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Revision of Palearctic Trissolcus Ashmead (Hymenoptera, Scelionidae)
expand article infoElijah J. Talamas, Matthew L. Buffington, Kim Hoelmer§
‡ Smithsonian Institution, Washington, DC, United States of America
§ United States Department of Agriculture, Newark, United States of America
Open Access

Abstract

Species of Trissolcus Ashmead are potent natural enemies of stink bugs (Pentatomidae). Research on biological agents to control the brown marmorated stink bug, Halyomorpha halys (Stål) (BMSB), in Western Europe requires reliable taxonomic resources for identification of Trissolcus wasps. To aid this research endeavor, we present a species identification key to females of Palearctic Trissolcus. Morphological characters and concepts of the genus and species groups are discussed. We discovered a number of nomenclatural and identification issues that we here rectify.

Keywords

Halyomorpha halys, Platygastroidea, biological control, identification key, egg parasitoid

Introduction

The introduction of the brown marmorated stink bug (BMSB) (Halyomorpha halys Stål; Hemiptera: Pentatomidae) into the Eastern United States, followed by invasion on the West Coast, prompted interest in Trissolcus as a classical biological control agent with emphasis on two species, T. cultratus (Mayr) and T. japonicus (Ashmead). Recent work in this genus (Talamas et al. 2015a) provided identification tools to separate these species from the Nearctic fauna. This work followed relatively recent revisions of Nearctic Trissolcus (Johnson 1984a, 1985ab) that made production of these tools a straightforward task. The second taxonomic component of this project is species-level revision of Trissolcus to enable identification of T. japonicus and T. cultratus in their native ranges. The expansive distributions of some species, including T. cultratus, led us to revise the genus for the entire Palearctic region. Some species extend to Southeast Asia and we here include species from this region for which we directly examined primary types.

BMSB has recently become established in Europe (Wermelinger et al. 2008) where its range continues to expand (Haye et al. 2015). In light of this, revision of Trissolcus from this region has assumed greater urgency. With this publication we provide Europe, Asia, and the Middle East with the basic tools of species identification that facilitate molecular diagnostics and research on host selection, species evolution, and biological control.

Criticism of previous taxonomy

Our quest to identify species of the Eastern Palearctic revealed that multiple species range from Western Europe to East Asia, requiring evaluation of material from the entire Palearctic region. In the process we discovered deep-rooted problems in the taxonomy of the group, many of which stem from insufficient or nonexistent examination of primary types. Species descriptions have typically been based on a small number of specimens from a restricted geographic area, and were not part of a thorough revisionary effort. As a result, most of the available names in the Palearctic are synonyms. The detrimental consequences of this are significant and difficult to overemphasize, as they put an incredible strain on taxonomists that are already faced with a formidable array of biodiversity to describe and understand. To be sure, it forces the time consuming loan of type specimens, creates literature with confusing or erroneous name usage, and prevents an accurate understanding of the true diversity within the genus and the distributions of its constituent species. Evaluation of Trissolcus for biological control of BMSB in Europe simply cannot progress without the ability to properly identify the species under study.

The above mentioned issues in Trissolcus are directly relevant to establishing proper name usage for T. japonicus, the primary candidate for biological control in the United States. Trissolcus halyomorphae was carefully characterized in the description by Yang (2009), but its identity as T. japonicus (Ashmead) was not recognized because no primary types were examined. Buhl (1996) apparently examined a single specimen of Trissolcus, which he correctly keyed to the couplet of T. cercus and T. pontus in Kozlov (1978), but nonetheless described the species, T. dobashii, as new. We here treat all three (T. cercus, T. pontus and T. dobashii) as junior synonyms of T. japonicus. Kononova (2014) and her concept of the flavipes group was produced seemingly unaware of Talamas et al. (2013) or Yang (2009) as she redescribed T. japonicus as T. mirus.

Reliance on erroneous literature alone, without reassessment of type material, has compounded taxonomic problems in Palearctic Trissolcus. For example, Trissolcus cultratus (Mayr), originally described as Microphanurus cultratus Mayr, was erroneously treated by Kozlov (1968) as a junior synonym of Trissolcus flavipes. Kozlov subsequently perpetuated the concept of T. cultratus, a distinctive and easily recognized species, under the name T. flavipes. The concept of T. flavipes that we provide here is a trans-Palearctic species which was redescribed as T. circus by Kozlov and Lê (1976) and as T. crassus by Kononova (2014).

