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) Author Gibson, Gary A. P. text Zootaxa 2013 3612 1 1 85 journal article 10.11646/zootaxa.3612.1.1 de4a8a7e-1d54-4c56-bf2f-f72305327c18 1175-5326 283321 FEE56A44-B572-4A95-BC11-2FA9D1187AF8 Catolaccus Thomson Pteromalus ( Catolaccus ) Thomson, 1878: 152 . Type species: Pteromalus cavigena Thomson , by monotypy (Ƥ lectotype , LUZN, not examined). Gender: masculine. Catolaccus Thomson ; Ashmead, 1904: 320, 322. Merisoides Masi, 1911: 141 . Type species: Merisoides crassiceps Masi , by monotypy (Ƥ holotype , MCSN, not examined). Synonymy by Delucchi (1956: 230). Hortobagyia Szelényi, 1981: 403 –404. Type species: Hortobagyia crassiceps Szelényi , by original designation and monotypy (Ƥ holotype , HNHM, not examined). Synonymy by Szelényi (1982: 385). Included regional species. Catolaccus aeneoviridis (Girault 1911) , C. cyanoideus Burks (1954) , C. kansensis (Girault 1917c) , C. victoria Burks (1954) . Diagnosis. Metapleuron entirely sculptured and with anterior margin abutting posterior margin of mesopleuron on same level ( Figs 11, 12 ). Flagellum of both sexes with 2 basal ringlike articles lacking mps and 6 funicular articles with mps ( Figs 1, 2, 4 ). Fore wing ventrally with 3 or more rows of admarginal setae not longer than dorsal discal setae and with speculum often extending only to base of marginal vein ( Fig. 10 ), though sometimes to base of stigmal vein. Head with arch-like malar depression extending about one-third to one-half distance to lower orbit ( Figs 4, 7 ). Head and mesosoma sometimes with metallic blue or dark green lustre ( Figs 3–6 ), but setae hairlike and not contrasting conspicuously with cuticle ( Figs 3–9 ). Propodeum with variably developed plicae extending posteriorly to nucha, but without costula or transverse carina within paraspiracular region ( Figs 11, 12 ). Mandibular dentition variable, but at least left mandible often tridentate ( Fig. 6 ) or middle tooth with additional shorter, ventral, subapical tooth ( Fig. 6 : right mandible). Description. Head and mesosoma black or with bluish to dark green lustre, with setae sometimes white but not contrasting conspicuously with cuticle and not suberrect and bristlelike if dark ( Figs 3–9 ). Eye bare or at least superficially bare with at most exceedingly short, sparse, inconspicuous setae ( Figs 4–7 ). Mandibles with 3 or 4 teeth, the left mandible more commonly tridentate ( Fig. 6 ), but both mandibles sometimes with shorter, subapical, ventral dent on middle tooth representing fourth tooth ( Fig. 7 ). Head in frontal view ( Fig. 6 ) subcircular; antenna inserted at or slightly above lower margin of eyes below middle, but at least within about ventral third of face; tentorial pits not evident; clypeus often appearing somewhat protuberant or at least distinctly differentiated from lower face by deep lateral angulation, and partly smooth to coriaceous or vertically alutaceous to somewhat reticulate-striate; lower face sometimes with more distinct, radiating striae lateral to clypeus, but usually meshlike coriaceous to reticulate, and upper face more distinctly though often finely meshlike reticulate, without tiny bump or smoother and shinier spot at ocular margin at midheight of eye ( Figs 6, 7 ). Head in dorsal view ( Fig. 5 ) variably distinctly transverse, but vertex broadly rounded ( Fig. 4 ) and with ( Fig. 8 ) or without dorsally margined occiput. Head in lateral view with arch-like malar depression extending about one-third to one-half distance to lower orbit ( Fig. 7 ); malar space at least 0.5× length of comparatively small eye. Antenna with scape not quite extending to anterior ocellus, at least in female ( Fig. 6 ); flagellum with 2 anelli and 6 funiculars in both sexes ( Figs 1, 2, 4 ), the second anellus, at least in female, noticeably longer than first anellus ( Fig. 2 : insert) and sometimes obviously longer than wide; clava tapered to apex with small, ventroapical, encircled, oval micropilose sensory region; flagellum of male ( Figs 1, 2 ) with funiculars closely abutting. Pronotum with collar smoothly rounded into or abruptly angled relative to steeply angled collum ( Figs 3, 5 ), but not separated from collum by smooth and shiny carina. Mesonotum ( Fig. 8 ) meshlike reticulate; mesoscutum with incomplete notauli; scutellum widely truncate anteriorly, the axillae separated by distance similar to width of axilla, comparatively low convex and broad, without frenal line dorsally, and usually with abruptly reflexed marginal rim ( Fig. 9 : arrow); mesopectus without mesosternal shelf; upper mesepimeron extensively smooth and shiny. Fore wing ( Figs 3 , 10 ) hyaline or rarely variably distinctly infumate behind marginal and stigmal veins; marginal vein not thickened and at most about 1.3× as long as stv and 1.25× as long as pmv ( Fig. 3 ); stigma not distinctly capitate; costal cell comparatively wide, with ( Fig. 10 ) or without setae dorsoapically but with setae extending length of cell ventrally; basal cell and basal and mediocubital folds bare or variably extensively setose ( Fig. 10 ), but at least vannal area and small region on disc adjacent to parastigma bare; disc ventrally with at least 3 rows of admarginal setae of similar length as dorsal setae and