Pteromalus grisselli, Gibson, Gary A. P., 2013

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 : 19-27

Pteromalus grisselli n. sp.

Figs 49–53 , 55–60

Material examined. Holotype Ƥ (USNM): 5 mi. W. Clarkston, Asotin Co., WASH., VI-3-82 / G.S. Wheeler Collector / Reared from Dictyna egg sac, #13 / Catolaccus n. sp., det. E. Grissell, 1983, ID 82-7513 / Holotype Pteromalus grisselli Gibson.

Paratypes (29Ƥ, 143). CANADA. Ontario: Ottawa, 2.VI.40, O. Peck (1Ƥ CNC). Yukon Territory: Klo-Kut, Old Crow area between Porcupine R. & Small lake, edge of open area, 3-7VII.75, REM, 7-7-75-1 (1Ƥ CNC, Photo 2011-97). USA. Idaho: Boise Co., Garden Valley, 11 mi. E, 5.VIII.82, reared from Dictyna egg sacs (13 WFBM). Kootenai Co., Atol, 5 mi. E, 10.VI.82, G.S. Wheeler, web sacs of Dictyna sp. (1Ƥ USNM). Reared from egg sacs of Dictyna sp., Centaurea maculosa, G.S. Wheeler—Athol , 1 mi. E, 25.VI.82, 6,11, 13, 29.VII.82 (3Ƥ, 23 CNC; 1Ƥ USNM; 9Ƥ, 33 WFBM (CNC Photo number 2011-117, 161, 163)); Farragut St. Pk., 25.VI.82 (2Ƥ, 13 WFBM); Rathdrum, 3 mi. S, 25.VI.82 (1Ƥ USNM; 13 WFBM); Salmon, 8.6 mi. S, 24.VII.82 (1Ƥ WFBM). Montana: Wheatland Co., Deadman’s Basin St. Rec. Area, 3600’, 5.VI.82, G. Gibson (2Ƥ CNC). Washington: Asotin Co., same data as holotype except some reared 3, 30.V, 6, 9, 15.VI and some labelled Sisymbrium altissimum (6Ƥ, 43, USNM; 1Ƥ, 23 WFBM (CNC Photo 2011-135, 136, 162)).

Etymology. Named after Dr. Eric Grissell, who first realized the similarity between this species and those of Catolaccus .

Description. FEMALE (habitus: Figs 55, 57). Length = 1.9–3.1mm. Head dark metallic green to bluish-green or bluish-purple ( Figs 49, 50 ) and often with diffuse coppery lustre under some angles of light; setae mostly brownish on vertex, but more white on face and genae ( Figs 49, 50 ); mandible yellow except for reddish-brown teeth; palpi yellowish-brown. Antenna with scape and ventral surface of pedicel yellow, but remainder of pedicel and flagellum dark brown ( Fig. 53 ). Mesosoma (Figs 55, 57) similar in colour to head except tegula yellow to yellowish-brown; setae mostly brownish similar to vertex, but propodeal callus sometimes with more whitish setae. Legs (Fig. 57) with coxae similar in colour to mesosoma or dark brown with variably distinct metallic lustre; femora extensively dark brown, sometimes with slight metallic lustre, but lighter yellowish apically; tibiae or at least protibia sometimes yellowish similar to femora apically, but meso- and metatibiae usually variably distinctly brown medially except narrowly basally and somewhat more broadly yellowish apically; tarsi yellow except apical one or two tarsomeres brown. Gaster dark brown except usually for metallic green lustre basally and laterally under some angles of light (Fig. 57). Fore wing hyaline with venation yellow.

Head in dorsal view transverse-rectangular, about 2× as wide as long; occiput not margined; OOL: POL: LOL: MPOD about 3.0: 3.4: 1.6: 1.0. Head in frontal view ( Fig. 49 ) about 1.2× as wide as high; distance between eyes about 1.7× height of eye; face meshlike reticulate, including immediately ventral of toruli, but clypeus vertically striate and striae extending at least half distance to toruli dorsomesally and obliquely toward lower orbit somewhat more extensively lateral of toruli ( Figs 49, 51 ); clypeus with lateral margin indicated by incision about as deep as emargination of anterior margin, hence more or less bilobed, but not protuberant, the clypeal lobes at same level as mouth laterally ( Figs 49, 51 ). Head in lateral view (Fig. 57) with malar space about 0.7× height of eye, abruptly incurved along oral margin only along narrow band above base of mandible anterior to obscure malar sulcus ( Fig. 51 ). Eye (Fig. 57) about 1.6× as high as wide, with very short and sparse, inconspicuous setae (best seen from strongly oblique view). Antenna ( Fig. 53 ) with length of pedicel + flagellum about 0.75× width of head; scape about 0.9× height of eye with apex separated from ventral margin of anterior ocellus by about one ocellar diameter if appressed to head ( Fig. 49 ); pedicel about 2× as long as apical width and almost as long as combined length of fl1–fl3; fl1strongly transverse, ringlike; fl2 transverse and about 1.5× as long as fl1; fl3 and fl4 slightly longer than wide, fl5 quadrate or fl5–fl8 slightly wider than long, and clava about 2× as long as wide.

