JHR 43: 45-l I 0 (20 I 5) ore JOURNAL OF A peer-reviewed open-access Journal Sirians (4) Hymenoptera http://jhr.pensoft.net The insertional Society of Hymenoptersts. RESEARCH Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae), natural enemies of native and invasive stink bugs (Hemiptera, Pentatomidae) Elijah J. Talamas', Norman F. Johnson’, Matthew Buffington' I Systematic Entomology Laboratory, USDA/ARS clo NMNH, Smithsonian Institution, Washington DC, USA 2 Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Columbus, OH 43212, USA Corresponding author: Elijah J. Talamas (elijah.talamas@ars.usda.gov) Academic editor: /. Yoder | Received 5 September 2014 | Accepted 10 March 2015 | Published 27 March 2015 http://zoobank. org/400COA04-5 BBO-4653-9A87-535B5CA22D0C Citation: Talamas EJ, Johnson NF, Buffington M (2015) Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae), natural enemies of native and invasive stink bugs (Hemiptera, Pentatomidae). Journal of Hymenoptera Research 43: 45-110. doi: 10.3897/JHR.43.8560 Abstract Trissolcus japonicus (Ashmead) and T. cultratus (Mayr), comb. rev. are under study as classical biological agents to control the brown marmorated stink bug Halyomorpha halys (Stal) in North America. Here we present diagnoses for all Nearctic species of Trissolcus, including T japonicus and T. cultratus comb. rev., and identification keys to enable separation of these species from the existing fauna. Trissolcus cultratus comb. rev. is removed from synonymy with 7. flavipes. Two new species are described, Trissolcus valkyria sp. n. and 7. zakotos sp. n. A neotype is designated for 7’ brochymenae and a lectotype is designated for T’ basalis. Keywords Trissolcus japonicus, Trissolcus cultratus, Trissolcus flavipes, Halyomorpha halys, Trissolcus, Scelionidae, biological control, identification key, egg parasitoid Copyright Elijah J. Talamas et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 46 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Introduction A decade after its introduction into the United States in 1999, the economically de- structive brown marmorated stink bug (BMSB), Halyomorpha halys Stal (Heteroptera: Pentatomidae), has been detected in 39 US states and the District of Columbia, as well as Canada, Switzerland, Germany, France, and Italy, and has been intercepted in New Zealand (Xu et al. 2013). BMSB has an extraordinarily wide host range in both its na- tive range (Asia) and invaded countries where it feeds on over 200 species of tree fruits, vegetables, field crops, ornamental plants, and native vegetation (Hoebeke and Carter 2003; Leskey et al. 2012). Some notable crops attacked include fruit trees (especially apples and pears), corn, wheat, soybean, and grape. In the US in 2010, $37 million in losses to mid-Atlantic apples was recorded; in some pear orchards 100% loss was ob- served (Leskey et al. 2012). Further, BMSB is a well-known nuisance species, invading homes and businesses in the mid-Atlantic region, with over 25,000 individuals being recorded from a single household (Inkley 2012). BMSB is difficult to manage with pesticides because it feeds on interior plant tis- sues via its proboscis, bypassing ingestion of pesticides that are deposited on the sur- faces of plant tissues. As a result, increased pesticide applications to combat BMSB dis- rupt ecosystem services, resulting in secondary pest outbreaks (Leskey et al. 2012). Xu et al. (2013) determined that a single introduction to North America from the Beijing area of China, with secondary migration to the West Coast, is responsible for the pres- ence of this destructive pest. Due to the difficulty and potential non-target effects of controlling this pest with pesticides, foreign exploration of natural enemies of BMSB began in earnest in 2008, with the Beijing area of China as a focal point of collections, followed by additional collecting in South Korea and Japan (Xu et al. 2013). These col- lections have identified two species with potential as classical biological control agents. Both species were initially referred to by incorrect names, and through the examination of primary type specimens, we have identified them as T7issolcus japonicus (Ashmead) and Trissolcus cultratus (Mayr), comb. rev. A community of extension agents, field scientists, and ARS scientists (at the Beneficial Insects Introduction Research Unit and Systematic Entomology Laboratory), are presently studying the host preferences of T: japonicus, T. cultratus, and native species of Trissolcus, and the ability of these species to successfully develop in BMSB eggs. However, such studies are ineffective unless the species can be distinguished reliably — a task that may be challenging for non-experts due to the small size of these insects and their defining characters, as well as historical taxonomic confusion of species. This paper is presented as an updated synthesis of the works of Norman Johnson (1984, 1985a,b) with the addition of four species, 7: japonicus, T. cultratus comb. rev., I. valkyria sp. n. and T. zakotos sp. n., and previously unexplored or unutilized character systems. Following the keys to species, an updated and expanded diagno- sis section provides more nuanced discussions on sexual dimorphism and phenotypic plasticity. The authors hope that this publication will demonstrate the long-term rel- evance of primary taxonomic research. The concepts of the previously described spe- Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 47 cies presented here are based primarily on the works of Johnson (1984, 1985a,b), as are many of the characters used in the identification keys and character data reported in the diagnoses. Without these publications, preparation for the introduction of an exotic species would require revision of the Nearctic fauna to establish the characters by which the introduced species could be recognized. Production of the identification tools provided here would not have been possible within the time frame of the USDA- ARS biocontrol release project, nor is there sufficient funding for revisionary work on Trissolcus in both the Nearctic and the Eastern Palearctic, from which T’ japonicus and T. cultratus comb. rev. originate. The revision of the latter is underway. Although it may be impossible to fully predict which species will become intro- duced pests, educated decisions can be made about which species have the greatest po- tential, and similarly, which species have potential as biological control agents. While this paper was in review, a wild population of T. japonicus was discovered in Beltsville, Maryland (Talamas et al 2015), which we were able to rapidly identify with the tools we produced for exactly this purpose. We believe that this emphasizes the utility of alpha taxonomy and the need for continued revisionary work in Platygastroidea. The identification keys of Johnson (1984, 1985a,b) remain relevant for the Nearc- tic species and the dichotomous key presented here largely follows their structure. The goals of this publication are to document the Nearctic fauna as completely as possible, and to provide identification tools with high resolution color illustrations that should greatly facilitate species-level identification. Two formats for this identification key are given: a traditional dichotomous key, and a multi-choice Lucid key’. The contributions of the authors are as follows: E.J. Talamas: character definition and coding, imaging, manuscript preparation; N.F. Johnson: character definition and coding, manuscript preparation; M. Buffington: manuscript preparation, project coordination. Materials and methods The locality data reported for primary types are not literal transcriptions of the labels: some abbreviations are expanded; additional data from the collectors are also included. The numbers prefixed with “USNMENT” or “OSUC ” are unique identifiers for the individual specimens (note the blank space after some acronyms). Details on the data associated with these specimens may be accessed at the following link, http://purl.ocle. org/NET/hymenoptera/hol, and entering the identifier in the form. The taxonomic synopsis was generated by the Hymenoptera Online Database (http://hol.osu.edu). Persistent URIs for each taxonomic concept were minted by xBio:D in accordance with best practices recommend by Hagedorn et al (2013). Morphological terms were matched to concepts in the Hymenoptera Anatomy Ontology using the text analyzer function. A table of morphological terms and URI links is provided in Suppl. material 1. Photographs were captured with a Z16 Leica®™ lens with a JVC KY-F75U digital cam- @T™M era using Cartograph®™ software, or a Leica DMRB compound microscope with a GT- 48 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Vision®™ Lw11057C-SCI digital camera attached. In both systems, lighting was achieved using techniques summarized in Buffington et al. (2005), Kerr et al. (2009) and Buffing- ton and Gates (2009). Single montage images were produced from image stacks with the program CombineZP*™. In some cases, multiple montage images were stitched together in Photoshop® to produce larger images at high resolution and magnification. Full resolu- tion images are archived at the image database at The Ohio State University (http://purl. oclc.org/NET/hymenoptera/specimage), MorphBank (http://www.morphbank.net), and Hymenoptera Holotypes of the Smithsonian Institution (http://usnmhymtypes.com). High quality optics and bright, diffuse lighting are critical for observing the char- acters in this key. The authors recommend fluorescent desk lamps, or fiber optic lamps with mylar sleeves afhxed to the tips of the light pipes, or a mylar ‘shield’ between the tip of the light pipes and the specimen. Direct illumination of the specimen should be avoided. Additionally, some characters are better observed with appendages moved (especially the legs in couplet 5 and the wings in couplet 6 of the 77issolcus species key). Fine forceps or a minuten pin achieve this effectively. Collections This work is based on specimens deposited in the following repositories with abbrevia- tions used in the text: BMNH _ Natural History Museum, London, England CNCI Canadian National Collection of Insects, Ottawa, Canada NHMW Naturhistorisches Museum Wien, Vienna, Austria NHRS Naturhistoriska riksmuseet, Stockholm, Sweden OSUC C.A. Triplehorn Insect Collection, Columbus, USA USNM — Smithsonian National Museum of Natural History, Washington DC, USA UANL Facultad de Ciencias Forestales, Linares, Mexico LACM Los Angeles County Museum of Natural History, Los Angeles, USA UCRC Entomology Research Museum, Riverside, USA MEMU Mississippi State University MSWC M.S. Wasbauer Collection, Sacramento, USA ANIC Australian National Insect Collection, Canberra City, Australia RMCA Musee Royal de l’Afrique Centrale, Tervuren, Belgium FSCA Florida State Collection of Arthropods, Gainesville, USA Character discussion Axillar crescent We coin this term to refer to the structure formed by the transaxillar, axillar, and axil- lular carinae located posterodorsal to the wing base (see Figs 1, 17, 19-20, 60, 62). The Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 49 transaxillar and axillar carinae are fused in Trissolcus and form the anterodorsal part of the axillar crescent. The axillular carina forms the posterior and ventral portion. Proper examination of this character may require removal or adjustment of the wings. Clypeal setae In the Nearctic fauna, species in the basalis and thyantae species groups have 6 clypeal setae (Fig. 27). Trissolcus cultratus and native species of the flavipes group have 2 (Fig. 23), making it easy to separate 7’ japonicus, which has 4 clypeal setae (Fig. 25). Rarely, and usually in males, superfluous clypeal setae exist. These are typically much smaller and arise near the base of one of the “true” clypeal setae. We consider the number of these setae to be extremely useful for identification of 7’ japonicus, but this character is variable. Episternal foveae The episternal foveae of the thyantae group are clearly defined; they extend from the dorsal limit of the acetabular carina to the mesopleural pit and are typically antero- posteriorly elongate. In the dasalis group, the episternal foveae are often distinctly separate from the mesopleural pit, and with the exception of some J! cosmopeplae, are distinctly separate from the dorsal limit of the postacetabular sulcus. Nearctic species of the flavipes group tend to be variable in the external expression of this character. In most cases, the foveae are irregularly shaped and are at varying distances from both the mesopleural pit and acetabular carina. In the Eastern Palearctic species of the flavipes group the episternal foveae often appear as a continuation of the postacetabular sulcus and extend dorsally to the mesopleural pit as in 7’ japonicus (Fig. 70) and 7. cultratus. Facial striae The presence of striae on the frons is typically weakly indicated or entirely absent, with a few exceptions. In some species the striae are present as shallowly incised short lines arising from the anterior articulation of the mandible (eg. 7’ cultratus, Fig. 54), and in a few, T’ radix, T. solocis and T: zakotos, the striae exist as rugulae that extend further toward the compound eye (Fig. 112). Mesopleural carina The mesopleural carina was used more extensively for species identification in the key of Johnson (1984) than it is here. Specifically, we observed that in 7’ edessae this carina may be present (Fig. 58) and we no longer use its absence to separate this species. In the thyantae and basalis groups this character exhibits far less intraspecific variability than in the flavipes group and we use it for the identification and delimitation of LT valkyria. Mesoscutal humeral sulcus Among the published descriptions and diagnoses, we have not encountered previous use of this character for species-level delimitation in 77issolcus. In all but one species, 50 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) T. cosmopeplae, the form of this character is fixed. As stated by Johnson (1985), T. cos- mopeplae, as currenly understood, is a highly variable species. We point out that most specimens of 7! cosmopeplae examined for this key have a mesoscutal humeral sulcus present as a smooth furrow, and that in the holotype specimen this sulcus is comprised of distinct cells. aoc Figures |-2. | 77issolcus, lateral view 2 Trissolcus, dorsal view. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... Characters annotations Al—12 ac aem anfo aoc antennomeres 1—12 (Fig. 1) acetabular carina (Figs 43-44, 70) anteroventral extension of metapleuron (Figs 1, 43, 58, 82, 88) antennal foramen (Fig. 1) anterior ocellus (Fig. 2) antennal scrobe (Figs 13, 26, 54) antecostal sulcus (Figs 2, 29, 61) postacetabular sulcus (Figs 42, 70, 82) axillar crescent (Figs 1, 17, 19-20, 60, 62) basiconic sensilla (Figs 8—9) clypeal setae (Fig. 23, 25, 27) central keel (Figs 7, 10, 79) procoxa (Fig. 1) mesocoxa (Fig. 1) metacoxa (Fig. 1) episternal foveae (Figs 35, 43, 48, 70, 82) facial striae (Figs 7, 10, 54, 112) genal carina (Figs 48, 112) gena (Fig. 1) hyperoccipital carina (Figs 2, 19, 29) interantennal process (Fig. 1) lateral ocellus (Figs 1—2) laterotergite(s) (Figs 2, 11-12) mesopleural carina (Fig. 43, 45, 58, 107) mandible (Fig 1) median mesoscutal carina (Fig. 39, 63) median mesoscutal sulcus (Fig. 93) mesopleural pit (Figs 35, 43, 82) malar sulcus (Figs 1, 24, 26, 28) mesoscutum (Figs 1—2) metascutellum (Figs 1-2, 21—22) mesoscutal humeral sulcus (Figs 29, 30, 46, 49) mesopleuron (Fig. 1) metanotum (Figs 1-2, 21—22) metapleuron (Fig. 1) metapostnotum (Figs 1-2, 21-22) netrion sulcus (Figs 44, 48) notaulus (Figs 2, 29) orbital furrow (Figs 1, 24, 26, 28-29, 71) paracoxal sulcus (Figs 82, 109) Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) propodeum (Figs 1—2, 21—22) pronotum (Figs 1—2) propleuron (Fig. 1) posteroventral portion of metapleuron (Figs 17, 18, 20, 74, 106) radicle (Figs 1, 37, 80, 112) mesoscutellum (Figs 1—2) sublateral seta (Figs 2, 30, 89) mediotergite (Figs 1-2) tegula (Figs 1—2) Key to genera of Nearctic Platygastroidea known to attack pentatomoid eggs The following key includes platygastroids with host records indicating emergence from pentatomoid eggs. More associations are certain to exist, particularly in Zélenomus, which contains many species with undocumented biology, and many undescribed species. 1 Metasoma with laterotergites tightly appressed to sternites, forming a sharply angled lateral margin (Fig. 11); female antenna with 12 antennomeres CEI Pilsner rterecanceee settee ner stems meets ier Gryon obesum Masner Metasoma with laterotergites wide and loosely attached to sternites, meta- soma without sharp lateral margin (Fig. 12); female antenna with 10 or 11 AN LOMO CLES 1 kt3. hae casving Soavtaeathertes hosetatachephacets Bua sun asbinnn ete: pha Ricetaatin: oat 2 Frons with central keel extending from interantennal process to anterior ocel- lus (Figs 7, 10); frons with facial striae distinct, striae often extending along inner orbit.ofcompound. cye (Pigs 73 VO) ce scgctes coves dasa uncoseabhatiecackssesaensts 3 Frons without central keel or keel short, not extending to anterior ocellus (Figs 54, 79); frons without facial striae or, if present, sinuate and usually at- tenuating before reaching inner orbit of compound eye (Figs 54, 112)....... 4 Mesoscutum with notauli (Fig. 5); base of metasoma usually yellow-orange and contrasting with dark color of posterior metasoma (Figs 3, 5); in lateral view, procoxa distinctly separated from mesocoxa (Fig. 3) ....ceseseeseeseceeeeees Beek hath bie 2, SP iey Brg TE x doy UE hte nce Be Paratelenomus saccharalis (Dodd) Mesoscutum without notauli (Fig. 6); base of metasoma never yellow-orange (Fig. 4); in lateral view, procoxa contiguous with mesocoxa (Fig. 4) «0.0.0.0... AE) EL Mee Oe Ae ON e, SEM AE A Ne eid NER AE Psix tunetanus (Mineo & Szab6) T2 longer than wide (Fig. 14); frons predominantly smooth and shining (Fig. 13); female antenna with basiconic sensilla on apical 4 (rarely 5 or 6) anten- MOULETES (Hig. )) aikvusaecesl cee Telenomus (T. astrictus, T. calvus, T. golia- thus, T. grenadensis, T. persimilis, T. podisi, T. sanctiventris, T. scaber) T2 wider than long (Fig. 16); frons with microsculpture throughout, often superimposed on coarse surface sculpture (Fig. 15); female antenna with ba- siconic sensilla on apical 5 antennomeres (Fig. 8) .......ceeeeeseeeeees Trissolcus Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... Se) S . “y i ai aT , a) - ‘ Oa 0.1 Figures 3-4. *' 3 Paratelenomus saccharalis, female (USNMENT00896342), head, mesosoma, meta- soma, lateral view 4 Psix tunetanus (USNMENT00989625), head, mesosoma, metasoma, lateral view. Scale bars in millimeters. Key to species of Nearctic Trissolcus (males and females) 1 Metapleuron with posteroventral portion glabrous (Figs 17, 19—20)........... 2 Metapleuron with posteroventral portion setose (Figs 18, 73-74, 76, 106)... Pee Ce COR nr CRS, Oat (thyantae group) 15 Vertex with hyperoccipital carina (Figs 19, 29, 64); mesoscutum with notauli (Figs 21, 29); clypeus with 4 or fewer setae (Figs 23, 25); inner margin of eye with orbital furrow not uniform in width, usually expanded near malar sulcus si cecal De 02 8) bane cuneate Mes tn cael ti (flavipes group) 3 Vertex without hyperoccipital carina (Fig. 30); mesoscutum usually without notauli (Fig. 30); clypeus with 6 setae (Fig. 27); inner margin of eye with 54 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 5-10.” 5 Paratelenomus saccharalis, female (USNMENT00896342), female, head, mesosoma, metasoma, dorsal view 6 Psix tunetanus, female (USNMENT00989625), head and mesosoma, dorsal view 7 Psix tunetanus, female (USNMENT00877258), head, anterior view 8 Tyissolcus strabus, female (USNMENT00954423), antennal clava, ventral view 9 Telenomus sp., female (OSUC 523925), antennal clava, ventral view 10 Paratelenomus saccharalis, female (USNMENT00896364), head and mesosoma, anterolateral view. Scale bars in millimeters. orbital furrow uniform in width, not expanded near malar sulcus (Fig. 24) .. Bessa es oesase ct eee ee ss Es AU VS Dc (basalis group) 8 3 Frons between antennal scrobe and anterior ocellus with parallel, arched ru- gabon] Sika SNES 129 Eee RE Oty Ree OE T. cultratus (Mayr), comb. rev. ~ Frons between antennal scrobe and anterior ocellus smooth or with rugae thatrarenotparalleland arched (Rigs 26s 405 9559526) eo scue-psesere enacts 4 Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 55 Figures | 1-12.” 11 Gryon obesum, female paratype (USNMENT00989078), head, mesosoma, meta- soma, anterolateral view 12 Tvissolcus euschisti, female (OSUC 404912), head, mesosoma, metasoma, ventrolateral view. Scale bars in millimeters. 4 Inner margin of eye with orbital furrow constricted ventrally (Fig. 26); mes- oscutellum rugose, at least laterally and usually throughout (Fig. 32); first laterotergite usually with setae (Fig. 92); mesoscutum often with median WE SOSCUEAIPS UM CUES UPC.) art Oe cage ERE A sees TOBA E T. strabus Johnson 56 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) — Figures 13-16. 13 Telenomus sp., female (USNMENT00872628), head, anterolateral view 14 Tele- nomus sp., female (USNMENT00903997), metasoma, dorsal view 15 Trissolcus urichi, female holotype (USNMENT00989070), head, anterolateral view 16 77issolcus urichi, female (USNMENT00896405), metasoma, dorsal view. Scale bars in millimeters. = Inner margin of eye with orbital furrow expanded near intersection with ma- lar sulcus (Figs 12, 15, 28); mesoscutellum smooth (Fig. 34) or with coria- ceous microsculpture (Fig. 33); first laterotergite without setae (Figs 62, 68); mesoscutum without median mesoscutal sulcus Fig. 69)... eee eeeeseeeeeeee 5 5 Clypeus with 4 setae (Fig. 25); mesopleuron with episternal foveae well-de- fined and deep, forming a continuous line of cells from postacetabular sulcus to mesopleural pit (Fig. 70); mesoscutum without median mesoscutal carina CE LOAD) Ea Ie at de et a ais a canes UE ord T. japonicus (Ashmead) — Clypeus with 2 setae (Fig. 23); mesopleuron with episternal foveae poorly defined, often shallow, irregular (Figs 42-45) and typically distant from postacetabular sulcus; mesoscutum often with median mesoscutal carina (Figs 39, 63) 0... 6 6 Female with antennal flagellum distinctly bicolored: A3—A6 yellow, A7—A11 dark brown (Fig. 59, as in Fig. 68); area bounded by axillar crescent (axcr Fig. TEstriareg smite POSH 8S) Vc oseon atl aicnessios a > ; 22 + Se a Figures 17-22.” 17 7. japonicus, female (USNMENT00675989), mesosoma, lateral view 18 T. oc- ciduus, female (USNMENT00764995), mesosoma, lateral view 19 Tvissolcus cultratus, female (USN- MENT00764850), head and mesosoma, dorsolateral view 20 7: euschisti, female (OSUC 523871), mesosoma, lateral view 21 T. japonicus, female (USNMENT00675989), mesosoma, posterolateral view 22 T. hullensis, female (OSUC 523881), mesosoma, dorsolateral view. Scale bars in millimeters. 7 Mesopleuron with anteroventral portion rugulose (Figs 41, 44-45)... Dain Mires ges as ae ae eas ana eee T. brochymenae (Ashmead) — Mesopleuron with anteroventral portion smooth or with shallowly impressed microseulpuure (Figs 2, 42-43) re.ccuue. rane canator. T. euschisti (Ashmead) 8 Mesoscutellum coarsely rugose (Figs 80-81, 90); vertex sharply angled onto OS eI IE NY) etann.8.84r.t0e Shar ashen sae tee damn Ete ee eke Asn. yA 9 58 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) 57} rad : ape" , -) Figures 23-28. *° 23 7) strabus, female (BMSB 1203), mouthparts, anterior view 24 T: hullensis, fe- male, (OSUC 523881), head and mesosoma, anterolateral view 25 T: japonicus, female holotype, (US- NMENT00831865), mouthparts, anterior view 26 T° strabus, female, (BMSB 1202), head mesosoma, anterolateral view 27 T. erugatus, female, (OSUC 523929), head mesosoma, anterolateral view 28 T° eus- chisti, female, (BMSB 1223), head mesosoma, anterolateral view. Scale bars in millimeters. — Mesoscutellum smooth or with coriaceous microsculpture (Figs 33-34); ver- exer Oulmdedhonto-oeciputn(h ies. 30s 66) ws loouaise nes msennsenpsnserorsicacest cea 10 9 Radicle yellow (Fig. 80, 83); metapleuron with paracoxal sulcus indicated by lineof distincyfoveaciniventraldnali (Fie."82)).c2- Prenat: T. radix Johnson — Radicle dark brown to black (Fig. 91); metapleuron with paracoxal sulcus absent or obscured by rugae in ventral half (Fig. 88)........ T. solocis Johnson Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... ob 30 A 0.2 Figures 29-30. °’ 29 7. strabus, female (BMSB 1202), head mesosoma, metasoma, dorsolateral view 30 7: basalis, female, (USNMENT00954022), head, mesosoma, metasoma, dorsolateral view. Scale bars in millimeters. 10 Metapleuron with paracoxal sulcus visible in ventral half (Fig. 109); frons with facial striae extending as rugulae from anterior mandibular articulation toward compoundicye (Fics 2)yscsetereaaws, T. zakotos Talamas, sp. n. — Metapleuron with paracoxal sulcus absent or obscured by coarse rugae in ventral half (Figs 35, 46, 67); frons without facial striae (Fig. 37) ............. 11 11 T2 smooth or with faintly impressed striation posterior to antecostal sulcus Ee SRON PIO) pt. cnn cascurmmnscunheas ensue toads ctoetueutiasensennelta sath axanmmeis 12 — T2 with pronounced striae posterior to antecostal sulcus (Figs 30, 51, 101)... 13 12 Metapostnotum invaginated near propodeal spiracle, not separating propodeum from metanotum near metascutellum (Figs 22, 66)..... 72 bullensis (Harrington) 60 186 14 i) 16 LF 18 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Metapostnotum invaginated near metascutellum, separating propodeum from metanotum near metascutellum (as in Fig. 21).... T. erugatus Johnson Mesoscutellum with distinct coriaceous microsculpture and setal bases usually pustulate (Fig. 31); mesopleuron with episternal foveae shallowly impressed and distinctly separated from mesopleural pit (Fig. 35); netrion sulcus incom- plete (as ti Fig AA). cee atin eens sen cdaet Bodiuinssdea ds auc teeeae T. basalis (Wollaston) Mesoscutellum entirely smooth and setal bases not strongly raised (Fig. 34); mesopleuron with episternal foveae extending dorsally to proximity of meso- pleural pit (Fig. 48); netrion sulcus complete (Fig. 48) 0... eeeeeeeeeeeee 14 Gena in lateral view bulging (Fig. 102), without genal carina (Fig. 102); mes- oscutum without notauli (Fig. 100); anteroventral extension of metapleuron short, not extending to mesocoxa in lateral view (Fig. 102); lateral mesoscu- tum with mesoscutal humeral sulcus present as a smooth furrow (as in Fig. 6 (Uh adden te BNE i 2 De hae Ae es 8 oa Be Ba EE ER, nx bl T. utahensis (Ashmead) Gena in lateral view narrow, often with genal carina extending dorsally from base of mandible (Fig. 48); mesoscutum with notauli sometimes indicated; anteroventral extension of metapleuron usually long and extending to base of mesocoxa in lateral view (Fig. 46, as in Fig. 43); lateral mesoscutum with mesoscutal humeral sulcus present as a smooth furrow (Fig. 49) or comprised OiCell SACHS. AG) eet teeta aa earn ae teat T. cosmopeplae (Gahan) Mesoscutellum covered with shallowly impressed coriaceous microsculpture (EEG SS) otra anc iS acten setts Mew Meanterec eral icone tateeaee te ucte tees a wees sedanisnds yee xd 16 Mesoscutellum entirely smooth, without microsculpture (Fig. 34) ........... 17 Frons outside of antennal scrobes with raised, irregular rugulae (Fig. 86); mesoscutum between notauli often with longitudinal rugulae (Fig. 87) ........ aise eese tet Cte ice, ope dere oo ieee art eal. ee T. ruidus Johnson Frons outside of antennal scrobes coriaceous, without raised rugulae but with more or less well-defined setigerous punctures, (Fig. 79); mesoscutum with- out longitudinal elements in sculpture (Fig. 77)............. T. parma Johnson Gena in lateral view bulging (Fig. 74)... ee eeeeeeeeeee T. occiduus Johnson Genavinidlateral viewmarrow CEIgsi9 9). LOG). a. oncesssncesenchndsorducebenpentbennmntesnat 18 Mesopleural carina absent ventrally (Fig. 97)............. T. thyantae Ashmead Mesopleural carina. coriplete (Eig 107) 100 eseccs.taisetenstictebecsoens ededoarcdseneseseete A ee oe ee ee Wey ao eee T. valkyria Johnson & Talamas, sp. n. Multi-choice Lucid key Lucid Key Server edition (only web browser required): http://keys.lucidcentral.org/key-server/key.jsp?keyIld=127 Applet edition (requires installation of Java Runtime Environment): hetp://keys.lucidcentral.org/keys/v3/Nearctic_Trissolcus/ Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 61 Taxonomy Trissolcus basalis (Wollaston) http://bioguid.osu.edu/osuc_concepts/3 189 Figures 30-31, 35-37; Morphbank* Telenomus Maderensis Wollaston, 1858: 25 (original description, synonymized by Nixon (1935)). Telenomus basalis Wollaston, 1858: 25 (original description); Kieffer 1926: 39 (description). Telenomus megacephalus Ashmead, 1894: 203, 212 (original description, synonymized by Nixon (1935)); Ashmead 1896: 790 (keyed); Ashmead 1900: 326 (distribution); Nixon 1935: 100 (junior synonym of Microphanurus basalis (Wollaston)). Telenomus megalocephalus Schulz: Schulz 1906: 152 (emendation). Telenomus piceipes Dodd, 1920: 354 (original description, synonymized by Nixon (1935)); Nixon 1935: 100 (junior synonym of Microphanurus basalis (Wollaston)). Liophanurus megacephalus (Ashmead): Kieffer 1926: 65, 76 (description, generic transfer, keyed). Telenomus maderensis Wollaston: Kieffer 1926: 39 (description); Nixon 1935: 100 (junior synonym of Microphanurus basalis (Wollaston)). Microphanurus basalis (Wollaston): Nixon 1935: 96, 100 (description, generic transfer, syno- nymy, keyed); Nixon 1943: 138 (keyed); Risbec 1950: 570, 571 (variation, keyed). Asolcus basalis (Wollaston): Delucchi 1961: 44, 57 (description, keyed); Voegelé 1962: 155 (variation, diagnosis); Voegelé 1964: 28 (keyed); Voegelé 1965: 96, 108 (variation, diagnosis, keyed); Voegelé 1969: 151 (keyed). Trissolcus basalis (Wollaston): Masner 1965: 125 (type information, generic transfer); Safavi 1968: 415 (keyed); Fabritius 1972: 31 (keyed); Kozlov and Lé 1977: 516 (keyed); Kozlov 1978: 637 (description); Kozlov and Kononova 1983: 121 (de- scription); Graham 1984: 100 (variation); Johnson 1985b: 432, 434 (description, keyed); Johnson 1991: 212, 213, 214, 216 (diagnosis, keyed); Ghahari, Buhl and Kocak 2011: 594 (listed); Mao, Valerio, Austin, Dowton and Johnson 2012: 194 (presentation of mitochondrial genome, phylogenetic position); Fusu, Bin and Popovici 2013: 263 (description of chromosomes). Trissolcus maderensis (Wollaston): Masner 1965: 126 (type information, generic transfer). Trissolcus piceipes (Dodd): Masner 1965: 127 (type information, generic transfer). Trissolcus megacephalus (Ashmead): Johnson 1983: 448 (type information). Lectotype designation. Masner (1965) did not mention the type status of the spec- imen labeled “Type H.T.” in his treatment of the types in BMNH, and Johnson (1985) referred to this specimen of the holotype, although it was originally part of a syntype series. Consequently, a lectotype was not actually designated for T: basalis. We here designate the specimen mentioned by Masner (1965) (B.M. TYPE HYM. 9.304) as the lectotype of this species. 