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Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae), natural enemies of native and invasive stink bugs (Hemiptera, Pentatomidae)
expand article infoElijah J. Talamas, Norman F. Johnson§, Matthew Buffington
‡ Smithsonian Institution, Washington, DC, United States of America
§ Ohio State University, Columbus, OH, United States of America
Open Access

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 (Stål) 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 T. flavipes. Two new species are described, Trissolcus valkyria sp. n. and T. zakotos sp. n. A neotype is designated for T. 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

Introduction

A decade after its introduction into the United States in 1999, the economically destructive brown marmorated stink bug (BMSB), Halyomorpha halys Stål (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 native 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 observed (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 tissues via its proboscis, bypassing ingestion of pesticides that are deposited on the surfaces of plant tissues. As a result, increased pesticide applications to combat BMSB disrupt 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 presence 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 collections 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 Trissolcus 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, T. japonicus, T. cultratus comb. rev., T. valkyria sp. n. and T. zakotos sp. n., and previously unexplored or unutilized character systems. Following the keys to species, an updated and expanded diagnosis section provides more nuanced discussions on sexual dimorphism and phenotypic plasticity. The authors hope that this publication will demonstrate the long-term relevance of primary taxonomic research. The concepts of the previously described species 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 introduced pests, educated decisions can be made about which species have the greatest potential, 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 Nearctic 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 key1.

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.oclc.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 camera using Cartograph®™ software, or a Leica®™ DMRB compound microscope with a GT-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 Buffington 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 resolution 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 characters in this key. The authors recommend fluorescent desk lamps, or fiber optic lamps with mylar sleeves affixed 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 Trissolcus species key). Fine forceps or a minuten pin achieve this effectively.

Collections

This work is based on specimens deposited in the following repositories with abbreviations 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 axillular carinae located posterodorsal to the wing base (see Figs 1, 17, 19–20, 60, 62). The 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.

Figures 1–2.

1 Trissolcus, lateral view 2 Trissolcus, dorsal view.

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 T. 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 T. 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 basalis group, the episternal foveae are often distinctly separate from the mesopleural pit, and with the exception of some T. 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 T. japonicus (Fig. 70) and T. 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. T. 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 T. 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 T. valkyria.

Mesoscutal humeral sulcus

Among the published descriptions and diagnoses, we have not encountered previous use of this character for species-level delimitation in Trissolcus. In all but one species, T. cosmopeplae, the form of this character is fixed. As stated by Johnson (1985), T. cosmopeplae, as currenly understood, is a highly variable species. We point out that most specimens of T. 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.

Characters annotations

A1–12 antennomeres 1–12 (Fig. 1)

ac acetabular carina (Figs 43–44, 70)

aem anteroventral extension of metapleuron (Figs 1, 43, 58, 82, 88)

anfo antennal foramen (Fig. 1)

aoc anterior ocellus (Fig. 2)

as antennal scrobe (Figs 13, 26, 54)

atcs antecostal sulcus (Figs 2, 29, 61)

ats postacetabular sulcus (Figs 42, 70, 82)

axcr axillar crescent (Figs 1, 17, 19–20, 60, 62)

bs basiconic sensilla (Figs 8–9)

cs clypeal setae (Fig. 23, 25, 27)

ctk central keel (Figs 7, 10, 79)

cx1 procoxa (Fig. 1)

cx2 mesocoxa (Fig. 1)

cx3 metacoxa (Fig. 1)

eps episternal foveae (Figs 35, 43, 48, 70, 82)

fs facial striae (Figs 7, 10, 54, 112)

gc genal carina (Figs 48, 112)

gen gena (Fig. 1)

hoc hyperoccipital carina (Figs 2, 19, 29)

iap interantennal process (Fig. 1)

loc lateral ocellus (Figs 1–2)

lt(s) laterotergite(s) (Figs 2, 11–12)

mc mesopleural carina (Fig. 43, 45, 58, 107)

mdb mandible (Fig. 1)

mmc median mesoscutal carina (Fig. 39, 63)

mms median mesoscutal sulcus (Fig. 93)

mpp mesopleural pit (Figs 35, 43, 82)

ms malar sulcus (Figs 1, 24, 26, 28)

mscm mesoscutum (Figs 1–2)

msct metascutellum (Figs 1–2, 21–22)

mshs mesoscutal humeral sulcus (Figs 29, 30, 46, 49)

mspl mesopleuron (Fig. 1)

mtnm metanotum (Figs 1–2, 21–22)

mtpl metapleuron (Fig. 1)

mtpm metapostnotum (Figs 1–2, 21–22)

nes netrion sulcus (Figs 44, 48)

not notaulus (Figs 2, 29)

of orbital furrow (Figs 1, 24, 26, 2829, 71)

pcxs paracoxal sulcus (Figs 82, 109)

ppm propodeum (Figs 1–2, 21–22)

prnm pronotum (Figs 1–2)

prpl propleuron (Fig. 1)

pvm posteroventral portion of metapleuron (Figs 17, 18, 20, 74, 106)

r radicle (Figs 1, 37, 80, 112)

scu mesoscutellum (Figs 1–2)

ss sublateral seta (Figs 2, 30, 89)

T1–6 mediotergite (Figs 1–2)

tga 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 Telenomus, 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 (Fig. 11) Gryon obesum Masner
Metasoma with laterotergites wide and loosely attached to sternites, metasoma without sharp lateral margin (Fig. 12); female antenna with 10 or 11 antennomeres 2
2 Frons with central keel extending from interantennal process to anterior ocellus (Figs 7, 10); frons with facial striae distinct, striae often extending along inner orbit of compound eye (Figs 7, 10) 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 attenuating before reaching inner orbit of compound eye (Figs 54, 112) 4
3 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) 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) Psix tunetanus (Mineo & Szabó)
4 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) antennomeres (Fig. 9) Telenomus (T. astrictus, T. calvus, T. goliathus, 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 basiconic sensilla on apical 5 antennomeres (Fig. 8) Trissolcus
Figures 3–4.