Mesosoma dorsally as in Fig. 55, and laterally as in Fig. 57. Fore wing with discal setae not reduced beyond speculum and with marginal fringe; costal cell dorsally with line of setae along anterior margin over about apical half and ventrally with 2 lines of setae along anterior margin over at least basal half, the setae becoming more numerous apically such that about apical third of cell completely setose (Fig. 59); basal fold with line of 2–10 setae, but at most with 1 seta apically within basal cell; mediocubital fold bare basal and distal to basal fold; disc with 1–9 setae on ventral surface adjacent to basal fold; smv: mv: stv: pmv about 3.0: 1.3: 1.0: 1.5; disc ventrally with about 5 or 6 rows of admarginal setae; speculum extending only slightly beyond base of marginal vein so as to mostly cover admarginal setae (Fig. 59). Propodeum (Fig. 60) with sinuate, carinate plica extending posteriorly to nucha, sometimes continuous through nuchal furrow but usually variably distinctly interrupted as separate carina within nuchal furrow; plical region and nucha similarly meshlike reticulate, the plical region with short longitudinal carinae along anterior margin and with variably developed, only sometimes complete median carina; paraspiracular area sparsely setose posteriorly (not visible in Fig. 60).

Gaster ovate (Fig. 55) to subcircular, but pointed apically, about 1.2–1.4× as long as wide and only about 0.8–0.85× length of mesosoma (air-dried specimens, excluding ovipositor sheaths).

MALE (habitus: Figs 56, 58). Length = 1.5–3.0 mm. Similar to female except as follows. Head and mesosoma often brighter green ( Figs 50, 52 , 58); pedicel sometimes and anelli usually yellowish, and funiculars often with at least extreme apices indistinctly yellowish (in smaller males) to distinctly bicolored, brown basally and yellow apically ( Figs 51 , 58); legs of smaller males somewhat lighter in colour though similar to female (Figs 56, 58), but entirely yellow beyond coxa in larger males except metafemur dark with metallic lustre; gaster with subbasal, transverse dorsal band and often larger yellowish-orange region ventrally. Sculpture similar to female except striae sometimes distinct only on clypeus. Structure similar to female except scape extending fully to ventral margin of anterior ocellus ( Fig. 50 ); oral margin flat lateral to clypeus, and rounded into gena on same level ( Fig. 52 ).

Distribution. Map 1.

MAP 1. Distribution of Pteromalus grisselli Gibson.

Biology. The biology of this species was studied by Wheeler and McCaffrey (1989) who reared most of the specimens included in the type series as egg predators from the egg sacs of Dictyna coloradensi Chamberlin ( Araneae : Dictynidae ).

Discussion. Evidence is lacking for the monophyly of Pteromalus , and P. grisselli and similar European species such as P. platyphilus Walker (1874) and P. crassicornis (Zetterstedt 1838) may eventually be shown to be more closely related to species of Catolaccus than to most other species currently classified in Pteromalus . However, I am describing it in Pteromalus based on current generic concepts. Pteromalus grisselli is similar to species classified in Catolaccus in several respects, including overall habitus (cf. Figs 3 , 57). The head is comparatively thick with similar relative dimensions as for Catolaccus , the antenna is similar, including the scape not extending to the anterior ocellus in females (cf. Figs 6 , 49 ), the right mandible is quadridentate and the left mandible tridentate (cf. Figs 6 , 52 ) (though not all Catolaccus have this mandibular structure), the propodeum has a similar structure and sculpture pattern as for most Catolaccus (cf. Figs 11 , 60), and the fore wings have a similarly broad costal cell, the same relative ratios of the veins, and a setal pattern typical of various Catolaccus (cf. Figs 10 , 59). Individuals, particularly males of P. grisselli , are more distinctly metallic than typical Catolaccus , but individuals of C. cyanoideus are at least obscurely dark metallic blue ( Figs 3–6 ) and some specimens have quite a distinct green lustre under some angles of light. Under current concepts, P. grisselli is differentiated from Catolaccus primarily by the absence of a broad, arch-like malar depression (cf. Figs 7 , 51 ), though there is a very slender incurved region along the oral margin above the base of the mandible (cf. Fig. 135 ) (see further below). Different species I classify in Eurydinoteloides lack a malar depression entirely ( Fig. 136 ) or have a variably slender ( Fig. 135 ) to broad, arch-like depression ( Figs 133, 134 ), and species I include in Lariophagus usually lack ( Fig. 26 ) though sometimes have ( Fig. 25 ) an arch-like malar depression. Hence, character-state distribution suggests that presence or absence and size of a malar depression can be a species rather than a generic feature. Pteromalus grisselli also differs from Catolaccus species in having the clypeus strongly striate and, even though distinctly bidentate ( Figs 49–52 ), it is less distinctly protuberant than for Catolaccus species (cf. Figs 6 , 49 ). Further, unlike Catolaccus , males of P. grisselli have a large subbasal yellowish region on the gaster (Figs 56, 58) like some, though not all, species classified in Pteromalus .

