Eupelmus (Macroneura) rameli, Fusu, 2017

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 : 539-543

EUPELMUS (MACRONEURA) WALKER

1.

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2(1).

– KEY TO EUROPEAN SPECIES OF EUPELMUS (MACRONEURA) Female……....……...……....……....……...……....……....……...……....……....…….........……...……...… 2 Male (males of E. aseculatus and E. rameli are unknown but probably exist; E. messene and most likely E. maculatus are represented by females only)…….......…….....……..............………...………..13 Fore wing rudiment conspicuous, with a translucent base adpressed to mesosoma and a brownish apical part abruptly bent upward ( Figs 56–64 ) [if apical part broken, remaining basal part truncated apically and apex of hind wing visible beneath ( Fig. 62 )]. Hind wing present, concealed beneath fore wing and apically reflexed and infuscate ( Fig. 60 ). Pronotal ridge with ( Figs 56–58, 60–64 ) or without ( Fig. 59 ) a line of erect setae. Acropleuron alutaceous to slightly reticulate, with at most faintly imbricate sculpture mesally ( Figs 88–90 ) ..............…….….....…….....….……........…….....…….....……................…….....…….....……........……….3 Fore wing rudiment inconspicuous, scale-like, apically pointed, and adpressed to mesosoma ( Figs 65–67 ). Hind wing reduced to the humeral sclerite, with a minute white disc adpressed to dorsellum only in largest specimens. Pronotal ridge with a pair of tufts of differentiated setae ( Figs 47 , 65–67 ). Acropleuron semicircularly strigose in dorsal half, imbricate-alutaceous to strigose ventrally and with effaced sculpture mesally ( Fig. 91 )……………………………………………………………………..10

3(2). –

4(3).

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5(4).

– 6(3).

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7(6).

– Mesoscutal plate sparsely setose only anteriorly, with posteromedial concave region asetose or with one or two setae, mostly non-metallic, yellowish to brownish, with purple metallic luster posteriorly under some angles of light ( Figs 56–58 ). Scutellum and axillae asetose mesally, with only a few setae on sides ( Fig. 94 ). Fore wing sinuously pointed at apex ( Figs 116, 117 ). Frontovertex alutaceous to coriaceous, shining ( Figs 56–58 ). Acropleuron finely coriaceous to alutaceous, with almost completely effaced sculpture medially ( Figs 88, 89 ). Gt5 coriaceous ( Fig. 68 )….………………………………….....4 Mesoscutal plate more evenly and comparatively densely setose except anteriorly and posteriorly narrowly asetose, with strong metallic luster at least posteriorly ( Figs 59–64 ). Scutellum uniformly setose ( Figs 92, 95 ). Fore wing with rounded apex ( Figs 111–115, 118–121 ) or if sinuously pointed then very long, with apical part at least 4× as long as basal part ( Fig. 39 ). Frontovertex at least slightly roughened, coriaceous-imbricate to reticulate. Acropleuron imbricate to slightly reticulate, with microsculptured region medially ( Fig. 90 ). Gt5 alutaceous-granular ( Fig. 75 )………………….....6 Fore wing rudiment short, with apical part at most 1.4× as long as basal part and with straight leading margin ( Fig. 116 ). Body almost uniformly yellowish to brownish with very faint metallic luster ( Fig.36 ) and mesobasitarsus with at least seven dark pegs ( Fig. 77 )

..........................…………………………………………………………………………… E. (M.) rameli sp. nov.

Fore wing rudiment longer, with apical part at least 1.8× as long as basal part and with outcurved leading margin ( Fig. 117 ). Body tricoloured, with a metallic green head, reddish-brown or brownish-yellow mesosoma and brown metasoma ( Fig. 37 ) or mesotarsus with at most one inconspicuous peg (cf. Fig. 84 )

