Lyrcus Walker

Gibson, Gary A. P., 2013, Revision of the species of Jaliscoa Bouček within a review of the identity, relationships and membership of Jaliscoa, Catolaccus Thomson, Eurydinoteloides Girault, Lyrcus Walker and Trimeromicrus Gahan (Hymenoptera: Pteromalidae), Zootaxa 3612 (1), pp. 1-85 : 63-71

Lyrcus Walker

Lyrcus Walker, 1842: 114 –115. Type-species: Lyrcus origo Walker , by monotypy (Ƥ lectotype, BMNH, examined). Gender: masculine.

Zatropis Crawford, 1908: 159 . Type-species: Zatropis catalpae Crawford , by original designation (Ƥ holotype, USNM, examined). Synonymy by Bouček (1993: 1250).

Oluspa Cameron, 1913: 129. Type-species: Oluspa albopilosella Cameron, by monotypy. (Ƥ lectotype, BMNH, examined). Synonymy by Bouček (1993: 1250).

Neocylus Bouček, 1988: 235 , 420. Type-species: Polycysteloides nigraeneus Girault (1913b) , by original designation (Ƥ holotype, QMBA, examined). N. syn.

Included species. L. albiclavus (Girault 1917c) n. comb., L. albopilosella ( Cameron 1913) , L. capitis (Burks 1955) n. comb., L. catalpae (Crawford 1908) , L. chalcis (Burks 1955) n. comb., L. coeliodis (Ashmead 1896) n. comb., L. cyaneus (Girault 1911) n. comb., L. deuterus (Crawford 1911) n. comb., L. golbachi ( De Santis in De Santis & Armesto 1983), Lyrcus helice (Walker 1843) n. comb., L. justicia (Girault 1917) , L. nigraeneus (Girault 1915) n. comb., L. nigroaeneus (Ashmead 1894a) n. comb., L. origo Walker 1842 , L. rosaecolis (Burks 1955) n. comb.

Diagnosis. Metapleuron completely though often finely sculptured and with anterior margin abutting posterior margin of mesopleuron on same level ( Figs 19 , 174 ). Flagellum of female with at least 3 basal ringlike articles lacking mps and at least 4 (usually 5) funicular articles with mps (Fig. 163); male of New World species with third flagellomere lacking mps and usually noticeably shorter than fourth flagellomere, but variable in length and setation (Figs 164, 166). Fore wing usually with 1 ( Figs 182, 183 ) or sometimes 2 ( Fig. 185 ) or extremely rarely 3 ( Fig. 184 ) partial rows of admarginal setae that usually are quite obviously longer than dorsal setae and most often at least partly exposed because of extensive speculum, though extent of speculum variable ( Figs 182–187 ). Head ( Figs 14 , 35 , 173 ) only extremely rarely with distinct malar depression, though sometimes narrowly depressed along base of mandible. Head and mesosoma brown to black or with variably distinct metallic lustre, sometimes as numerous small green to blue spots from which setae originate ( Figs 171, 173 ), and with setae short, brown and hairlike (Figs 176–179) to white and variably long and broadly lanceolate so as to then contrast conspicuously with cuticle ( Figs 16 , 170–173, 175 ). Propodeum with plical carina at least posteriorly within nuchal furrow ( Figs 20 , 36 , 190–197 ) and often with transverse carina on paraspiracular region or within paraspiracular furrow ( Figs 190–194, 196 : psc), but only rarely with complete costula ( Figs 190, 191 ). Both mandibles with four similar teeth.