The specimen of P. grisselli from Yukon (Map 1) might indicate a Holarctic species and I describe P. grisselli as a new species with some hesitation because at least females appear to be very similar to P. platyphilus . This latter species is widely distributed throughout the Palaearctic region (Noyes 2012) and is also a predator in spider egg sacs, including those of Dictyna species (see Noyes 2012 for references). Graham (1969) included P. platyphilus along with P. crassicornis in his platyphilus species-group of Pteromalus (see Graham 1969, couplet 49 for differential features of group). I have not seen any specimen identified as P. crassicornis , but the CNC has a single female from Turkey that is identified as P. platyphilus (by Z. Bouček, 1989) and a female from Spain identified as Pteromalus cf. platyphilus (by Z. Bouček, 1993). Graham (1969: 508) stated that the “striations of the clypeus extend well up the genae, nearly to the lower corners of the eyes”, in both P. platyphilus and P. crassicornis . This is similar to the facial sculpture of P. grisselli ( Figs 49–52 ), though Dzhanokmen (2001) described the clypeus of P. platyphilus as finely rugose. Neither female identified by Bouček has the lower face as strongly striate as for females of P. grisselli or described for the platyphilus species-group by Graham (1969), though the lower face lateral to the clypeus is obliquely reticulate-striate partly toward the lower orbit. Similar to P. grisselli , both females have a slender malar depression along the oral margin above the base of the mandible. Graham (1969: 514) differentiated P. crassicornis from P. platyphilus in part by the gena having “a narrow hollow above the base of mandible”, though the original description of Catolaccus pappi Szelényi (1982: 386) , which was synonymized under P. platyphilus by Bouček and Rasplus (1991), stated the head “near mandible bases on both sides with a deep hole”. The female from Turkey has its broad costal cell bare dorsally but setae on the mediocubital fold proximal to the basal fold, unlike P. g r i s s e l l i and the female from Spain, though the latter, like P. grisselli , has two rows of setae along the ventral surface of the costal cell, unlike the single row described for P. platyphilus by Dzhanokmen (2001). Further, Szelényi (1982) described the speculum in C. pappi as being closed below, unlike P. grisselli or the female from Spain or Turkey. Both of the latter females are dark, similar to the description of P. platyphilus by Dzhanokmen (2001), though the female from Spain has a slight green lustre under some angles of light. The female from Turkey certainly represents a different species from P. grisselli , but observed differences between the female from Spain and those from North America might represent nothing more than infraspecific variation. My decision to describe P. grisselli as a new species is based primarily on males. The CNC has five males from Oman identified as P. platyphilus by Z. Bouček in 1991. These have bicolored funiculars (apices narrowly white) and a large subbasal white region on the gaster similar to males of P. grisselli . However, all the Oman males have a distinctive head structure and sculpture. The oral margin is broadly transverse with the region lateral to the clypeus being slightly produced and reflexed relative to the clypeus, and the clypeus and face meshlike reticulate rather than striate as for males of P. grisselli (cf. Figs 52, 54 ). Graham (1969) noted that males of P. platyphilus were unknown at that time, but tentatively keyed them based on specimens identified as such by Bouček. Graham (1969) did not mention facial sculpture, but the first differential feature used was “edge of oral fossa, on either side of the clypeus, with a projecting flange rather like that of Rohatina inermis ” (Graham 1969: 514; fig. 572). Consequently, males of P. grisselli are different from those identified as P. platyphilus by Bouček. Given the wide distribution known for P. platyphilus and some apparent discrepancies between the observations of Bouček and the descriptions of Graham (1969), Szelényi (1982) and Dzhanokmen (2001), future revision of the platyphilus species-group is required to more confidently assess species limits and sex associations. Further studies that including molecular techniques are also required to assess whether platyphilus -group species might be more closely related to species classified in Catolaccus than to others classified in Pteromalus .