….….……………………………………………………………………………………………………….........5 Body almost uniformly yellowish to brownish with very faint metallic luster ( Fig. 35 ). Mesotarsus without pegs or rarely with single inconspicuous peg basally on anterior margin (cf. Fig. 84 ). Apical part of fore wing rudiment about 1.8–2.0× as long as basal part ( Fig. 117 ). …....…………..……………………………………….…….. E. (M.) aseculatus (Kalina) Body tricoloured, with head metallic green, mesosoma reddish-brown or brownish-yellow, and metasoma brown ( Fig. 37 ). Mesotarsus with numerous dark pegs ( Fig. 76 ). Apical part of fore wing rudiment longer, 2.0–2.4× as long as basal part…..……….......…....…....…….. E. (M.) seculatus (Ferrière) Mesotarsus without pegs but with one to three longer seta-like spines ( Fig. 85 ). Ovipositor sheaths nearly uniformly dark brown and apparently very short because gaster extending over about basal half of third valvula ( Fig. 39 ). Fore wing with basal cell asetose ( Figs 59 , 92 ) and apical part very long and sinuously pointed ( Fig. 39 ), at least 4× as long as basal part. Pronotal ridge with recumbent, undifferentiated setae ( Fig. 59 ). Axillae and scutellum with about anterior third Λ- like angulate, with dense, lanceolate, reflective setae ( Fig. 92 ) ……………....…..…..….…..…..…..…..…..…..…..…..…..…..…..…..…..…. E. (M.) falcatus (Nikol’skaya) Mesotarsus with numerous dark pegs, with asymmetrical peg pattern on basitarsus ( Figs 78–83 ). Ovipositor sheaths with a pale median band or spot and gaster usually extending at most to apex of second valvifer ( Figs 41 , 43–45 ). Fore wing with basal cell densely setose dorsally ( Figs 60–64 , 95 ) and apical part shorter and with rounded apex ( Figs 111, 118 ). Pronotal ridge with line of erect, comparatively long setae ( Fig. 64 ). Axillae and scutellum weakly convex, with sparse, hair-like setae ( Fig. 95 ) [ vesicularis complex]…………………………………………………………………………………………………………...... 7 Mesoscutalplatewithconcavepartonlynarrowlyvioletnearhindmargin( Fig.63 )orrarelywithanarrow purple to light violet median stripe bordered by bluish-green ( Fig. 64 ). Body tricoloured ( Fig. 43 ) or dark brown with mostly dark-green metallic luster ( Fig. 45 ), with Gt1 completely yellowish to reddish-brown or with indiscernible metallic hue ( Figs 72–74 ), and at most with a pair of darker subbasal spots ( Fig. 73 ). Fore wing rudiment short, with apical part 1.3–1.69× times longer than basal part ( Figs 118–120 ).….….……………………………………………………………………………………………………………….8 Mesoscutal plate with concave part having a large triangular violet spot narrowly bordered on outer sides by blue and bluish-green ( Figs 60, 61 ). Body dark brown ( Figs 41 , 44 ) with Gt1 translucent in distal half, yellowish to light brownish, and dark basally with strong metallic luster ( Fig. 70 ). Fore wing rudiment long, with apical part 1.71–2.5× times longer than basal part ( Figs 111–113 )…………………………………………………………………………………………………………..9

8(7).

– Metasoma with short, sparse, brownish setae, the setae on Gt2–Gt4 usually about equal in length to distance between sockets of two adjacent setae ( Figs 72 , 98, 100 ); Gt1 whitish to brownish-yellow ( Fig. 72 ), dorsally with 5–15 inconspicuous setae arranged into one distinct row and sometimes at most 6 setae offset to form a second row ( Fig. 98 ). Body tricoloured, head metallic green, mesosoma mostly reddish-brown or brownish-yellow, and metasoma brown ( Fig. 43 ), but some smaller specimens almost uniformly yellowish. Mesoscutal plate brownish on outer sides, the rest bronze-green to coppery except concave part bluish-green medially and usually narrowly violet near hind margin or rarely with a narrow purple to light violet median stripe bordered by bluish-green ( Fig. 64 ).......... ........................................................................................................….. E. (M.) messene Walker stat. rev. Metasoma with dense, reflective setae, on Gt2–Gt4 distance between sockets of two adjacent setae usually shorter than seta length ( Figs 73, 74 , 97, 101 ); Gt1 darker, yellowish to reddish-brown ( Fig. 73 ), dorsally with 14–30setae arranged into 1–3rows( Fig.97 ). Body dark brown with mostly dark-green metallic luster( Fig.45 ). Mesoscutal plate almost uniformly dark green to bluish-green with strong coppery reflections and concave part bluish-green medially, only rarely narrowly violet near hind margin ( Fig. 63 ) ……..........................................................................................................….. E. (M.) vesicularis (Retzius)

9(7).