Description. Head and mesosoma uniformly brown to black or dark with variably distinct metallic blue or green lustre ( Figs 15 , 33, 34 , 176–179) or with metallic green to blue spots from which broadly lanceolate setae arise ( Figs 171, 173 ); head and mesosoma with setae most commonly comparatively inconspicuous, short, brown and hairlike ( Figs 33, 34 , 176–179), but sometimes white and then usually elongate-slender ( Figs 16 , 175 ) to broadly lanceolate ( Figs 170–173 ) and contrasting conspicuously with cuticle. Eye bare or at least superficially bare with at most exceedingly short, sparse, inconspicuous setae. Mandibles quadridentate, with four similarly acute teeth or with three ventral teeth and rounded dorsal lobe or narrow truncation, but dorsal dent separated from third tooth by narrow, acutely angled incision. Head in frontal view transverse-oval to subcircular; antenna inserted obviously above lower margin of eyes near middle of face; tentorial pits not evident; clypeus with ventral margin usually transverse to slightly incurved, but very rarely bidentate, and variably conspicuously vertically striate; 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 ( Figs 14 , 35 , 173 ) or without distinct malar sulcus/line, but only extremely rarely with distinct malar depression; malar space about 0.15–0.6× height of eye. Antenna with scape usually extending dorsally only to about level of anterior ocellus, but rarely to about level of vertex; flagellum of female (Fig. 163) and usually male (Fig. 164) with 3 or very rarely 4 (Fig. 165) transverse basal flagellomeres lacking mps, but third anellus of male sometimes similar in appearance to subsequent flagellomeres except for lack of mps (Fig. 166) and one Australasian species with mps on third flagellomere (Fig. 167); clava with apical clavomere uniformly conical without micropilose sensory region.

FIGURES 176–181. 176–179, Lyrcus spp. Ƥ: 176, dorsal (18); 177, lateral (186); 178, lateral (167); 179, lateral (166). 180, Mesopolobus verditer Ƥ, lower face and mandibles (141). 181, Mesopolobus sp., Ƥ habitus, lateral (142). No. in parenthesis = CNC 2011 photo no.

Pronotum with collar usually smoothly to abruptly margined relative to steeply angled collum, but sometimes separated by smooth, shiny carina. Mesonotum meshlike reticulate to coriaceous-reticulate in small specimens; 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, but not reflexed into distinct marginal rim. Fore wing usually hyaline ( Figs 17 , 182 ) but one known species with brownish region behind parastigma and base of marginal vein (Fig. 179); marginal vein not thickened and about 1.2–2.3× length of stigmal vein and about 0.9–1.5× length of postmarginal vein; stigma small, not distinctly capitate; costal cell comparatively narrow, without setae dorsally but variably setose ventrally, most often with row of setae only within about apical third but sometimes with setae apically and basally or with row along entire length; basal cell, basal fold and mediocubital fold often bare but sometimes with up to 2 lines of setae along basal fold; disc ventrally usually with 1 transverse or 2 partial or irregular rows of comparatively long admarginal setae relative to length of dorsal setae ( Figs 182, 183 ), though extremely rarely in up to 3 partial rows ( Fig. 184 ) and sometimes not distinctly differentiated in length from dorsal setae; disc dorsally uniformly setose beyond speculum; speculum extending at least to base of marginal vein ( Figs 18 , 186, 187 ) and usually partly ( Figs 184, 185 ) or completely ( Figs 182, 183 ) to stigmal vein. Metapleuron completely though often finely sculptured, and with anterior margin on same level as and abutting mesopleuron ( Figs 19 , 174 ). Metacoxa bare dorsobasally; metatibia with single tibial spur. Propodeum usually with vertical carina or stronger flange posterolaterally on callus (cf. Fig. 103 : arrow 2), though in dorsal view only very rarely projecting laterally as tiny denticle ( Fig. 197 : arrow); usually with transverse-rectangular or more or less Λ-shaped coriaceous-reticulate nucha ( Figs 190–194 ), though propodeum sometimes variably strongly transverse ( Figs 194–196 ) and then sometimes comparatively smooth and shiny ( Fig. 196 ) with posteromedian region delineated anteriorly by anteromesally convergent ridges or carinae so as to more or less resemble an adpetiolar strip ( Figs 195, 196 ), the convergent carinae rarely extending almost to anterior margin or, more commonly, forming inverted Y-shaped carinal complex in combination with median carina ( Fig. 196 ); plica at least indicated by outer margin of anterolateral plical depression and carina within nuchal furrow, and if complete then usually quite distinctly sinuate at about midlength ( Figs 190–193, 197 ); plical region usually reticulate to coriaceous-reticulate, with or without median carina, and sometimes with transverse carina crossing median carina (then appearing +-like) or raised medially so as to be transversely angulate, but only rarely with complete costula between plica and median carina ( Figs 190, 191 ); paraspiracular region sometimes with transverse carina in paraspiracular furrow ( Figs 193, 194, 196 : psc) or extending mesally to plica ( Figs 190–192 : psc).

