Eurydinoteloides Girault

Eurydinoteloides Girault, 1913a: 55 . Type-species: Eurydinoteloides americana Girault, by monotypy (Ƥ holotype, ZMHB, examined). Gender: feminine (ICZN: 30.1.4.4).

Aeronea Cameron, 1913: 127–128. Type-species: Aeronea laticeps Cameron, by monotypy (Ƥ syntype, BMNH, examined). Junior homonym of Aeronea Agassiz (1846) discovered by Ghesquière (1946: 370). Synonymy by Bouček (1988: 418). Aeroneisca Ghesquière, 1946: 370. Replacement name for Aeronea Cameron nec Agassiz. Synonymy by Bouček (1993: 1250). Protolaccus Burks, 1954: 15 –16. Type-species: Neocatolaccus syrphidis Girault (1916), by original designation (Ƥ and 33 syntypes, USNM, examined). Synonymy by Bouček (1988: 418).

Included species. Eurydinoteloides americana Girault (1913), E. bacchadis (Burks 1954), E. dymnus (Walker 1847) n. comb., E. eudubia (Özdikmen 2011) n. comb., E. hermeas (Walker 1847) n. comb., E. incerta (Ashmead 1893) n. comb., E. laticeps (Cameron 1913), E. orontas (Walker 1847) n. comb., E. perdubia (Girault 1916) n. comb., E. platensis (De Santis in De Santis et al. 1979) n. comb., E. syrphidis (Girault 1916), E. tepicensis (Ashmead 1895) n. comb., E. timaea (Walker 1847) n. comb., E. tortricidis (Crawford 1921) n. comb.

Diagnosis. Metapleuron variable, but at least partly smooth and shiny and with anterior margin raised at least slightly above level of posterior margin of mesopleuron (e.g. Figs 114–116, 137–142). Flagellum of female with 3 basal ringlike articles lacking mps and 5 funicular articles with mps (cf. Fig. 163); male with third flagellomere usually more closely resembling subsequent funiculars than basal two flagellomeres (Figs 129–132), but variable in length and setation, and with (Figs 131, 132) or without (Figs 129, 130) mps. Fore wing with length and number of rows of admarginal setae and extent of speculum variable (Figs 143–147). Head with or without distinct malar depression (Figs 133–136). Head and mesosoma dark or with variably distinct metallic lustres and almost always with variably conspicuous white setae on at least head of male and on head and mesosoma of female (e.g. Figs 119–122). Propodeum with (Figs 152, 154) or without (Figs 124, 156) plical carina posteriorly within nuchal furrow and with or without complete or partial costula, but without transverse carina within paraspiracular region. Both mandibles with four similar teeth (Figs 111, 133, 136).

Description. Head and mesosoma brown, black or variably distinctly metallic green to blue or red, and usually with comparatively long, slender, almost parallel-sided white setae (Fig. 125) at least on head (some males) or both head and mesosoma (females and most males), though setae rarely conspicuously widened over about basal two thirds and abruptly narrowed to point in apical third (cf. Fig. 171) or very rarely (Neotropical species) with white setae on mesoscutum and brown setae on scutellar-axillar complex or all setae dark and hairlike. Eye bare or at least superficially bare with at most exceedingly short, sparse, inconspicuous setae. Mandibles quadridentate, with four similarly acute teeth (Figs 111, 133, 136). Head in frontal view transverse-oval to subcircular (Fig. 111); antenna inserted above lower margin of eyes near middle of face; tentorial pits not evident; clypeus with ventral margin virtually transverse to variably distinctly incurved and variably conspicuously vertically striate to coriaceous; face usually shallowly meshlike reticulate, without tiny bump or smoother and shinier spot at ocular margin at midheight of eye. Head in dorsal view abruptly declined immediately behind posterior ocelli, hence strongly transverse with vaulted vertex. Head in lateral view with or without malar depression extending up to about half distance to lower orbit (Figs 133–136); malar space about 0.4–0.66× height of eye in female, but sometimes only about 0.3× in male; with or without malar sulcus/line. Antenna with scape most often extending dorsally only to about level of anterior ocellus, but rarely in female attaining and sometimes in male exceeding (Fig. 131) level of vertex; flagellum of female with 3 anelli and 5 funiculars with mps (cf. Fig. 163); flagellum of male with at least 2 very strongly transverse and similarly ringlike anelli (Figs 129–132) but third flagellomere variable in length, setation, and most often with reduced number (Fig. 131) or without (Figs 129, 130) mps so as to variably closely resemble subsequent flagellomeres; clava with apical clavomere uniformly conical without micropilose sensory region; flagellum of male usually with compacted articles, but sometimes at least apical flagellomeres separated by short tubular extension.