213 Paratelenomus saccharalis, female (USNMENT00896342), head, mesosoma, metasoma, lateral view 4 Psix tunetanus (USNMENT00989625), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 5–10.

225 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 Trissolcus 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.

Figures 11–12.

2311 Gryon obesum, female paratype (USNMENT00989078), head, mesosoma, metasoma, anterolateral view 12 Trissolcus euschisti, female (OSUC 404912), head, mesosoma, metasoma, ventrolateral view. Scale bars in millimeters.

Figures 13–16.

2413 Telenomus sp., female (USNMENT00872628), head, anterolateral view 14 Telenomus sp., female (USNMENT00903997), metasoma, dorsal view 15 Trissolcus urichi, female holotype (USNMENT00989070), head, anterolateral view 16 Trissolcus urichi, female (USNMENT00896405), metasoma, dorsal 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) (thyantae group) 15
2 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 (Figs 1, 12, 15, 28) (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 orbital furrow uniform in width, not expanded near malar sulcus (Fig. 24) (basalis group) 8
3 Frons between antennal scrobe and anterior ocellus with parallel, arched rugae (Figs 52, 54) T. cultratus (Mayr), comb. rev.
Frons between antennal scrobe and anterior ocellus smooth or with rugae that are not parallel and arched (Figs 26, 40, 55, 59, 65) 4
4 Inner margin of eye with orbital furrow constricted ventrally (Fig. 26); mesoscutellum rugose, at least laterally and usually throughout (Fig. 32); first laterotergite usually with setae (Fig. 92); mesoscutum often with median mesoscutal sulcus (Fig. 93) T. strabus Johnson
Inner margin of eye with orbital furrow expanded near intersection with malar sulcus (Figs 12, 15, 28); mesoscutellum smooth (Fig. 34) or with coriaceous microsculpture (Fig. 33); first laterotergite without setae (Figs 62, 68); mesoscutum without median mesoscutal sulcus Fig. 69) 5
5 Clypeus with 4 setae (Fig. 25); mesopleuron with episternal foveae well-defined and deep, forming a continuous line of cells from postacetabular sulcus to mesopleural pit (Fig. 70); mesoscutum without median mesoscutal carina (Fig. 69) 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) 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. 1) striate (as in Figs 17, 19) T. edessae Fouts
Female with antennal flagellum (A3–A11) infuscate throughout (Fig. 38); area bounded by axillar crescent (axcr Fig. 1) with cells visible only along dorsal margin (Figs 20, 62) 7
7 Mesopleuron with anteroventral portion rugulose (Figs 41, 44–45) T. brochymenae (Ashmead)
Mesopleuron with anteroventral portion smooth or with shallowly impressed microsculpture (Figs 12, 42–43) T. euschisti (Ashmead)
8 Mesoscutellum coarsely rugose (Figs 8081, 90); vertex sharply angled onto occiput (Fig. 81) 9
Mesoscutellum smooth or with coriaceous microsculpture (Figs 33–34); vertex rounded onto occiput (Figs 30, 66) 10
9 Radicle yellow (Fig. 80, 83); metapleuron with paracoxal sulcus indicated by line of distinct foveae in ventral half (Fig. 82) 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
10 Metapleuron with paracoxal sulcus visible in ventral half (Fig. 109); frons with facial striae extending as rugulae from anterior mandibular articulation toward compound eye (Fig. 112) 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 (Figs 61, 66) 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) T. hullensis (Harrington)
Metapostnotum invaginated near metascutellum, separating propodeum from metanotum near metascutellum (as in Fig. 21) T. erugatus Johnson
13 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 incomplete (as in Fig. 44) T. basalis (Wollaston)
Mesoscutellum entirely smooth and setal bases not strongly raised (Fig. 34); mesopleuron with episternal foveae extending dorsally to proximity of mesopleural pit (Fig. 48); netrion sulcus complete (Fig. 48) 14
14 Gena in lateral view bulging (Fig. 102), without genal carina (Fig. 102); mesoscutum without notauli (Fig. 100); anteroventral extension of metapleuron short, not extending to mesocoxa in lateral view (Fig. 102); lateral mesoscutum with mesoscutal humeral sulcus present as a smooth furrow (as in Fig. 30) 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 of cells (Fig. 46) T. cosmopeplae (Gahan)
15 Mesoscutellum covered with shallowly impressed coriaceous microsculpture (Fig. 33) 16
Mesoscutellum entirely smooth, without microsculpture (Fig. 34) 17
16 Frons outside of antennal scrobes with raised, irregular rugulae (Fig. 86); mesoscutum between notauli often with longitudinal rugulae (Fig. 87) T. ruidus Johnson
Frons outside of antennal scrobes coriaceous, without raised rugulae but with more or less well-defined setigerous punctures, (Fig. 79); mesoscutum without longitudinal elements in sculpture (Fig. 77) T. parma Johnson
17 Gena in lateral view bulging (Fig. 74) T. occiduus Johnson
Gena in lateral view narrow (Figs 95, 106) 18
18 Mesopleural carina absent ventrally (Fig. 97) T. thyantae Ashmead
Mesopleural carina complete (Fig. 107) T. valkyria Johnson & Talamas, sp. n.
Figures 17–22.

2517 T. japonicus, female (USNMENT00675989), mesosoma, lateral view 18 T. occiduus, female (USNMENT00764995), mesosoma, lateral view 19 Trissolcus cultratus, female (USNMENT00764850), head and mesosoma, dorsolateral view 20 T. 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.

Figures 23–28.

2623 T. strabus, female (BMSB 1203), mouthparts, anterior view 24 T. hullensis, female, (OSUC 523881), head and mesosoma, anterolateral view 25 T. japonicus, female holotype, (USNMENT00831865), 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. euschisti, female, (BMSB 1223), head mesosoma, anterolateral view. Scale bars in millimeters.

Figures 29–30.

2729 T. strabus, female (BMSB 1202), head mesosoma, metasoma, dorsolateral view 30 T. basalis, female, (USNMENT00954022), head, mesosoma, metasoma, dorsolateral view. Scale bars in millimeters.

Figures 31–34.