Gaster of female lanceolate (Figs 176–179), variable in length relative to combined length of head and mesosoma with hypopygium extending at most about two-thirds length of gaster; gaster of male (at least Nearctic species) usually without evident paler 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 and Australasian region ( Australia, Papua New Guinea).

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

Discussion. Bouček (1988) described his new genus Neocylus based on Polycysteloides nigraeneus Girault (1913b) from Australia. In his key to the genera of Pteromalidae he keyed it and compared it to Neocatolaccus (couplet 224) because of the presence of a costula (couplet 221). He also stated that there were two species of Neocylus . I examined the type specimens of N. nigraeneus and its junior synonym N. reticulata (Dodd in Girault 1915) (QMBA), plus four females from Australia (two each in BMNH and QMBA) and four females and two males from Papua New Guinea (BMNH) identified as Neocylus by Bouček. Based on a manuscript name on one of the females, he considered the New Guinea specimens as the second species. This was likely because of a tibial colour difference, but based on additional specimens I believe the New Guinea and Australian specimens comprise a single species. All the specimens have a carinate costula ( Fig. 190 : cos) that is continuous with a transverse carina on the paraspiracular region ( Fig. 190 : psc). The paraspiracular carina extends over the callus about midway between the spiracle and posterior margin of the propodeum to more or less connect with the posterolateral, vertical carina on the callus (cf. Fig. 103 : arrow 1). The propodeum also has a distinct nucha delimited laterally by parallel posterior portions of complete, strongly sinuate plical carinae. Posterior to the costula the median carina variably distinctly bifurcates so that anteriorly the nucha is delimited by an inverted Y-shaped carinal complex ( Fig. 190 ), which is somewhat intermediate in structure between a typical nucha and a typical adpetiolar strip. Females are similar to New World Lyrcus in all other features, including having three anelli, quadridentate mandibles, an unmodified metapleuron, and the speculum extending along the marginal vein so that one to two rows of admarginal setae are clearly exposed behind at least the basal half of the marginal vein ( Fig. 185 ). The admarginal setae usually are obviously longer than the discal setae, though they are only about as long as the discal setae in small specimens.

The two Neocylus males have a similar propodeal sculpture pattern and fore wing setal pattern as for females, except a few setae on the basal fold differentiate the apex of the basal cell from the speculum. More conspicuously different, males have only two ringlike anelli and six elongate funiculars that are similar in length, setation, and presence of mps (Fig. 167). All known New World males of Lyrcus lack mps from the third flagellomere, which usually (Fig. 164) resembles that of females (Fig. 163), though sometimes it more closely resembles the subsequent flagellomeres except for the absence of mps (Fig. 166). Males of one such species from Ecuador (CNC) are associated with females that have quite a similar propodeal sculpture pattern to N. nigraenus (cf. Figs 190, 191 ).

