Revision of Lutnes Cameron (Hymenoptera: Eupelmidae) Author Gibson, Gary A. P. text Zootaxa 2018 2018-04-30 4415 2 330 356 journal article 30149 10.11646/zootaxa.4415.2.5 efc997e8-29bf-4b9d-8dc8-488626b30627 1175-5326 1242004 862FFD70-CB4F-45EA-BF25-FED1B621842F Lutnes testaceus (Cameron) n. comb. Figs 46–59 Eupelmus testaceus Cameron, 1884 : 117 . Argaleostatus testaceus ; Gibson, 1995 : 149 ; figs 13, 14, 123, 124, 235, 236, 289, 472, 511. Material examined. Type material. Syntype ♀ (BMNH): “V. de Chiriqui, | 3-4000 ft . | Champion, / Type / Eupelmus | testaceus Cam. type | BCa Hy i, 112 [hand written] / B.C.A. Hymen. I. | Eupelmus | testaceus , | Cam. / B.M. TYPE | HYM. | 5.989 / NHMUK010749026. Cameron (1884) described length of the species as “6–7 millim”. Nothing else in the description indicates he had more than one specimen, but because of the variation in length given I interpret the BMNH type specimen as a syntype . It is card mounted by its venter ( Fig. 49 ), with some parts partly covered by a film of glue, and is contorted but is mostly complete except the following are missing: right flagellum beyond fl6, apices of left fore wing and right hind wing, and left fore leg and both hind legs beyond coxae. The BMNH specimen from Panama , Bugaba, was also collected by Champion and appears to be from the Cameron collection; it could be a second syntype except that only Volcan de Chiriquí was given for the type locality in the original description. Additional material examined. Bolivia . Beni , Rurrenabaque , 270 m , 18-23.VII.1979 , M. Cooper (1 b BMNH ). Brazil . Goias , Jatai , XI.1972 , Oliveira ( 1 m BMNH ). Costa Rica . Heredia Pr ., La Selva Biol. Sta. , 3 km S Pto. Viejo , 10°26’N 84°01’W , 18.V. 1990 . H.A. Hespenheide ( 1m ♀, CNC). Puntarenas , Pen-Osa , 23 km N Puerto Jimenez , La Palma , 10 m , II-III.1993, P. Hanson (1 b CNC ). Ecuador . NAPO , Res. Ethnica Waorani , 1 km S Onkone Gare Camp , Trans. Ent. , 220 m , 00°39'10"S 076°26'W , 220 m , 21.VI.1994 (at 7 x-trans, 19 m mark, Lot# 693: 1 m ♀ CNC, CNC Photo 2017-11; at 7 x-trans, 73 m mark, Lot# 697: 1 m ♀ USNM; at 9 x-trans, 15 m mark, Lot# 711: 1 m ♀ USNM, CNC Photo 2017-12), 9.X.1994 (at trans 6, Sta. 2, Lot# 921: 1 m ♀ CNC), 10.X.1994 (at trans 4, Sta. 3, Lot# 942: 1 m ♀ USNM), T.L. Erwin et al., insecticidal fogging of mostly bare green leaves, some with covering of lichenous or bryophytic plants in terre firme forest, Project MAXUS . Orellana , Tiputini , Biodiversity Stn. , 00°37'55"S , 076°08'39"W , 216 m , 7.II.1999 (Transect 6, Lot# 2057: 1 m ♀ CNC; 1 m ♀ USNM), 8.II.1999 (Transect 4, Lot# 2037: 1 m ♀ USNM), 4.VIII.1998 (Transect 7, Lot# 1862: 1 m ♀ USNM; Transect 7, Lot# 1864: 1 m ♀ CNC), 22.X.1998 (Transect 7, Lot# 1965: 1 m ♀ USNM), 23.X.1999 (Transect 2, Lot# 1915: 1 m ♀ CNC), T.L. Erwin et al., insecticidal fogging of mostly bare green leaves, some with covering of lichenous or bryophytic plants. Pichincha Prov., Rio Palenque Stn., 47 km S Santo Domingo , 1-4.V.1983 , L.D. Coote (1 b CNC ). Guyana . Kourou , Savane , II.2013 , G.L. Giuglaris , window pan trap ( 1 m CNC ). Panama . Bugaba, Champion (1 b BMNH ). Canal Zone , Barro Colorado Is., 9°10'N 79°50'W— 3.VI.1970 , H. Hespenheide (1 b CNC ) ; 13-20.I.1993 (2 b ♀ CNC, CNC Photo 2017-9), 26-27.I.1993 ( 1 m ♀ CNC), 10- 17.II.1993 (1 b ♀ CNC), 17-24.III.1993 (1 b ♀ CNC, CNC Photo 2017-8), 28.VII-4.VIII.1993 (2 b ♀ CNC, CNC Photo 2017-10), J. Pickering. Canal Zone, Pipeline Road, 20.III.1982 , W.J. Hanson (1 b EMUS ). Chiriquí Prov. , Concepcion to Volcan , 3.VI.1977 , 500- 1500 m , S.&J. Peck (1 b CNC ) . Colón Prov., 2 km S Sabanitas , 9°19'19"N 79°47'54"W , 120 m , 15-19.VII.1999 , A. Gillogly & J. Woolley , YPT 99/027 (1b TAMU ). Venezuela . Zulia , El Tucuco , 200 m , 23.IV.1981 , L. Masner (1 b CNC ). Description. FEMALE (habitus: Figs 48–51 ). Length = 3.5–4.5 mm . Head ( Figs 46–51 , 54, 55 ) sometimes almost entirely bright green ( Figs 47 , 55 ), but usually at least interantennal prominence and frontovertex with variably extensive and distinct reddish-violaceous, coppery, bluish or purple lusters, and sometimes head almost entirely dark reddish-violaceous ( Figs 46, 49 , 54 ) except for more greenish lower face. Head in lateral view ( Figs 50, 51 ) with frontal surface variably convex, about 1.5–1.9× as high as long; malar space about 0.33–0.45× eye height. Head in frontal view ( Figs 46, 47 ) without differentiated ocellocular line between posterior ocellus