2831 T. basalis, female (USNMENT00954023), mesoscutellum, dorsal view 32 T. strabus, female (OSUC 523850), mesoscutellum, dorsal view 33 T. ruidus, (OSUC 76431), mesoscutellum, dorsal view 34 T. occiduus, female (OSUC 76430), mesoscutellum, dorsal view. Scale bars in millimeters.

Figures 35–37.

29 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.

Figures 38–41.

30 Trissolcus brochymenae, female neotype (USNMENT00954611) 38 lateral habitus 39 dorsal habitus 40 head, anterior view 41 head and mesosoma, anterolateral view. Scale bars in millimeters.

Figures 42–45.

3142 T. 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.

Figures 46–49.

32 Trissolcus cosmopeplae 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 50–51.

33 Trissolcus cultratus 50 female lectotype (NHMW 0008A), head, mesosoma, metasoma, lateral view 51 female (USNMENT00764850), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 52–55.

3452 T. cultratus, female lectotype (NHMW 0008A), head, anterolateral view 53 T. flavipes, female lectotype (NHRS-HEVA 000002617), head, anterolateral view 54 T. cultratus, female (USNMENT00675734), head, anterior view 55 T. flavipes, female lectotype (NHRS-HEVA 000002617), head, anterior view. Scale bars in millimeters.

Figures 56–59.

35 Trissolcus edessae 56 female holotype (USNMENT00872412) head, mesosoma, metasoma, lateral view 57 female holotype (USNMENT00872412) head, mesosoma, dorsal view 58 (USNMENT00764981), mesopleuron, metapleuron, lateral view 59 female holotype (USNMENT00872412), head, anterior view. Scale bars in millimeters.

Figures 60–61.

36 Trissolcus erugatus female paratype (USNMENT00903009) 60 head, mesosoma, metasoma, lateral view 61 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 62–65.

37 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, anterior view. Scale bars in millimeters.

Figures 66–67.

38 Trissolcus hullensis 66 female (OSUC 523881), head, mesosoma, metasoma, dorsal view 67 female (USNMENT00903008), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 68–71.

39 Trissolcus japonicus 68 female (USNMENT00872402), head, mesosoma, metasoma, lateral view 69 female (USNMENT00675989), head, mesosoma, metasoma, dorsal view 70 female (USNMENT00675989), head, mesosoma, ventral view 71 female (USNMENT00872402), head, anterolateral view. Scale bars in millimeters.

Figures 72–75.

40 Trissolcus occiduus 72 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.

Figures 76–79.

41 Trissolcus parma 76 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.

Figure 80.

42 Trissolcus radix female (USNMENT00764955), head, mesosoma, metasoma, lateral view. Scale bar in millimeters.

Figures 81–83.

43 Trissolcus radix 81 female paratype (USNMENT00764955), head, mesosoma, metasoma, dorsal view 82 female (USNMENT00764955), head and mesosoma, lateral view 83 female (OSUC 76271), head, anterolateral view. Scale bars in millimeters.

Figures 84–87.

44 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.

Figures 88–91.

45 Trissolcus 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, anterolateral view. Scale bars in millimeters.

Figures 92–93.

46 Trissolcus strabus, female (BMSB 1203) 92 head, mesosoma, metasoma, lateral view 93 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 94–98.

47 Trissolcus thyantae 94 female holotype (USNMENT00989048), head, mesosoma, metasoma, dorsal view 95 female holotype (USNMENT00989048), head, mesosoma, lateral view 96 female (OSUC 76325), head and mesosoma, dorsolateral view 97 female (USNMENT00764991), mesosoma, ventrolateral view 98 female holotype (USNMENT00989048), head, anterolateral view. Scale bars in millimeters.

Figures 99–103.

48 Trissolcus utahensis 99 female (USNMENT00872111), head, mesosoma, metasoma, lateral view 100 female lectotype (USNMENT00), head and mesosoma, dorsal view 101 lectotype female (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.

Figures 104–108.

49 Trissolcus valkyria, female holotype (OSUC 542457) 104 head, mesosoma, metasoma, 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.

Figures 109–112.

50 Trissolcus zakotos 109 female paratype (USNMENT00954596), head, mesosoma, metasoma, lateral view 110 female holotype (USNMENT00903008), head, mesosoma, metasoma, dorsal view 111 female holotype (USNMENT00903008), mesosoma, dorsolateral view 112 female paratype (USNMENT00954600), head, anterolateral view. Scale bars in millimeters. 50

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Taxonomy

Trissolcus basalis (Wollaston)

Figures 30, 31, 35–37; Morphbank 2

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, synonymy, 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 Lê 1977: 516 (keyed); Kozlov 1978: 637 (description); Kozlov and Kononova 1983: 121 (description); 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 specimen 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.