Within North America, the only other pteromalids known to be predators in egg sacs of spiders are Arachnopteromalus dasys Gordh (1976) and Epipteromalus algonquinensis Ashmead (1904) . Arachnopteromalus is monotypic, but Sureshan and Talukdar (2009) described a second species of Epipteromalus , E. bengalensis from spider egg sacs in India. Epipteromalus algonquinensis was differentiated in part by “strong striation radiating from mouth margin and extending above toruli” (Bouček and Heydon 1997: 597; fig. 446) similar to at least P. grisselli , whereas E. bengalensis was described as having the striae hardly reaching beyond the clypeal margins. Individuals of E. algonquinensis also have a very slender, incurved region above the base of the mandible and very sparse, short, and inconspicuous setae on the eyes similar to P. grisselli , and males have a subbasal yellow region on the gaster. However, unlike P. grisselli , platyphilus -group species or Catolaccus species, both mandibles of E. algonquinensis have four similar teeth, the marginal vein is longer (1.5× length of stigmal vein), the propodeum has an anteriorly margined adpetiolar strip rather than a nucha, and the scutellum is strongly convex and anteriorly narrowed such that the axillae are separated by a distance equal to only about half the width of an axilla. Arachnopteromalus dasys is similar to E. algonquinensis and P. grisselli in having a slender incurved region along the oral margin above the base of the mandible, but is distinguished by being generally much more densely and conspicuously setose, including having the fore wings entirely setose, and having longer eyes such that the toruli are distinctly above the lower margins of the eyes and the malar space is comparatively short. It also has a reticulate nucha and complete, carinate plicae similar to platyphilus- group and Catolaccus species, but has quadridentate mandibles and a marginal vein that is almost 1.5× as long as the stigmal vein; the male has a uniformly dark metasoma.

Jaliscoa Bou č ek

Jaliscoa Bouček, 1993: 1281 –1282. Type species: Jaliscoa nudipennis Bouček , by original designation (USNM). Gender: feminine.

Heterolaccus ; Burks, 1954 (in part).

Included species. Jaliscoa grandis (Burks 1954) n. comb., J. hunteri (Crawford 1908) n. comb., J. nudipennis Bouček (1993) , J. townsendi (Crawford 1912) n. comb., J. vulgaris (Ashmead 1894b) n. comb.

Diagnosis. Metapleuron extensively smooth and shiny with anterior margin evenly curved, unsculptured, and raised above and slightly over posterior margin of mesopleuron ( Figs 66 , 71 , 84 , 92 , 96, 102, 103 ). Flagellum of both sexes with 2 basal ringlike articles lacking mps and 6 funicular articles with mps ( Figs 68 , 79 , 85, 86 ). Fore wing ventrally usually with 1 or 2 rows of admarginal setae (sometimes up to 3 irregular rows distally) of similar length to discal setae when latter not reduced in density behind marginal vein, and with speculum extending behind marginal vein at least to near base of stigmal vein ( Figs 72 , 80 , 93, 94 , 104 , 110 ). Head with bare, arch-like malar depression extending about half distance to lower orbit. Head and mesosoma black or at most with limited bluish lustre under some angles of light except propodeum sometimes more conspicuously blue to green, and with parallel-sided, white setae contrasting conspicuously with cuticle (e.g. Figs 77 , 81 , 85, 89 ). Propodeum with plical carina present ( Figs 95, 96 , 103 , 109 ) or absent ( Figs 71 , 83, 84 ) posteriorly in nuchal furrow and without or with variably complete, carinate costula ( Figs 71 , 83, 84 , 96 , 103 ), but without transverse carina within paraspiracular region. Both mandibles with four similar teeth ( Figs 67, 68 , 85–87 , 98 ).

Description. Head and mesosoma black ( Figs 81, 82 , 90 , 105, 106 ) or at most with limited bluish lustre ( Figs 66 , 102 ) except propodeum often more conspicuously green to bluish ( Figs 83 , 109 ), but with conspicuously contrasting, very slightly lanceolate (almost parallel-sided) white setae (e.g. Figs 77 , 81 , 85, 89 ). Eye bare or at least superficially bear with at most exceedingly short, sparse, inconspicuous setae. Mandibles quadridentate, with four similarly acute teeth ( Figs 67, 68 , 85–87 , 98 ). Head in frontal view ( Figs 67, 68 , 77 , 85 , 86, 98 ) transverseoval; antenna inserted obviously above lower margin of eyes near middle of face; tentorial pits not evident; clypeus incurved, vertically finely strigose ( Fig. 87 ) to mostly coriaceous- or reticulate-striate; supraclypeal region, lower face toward inner orbit, and scrobal depression with isodiametric meshlike sculpture ( Figs 88 , 105 ), but parascrobal region above level of toruli with more vertically aligned and elongated sculpture, and without tiny bump or smoother and shinier spot at ocular margin at midheight of eye ( Figs 67, 68 , 77 ). Head in dorsal view abruptly declined immediately behind posterior ocelli ( Figs 73, 75 , 89 , 99 ), hence strongly transverse with vaulted vertex, but occiput not margined. Head in lateral view with arch-like malar depression extending almost half distance to lower orbit ( Figs 69 , 78 , 88 , 105 ); malar space at most 0.5× height of eye and with ( Fig. 69 ) or without ( Fig. 88 ) malar sulcus. Antenna with scape extending about to level of ventral margin of anterior ocellus ( Fig. 79 ); flagellum with 2 anelli and 6 funiculars with mps in both sexes ( Figs 68 , 79 , 85, 86 ); clava with apical clavomere uniformly conical with tiny, terminal, encircled, micropilose sensory region; flagellum of male with funiculars closely abutting.