Though somewhat variable (apparently correlated with body size), at least larger females have a distinct, carinate costula ( Fig. 191 : cos) that extends more or less completely to the plical carina as well as a transverse carina on the paraspiracular region ( Fig. 191 : psc) that is continuous with the plical carina. Both sexes of this species also have the speculum extending only to the base of the marginal vein such that the dorsal discal setae overlie a mostly single row of admarginal setae that are obviously longer than the discal setae ( Fig. 186 ). Other New World Lyrcus I have seen lack a complete, carinate costula, though some species have an incomplete costula ( Fig. 192 : cos) or transverse ridge across the plical region and/or a paraspiracular carina ( Figs 193, 194, 196 : psc) within the paraspiracular furrow or even across the paraspiracular region from the plical carina ( Fig. 192 : psc). I interpret the propodeal sculpture patterns illustrated by figures 190–193 to constitute a structural transformation series, though with uncertain polarity. A complete, carinate costula that is continuous with a paraspiracular carina ( Figs 190, 191 : cos) could be apomorphic and indicate a sister-group relationship between N. nigraeneus and the species from Ecuador. Equally possible is that it is a symplesiomorphic pattern from which other New World Lyrcus propodeal sculptural patterns were derived through reduction and secondary loss of the costula and, sometimes, the paraspiracular carina ( Figs 190–194, 196 : psc). The other principal genera treated in this work ( Catolaccus , Jaliscoa , Eurydinoteloides and Trimeromicrus ) lack a paraspiracular carina. However, almost all Neocatolaccus ( Fig. 160 ) and at least some Heteroschema ( Fig. 158 ) possess both a complete costula and a paraspiracular carina. If a paraspiracular carina is part of the groundplan propodeal sculpture pattern for Lyrcus , then it suggests a possible common ancestor for Lyrcus and one or both of these latter two genera. Both Neocatolaccus and Heteroschema are further characterized by the presence of three anelli in females and two anelli in males similar to N. nigraeneus and, as discussed above, both mandibles of Neocatolaccus are more or less quadridentate ( Fig. 161 ). I have also seen a single female (CNC) of an undescribed species of Neocatolaccus from Alabama ( USA) that could easily be mistaken for a species of Lyrcus except for the presence of slender white setae (but also possessed by rare Lyrcus ) and the presence of two metatibial spurs. This female is comparatively small for a Neocatolaccus , only about 2 mm in length, and perhaps because of its small body size lacks a complete costula ( Fig. 162 ). There is an angulation across the plical region so that indistinct anterior and posterior panels are differentiated, but the angulation is carinate only laterally near each plica and is continuous with a somewhat stronger carina on the paraspiracular region ( Fig. 162 : psc). Unlike the fore wing of other known Neocatolaccus (Fig. 149), there is only a single row of admarginal setae (Fig. 150), though like other species these are covered by dorsal discal setae over about the apical half of the marginal vein and they are only as long as the discal setae (Figs 149, 150). Species of Lyrcus with either slender or broadly lanceolate white setae display the typical fore wing setal pattern for Lyrcus , that is, only a single row of conspicuously long and completely exposed admarginal setae ( Figs 182, 183 ).

The features discussed above may or may not reflect a relationship among Heteroschema , Neocatolaccus , Neocylus nigraeneus and New World Lyrcus . However, the only feature differentiating Neocylus and New World Lyrcus is the presence of two anelli in males of N. nigraeneus and three anelli in males of Lyrcus . Males of Eurydinoteloides are highly variable in structure of the third flagellomere, some species lacking mps from two and others from the basal three flagellomeres, and structure of the third flagellomere varies from more or less anelliform to elongate and similarly setose as more distal flagellomeres ( Figs 129–132 ). Structure of the third flagellomere also varies in New World Lyrcus (Figs 164, 166), though mps are always lacking. I do not consider presence or absence of mps on the third flagellomere in males sufficient for maintaining Neocylus and Lyrcus as separate genera. I therefore synonymize Neocylus under Lyrcus and consider two anelli in males as the likely groundplan structure for Lyrcus . Propodeal sculpture pattern and the different number of anelli between the sexes may indicate L. nigraeneu s as the basal lineage of Lyrcus , and represent symplesiomorphies that are shared with some other genera such as Neocatolaccus and Heteroschema . If so, Lyrcus did not originate in the New World as thought previously.

