EUPELMUS (MACRONEURA)

Fusu, Lucian, 2017, An integrative taxonomic study of European Eupelmus (Macroneura) (Hymenoptera: Chalcidoidea: Eupelmidae), with a molecular and cytogenetic analysis of Eupelmus (Macroneura) vesicularis: several species hiding under one name for 240 years, Zoological Journal of the Linnean Society 181 (3), pp. 519-603 : 569-574

EUPELMUS (MACRONEURA)

VESICULARIS (RETZIUS)

FIGS 45 , 63 , 73, 74, 83 , 97, 101, 102 (♀), 62, 75, 95 ( CLADE B ♀) , 19, 27, 46, 55, 105, 123 (♂)

Ichneumon non-ailé à deux vessies mobiles – De Geer, 1771: 909–911. Unavailable name discovered by Retzius, 1783: 70.

Ichneumon vesicularis Retzius, 1783: 70 . Neotype ♀ (present designation), NHRS, examined. Type locality: Sweden, Uppsala, Almunge.

Diplolepis vesicularis – Spinola, 1808: 161.

Cleonymus hemipterus Spinola, 1811: 149 . Nomen nudum discovered by Nees ab Esenbeck, 1834: 76.

Eupelmus DeGeeri Dalman, 1820: 379–381 . Lectotype ♀ (present designation), NHRS, examined. Type locality: Sweden. Synonymy by Nees ab Esenbeck, 1834: 77.

Eupelmus Geeri – Nees ab Esenbeck, 1834: 76. Unjustified emendation discovered by Dalla Torre, 1898: 275.

Macroneura maculipes Walker, 1837: 354 . Lectotype ♂, designated by Graham, 1969: 93, BMNH, examined. Type locality: United Kingdom. Synonymy with Eupelmus Geeri by Reinhard, 1857: 80.

Eupelmus maculipes – Giraud, 1863: 1270 (combination by inference in context).

Eupelmus albitarsis Costa, 1883: 101. Syntypes ♀, UDSN, examined. Type locality: Italy, Sardinia. – Costa, 1884: 332 (subsequent description). Synonymy by Ruschka, 1921: 302.

Euryscapus saltator Lindeman, 1887: 190–191 . Syntypes ♀ ♂, ZMMU, not examined. Type locality: Russia: Moscow, Vladimir, Nizhniy-Novgorod and Tula. Synonymy by Ruschka, 1921: 302.

Mira saltator – Dalla Torre, 1898: 238.

Eupelminus saltator – McConnell, 1918: 168–169.

Eupelmella Degeeri – Masi, 1919: 307.

Eupelmus vesicularis – Ruschka, 1921: 301.

Eupelmella vesicularis – Gahan, 1933: 52.

Eupelmella saltator – Procter, 1938: 425.

Macroneura vesicularis – Kloet & Hincks, 1945: 286.

Macroneura (Macroneura) vesicularis – Kalina, 1981: 94.

Eupelmus (Macroneura) vesicularis – Gibson, 1995: 197, 318.