Pronotum with collar smoothly rounded or transversely ridged ( Fig. 82 : arrow) relative to steeply angled collum ( Figs 66 , 74, 76 , 91 97 ), but not separated from collum by smooth and shiny carina. Mesonotum meshlike-reticulate ( Figs 81 , 90 , 100 ); mesoscutum with incomplete notauli; scutellum widely truncate anteriorly, the axillae separated by distance about equal to width of axilla, moderately convex and broad, without frenal line, and with posterior margin vertical above metanotum, not reflexed into distinct marginal rim; mesopectus sometimes with anterior margin of femoral depression carinately margined and continuous ventrally as transverse carina anterior to mesocoxa ( Figs 101, 102 : mpc) differentiating mesosternal shelf; upper mesepimeron extensively smooth and shiny ( Figs 82 , 92 , 101, 102 ). Fore wing ( Figs 72 , 80 , 93, 94 , 104 , 110 ) hyaline; marginal vein not thickened and about 2–3× length of stigmal vein and about 1.3–1.8× length of postmarginal vein; stigma not distinctly capitate; costal cell variably wide, but dorsally bare and ventrally either bare or with variable number of setae over at most about apical third; basal cell including basal and mediocubital folds bare; disc ventrally usually with 1 or 2 rows of admarginal setae (sometimes up to 3 irregular rows distally) of similar length as dorsal setae when these not reduced; disc sometimes with setae variably reduced in density and length beyond speculum, at least behind stigmal and/or marginal veins ( Figs 72 , 94 , 104 ), and then sometimes marginal fringe absent ( Fig. 94 ); speculum extending behind marginal vein at least to level near base of stigmal vein such that admarginal setae fully exposed. Metapleuron extensively smooth and shiny with anterior margin uniformly curved, unsculptured, and raised slightly above and over posterior margin of mesopleuron ( Figs 66 , 71 , 92, 96 , 102, 103 ). Metacoxa bare dorsobasally; metatibia with single tibial spur. Propodeum ( Figs 71 , 83, 84 , 95, 96 , 103 , 109 ) with vertical flange posterolaterally on callus ( Fig. 103 : arrow 1), but in dorsal view not developed as distinct, laterally projecting denticle; with Λ-shaped to transverserectangular, finely meshlike 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 basolateral depression (e.g. Figs 71 , 83, 84 ) and sometimes with complete plical carina extending from depression to nucha ( Figs 96 , 103 ); plical region meshlike coriaceous-reticulate similar to nucha, with median carina and variably developed costula, sometimes as short, irregular, transverse carina crossing median carina at about midlength so as to appear more or less +-like (e.g. Fig. 71 ) or as variably conspicuous, convex angulation (e.g. Figs 83, 84 ) or as uniform carina recurved from plical carina and directed anteromesally to median carina to differentiate anterior and posterior, transverse panels ( Fig. 103 ); paraspiracular region without transverse carina, though sometimes callus with carina extending posterolaterally from spiracle ( Fig. 103 : arrow 2).

Gaster of female lanceolate with hypopygium extending at most about two-thirds length of gaster (e.g. Figs 64, 65 ); gaster of male with pale region basally (e.g. Figs 70 , 75 ); 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. Parasitoids of Bruchinae ( Coleoptera : Chrysomelidae ) and Curculioninae ( Coleoptera : Curculionidae ).

Discussion. When Bouček (1993) established Jaliscoa he stated that the genus belonged to an otherwise numerous group of Pteromalinae characterized by a propodeum with a costula delimiting two depressed anterior areas ( Fig. 103 ), but differed by having reduced fore wing pilosity ( Fig. 104 ) and a transverse carina on the ventral part of the mesopleuron delimiting a mesosternal shelf ( Figs 101, 102 : mpc). He also noted the presence of only 2 anelli and a “hollowed” gena, and although not discussed further included in the description the statement that the metapleuron had a “sharp and raised anterior edge” (Bouček 1993: 1281). Because of the presence of a mesosternal shelf, Bouček and Heydon (1997) keyed Jaliscoa through couplet 151 along with Perilampidea Crawford (1913) and Halticopteroides Girault (1913a) . Perilampidea and Halticopteroides both contain only two described species (Noyes 2012). In addition to the differential features given in Bouček and Heydon (1997), members of the latter two genera have a metapleuron that is completely sculptured and abutting the mesopleuron on the same level, three similar teeth or two teeth and a dorsal truncation on the left mandible, lack a malar depression, usually have two metatibial spurs, and the costal cell is often at least partly setose dorsally. Females of some species of Halticopteroides have three anelli. Relationships and limits of Perilampidea and Halticopteroides relative to each other require further study, as do their relationships to other genera such as Neocatolaccus Ashmead (1904) and Heteroschema Gahan (1919) , which lack a mesosternal shelf, but typically have two metatibial spurs, similar propodeal structures, and three anelli in females but only two anelli in males.