– Metasoma with very dense reflective setae, on Gt2–Gt4 distance between sockets of two adjacent setae much less than seta length ( Figs 70, 71 , 96, 99 ). Gt1 dorsally extensively setose, with two to four rows of overlapping setae ( Fig. 96 ). Frontovertex coriaceous-imbricate to reticulate, dull ( Fig. 61 ). Mesoscutum with posteromedial concave region finely coriaceous, evenly and comparatively densely setose except narrowly asetose posteriorly ( Fig. 61 ). Head 1.0–1.1× as wide as high ……........………... E. (M.) barai sp. nov. Metasoma with short and very sparse hair-like setae, on Gt2–Gt4 distance between sockets of two adjacent setae greater than seta length ( Fig. 69 ). Gt1 dorsally asetose or with only three to nine setae in single row ( Fig. 69 ). Vertex coriaceous, appearing shiny under low magnification ( Fig. 60 ), and frons mostly coriaceous-imbricate to slightly reticulate. Mesoscutum with posteromedial concave region mostly smooth and shiny or with effaced sculpture, with setation progressively sparser posteriorly and broadly asetose in front of scutellum ( Fig. 60 ). Head 1.1–1.2× as wide as high …...................................... ..............................................................................................................................… E. (M.) balcanicus sp. nov.

10(2).

– Mesotarsus without pegs ( Fig. 84 ). Gaster extending over base of third valvula, hence ovipositor sheaths apparently very short ( Fig. 51, 53 ). Ovipositor sheaths lighter, only narrowly dark-brown basally, otherwise almost uniformly pale or more often gradually darker to light brown apically and along ventral margin, and with apical brownish band not reaching dorsal margin of sheath ( Figs 51, 53 ), although rarely as described below. Pronotal ridge with tufts of setae about 0.8× as long as pronotal collar ( Fig. 53 ). Mesoscutal plate with posterior depressed area with a wide-triangular violet spot bordered anteriorly by blue and greenish-blue covering about one-fourth mesoscutum length ( Fig. 66 ) ……………..........................................………………………………………………………………11 Mesotarsus typically with at least one peg and usually more ( Figs 86, 87 ), although extremely rarely without. Gaster usually extending at most to apex of second valvifer, hence ovipositor sheaths appearing longer ( Figs 47, 49 ). Ovipositor sheaths darker, comparatively abruptly dark brown in about apical third to half, more or less strongly along ventral margin and basally, and with a yellowish elongate spot ( Fig. 49 ) or almost complete transverse band ( Fig. 47 ). Pronotal ridge with tufts of setae about as long as or slightly longer than pronotal collar ( Fig. 47 ). Mesoscutal plate sometimes without a delimited violet spot, at most violaceous under some angles of light.....................………12 11(10). Tegula white to yellowish white, contrasting with surrounding sclerites ( Fig. 66 ). Body almost uniformly dark, with hardly lighter mesosoma and comparatively strong metallic luster ( Fig. 53 )……… ……………………………………………………………….....……….……….. E. (M.) maculatus (Ferrière)

– Tegula yellowish-brown to reddish-brown, concolorous with mesoscutum ( Fig. 65 ). Body somewhat tricoloured, head metallic green, mesosoma reddish-brown or brownish-yellow, and metasoma brown ( Fig. 51 )……………………………………………………...............……….. E. (M.) impennis (Nikol’skaya) 12(10). Basitarsus with one to three (most frequently two) basal pegs on either side and with total number of basal pegs not less than three; basitarsus and second tarsomere both with one apical peg on either side ( Fig. 86 ). Mesoscutal plate with posterior depressed area having a wide-triangular violet spot bordered anteriorly by blue and greenish-blue and covering about one-fourth mesoscutum length (cf. Fig. 66 )…………….….….….….….…......................….….….….….….….…… E. (M.) muellneri Ruschka

– Basitarsus without pegs or with at most a single peg basally on posterior margin near base of spur, and sometimes with one apical peg on either side although rarely one of the two apical pegs missing; second tarsomere rarely with one or both apical pegs missing ( Fig. 87 ). Mesoscutal plate with posterior depressed area having a very faint purple to violet luster under only some angles of light ( Fig. 67 ) …………………………………………............…………………………………….. E. (M.) vladimiri sp. nov.