62 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) * ‘ 3 enh ‘ey Figures 31-34." 31 7° basalis, female (USNMENT00954023), mesoscutellum, dorsal view 32 T. stra- bus, female (OSUC 523850), mesoscutellum, dorsal view 33 7: ruidus, (OSUC 76431), mesoscutellum, dorsal view 34 T. occiduus, female (OSUC 76430), mesoscutellum, dorsal view. Scale bars in millimeters. Diagnosis. Within the New World species of the basalis group, the combination of the broadly rounded vertex, wide gena, and rugose T2 is found only in T° basalis and T. utahensis. Trissolcus basalis may be distinguished by its coriaceous mesoscutellum, incomplete netrion sulcus and weakly developed episternal foveae. Trissolcus basalis may be dark in color, but typically can be distinguished by the yellow scape (sharply contrasting in color with the dark radicle) and abruptly bicolored antennae. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3189] Associations. Emerged from egg of Aelia Fabricius: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Aelia acuminata (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Ae- lia cognata Fieber: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Aelia germari Kiister: [Hemiptera: Heteroptera: Pentatomoidea: Pentato- midae]; emerged from egg of Agonoscelis rutila (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Asopinae Spinola: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Calidea Laporte: [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; emerged from egg of Calidea dregeii Germar: [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; emerged from egg of Carpocoris fuscispinus (Boheman): [Hemiptera: Heteroptera: Pentato- Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 63 moidea: Pentatomidae]; emerged from egg of Coleotichus blackburniae: |Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; emerged from egg of Cuspicona simplex Walker: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Dolicoris baccharum (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pen- tatomidae]; emerged from egg of Dolycoris baccarum (Linnaeus): [Hemiptera: Het- eroptera: Pentatomoidea: Pentatomidae]; emerged from egg ofEurydema ornata (Lin- naeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Eurygaster austriaca (Schrank): [Hemiptera: Heteroptera: Pentatomoidea: Scutel- leridae]; emerged from egg of Eurygaster integriceps Puton: [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; emerged from egg of Euschistus Dallas: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Euschistus servus (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Euthyrhynchus floridanus (Linnaeus): [Hemiptera: Heteroptera: Pentato- moidea: Pentatomidae]; emerged from egg of Graphosoma semipunctata (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Ha- lyomorpha annulicornis (Signoret): [Hemiptera: Heteroptera: Pentatomoidea: Pentato- midae]; emerged from egg of Nezara Amyot & Serville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Nezara Amyot & Serville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; egg parasite of Negara vir- idula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentato- midae]; emerged from egg of Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; parasite of Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Odontotarsus grammicus (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; egg ectoparasite of Oechalia Stal: [He- miptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Oechalia schellenbergi Guérin-Méneville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomi- dae]; emerged from egg of Pentatomidae: |Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Pentatomidae: [Hemiptera: Heteroptera: Pen- tatomoidea: Pentatomidae]; emerged from egg of Piezodorus hybneri (Gmelin): [He- miptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Plautia af- finis (Dallas): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from ege of Raphigaster Laporte: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Solanum nigrum L.: [Solanales: Solanaceae]; collected on bokhara: [Fa- bales: Fabaceae]; collected on buchan: [Capparales: Brassicaceae]; collected on cotton: [Malvales: Malvaceae]; collected on hore: [Lamiales: Lamiaceae]; collected on mint: [Lamiales: Lamiaceae]; collected on mung: [Fabales: Fabaceae]; collected in drilled soybean: [Fabales: Fabaceae]; collected on soybean: [Fabales: Fabaceae] Material examined. Lectotype, female, 7’ basalis:s PORTUGAL: Madeira Reg. Auté., Madeira Island, VII-1855, Wollaston, B.M. TYPE HYM. 9.304 (deposited in 64 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 35-37. ° Trissolcus basalis 35 female (USNMENT00903007), head, mesosoma, metasoma, lateral view 36 female (USNMENT00872109), head and mesosoma, dorsal view 37 female (OSUC 75271), head, anterior view. Scale bars in millimeters. BMNH). Holotype, female, 7: megacephalus: SAINT VINCENT AND THE GREN- ADINES: Saint Vincent Island, no date, H. H. Smith, USNM Type No. 2525 (deposited in USNM). Other material: (58 females, 15 males, 393 sex unrecorded) AUSTRALIA: 8 females, 4 males, 149 sex unrecorded, ANIC DB 32-020991, 32-020992, 32-020993, 32-020994, 32-020996—32-020999 (ANIC); OSUC 17738 (BMNH); OSUC 75398- 75424 (OSUC); OSUC 145814, 78027-78147 (QDPC); USNMENT00872088, USNMENT00872089, USNMENT00872090, USNMENT00903007 (USNM). BRAZIL: 67 sex unrecorded, OSUC 75299-75365 (OSUC). CHINA: 10 sex un- recorded, OSUC 75366, 75389-75397 (OSUC). DEMOCRATIC REPUBLIC OF THE CONGO: 3 females, OSUC 182081-182083 (RMCA). DOMINICAN RE- PUBLIC: 3 sex unrecorded, OSUC 398658—398659, 398667 (CNCI). EGYPT: 2 females, 4 sex unrecorded, OSUC 144795—144796, USNMENT00872006, USN- MENT00872007, USNMENT00872008, USNMENT00872009 (USNM). ERI- TREA: 1 female, OSUC 17736 (BMNH). FIJI: 16 sex unrecorded, OSUC 77661- 77676 (BPBM). FRENCH POLYNESIA: 2 sex unrecorded, OSUC 77659-77660 (BPBM). GREECE: 1 sex unrecorded, OSUC 398669 (CNCI). IRAN: 1 sex unre- Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 65 corded, OSUC 144797 (CNCI). ITALY: 2 females, 1 male, OSUC 173847-173849 (OSUC). JAMAICA: 2 sex unrecorded, OSUC 398660—398661 (CNCI). JAPAN: 1 sex unrecorded, OSUC 144391 (CNCI). MONTSERRAT: 12 sex unrecorded, OSUC 398662 (CNCI); OSUC 145281 (FSCA); OSUC 75289-75298 (OSUC). MOROC- CO: 1 sex unrecorded, OSUC 17743 (BMNH). NEW CALEDONIA: 1 sex unre- corded, OSUC 77624 (BPBM). OCEANIA: 5 sex unrecorded, OSUC 77625-77628 (BPBM); OSUC 75425 (OSUC). SAINT VINCENT AND THE GRENADINES: 3 sex unrecorded, OSUC 143816—-143818 (LACM). SENEGAL: 1 female, OSUC 17737 (BMNH). SOUTH AFRICA: 6 sex unrecorded, OSUC 145553, 75384-75388 (OSUC). TANZANIA: 1 sex unrecorded, OSUC 17741 (BMNH). TONGA: 31 sex unrecorded, OSUC 77629-77658 (BPBM); OSUC 75427 (OSUC). TRINIDAD AND TOBAGO: 2 sex unrecorded, USNMENT00764950, USNMENT0076495 1 (USNM). TURKEY: 3 females, OSUC 17739-17740, 17742 (BMNH). UNITED STATES: 38 females, 9 males, 49 sex unrecorded, ANIC DB 32-020995 (ANIC); OSUC 398668 (CNCI); OSUC 131149-131186, 154353, 157486—157487, 157542— 157549, 157563-157566, 7339, 75256-75288 (OSUC); USNMENT00872103, USNMENT00872104, USNMENT00872105, USNMENT00872106, USN- MENT00872107, USNMENT00872108, USNMENT00872109 (USNM). VA- NUATU: 1 male, 1 sex unrecorded, ANIC DB 32-020997 (ANIC); OSUC 75426 (OSUC). ZIMBABWE: 17 sex unrecorded, OSUC 75367-75383 (OSUC). Trissolcus brochymenae (Ashmead) http://bioguid.osu.edu/osuc_concepts:3 195 Figures 38-41, 44-45; Morphbank’° Telenomus Crochymenae Ashmead, 1881: 181 (original description, spelling error). Telenomus brochymenae Ashmead: Ashmead 1887: 118 (emendation). Trissolcus brochymenae (Ashmead): Ashmead 1893: 162, 164 (generic transfer, descrip- tion, keyed); Brues 1916: 549, 550 (description, keyed); Kieffer 1926: 127, 129 (description, keyed); Masner 1964: 146 (variation); Masner and Muesebeck 1968: 72 (lectotype designation); Johnson 1984: 799 (description, synonymy, keyed); Johnson 1987: 289, 298 (diagnosis, variation, synonymy, keyed). Trissolcus murgantiae Ashmead, 1893: 162, 163 (original description, keyed, synonymized by Johnson (1984)); Brues 1916: 549, 550 (description, keyed); Kieffer 1926: 127, 128 (description, keyed); Masner and Muesebeck 1968: 73 (lectotype designation); Johnson 1984: 799 (junior synonym of Trissolcus brochymenae (Ashmead)). Trissolcus rufiscapus Ashmead, 1893: 162, 163 (original description, keyed, synonymized by Johnson (1984)); Kieffer 1926: 127, 129 (description, keyed); Masner and Muesebeck 1968: 73 (type information); Johnson 1984: 799 (junior synonym of Trissolcus brochymenae (Ashmead)). Trissolcus laticeps Ashmead, 1894: 212 (original description, synonymized by Johnson (1987)); Ashmead 1900: 326 (distribution); Kieffer 1926: 127, 130 (descrip- 66 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) tion, keyed); Masner 1965: 126 (type information); Johnson 1983: 448 (lecto- type designation); Johnson 1987: 298 (junior synonym of Trissolcus brochymenae (Ashmead)). Neotype designation. The last known examination of the lectotype of 7’ brochymenae was by Johnson (1984) in his revision of the flavipes species group. The specimen was returned to USNM intact but presently consists of a pin, labels, and an empty point. Trissolcus brochymenae is the type species of Trissolcus and we consider the designation of a neotype to be important for the stability of both the genus and species names. Additionally, 7’ brochymenae is a morphologically variable species with a geographic distribution that spans the United States. A case study of cryptic species within 77is- solcus was recently presented by Matsuo et al. (2014) and a similar phenomenon may exist in other species, including 7’ brochymenae. Trissolcus brochymenae is morphologi- cally very close to 7! euschisti, separable by only a few characters, and in our opinion this increases the need for an objective neotype. The specimen selected for this pur- pose was originally a syntype, reared from the same egg mass as the lectotype and is consistent with Ashmead’s original description and the most thorough treatment of the species (Johnson 1984). In accordance with article 75 of The Code we hereby designate specimen USNMENT00965611 (Figs 38-41) as the neotype of Trissolcus brochymenae, deposited in the insect collection of the National Musuem of Natural History (USNM). Diagnosis. Trissolcus brochymenae is most similar to T. euschisti and may be distin- guished from it by the strongly rugulose ventral portion of the mesepisternum anterior to the mesopleural carina (Figs 44-45). This species is also similar to 7° euschisti in that it shows a great deal of variability, presumably in association with its wide geographic distribution and host range. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3195] Associations. emerged from Acrosternum hilare (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Acrosternum impicticorne (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Bro- chymena arborea (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Hydrangea L.: (Rosales: Hydrangeaceae]; emerged from Murgantia his- trionica (Hahn): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Murgantia histrionica (Hahn): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Piezodorus guildini (Westwood): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Piezodorus guildi- nii (Westwood): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from sentinel egg mass of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pen- tatomoidea: Pentatomidae]; emerged from egg of Podisus nigrolimbatus (Spinola): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Prosopis L.: [Fabales: Fabaceae]; collected on cotton: [Malvales: Malvaceae]; collected on rose: [Ro- sales: Rosaceae]; collected on soybean: [Fabales: Fabaceae]; living in soybean: [Fabales: Fabaceae]; emerged from egg of stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 67 Figures 38-41. °° 7rissolcus brochymenae, female neotype (USNMENT00954611) 38 lateral habitus 39 dorsal habitus 40 head, anterior view 41 head and mesosoma, anterolateral view. Scale bars in mil- limeters. 