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

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 Aelia cognata Fieber: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Aelia germari Küster: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; 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: Pentatomoidea: 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: Pentatomidae]; emerged from egg of Dolycoris baccarum (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg ofEurydema ornata (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Eurygaster austriaca (Schrank): [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; 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: Pentatomoidea: Pentatomidae]; emerged from egg of Graphosoma semipunctata (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Halyomorpha annulicornis (Signoret): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; 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 Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Nezara viridula (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; 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 Stål: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Oechalia schellenbergi Guérin-Méneville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; solitary egg parasitoid of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Piezodorus hybneri (Gmelin): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Plautia affinis (Dallas): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Raphigaster Laporte: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Solanum nigrum L.: [Solanales: Solanaceae]; collected on bokhara: [Fabales: 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, T. basalis: PORTUGAL: Madeira Reg. Autó., Madeira Island, VII–1855, Wollaston, B.M. TYPE HYM. 9.304 (deposited in BMNH). Holotype, female, T. megacephalus: SAINT VINCENT AND THE GRENADINES: 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 7539875424 (OSUC); OSUC 145814, 7802778147 (QDPC); USNMENT00872088, USNMENT00872089, USNMENT00872090, USNMENT00903007 (USNM). BRAZIL: 67 sex unrecorded, OSUC 7529975365 (OSUC). CHINA: 10 sex unrecorded, OSUC 75366, 7538975397 (OSUC). DEMOCRATIC REPUBLIC OF THE CONGO: 3 females, OSUC 182081182083 (RMCA). DOMINICAN REPUBLIC: 3 sex unrecorded, OSUC 398658398659, 398667 (CNCI). EGYPT: 2 females, 4 sex unrecorded, OSUC 144795144796, USNMENT00872006, USNMENT00872007, USNMENT00872008, USNMENT00872009 (USNM). ERITREA: 1 female, OSUC 17736 (BMNH). FIJI: 16 sex unrecorded, OSUC 7766177676 (BPBM). FRENCH POLYNESIA: 2 sex unrecorded, OSUC 7765977660 (BPBM). GREECE: 1 sex unrecorded, OSUC 398669 (CNCI). IRAN: 1 sex unrecorded, OSUC 144797 (CNCI). ITALY: 2 females, 1 male, OSUC 173847173849 (OSUC). JAMAICA: 2 sex unrecorded, OSUC 398660398661 (CNCI). JAPAN: 1 sex unrecorded, OSUC 144391 (CNCI). MONTSERRAT: 12 sex unrecorded, OSUC 398662 (CNCI); OSUC 145281 (FSCA); OSUC 7528975298 (OSUC). MOROCCO: 1 sex unrecorded, OSUC 17743 (BMNH). NEW CALEDONIA: 1 sex unrecorded, OSUC 77624 (BPBM). OCEANIA: 5 sex unrecorded, OSUC 7762577628 (BPBM); OSUC 75425 (OSUC). SAINT VINCENT AND THE GRENADINES: 3 sex unrecorded, OSUC 143816143818 (LACM). SENEGAL: 1 female, OSUC 17737 (BMNH). SOUTH AFRICA: 6 sex unrecorded, OSUC 145553, 7538475388 (OSUC). TANZANIA: 1 sex unrecorded, OSUC 17741 (BMNH). TONGA: 31 sex unrecorded, OSUC 7762977658 (BPBM); OSUC 75427 (OSUC). TRINIDAD AND TOBAGO: 2 sex unrecorded, USNMENT00764950, USNMENT00764951 (USNM). TURKEY: 3 females, OSUC 1773917740, 17742 (BMNH). UNITED STATES: 38 females, 9 males, 49 sex unrecorded, ANIC DB 32-020995 (ANIC); OSUC 398668 (CNCI); OSUC 131149131186, 154353, 157486157487, 157542157549, 157563157566, 7339, 7525675288 (OSUC); USNMENT00872103, USNMENT00872104, USNMENT00872105, USNMENT00872106, USNMENT00872107, USNMENT00872108, USNMENT00872109 (USNM). VANUATU: 1 male, 1 sex unrecorded, ANIC DB 32-020997 (ANIC); OSUC 75426 (OSUC). ZIMBABWE: 17 sex unrecorded, OSUC 7536775383 (OSUC).

Trissolcus brochymenae (Ashmead)

Figures 38–41, 44–45; Morphbank 3

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, description, 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 (description, keyed); Masner 1965: 126 (type information); Johnson 1983: 448 (lectotype designation); Johnson 1987: 298 (junior synonym of Trissolcus brochymenae (Ashmead)).

Neotype designation

The last known examination of the lectotype of T. 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, T. brochymenae is a morphologically variable species with a geographic distribution that spans the United States. A case study of cryptic species within Trissolcus was recently presented by Matsuo et al. (2014) and a similar phenomenon may exist in other species, including T. brochymenae. Trissolcus brochymenae is morphologically very close to T. euschisti, separable by only a few characters, and in our opinion this increases the need for an objective neotype. The specimen selected for this purpose 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 distinguished 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 T. 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

Associations

emerged from Acrosternum hilare (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Acrosternum impicticorne (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Brochymena arborea (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Hydrangea L.: [Rosales: Hydrangeaceae]; emerged from Murgantia histrionica (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 guildinii (Westwood): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from sentinel egg mass of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: 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: [Rosales: Rosaceae]; collected on soybean: [Fabales: Fabaceae]; living in soybean: [Fabales: Fabaceae]; emerged from egg of stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on turnip greens: [Capparales: Brassicaceae]; collected on wax myrtle: [Myricales: Myricaceae]

Material examined

Neotype, female, T. brochymenae: UNITED STATES: FL, Duval Co., Jacksonville, no date, reared from egg, W. H. Ashmead, USNMENT00965611 (deposited in USNM). Lectotype, female, T. laticeps: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, no date, H. H. Smith, USNM Type No. 2526 (deposited in USNM). Lectotype, female, T. murgantiae: UNITED STATES: LA, East Baton Rouge Parish, Baton Rouge, no date, reared from egg, H. A. Morgan, USNMENT00989032 (deposited in USNM). Holotype, female, T. 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 398724398725 (CNCI); OSUC 373344373345, 495206495305, 7544575499 (OSUC). COLOMBIA: 1 sex unrecorded, OSUC 398719 (CNCI). COSTA RICA: 7 sex unrecorded, OSUC 398702398703, 398714 (CNCI); OSUC 142482142485 (OSUC). DOMINICAN REPUBLIC: 4 sex unrecorded, OSUC 398710398712, 398716 (CNCI). GUATEMALA: 1 sex unrecorded, OSUC 398718 (CNCI). HONDURAS: 1 sex unrecorded, OSUC 398717 (CNCI). JAMAICA: 3 sex unrecorded, OSUC 398708398709, 398713 (CNCI). TRINIDAD AND TOBAGO: 2 sex unrecorded, OSUC 398706398707 (CNCI). UNITED STATES: 71 females, 2 males, 45 sex unrecorded, OSUC 17821 (BMNH); OSUC 398679398688 (CNCI); OSUC 436701 (LACM); OSUC 145555, 157494157503, 266797, 413700413703, 413709413713, 523852523855, 523857523861, 523864523865, 523904523923, 523931, 523933, 523935523937, 523940, 523942, 523944, 542440, 542442, 542445, 542450, 542453542454, 62796, 7046470465, 7543475444, 7642576426 (OSUC); BMSB 1216–1217, OSUC 145648, USNMENT00872091, USNMENT00872092USNMENT00872095, USNMENT00989146USNMENT00989149, USNMENT00989160USNMENT00989170, USNMENT00989173 (USNM). VENEZUELA: 5 sex unrecorded, OSUC 398704, 398720398723 (CNCI). VIRGIN ISLANDS: 2 sex unrecorded, OSUC 398705, 398715 (CNCI).