Pronotum with collar abruptly margined to steeply angled collum, but without carina. Mesonotum meshlike reticulate; mesoscutum with incomplete notauli; scutellum without frenum, in lateral view almost flat and in same plane as dorsal margin of metanotum or variably distinctly convex with apex curved to vertical posterior margin above metanotum (Fig. 121) or very rarely in lateral view with marginal rim reflexed posteriorly as short denticle (Fig. 120). Fore wing (Figs 126–128, 143–147) hyaline; marginal vein not thickened and about 1.3–2.2× length of stigmal vein and about 1.0–1.5× length of postmarginal vein; stigma small, not distinctly capitate; costal cell comparatively narrow, without setae dorsally but variably setose ventrally, usually with 1 or 2 rows of setae within at most about apical third but some Neotropical species with 1 or 2 setae also basally or rarely setose along length; basal cell, basal fold and mediocubital fold usually bare but some Neotropical species with setae on basal fold or apically within basal cell (Fig. 147); disc ventrally usually with 1 transverse or 2 irregular or partial rows of admarginal setae that often are longer than dorsal discal setae (Fig. 146), though sometimes in up to 3 or 4 partial rows (Fig. 145) and/or not noticeably longer than dorsal setae or hyaline and then less conspicuous in some species (Fig. 144); disc dorsally usually uniformly setose beyond speculum (Figs 143, 147), but rarely setae reduced in density and length and then sometimes marginal fringe absent (Fig. 144); speculum usually extending to level about equal with middle of stigmal vein such that admarginal setae completely exposed (Figs 127, 128, 143–146) though some species south of USA with discal setae partly to completely (Fig. 147) overlying admarginal setae behind marginal vein. Metapleuron at least partly smooth and shiny and with anterior margin raised at least slightly above level of posterior margin of mesopleuron, but otherwise variable, sometimes only slightly angulate anteromesally and then margin variably distinctly sculptured (Figs 114–116, 137, 138) or with margin uniformly curved and smooth and then either recurved outward at abrupt angle relative to mesopleuron (Figs 141, 142) or with convex surface extending variably conspicuously over posterior margin of mesopleuron (Figs 48, 139, 140). Metacoxa bare dorsobasally; metatibia with single tibial spur. Propodeum with vertical carina or angulation posterolaterally (cf. Fig. 103: arrow 1), the ridge extending through furrow that differentiates metacoxal flange above metacoxa and in dorsal view variably distinctly angulate so as sometimes to project posteriorly to posterolaterally as short denticle (Fig. 151); with Λ-shaped to transverse-rectangular, coriaceous-reticulate nucha, the furrow delineating nucha without longitudinal carinae except usually for posterior continuation of median carina and sometimes plical carinae; plica at least indicated by outer margin of anterolateral plical depression and often with plical carina within nuchal furrow or variably conspicuous, sinuate convexity or carina posterior of depression (Figs 151–154); paraspiracular region without transverse carina; median region meshlike coriaceous-reticulate, usually convex and often with transverse carina crossing median carina, hence often appearing +-like (Figs 151, 152) or more distinctly raised medially so as to be transversely angulate and sometimes with angulation (Figs 123, 124) extending partly or entirely between plica and median carina at about midlength to differentiate variably distinctly concave anterior panel from posterior convex panel, though angulation not distinctly carinate (cf. Figs 158, 160).

Gaster of female (Figs 119–122) variably elongate-lanceolate with hypopygium extending at most about twothirds length of gaster; gaster of male usually with pale or at least lighter brown region basally; petiole very short, transverse, smooth and shiny, and not braced ventrally by extension of first gastral sternite; cercal setae all of similar length.

Distribution. New World.

Hosts. See Noyes (2012) for included species listed above.