The key to species by Kozlov (1978) treats most of the Palearctic fauna, but ignores pleural characters entirely and in some cases the couplets do not match specimens from the type series. We suspect that many of the identification keys are simply regurgitations of previous works without contributing new knowledge to an understanding of the group. Specimen size and distribution were frequently used for identification, precluding the possibility of large distributions or size differences caused by emergence from different host species. Based on our experience comparing the identification keys of Kozlov, Petrov (2013), and Kononova to primary types and specimens from a broad geographic distribution, we recommend that these publications be avoided, as they do not accurately enable species identification. We suspect that all of the species described by Kononova (2014) will be treated as junior synonyms once the material is fully assessed.

Our experience with Trissolcus leads us to believe that other groups in Platygastroidea (esp. Platygastrinae and Telenomus) are likely to harbor large numbers of trans-Palearctic species that have been described multiple times.

The contributions of the authors are as follows: E.J. Talamas: character definition and coding, species concept development, imaging, manuscript preparation; M.L. Buffington: project co-supervisor, manuscript preparation; K. Hoelmer: project co-supervisor, fieldwork coordinator and specimen acquisition.

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 “ZMAS” 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, purl.oclc.org/NET/hymenoptera/hol, and entering the identifier in the form. The taxonomic synopsis was generated by the Hymenoptera Online Database (hol.osu.edu).

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) and MorphBank (http://www.morphbank.net).

The specimen(s) that formed the bases for concepts of species that we here treat as junior synonyms are provided in Morphbank collections following the act of synonymy for each species. The Morphbank collection for the valid name contains all images of that species, including the types of junior synonyms.

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.

Abbreviations and characters annotated in the figures

A1A11 antennomeres 1–11 (Figures 4, 6)

aem anteroventral extension of metapleuron (Figures 4, 52, 58, 99, 118, 128, 189)

anfo antennal foramen (Figures 4, 6)

aoc anterior ocellus (Figure 3)

as antennal scrobe (Figures 150, 191, 204)

asu acropleural sulcus (Figures 5, 153)

atc acetabular carina (Figures 6)

atcs antecostal sulcus (Figures 3)

ats postacetabular sulcus (Figures 56, 37, 74, 170, 202)

axcr axillar crescent (Figures 4–5)

bs basiconic sensilla (Figures 6, 80, 141, 142)

bc basal costae (Figure 3)

cs clypeal setae (Figures 6, 26, 39, 221)

cx1–cx3 coxae (Figures 4, 6, 58, 99, 118, 128)

epc epomial carina (Figures 5, 225)

eps episternal foveae (Figures 5, 6, 37, 50, 74, 105, 154, 170, 202)

ff felt field (Figure 6)

fs facial striae (Figures 37, 75, 76, 193)

gc genal carina (Figures 5, 25, 37)

gen gena (Figures 4, 6)

hoc hyperoccipital carina (Figures 3, 24, 41, 47)

iap interantennal process (Figures 4, 6)

lbr labrum (Figures 6, 50)

loc lateral ocellus (Figures 34)

lpT1 lateral patch on T1 (Figures 5, 146, 149)

lt1–lt6 laterotergite 1–6 (Figures 4, 6)

mc mesopleural carina (Figures 25, 37, 189)

mdb mandible (Figures 4, 6)

mees mesepimeral sulcus (Figures 56, 25, 185)

meps metapleural epicoxal sulcus (Figures 5, 52)

mes mesopleural epicoxal sulcus (Figures 56)

mmc median mesoscutal carina (Figures 47, 50, 215)

mms median mesoscutal sulcus (Figure 3, 98, 100)

mns metanotal trough (Figure 3, 5)

mpit metapleural pit (Figures 64, 94, 133)

mpp mesopleural pit (Figures 56)

ms malar sulcus (Figures 46)

mst malar striae (Figures 193)

mscm mesoscutum (Figures 34)

mshs mesoscutal humeral sulcus (Figures 3, 5, 81, 138, 169)