Description: Female. Length = 1.8–3.2 mm. Body dark brown with mostly dark green metallic luster and even small or medium size specimens quite dark ( Fig. 45 ). Head brownish-black with dark green to bluish-green luster; vertex and parascrobal area with coppery or bronze metallic luster, lower face and gena with bronze metallic luster, sometimes interantennal area coppery-red and scrobal depression golden-green; frons sometimes distinctly tricoloured with dark bluish-green luster along inner orbits and mesally bronze to coppery below level of anterior ocellus. Antenna as described for E. balcanicus . Mesosoma similar in colour to head, with pronotum, mesoscutum, scutellum, acropleuron, and metapleuron almost black with bluish-green metallic luster and some coppery reflections and the median line and lateral panels of pronotum, prepectus, tegula, and axillae lighter, brown to dark brown. In some lighter coloured specimens mesosoma yellowish-brown with bluish-green metallic luster and usually with axillae, parts of pronotum and especially prepectus lighter, but acropleuron always rather dark; mesoscutal plate evenly and comparatively densely setose except narrowly asetose posteriorly, dark green to bluish-green with strong coppery reflections and concave part bluish-green medially, sometimes narrowly violet near hind margin; axillae more or less uniformly brown to dark brown with faint green luster anteriorly, but at least slightly contrasting with the greenish-black scutellum; scutellar-axillar complex with brownish hair-like setae only on axillae and base of scutellum and at least apical half of scutellum with whitish slightly lanceolate setae ( Fig. 63 ). Legs with a pattern of light areas as described in detail for E. balcanicus except coxae without violet luster; in some specimens middle leg and hind tibia almost entirely brownish-yellow. Metasoma with dense reflective whitish setae, on Gt2–Gt4 distance between sockets of two adjacent setae usually shorter than seta length ( Figs 97, 101 ); Gt1 dorsally with at least 14–18 setae arranged into a row and with at most 6 setae offset to form an indistinct second row ( Fig. 97 ), but sometimes with up to 30 setae arranged into two to three rows; dark brown with faint metallic luster, except Gt1 translucent, yellowish to reddish-brown (a colour similar to that of shellac flakes) and at most with a pair of darker subbasal spots with indistinct metallic hue ( Figs 73, 74 ). Ovipositor sheaths as described for E. messene , but frequently rather extensively darkened along ventral margin ( Fig. 45 ), hence with a yellowish elongate spot or a complete transverse band.

Head in lateral view hemispherical, in dorsal view 1.8–2.0× as broad as long, in frontal view 1.1–1.3× as broad as high. Frontovertex coriaceous-imbricate to reticulate. Pedicel plus flagellum 1.1–1.4× head width. Pronotal ridge with erect setae shorter than pronotal collar. Mesoscutal plate with flat, V-shaped anterior region differentiated by minute reticulate sculpture and posteromedial region evidently reticulate mesally and shallowly concave. Scutellum and axillae weakly convex ( Fig. 63 ). Acropleuron imbricate to slightly reticulate, with microsculptured region medially. Fore wing base extended to near petiole; basal cell densely setose dorsally; apical part normally abruptly bent upward, comparatively short, 1.3–1.7× as long as basal part, with marginal and postmarginal veins extended along straight leading margin and without a trace of a stigmal vein, posterior and leading margins obliquely angled to rounded or obtuse apex (cf. Figs 118–120 ). Hind wing concealed beneath fore wing and apically reflexed. Middle leg with row of four to seven mesotibial apical pegs; mesotarsus with asymmetrical peg pattern on basitarsus, anterior margin with 6–11 pegs in single row or sometimes differentiated into two medially slightly overlapping rows and posterior margin with two to four pegs (usually three and exceptionally four) within basal half and with or without one peg apically, second tarsomere with one to three pegs on anterior margin and one peg on posterior margin, and third and fourth tarsomeres with one apical peg on either side ( Fig. 83 ). Metasoma ovoidal and comparatively narrow, 1.8–3.0× as long as wide, Gt5 alutaceous-granular. Syntergum and anal plate forming truncate to somewhat obliquely inclined surface above ovipositor sheaths and gaster extending to about apex of second valvifer. Ovipositor sheaths 0.5–0.6× as long as metatibia, 0.3–0.4× as long as metasoma, and 0.9–1.1× as long as head.

Male. Length = 1.2–2.3 mm. Body metallic dark green ( Fig. 46 ). Head as described for E. barai except antenna with fl2 to fl8 with dense, strongly curved and more adpressed pale setae such as about apical half of seta parallel with segment surface; fl2, fl3 and rarely fl4 with a group of differentiated setae ventrally ( Fig. 105 ). Mesosoma with stronger metallic luster than head, uniformly dark green with coppery and bronze reflection except usually propodeum and rarely mesoscutum with comparatively strong green to bluish-green luster ( Fig. 55 ). Fore wing ( Figs 19 , 27 ) variably conspicuously and uniformly infuscate beyond parastigma, without a differentiated oval brownish area, hyaline only in smallest specimens, with setation and WIP as in E. barai . Legs and metasoma as described for E. barai .