Trissolcus cosmopeplae (Gahan)

Figures 46–49; Morphbank 4

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 (T. erugatus, T. hullensis, T. 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

Associations

Emerged from egg of Cosmopepla bimaculata (Thomas): [Hemiptera: 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 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: [Cornales: Cornaceae]

Material examined

Holotype, female, T. cosmopeplae: UNITED STATES: IL, Champaign Co., Urbana, 8.VIII.1925, reared from egg, USNMENT00989096 (deposited in USNM). Other material: (9 females, 1 male, 74 sex unrecorded) CANADA: 16 sex unrecorded, OSUC 145181, 398732398743 (CNCI); OSUC 145556, 7561275613 (OSUC). UNITED STATES: 9 females, 1 male, 58 sex unrecorded, OSUC 398744398747 (CNCI); OSUC 413941, 7560675611, 76429, 7712277177 (OSUC).

Trissolcus cultratus (Mayr), comb. rev.

Figures 19, 50–51, 52, 54; Morphbank 5

Telenomus cultratus Mayr, 1879: 699, 701, 703 (original description, keyed, synonymized 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); Szabó 1975: 266, 267 (description, lectotype designation, keyed).

Diagnosis

Trissolcus cultratus is easily distinguished from other members of the flavipes 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 T. brochymenae, T. edessae, T. euschisti, and T. japonicus.

Link to distribution map

Associations

Emerged from egg of Carpocoris pudicus (Poda): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Eurygaster Laporte: [Hemiptera: 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, USNMENT00979613 (CUIC); OSUC 7576575767 (OSUC); USNMENT00675943, USNMENT00675944 (USNM). CHINA: 2 females, UCRC ENT 142635, 143817 (UCRC). CZECH REPUBLIC: 1 female, 3 males, USNMENT00896311, USNMENT00896312, USNMENT00896313, USNMENT00896314 (CNCI). FRANCE: 4 females, OSUC 7575375756 (OSUC). HUNGARY: 3 females, 1 sex unrecorded, OSUC 7577175773, 75783 (OSUC). JAPAN: 32 females, 5 males, OSUC 144472144480, 542363, 542374, 542412, 542415, USNMENT00896136, USNMENT00896138, USNMENT00896140, USNMENT00896305, USNMENT00896307USNMENT00896309, USNMENT00896315, USNMENT00896339, USNMENT00896341 (CNCI); OSUC 75784, 7578675788 (OSUC); UCRC ENT 297012 (UCRC); USNMENT00675730USNMENT00675737, USNMENT00764849 (USNM). RUSSIA: 34 females, USNMENT00896048, USNMENT00896049, USNMENT00896050USNMENT00896054, USNMENT00896074, USNMENT00896075, USNMENT00979282USNMENT00979286, 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 144470144471, USNMENT00896011, USNMENT00896015, USNMENT00896016, USNMENT00896018, USNMENT00896019, USNMENT00896029, USNMENT00896032, USNMENT00896044USNMENT00896046, USNMENT00896112, USNMENT00896113USNMENT00896116, USNMENT00896118, USNMENT00896119, USNMENT00896121, USNMENT00896122, USNMENT00896134, USNMENT00896135, USNMENT00896157, USNMENT00979237, USNMENT00979246USNMENT00979250, USNMENT00979253, USNMENT00979280 (CNCI). SWITZERLAND: 4 females, 1 male, USNMENT00979222USNMENT00979226 (CNCI). TAIWAN: 1 female, UCRC ENT 112210 (UCRC). UNITED KINGDOM: 1 female, USNMENT00916251 (BMNH).

Comments

Kozlov (1968) designated a lectotype for T. flavipes and simultaneously treated T. cultratus as a junior synonym. However, the concept of T. flavipes presented in the key and description of his publication was that of T. cultratus, and not of T. 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. cultratus, contrasting with absence of large rugae and presence of a circular impression on the frons of T. 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.

Trissolcus edessae Fouts

Figures 56–59; Morphbank 6

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 Nearctic Trissolcus in the flavipes group (T. brochymenae, T. euschisti, and T. strabus) by the abruptly bicolored female antennae. It may be separated from T. japonicus by the presence 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 T. cultratus by the absence of parallel arched rugae on the frons. In T. edessae a median mesoscutal carina is often present, and this is absent in T. cultratus and T. japonicus.

Johnson (1984) used the absence of a mesopleural carina in T. edessae as a diagnostic 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

Associations

Emerged from egg of Acrosternum hilare (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Edessa bifida (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; parasite of Edessa bifida (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Euschistus Dallas: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pachycoris torridus (Scopoli): [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]

Material examined

Holotype, female: UNITED STATES: LA, Orleans Parish, New Orleans, 23.VII.1919, C. E. Smith, USNMENT00872412 (deposited in USNM). Other material: (2 females, 1 male, 29 sex unrecorded) EL SALVADOR: 3 sex unrecorded, USNMENT00764980, USNMENT00764981, USNMENT00764993 (USNM). NICARAGUA: 2 sex unrecorded, OSUC 398762398763 (CNCI). UNITED STATES: 2 females, 1 male, 24 sex unrecorded, OSUC 17814 (BMNH); OSUC 398760398761 (CNCI); OSUC 523872 (MEMU); OSUC 145559, 542444, 7561775636 (OSUC); OSUC 145649 (USNM).

Trissolcus erugatus Johnson

Figures 27, 60–61; Morphbank 7

Trissolcus erugatus Johnson, 1985b: 433, 436 (original description, keyed); Sarazin 1986: 980 (type information).