and inner orbit; punctate-reticulate with interantennal prominence often somewhat more minute reticulate-rugulose; with whitish-translucent, lanceolate setae on interantennal prominence, lower face below toruli, clypeus, and about lower one-third of parascrobal region though more extensively along scrobal margin, but with longer, light brownish, hairlike setae on lower face below lanceolate setae lateral of clypeus and white hairlike setae on parascrobal region above lanceolate setae, with setae on vertex entirely white and undifferentiated from setae on upper parascrobal region or with dark, longer and/or stouter setae at least posteriorly at confluence with occiput variably distinctly differentiated from white setae on upper parascrobal region; scrobal depression bell-shaped, higher than wide with carinate lateral margin extending near anterior ocellus, but dorsally not carinately margined below ocellus and ventrally with lateral margin recurved toward lower inner orbit lateral to torulus, though comparatively inconspicuously so; interorbital distance about 0.3–0.4× head width; OOL about 0.6–1.0×, POL about 1.6–3.0×, and LOL about 0.9–1.5× MPOD, respectively. Labiomaxillary complex yellowish. Eye moderately setose ( Figs 46, 47 ); about 1.25–1.4× as high as long; height 1.35–2.0× and length 1.0–1.6× interocular distance, respectively. Antenna sometimes almost entirely yellowish except clava dark at least apically ( Figs 49, 51 ), but often apical 1–3 funiculars ( Fig. 50 ) and sometimes scape basally and apically dark, and if flagellum partly dark basally then at least flagellomeres paler dorsally; fl1 slightly transverse to subquadrate, with dorsal length about 0.25–0.4× dorsal length of pedicel and about 0.3–0.4× dorsal length of fl2; fl2 the longest funicular, about 1.9–2.5× as long as wide, and subsequent funiculars gradually decreasing in length such that fl8 quadrate to slightly transverse; clava about 2.3–2.7× as long as wide and about as long as combined length of fl8 + fl7 + about apical half of fl6. Mesosoma usually entirely orangish-yellow to orange except for dark mark on pronotum anterior to spiracle ( Figs 49, 51 , 55 ) but sometimes mesopectus and acropleuron darker brown anteriorly to about level of apex of tegula ( Fig. 56 ), and rarely mesoscutum similarly dark as head ( Figs 48 , 54 ). Pronotum variably setose, though usually with comparatively long, dark hairlike setae at least posterolaterally ( Figs 54–57 ); propleuron and prosternum with white setae. Mesoscutum ( Figs 54, 55 ) mostly punctate-reticulate except lateral flange more finely meshlike coriaceous-reticulate to transversely strigose, with lateral lobe longitudinally carinate over less than posterior half; quite densely setose with brownish to dark setae intermixed with variably numerous but more inconspicuous white setae within depressed median region ( Figs 55, 56 ). Scutellar-axillar complex ( Figs 54, 55 ) similarly punctate-reticulate as mesoscutum or more longitudinally reticulate-strigose, but with entirely dark setae. Prepectus setose ( Figs 56, 57 ), with some dark and/or white setae, and subdivided by vertical line near mid-length. Mesopleurosternum at least mostly with conspicuous white setae on acropleuron and/or mesopectus ( Fig. 56 ), but often with variably numerous and conspicuous dark setae on acropleuron dorsally and sometimes on mesopectus dorsally below tegula ( Fig. 57 ); acropleuron mostly meshlike reticulate with smaller reticulations mesally. Macropterous ( Figs 48, 50 ) to variably obviously brachypterous ( Figs 49, 51 ). Fore wing of macropterous individual extending at least to near posterior margin of Gt6 ( Figs 48, 50 ) and of normal shape ( Fig. 52 ) such that length about 3.0–3.5× width, with ratio of cc: mv: stv: pmv = 6.5–9.7: 3.7–5.7: 1.0: 2.6–3.0; costal cell dorsally densely setose with orangish to somewhat darker brown setae; basal cell usually conspicuously though variably dark orangish with dense, comparatively thick orange setae similar to costal cell except often for darker brown setae posterior of mediocubital fold to basal fold, and apically behind apex of mv hyaline with white hairlike setae; discal region orangish with orangish lanceolate setae from basal fold to about apex of pmv but more hyaline with somewhat darker brown hairlike setae beyond pmv, except for anterior hyaline region with white hairlike setae behind marginal vein within anterior half apically separated from or