68 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 42-45. °' 42 7. euschisti female (OSUC 523902), mesosoma, ventrolateral view 43 T° euschisti female (OSUC 404912), mesosoma, ventrolateral view 44 T. brochymenae (BMSB 1216), mesosoma, anterolateral view 45 T: brochymenae (USNMENT00896401), mesosoma, ventrolateral view. Scale bars in millimeters. Pentatomidae]; collected on turnip greens: [Capparales: Brassicaceae]; collected on wax myrtle: [Myricales: Myricaceae] Material examined. Neotype, female, 7. brochymenae. UNITED STATES: FL, Duval Co., Jacksonville, no date, reared from egg, W. H. Ashmead, USN- MENT00965611 (deposited in USNM). Lectotype, female, 7: /aticeps: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, no date, H. H. Smith, USNM Type No. 2526 (deposited in USNM). Lectotype, female, 7) mur- gantiae. UNITED STATES: LA, East Baton Rouge Parish, Baton Rouge, no date, reared from egg, H. A. Morgan, USNMENT00989032 (deposited in USNM). Holotype, female, 7. rufiscapus: UNITED STATES: Washington, 12.IV.1885, USNMENT00989047 (deposited in USNM). Paralectotype: UNITED STATES: 1 female, USNM Type No. 2231 PLT (USNM). Other material: (71 females, 2 males, 236 sex unrecorded) BRAZIL: 159 sex unrecorded, OSUC 398724—398725 (CNC]T; OSUC 373344—-373345, 495206-495305, 75445-75499 (OSUC). COLOMBIA: 1 sex unrecorded, OSUC 398719 (CNCI). COSTA RICA: 7 sex unrecorded, OSUC 398702-398703, 398714 (CNCI); OSUC 142482-142485 (OSUC). DOMINICAN REPUBLIC: 4 sex unrecorded, OSUC 3987 10-3987 12, Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 69 398716 (CNCI). GUATEMALA: 1 sex unrecorded, OSUC 398718 (CNCI). HONDURAS: 1 sex unrecorded, OSUC 398717 (CNCI). JAMAICA: 3 sex un- recorded, OSUC 398708-398709, 398713 (CNCI). TRINIDAD AND TOBA- GO: 2 sex unrecorded, OSUC 398706—398707 (CNCI). UNITED STATES: 71 females, 2 males, 45 sex unrecorded, OSUC 17821 (BMNH); OSUC 398679-— 398688 (CNCI); OSUC 436701 (LACM); OSUC 145555, 157494-157503, 266797, 413700-413703, 413709-413713, 523852-523855, 523857-523861, 523864—-523865, 523904—-523923, 523931, 523933, 523935-523937, 523940, 523942, 523944, 542440, 542442, 542445, 542450, 542453-542454, 62796, 70464-70465, 75434-75444, 76425-76426 (OSUC); BMSB 1216-1217, OSUC 145648, USNMENT00872091, USNMENT00872092-USNMENT00872095, USNMENT00989146—USNMENT00989149, USNMENT00989160—USN- MENT00989170, USNMENT00989173 (USNM). VENEZUELA: 5 sex unre- corded, OSUC 398704, 398720—398723 (CNCI). VIRGIN ISLANDS: 2 sex un- recorded, OSUC 398705, 398715 (CNCI). Trissolcus cosmopeplae (Gahan) http://bioguid.osu.edu/osuc_concepts:3206 Figures 46—49; Morphbank* Telenomus cosmopeplae Gahan, 1926: 67 (original description). Trissolcus cosmopeplae (Gahan): Krombein and Burks 1967: 297 (generic transfer); Masner and Muesebeck 1968: 72 (type information); Johnson 1985b: 432, 436 (description, keyed). Diagnosis. Trissolcus cosmopeplae may be distinguished from other species that have sublateral setae and a narrow gena (7! erugatus, I; hullensis, 1: radix, T; solocis, and T: zakotos) by the presence of extensive rugulae on T2 and the mesoscutellum without macrosculpture. This is also the only New World species outside the thyantae and flavipes groups in which notauli may be visible. All other species with sublateral setae and a narrow gena usually have the posterior region of the mesoscutum longitudinally rugulose and the notauli, if present, are thus obscured. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3206] Associations. Emerged from egg of Cosmopepla bimaculata (Thomas): [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Euschistus conspersus Uhler: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: |Hemiptera: Heteroptera: Pentatomoidea: Pentatomi- dae]; emerged from egg of Perillus bioculatus (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on alfalfa: [Fabales: Fabaceae]; collected on bitterbrush: [Rosales: Rosaceae]; collected on blackberry: [Rosales: Rosaceae]; collected on brome: [Cyperales: Poaceae]; collected on red-osier dogwood: [Cor- nales: Cornaceae] 70 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) - ) ° fiw 4 jj se ae, We iy ~~ vd lae 46 female holotype (USNMENT00989096), head, mesosoma, metasoma, lateral view 47 female (OSUC 77129), head, mesosoma, metasoma, dorsal view 48 female (OSUC 77129), head and mesosoma, ventrolateral view 49 female (OSUC 77129), head and mesosoma, dorsolateral view. Scale bars in millimeters. Figures 46-49.» Trissolcus cosmopep Material examined. Holotype, female, 7: cosmopeplae: UNITED STATES: IL, Champaign Co., Urbana, 8.VIII.1925, reared from egg, USNMENT00989096 (de- posited in USNM). Other material: (9 females, 1 male, 74 sex unrecorded) CANA- DA: 16 sex unrecorded, OSUC 145181, 398732—398743 (CNCI); OSUC 145556, 75612-75613 (OSUC). UNITED STATES: 9 females, 1 male, 58 sex unrecorded, OSUC 398744-—398747 (CNCI); OSUC 413941, 75606-75611, 76429, 77122-— Ti V/PLOSUGE), Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... | Trissolcus cultratus (Mayr), comb. rev. http://bioguid.osu.edu/osuc_concepts:13182 Figures 19, 50-52, 54; Morphbank° Telenomus cultratus Mayr, 1879: 699, 701, 703 (original description, keyed, syn- onymized by Kozlov (1968)); Kozlov 1968: 200 (junior synonym of Trissolcus flavipes (Thomson)). Aphanurus Cultratus (Mayr): Kieffer 1912: 70 (description, generic transfer). Microphanurus cultratus (Mayr): Kieffer 1926: 91, 95 (description, generic transfer, keyed); Nixon 1939: 130, 133 (description, keyed); Rjachovsky 1959: 83 (keyed). Asolcus cultratus (Mayr): Masner 1959: 378 (diagnosis, variation); Delucchi 1961: 44, 51 (description, keyed). Trissolcus cultratus (Mayr): Safavi 1968: 414 (keyed); Szabé6 1975: 266, 267 (description, lectotype designation, keyed). Diagnosis. Trissolcus cultratus is easily distinguished from other members of the flavi- pes group treated here by the parallel arched rugae on the frons between the anterior ocellus and the antennal scrobe. This species also lacks a well-developed orbital furrow near the malar sulcus, and by this character it may be separated from 7. brochymenae, LT. edessae, T: euschisti, and T. japonicus. Link to distribution map. [http://hol.osu.edu/map-large.html?id=13182] Associations. Emerged from egg of Carpocoris pudicus (Poda): [Hemiptera: Heter- optera: Pentatomoidea: Pentatomidae]; emerged from egg of Eurygaster Laporte: [He- miptera: Heteroptera: Pentatomoidea: Scutelleridae]; collected near eggs of Raphigaster nebulosa (Poda): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Raphigaster nebulosa (Poda): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; on leaf of maple: [Sapindales: Aceraceae]; collected near mulberry: [Urticales: Moraceae] Material examined. Lectotype, female: Other material: (122 females, 13 males, 4 sex unrecorded) AUSTRIA: 5 females, 2 sex unrecorded, USNMENT00979612, US- NMENT00979613 (CUIC); OSUC 75765-75767 (OSUC); USNMENT00675943, USNMENT00675944 (USNM). CHINA: 2 females, UCRC ENT 142635, 143817 (UCRC). CZECH REPUBLIC: 1 female, 3 males, USNMENT00896311, USN- MENT00896312, USNMENT00896313, USNMENT00896314 (CNCI). FRANCE: 4 females, OSUC 75753-75756 (OSUC). HUNGARY: 3 females, 1 sex unrecord- ed, OSUC 75771-75773, 75783 (OSUC). JAPAN: 32 females, 5 males, OSUC 144472-144480, 542363, 542374, 542412, 542415, USNMENT00896136, USN- MENT00896138, USNMENT00896140, USNMENT00896305, USNMENT- 00896307-—USNMENT00896309, USNMENT00896315, USNMENT00896339, USNMENT00896341 (CNCI); OSUC 75784, 75786-75788 (OSUC); UCRC ENT 297012 (UCRC); USNMENT00675730—USNMENT00675737, USNMENT00764849 (USNM). RUSSIA: 34 females, USNMENT00896048, USNMENT00896049, USNMENT00896050-USNMENT00896054, 72 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 50-51. °° 7rissolcus cultratus 50 female lectotype (NHMW 0008A), head, mesosoma, meta- soma, lateral view 51 female (USNMENT00764850), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters. USNMENT00896074, USNMENT00896075, USNMENT00979282- USNMENT00979286, USNMENT00979289 (CNCI); UCRC ENT 110944, 110951, 110963, 110983, 110985, 110992, 111001—111003, 111009, 111011, 111066, 111078, 133622, 297001—297003, 297009, 297013 (UCRC). SOUTH KOREA: 29 females, 3 males, OSUC 144470-144471, USNMENT00896011, USNMENT00896015, USNMENT00896016, USNMENT00896018, USNMENT00896019, USNMENT00896029, USNMENT00896032, USNMENT00896044 — USNMENT00896046, USNMENT00896112, USNMENT00896113 — USNMENT00896116, USNMENT00896118, USNMENT00896119, USNMENT00896121, USNMENT00896122, USNMENT00896134, Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... is vipes, female lectotype (NHRS-HEVA 000002617), head, anterolateral view 54 T° cultratus, female (US- NMENT00675734), head, anterior view 55 T° flavipes, female lectotype (NHRS-HEVA 000002617), head, anterior view. Scale bars in millimeters. USNMENT00896135, USNMENT00896157, USNMENT00979237, USNMENT- 00979246-USNMENT00979250, USNMENT00979253, USNMENT00979280 (CNCD. SWITZERLAND: 4 _ females, 1 male, USNMENT00979222-— USNMENT00979226 (CNCI). TAIWAN: 1 female, UCRC ENT 112210 (UCRC). UNITED KINGDOM: 1 female, USNMENT00916251 (BMNH). Comments. Kozlov (1968) designated a lectotype for 7. flavipes and simultane- ously treated 7: cultratus as a junior synonym. However, the concept of 7: flavipes presented in the key and description of his publication was that of 7. cultratus, and not of 7° flavipes, which in our assessment is a distinctly different species; the two may easily be separated by the presence of parallel arched rugae on the frons of T! cultra- tus, contrasting with absence of large rugae and presence of a circular impression on the frons of 7: flavipes (see Figs 52-55). The arched rugae on the frons of T° cultratus make the species particularly easy to identify, and the erroneous use of this character to identify T. flavipes was propagated throughout subsequent literature because Kozlov’s treatment was followed, and the primary type of T° cultratus was not re-examined. An unfortunate consequence of this error is that undoubtedly most, if not all, specimens of T. cultratus and T:. flavipes have been misidentified. 74 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Trissolcus edessae Fouts http://bioguid.osu.edu/osuc_concepts:3221 Figures 56-59; Morphbank® Trissolcus edessae Fouts, 1920: 65 (original description); Masner and Muesebeck 1968: 72 (type information); Johnson 1984: 799, 801 (description, keyed); Johnson 1987: 289, 300 (diagnosis, keyed). Diagnosis. Trissolcus edessae may be distinguished from the native species of Nearc- tic Trissolcus in the flavipes group (ZT! brochymenae, T: euschisti, and T. strabus) by the abruptly bicolored female antennae. It may be separated from 7! japonicus by the pres- ence of 2 clypeal setae and the episternal foveae that do not form a continuous line from the postacetabular sulcus to the mesopleural pit. It may be separated from 7 cultratus by the absence of parallel arched rugae on the frons. In 7! edessae a median mesoscutal carina is often present, and this is absent in 7! cultratus and T japonicus. Johnson (1984) used the absence of a mesopleural carina in T! edessae as a diagnos- tic character. Our examination included a specimen in which the mesopleural carina is present (Fig. 58) and thus we prefer not to use this character for identification. A result of this is that unambiguous identification of male specimens may require movement or removal of the wings to properly evaluate the surface sculpture within the axillar crescent. Link to distribution map. [http://hol.osu.edu/map-large.html?id=322 1] Associations. Emerged from egg of Acrosternum hilare (Say): [Hemiptera: Heter- optera: Pentatomoidea: Pentatomidae]; emerged from egg of Edessa bifida (Say): [He- miptera: Heteroptera: Pentatomoidea: Pentatomidae]; parasite of Edessa bifida (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Eus- chistus Dallas: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pachycoris torridus (Scopoli): [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae] Material examined. Holotype, female: UNITED STATES: LA, Orleans Par- ish, New Orleans, 23.VII.1919, C. E. Smith, USNMENT00872412 (deposited in USNM). Other material: (2 females, 1 male, 29 sex unrecorded) ELSALVADOR: 3 sex unrecorded, USNMENT00764980, USNMENT00764981, USNMENT00764993 (USNM). NICARAGUA: 2 sex unrecorded, OSUC 398762-—398763 (CNCI). UNITED STATES: 2 females, 1 male, 24 sex unrecorded, OSUC 17814 (BMNH); OSUC 398760-398761 (CNCI); OSUC 523872 (MEMU); OSUC 145559, 542444, 75617-75636 (OSUC); OSUC 145649 (USNM). Trissolcus erugatus Johnson http://bioguid.osu.edu/osuc_concepts:3228 Figures 27, 60-61; Morphbank’ Trissolcus erugatus Johnson, 1985b: 433, 436 (original description, keyed); Sarazin 1986: 980 (type information). Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... (= Figures 56-59. Trissolcus edessae 56 female holotype (USNMENT00872412) head, mesosoma, meta- soma, lateral view 57 female holotype (USNMENT00872412) head, mesosoma, dorsal view 58 (USN- MENT00764981), mesopleuron, metapleuron, lateral view 59 female holotype (USNMENT00872412), head, anterior view. Scale bars in millimeters. Diagnosis. Trissolcus erugatus may be distinguished from the most common South- western species of Trissolcus discussed here, 7) utahensis, by its strongly narrowed gena, angulate vertex, and the lack of rugulae on T2 (occasionally rugulae are present, but these are very short in comparison with those of 7’ utahensis). It may be distinguished from T’ hullensis by the following characters: metapostnotum invaginated near metas- cutellum and separating metanotum from propodeum, anterior extension of metapleu- ron short, not reaching mesocoxa, mandibular teeth shallowly incised; mesopleural 76 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 60-61. °° 7rissolcus erugatus female paratype (USNMENT00903009) 60 head, mesosoma, metasoma, lateral view 61 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters. carina absent; legs and Al—A6 usually yellow. Trissolcus cosmopeplae may usually be separated from 7! erugatus by the strong development of rugulae on T2 and the long anteroventral extension of the metapleuron toward the mesocoxa in the former species. Trissolcus erugatus seems to be a rather isolated species within the New World fauna of the genus. The narrowed gena allies it with 7: Aullensis, T. solocis, T. radix, and T° cosmopeplae, but the condition of the metapostnotum, mandibular teeth, and metapleural extension usually distinguish it quite clearly. Specimens from the South- west are easily identifiable, but variation in color and sculpture in the northern part of its range may result in confusion between this species and 7: cosmopeplae. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... Ta Link to distribution map. [http://hol.osu.edu/map-large.html?id=3228] Associations. collected on Larrea tridentata (Sessé & Moc. ex DC.) Coville: [Sap- indales: Zygophyllaceae]; emerged from egg of Thyanta custator (Fabricius): [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on alfalfa: [Fabales: Fa- baceae]; collected on lodgepole pine: [Pinales: Pinaceae] Material examined. Paratypes: (1 female, 1 male, 11 sex unrecorded) CANADA: 1 female, OSUC 17813 (BMNH). UNITED STATES: 1 male, 11 sex unrecorded, OSUC 398779-398780 (CNCI); OSUC 77860-77862 (MSWC); OSUC 145560, 75668-75672 (OSUC); USNMENT00903009 (USNM). Other material: (3 females, 2 males, 9 sex unrecorded) CANADA: 4 sex unrecorded, OSUC 398784—398787 (CNCI). UNITED STATES: 3 females, 2 males, 5 sex unrecorded, OSUC 39878 1-— 398783, 398788 (CNCI); OSUC 436700 (LACM); OSUC 413943, 523926- 525927,.523929; 75667. OSUG) Trissolcus euschisti (Ashmead) http://bioguid.osu.edu/osuc_concepts:3232 Figures 12, 20, 28, 42-43, 62-65; Morphbank* Telenomus euschristus Ashmead, 1888: ii (original description, spelling error). Trissolcus euschisti (Ashmead): Ashmead 1893: 161, 162 (emendation, description, ge- neric transfer, keyed); Harrington 1900: 183 (variation); Brues 1916: 549, 550 (description, keyed); Kieffer 1926: 127, 129 (description, keyed); Johnson 1984: 799, 801 (lectotype designation, synonymy, description, keyed); Johnson 1987: 289, 301 (diagnosis, keyed). Trissolcus podisi Ashmead, 1893: 161, 162 (original description, keyed, synonymized by Johnson (1984)); Brues 1916: 550 (description, keyed); Kieffer 1926: 127, 129 (description, keyed); Masner and Muesebeck 1968: 73 (lectotype designation); Johnson 1984: 801 (junior synonym of Trissolcus euschisti (Ashmead)). Trissolcus rufitarsis Kieffer, 1906: 262 (original description, synonymized by John- son (1984)); Kieffer 1926: 127, 128 (description, keyed); Hoebeke 1980: 27 (type information); Johnson 1984: 801, 803 (lectotype designation, junior syn- onym of Trissolcus euschisti (Ashmead)); Zuparko and Hamai 1994: 314 (type information). Trissolcus euchisti (Ashmead): Brues 1908: 11 (emendation); Golin, Loiacono, Margaria and Aquino 2011: 618 (host association). Diagnosis. Trissolcus euschisti may be distinguished from the similar 7’ brochymenae by the smooth or shallowly impressed sculpture on the ventral portion of the meso- pleuron anterior to the mesopleural carina. The smaller specimens of 7! euschisti are often quite distinct from the larger ones in the following characters: number of lateral setae on Tl, extent of fine wrinkles on T2, extent of rugae on S2, extent of S1 seta- tion, number of setae on the mesopleuron above the mesocoxa, sculpture of the upper 78 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) hg! My fe. A @ 5 Ee y | 77 7} —_x- “ Uf 4 NM a Figures 62-65. °” Trissolcus euschisti 62 female (USNMENT00872098), head, mesosoma, metasoma, lateral view. 63 female (USNMENT00872098), head, mesosoma, metasoma, dorsal view 64 female (OSUC 334007), head and mesosoma, dorsolateral view 65 female (USNMENT00872098), head, an- terior view. Scale bars in millimeters. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 79 portion of the frons, extent of transverse striae within the antennal scrobe, and the presence of a shallow groove below the anterior ocellus. The separation of 7! euschisti and T. brochymenae may be difficult with specimens that exhibit an intermediate state of faint rugosity on the anteroventral mesopleuron. These specimens are not common in our experience and the situation reflects the need for molecular data to further test the hypotheses of species delimitation presented here and in the revisions of Nearctic Trissolcus by Johnson (1984, 1985a,b). Link to distribution map. [http://hol.osu.edu/map-large.html?id=3232] Associations. Collected on Acer saccharum Marshall: [Sapindales: Aceraceae]; emerged from egg of Acrosternum hilare (Say): [Hemiptera: Heteroptera: Pentato- moidea: Pentatomidae]; on leaf of Catalpa Scop.: [Scrophulariales: Bignoniaceae]; emerged from egg of Edessa meditabunda (Fabricius): [Hemiptera: Heteroptera: Pen- tatomoidea: Pentatomidae]; emerged from egg of / host egg of Euschistus Dallas: [He- miptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Euschistus conspersus Uhler: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: |Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of / host egg of Pentatomidae: |Hemiptera: Heteroptera: Pentato- moidea: Pentatomidae]; unspecified association Pentatomidae: [Hemiptera: Heterop- tera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from sentinel egg mass of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pen- tatomidae]; parasite of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pen- tatomoidea: Pentatomidae]; unspecified association Podisus maculiventris (Say): [He- miptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus serieventris Uhler: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus spinosus (Dallas): [Hemiptera: Heteroptera: Pentatomoidea: Pen- tatomidae]; collected on Quercus agrifolia Nee.: [Fagales: Fagaceae]; emerged from sentinel egg mass of Thyanta accerra custator (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on alder: [Fagales: Betulaceae]; on young maple: [Sapindales: Aceraceae]; collected on pecan: [Juglandales: Juglandaceae]; col- lected on red pine: [Pinales: Pinaceae]; collected on wax myrtle: [Myricales: Myri- caceae] Material examined. Lectotype, female, T° euschristus: UNITED STATES: Riley Co., VI, Marlatt, OSUC 256926 (deposited in OSUC). Lectotype, female, T° rufitar- sis: UNITED STATES: Ormsby Co., VII, Baker, Cornell U. No. 388.1 (deposited in CUIC). Lectotype, female, 7! rufitarsis: UNITED STATES: Ormsby Co., VII, Baker, USNMENT00979614 (deposited in CUIC). Lectotype, female, T° podisi: UNITED STATES: PA, Philadelphia Co., Philadelphia, no date, Cresson, USNMENT00989033 (deposited in USNM). Paratypes:s UNITED STATES: 2 sex unrecorded, OSUC 398789, 398823 (CNCI). Other material: (71 females, 10 males, 243 sex unrecorded) BRAZIL: 3 sex unrecorded, OSUC 75746-75748 (OSUC). CANADA: 1 female, 1 male, 32 sex unrecorded, OSUC 145426-145427, 398793—398798, 398800—398803, 398805-—398817, 398836—398840, 398844—398847 (CNCI). FRENCH GUIANA: 80 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) 1 sex unrecorded, OSUC 248138 (OSUC). MEXICO: 1 sex unrecorded, OSUC 75745 (OSUC). NORTH AMERICA: 1 female, OSUC 398799 (CNCI). UNITED STATES: 69 females, 9 males, 201 sex unrecorded, OSUC 17808 (BMNH); OSUC 145178, 145409-145410, 398790-398792, 398818-398822, 398824-398835, 398841-398843 (CNCI); USNMENT00979600, USNMENT00979603, USN- MENT00979605 (CUIC); OSUC 436702 (LACM); OSUC 523870—523871, 523874 (MEMU); OSUC 145411-145418, 145421-145425 (MSWC); IRREC 1794, IRREC834, OSUC 143837, OSUC 143838—OSUC 143850, OSUC 145177, OSUC 145561, OSUC 157488—-OSUC 157493, OSUC 181546, OSUC 248134, OSUC 248139, OSUC 334007, OSUC 402728, OSUC 404912, OSUC 409995, OSUC 413680, OSUC 413681, OSUC 413682—OSUC 413699, OSUC 413729- OSUC 413748, OSUC 413940, OSUC 523862, OSUC 523863, OSUC 523866— OSUC 523868, OSUC 523883-OSUC 523903, OSUC 523924, OSUC 523928, OSUC 523934, OSUC 523939, OSUC 523941, OSUC 542439, OSUC 542441, OSUC 542443, OSUC 70463, OSUC 75678—OSUC 77202, OSUC 79805 (OSUC); OSUC 145419-145420 (UCRC); BMSB 1218, 1220-1230, 1232, OSUC 523851, USNMENT00872096—USNMENT00872102, © USNMENT00989171, © USN- MENT00989172, USNMENT00989174-USNMENT00989179 (USNM). Trissolcus hullensis (Harrington) http://bioguid.osu.edu/osuc_concepts:3244 Figures 22, 24, 66-67; Morphbank” Telenomus hullensis Harrington, 1900: 182 (original description); Kieffer 1926: 27, 40 (description, keyed). Trissolcus hullensis (Harrington): Johnson 1984: 10 (generic transfer); Johnson 1985b: 433, 438 (description, keyed); Sarazin 1986: 981 (type information). Diagnosis. Trissolcus hullensis is most closely related to 7! solocis, T’ radix and T: zakotos. Trissolcus hullensis may be distinguished from these by the anteriorly invaginated metapostnotum. Additional characters useful for identification are: the paracoxal sulcus in the ventral half of the metapleuron, absent in 7’ /ullensis, present in T! radix and T; zakotos; sculpture of the mesoscutellum, coriaceous or smooth in southern specimens of 7 hullensis, coarsely areolate in T. solocis and T. radix; the color of the radicle, black in 7. hullensis, T: zakotos and T. solocis, yellow in T: radix; and the rounded vertex, sharply angled in 7’ radix and T’ solocis. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3244] Associations. Emerged from Euschistus servus (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: [Hemiptera: Het- eroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Recur- varia milleri Busck: [Lepidoptera: Glossata: Gelechioidea: Gelechiidae]. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 81 Figures 66-67. °° Trissolcus hullensis 66 female (OSUC 523881), head, mesosoma, metasoma, dorsal view 67 female (USNMENT00903008), head, mesosoma, metasoma, lateral view. Scale bars in millimeters. Material examined. Non-type:. UNITED STATES: 1 female, OSUC 266782 (OSUC). Other material: (17 females, 1 male, 64 sex unrecorded) CANADA: 1 female, 5 sex unrecorded, OSUC 17815 (BMNH); OSUC 145179, 145392-145393, 398853 (CNCI); OSUC 75837 (OSUC). MEXICO: 11 sex unrecorded, OSUC 398854 (CNCI); OSUC 77870-77877 (MSWC); OSUC 75838-75839 (OSUC). UNITED STATES: 16 females, 1 male, 48 sex unrecorded, OSUC 145391, 398855—398857, 542438 (CNCI); OSUC 523873, 523875—-523882 (MEMU); OSUC 77865-77869 (MSWC); OSUC 142487-142491, 143851, 145369-145373, 145389, 145562, 523856, 523946, 542456, 62453, 70529, 75826-75836, 76427-76428 (OSUC); OSUC 145374-145378, 145380—-145388, 145390 (UCRC). 82 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Trissolcus japonicus (Ashmead) http://bioguid.osu.edu/osuc_concepts:3249 Figures 17, 21, 25, 68-71; Morphbank"® Dissolcus japonicus Ashmead, 1904: 73 (original description); Kieffer 1926: 124, 125 (description, keyed). Trissolcus japonicus (Ashmead): Masner and Muesebeck 1968: 72 (type information, generic transfer); Hirashima and Yamagishi 1981: 153 (description, synonymy); Ryu and Hirashima 1984: 37, 43 (description, keyed); Talamas, Buffington and Hoelmer 2013: 114 (description, synonymy, type information). Trissolcus halyomorphae Yang: Qiu, Yang and Tao 2007: 62 (unavailable: nomen nu- dum); Yang, Yao, Qiu and Li 2009: 40 (original description); Talamas, Buffington and Hoelmer 2013: 114 (junior synonym of Trissolcus japonicus (Ashmead)). Diagnosis. As previous authors have stated (Yang et al. 2009), 7’ japonicus belongs to the flavipes species group, first recognized by Kozlov and Lé (1976) and refined by Johnson (1984). Trissolcus japonicus may be separated from other species of the flavipes group Jrissolcus in the Nearctic by the following characters: orbital furrow expanded near intersection with malar sulcus; postacetabular and mesopleural epicoxal sulci formed by lines of closed cells (Fig. 70); episternal foveae extending from dorsal apex of postacetabular carina to mesopleural pit (Fig. 68); 4 clypeal setae (Fig. 25). Link to distribution map. [http://hol.osu.edu/map-large.html?id=3249] Associations. Emerged from egg of Halyomorpha halys (Stal): [Hemiptera: Het- eroptera: Pentatomoidea: Pentatomidae]; emerged from Plautia stali Scott: [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on mulberry: [Urticales: Moraceae]; emerged from stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Pen- tatomidae]; emerged from egg of stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae] Material examined. Holotype, female, D. japonicus: JAPAN: Kanagawa Pref., Ashi- garashimo Dist., Hakone Town, no date, Koebele, USNMENT00831865 (deposited in USNM). Paratypes: CHINA: 2 females, USNMENT00872401, USNMENT00872402 (USNM). Other material: (44 females, 16 males, 12 sex unrecorded) CHINA: 32 fe- males, 11 males, 1 sex unrecorded, USNMENT00979190, USNMENT00979191, USNMENT00979192-USNMENT00979198, = USNMENT00979200, — US- NMENT00979201-USNMENT00979221 = (CNCI); © USNMENT00675704, USNMENT00675738, | USNMENT00675739, © USNMENT00675743, —US- NMENT00675747, © USNMENT00675925, © USNMENT00764940, USN- MENT00764941, © USNMENT00764944, § USNMENT00764948, = USN- MENT00764949, USNMENT00764984, USNMENT00916255 (USNM). JA- PAN: 8 females, 4 males, 10 sex unrecorded, OSUC 144481-144482, 398858, USNMENT00896340 (CNCI); OSUC 145632, 75843-75848 (OSUC); US- Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 83 Figures 68-71. °° Trissolcus japonicus 68 female (USNMENT00872402), head, mesosoma, metasoma, lateral view 69 female (USNMENT00675989), head, mesosoma, metasoma, dorsal view 70 female (USN- MENT00675989), head, mesosoma, ventral view 71 female (USNMENT00872402), head, anterolateral view. Scale bars in millimeters. NMENT00675755, USNMENT00675770, USNMENT00872125—-USN- MENT00872133 (USNM). RUSSIA: 1 female, USNMENT00979287 (CNCI). SOUTH KOREA: 3 females, 1 male, USNMENT00979251, USNMENT00979254 (CNCI); USNMENT00675705, USNMENT00675708 (USNM). 84 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Trissolcus occiduus Johnson http://bioguid.osu.edu/osuc_concepts:3275 Figures 18, 34, 72-75; Morphbank"! Trissolcus occiduus Johnson, 1985a: 109, 111 (original description, keyed). Diagnosis. This species may be distinguished from other species in the thyantae group by the expanded gena. It may also be separated from 7’ thyantae by the complete meso- pleural carina, and from 7 parma and T. ruidus by the entirely smooth mesoscutellum and absence of a genal carina. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3275] Associations. Collected on Abronia maritima Nutt. ex S.Watson: [Caryophyllales: Nyctaginaceae]; emerged from egg of Chlorochroa norlandorum Buxton & Thomas: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Chlo- rochroa sayi (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatoma sayii (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae] Material examined. Holotype, female: UNITED STATES: CA, Ventura Co., area 2, Point Mugu Naval Air Station, 31.VU—24.VIII.1981, C. D. Nagano & J. N. Hogue, CNCI 0004 (deposited in CNCI). Paratypes: UNITED STATES: 4 fe- males, 5 sex unrecorded, OSUC 17811 (BMNH); OSUC 145180 (CNCI); OSUC 143814 (LACM); OSUC 77864 (MSWC); OSUC 145564, 76216-76217 (OSUC); USNMENT00764995, USNMENT00877675 (USNM). Other material. UNITED STATES: 3 females, 1 male, 1 sex unrecorded, OSUC 145365, 76430 (OSUC); US- NMENT00954754, USNMENT00979294, USNMENT00979295 (USNM). Trissolcus parma Johnson http://bioguid.osu.edu/osuc_concepts:3284 Figures 76-79; Morphbank”* Trissolcus parma Johnson, 1985a: 110, 111 (original description, keyed); Sarazin 1986: 981 (type information). Diagnosis. Trissolcus parma may be distinguished from 7’ ruidus by the lack of rugulae outside of the antennal scrobe and the lack of longitudinal elements in the sculpture of the posterior portion of the mesoscutum. It may be separated from the other members of the thyantae group by the presence of microsculpture on the mesoscutellum. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3284] Associations. Collected on Medicago sativa L.: [Fabales: Fabaceae]; collected un- der Vaccinium uliginosum L..: [Ericales: Ericaceae] Material examined. Holotype, female: CANADA: AB, Scandia, 2.VII.1956, sweeping, O. Peck, CNC No. 18339 (deposited in CNCI). Paratypes: (1 female, 1 male, 1 sex unrecorded)CANADA: 1 female, 1 sex unrecorded, OSUC 17809 (BMNH); Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 85 ai Figures 72-75. * Trissolcus occiduus 712. female (USNMENT00877675), head, mesosoma, metasoma, dorsal view 73 female (USNMENT00764995), head, mesosoma, metasoma, lateral view 74 female (OSUC 76126), head and mesosoma, lateral view 75 female (USNMENT00764995), head, anterior view. Scale bars in millimeters. OSUC 145565 (OSUC). UNITED STATES: 1 male, USNMENT00764990 (USNM). Other material: (2 females) CANADA: 1 female, OSUC 76264 (OSUC). UNITED STATES: 1 female, OSUC 62481 (OSUC). 