley 1964; Petersen 1970; Gupta & Jonathan 2003) ( Fig. 24 View FIGURE 24 ).

Biology. The only known hosts are Oryctes rhinoceros ( Linnaeus, 1758) ( Richards 1919) and Chalcosoma atlas Linnaeus, 1758 ( Piek 1986a). In Singapore, many specimens were observed foraging on the coastal and native shrubs Premna serratifolia ( Lamiaceae ) ( Ming & Soh 2020).

Remarks. Numerous subspecies, varieties and forms have been described: ssp. bimaculata ( Gribodo, 1893) from The Philippines ( Palawan), ssp. javanensis Betrem, 1964 from Indonesia ( Java), spp. nigriventris ( Mantero, 1903) from Indonesia (Nias), spp. procer ( Illiger, 1802) from Myanmar, Thailand, Malaysia, Singapore and Indonesia (Borneo, Java, Sumatra), spp. procer form sumatrensis ( Gribodo, 1893) from Indonesia ( Sumatra) and ssp. procer variety sarawakensis Betrem, 1928 from Malaysia ( Sarawak) ( Betrem & Bradley 1964; Osten 2005b). As coloration is highly variable, the validity of these taxa remains doubtful, as they are mainly characterized by the extent of yellow coloration or its absence on the various tagmas. All forms and varieties are considered invalid here, and only three subspecies are considered valid, for the reasons set out below.

M. procer procer form sumatrensis syn. nov., known only from the male, differs from the typical form only by the third metasomal segment, which is black and immaculate ( Betrem 1928). In this study, specimens belonging to the sumatrensis form and others belonging to the typical form were examined from the same locality. We therefore consider this form as invalid.

M. procer procer var. sarawakensis syn. nov. is known only from the male, whose coloration is similar to the nominotypical subspecies, but with additional yellow maculae on parts of the scutum, dorso-lateral and dorso-median areas of propodeum ( Betrem 1928). For the same reasons as the previous form, this variety must be considered as a mere intraspecific variation.

M. procer javanensis syn. nov. is recognizable only in female and differs from the nominotypical subspecies by the ocellar triangle and the vertex, which are more extensively marked with black ( Betrem & Bradley 1964). This subspecies is based solely on the female and on head coloration, which is highly variable. We therefore synonymize this subspecies with the nominotypical subspecies.

Only the ssp. bimaculata ( Figs 20 View FIGURE 20 , 22 View FIGURE 22 ), ssp. nigriventris ( Fig. 25 View FIGURE 25 ), and ssp. procer ( Figs 19 View FIGURE 19 , 21 View FIGURE 21 , 23 View FIGURE 23 ) are considered valid in this study. M. procer bimaculata ( Gribodo, 1893) was previously known only from one female and one male from Palawan (The Philippines) ( Gribodo 1893; Betrem 1928; Bradley & Betrem 1964; Petersen 1970). The female differs from the nominal form by having the mesosoma entirely black, although the scutellum may sometimes be faintly marked with yellow, the first metasomal segment is completely black, and the pronotal tubercles are more blunt. The male differs by having the clypeus largely black, a black mesosoma (sometimes with faint yellow markings on the pronotum and/or scutellum), and the first metasomal segment entirely black. The examination of twenty-four specimens from Palawan shows that this coloration appears to be stable. Furthermore, no specimens of the nominotypical subspecies are known from The Philippines. This subspecies is therefore considered valid.

M. procer nigriventris ( Mantero, 1903) , known only from a single female from Nias island ( Indonesia) ( Fig. 25 View FIGURE 25 ), differs from the typical form by having the mesosoma and metasoma entirely black, with only the head marked with yellow ( Betrem 1928). Mantero does not provide a proper description, merely stating: “L’esemplare ( ♀) di Nias, lungo circa 43 milimetri credo sia riferibile alla var. 2 di Gribodo (L. cit. p. 156) che si potrebbe chiamare nigriventris.” The “variety 2” of procer mentioned by Mantero is likely a female of M. velutina , entirely black with yellow markings on the frons. Betrem & Bradley (1964) treated nigriventris as a subspecies. Examination of the holotype confirms that this taxon clearly belongs to M. procer , particularly based on the characteristic punctation of the pronotum and scutellum. The specimen exhibits a completely black mesosoma and metasoma, but the areas that are usually yellow are brownish. Further examination of additional specimens is necessary to clarify the status of this taxon; this atypical coloration may be the result of intraspecific variability.

Key to subspecies of Megascolia procer

(unknown in M. procer nigriventris )

1. Mesosoma View in CoL entirely melanistic (exceptionally small yellow spot on scutellum); T1 entirely black ( Figs 20 View FIGURE 20 , 22 View FIGURE 22 , 25 View FIGURE 25 ) …2

- Mesosoma largely marked with yellow on pronotum, scutellum and metanotum; T1 largely marked with yellow ( Figs 19 View FIGURE 19 , 21 View FIGURE 21 , 23 View FIGURE 23 ) … ssp. procer (Illiger)

2. Metasoma entirely melanistic ( Fig. 25 View FIGURE 25 ). Nias island... ssp. nigriventris (Mantero)

- Metasoma with small lateral yellow spots on T3 ( Figs 20 View FIGURE 20 , 22 View FIGURE 22 ). The Philippines... ssp. bimaculata (Gribodo)