extending to base of stigmal vein (mv anterior to hyaline region more distinct than elsewhere because setae pale), somewhat smaller posterior hyaline region with white hairlike setae behind medial fold, and often darker brown lanceolate setae along basal and sometimes apical margins of anterior hyaline region and sometimes basal to posterior hyaline region. Fore wing of brachypterous individual extending at least to about middle of gaster ( Fig. 51 ), sometimes with leading and posterior margins diverging quite distinctly and disc longer than basal cell so as to resemble unreduced fore wing except for length and shorter stv, but usually wing obviously elongate-narrow, about 5.5–6.0× as long as wide, with subparallel leading and posterior margins ( Figs 49, 53 ), basal region and disc subequal in length, stv often almost subsessile, anterior and posterior hyaline regions often more extensively bordered basally and apically by dark brown lanceolate setae than for unreduced wing, and posterior hyaline region often very small or not apparent except for a few white setae along posterior margin of wing. Legs usually yellowish-orange similar to mesosoma except metafemur with longitudinal dark stripe on inner surface dorsally ( Fig. 51 ) and sometimes following dark: outer surface of mesotibia subbasally, outer surface of metatibia over about dorsal half, and meta- and/or mesotarsi; dorsal surface of protibia and mesotibia similarly convex. Propodeum ( Fig. 58 ) with medial length about 0.5–0.6× length of scutellum; with variably distinctly transversely strigose or strigose-rugulose furrows along foramen converging anteriorly to delineate complete though often somewhat irregular carina medially and slender lunate region posteromedially; callar region subdivided by oblique to sinuous postspiracular groove, with region anteromesal of groove distinctly meshlike coriaceous to reticulate and region posterolateral of groove smooth and shiny. FIGURES 46–53. Lutnes testaceus (Cameron) , ♀. 46 & 47 , head, frontal: 46 , 2017-11 (m); 47 , 2017-9 (b). 48 & 49 , dorsal habitus: 48 , 2017-11 (m); 49 , BMNH syntype [insert: enlargement of head] (b). 50 & 51 , lateral habitus: 50 , 2017-11 (m); 51 , 2017-8 (b). 52 & 53 , fore wing: 52 , 2017-12 (m), 53 , 2017-9 (b) [b = brachypterous, m = macropterous]. Gaster usually similarly orangish-yellow to orange as mesosoma ( Figs 49, 51 ) though sometimes variably extensively darker brown ( Figs 48, 50 ) except Gt2 and Gt1 at least apically paler, usually more translucent yellowish-white to white than subsequent tergites ( Figs 48–51 ), with Gt1 sometimes entirely pale dorsally except brownish dorsolaterally near middle; ventrally white or at least distinctly paler than apical or sometimes apical two orangish-yellow to light brownish sternites ( Figs 50, 51 ); Gt1 and Gt2 smooth and shiny but subsequent tergites increasingly distinctly meshlike reticulate ( Fig. 59 ); Gt1 bare and Gt2 at most with single, sparse line of setae dorsally, but subsequent tergites extensively setose with pale hairlike setae; syntergum obviously transverse, but about two-thirds as long as Gt6. Distribution. Neotropical: Bolivia , Brazil , Costa Rica , Ecuador , Guyana , Panama , Venezuela . Remarks. The above description is based on 16 macropterous and 14 brachypterous females, excluding the syntype . There is some variation in relative length and shape of the fore wing such that some females are less readily assigned to one or the other form, but obviously brachypterous females have distinctly elongate-slender fore wings with subparallel margins (about 4.5–6× as long as wide, Fig. 53 ) compared to macropterous females (fore wing at most about 3.5× as long as wide, Fig. 52 ). As noted in the introduction, wing polymorphism is known for at least two species of Anastatus from the Palaearctic region as well as for Merostenus melinus (Yoshimoto) ( Gibson 2017a ) and Eupelmus atropurpureus Walker ( Gibson & Fusu 2016 ) within Eupelminae . Additional examples may well exist unrecognized in other genera or regions where the species are less well known. In examining available material I observed one structural and two setal features that tended to correlate with brachyptery versus macroptery. Macropterous females typically have a broader interorbital region than brachypterous females ( cf . Figs 46, 47 ). Measured interorbital region width varied between 0.3× ( Fig. 47 ) and 0.39× ( Fig. 46 ) head width for all