86 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) A , i a A % — \ t / i>, Figures 76-79. “' Trissolcus parma 16 female (OSUC 76432), head, mesosoma, metasoma, lateral view 77 female (OSUC 62481), head, mesosoma, metasoma, dorsal view 78 female (USNMENT00765990), head and mesosoma, ventral view 79 female (USNMENT00765990), head, anterior view. Scale bars in millimeters. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 87 0.2 ~ 80 Figure 80.” Trissolcus radix female (USNMENT00764955), head, mesosoma, metasoma, lateral view. Scale bar in millimeters. Trissolcus radix Johnson http://bioguid.osu.edu/osuc_concepts:3295 Figures 80-83; Morphbank’” Trissolcus radix Johnson, 1985b: 432, 440 (original description, keyed). Diagnosis. Trissolcus radix is most closely related to 1 hullensis, T; solocis, and T’ zako- tos, from which it may be distinguished by the bright yellow radicle. The well defined paracoxal sulcus in the ventral half of the metapleuron serves to separate this species from T. hullensis and T: solocis, and the rugose sculpture of the mesoscutellum will separate it from 77 ullensis and T. zakotos. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3295] Associations. Emerged from egg of Euthyrhynchus floridanus (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on coffee: [Rubiales: Rubiaceae] Material examined. Paratypes: (1 female, 4 sex unrecorded) COSTA RICA: 1 sex unrecorded, OSUC 76272 (OSUC). GUATEMALA: | sex unrecorded, USN- MENT00764955 (USNM). UNITED STATES: 1 female, 2 sex unrecorded, OSUC 145567, 76270-76271 (OSUC). Other material: MEXICO: 2 sex unrecorded, USN- MENT00896395, USNMENT00896396 (UANL). 88 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 81-83. Trissolcus radix 8\ female paratype (USNMENT00764955), head, mesosoma, metas- oma, dorsal view 82 female (USNMENT00764955), head and mesosoma, lateral view 83 female (OSUC 76271), head, anterolateral view. Scale bars in millimeters. Trissolcus ruidus Johnson http://bioguid.osu.edu/osuc_concepts:3299 Figures 33, 84-87; Morphbank"* Trissolcus ruidus Johnson, 1985a: 111 (original description, keyed); Sarazin 1986: 981 (type information). Diagnosis. Trissolcus ruidus may be separated from T’ parma by the presence of rugae on the lateral frons (Fig. 86) and longitudinal elements that are often present in the sculpture of the mesoscutum between the notauli. Like 7’ parma, it may be separated Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 89 : fs 7 ™ r ee ot, | i i ie E aie ia E nits Figures 84-87. “ Trissolcus ruidus female paratype (OSUC 145568) 84 head, mesosoma, metasoma, ventrolateral view 85 head, mesosoma, metasoma, dorsal view 86 head mesosoma, anterolateral view 87 head, anterior view. Scale bars in millimeters. from the other members of the thyantae group by the presence of microsculpture on the mesoscutellum. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3299] Material examined. Holotype, female: UNITED STATES: AZ, Cochise Co., Portal, Southwestern Research Station (SWRS), 19.X.1978, Masner & Gibson, CNC 90 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) No. 18341 (deposited in CNCI). Paratype. UNITED STATES: 1 sex unrecorded, OSUC 145568 (OSUC). Other material. UNITED STATES: 2 females, 1 male, OSUC 76431-76432 (OSUC); OSUC 144847 (USNM). Trissolcus solocis Johnson http://bioguid.osu.edu/osuc_concepts:331 1 Figures 88—91; Morphbank” Trissolcus solocis Johnson, 1985b: 433, 441 (original description, keyed). Diagnosis. Trissolcus solocis may be distinguished from the closely related 7’ hullensis and 7° zakotos by the coarse sculpture of the mesoscutellum. From 7) radix it may be most easily separated by its black radicle and the absence of a well-defined paracoxal sulcus in the ventral half of the metapleuron. Link to distribution map. [http://hol.osu.edu/map-large.html?id=33 1 1] Associations. Emerged from egg of Acrosternum marginatum (Palisot): [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Alcaeor- rhynchus grandis (Dallas): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Hemiptera: [Hemiptera] Material examined. Paratypes: (1 female, 1 male, 9 sex unrecorded) MEXICO: 4 sex unrecorded, UINMENT00764956, USNMENT00764957, USNMENT00764958, USNMENT00764959 (USNM). UNITED STATES: 1 female, 1 male, 5 sex unre- corded, OSUC 398866 (CNCI); OSUC 145569, 76309-76313 (OSUC). Trissolcus strabus Johnson http://bioguid.osu.edu/osuc_concepts:3313 Figures 8, 23, 26, 29, 32, 92-93; Morphbank’® Trissolcus strabus Johnson, 1984: 798, 806 (original description, keyed); Sarazin 1986: 981 (type information); Johnson 1987: 286, 296 (diagnosis, keyed). Diagnosis. Trissolcus strabus may be distinguished from species of the flavipes group in the Nearctic by the ventral constriction of the orbital furrow and the relatively coarse sculpture of the mesoscutellum. Most specimens have setae present on the first later- otergite, a character found among some flavipes group species of the Neotropics, but not elsewhere in the Nearctic. The rugose mesoscutellum can be used as a diagnostic character in most cases, but the degree of rugosity is variable. In some specimens the mesoscutellum is almost completely smooth with faint hints of rugae along the ante- rior margin. In others, the rugosity is confined to the lateral portions of the sclerite. In the latter case, rugose sculpture exists where there is setation, and in specimens Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 91 Figures 88-91. © T7rissolcus solocis female paratype (OSUC 76312) 88 head, mesosoma, metasoma, lateral view 89 head, mesosoma, metasoma, dorsal view 90 head and mesosoma, dorsolateral view 91 head, ante- rolateral view. Scale bars in millimeters. 92 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 92-93. “° Trissolcus strabus, female (BMSB 1203) 92 head, mesosoma, metasoma, lateral view 93 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters. with an entirely rugose mesoscutellum, the entire surface is setose. This leads us to hypothesize that, at least on the mesoscutellum of 7’ strabus, the rugose sculpture and setation are linked. The specimens with reduced macrosculpture on the mesoscutel- lum also have reduced sculpture on the lateral mesoscutum (lateral of the notaulus), revealing coriaceous microsculpture. Link to distribution map. [http://hol.osu.edu/map-large.html?id=33 13] Associations. Emerged from egg of / host egg of Brochymena Amyot & Serville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; on leaf of apple: [Rosales: Rosaceae] Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 93 Material examined. Holotype, female: CANADA: ON, Hamilton, 31.VII.1980, malaise trap, M. Sanborne, CNC No. 18342 (deposited in CNCI). Paratypes: (1 female, 3 sex unrecorded) CANADA: 1 sex unrecorded, OSUC 145570 (OSUC). UNITED STATES: 1 female, 2 sex unrecorded, OSUC 17810 (BMNH); OSUC 76314 (OSUC); USNMENT00764998 (USNM). Non-type: UNITED STATES: 1 female, OSUC 248187 (OSUC). Other material: (14 females, 1 male, 13 sex unrecord- ed) UNITED STATES: 14 females, 1 male, 12 sex unrecorded, IRREC 1469-1470, 1472, 1521, 1587, 1595, 1787, 1789, 1797, IRREC1582, IRREC1584 (OSUC); BMSB 1202-1215, OSUC 145650, OSUC 523850 (USNM). Trissolcus thyantae Ashmead http://bioguid.osu.edu/osuc_concepts:3321 Figures 94-98; Morphbank"” Trissolcus thyantae Ashmead, 1893: 162, 163 (original description, keyed); Brues 1916: 550 (description, keyed); Kieffer 1926: 127, 128 (description, keyed); Masner and Muesebeck 1968: 74 (lectotype designation); Johnson 1985a: 108, 111 (descrip- tion, keyed). Diagnosis. Trissolcus thyantae is most similar to T’ occiduus and T. valkyria. It may be separated from J’ occiduus by the narrow malar region and from both by the lack of a complete mesopleural carina. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3321] Associations. Emerged from egg of Euschistus Dallas: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Euschistus variolarius (Palisot de Beauvois): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from ege of Panthea furcilla (Packard): [Lepidoptera: Glossata: Noctuoidea: Noctuidae]; emerged from egg of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pen- tatomidae]; emerged from egg of Pinaceae: [Pinales: Pinaceae]; emerged from egg of Thyanta custator (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentato- midae]; collected on pine: [Pinales: Pinaceae]; emerged from egg of pine: [Pinales: Pinaceae]; collected on soybean: [Fabales: Fabaceae] Material examined. Lectotype, female: UNITED STATES: AL, Dallas Co., Selma, [X-1880, reared from egg, E. A. Schwarz, USNMENT00989048 (deposited in USNM). Non-type: UNITED STATES: 1 female, OSUC 266773 (OSUC). Orh- er material: (2 females, 27 sex unrecorded) CANADA: 1 female, 6 sex unrecorded, OSUC 17812 (BMNH); OSUC 145196, 145368, 398870—398871 (CNCI); OSUC 145572, 76328 (OSUC). UNITED STATES: 1 female, 21 sex unrecorded, OSUC 157505-157506, 157512-157520, 76320-76327 (OSUC); USNMENT00764991, USNMENT00764994, USNMENT00979296 (USNM). 94 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) metasoma, dorsal view 95 female holotype (USNMENT00989048), head, mesosoma, lateral view 96 fe- male (OSUC 76325), head and mesosoma, dorsolateral view 97 female (USNMENT00764991), meso- soma, ventrolateral view 98 female holotype (USNMENT00989048), head, anterolateral view. Scale bars in millimeters. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 95 Trissolcus utahensis (Ashmead) http://bioguid.osu.edu/osuc_concepts:3327 Figures 99-103; Morphbank’® Telenomus utahensis Ashmead, 1893: 143, 145, 148 (original description, keyed). Hadronotus mesillae Cockerell, 1897: 25 (original description, synonymized by Muese- beck & Walkley (1951)); Brues 1910: 47 (keyed); Kieffer 1926: 454, 464 (descrip- tion, keyed); Muesebeck and Walkley 1951: 694 (junior synonym of Télenomus utahensis Ashmead). Telenomus ashmeadi Morrill, 1907: 419 (original description, synonymized with Te/- enomus mesillae (Cockerell) by Gahan (1932)); Kieffer 1926: 27, 48 (description, keyed); Gahan 1932: 757 (junior synonym of Telenomus mesillae (Cockerell)); Mani 1936: 335 (description of misidentified Indian specimen). Liophanurus utahensis (Ashmead): Kieffer 1926: 65, 73 (description, generic transfer, keyed). Telenomus mesillae (Cockerell): Gahan 1932: 757 (generic transfer, synonymy). Trissolcus utahensis (Ashmead): Krombein and Burks 1967: 297 (generic transfer); Masner and Muesebeck 1968: 74 (type information); Johnson 1985b: 432, 441 (description, keyed). Trissolcus ashmeadi (Morrill): Masner and Muesebeck 1968: 71 (lectotype designation). Trissolcus mesillae (Cockerell): Masner and Muesebeck 1968: 73 (type information). Diagnosis. Trissolcus utahensis is a relatively dark-colored species, though some spec- imens from the southern part of its range have lighter-colored appendages. In the Nearctic region it is most similar to 7’ basalis. The two may be distinguished by the color of Al, usually dark, concolorous with the radicle in 7 utahensis, and yellow, sharply contrasting with the dark radicle in 7’ basalis; and the mesoscutellar sculpture, smooth in 7) utahensis, coriaceous in 7, basalis. Link to distribution map. [http://hol.osu.edu/map-large.html?id=3327] Associations. Emerged from Chlorochroa sayi (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Chlorochroa sayi (Stal): [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Chlorochroa uhleri (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Fremon- tia TYorr.: [Malvales: Malvaceae]; emerged from egg of Pentatoma ligata Say: [Hemip- tera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatoma sayii (Stal): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; egg parasite of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pen- tatomidae: |Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; unspecified asso- ciation Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; col- lected on Pinus ponderosa P. & C. Lawson: [Pinales: Pinaceae]; collected on Prosopis L.: [Fabales: Fabaceae]; living with Rhyacionia neomexicana (Dyar): [Lepidoptera: Glossata: Tortricoidea: Tortricidae]; collected on Russian thistle: [Caryophyllales: Chenopodi- 96 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Figures 99-103. Trissolcus utahensis 99 female (USNMENT00872111), head, mesosoma, metasoma, lateral view 100 female lectotype (USNMENT00), head and mesosoma, dorsal view 101 lectotype fe- male (USNMENT00989049), metasoma, dorsal view 102 female lectotype (USNMENT00989049), head and mesosoma, lateral view 103 female lectotype(USNMENT00989049), head and mesosoma, anterolateral view. Scale bars in millimeters. aceae]; emerged from Thyanta pallidovirens (Stal): [Hemiptera: Heteroptera: Pentato- moidea: Pentatomidae]; collected on tomato: [Solanales: Solanaceae]; on leaf of tomato: [Solanales: Solanaceae]; collected on wild carrot: [Apiales: Apiaceae]. Material examined. Lectotype, female, 7. utahensis: UNITED STATES: Wasatch Range, 27.V1.1891, E. A. Schwarz, USNMENT00989049 (deposited in USNM). Paralectotype. UNITED STATES: 1 male, USNMENT00764992 (USNM). Lecto- Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 97 type, female, 7° ashmeadi: UNITED STATES: TX, Ward Co., Barstow, 12.1X.1905, reared from egg, A. W. Morrill, USNM Type No. 10364 (deposited in USNM). Holotype, female, H. mesillae. UNITED STATES: NM, Dofa Ana Co., Las Cruces, no date, reared from egg, T. D. A. Cockerell, USNM Type No. 3696 (deposited in USNM). Other material: (10 females, 3 males, 142 sex unrecorded) CANADA: 5 sex unrecorded, OSUC 145192—145193, 398862 (CNCI); OSUC 76416-76417 (OSUC). UNITED STATES: 10 females, 3 males, 136 sex unrecorded, OSUC 17807 (BMNH); OSUC 143819-143823, 436690-436699 (LACM); OSUC 77878-77930 (MSWC); OSUC 145230-145252, 145635, 405748, 413942, 542448542449, 542451—-542452, 542455, 76383-76415, 77203-77212 (OSUC); OSUC 205760 (UCDC); USNMENT00872110—USNMENT00872114 (USNM). Trissolcus valkyria Johnson & Talamas, sp. n. http://zoobank.org/4DB6B48B-9DA2-47E0-98A5-A28 103C42C15 http://bioguid.osu.edu/osuc_concepts:344497 Figures 104—108; Morphbank” Description. Female body length: 0.97—1.11 mm (n=6). Color of radicle: yellow; brown; pale brown. Number of mandibular teeth: three. Number of clypeal setae: 6. Facial striae: absent. Shape of gena in lateral view: receding posteriorly. Genal carina: extending dorsally from base of mandible. Macrosculpture of frons outside of antennal scrobe: absent; irregularly rugose. Orbital furrow: narrow to absent near malar sulcus. Hyperoccipital carina: absent. Preocellar pit: present. Epomial carina: present. Netrion sulcus: complete. Mesoscutal suprahumeral sulcus: indicated by cells. Mesoscutal humeral sulcus: indicated by cells. Pattern of mesoscutal microsculpture: antero-posteriorly uniform. Macrosculpture of mesos- cutum: coriacious. Area bounded by axillar crescent: smooth. Parapsidal line: ab- sent. Notaulus: present. Median mesoscutal carina: absent; present. Median mesos- cutal sulcus: absent. Sculpture of mesoscutellum: smooth. Postacetabular sulcus: comprised of cells. Shape of episternal foveae: round; antero-posteriorly elongate. Number of episternal foveae: 3-5. Course of episternal foveae ventrally: abutting cells of postacetabular sulcus. Course of episternal foveae dorsally: extending dorsally to mesopleural pit. Sculpture of anterior mesepisternum: smooth or with shallowly impressed micros- culpture. Mesopleural epicoxal sulcus: comprised of cells. Mesopleural carina: com- plete. Speculum: transversely striate. Paracoxal sulcus in ventral half of metapleuron: absent or indistinguishable from sculpture. Anteroventral extension of metapleuron: short, not reaching mesocoxa. Line of pits along metapleural carina: present. Se- tation of metapleuron: present. Metapostnotum: invaginated near edge of metas- cutellum and separating metanoum from propodeum. Color of legs beyond coxae: yellow; femora brown, otherwise variably yellow to brown. Metasomal depression: punctate or crenulate dorsally. 