Diagnosis

Trissolcus erugatus may be distinguished from the most common Southwestern species of Trissolcus discussed here, T. 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 T. utahensis). It may be distinguished from T. hullensis by the following characters: metapostnotum invaginated near metascutellum and separating metanotum from propodeum, anterior extension of metapleuron short, not reaching mesocoxa, mandibular teeth shallowly incised; mesopleural carina absent; legs and A1–A6 usually yellow. Trissolcus cosmopeplae may usually be separated from T. 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 T. hullensis, 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 Southwest are easily identifiable, but variation in color and sculpture in the northern part of its range may result in confusion between this species and T. cosmopeplae.

Link to distribution map

Associations

collected on Larrea tridentata (Sessé & Moc. ex DC.) Coville: [Sapindales: Zygophyllaceae]; emerged from egg of Thyanta custator (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on alfalfa: [Fabales: Fabaceae]; 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 398779398780 (CNCI); OSUC 7786077862 (MSWC); OSUC 145560, 7566875672 (OSUC); USNMENT00903009 (USNM). Other material: (3 females, 2 males, 9 sex unrecorded) CANADA: 4 sex unrecorded, OSUC 398784398787 (CNCI). UNITED STATES: 3 females, 2 males, 5 sex unrecorded, OSUC 398781398783, 398788 (CNCI); OSUC 436700 (LACM); OSUC 413943, 523926523927, 523929, 75667 (OSUC).

Trissolcus euschisti (Ashmead)

Figures 12, 20, 28, 42–43, 62–65; Morphbank 8

Telenomus euschristus Ashmead, 1888: ii (original description, spelling error).

Trissolcus euschisti (Ashmead): Ashmead 1893: 161, 162 (emendation, description, generic 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 Johnson (1984)); Kieffer 1926: 127, 128 (description, keyed); Hoebeke 1980: 27 (type information); Johnson 1984: 801, 803 (lectotype designation, junior synonym of Trissolcus euschisti (Ashmead)); Zuparko and Hamai 1994: 314 (type information).

Trissolcus euchisti (Ashmead): Brues 1908: 11 (emendation); Golin, Loiácono, Margaría and Aquino 2011: 618 (host association).

Diagnosis

Trissolcus euschisti may be distinguished from the similar T. brochymenae by the smooth or shallowly impressed sculpture on the ventral portion of the mesopleuron anterior to the mesopleural carina. The smaller specimens of T. euschisti are often quite distinct from the larger ones in the following characters: number of lateral setae on T1, extent of fine wrinkles on T2, extent of rugae on S2, extent of S1 setation, number of setae on the mesopleuron above the mesocoxa, sculpture of the upper 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 T. 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

Associations

Collected on Acer saccharum Marshall: [Sapindales: Aceraceae]; emerged from egg of Acrosternum hilare (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; on leaf of Catalpa Scop.: [Scrophulariales: Bignoniaceae]; emerged from egg of Edessa meditabunda (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of / host egg of Euschistus Dallas: [Hemiptera: 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: Pentatomoidea: Pentatomidae]; unspecified association Pentatomidae: [Hemiptera: Heteroptera: 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: Pentatomidae]; parasite of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; unspecified association Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus serieventris Uhler: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus spinosus (Dallas): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; 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]; collected on red pine: [Pinales: Pinaceae]; collected on wax myrtle: [Myricales: Myricaceae]

Material examined

Lectotype, female, T. euschristus: UNITED STATES: Riley Co., VI, Marlatt, OSUC 256926 (deposited in OSUC). Lectotype, female, T. rufitarsis: UNITED STATES: Ormsby Co., VII, Baker, Cornell U. No. 388.1 (deposited in CUIC). Lectotype, female, T. 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: UNITED STATES: 2 sex unrecorded, OSUC 398789, 398823 (CNCI). Other material: (71 females, 10 males, 243 sex unrecorded) BRAZIL: 3 sex unrecorded, OSUC 7574675748 (OSUC). CANADA: 1 female, 1 male, 32 sex unrecorded, OSUC 145426145427, 398793398798, 398800398803, 398805398817, 398836398840, 398844398847 (CNCI). FRENCH GUIANA: 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, 145409145410, 398790398792, 398818398822, 398824398835, 398841398843 (CNCI); USNMENT00979600, USNMENT00979603, USNMENT00979605 (CUIC); OSUC 436702 (LACM); OSUC 523870523871, 523874 (MEMU); OSUC 145411145418, 145421145425 (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 145419145420 (UCRC); BMSB 1218, 1220–1230, 1232, OSUC 523851, USNMENT00872096USNMENT00872102, USNMENT00989171, USNMENT00989172, USNMENT00989174USNMENT00989179 (USNM).

Trissolcus hullensis (Harrington)

Figures 22, 24, 66–67; Morphbank 9

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 T. 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 T. hullensis, present in T. radix and T. zakotos; sculpture of the mesoscutellum, coriaceous or smooth in southern specimens of T. hullensis, coarsely areolate in T. solocis and T. radix; the color of the radicle, black in T. hullensis, T. zakotos and T. solocis, yellow in T. radix; and the rounded vertex, sharply angled in T. radix and T. solocis.

Link to distribution map

Associations

Emerged from Euschistus servus (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Podisus maculiventris (Say): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Recurvaria milleri Busck: [Lepidoptera: Glossata: Gelechioidea: Gelechiidae].

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, 145392145393, 398853 (CNCI); OSUC 75837 (OSUC). MEXICO: 11 sex unrecorded, OSUC 398854 (CNCI); OSUC 7787077877 (MSWC); OSUC 7583875839 (OSUC). UNITED STATES: 16 females, 1 male, 48 sex unrecorded, OSUC 145391, 398855398857, 542438 (CNCI); OSUC 523873, 523875523882 (MEMU); OSUC 7786577869 (MSWC); OSUC 142487142491, 143851, 145369145373, 145389, 145562, 523856, 523946, 542456, 62453, 70529, 7582675836, 7642776428 (OSUC); OSUC 145374145378, 145380145388, 145390 (UCRC).