For the reasons discussed under Catolaccus , I transfer to Jaliscoa the four species with a modified metapleuron that Burks (1954) included in Heterolaccus . The generic key of Bouček and Heydon (1997) is constructed primarily for females, but both sexes of those Jaliscoa species without a mesosternal shelf key to Lyrcus (couplet 263) because of the presence of a malar depression and a flagellum with two anelli and six funiculars. Females of Lyrcus as defined herein do not key to couplet 263 because they have at least three distinct anelli. Males of Lyrcus , and particularly males of Eurydinoteloides , are variable in structure of the third flagellomere (see under respective genera). However, regardless of the number of anelli in either sex, an unmodified metapleuron ( Figs 19 , 174 ) differentiates Lyrcus from Jaliscoa . All species of Jaliscoa have the metapleuron extensively smooth and shiny, convex anteriorly, and with the anterior margin evenly curved, unsculptured, and raised slightly above and over the posterior margin of the mesopleuron ( Figs 66 , 71 , 84 , 92 , 96, 102, 103 ). This derived structure is also shared with some species of Eurydinoteloides ( Figs 48 , 139, 140 ). Other species of Eurydinoteloides have a similar structure except that the anterior margin is recurved outward at an abrupt angle relative to the mesopleuron ( Figs 141, 142 ), whereas others have a less conspicuously modified metapleuron with only a comparatively small, smooth and shiny region anteromesally and with the anterior margin variably distinctly angulate with some fine sculpture, and sometimes only slightly separated from the mesopleuron ( Figs 114–116 , 118 , 137, 138 ). This latter structure may well be the groundplan metapleural structure for Eurydinoteloides because it is more similar to the presumed plesiomorphic metapleural structure of Lyrcus ( Figs 19 , 174 ) and most other pteromalids ( Figs 27, 28 , 212, 213 ). If so, the similar metapleural structures shared by Jaliscoa and some Eurydinoteloides must either have evolved convergently or the common structure indicates Jaliscoa renders Eurydinoteloides paraphyletic. Independent evolution might be supported by the number of anelli in females of the two genera (see below) and by a similar metapleural structure also being possessed by specimens (mostly in CNC) representing several undescribed Neotropical species of uncertain generic status. The species (e.g. Fig. 61 ) are morphologically similar to the only two species currently classified in Toxeumelloides Girault (1913a) , but both described species of Toxeumelloides have a plesiomorphic metapleural structure as well as a flagellum with two anelli (Bouček 1993, fig. 80). The undescribed species have a derived metapleural structure ( Fig. 62 ) very similar to that of Jaliscoa and some Eurydinoteloides species, as well as three anelli in both sexes ( Fig. 61 : insert). Mandibles are concealed in most available specimens, but it appears that at least the left mandible is tridentate in Toxeumelloides species, whereas the left mandible is quadridentate in the undescribed species similar to Jaliscoa . Otherwise, the undescribed species resemble described species of Toxeumelloides . The fore wing has one or more rows of admarginal setae that are covered by the discal setae behind at least the apical half of the marginal vein and that are not conspicuously longer than the discal setae (Bouček 1993, fig. 79), and the costal cell is setose along its length. However, the two most distinctive features that distinguish them from Jaliscoa and Eurydinoteloides is an elongate, smooth and shiny tubular petiole in both sexes ( Figs 61, 63 ; Bouček 1993, figs 81, 82), and dark hairlike setae that do not contrast with the cuticle. The described species of Toxeumelloides and the undescribed Neotropical species also have a variably developed malar depression, either abruptly reflexed along the oral margin above the base of the mandible similar to P. grisselli and some Eurydinoteloides species ( Fig. 135 ) or more arch-like, similar to Jaliscoa and some Eurydinoteloides ( Figs 112, 113 , 133 ) species. Based on similarities, it seems likely that the undescribed species are closely related to Toxeumelloides , but comprise a clade with a secondarily modified metapleuron and three anelli. Recognition of the undescribed species as a separate genus based on these two differences could render Toxeumelloides paraphyletic. However, if the difference in mandibular dentition noted above is shown to be accurate, then the presence of only three teeth on the left mandible might be evidence for a monophyletic Toxeumelloides that is the sister group of the undescribed group of species.