98 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) s t Figures 104-108. ” 77issolcus valkyria, female holotype (OSUC 542457) 104 head, mesosoma, meta- soma, dorsal view 105 lateral habitus 106 head and mesosoma, lateral view 107 head and mesosoma, ventrolateral view 108 head, anterolateral view. Scale bars in millimeters. Sublateral setae on T1: absent. Setation of laterotergite 1: absent. Sculpture of T2 posterior to antecostal sulcus: smooth or with very faintly impressed striation; dis- tinctly striate posterior to basal costae. Diagnosis. 77issolcus valkyria is most similar to T: thyantae with which it shares a mesoscutellum without microsculpture and a narrow gena. T7issolcus valkyria may be separated T° thyantae and T: occiduus by the presence of a complete and well defined mesopleural carina. From 7: occiduus it may also be separated by the narrow gena. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 99 Etymology. The epithet “valkyria” is Old Norse for “chooser of the slain” and refers to the female figures in Norse mythology that selected which soldiers would die in battle. The name is to be treated as a noun in apposition. Link to distribution map. [http://hol.osu.edu/map-large.html?id=344497] Material examined. Holotype, female: UNITED STATES: WI, Juneau Co., North Rynearson site, Necedah National Wildlife Refuge, 21.VI-11.VII.1996, flight intercept trap, R. H. Williams, OSUC 542457 (deposited in OSUC). Paratypes: UNITED STATES: 5 females, OSUC 405747, 76433 (OSUC); OSUC 144848— 144849, 145646 (USNM). Comments. 77issolcus valkyria, was previously recognized by Johnson but re- mained undescribed due to a dearth of specimens. A small number of additional speci- mens are now known, providing in our opinion a sufficient basis for the description of this species. Trissolcus zakotos Talamas, s http://zoobank.org/4E138794- 6517. 42FE-9AC7-B7D92CDB04B6 http://bioguid.osu.edu/osuc_concepts:345034 Figures 109-112; Morphbank”’ Description. Female body length: 1.28—1.41 mm (n=20). Male body length: 1.18 mm (n=1). Color of radicle: brown. Number of mandibular teeth: three. Number of clypeal setae: 6. Facial striae: present as 3 or more rugulae extending onto lateral frons. Shape of gena in lateral view: receding posteriorly. Genal carina: extending dorsally from base of mandible. Macrosculpture of frons outside of antennal scrobe: irregularly rugose. Orbital furrow: narrow to absent near malar sulcus. Hyperoccipital carina: absent. Preocellar pit: present. Epomial carina: present. Netrion sulcus: complete. Mesoscutal suprahumeral sulcus: indicated by cells. Mesoscutal humeral sulcus: indicated by cells. Pattern of mesoscutal microsculpture: antero-posteriorly uniform. Macrosculpture of mesoscutum: reticulate anteriorly, longitudinally rugulose posteriorly. Area bounded by axillar crescent: smooth. Parapsidal line: absent. Notaulus: absent. Median mesoscutal carina: absent. Median mesoscutal sulcus: absent. Sculpture of mesoscutellum: coriaceous. Postacetabular sul- cus: comprised of cells. Shape of episternal foveae: irregular; round. Number of episternal foveae: 1-2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separated from mesopleural pit. Sculpture of anterior mesepisternum: faintly rugulose; finely reticulate. Mesopleural epi- coxal sulcus: present as a smooth furrow; comprised of cells. Mesopleural carina: com- plete; well defined in anterior half, posterior half poorly defined to absent. Speculum: transversely striate. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of distinct foveae. Anteroventral extension of metapleuron: long, extending to mesocoxa. Line of pits along metapleural carina: present. Setation of metapleuron: absent. Meta- postnotum: invaginated near edge of metascutellum and separating metanoum from 100 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) : = “ee 7 112 Figures 109-112. °° 77rissolcus zakotos 109 female paratype (USNMENT00954596), head, mesosoma, metasoma, lateral view 110 female holotype (USNMENT00903008), head, mesosoma, metasoma, dor- sal view 111 female holotype (USNMENT00903008), mesosoma, dorsolateral view 112 female paratype (USNMENT 00954600), head, anterolateral view. Scale bars in millimeters. *° propodeum. Color of legs beyond coxae: femora and tibiae brown, otherwise variably yellow to brown. Metasomal depression: punctate or crenulate dorsally. Sublateral setae on T1: absent; present. Setation of laterotergite 1: absent. Sculpture of T2 posterior to antecostal sulcus: smooth or with very faintly impressed striation. Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 101 Diagnosis. Trissolcus zakotos is closest to T. radix, with which it shares a well de- fined paracoxal sulcus. The two may be separated by the presence of of bright yellow radicle and coarse sculpture of the mesoscutellum in T: radix. In T: zakotos the radicle is brown and the mesoscutellum is covered by microsculpture, but without additional rugae. Additionally, 7. zakotos has numerous (3-5) rugae radiating from the lateral edge of the clypeus. This character is present is both 7: radix and T. solocis but is less pronounced and the number of rugae is smaller (1—2). Etymology. The epithet “zakotos” is Greek for “angry” and is applied to this spe- cies because of the appearance of its frons. The name is treated as an appositional noun. Link to distribution map. [http://hol.osu.edu/map-large.html?id=345034] Associations. Emerged from Apateticus bracteatus (Fitch): [Hemiptera: Heterop- tera: Pentatomoidea: Pentatomidae] Material examined. Holotype, female: UNITED STATES: MT, Ravalli Co., Hamilton, V-1972, W. L. Jellison, USNMENT00903008 (deposited in USNM). Paratype: UNITED STATES: 22 females, 1 male, USNMENT00954588—USN- MENT00954589 (CNCI); USNMENT00954586—USNMENT00954587 (OSUC); USNMENT00903005, USNMENT00903006, USNMENT00954590—USN- MENT00954606 (USNM). Acknowledgements We are grateful to: Luciana Musetti (OSUC), Sara Hemly (OSUC), Manuela Vizek (NHMW), Hege Vardal (NHRS), Lubomir Masner (CNCI), Andy Bennett (CNCI), Tim Haye (CABI), Kim Hoelmer (BIIRU) and Christine Dieckhoff (BIIRU) for loans and specimens deposited in USNM; David Notton (BMNH) for specimen loans and commentary on nomenclature, Joe Cora (OSUC) for critical database support and making taxonomic literature available; Istvan Miko for commentary on morphological characters; and to Ian Realo and Samantha Fitzsimmons-Schoenberger for photog- raphy and transcribing label data. This work was made possible by funding from the Systematic Entomology Lab, USDA-ARS, and the Beneficial Insect Introduction Re- search Laboratory. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recom- mendation or endorsement by the USDA; USDA is an equal opportunity provider and employer. References Ashmead WH (1881) On a parasite bred from the eggs of the orange tree plant bug; being another insect friend of the orange-grower. Florida Agriculturist 4: 181-181. *! Ashmead WH (1887) Studies on the North American Proctotrupidae, with descriptions of new species from Florida. Entomologica Americana 3: 73-119. 102 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Ashmead WH (1888) Descriptions of some unknown parasitic Hymenoptera in the collection of the Kansas State Agricultural College, received from Prof. E. A. Popenoe. Bulletin of the Kansas Agricultural Experiment Station Appendix 3: i—viii. * Ashmead WH (1893) A monograph of the North American Proctotrypidae. Bulletin of the United States National Museum 45: 1-472. * doi: 10.5479/si.03629236.45.1 Ashmead WH (1894) Report on the parasitic Cynipidae, part of the Braconidae, the Ichneu- monidae, the Proctotrypidae, and part of the Chalcidinae. Part HI. Zoological Journal of the Linnean Society of London 25: 188-254. ” Ashmead WH (1896) Report on the parasitic Hymenoptera of the island of Grenada, comprising the families Cynipidae, Ichneumonidae, Braconidae, and Proctotrypidae. Proceedings of the Zoological Society of London 1895: 742-812. °° Ashmead WH (1900) Report upon the aculeate Hymenoptera of the islands of St. Vincent and Gre- nada, with additions to the parasitic Hymenoptera and a list of the described Hymenoptera of the West Indies. Transactions of the Royal Entomological Society of London 1900: 207-367. ”” Ashmead WH (1904) Descriptions of new Hymenoptera from Japan — I. Journal of the New York Entomological Society 12: 65-84. * Brues CT (1908) Hymenoptera. Fam. Scelionidae. Genera Insectorum 80: 1-59. ” Brues CT (1910) Notes and descriptions of North American parasitic Hymenoptera. VIII. Bulletin of the Wisconsin Natural History Society 8: 45-52. °° Brues CT (1916) Serphoidea (Proctotrypoidea). In: Viereck. The Hymenoptera or, wasp-like insects, of Connecticut. Guide to the Insects of Connecticut, Part HI. Bull. State Geol. Nat. Hist. Surv. No. 22, 529-577. ° Buffington ML, Gates M (2009) Advanced imaging techniques I: using a compound micro- scope for photographing point-mount specimens. American Entomologist 54: 222-224. doi: 10.1093/ae/54.4.222 Buffington ML, Burks R, McNeil L (2005) Advanced techniques for imaging microhymenoptera. American Entomologist 51: 50-54. doi: 10.1093/ae/51.1.50 Cockerell TDA (1897) A parasite of hemipterous eggs. The Canadian Entomologist 29: 25-26. ° doi: 10.4039/Ent2925b-2 Delucchi VL (1961) Le complexe des Asolcus Nakagawa (Microphanurus Kieffer) (Hymenoptera, Proctotrupoidea) parasites oophages des punaises des cereales au Maroc et au Moyen-Orient. Cahiers de la Recherche Agronomique 14: 41-67. °° Dodd AP (1920) Notes on the exotic Proctotrupoidea in the British and Oxford University Museums, with descriptions of new genera and species. Transactions of the Entomological Society of London 1919: 321-382. “ Fabritius K (1972) Genul Tvissolcus Ashmead 1893 (Hymenoptera,Scelionidae) in Romania si perspectivele utilizarii acestui gen de entomofagi in combaterea biolgica si integrata a plosnitelor cerealelor. Lucrari Stiintifice - Zoologie, Constanta 1972: 27-42. ° Fouts (1920) Some new parasites, with remarks on the genus Platygaster (Hymenoptera). Pro- ceedings of the Entomological Society of Washington 22: 61-72. % Fusu L, Bin F, Popovici OA (2013) First report of chromosomes of the parasitoid wasp Trissolcus basalis (Wollaston) (Hymenoptera: Platygastridae: Telenominae). Entomological Science 16: 263-265. ° doi: 10.1111/ens.12011 Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 103 Gahan AB (1926) A new egg-parasite (Hymenoptera: Serphoidea). Proceedings of the Ento- mological Society of Washington 28: 67. ® Gahan AB (1932) Miscellaneous descriptions and notes on parasitic Hymenoptera. Annals of the Entomological Society of America 25: 736-757. © doi: 10.1093/aesa/25.4.736 Ghahari H, Buhl PN, Kocak E (2011) Checklist of Iranian Trissolcus Ashmead (Hymenoptera: Platygastroidea: Scelionidae: Telenominae). International Journal of Environmental Studies, 68: 593-601. ”° Golin V, Loiacono MS, Margaria CB, Aquino DA (2011) Natural incidence of egg parasitoids of Edessa meditabunda (F.) (Hemiptera: Pentatomidae) on Crotalaria spectabilis in Campo Novo do Parecis, MT, Brazil. Neotropical Entomology 40: 617-618. ” Graham MWR de V (1984) Madeira insects, mainly Hymenoptera Proctotrupoidea, Cer- aphronoidea, and Bethyloidea. Boletim do Museu Municipal do Funchal 36: 83-110. ” Harrington WH (1900) Catalogue of Canadian Proctotrypidae. Transactions of the Royal Society of Canada (2)5(4): 169-206. ” Harris AC (2010) Halyomorpha halys (Hemiptera: Pentatomidae) and Protaetia brevitarsis (Coleoptera: Scarabeidae: Cetoniinanae) intercepted in Dunedin. The Weta 40: 42-44. Hagedorn G, Catapano T, Giintsch A, Mietchen D, Endresen D, Sierra S, Groom Q, Biserkov J, Glockler F, Morris R (2013) Best practices for stable URIs. “4 Hirashima Y, Yamagishi K (1981) Redescriptions of the types of some Japanese Scelionidae preserved in the United States National Museum (Hymenoptera, Proctotrupoidea). Journal of the Faculty of Agriculture, Kyushu University 25: 153-159.” Hoebeke ER (1980) Catalogue of the Hymenoptera types in the Cornell University Insect Col- lection. Part I: Symphyta and Apocrita (Parasitica). Search: Agriculture; Cornell University Agricultural Experiment Station 9: 1-36. ”° Hoebeke ER, Carter ME (2003) Halyomorpha halys (Stal) (Heteroptera: Pentatomidae): A polyphagous plant pest from Asia newly detected in North America. Proceedings of the Entomological Society of Washington 105: 225-237. Inkley DB (2012) Characteristics of home invasion by the brown marmorated stink bug (He- miptera: Pentatomidae). Journal of Entomological Science 47: 125-130. Johnson NF (1983) Types of Neotropical Telenominae described by W. H. Ashmead and P. Cameron (Hymenoptera: Scelionidae). Proceedings of the Entomological Society of Washington 85: 439-449. ”” Johnson NF (1984) Revision of the Nearctic species of the Trissolcus flavipes group (Hymenop- tera: Scelionidae). Proceedings of the Entomological Society of Washington 86: 797-807. ” Johnson NF (1984) Systematics of Nearctic Telenomus: classification and revisions of the podisi and phymatae species groups (Hymenoptera: Scelionidae). Bulletin of the Ohio Biological Survey 6: 1-113.” Johnson NF (1985a) Revision of the New World species of the thyantae group of Trissolcus (Hymenoptera: Scelionidae). The Canadian Entomologist 117: 107-112. *° doi: 10.4039/ Ent117107-1 Johnson NF (1985b) Systematics of New World Trissolcus (Hymenoptera: Scelionidae): spe- cies related to 