Antenna ( Fig. 105 ) short, flagellum cylindrical, clava elongate with narrow ventral micropilose sensory region, pedicel plus flagellum 1.4–1.6× head width. Fl1 inconspicuous, strongly discoidal. Basal funiculars barrel-shaped, fl2 1.3–1.6, fl3 1.4–1.6, and fl8 1.0–1.3× as long as wide. Funiculars with dense MPS in two or three rows ( Fig. 123 ). Mesosoma 1.75–2.1× as long as broad. Fore wing 2.3–2.45× as long as broad. Propodeum coriaceous with percurrent median carina.

Comparative diagnosis and variability: The above description and the distribution map do not include three females that are provisionally identified as E. vesicularis but separated as Clade B. The three females form a distinct clade in the analysis based on COI and are identical at the sequenced 28S locus with specimens of E. barai from the Eastern part of the range of this latter species. One possible explanation is a historical introgression between E. barai and some other species morphologically similar to E. vesicularis (see under results: Molecular analyses). The three specimens originate in South France, Belgium, and Canada (Appendix 1). Because this is the only record from North America of a female that is morphologically identified as E. vesicularis , this could be an introduction not followed by establishment, or a mislabelling. These three females basically differ from all other examined E. vesicularis females only in having the axillae very dark and concolorous with the scutellum ( Fig. 62 , 95 ); otherwise they have the colour, the fore wing rudiment, and the pubescence of the gaster ( Fig. 75 ) as for females from the main clade of E. vesicularis , which includes also specimens from the type localities of Ichneumon vesicularis and Macroneura maculipes . I have seen further specimens with the axillae concolorous with the scutellum from S France (MNHN), N Spain (BMNH), N Italy (NHMV), and Switzerland (CNC) (Appendix 1).

Because of their dark colour, females of E. vesicularis can be confused with those of E. barai , but are distinguished by having the concave part of the mesoscutum bluish-green medially, without violet luster or sometimes only narrowly violet near the hind margin, while females of E. barai have a large triangular violet spot in the concave part. The reflexed dark part of the fore wing is shorter than in E. barai , about 1.5× the length of the basal part. Setae on the metasoma, although dense, are thinner and sparser than in E. barai (cf. Figs 70, 71 , 96, 99 with 73, 74, 97, 101), and even if reflective do not form a compact silvery white surface when examined under oblique angle at low magnification. Gt1 is usually less setose dorsally in E. vesicularis with setae arranged into a single row with some setae offset to form an indistinct second row ( Fig. 97 ), although sometimes similarly setose as in E. barai , with up to three rows of setae. The body of E. vesicularis is mainly dark with green reflections, while E. barai females additionally have dark violet reflections that frequently are predominant.

In addition to the characters mentioned in the key that are useful to separate males of E. barai from those of E. vesicularis (the distribution of the two species overlap in part, especially in the Carpathian Basin, Fig. 124 ), there is also a subtle difference in the colour of the mesonotum between most males of the two species. In E. barai , the scutellar-axillar complex is almost always differently coloured than mesoscutum (scutellum and axillae dark with coppery and bronze reflections and mesoscutum dark bluish-green with less coppery and bronze luster), whereas in E. vesicularis the mesonotum is rather uniformly dark green with coppery and bronze reflections, except rarely the mesoscutum is comparatively stronger green to bluish-green and then more similar to E. barai (cf. Figs 54 with 55).

Females of E. vesicularis are usually easily differentiated from those of E. messene because of their different body colour as described above. However, some lighter females of E. vesicularis can appear very similar to rare darker E. messene , because they have pale axillae that contrast rather conspicuously with the scutellum (axillae are light and reddish in most E. messene , cf. Figs 63 with 64). However, at least the acropleuron is always lighter in E. messene than in E. vesicularis , being brownish-yellow to reddish-brown with at most a very faint metallic luster (cf. Figs 43 with 45). Setation on the metasoma can also be used to separate females of the two species, because in E. messene the setae are brownish, nonreflective, and rather inconspicuous, while in E. vesicularis they are reflective and slightly thicker. The setae are usually also shorter and sparser in E. messene , although some specimens have them rather dense similar to some E. vesicularis with sparser pubescence (cf. Figs 72 with 73). Even in this situation, the metasoma of E. vesicularis appears more setose because the setae are more conspicuous ( Figs 73, 74 , 97 ). Also in E. messene, Gt 1 is much less setose dorsally, with single row of 5–15 inconspicuous setae arranged into one distinct row, sometimes with at most six setae offset to form a second indistinct row ( Fig. 98 ). In E. vesicularis, Gt 1 dorsally with at least 14–18 and up to 30 setae arranged into one to three rows ( Fig. 97 ). The number of basal pegs on the posterior margin of the basitarsus is reduced in E. vesicularis (two to four pegs with most specimens having three and very few four) compared to E. messene (three to six pegs with most specimens having four, although those with three or five pegs are rather frequent).