Trissolcus japonicus (Ashmead)

Figures 17, 21, 25, 68–71; Morphbank 10

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 nudum); 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), T. japonicus belongs to the flavipes species group, first recognized by Kozlov and Lê (1976) and refined by Johnson (1984). Trissolcus japonicus may be separated from other species of the flavipes group Trissolcus 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

Associations

Emerged from egg of Halyomorpha halys (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Plautia stali Scott: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on mulberry: [Urticales: Moraceae]; emerged from stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of stink bug: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female, D. japonicus: JAPAN: Kanagawa Pref., Ashigarashimo 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 females, 11 males, 1 sex unrecorded, USNMENT00979190, USNMENT00979191, USNMENT00979192USNMENT00979198, USNMENT00979200, USNMENT00979201USNMENT00979221 (CNCI); USNMENT00675704, USNMENT00675738, USNMENT00675739, USNMENT00675743, USNMENT00675747, USNMENT00675925, USNMENT00764940, USNMENT00764941, USNMENT00764944, USNMENT00764948, USNMENT00764949, USNMENT00764984, USNMENT00916255 (USNM). JAPAN: 8 females, 4 males, 10 sex unrecorded, OSUC 144481144482, 398858, USNMENT00896340 (CNCI); OSUC 145632, 7584375848 (OSUC); USNMENT00675755, USNMENT00675770, USNMENT00872125USNMENT00872133 (USNM). RUSSIA: 1 female, USNMENT00979287 (CNCI). SOUTH KOREA: 3 females, 1 male, USNMENT00979251, USNMENT00979254 (CNCI); USNMENT00675705, USNMENT00675708 (USNM).

Trissolcus occiduus Johnson

Figures 18, 34, 72–75; Morphbank 11

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 T. thyantae by the complete mesopleural carina, and from T. parma and T. ruidus by the entirely smooth mesoscutellum and absence of a genal carina.

Link to distribution map

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 Chlorochroa sayi (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatoma sayii (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female: UNITED STATES: CA, Ventura Co., area 2, Point Mugu Naval Air Station, 31.VII–24.VIII.1981, C. D. Nagano & J. N. Hogue, CNCI 0004 (deposited in CNCI). Paratypes: UNITED STATES: 4 females, 5 sex unrecorded, OSUC 17811 (BMNH); OSUC 145180 (CNCI); OSUC 143814 (LACM); OSUC 77864 (MSWC); OSUC 145564, 7621676217 (OSUC); USNMENT00764995, USNMENT00877675 (USNM). Other material: UNITED STATES: 3 females, 1 male, 1 sex unrecorded, OSUC 145365, 76430 (OSUC); USNMENT00954754, USNMENT00979294, USNMENT00979295 (USNM).

Trissolcus parma Johnson

Figures 76–79; Morphbank 12

Trissolcus parma Johnson, 1985a: 110, 111 (original description, keyed); Sarazin 1986: 981 (type information).

Diagnosis

Trissolcus parma may be distinguished from T. 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

Associations

Collected on Medicago sativa L.: [Fabales: Fabaceae]; collected under 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); 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).

Trissolcus radix Johnson

Figures 80, 81–83; Morphbank 13

Trissolcus radix Johnson, 1985b: 432, 440 (original description, keyed).

Diagnosis

Trissolcus radix is most closely related to T. hullensis, T. solocis, and T. zakotos, 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 T. hullensis and T. zakotos.

Link to distribution map

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: 1 sex unrecorded, USNMENT00764955 (USNM). UNITED STATES: 1 female, 2 sex unrecorded, OSUC 145567, 7627076271 (OSUC). Other material: MEXICO: 2 sex unrecorded, USNMENT00896395, USNMENT00896396 (UANL).

Trissolcus ruidus Johnson

Figures 33, 84–87; Morphbank 14

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 T. parma, it may be separated from the other members of the thyantae group by the presence of microsculpture on the mesoscutellum.

Link to distribution map

Material examined

Holotype, female: UNITED STATES: AZ, Cochise Co., Portal, Southwestern Research Station (SWRS), 19.X.1978, Masner & Gibson, CNC No. 18341 (deposited in CNCI). Paratype: UNITED STATES: 1 sex unrecorded, OSUC 145568 (OSUC). Other material: UNITED STATES: 2 females, 1 male, OSUC 7643176432 (OSUC); OSUC 144847 (USNM).

Trissolcus solocis Johnson

Figures 88–91; Morphbank 15

Trissolcus solocis Johnson, 1985b: 433, 441 (original description, keyed).

Diagnosis

Trissolcus solocis may be distinguished from the closely related T. hullensis and T. zakotos by the coarse sculpture of the mesoscutellum. From T. 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

Associations

Emerged from egg of Acrosternum marginatum (Palisot): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Alcaeorrhynchus 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, USNMENT00764956, USNMENT00764957, USNMENT00764958, USNMENT00764959 (USNM). UNITED STATES: 1 female, 1 male, 5 sex unrecorded, OSUC 398866 (CNCI); OSUC 145569, 7630976313 (OSUC).

Trissolcus strabus Johnson

Figures 8, 23, 26, 29, 32, 92–93; Morphbank 16

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 laterotergite, 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 anterior 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 with an entirely rugose mesoscutellum, the entire surface is setose. This leads us to hypothesize that, at least on the mesoscutellum of T. strabus, the rugose sculpture and setation are linked. The specimens with reduced macrosculpture on the mesoscutellum also have reduced sculpture on the lateral mesoscutum (lateral of the notaulus), revealing coriaceous microsculpture.

Link to distribution map

Associations

Emerged from egg of / host egg of Brochymena Amyot & Serville: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; on leaf of apple: [Rosales: Rosaceae]

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 unrecorded) 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

Figures 94–98; Morphbank 17

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 (description, keyed).

Diagnosis

Trissolcus thyantae is most similar to T. occiduus and T. valkyria. It may be separated from T. occiduus by the narrow malar region and from both by the lack of a complete mesopleural carina.