Oaxa Bouček also shares some important differential features with Lyrcus , including quadridentate mandibles and three anelli in females (Fig. 168). The fore wing setal pattern is also similar to some Lyrcus (e.g. Fig. 186 ), having one to two closely set rows of admarginal setae that, although largely overlain by a band of discal setae along the marginal vein, are conspicuously longer than the discal setae ( Fig. 188 ). When Bouček (1993) described O. albiclava , supposedly from a single female from Mexico, he stated that the species “may be regarded as similar to certain small Lyrcus ”. This comment probably was based at least in part on the similar fore wing setal patterns. Bouček (1993) differentiated Oaxa based on large eyes (malar space 0.26× eye height), rather deep scrobes, and a deeply emarginate clypeus (Bouček 1993, fig. 64), cercus with one curved seta more than twice as long as any other seta (Bouček 1993, fig. 65), strongly modified pronotum (essentially a vertical collum without horizontal collar, but subdorsally with a fine transverse carina and line of setae differentiating an almost vertical, strongly transverse, shiny collar), and propodeum ( Fig. 203 ) without any trace of a transverse swelling or costula (also with nucha abruptly, Λ-like margined anteriorly so as to somewhat resemble an adpetiolar strip, and paraspiracular region without a paraspiracular carina). Bouček and Heydon (1997) used a combination of these features to key Oaxa in couplet 298 prior to keying Lyrcus in couplet 303. As its name implies, females of O. albiclava also have a white clava (Fig. 168). In addition to the holotype (stated as deposited in TAMU, but now in USNM), I saw another female (TAMU) collected by the same individuals one day earlier in the same state (Oaxaca) but from a different locality than the holotype. This female is labelled as a “ paratype ” of O. albiclava , apparently by Bouček in 1993, though the original description provides no indication that the type series consisted of more than the holotype. The CNC also has one female, but collected in Chiapas, and TAMU has a long series of the undescribed males of O. albiclava collected from both Oaxaca localities on the same days as the females. Males have quite a different flagellar structure than females (cf. Figs 168, 169), the flagellum being filiform with two ringlike anelli and six similarly long and setose funiculars, the third flagellomere (first funicular) usually having a single mps on either side (Fig. 169). I also saw a single female and eight males from French Guiana (CNC) that are very similar to O. albiclava except the female has superficial coriaceous rather than raised-reticulate mesonotal sculpture, and only about the apical half of the admarginal setae are covered by the dorsal discal setae. Another female from Brazil (CNC) resembles Oaxa in having a white clava, large eyes (malar space about 0.28× eye height), and a similar propodeal structure though the Λ-like posteromedian region is not as abruptly delimited and is slightly more convex. However, the flagellum has only two anelli and six funicular segments, the cercus lacks a conspicuously long seta, the clypeus is only very shallowly emarginate, the fore wing has a single row of conspicuously long and mostly exposed admarginal setae, and the pronotum has a more distinct, horizontal, sculptured collar. Mandibular dentition is not visible. Another female from Ecuador (CNC) with a white clava and even larger eyes (malar space only about 0.17× eye height), has three anelli and a single row of conspicuously long admarginal setae that are completely covered by the dorsal discal setae, but differs from O. albiclava by a having a shallowly incurved clypeus, a distinct, horizontal, sculptured pronotal collar, cercal setae that do not appear to be conspicuously differentiated in length, and a distinct transverse carina within the paraspiracular region. Again, mandibular dentition is not visible. I am uncertain of the generic identity of the Brazilian female because of its flagellar structure, but the female from Ecuador likely belongs to Lyrcus based on its fore wing setal pattern and the presence of three anelli and a paraspiracular carina. Some species of Lyrcus have a lighter coloured clava and I have seen females of an undescribed species from Florida (CNC) with a deeply incised clypeus similar to O. albiclava and with comparatively large eyes, the malar space being only about one-third the height of an eye. It is quite possible that O. albiclava is nothing more than an unusually modified Lyrcus , but if so it is characterized by at least one postulated symplesiomorphy (third flagellomere of male with mps) and one apomorphy (one cercal seta much longer than others). I therefore prefer to retain Oaxa as separate from Lyrcus . Further study of Neotropical pteromalines is necessary to assess more confidently the morphological limits of Oaxa relative to Lyrcus and determine the correct generic placement for the species from Brazil.