Biology: Polyphagous species; I have seen specimens reared from four insect orders. Coleoptera : Curculionidae : Ceutorhynchus obstrictus (Marsham) , Ischnopterapion loti (W. Kirby) , Xylocleptes bispinus (Duftschmid) . Diptera : Anthomyiidae : Botanophila seneciella (Meade) ; Cecidomyiidae : Asphondylia ulicis Trail , Giraudiella inclusa (Frauenfeld) , Lasioptera carophila F. Low , L. rubi (Schrank) ; Opomyzidae : Opomyza florum (Fabricius) . Hymenoptera : Cynipidae : Aulacidea follioti Barbotin , A. hieracii (Bouché) , Diplolepis mayri (Schlechtendal) , D. spinosissimae (Giraud) , Diplolepis sp. , Liposthenes glechomae (Linnaeus) , Phanacis caulicola (Hedicke) , P. centaureae Förster , P. hypochoeridis (Kieffer) , Timaspis lusitanica Tavares ; Diprionidae : Diprion pini (Linnaeus) ; Eurytomidae : Eurytoma noxialis (Portschinsky) , Tetramesa hyalipennis (Walker) , T. linearis (Walker) , T. maderae (Walker) , Tetramesa sp. ; Tenthredinidae : Pontania collactanea (Förster) . Lepidoptera : Coleophoridae : Coleophora discordella Zeller ; Glyphipterigidae : Glyphipterix simpliciella (Stephens) ; Gracillariidae : Phyllonorycter quinqueguttella (Stainton) ; Tortricidae : Grapholita delineana Walker , Rhyacionia buoliana ([Denis & Schiffermüller]).

Distribution: European (for ascertained records see under non-type material examined). All previous records of E. vesicularis from North America and Australasia are based on specimens of E. messene , those from other areas need confirmation. Based on the distribution data, it seems to prefer a more humid and cooler climate. Distributed mostly in the Boreal and Atlantic bioregions and also in the North of the Continental bioregion ( Fig. 124 ). Judging from label data, outside this area the species is present in mountains ( Bulgaria, Greece and Romania) or near wetlands ( Hungary). The single record from outside Europe is from Kyrgyzstan (in Tian Shan mountains at 1960 m altitude). Eupelmus vesicularis is likely a relict species in the part of its range where it is found mainly in mountains.

Remarks: Eupelmus vesicularis was described from Sweden by Retzius (1783) as Ichneumon vesicularis in his Caroli De Geer genera et species Insectorum , with reference to figure 22, plate 31 of De Geer’s (1771) work. This is reproduced here as Fig. 102 . Graham (1969) did not find any type specimen in the De Geer collection in Stockholm when he examined it in 1959. G. Gibson (personal communication) confirmed that no specimen and no pin hole is present in the bottom of the box under the label in this collection, suggesting that the type was lost prior to the De Geer collection being arranged in the present drawers by Dalman in 1820. My searches lead to the same result ( Fig. 102 ). However, the drawing in De Geer (1771), to which Retzius (1783) made reference, is a very realistic and vivid representation ( Fig. 102 ), so there is no doubt that what Retzius described is E. vesicularis s.l.

The three more common species confused under E. vesicularis ( E. vesicularis s.s., E. messene , and E. barai ) differ in very few morphological characters, of which all except colour are very inconspicuous and colour is quite variable. The examination of a large number of specimens from Sweden showed that a single species is present in the type locality, with all specimens quite similar except for variation in the degree of melanization and the number of mesotarsal pegs. In order to secure stability of nomenclature, a neotype is designated below to establish the taxon Ichneumon vesicularis objectively. The selected specimen is from near Almunge, in Uppsala lan, about 60 km south from Lövstabruk where De Geer lived. It agrees well with the description given by Retzius (1771: 909).