Link to distribution map

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 egg of Panthea furcilla (Packard): [Lepidoptera: Glossata: Noctuoidea: Noctuidae]; emerged from egg of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pinaceae: [Pinales: Pinaceae]; emerged from egg of Thyanta custator (Fabricius): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; 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, IX-1880, reared from egg, E. A. Schwarz, USNMENT00989048 (deposited in USNM). Non-type: UNITED STATES: 1 female, OSUC 266773 (OSUC). Other material: (2 females, 27 sex unrecorded) CANADA: 1 female, 6 sex unrecorded, OSUC 17812 (BMNH); OSUC 145196, 145368, 398870398871 (CNCI); OSUC 145572, 76328 (OSUC). UNITED STATES: 1 female, 21 sex unrecorded, OSUC 157505157506, 157512157520, 7632076327 (OSUC); USNMENT00764991, USNMENT00764994, USNMENT00979296 (USNM).

Trissolcus utahensis (Ashmead)

Figures 99–103; Morphbank 18

Telenomus utahensis Ashmead, 1893: 143, 145, 148 (original description, keyed).

Hadronotus mesillae Cockerell, 1897: 25 (original description, synonymized by Muesebeck & Walkley (1951)); Brues 1910: 47 (keyed); Kieffer 1926: 454, 464 (description, keyed); Muesebeck and Walkley 1951: 694 (junior synonym of Telenomus utahensis Ashmead).

Telenomus ashmeadi Morrill, 1907: 419 (original description, synonymized with Telenomus 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 specimens from the southern part of its range have lighter-colored appendages. In the Nearctic region it is most similar to T. basalis. The two may be distinguished by the color of A1, usually dark, concolorous with the radicle in T. utahensis, and yellow, sharply contrasting with the dark radicle in T. basalis; and the mesoscutellar sculpture, smooth in T. utahensis, coriaceous in T. basalis.

Link to distribution map

Associations

Emerged from Chlorochroa sayi (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Chlorochroa sayi (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Chlorochroa uhleri (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on Fremontia Torr.: [Malvales: Malvaceae]; emerged from egg of Pentatoma ligata Say: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatoma sayii (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; egg parasite of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; unspecified association Pentatomidae: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected 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: Chenopodiaceae]; emerged from Thyanta pallidovirens (Stål): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; collected on tomato: [Solanales: Solanaceae]; on leaf of tomato: [Solanales: Solanaceae]; collected on wild carrot: [Apiales: Apiaceae].

Material examined

Lectotype, female, T. utahensis: UNITED STATES: Wasatch Range, 27.VI.1891, E. A. Schwarz, USNMENT00989049 (deposited in USNM). Paralectotype: UNITED STATES: 1 male, USNMENT00764992 (USNM). Lectotype, female, T. ashmeadi: UNITED STATES: TX, Ward Co., Barstow, 12.IX.1905, reared from egg, A. W. Morrill, USNM Type No. 10364 (deposited in USNM). Holotype, female, H. mesillae: UNITED STATES: NM, Doña 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 145192145193, 398862 (CNCI); OSUC 7641676417 (OSUC). UNITED STATES: 10 females, 3 males, 136 sex unrecorded, OSUC 17807 (BMNH); OSUC 143819143823, 436690436699 (LACM); OSUC 7787877930 (MSWC); OSUC 145230145252, 145635, 405748, 413942, 542448542449, 542451542452, 542455, 7638376415, 7720377212 (OSUC); OSUC 205760 (UCDC); USNMENT00872110USNMENT00872114 (USNM).

Trissolcus valkyria Johnson & Talamas, sp. n.

Figures 104–108; Morphbank 19

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 mesoscutum: coriacious. Area bounded by axillar crescent: smooth. Parapsidal line: absent. Notaulus: present. Median mesoscutal carina: absent; present. Median mesoscutal 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 microsculpture. Mesopleural epicoxal sulcus: comprised of cells. Mesopleural carina: complete. 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. Setation of metapleuron: present. Metapostnotum: invaginated near edge of metascutellum and separating metanoum from propodeum. Color of legs beyond coxae: yellow; femora brown, otherwise variably yellow to brown. Metasomal depression: punctate or crenulate dorsally.

Sublateral setae on T1: absent. Setation of laterotergite 1: absent. Sculpture of T2 posterior to antecostal sulcus: smooth or with very faintly impressed striation; distinctly striate posterior to basal costae.

Diagnosis

Trissolcus valkyria is most similar to T. thyantae with which it shares a mesoscutellum without microsculpture and a narrow gena. Trissolcus valkyria may be separated T. thyantae and T. occiduus by the presence of a complete and well defined mesopleural carina. From T. occiduus it may also be separated by the narrow gena.

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

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 144848144849, 145646 (USNM).

Comments

Trissolcus valkyria, was previously recognized by Johnson but remained undescribed due to a dearth of specimens. A small number of additional specimens are now known, providing in our opinion a sufficient basis for the description of this species.

Trissolcus zakotos Talamas, sp. n.

Figures 109–112; Morphbank 20

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 sulcus: 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 epicoxal sulcus: present as a smooth furrow; comprised of cells. Mesopleural carina: complete; 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. Metapostnotum: invaginated near edge of metascutellum and separating metanoum from 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.

Diagnosis

Trissolcus zakotos is closest to T. radix, with which it shares a well defined 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, T. zakotos has numerous (3–5) rugae radiating from the lateral edge of the clypeus. This character is present is both T. 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 species because of the appearance of its frons. The name is treated as an appositional noun.

Link to distribution map

Associations

Emerged from Apateticus bracteatus (Fitch): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female: UNITED STATES: MT, Ravalli Co., Hamilton, V-1972, W. L. Jellison, USNMENT00903008 (deposited in USNM). Paratypes: UNITED STATES: 22 females, 1 male, USNMENT00954588USNMENT00954589 (CNCI); USNMENT00954586USNMENT00954587 (OSUC); USNMENT00903005, USNMENT00903006, USNMENT00954590USNMENT00954606 (USNM).

Acknowledgements

We are grateful to: Luciana Musetti (OSUC), Sara Hemly (OSUC), Manuela Vizek (NHMW), Hege Vårdal (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 photography and transcribing label data. This work was made possible by funding from the Systematic Entomology Lab, USDA-ARS, and the Beneficial Insect Introduction Research Laboratory. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.

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Endnotes