Although the shared metapleural structure of Jaliscoa and some Eurydinoteloides could be convergent, a close relationship between the two taxa is strongly suggested by some species sharing very similar morphologies, particularly those Eurydinoteloides with a similarly broad, arch-like malar depression. Some species of Eurydinoteloides from the Neotropical region with a broad malar depression are also completely black and therefore closely resemble Jaliscoa species, particularly J. hunteri or J. bouceki (cf. Figs 91 , 121 ). However, in addition to having three anelli, such species have a metapleuron whose anterior margin is anteromesally angulate and partly sculptured ( Figs 115, 116 ) rather than evenly curved and smooth (see further under Eurydinoteloides ). Some other species of Eurydinoteloides not only have a broad malar depression ( Fig. 133 ) but also have the anterior margin of the metapleuron evenly curved and smooth ( Figs 139, 140 ) similar to Jaliscoa , but these are differentiated from Jaliscoa by the presence of three anelli and a more distinct metallic lustre ( Figs 122 , 133 , 139 ). The latter colour feature is less reliable because, as noted in methods, different light sources affect the perception and intensity of observed metallic lustre. Males of an undescribed species of Eurydinoteloides from Florida are particularly similar to Jaliscoa males because they not only have an arch-like malar depression ( Fig. 133 ), but also a flagellum with two anelli and six similar funiculars encircled by mps ( Fig. 132 ). However, like females, those Eurydinoteloides males with an arch-like malar depression also have quite an obvious metallic lustre on the head and mesosoma.

If Jaliscoa and Eurydinoteloides represent a monophyletic lineage, either as monophyletic sister taxa or with Jaliscoa rendering Eurydinoteloides paraphyletic, then a single genus might be recognized based on the metapleuron being partly shiny with the anterior margin at least slightly separated from the mesopleuron. Two subgenera within this genus might then be recognized based on females having either two or three anelli. Species with either two or three anelli in females are classified together in some other genera such as Mesopolobus Westwood (1833) , Psilocera Walker (1833) , and Lariophagus . However, intermediate structures between a large third flagellomere with mps and a smaller flagellomere without or with at least a reduced number of mps are evident in Psilocera and Lariophagus . Such intermediate structures are not evident for Jaliscoa or Eurydinoteloides in which females have, respectively, either two or three distinct anelli. Further, three compared to two anelli presumably represents a derived state within Pteromalinae . Consequently, this transformation series is opposite of the transformation series proposed above for the metapleuron in Eurydinoteloides and Jaliscoa . If these two genera do form a monophyletic group based on shared metapleural structure, then this suggests either that the three anelli shared by Eurydinoteloides and Lyrcus females is convergent or the two anelli of Jaliscoa females represents a secondary reversal. As noted above, some species of Eurydinoteloides are very similar to Jaliscoa . However, other species of Eurydinoteloides are very similar to Lyrcus , as discussed under this latter genus. I therefore prefer to retain Jaliscoa and Eurydinoteloides as separate genera until a rigorous phylogenetic analysis of Pteromaline is conducted to better resolve generic relationships.

Figures

FIGURES 49 – 54. Pteromalus spp. 49 – 53, Pteromalus grisselli: 49, Ƥ head, frontal (136); 50, 3 head and antenna, frontal (162); 51, Ƥ lower face and mandibles, frontolateral (136); 52, 3 lower face and mandibles, frontal (162); 53, Ƥ antenna (holotype) [insert: pedicel – fl 3]. 54, Pteromalus? platyphilus, 3 lower face and mandibles (175). No. in parenthesis = CNC 2011 photo no.

FIGURES 1 – 8. Catolaccus spp. 1, C. aeneoviridis, 3 antenna (132). 2 – 6, C. cyanoideus: 2, 3 antenna (188) [insert: pedicel – fl 3]; 3, Ƥ habitus, lateral (21); 4, Ƥ head and antenna, lateral (21); 5, Ƥ head and pronotum, dorsal (21); 6, Ƥ head, frontal (20). 7 and 8, C. aeneoviridis Ƥ: 7, head, frontolateral (34); 8, head and mesosoma, dorsal (131). No. in parenthesis = CNC 2011 photo no.

FIGURES 9 – 12. Catolaccus spp. 9, C. aeneoviridis Ƥ, mesoscutum – propodeum, lateral (33) [arrow points to reflexed marginal rim of scutellum]. 10, C. cyanoideus Ƥ, fore wing (19). 11 and 12, Ƥ propodeum, posterolateral (SEM): 11, C. aeneoviridis (115); 12, C. kansensis (96). 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 21 – 28. Lariophagus spp. 21 and 22, Ƥ habitus, dorsal: 21, L. fragariae (PT) (20); 22, Lariophagus sp. (15). 23 and 24, Ƥ scape – fl 5: 23, L. fragariae (PT) (20); 24, Lariophagus sp. (14). 25 and 26, Ƥ lower face and mandibles, frontolateral: 25, L. fragariae (17); 26, Lariophagus sp. (14). 27, Lariophagus sp., Ƥ meso- and metapleuron (13). 28, L. fragariae, Ƥ metapleuron and propodeal denticle [arrow] (160). No. in parenthesis = CNC 2011 photo no.