Eupelmus degeeri : This was described from a series of females from various localities in Sweden (Vestrogothia, Smolandia, Scania, Gottlandia & c. according to the original description), also with reference to the same drawing of De Geer (1771) as in the description of I. vesicularis . Apparently, Dalman (1820) was unaware of the description published 37 years earlier by Retzius. There are seven female syntypes in NHRS, all conspecific with the neotype of I. vesicularis and with specimens from all over Sweden, so I have no doubts that Eupelmus degeeri is a synonym of I. vesicularis . One specimen agrees well with Dalman’s description, is entire, uncontorted, and pinned laterally through metasoma in such a way that all useful characters are visible. It bears a small white label with ‘164’ on it, a small red rectangle, and a small purple rectangle. This specimen is here designated lectotype and other six specimens are considered paralectotypes.

Macroneura maculipes : Graham (1969) examined the type series and designated the male lectotype of M. maculipes . I concur with Graham that it is the male of E. vesicularis .

Eupelmus atrocoeruleus : This is a misspelling of E. atropurpureus Dalman. In his description of both sexes Thomson (1876: 106) indicated ‘Dalm. l. c. 381. 4’, referring to the fourth species of Eupelmus on page 381 of Dalman (1820), that is E. atropurpureus . The description of the female refers undoubtedly to E. atropurpureus . Ruschka (1921) considered the male was the male of E. vesicularis and listed E. atrocoeruleus in part as its synonym. Gahan (1933) considered it to be the male of E. atropurpureus , mainly because he considered that E. vesicularis (s.l.) males do not exist (only E. messene is present in North America and it is thelytokous).

Eupelmus albitarsis : This was considered a synonym of E. vesicularis by Ruschka (1921). The type was found at my request by Dr Nicola Maio in Costa’s collection deposited in UDSN. I received a photograph of the unique female type with the label ‘Tissi’ (comune in Sassari province, Sardinia) printed on a small white rectangle and a handwritten label ‘ Eupelmus albitarsis ’. Only one hind leg remains glued to the card rectangle and it is impossible to ascertain the identity of the species. Following Ruschka (1921), I consider it as a synonym of I. vesicularis . Although fascicle 2 of Atti della Reale Accademia delle Scienze Fisiche e Matematiche in Naples containing Costa’s paper ( Costa, 1883) is dated 30 June 1883, the whole volume 1 is dated 1888 (see also Gahan, 1933). It is not known to me if the different fascicles were published separately before the whole volume was printed in 1888. If not, the name E. albitarsis was made available from Costa (1884). Costa (1884) published the description of E. albitarsis a second time in a reprint containing only the descriptions of new species of his 1883 paper. Volume 15 for 1883 of Bullettino della Società Entomologica Italiana containing Costa’s paper ( Costa, 1884) was not published until 1884, as follows from the report of the treasurer on page xvii dated 5 June 1884.

Euryscapus saltator : This was described from Russia as a parasitoid of the Hessian fly and synonymized with E. vesicularis by Ruschka (1921). It was not possible to examine the type material of E. saltator , but because the most obvious character useful to separate the three common species in the vesicularis complex is body colour, even laconic species descriptions in the style of the 19th century can be interpreted to accurately review the synonyms. The species was described as black with green metallic luster, yellow legs, and first gastral tergite reddish-brown (characteristic colour of E. vesicularis ). Because of the large number of localities given in the original description, most likely Lindeman had a mixed type series including also E. messene .

Type material examined: SWEDEN: Neotype ♀ of Ichneumon vesicularis Retzius (present designation), labelled: Almunge sin / Harparbol / Upl., 2/5–43 / O. Lundbald ; Neotypus Ichneumon vesicularis Retzius, 1783 Det. Fusu L. 2008 [red label] ( NHRM) . Lectotype ♀ of Eupelmus degeeri Dalman (present designation), with following labels: ‘164’ [small white label]; small red rectangle; small purple rectangle; Lectotypus Eupelmus degeeri Dalman, 1820 Det. Fusu L. 2008 [red label]. Paralectotypes of Eupelmus degeeri , 6♀: two labelled with small yellow rectangle, four with small green rectangle ( NHRM) . UNITED KINGDOM: Lectotypus ♂ of Macroneura maculipes Walker, 1837 ( BMNH) .