13(1).

– Basal cell similarly setose as disc ( Figs 27, 29, 31, 33 ). Flagellum cylindrical ( Figs 103–107 ), clava elongate with narrow ventral micropilose sensory region; fl1 inconspicuous, strongly discoidal, with at most a couple of short setae along extreme apical margin ( Figs 103–107 ); fl2, fl3 and sometimes fl4 with a group of differentiated setae ventrally, visible in lateral view as short, dark tufts ( Figs 104, 105 ). Mid- and hind tibiae with light and dark contrasting pattern, yellow to yellowish-white basally and comparatively abruptly brown in distal half to one-fourth ( Figs 38, 40, 42 , 46 ); hind tibia sometimes extensively pale with dark markings reduced to apical spots. Fore wing variably conspicuously infuscate, from almost hyaline ( Fig. 33 ) to almost uniformly dusky beyond parastigma ( Fig. 27 ) or with a diffuse oval infuscate area behind marginal and stigmal veins ( Fig. 29 )……………………........................…………………………………….……………………...14 Basal cell bare ( Figs 30, 34 ) or with only a few minute setae ventrally and at most one to five stout setae dorsally ( Figs 28, 32 ), but mediocubital and basal folds with stout dark setae, larger than setae on disc. Flagellum at least indistinctly subclavate ( Fig.108 ) to clavate ( Fig.109 ), with flagellomeres gradually wider and shorter toward apex and clava comparatively broad, with broad ventral micropilose sensory region; fl1 sometimes subquadrate or only slightly transverse and setose as subsequent flagellomeres, with two rows of setae ( Figs 109, 110 ); fl2–fl4 without differentiated setae ventrally. Mid-tibia with light and dark parts mostly not contrasting, almost uniformly dark or uniformly light to gradually darker from about middle; hind tibia mostly as above, but with dark region more extensive ( Figs 48, 50 , 52 ). Fore wing with conspicuous oval brown spot behind marginal and stigmal veins ( Figs 28, 30, 34 ) except in smallest specimens of about 1 mm where spot faint and diffuse ( Fig. 32 )…………….........................…………………………….. 17 14(13). Legs extensively yellow: front femur yellow except sometimes with a brownish spot medially on posterior surface; mesofemur variably extensively yellow basally to almost entirely yellow with only a brownish stripe along posterior surface ( Figs 38, 40 ). Scape usually with up to basal third yellowish on inner and outer surfaces plus yellow sensory region ( Figs 103, 104 )……..................……………. 15

– Legs with anterior and middle femora dark except narrowly pale basally ( Figs 42 , 46 ). Scape, excluding yellow sensory region, usually only narrowly yellowish basally ( Figs 105, 106 ) [ vesicularis complex] …....….……….….….….….….….….….….….….….….….….….….….….…….........…………................ 16 15(14). Tegula bicoloured, opaque yellowish-white to yellowish-brown along inner margin and variably broadly translucent brownish apically. Fore wing elongate, 2.6–2.8× as long as broad, with a diffuse oval infuscate area behind marginal and stigmal veins; costal cell with a dorsal line of less than five setae for at most parastigma length; speculum broadly open; basal cell setation sparser than on disc ( Fig. 31 ). Antenna long ( Fig. 103 ), pedicel plus flagellum 1.8– 1.9× head width. Body mostly bright bluish-green, head concolorous with mesosoma ( Fig. 40 ) ……………………………………….................................…………………… E. (M.) falcatus (Nikol’skaya)