Generic synonymy. Girault (1913a) stated that the minutien-pinned holotype of E. americana lacked the gaster and had the head, hind legs and antennae on a slide. I did not examine the parts on the slide, but did examine the remaining mesosoma of the holotype (Fig. 117). The left hind wing, apex of the left fore wing, and both right wings are missing, as are the legs except for the left front and middle legs. The original description states that it was “olive green, the abdomen and metathorax metallic”, but the thorax and propodeum medially are black and there is only a very slight greenish lustre elsewhere. The colour of the tegula and remaining legs is as described, though the “brown” femora may have faded over time, in particular the mesofemur being only slightly darker, more orange than brown. The metapleuron has only a comparatively small smooth and shiny region anteromesally, being quite distinctly reticulate dorsally, posteriorly and ventrally, with the anterior margin angulate and raised only very slightly above and over the posterior margin of the mesopleuron (Fig. 118). The propodeum has distinct plical carinae extending posteriorly to the nucha and a slightly sinuate, fine but complete, ridge-like costula dividing the plical region into anterior and posterior panels (Fig. 123). Although artefacts obscure setal pattern of the remaining left fore wing, the speculum extends to about the basal third of the stigmal vein (single broken seta within speculum near apex of marginal vein) and the dorsal discal setae are not reduced in length or density. There are 17 admarginal setae (Fig. 126: circles) arranged into 2–3 indistinct rows (most obviously behind about basal half of marginal vein) and the admarginal setae are not conspicuously longer than the discal setae (Fig. 126: rectangles).

Bouček (1988: 418) did not provide any explanation for his synonymy of Aeronea and Protolaccus under Eurydinoteloides except that his actions were based on study of the type specimen of “ E. americanus Girault ”1. Bouček (1993: 1250) was more explicit in stating that the synonymy was based on “notes and sketches from the headless holotype ”, suggesting that he also did not see the slide with the head. Girault (1913a) did not mention the presence of a distinct malar depression in the original description of E. americana . However, Burks (1954) established Protolaccus for two species with an arch-like malar depression, E. bacchadis (Burks 1954) (Fig. 112) and E. syrphidis (Girault 1916) . The type species of Aeronea, E. laticeps (Cameron 1913), also has an arch-like malar depression (Fig. 113). Eurydinoteloides bacchadis, E. syrphidis and E. laticeps are all very similar to each other (cf. Figs 111–116, 119, 120, 127, 128), but differ from E. americana in several features, including fore wing setal pattern and propodeal structure. Type females of the three putative species have the discal setae reduced in length and density behind the marginal vein and there is only a single row of comparatively long admarginal setae behind the length of the marginal vein (Figs 127, 128). Further, the propodea lack plical carinae posterior to the anterolateral plical depression and the costulae are directed mesally from the posterior margin of each depression toward the median carina so that paraspiracular regions and relatively large posterior plical panels form a continuous U-shaped region (Fig. 124). Based on the above observations, I consider it likely that E. americana has an arch-like malar depression similar to E.

1. Bouček (1993) emended americana to americanus because based on Article 30(b) of the 1985 International Code of Zoological Nomenclature “ oides ” is masculine. However article 30(b) was superseded by Article 30.1.4.4. of ICZN (2000), which states that “ oides ” is to be treated as masculine unless the author, when the name was established, “stated that it had another gender or treated it as such by combining it with an adjectival species-group name in another gender form”. Consequently, based on current ICZN rules, Eurydinoteloides should be treated as feminine.

bacchadis, E. laticeps and E. syrphidis, and therefore concur with the generic synonymy proposed by Bouček (1988). Further, although E. americana represents a distinct species, E. bacchadis, E. laticeps and E. syrphidis are all very similar except for colour of the femora. The type specimens of E. bacchadis have yellow femora (Fig. 120), whereas E. laticeps has brown femora (Fig. 119) similar to E. syrphidis . It is therefore very possible that at least E. syrphidis is a junior synonym of E. laticeps, but confident synonymy is not possible until the Neotropical species of Eurydinoteloides are revised and leg colour can be assessed as a species character.