7. basalis. The Canadian Entomologist 117: 431-445. *' doi: 10.4039/ Ent117431-4 104 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Johnson NF (1987) Systematics of New World Trissolcus, a genus of pentatomid egg-par- asites (Hymenoptera: Scelionidae). Journal of Natural History 21: 285-304. ** doi: 10.1080/00222938700771021 Johnson NF (1991) Revision of Australasian T7issolcus species (Hymenoptera: Scelionidae). Invertebrate Taxonomy 5: 211-239. * doi: 10.1071/IT9910211 Kerr PH, Fisher EM, Buffington ML (2008) Dome lighting for insect imaging under a microscope. American Entomologist 54: 198-200. doi: 10.1093/ae/54.4.198 Kieffer JJ (1906) Beschreibung neuer Proctotrypiden aus Nord- und Zentralamerika. Berliner Entomologische Zeitschrift 50: 237-290. * Kieffer JJ (1912) Proctotrypidae (3e partie). Species des Hyménoptéres d’Europe et d’Algérie, 11: 1-160. ® Kieffer JJ (1926) Scelionidae. Das Tierreich. Vol. 48. Walter de Gruyter & Co., Berlin, 885 pp. *° Kononova SV (1995) [25. Fam. Scelionidae]. In: Lehr. [Key to insects of Russian Far East in six volume. vol. 4. Neuropteroidea, Mecoptera, Hymenoptera. Part 2. Hymenoptera]. Dal’nauka, Vladivostok, 57-121. * Kozlov MA (1968) [Telenomines (Hymenoptera, Scelionidae, Telenominae) of the Caucasus — egg parasites of the sunn pest (Eurygaster integriceps Put.) and other grain bugs]. Trudy Vsesoyuznogo Entomologicheskogo Obshchestva 52: 188-223. ** Kozlov MA (1978) [Superfamily Proctotrupoidea]. In: Medvedev. [Determination of insects of the European portion of the USSR]. Vol. 3, part 2. Nauka, Leningrad, 538-664. * Kozlov MA, Kononova SV (1983) [Telenominae of the fauna of the USSR]. Nauka, Leningrad, 336 pp. ”° Kozlov MA, Lé XH (1976) [Palearctic species of the Trissolcus flavipes Thomson group (Hymenoptera, Proctotrupoidea, Scelionidae)]. Entomologicheskoye Obozreniye 55: 657-667. *' Kozlov MA, Lé XH (1977) [Palearctic species of egg parasites of the genus Trissolcus Ashmead, 1893 (Hymenoptera, Scelionidae, Telenominae)]. Insects of Mongolia 5: 500-525. ** Krombein KV, Burks BD (1967) Hymenoptera of America north of Mexico. Synoptic Catalog (Agriculture Monograph No. 2). Second supplement. United States Government Printing Office, Washington, 584 pp. ”° Leskey TC, Hamilton GC, Nielsen AL, Polk DF, Rodriguez-Saona C, Bergh JC, Her- bert DA, Kuhar TP, Pfeiffer DR, Dively GP, Hooks C, Raupp MJ, Shrewsbury PM, Krawczyk G, Shearer PW, Whalen J, Koplinka-Loehr C, Myers E, Inkley D, Hoelmer KA, Lee D-H, Wright SE (2012) Pest status of the brown marmorated stink bug, Ha- lyomorpha halys (Stal), in the USA. Outlooks on Pest Management 23: 218-226. doi: 10.1564/230ct07 Mani MS (1936) Some new and little known parasitic Hymenoptera from India. Record of the Indian Museum, Calcutta 38: 333-340. ” Mao M, Valerio A, Austin AD, Dowton M, Johnson NF (2012) The first mitochondrial ge- nome for the wasp superfamily Platygastroidea: the egg parasitoid Trissolcus basalis. Ge- nome 55: 194-204. ”° doi: 10.1139/g2012-005 Masner L (1959) Some problems of the taxonomy of the subfamily Telenominae (Hym. Sceli- onidae). Trans. I. Int. Conf. Insect Pathology and Biol. Control, Prague 1958: 375-382. °° Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 105 Masner L (1964) A comparison of some Nearctic and Palearctic genera of Proctotrupoidea (Hymenoptera) with revisional notes. Casopis Ceskoslovenské Spolecnosti Entomologické 61: 123-155. ”” Masner L (1965) ‘The types of Proctotrupoidea (Hymenoptera) in the British Museum (Natu- ral History) and in the Hope Department of Entomology, Oxford. Bulletin of the British Museum (Natural History) Entomology Supplement 1: 1-154. °° Masner L, Muesebeck CFW (1968) The types of Proctotrupoidea (Hymenoptera) in the United States National Museum. Bulletin of the United States National Museum 270: 1-143. ” doi: 10.5479/si.03629236.270 Mayr GL (1879) Ueber die Schlupfwespengattung Telenomus. Verhandlungen der Zoologisch- Botanischen Gesellschaft in Wien 29: 697-714. '°° Mik6 I, Vilhelmsen L, Johnson NF, Masner L, Pénzes Z (2007) Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa 1571: 1—78.'°' Morrill AW (1907) Description of a new species of Telenomus with observations on its habits and life history. American Naturalist 41: 417-430. '” doi: 10.1086/278806 Muesebeck CFW, Walkley LM (1951) Superfamily Proctotrupoidea. In: Muesebeck CFM, Krombein KV, Townes HK. Hymenoptera of America north of Mexico — Synoptic Cata- log. U.S. Dept. Agriculture Monograph No. 2, 655-718. '° Nixon GEJ (1935) A revision of the African Telenominae (Proctotrupoidea, fam. Scelionidae). Transactions of the Royal Entomological Society of London 83: 73-103. '‘ doi: 10.1111/ j.1365-2311.1935.tb00416.x Nixon GEJ (1939) Parasites of hemipterous grain-pests in Europe (Hymenoptera: Proc- totrupoidea). Arbeiten tiber Morphologische und Taxonomische Entomologie aus Berlin- Dahlem 6: 129-136. !° Nixon GE] (1943) A synopsis of the Ethiopian and Indo-Malayan species of Microphanurus (Serphoidea, Scelionidae). Bulletin of Entomological Research 34: 135-144. '°° Qiu LF, Yang ZQ, Tao WQ (2007) [Biology and population dynamics of Trissolcus halyo- morphae]. Scientia Silvae Sinicae 43: 62-65. '°” Risbec J (1950) Contribution a [étude des Proctotrupidae (Serphiidae). (II). In: Risbec. Travaux du Laboratoire d’Entomologie du Secteur Soudanis de Recherches Agronomiques. Gouvernement Général de Afrique Occidentale Frangaise, Paris, 511-639. '°° Rjachovsky VV (1959) [Egg parasites of the sunn pest in the Ukrainian SSR]. Ukrainskii Nauchno-Issledovatel’skii Institut Zashchity Rastenii 8: 76-88. '° Ryu J, Hirashima Y (1984) Taxonomic studies on the genus T7issolcus Ashmead of Japan and Korea (Hymenoptera, Scelionidae). Journal of the Faculty of Agriculture, Kyushu University 29: 35-58. 11° Safavi M (1968) Etude biologique et ecologique des hymenopteres parasites des oeufs des pu- naises de cereales. Entomophaga 13: 381-495, '"! Sarazin MJ (1986) Primary types of Ceraphronoidea, Evaniodea, Proctotrupoidea, and Trigona- loidea (Hymenoptera) in the Canadian National Collection. The Canadian Entomologist 118: 957-989. ! doi: 10.4039/Ent118957-10 Schulz WA (1906) Spolia Hymenopterologica. Junfermannschen Buchhandlung, Paderborn, 355-pps"” 106 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Szabé JB (1975) Neue Gattungen und Arten der palaearktischen Telenominen (Hymenoptera, Scelionidae). Annales Historico-Naturales Musei Nationalis Hungarici 67: 265-278. |“ Talamas EJ, Buffington M, Hoelmer L (2013) New synonymy of Trissolcus halyomorphae Yang. Journal of Hymenoptera Research 33: 113-117. '” doi: 10.3897/jhr.33.5627 Talamas EJ, Herlihy MV, Dieckhoff C, Hoelmer KA, Bufington ML, Bon M-C, Weber DC (2015) Trissolcus japonicus (Ashmead) emerges in North America. Journal of Hymenoptera Research 43: 119-128. doi: 10.3897/JHR.43.4661 Voegelé J (1962) Isolement d’une espece jumelle d’Asolcus basalis Wollaston (Hymenoptera, Proctotrupoidea). Al Awamia 4: 155-161. 11° Voegelé J (1964) Contribution a la connaissance des stades larvaires des especes du genre Asolcus Nakagawa (Microphanurus Kieffer) (Hymenoptera, Proctotrupoidea). Al Awamia 10: 19-31. 1” Voegelé J (1965) Nouvelle methode d’etude systematique des especes du genre Asolcus. Cas d’Asolcus rungsi. Al Awamia 14: 95-113. '"° Voegelé J (1969) Les hyménopteéres parasites oophages des Aelia. Al Awamia 31: 137-323. '” Watanabe C (1954) Discovery of four new species of Telenominae, egg parasites of pentatomid and plataspid bugs, in Shikoku, Japan. Transactions of the Shikoku Entomological Society cea MES) Sar Wermelinger B, Wyniger D, Forster B (2008) First records of an invasive bug in Europe: Halyomorpha halys (Stal) (Heteroptera: Pentatomidae), a new pest on woody ornamentals and fruit trees? Mitteilungen der Schweizerischen Entomologischen Gesellschaft 81: 1-8. Wollaston TV (1858) Brief diagnostic characters of undescribed Madeiran insects. Annals and Magazine of Natural History 1: 18-125. ™! Xu J, Fonseca D, Hamilton G, Hoelmer K, Nielson (2013) Tracing the origin of brown mar- morated stink bugs, Halymorpha halys. Biological Invasions 16: 153-166. doi: 10.1007/ s10530-013-0510-3 Yang ZQ, Yao XY, Qiu LF, Li ZX (2009) A new species of Trissolcus (Hymenoptera: Scelio- nidae) parasitizing eggs of Halyomorpha halys (Heteroptera: Pentatomidae) in China with comments on its biology. Annals of the Entomological Society of America 102: 39-47. '” doi: 10.1603/008.102.0104 Yoder MJ, Miko I, Seltmann K, Bertone MA, Deans AR (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. !* Zhu G, Bu W, Gao Y, Liu G (2012) Potential Geographic Distribution of Brown Marmorated Stink Bug Invasion (Halyomorpha halys). PLoS ONE 7(2): e31246. ' doi: 10.137 1/jour- nal.pone.0031246 Zuparko RL, Hamai J (1994) Depositions of parasitic Hymenoptera (Insecta) types from the University of California, Berkeley. Pan-Pacific Entomologist 70: 313-317. '” Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... 107 Endnotes http://keys.lucidcentral.org/keys/v3/Nearctic_Trissolcus/ http://morphbank.net/?id=836557 http://morphbank.net/?id=83644 1 http://morphbank.net/?id=836569 http://morphbank.net/?id=836661 http://morphbank.net/?id=836457 http://morphbank.net/?id=836577 http://morphbank.net/?id=836477 9 http://morphbank.net/?id=836585 10 http://morphbank.net/?id=83665 1 11 http://morphbank.net/?id=836508 12 http://morphbank.net/?id=836520 13. http://morphbank.net/?id=836710 14 http://morphbank.net/?id=836536 15. http://morphbank.net/?id=836613 16 http://morphbank.net/?id=836442 17. http://morphbank.net/?id=836494 18 — http://morphbank.net/?id=836630 19 http://morphbank.net/?id=836687 20 http://morphbank.net/?id=836672 21 http://morphbank.net/?id=840334 22. http://morphbank.net/?id=840339 23 http://morphbank.net/?id=836716 24 — http://morphbank.net/?id=840385 25. http://morphbank.net/?id=836718 26 http://morphbank.net/?id=836719 27 http://morphbank.net/?id=836720 28 — http://morphbank.net/?id=836721 29 http://morphbank.net/?id=836722 30. http://morphbank.net/?id=836725 31 http://morphbank.net/?id=836726 32. http://morphbank.net/?id=836727 33 http://morphbank.net/?id=836728 34 — http://morphbank.net/?id=840343 35. http://morphbank.net/?id=836729 36. _http://morphbank.net/?id=836730 37 http://morphbank.net/?id=83673 1 38 — http://morphbank.net/?id=836732 39 — http://morphbank.net/?id=836733 40 http://morphbank.net/?id=836734 41 http://morphbank.net/?id=836735 CON WWM KR OD NH Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) http://morphbank.net/?id=836736 http://morphbank.net/?id=836737 http://morphbank.net/?id=836738 http://morphbank.net/?id=836739 http://morphbank.net/?id=836740 http://morphbank.net/?id=83674 1 http://morphbank.net/?id=836742 http://morphbank.net/?id=836743 http://morphbank.net/?id=836744 urn:lsid:biosci.ohio-state.edu:osuc_pubs:64 urn:|sid:biosci.ohio-state.edu:osuc_pubs:68 urn:lsid:biosci.ohio-state.edu:osuc_pubs:72 urn:lsid:biosci.ohio-state.edu:osuc_pubs:74 urn:|sid:biosci.ohio-state.edu:osuc_pubs:75 urn:lsid:biosci.ohio-state.edu:osuc_pubs:79 urn:|sid:biosci.ohio-state.edu:osuc_pubs:84 urn:lsid:biosci.ohio-state.edu:osuc_pubs:97 urn:lsid:biosci.ohio-state.edu:osuc_pubs:175 urn:lsid:biosci.ohio-state.edu:osuc_pubs:184 urn:lsid:biosci.ohio-state.edu:osuc_pubs:192 urn:lsid:biosci.ohio-state.edu:osuc_pubs:1094 urn:lsid:biosci.ohio-state.edu:osuc_pubs:1033 urn:lsid:biosci.ohio-state.edu:osuc_pubs:409 urn:|sid:biosci.ohio-state.edu:osuc_pubs:1022 urn:lsid:biosci.ohio-state.edu:osuc_pubs:669 urn:lsid:biosci.ohio-state.edu:osuc_pubs:24023 urn:lsid:biosci.ohio-state.edu:osuc_pubs:730 urn:|sid:biosci.ohio-state.edu:osuc_pubs:756 urn:lsid:biosci.ohio-state.edu:osuc_pubs:23605 urn:lsid:biosci.ohio-state.edu:osuc_pubs:23609 urn:|sid:biosci.ohio-state.edu:osuc_pubs:676 urn:lsid:biosci.ohio-state.edu:osuc_pubs:39 1 http://wiki.pro-ibiosphere.eu/wiki/Best_practices_for_stable_URIs urn:lsid:biosci.ohio-state.edu:osuc_pubs:944 urn:|sid:biosci.ohio-state.edu:osuc_pubs:42 1 urn:|sid:biosci.ohio-state.edu:osuc_pubs:718 urn:|sid:biosci.ohio-state.edu:osuc_pubs:1030 urn:lsid:biosci.ohio-state.edu:osuc_pubs:550 urn:lsid:biosci.ohio-state.edu:osuc_pubs:655 urn:|sid:biosci.ohio-state.edu:osuc_pubs:656 urn:|sid:biosci.ohio-state.edu:osuc_pubs:637 urn:lsid:biosci.ohio-state.edu:osuc_pubs:633 84 85 86 87 88 89 90 Di 92 BS 94 95 96 Di. 98 99 100 101 102 103 104 105 106 107 108 109 110 ele 112 LDS 114 bes 116 ey 118 Me?) 120 121 122 123 124 125 Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae)... urn:lsid:biosci.ohio-state.edu:osuc_pubs:399 urn:lsid:biosci.ohio-state.edu:osuc_pubs:328 urn:|sid:biosci.ohio-state.edu:osuc_pubs:310 urn:|sid:biosci.ohio-state.edu:osuc_pubs:602 urn:|sid:biosci.ohio-state.edu:osuc_pubs:688 urn:lsid:biosci.ohio-state.edu:osuc_pubs:326 urn:lsid:biosci.ohio-state.edu:osuc_pubs:364 urn:lsid:biosci.ohio-state.edu:osuc_pubs:1029 urn:|sid:biosci.ohio-state.edu:osuc_pubs:597 urn:lsid:biosci.ohio-state.edu:osuc_pubs:494 urn:lsid:biosci.ohio-state.edu:osuc_pubs:794 urn:lsid:biosci.ohio-state.edu:osuc_pubs:23695 urn:lsid:biosci.ohio-state.edu:osuc_pubs:889 urn:lsid:biosci.ohio-state.edu:osuc_pubs:380 urn:|sid:biosci.ohio-state.edu:osuc_pubs:342 urn:|sid:biosci.ohio-state.edu:osuc_pubs:312 urn:lsid:biosci.ohio-state.edu:osuc_pubs:647 http://www.mapress.com/zootaxa/2007f/zt0 157 1p078.pdf urn:|sid:biosci.ohio-state.edu:osuc_pubs:1095 urn:lsid:biosci.ohio-state.edu:osuc_pubs:22 1 urn:|sid:biosci.ohio-state.edu:osuc_pubs:544 urn:lsid:biosci.ohio-state.edu:osuc_pubs:578 urn:|sid:biosci.ohio-state.edu:osuc_pubs:582 urn:|sid:biosci.ohio-state.edu:osuc_pubs:26594 urn:lsid:biosci.ohio-state.edu:osuc_pubs:412 urn:|sid:biosci.ohio-state.edu:osuc_pubs:649 urn:|sid:biosci.ohio-state.edu:osuc_pubs:558 urn:|sid:biosci.ohio-state.edu:osuc_pubs:506 urn:lsid:biosci.ohio-state.edu:osuc_pubs:357 urn:|sid:biosci.ohio-state.edu:osuc_pubs:853 urn:lsid:biosci.ohio-state.edu:osuc_pubs:548 doi: 10.3897/JHR.33.5627 urn:|sid:biosci.ohio-state.edu:osuc_pubs:1036 urn:lsid:biosci.ohio-state.edu:osuc_pubs:1037 urn:lsid:biosci.ohio-state.edu:osuc_pubs:1039 urn:lsid:biosci.ohio-state.edu:osuc_pubs:23728 urn:lsid:biosci.ohio-state.edu:osuc_pubs:616 urn:lsid:biosci.ohio-state.edu:osuc_pubs:646 http://dx.doi.org/10.1603/008.102.0104 http://dx.doi.org/10.1371/journal.pone.0015991 doi: 10.1371/journal.pone.0031246 urn:|sid:biosci.ohio-state.edu:osuc_pubs:1724 109 110 Elijah J. Talamas et al. / Journal of Hymenoptera Research 43: 45-110 (2015) Supplementary material | URI table of HAO morphological terms Authors: Elijah J. Talamas, Norman E Johnson, Matthew Bufhngton Data type: Microsoft Excel Spreadsheet (-xls) Explanation note: This table lists the morphological terms used in this publication and their associated concepts in the Hymenoptera Anatomy Ontology. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.