Non-type material: Austria ( CNC, NHMV), Bulgaria ( PUPB / IBER), Czech Republic ( BMNH, CNC), Denmark

( CNC, LUZN), Estonia ( AICF, ZMUH), Finland ( ZMUH), France ( AICF, BMNH, CNC, MNHN), Germany ( BMNH, CNC, ENHM, HNHM, LUZN, NHMV), Greece ( AICF), Hungary ( AICF, BMNH, CNC, HNHM), Netherlands ( AICF), Poland ( AICF, HNHM), Romania ( AICF, ANCO, HNHM), Russia ( NHMV), Slovakia ( AICF), Sweden ( AICF, CNC, HNHM, LUZN, NHRM), and United Kingdom ( AICF, BMNH, CNC, RRAC).

Extralimital material: Kyrgyzstan ( AICF). See Appendix 1.

THE CLADE ‘ EURONMACRA’

Females of the ‘Euronmacra’ clade have an inconspicuous fore wing rudiment that is scale-like, apically pointed and adpressed to mesosoma ( Figs 65, 67 ). The hind wing is reduced to the humeral sclerite, with a minute white disc adpressed to dorsellum only in largest specimens. The long, differentiated setae on the pronotal ridge are grouped in a pair of tufts ( Figs 47 , 66–67 ). The acropleuron is semicircularly strigose in dorsal half, imbricate-alutaceous to strigose ventrally and with effaced sculpture mesally ( Fig. 91 ). Axillae are low convex but scutellum with extreme anterior edge Λ- like angulate.

EUPELMUS (MACRONEURA) IMPENNIS (NIKOL’SKAYA)

Figs 51 , 65 , 84 , 93 (♀), 20, 28, 52, 109 (♂)

Eupelmella impennis Nikol’skaya, 1952: 494. Syntypes ♀, ZIN, examined. Type locality: Soviet Central Asia [according to label data Tajikistan: Aiwanj and Uzbekistan: Dangara] .

Macroneura impennis – Bouček, 1970: 83.

Macroneura (Euronmacra) impennis – Kalina, 1981: 93, 105 (misidentification of E. vladimiri ).

Eupelmus (Macroneura) impennis – Gibson & Fusu, 2016: 22.

Description: Female. Length = 1.9–3.1 mm. Body somewhat tricoloured, with a metallic green head, reddish-brown or brownish-yellow mesosoma and brown metasoma ( Fig. 51 ). Colour and setation as described for E. vladimiri , except legs lighter with slightly paler, light yellowish, apices of mid- and hind tibiae, and yellowish to rufous mesotibial pegs. Concave part of mesoscutal plate with a wide-triangular purple to light-violet spot bordered anteriorly by blue and greenish-blue, covering about one-fourth to one third of mesoscutum length ( Fig. 65 ). Ovipositor sheaths generally light, only narrowly dark brown basally, otherwise almost uniformly pale or more frequently gradually darker to light brown apically and along ventral margin; apical brownish band usually not reaching dorsal margin of sheath ( Fig. 51 ).

Head in lateral view hemispherical, 1.6–1.8× as high as long, transverse in dorsal view, 1.8–2.0× as broad as long. Frontovertex coriaceous-imbricate to reticulate. Pedicel plus flagellum 1.2–1.3× head width. Pronotal ridge with two paramedial tufts with three to five setae each and about 0.8× as long as pronotal collar. Mesoscutal plate with flat, V-shaped anterior region differentiated by minute reticulate sculpture and posteriorly reticulate with larger mesh size to smooth in distal third and posterior concave area declined to median. Scutellum and axillae weakly convex except scutellum within extreme anterior edge Λ- like angulate ( Fig. 93 ). Acropleuron semicircularly strigose in dorsal half, imbricate-alutaceous to strigose ventrally and with effaced sculpture mesally. Wings ( Fig. 65 ) as described for E. vladimiri . Middle leg with row of three or four mesotibial apical pegs; mesotarsus without pegs ( Fig. 84 ). Metasoma ovoidal and comparatively narrow, 2.0–2.3× as long as wide, Gt5 coriaceous. Posteroventral margins of syntergum obliquely angled inward between anal plate and ovipositor sheaths so in posterior view appearing depressed over sheath. Gaster extending over base of third valvula hence ovipositor appearing very short ( Fig. 51 ). Ovipositor sheaths 0.5–0.6× as long as metatibia and about 0.3× as long as metasoma.