Monophyly of Lyrcus remains unsubstantiated and different species intergrade in structure with those of other genera, particularly Eurydinoteloides and Mesopolobus (see below), but also Neocatolaccus . It is therefore uncertain whether white and variably conspicuously widened setae represent apomorphic setal structures derived convergently in some Lyrcus , Eurydinoteloides and several other pteromaline genera, or whether the presence of such setae in only a few species of Lyrcus indicates it is paraphyletic relative to Eurydinoteloides and/or other genera. Dark, hairlike setae that do not contrast conspicuously with the cuticle is undoubtedly symplesiomorphic for Pteromalinae . It perhaps is also the most likely groundplan state for Lyrcus based on commonality within the genus and the setal structure of L. nigraeneus . However, if Lyrcus and Neocatolaccus share a common ancestor, as discussed above, slender white setae ( Fig. 125 ) could be the groundplan setal structure for both Lyrcus and Eurydinoteloides . This would then indicate that the dark, hairlike setae shared by most Lyrcus species represents a secondary reversal, and that the more broadly spatulate white setae ( Fig. 171 ) of rare Eurydinoteloides and species formerly classified in Zatropis represent secondary, apomorphic modifications. If so, Zatropis represents a separate clade from the vast majority of Lyrcus that could be recognized as a separate taxon. However, exact relationships remain obscure and the different setal structures appear to form a continuum. I do not consider the recognition of separate genera or subgenera in Lyrcus warranted at present based on white versus dark or hairlike versus flattenedlanceolate or spatulate setae.

Heydon and Bouček (1992) specifically mentioned Acaenacis as being similar to Zatropis in having flattened white setae. Individuals of Acaenacis could be mistaken for some Lyrcus because they also have three anelli, an unmodified metapleuron, and one or two rows of admarginal setae that typically are mostly exposed although not conspicuously longer than the dorsal discal setae. The propodeum varies in structure in females, often being comparatively strongly transverse, though always with a carinately margined Λ-like adpetiolar strip and usually a median carina such that there is an inverted Y-shaped carinal complex. However, at least one species of Lyrcus from Trinidad (BMNH) with flattened white setae similar to L. nigroaeneus has a similar propodeal structure except the anteromesally convergent median region extends almost to the anterior margin of the propodeum. Heydon and Bouček keyed Acaenacis prior to Lyrcus using several features. Of those listed, the most distinctive is the unusually high placement of the toruli, within the upper third of the face. However, some Heteroschema have the antennae inserted similarly high on the face, particularly males ( Fig. 157 ). Individuals of Acaenacis also have comparatively deep, hole-like anterior tentorial pits, though this is a less obvious feature, and the flagellomeres are always elongate and sometimes quite conspicuously setose. Girault (1917a) described the mandibles of A. taciti (Girault) as being tridentate, but I have not seen specimens with exposed mandibles to confirm this as a valid generic feature.

Because of the absence of mps from the third flagellomere of both sexes of New World Lyrcus , species will key, respectively, to either couplet 303 ( Lyrcus ) or couplet 306 ( Meraporus Walker 1834 and Mesopolobus ) using Bouček and Heydon (1997) depending on whether the admarginal setae are exposed or extensively covered by the dorsal discal setae. Meraporus , like Lyrcus , has quadridentate mandibles, but the head and mesosoma are comparatively bright metallic green rather than dark and the propodeum has a reticulate-rugose to obliquely strigose median region in combination with an adpetiolar strip (cf. Fig. 198 ; Bouček and Heydon 1997, fig. 492). Fully winged individuals also have one or two rows of admarginal setae that are similar in length to and mostly overlain by the dorsal discal setae (cf. Fig. 147). Male Meraporus are differentiated further by a pattern of smooth lines on the head (Bouček and Heydon 1997, fig. 289; Graham 1969, fig. 325). Bouček and Heydon (1997) noted that the mesoscutum (and head) of Meraporus has irregular reticulation because of intermixed shallow setiferous punctures. The head and mesoscutum of Lyrcus are uniformly meshlike reticulate to coriaceous, though species of Mesopolobus classified in the subgenus Xenocrepis Förster (Gibson et al. 2006, fig. 14) are sculptured similar to Meraporus .