– Tegula brown. Fore wing comparatively short, 2.1–2.3× as long as broad, hyaline; costal cell with a dorsal line of more than seven setae over about apical half or less but clearly over more than parastigma length; speculum closed posteriorly by setae; basal cell setation similarly dense as on disc. Antenna short ( Fig. 104 ), pedicel plus flagellum 1.4–1.5× head width. Body with frontovertex mostly dark violet and face bright bluish-green, with these areas of different colour well delimited along upper limit of scrobal depression; mesosoma green to bluish-green with bronze reflections and pronotum dark violet dorsally ( Fig. 38 ).…………..............................……………………………...………….…………. E. (M.) seculatus (Ferrière) 16(14). Funiculars with MPS in single row ( Fig. 122 ) and with semierect slightly curved setae, with only apices of setae parallel with segment surface ( Figs 106, 107 ); basal funiculars subcylindrical ( Fig. 107 ). Antenna long, pedicel plus flagellum 1.6–1.8× head width. Fore wing with a variably conspicuous diffuse oval infuscate area behind marginal and stigmal veins ( Fig. 29 ) ……..…………………………...……………….…… E. (M.) barai sp. nov. and E. (M.) balcanicus sp. nov.

– FunicularswithnumerousMPSintwotothreerows( Fig.123 )andwithstronglycurvedandmoreadpressed setae such as about apical half of setae parallel with segment surface ( Fig. 105 ); basal funiculars barrel-shaped ( Fig. 105 ). Antenna short, pedicel plus flagellum 1.4–1.5× head width. Fore wing variably conspicuously infuscate beyond parastigma, without a clearly differentiated oval brownish area ( Fig. 27 ).………………………………………………………….………………………... E. (M.) vesicularis (Retzius) 17(13). Flagellum subclavate and longer, pedicel plus flagellum about 1.5–1.6× head width, fl8 usually longer than wide or at most subquadrate; fl1 inconspicuous, strongly discoidal, with at most a couple of short setae along extreme apical margin ( Fig. 108 ). Fore coxa largely nonmetallic, yellowish, and concolorous with femur ( Fig. 50 ). Costal cell ventrally with single row of setae along length except more setose distally in front of parastigma ( Fig. 30 ). Hind tibia with about basal half or slightly less yellowish to orangish and about distal half darker ( Fig. 50 ).............................................. E. (M.) vladimiri sp. nov.

– Flagellum clavate and short, pedicel plus flagellum about 1.3× or less head width, fl8 subquadrate to transverse; fl1 subquadrate or only slightly transverse and setose similar to subsequent flagellomeres, with two rows of setae ( Figs 109, 110 ). Fore coxa metallic, concolorous with mesosoma ( Fig. 52 ). Costal cell ventrally sometimes with two rows of setae along length except more setose distally in front of parastigma. Hind tibia sometimes with more than distal half darkened ( Fig. 48 ) ……………………………………………………....18 18(17). Fore femur dark brown; hind tibia with basal quarter yellowish-white ( Fig. 48 ). Cubital fold largely bare behind speculum, i.e. speculum open posteriorly ( Fig. 34 ). .. …………… E. (M.) muellneri Ruschka

– Fore femur yellow to orangish; hind tibia with about basal half or slightly less yellowish to orangish ( Fig. 52 ). Cubital fold setose along length, that is speculum closed posteriorly ( Fig. 28 )…………...........................……………………………………………….. E. impennis (Nikol’skaya)

SPECIES WITH APICAL PART OF FORE WING BROWNISH AND ABRUPTLY BENT UPWARD

[the clade E. ( Macroneura ) s.s. and E. falcatus species sola]

Females of all the species treated below have a fore wing with a translucent base adpressed to mesosoma and with a brownish apical part abruptly bent upward. Hind wing is present, concealed beneath the fore wing, apically reflexed, and infuscate. The articulation between the two parts of the fore wing is fragile and the apical part breaks easily. Even if the fore wing is broken, a specimen can still be recognized as belonging here because the remaining basal part is truncated apically and the apex of the hind wing is visible beneath ( Fig. 62 ). Sculpture on acropleuron is alutaceous to slightly reticulate, at most faintly imbricate mesally ( Figs 88–90 ).

Figures

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 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.

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 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 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 35–42. Habitus of Eupelmus (Macroneura) species: E. aseculatus ♀ (35), E. rameli ♀ holotype (36), E. seculatus ♀ (37) and ♂ (38) with the insert showing the characteristic colour of the head, Eupelmus falcatus ♀ (39) and ♂ (40), E. barai ♀ (41) and ♂ (42). Scale bars – 1 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 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 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 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.