females, but was 0.3–0.34× for brachypterous females and 0.34–0.39× for macropterous females (excluding female from Guyana discussed below). Based on measurements taken from a dorsal habitus image ( Fig. 49 ), the interorbital width of the syntype is at or near the highest measured value for a brachypterous female. Most observed macropterous females also have entirely white setae on the mesopectus and acropleuron anterodorsally ( Fig. 56 ), and the vertex has dark setae, at least posteriorly at the confluence with the occiput ( Fig. 56 ), which often are stouter and/or longer so as to be quite distinctly differentiated from the white setae on the upper parascrobal region. Most observed brachypterous females have at least a few, though sometimes quite inconspicuous, dark setae on mesopectus and/or acropleuron dorsally below the tegula ( Fig. 57 ), and when more numerous then often also on the prepectus, and the vertex has similarly short and white setae as the setae on the upper parascrobal region ( Figs 55, 57 ). However, a macropterous female from Ecuador (collected 9 October , 1994) has some dark pleural setae and white vertexal setae similar to most brachypterous females (interorbital distance: head width = 0.34). The macropterous female from Brazil has dark seta on the vertex similar to most macropterous females, but dark setae also on the mesopectus and acropleuron dorsally similar to most brachypterous females (interorbital distance: head width = 0.35). Finally, the female from Guyana has fore wings that might be somewhat reduced in length (comparison difficult because gaster abnormally inflated and distended because of critical-point drying), but the specimen is considered as macropterous because the fore wings are about 2.9× as long as wide with divergent anterior and posterior margins ( cf . Fig. 52 ). However, width of the interorbital region is similar to typical brachypterous females (0.32× head width) and there are mostly dark setae on the mesopectus dorsally as for most brachypterous females. Brachypterous females also tend to have a generally more distinctly green head with less extensive or distinct reddish-violaceous lusters ( Figs 47 , 55 ), though this is not true for the syntype ( Fig. 49 : insert) or the similar brachypterous female from Bugaba. The propodeum of brachypterous females usually also have the anteromesal sculptured callar region quite extensively though inconspicuously setose ( Fig. 58 ). Macropterous females more commonly have the head more extensively reddishviolaceous ( Figs 46 , 54 ) other than for the lower face and, like females of other species, the anteromesal sculptured callar region usually is setose only near its anterior margin to about the level of the posterior margin of the spiracle. This latter, variable setal feature is not described for the different species because it only differs within L. testaceus , though females of all species have the callar region variably densely setose along its lateral margin. Based on observed patterns of variation I currently interpret the differences discussed above as intraspecific variation within a single species rather than brachypterous females and macropterous females representing two separate species. FIGURES 54–59. Lutnes testaceus (Cameron) , ♀. 54 & 55 , head and mesosoma, dorsal: 54 , 2017-11 (m); 55 , 2017-10 (b). 56 & 57 , vertex and anterior of mesosoma, lateral: 56 , 2017-11 (m); 57 , 2017-8 (b). 58 , propodeum (2017-9, b). 59 , (2017-10, b) [b = brachypterous, m = macropterous]. Abbreviations: car = callar region, plr = plical region, psg = postspiracular groove. Body color pattern is sufficient to differentiate L. testaceus from other Lutnes females, uniquely having the meso- and metasoma mostly orangish-yellow in distinct contrast to a dark head, other than for a single macropterous female from Ecuador (collected July 21, 1994 , lot 693) that has an entirely dark mesoscutum ( Figs 48 , 54 ). The legs are also orangish-yellow similar to the mesosoma except for the inner surface of the metafemur having a longitudinal dark streak ( Fig. 51 ), sometimes the meta- and/or mesotarsi, often the outer surface of the metatibia over about its dorsal half, and sometimes the mesotibia subbasally being dark. The only other Lutnes species having orangish lanceolate setae on the fore wing disc is L. aurantimacula , but females of this species are distinguished by the features given in the key and as discussed under that species.