Discussion. Males of E. americana are not currently recognized, but those of E. bacchadis and E. syrphidis are atypical for Eurydinoteloides in having a comparatively long scape that extends to or exceeds the level of the vertex (Fig. 131) rather than just to the ventral margin of the anterior ocellus as for most species of the genus. This feature is correlated with six elongate funiculars with sparse mps, including on the third flagellomere (Fig. 131). However, males of an undescribed species from Florida with a distinct malar depression (Fig. 133) have six shorter, more robust funiculars encircled by mps (Fig. 132) as well as a more typical scape that extends only to about the level of the anterior ocellus. Further, males of a few other Neotropical Eurydinoteloides species without a malar depression have a scape that extends to the level of the vertex. Consequently, presence or absence of an elongate scape in males does not appear to correlate with presence or absence of a malar depression.

FIGURES 143–150. Ƥ fore wing. 143–147, Eurydinoteloides spp.: 143, (61); 144, (179); 145, (48); 146, (177), 147, (119). 148, Heteroschema sp. (149). 149, Neocatolaccus tylodermae (178). 150, Neocatolaccus sp. (125). No. in parenthesis = CNC 2011 photo no.

Development of a malar depression is highly variable in Eurydinoteloides . Some species have a distinct, archlike depression that extends about half way (Figs 112, 113, 133) or somewhat less (Fig. 134) to the lower orbit, others have the gena abruptly incurved only above the base of the mandible as a very short, transverse depression along the oral margin (Fig. 135), and still others lack an evident depression (Fig. 136). Development of the metapleuron is similarly variable. As noted above, individuals of E. americana, E. bacchadis, E. laticeps and E. syrphidis all have an anteromesally angulate metapleuron with the margin finely sculptured and raised only slightly above the posterior margin of the mesopleuron, and the surface with only a comparatively small smooth and shiny region (Figs 114–116, 118). In such instances, careful observation is necessary to observe the difference in structure from that of Lyrcus, in which the metapleuron is uniformly, though sometimes very faintly sculptured, and the anterior margin abuts the posterior margin of the mesopleuron on the same level (Figs 19, 174). Further, some Eurydinoteloides species without a broad malar depression have a similarly angulate metapleuron (Figs 137, 138). Others, with or without a distinct malar depression, have the metapleuron extensively smooth and shiny with the anterior margin evenly curved, unsculptured, and either convex and extending slightly over the posterior margin of the mesopleuron (Figs 48, 139, 140) or, less commonly, recurved outward at abrupt angle relative to the mesopleuron so that the latter abuts the inner surface (Figs 141, 142). Similar to presence or absence of a malar depression, the different metapleural structures appear to form a continuum and neither character appears to be valid for the recognition of subgenera within Eurydinoteloides . Based on a greater similarity to the metapleuron of most pteromalids (Figs 19, 27, 28, 174, 212, 213), the only slightly angulate metapleural structure of such species as E. americana likely is the groundplan structure for Eurydinoteloides . The groundplan structure of the gena, that is, with or without an arch-like malar depression, is more difficult to propose confidently.

As noted above, the admarginal setae are completely exposed and similar in length to the dorsal discal setae in E. americana (Fig. 126). All North American and most Neotropical species of Eurydinoteloides are characterized by one or two rows of clearly exposed but obviously longer admarginal setae (Figs 127, 128, 143, 146). However, a relatively few species from south of the USA do not have the admarginal setae differentiated in length from the dorsal discal setae and/or have up to three or four partial rows of admarginal setae (Fig. 145) and/or the dorsal discal setae extending variably extensively along the length of the marginal vein, very rarely to its base, so as to partly or completely obscure the admarginal setae (Fig. 147). Such species often also have the fore wing more extensively setose than for typical species, including one or more of the costal cell, basal fold, basal cell, and the triangular region between the stigmal and postmarginal veins having setae (Fig. 147). Although these latter setal patterns are relatively uncommon, they make confident hypothesis of the groundplan fore wing setal pattern difficult.