variably exposed depending on extent of speculum; disc dorsally variably densely setose beyond speculum and marginal fringe rarely absent if discal setae comparatively sparse; speculum often extending only to base of marginal vein ( Fig. 10 ), but sometimes partly or completely to base of stigmal vein. Metapleuron completely sculptured and with anterior margin on same level as and abutting posterior margin of mesopleuron ( Figs 11, 12 ). Metacoxa bare dorsobasally; metatibia with single tibial spur. Propodeum ( Figs 11, 12 ) in dorsal view rounded posterolaterally, without distinct angulation or denticle projecting laterally; with transverse-rectangular or more globose, sculptured nucha and with variably developed plicae extending from anterior margin to anterolateral margins of nucha; plical region meshlike reticulate ( Fig. 11 ) to transversely aciculate-coriaceous ( Fig. 12 ) and with or without median carina, but without costula; paraspiracular region without transverse carina. Gaster of female ( Fig. 3 ) ovate to distinctly elongate-lanceolate with hypopygium extending at most about two-thirds length of gaster; gaster of male uniformly dark, without pale 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. Generic limits. Burks (1954) provided a history of the interpretation of Catolaccus when he included it as one of five genera treated as the “ Catolaccus group in the Americas”. He also included Heterolaccus Masi (1937) in the group, but unfortunately misinterpreted this name. Bouček (1961) synonymized Heterolaccus as a subgenus of Pteromalus and because of this De Santis (1979) transferred to Pteromalus four of the five species that Burks (1954) included in Heterolaccus . However, in the same year Burks (1979) transferred three of the species that occur in North America north of Mexico to Catolaccus . Consequently, Noyes (2012) included two of the names ( townsendi and vulgaris ) in Pteromalus and three of the names ( fragariae , grandis and hunteri ) in Catolaccus . Burks (1954: 3) originally defined the Catolaccus -group using several features that are diagnostic of many genera, but primarily by the genae being “deeply excavated at the bases of the mandibles”. Heterolaccus sensu Burks and Catolaccus were differentiated from the other three Catolaccus -group genera by the presence of only two anelli in both sexes, and Heterolaccus sensu Burks was differentiated from Catolaccus by a “relatively narrow and asetose or nearly so” costal cell, and antennae inserted considerably above the ventral margin of the eyes ( cf . Figs 6 , 67 ). Bouček (1993: 1282) described the costal cell of J. nudipennis as being “very broad”, though bare ( Fig. 104 ) as for some Heterolaccus sensu Burks. Although none of the taxa included in Heterolaccus by Burks (1954) has a mesosternal shelf, four of the five ( grandis , hunteri , townsendi and vulgaris ) have a modified metapleuron similar to J. nudipennis . The metapleuron is partly smooth and shiny and its anterior margin is uniformly curved, convex, unsculptured and separated from the posterior margin of the mesopleuron ( Figs 92 , 102 ). Further, one species of Heterolaccus sensu Burk (1954) ( grandis ) was characterized by a strong pronotal carina that is distinctly emarginate sublaterally ( Fig. 81 : arrow), which is similar to the “laterally indented” structure described for J. nudipennis ( Fig. 100 ). Another species ( hunteri ) was partly characterized by reduced fore wing pilosity ( Fig. 94 ), and all the species have one or two rows of clearly exposed admarginal setae. I interpret the mesosternal shelf ( Fig. 101 ) and comparatively wide costal cell ( Fig. 104 ) of J. nudipennis as specific rather than generic features, and conclude that the four species with a modified metapleuron and two anelli in both sexes that Burks (1954) erroneously included in Heterolaccus are congeneric with J. nudipennis . In addition to having a plesiomorphic metapleural structure ( Figs 11, 12 ), Catolaccus is differentiated from Jaliscoa by the presence of three or more rows of admarginal setae that in most species are extensively overlain by the dorsal discal setae ( Fig. 10 ). Species also have inconspicuous hairlike setae on the head and mesonotum ( Figs 3–9 ) even if the setae are sometimes whitish. Another feature of most Catolaccus , although not distinctive for all species, is that the apical margin of the scutellum is reflexed into quite an evident marginal rim that in lateral view projects posteromedially as a short denticle or lip ( Fig. 9 : arrow) rather than being curved down in a single plane as is typical of Jaliscoa ( Figs 82 , 106 .). Individuals of Catolaccus also have the costal cell setose along its entire length ventrally ( Fig. 10 ), a shorter marginal vein ( Fig. 10 ; Dzhanokmen 1980, figs 7, 10), typically have a more broadly rounded vertex ( Figs 4, 5 ), and usually at least the left mandible is tridentate ( Fig. 6 ) or the middle tooth of one or both mandibles has an additional, though smaller, preapical tooth ( Figs 6, 7 ; Dzhanokmen 1980, figs 1, 5). My examination of type material determined that nine of 13 New World species classified in Catolaccus or originally described in Catolaccus and erroneously transferred to Pteromalus are incorrectly assigned (see below). The generic description given above is based primarily on New World species. Additional study of the five Palaearctic species included by Noyes (2012) in Catolaccus is required to verify the morphological limits and membership of the genus on a world level. Further study is also required to better establish phylogenetic relationships of Catolaccus to other genera, particularly Pteromalus . The USNM and CNC have specimens from the USA and Canada that E.E. Grissell (USNM, retired) identified as “ Catolaccus prob. n. sp.” in Wheeler and McCaffrey (1989: 371). Most of the specimens were reared from the egg sacs of Dictyna coloradensi Chamberlin ( Araneae : Dictynidae ). Although I do not treat Pteromalus in this work, I describe the new species below within Pteromalus based on current, though admittedly inadequate, generic concepts so that it is formally characterized and illustrated for comparison of Catolaccus with Pteromalus and other pteromalid species that are spider egg parasitoids. Species differentiation. Specimens of Catolaccus were not borrowed comprehensively from museums in order to revise the genus further to Burks (1954), but based on available material the following was noted. The first couplet of the key to Nearctic species of Catolaccus by Burks (1954) uses the presence of an occipital carina ( Fig. 8 : oc) to differentiate C. aeneoviridis and C. victoria from C. cyanoideus and C. kansensis . However, presence or absence of an occipital carina may not be visible if the posterior of the head is appressed to the pronotum. Fore wing setal pattern is usually a more readily visible feature to differentiate the two species pairs. Both C. cyanoideus ( Figs 3 , 10 ) and C. kansensis have setae dorsoapically within the costal cell and variably extensively on the basal and mediocubital folds (often also within the basal cell). Catolaccus aeneoviridis and C. victoria lack setae from the costal cell dorsally and from the basal cell, including the basal and mediocubital folds (also correlated with a more extensive speculum). Further, both C. aeneoviridis and C. victoria have a small, subapical, ventral dent on the middle tooth of both mandibles so that the mandibles appear more or less quadridentate ( Fig. 7 ), whereas C. cyanoideus and C. kansensis have the left mandible tridentate ( Fig. 6 ). Males of at least C. aeneoviridis also differ in antennal structure from those of C. cyanoideus and C. kansensis , having elongate funiculars and, at least in larger individuals, each funicular having multiple rows of closely aligned mps ( Fig. 1 ) compared to males of the latter two species ( Fig. 2 ). Correlated with the longer flagellum of C. aeneoviridis is a longer scape that, unlike males or females of other regional species, extends to the anterior ocellus. Additional study is required to better evaluate the species status of C. aeneoviridis relative to C. victoria , which Burks (1954) differentiated primarily on relative development of the median and plical carinae. FIGURES 1–8. Catolaccus spp. 1 , C. aeneoviridis , 3 antenna (132). 2–6 , C. cyanoideus : 2 , 3 antenna (188) [insert: pedicel–fl3]; 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. Catolaccus kansensis is differentiated from C. cyanoideus and the other two Nearctic species by a unique propodeal sculpture pattern. The propodeum completely lacks a median carina and is transversely aciculatecoriaceous ( Fig. 12 ) rather than being meshlike reticulate as for the other three described Nearctic species ( Fig. 11 ). However, I examined Old World specimens [ Austria , France (CNC); Hungary (INHS); Bulgaria , India , Israel , Italy (UCRC); Turkey (CNC, UCRC)] that I identify as C. crassiceps (Masi 1911) based on differentiation of C. crassiceps from C. ater (Ratzeburg) by Bouček (1970). The specimens are very similar to C. kansensis in colour pattern, propodeal sculpture pattern, mandibular dentition (Dzhanokmen 1980, fig. 8) and fore wing setal pattern (Bouček 1970, fig. 6; Dzhanokmen 1980, fig. 10). The CNC also has a female from Lethbridge, Alberta, Canada , identified as C. crassiceps by Bouček in 1989, which is C. kansensis sensu Burks (1954) . It is therefore quite possible that C. kansensis is a junior synonym of C. crassiceps . I do not formally propose the synonymy because I have not examined the fragmentary type remains of C. crassiceps (see Graham 1969 and Bouček 1970 for type details and comments on accuracy of the original description). Further, molecular analyses should be conducted prior to such synonymy to determine if a single species or a sibling species-complex occurs throughout the Nearctic, Palaearctic and Oriental regions. Catolaccus ater (Ratzeburg 1852) differs from C. crassiceps in having a more distinctly reticulate propodeum, more extensively and densely setose fore wings, and apparently a similar mandibular structure as C. cyanoideus (Dzhanokmen 1980, fig. 5). It is definitely a different species than C. cyanoideus because it has completely bare eyes, is dark non-metallic, and has even more extensively setose fore wings. Only the vannal area and a very slender inconspicuous band on the disc (adjacent to the parastigma and sometimes basally on the mediocubital fold basally) are bare (Dzhanokmen 1980, fig.7).