Male. Length = 1.5–1.7 mm. Body metallic bronze-green ( Fig. 52 ). Head dark green with bronze and coppery reflections mainly on scrobal depression, interantennal prominence, parascrobal area and frontovertex. Lower gena with one long seta. Maxillary palpus brown. Antenna with scape dark brown with dark greenish luster except narrowly yellow to light brown basally; outer surface similarly dark ventrally along entire length except without metallic luster; pedicel and flagellum brown; pedicel with line of three hooked setae ventrally and a straight distal seta much shorter than other three; fl2–fl8 with dense, curved and adpressed dark setae such as the apical half of seta parallel with segment surface; fl2–fl4 without differentiated black setae ventrally. Mesosoma similar in colour to head, sometimes with bluish-green luster on mesoscutum posteriorly between notauli. Tegula bicoloured, opaque yellowish-brown along inner margin, and variably broadly translucent light brownish apically. Fore wing translucent except with a diffuse oval infuscate area behind marginal and stigmal veins; WIP with a yellow to orange and reddish band in apical third ( Fig. 20 ). Setae behind submarginal vein darker and stouter than on disc; costal cell dorsally with single line of five or six setae near leading margin over at most parastigma length and ventrally with one or two rows of setae along most of its length except more setose distally in front of parastigma with setae in about three rows; basal cell bare, sometimes with a few minute setae ventrally and at most one to five stout setae dorsally, but mediocubital and basal folds with stout, long dark setae; cubital fold setose along length ( Fig. 28 ). Front leg yellow to orangish except apical tarsomeres brownish, coxa concolorous with mesosoma, and sometimes femur faintly brownish on posterior surface and tibia faintly brownish basally and ventrally. Middle leg similar to front leg except femur brown in distal half on posterior surface and tibia darker mesally. Hind leg with coxa metallic, femur brown except variably extensively pale basally, tibia yellowish to orangish in about basal half or less, and darker in about distal half but apical region graduated rather than abruptly delineated. Metasoma dark brown with faint metallic luster under some angles of light and with bronze-green to coppery luster basally on Gt1.

Antenna ( Fig. 109 ) comparatively short, flagellum clavate with flagellomeres gradually wider and shorter toward apex and clava comparatively broad with broad ventral micropilose sensory region along length, collapsed in air dried specimens; pedicel plus flagellum 1.2–1.3× head width. Fl1 subquadrate and setose as subsequent flagellomeres, with two rows of setae. Basal funiculars cylindrical, fl2 1.5–1.6, fl3 1.4, and fl8 0.8× as long as wide. Funiculars with MPS in single row. Mesosoma 1.85–2.0× as long as broad. Fore wing 2.6–2.65× as long as broad. Propodeum coriaceous to reticulate with percurrent median carina.

Comparative diagnosis and variability: Females of E. impennis are structurally very similar to those of E. maculatus , but they differ markedly in colour pattern (see under this species). The male of E. impennis is most similar to that of E. muellneri (see under this species).

Figures

Figure 124. Distribution map of the species in the Eupelmus vesicularis complex: E. vesicularis (green squares), E. messene (yellow stars), E. balcanicus (blue triangles), E. barai (white circles). Insert: enlarged map of the area with the most dense records (Eastern Austria, Hungary, Romania, and Moldova).

Figures 43–50. Habitus of Eupelmus (Macroneura) species: E. messene ♀ (43), E. balcanicus ♀ holotype (44), E. vesicularis ♀ (45) and ♂ (46), E. muellneri ♀ (47) and ♂ (48), E. vladimiri ♀ (49) and ♂ (50). Scale bars – 1 mm.