The male flagellar structure of the undescribed species from Florida with two strongly transverse anelli and six similar funiculars encircled by mps (Fig. 132) could represent the groundplan structure for Eurydinoteloides (see further under Lyrcus). However, males of most Eurydinoteloides species have structures of the third flagellomere that are intermediate between a typical funicular and an anellus. The mps can be reduced in number (Fig. 131) or completely lacking (Figs 129, 130), though the flagellomere usually is variably longer and more setose than the basal two anelli so as to more closely resemble the subsequent flagellomeres than the basal two anelli. If such males were interpreted as having only two anelli then they would key to Lyrcus at couplet 263 using Bouček and Heydon (1997). However, rather than keying Lyrcus, this couplet differentiates mostly Jaliscoa species. Eurydinoteloides females and those males with the third flagellomere interpreted as an anellus will key either to Eurydinoteloides (couplet 286) or Lyrcus (couplet 287) depending, respectively, on the presence or absence of a distinct malar depression.

FIGURES 163–169. Antenna. 163 and 164, Lyrcus nigroaeneus: 163, Ƥ (187); 164, 3 (172). 165–167, Lyrcus spp.: 165, Lyrcus sp. Ƥ (186) [insert: fl1–fl5]; 166, Lyrcus sp. 3 (138) [insert: pedicel–fl4]; 167, L. nigraeneus 3 (137) [insert: pedicel–fl3]. 168 and 169, Oaxa albiclava antenna: 168, Ƥ (118) [insert: fl1–fl4]; 169, 3 (140) [insert: pedicel–fl4]. No. in parenthesis = CNC 2011 photo no.

Although the presence of lanceolate, more or less parallel-sided white setae that contrast conspicuously with the cuticle (Fig. 125) certainly is not unique to Eurydinoteloides among pteromaline genera, such setae differentiate at least North American females of Eurydinoteloides from most Lyrcus species. Most Lyrcus lack contrasting setae or, if there are flattened white setae, then they usually are conspicuously widened over about the basal two-thirds and narrowed abruptly to a point in the apical third (Fig. 171) as well as typically originating from small blue to green spots (Figs 170, 173). However, L. cyaneus (Figs 14–16) as well as at least two other North America species have slender white setae (Fig. 175) similar to typical Eurydinoteloides . Rare Eurydinoteloides also have more conspicuously lanceolate setae, indicating this feature is homoplastic. Under high magnification, both types of flattened setae have a median rib-like thickening whether they are parallel-sided (Fig. 125) or more broadly lanceolate (Fig. 171), but studies of flattened setae in other genera have not been conducted to determine whether this similarity has any phylogenetic significance or is simply a functional feature. I also observed two females of Eurydinoteloides from Guatemala (CNC) with dark hairlike setae, and females and males from Costa Rica, Ecuador and Trinidad (CNC) that have white, slightly lanceolate setae on the head and mesoscutum, but brown setae on the scutellum similar to some species classified in Lariophagus . Males of Eurydinoteloides also sometimes lack conspicuous setae, having only very slender, more hairlike and comparatively inconspicuous white setae on the face and often laterally on the pronotum. Such males are easily mistaken for Lyrcus males if metapleural structure is not visible.

The propodeum of Eurydinoteloides sometimes has a variably distinctly differentiated transverse angulation or ridge across the plical region (Figs 123, 124, 151, 152) that is very similar to some species of Jaliscoa (Figs 71, 84, 96). Although then evident as a costula, the transverse ridge is usually less distinct than the more carinate costula that characterizes some other genera with conspicuous white setae as well as three anelli in females, such as Heteroschema (Fig. 158: cos) and Neocatolaccus (Fig. 160). In addition to having a plesiomorphic metapleural structure, both sexes of these latter two genera are characterized by two metatibial spurs and/or a comparatively long scape that extends to or beyond the level of the vertex (Fig. 157), though the scape is not always long and two metatibial spurs are not always evident. The inner metatibial spur is often very short and inconspicuous, and apparently even missing from some Heteroschema . At least described North American species of Heteroschema have the left mandible tridentate (Fig. 159), whereas Neocatolaccus have both mandibles more or less quadridentate (sometimes dorsal two teeth appearing more like a truncation, Fig. 161) similar to Eurydinoteloides (Fig. 111), Lyrcus and Jaliscoa (Fig. 87). Whether or not the different mandibular structures of Neocatolaccus and Heteroschema is a valid generic difference is questionable because I am uncertain of the limits of the two genera. Heteroschema differs from Neocatolaccus primarily by the presence of a distinct nucha (cf. figs 158, 160), but species with intermediate structures exist that might be assigned to either genus (see further under Lyrcus).