As noted above, structure and sculpture of the propodeum is quite variable in Lyrcus ( Figs 190–197 ). There always appears to be a plical carina at least posteriorly within the nuchal furrow, though sometimes this is not obvious in very small males or some individuals with a strongly transverse propodeum. Most species also have a convex, meshlike sculptured nucha ( Figs 190–195, 197 ) and usually the plical carinae are quite strongly sinuate near midlength of the propodeum so to be subparallel posteriorly ( Figs 192, 193, 197 ), though length:width ratio of the propodeum, shape of the plicae (cf. Figs 192, 196 ), and nucha versus an adpetiolar strip (cf. Figs 193, 196 ) are all variable. Correlated with a more strongly transverse propodeum is often more uniformly outcurved plical carinae and a more strongly transverse nucha, which sometimes is delineated anteriorly by a furrow with an abrupt margin ( Fig. 195 ) or even rarely a distinct Λ-like carina and is more or less flattened and/or not meshlike sculptured ( Fig. 196 ) so as to more closely resemble an adpetiolar strip than a convex nucha. Although longitudinal carinae may extend from the anterior margin of the propodeum partly through the plical region ( Figs 192, 194–196 ), the nuchal furrow lacks longitudinal carinae except for the plical carinae and sometimes a median carina ( Figs 190–196 ). Those species with a more transverse propodeum and particularly those with outcurved plicae and the posteromedian region resembling an adpetiolar strip can be mistaken for Mesopolobus , another genus that almost always is characterized by three anelli in both sexes. Most Mesopolobus have quite a smooth and shiny, slender, transverse-triangular or Λ-like margined adpetiolar strip in combination with the plical carinae being quite uniformly out-curved, only slightly sinuate or sinuate only posteriorly at the lateral margin of the adpetiolar strip ( Figs 198–202 ). The plical region is often also more coarsely sculptured than for typical Lyrcus , including some short longitudinal carinae within the nuchal furrow anterior to the adpetiolar strip in addition to the median and plical carinae ( Figs 198–200, 202 ), and/or more extensive oblique carinae, striae or rugosity on the panels ( Figs 198–200 ). However, all of these propodeal features are variable and there is no distinct division between structures possessed by Lyrcus and those by Mesopolobus . A much brighter metallic lustre (Figs 180, 181) is characteristic of many Mesopolobus species, though smaller-bodied individuals are often comparatively dark or dull-metallic similar to typical Lyrcus . Most Mesopolobus also have at least the left mandible tridentate, consisting of two ventral teeth and a broader dorsal truncation (Fig. 180), though the truncation can be shallowly concave and a few species have both mandibles quadridentate similar to Lyrcus . The single most reliable feature to differentiate members of the two genera is fore wing setal pattern. Although the number of rows of admarginal setae and the size of the speculum varies in Mesopolobus as for Lyrcus , the admarginal setae are always as short as or shorter than the dorsal discal setae even if there is two or rarely only a single row of admarginal setae and these are entirely or mostly visible because of an extensive speculum ( Fig. 189 ).

Trimeromicrus maculatus Gahan is intermediate in features between species I classify in Lyrcus and Mesopolobus and its correct relationships and classification remain problematic (see further under Trimeromicrus ). Exclusion of T. maculatus and species with a modified metapleuron from Lyrcus makes it less likely that Lyrcus is polyphyletic. However, even with removal of these species there is no evidence that Lyrcus is monophyletic or at least not paraphyletic relative to some other genera. The genus is differentiated not by autapomorphies but by a combination of features that, although unique in combination, are shared individually with members of other genera.

Figures

FIGURES 13 – 20. Lyrcus cyaneus, Ƥ syntypes (INHS). 13, syntypic series on card; 14, head, frontolateral; 15, habitus, lateral; 16, head and antennae, frontal; 17, fore wing; 1 8, fore wing admarginal area (upper surface / lower surface); 19, propodeum and metapleuron, lateral; 20, propodeum, posterior.

FIGURES 170 – 175. 170 and 171, Lyrcus nigroaeneus Ƥ: 170, head and mesosoma, dorsal (130); 171, mesonotal setae (left: photo (130), right: SEM (94 )). 172 – 174, Lyrcus albopilosella (Ƥ lectotype): 172, head and mesosoma, dorsal; 173, head, frontolateral; 174, metapleuron. 175, Lyrcus sp., Ƥ mesosoma, dorsal (168). No. in parenthesis = CNC 2011 photo no.