FIGURES 61 – 66. 61 – 63, undetermined species near Toxeumelloides: 61, 3 habitus, lateral (73) [insert: pedicel – fl 4]; 62, 3 meso- and metapleuron (73); 63, Ƥ propodeum and petiole, dorsal (170). 64 – 66, Jaliscoa bouceki Ƥ: 64, Ƥ habitus, dorsal (10); 65, Ƥ habitus, lateral (11); 66, Ƥ mesosoma, lateral (11). No. in parenthesis = CNC 2011 photo no.

FIGURES 67 – 72. Jaliscoa bouceki. 67, Ƥ head, frontal (154); 68, 3 head and antenna, frontal (156); 69, Ƥ head, frontolateral (88) [arrows point to malar sulcus]; 70, 3 habitus, lateral (155); 71, Ƥ propodeum, posterolateral (10: SEM); 72, Ƥ fore wing (116) [insert: admarginal setae]. No. in parenthesis = CNC 2011 photo no.

FIGURES 79 – 84. Jaliscoa grandis Ƥ. 79, antennae (6); 80, fore wing (27) [insert: admarginal setae]; 81, pro- and mesonotum, dorsal (27) [arrow points to pronotal emargination]; 82, mesosoma, lateral (12) [arrow points to pronotal ridge]; 83, propodeum, posterior (6); 84, propodeum, posterolateral (105: SEM). No. in parenthesis = CNC 2011 photo no.

FIGURES 91 – 96. Jaliscoa hunteri Ƥ. 91, habitus, lateral (8); 92, meso- and metapleuron (8); 93, fore wing (38); 94, fore wing (87); 95, propodeum, posterior (9); 96, propodeum, posterolateral (9: SEM). 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 85 – 90. Jaliscoa hunteri. 85, Ƥ head and antenna, frontal (7); 86, 3 head and antennae, frontal (157); 87, Ƥ clypeus and mandibles, frontal (7); 88, Ƥ lower face, frontolateral (158); 89, Ƥ habitus, dorsal (9); 90, Ƥ pro- and mesonotum, dorsal (28). No. in parenthesis = CNC 2011 photo no.

FIGURES 105 – 110. Jaliscoa vulgaris Ƥ. 105 and 106, Ƥ paratype (BMNH) (174): 105, head and antenna, frontolateral; 106, habitus, lateral. 107 – 110, Ƥ lectotype: 107, head, lateral; 108, habitus, dorsal; 109, propodeum, posterior [arrow points to plical carina within nuchal furrow]; 110, fore wing. No. in parenthesis = CNC 2011 photo no.

FIGURES 73 – 78. Jaliscoa grandis. 73, Ƥ habitus, dorsal (6); 74, Ƥ habitus, lateral (150); 75, 3 habitus, dorsal (151); 76, 3 habitus, lateral; 77, Ƥ head, frontal (6); 78, Ƥ lower face, frontolateral (169). No. in parenthesis = CNC 2011 photo no.

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 45 – 48. Eurydinoteloides tepicensis, Ƥ holotype. 45, habitus, dorsal; 46, habitus, lateral; 47, head and antennae, lateral; 48, meso- and metapleuron.

FIGURES 137 – 142. Eurydinoteloides spp., Ƥ metapleuron. 137 and 138, Eurydinoteloides sp.: 137, (91); 138, SEM. 139 and 140, Eurydinoteloides sp.: 139, (60); 140, (60: SEM). 141 and 142, Eurydinoteloides spp. No. in parenthesis = CNC 2011 photo no.

FIGURES 111 – 116. Eurydinoteloides spp. Ƥ. 111, E. bacchadis, head, frontal (53). 112, E. bacchadis (Ƥ paratype), lower face, frontolateral. 113 and 114, E. laticeps (Ƥ syntype): 113, head, lateral; 114, meso- and metapleuron. 115 and 116, E. bacchadis: 115, meso- and metapleuron (54); 116, metapleuron (54: SEM). No. in parenthesis = CNC 2011 photo no.

FIGURES 117 – 122. Eurydinoteloides spp. Ƥ. 117 and 118, E. americana (holotype): 117, mesosoma, lateral; 118, meso- and metapleuron. 119, E. laticeps (syntype), habitus, dorsal. 120, E. bacchadis (paratype), habitus, lateral. 121 and 122, habitus, lateral: 121, Eurydinoteloides sp. (164) [insert: pedicel – fl 4]; 122, Eurydinoteloides sp. (59). No. in parenthesis = CNC 2011 photo no.

FIGURES 210 – 215. Trimeromicrus maculatus Ƥ. 210, head and antenna, frontal (3); 211, head, frontolateral (SEM) [insert: inner orbit magnified]; 212, mesosoma, lateral (1); 213, meso- and metapleuron (SEM); 214, lower face and mandibles (4); 215, propodeum, posterolateral (109: SEM). [Arrow in Figs 210 and 211 points to smooth bump on inner orbit.] No. in parenthesis = CNC 2011 photo no.