Figures 56–67. Mesosoma in dorsal view in Eupelmus (Macroneura) females: E. aseculatus (56), E. rameli (57), E. seculatus (58), E. falcatus (59), E. balcanicus (60), E. barai (61), E. vesicularis clade B from Canada (62), E. vesicularis from Sweden (63), E. messene (64), E. impennis (65), E. maculatus (66), E. vladimiri (67). Scale bars – 0.2 mm.

Figures 68–87. Metasoma and mesotarsus in Eupelmus (Macroneura) females: E. rameli (68, 77), E. balcanicus (69, 78), E.barai from Romania (70, 79),E. barai from Spain (71, 82), E.messene (72, 80), E.vesicularis from Sweden (73, 83),E.vesicularis from Slovenia (74), E. vesicularis clade B from Canada (75), E. seculatus (76), E. messene (lectotype of Eupelminus coleopterophagus) (81), E. impennis (84), E. falcatus (85), E. muellneri (86), E. vladimiri (87). Scale bars – 0.2 mm.

Figures 96–102. Metasoma in dorsal view in females of: Eupelmus barai from Romania, reared air dried specimen (96), E. vesicularis from Sweden, collected in Malaise trap and chemically dried (97), and E. messene from Romania, reared air dried specimen (98). Detail of setation on the middle of Gt3 in E. barai (99), E. messene (100) and E. vesicularis (101). Reproduction of figure 22 of plate 31 in De Geer’s (1771) book and the empty space in De Geer’s collection showing no pin hole below the label for ‘I. non-ailé à 2 vesſies mobiles T.2 p. 909.’ (102). Scale bars – 0.2 mm for 96–98 and 20 µm for 99–101.

Figures 111–123. Details of Eupelmus (Macroneura) morphology. Fore wing rudiment in females: E. barai (111–115), E. rameli (116), E. aseculatus (117), E. messene (118–121). Antenna in males: E. barai (122) and E. vesicularis (123). Scale bar – 0.2 mm. wbL – fore wing base length, waL – fore wing apical part length. Numbers by the wings indicate the size of the specimen.

Figures 103–110. Antenna in Eupelmus (Macroneura) males: E. falcatus (103), E. seculatus with insert showing pedicel and fl1–fl4 (104), E. vesicularis with insert showing pedicel and fl1–fl3 (105), E. barai from Romania (106), E. barai from Spain (107), E. vladimiri (108), E. impennis (109), E. muellneri with insert showing pedicel, quadrate fl1 and base of fl2 (110). Scale bars – 0.2 mm. Black arrows in insert of Figures 104 and 105 point to the groups of differentiated setae on the ventral side of fl2 to fl4. White arrows in Figures 105 point to the MPS visible below the setae.

Figures 51–55. Habitus of Eupelmus (Macroneura) species: E. impennis ♀ (51) and ♂ (52), E. maculatus ♀ from Argentina (53), E. barai ♂ in dorsal view (54), E. vesicularis ♂ in dorsal view (55). Scale bars – 1 mm for 51–53 and 0.2 mm for 54, 55.

Figures 19–26. WIP on the wings of Eupelmus (Macroneura) males: E. vesicularis (19), E. impennis (20), E. barai (21), E. vladimiri – large specimen (22), E. falcatus (23), E. vladimiri – small specimen (24), E. seculatus (25), E. muellneri (26). Scale bars 0.2 mm.

Figures 27–34. Wings of Eupelmus (Macroneura) males: E. vesicularis (27), E. impennis (28), E. barai (29), E. vladimiri – large specimen (30), E. falcatus (31), E. vladimiri – small specimen (32), E. seculatus (33), E. muellneri (34). Scale bars – 0.2 mm. Arrows indicate the extent of the dorsal line of setae along leading margin of costal cell. Inserts: distal part of costal cell; dorsal setae are close to the leading wing margin, thicker, and pointing anteriorly.

Figures 88–95. Mesosoma in lateral view and scutellar-axillar complex in females of: Eupelmus aseculatus (88), holotype of E. rameli (89, 94), E. barai (90), E. muellneri (91), E. falcatus (92), E. impennis (93), E. vesicularis clade B (95). Scale bars – 0.2 mm for 88–91 and 50 µm for 92–95.