FIGURES 182 – 189. 182 – 187, Lyrcus spp., Ƥ fore wing: 182, L. albopilosella (lectotype); 183, Lyrcus sp. (128); 184, L. rosaecollis (129); 185, L. nigraeneus (127), 186, Lyrcus sp. (123); 187, Lyrcus sp. (124). 188, Oaxa albiclava, Ƥ fore wing (118). 189, Mesopolobus finlaysoni, Ƥ fore wing (201). No. in parenthesis = CNC 2011 photo no.

FIGURES 33 – 38. Lyrcus helice, Ƥ lectotype. 33, habitus, dorsal; 34, habitus, lateral; 35, head, frontolateral; 36, propodeum, posterior [arrow points to paraspiracular carina]; 37, left antenna; 38, admarginal area of left fore wing and hind wing below.

FIGURES 190 – 197. 190 – 196, Lyrcus spp., Ƥ propodeum, posterolateral (SEM): 190, L. nigraeneus (98); 191, Lyrcus sp. (99); 192, Lyrcus sp. (102); 193, Lyrcus sp. (104); 194, L. nigroaeneus (94); 195, Lyrcus sp. (108); 196, Lyrcus sp. (107). 197, Lyrcus sp., Ƥ propodeum, posterior (30) [arrow points to denticle]. No. in parenthesis = CNC 2011 photo no.

FIGURES 97 – 104. Jaliscoa nudipennis Ƥ. 97, habitus, lateral (holotype); 98, head, frontal (35); 99, habitus, dorsal (holotype); 100, pro- and mesonotum (32); 101, mesopectus, ventrolateral (25); 102, meso- and metapleuron (26); 103, propodeum, posterolateral (106: SEM) [arrow 1 points to vertical and arrow 2 points to oblique carina on callus]; 104, fore wing (36). No. in parenthesis = CNC 2011 photo no.

FIGURES 157 – 162. 157 – 159, Heteroschema spp.: 157, 3 head, lateral (139) [insert: pedicel – fl 3]; 158, Ƥ propodeum, posterolateral (95); 159, Ƥ lower face and mandibles, frontal (145). 160 and 161, Neocatolaccus tylodermae Ƥ: 160, propodeum, posterior (114: SEM); 161, clypeus and mandibles (146). 162, Neocatolaccus sp. Ƥ, propodeum, posterolateral (125: SEM). No. in parenthesis = CNC 2011 photo no.

FIGURES 129 – 136. Eurydinoteloides spp. 129 – 132, 3 antenna: 129, Eurydinoteloides sp. (190) [insert: pedicel – fl 5]; 130 Eurydinoteloides sp. (171) [insert: pedicel – fl 4]; 131, E. bacchadis (147) [insert: pedicel – fl 3]; 132, Eurydinoteloides sp. (189) [insert: pedicel – fl 3]. 133 – 136, Ƥ lower face, frontolateral: 133, Eurydinoteloides sp. (133); 134, Eurydinoteloides sp. (64); 135, Eurydinoteloides sp. (SEM); 136, Eurydinoteloides sp. (89). No. in parenthesis = CNC 2011 photo no.

FIGURES 198 – 203. 198 – 202, Mesopolobus spp., Ƥ propodeum, posterolateral (SEM): 198, M. tortricidis (110); 199, M. finlaysoni (111); 200, Mesopolobus sp. (112); 201, Mesopolobus sp. (113); 202, Mesopolobus sp. (39). 203, Oaxa albiclava, Ƥ propodeum, posterolateral (203: SEM). No. in parenthesis = CNC 2011 photo no.

FIGURES 123 – 128. Eurydinoteloides spp. Ƥ. 123, E. americana (holotype), propodeum, posterior; 124, E. bacchadis propodeum, posterolateral (101: SEM); 125, Eurydinoteloides sp., mesonotal setae (31) (left: photo, right: SEM); 126, E. americana (holotype), fore wing (circles = admarginal setae, rectangles = leading row of dorsal discal setae); 127, E. laticeps (syntype), fore wing; 128, E. bacchadis, fore wing (155). No. in parenthesis = CNC 2011 photo no.