mspl mesopleuron (Figures 4)

msct metascutellum (Figures 34)

mtnm metanotum (Figures 34)

mtpl metapleuron (Figure 4)

mtpm metapostnotum (Figure 34)

mtps metapleural sulcus (Figure 5)

mv marginal vein (Figures 79, 126)

nes netrion sulcus (Figures 56, 22, 140, 170)

net netrion (Figures 56)

not notaulus (Figure 3)

of orbital furrow (Figures 46, 37, 61, 110–111)

pcxs paracoxal sulcus (Figures 5, 52)

pdms posterodorsal metapleural sulcus (Figures 5, 139, 189)

pmma posterior mesepimeral area (Figure 6)

ppm propodeum (Figures 34)

prcs pronotal cervical sulcus (Figure 5)

prnm pronotum (Figures 34)

prpl propleuron (Figure 4)

pss pronotal suprahumeral sulcus (Figures 3, 5)

pssu prespecular sulcus (Figures 5, 153)

psu posterior scutellar sulcus (Figures 3, 5)

pv postmarginal vein (Figures 79, 126)

r radicle (Figures 4, 6, 26, 188)

S1S6 sternites 1–6 (Figures 6)

sasu subacropleural sulcus (Figure 5, 153)

scu mesoscutellum (Figure 34)

slt1 setation of laterotergite 1 (Figures 5, 110, 146, 149)

shms mesoscutal suprahumeral sulcus (Figures 3, 5)

spec speculum (Figures 5, 154, 185)

ss sublateral seta (Figure 3)

sss scutoscutellar sulcus (Figures 5)

sv stigmal vein (Figure 79, 126)

T1T7 mediotergites 1–7 (Figure 34)

tga tegula (Figures 35)

vplc ventral mesopleural carina (Figures 6, 25)

Collections

This revision is based on material from the following collections:

BMNH British Museum of Natural History, London, England

BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA

CNCI Canadian National Collection of Insects, Ottawa, Canada

EIHU Hokkaido University Museum of Entomology, Sapporo, Japan

EMBT Department of Agriculture, Bangkok, Thailand

HNHM Hungarian Natural History Museum, Budapest, Hungary

IEBR Institute of Ecology and Biological Resources, Hanoi, Vietnam

KUEC Entomology Laboratory of Kyushu University, Fukuoka, Japan

MCSN Museo Civico de Storia Naturale “Giacomo Doria”, Genoa, Italy

MNHN Muséum National d’Histoire Naturelle, Paris, France

MZUF Muséum National d’Histoire Naturelle, Paris, France

NHMW Naturhistorisches Museum Wien, Vienna, Austria

NHRS Naturhistoriska Riksmuseet - Entomology, Stockholm, Sweden

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany

OSUC C.A. Triplehorn Collection, The Ohio State University, USA

UCRC Entomology Research Museum, University of California, Riverside, California, USA

USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

ZIN Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia

ZMUC Zoological Museum, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark

Character discussions

Clypeal setae

The number of setae on the clypeus is at times a challenging character to observe, but it is very useful for delimiting the flavipes group and separating similar species within it. Consistent with Talamas et al. (2015a), species in the flavipes group have 4 or fewer clypeal setae, and those in the thyantae group have 6. We find that the number of clypeal setae in the basalis group is almost always 6, but we encountered a small number of specimens of T. comperei with 8 clypeal setae. Scanning electron microscopy revealed that in addition to the two prominent clypeal setae, T. cultratus has a pair of extremely small setae that are not visible with light microscopy. We here clarify that our treatment of these setae is based on brightfield microscopy, and note that the actual number of setae may vary when analyzed in an SEM.

Episternal foveae

In our revisionary efforts, we found no explicit mention of episternal foveae, nor are we aware of their use in any identification tools for Palearctic Trissolcus. The separation, or lack thereof, of the episternal foveae from the postacetabular sulcus and mesopleural pit, as well as their presence or absence, is relatively easy to interpret and has great diagnostic utility in Trissolcus.

Subacropleural sulcus

We here coin the name subacropleural sulcus for the line of foveae anterior to the prespecular sulcus that extends from the acropleural sulcus toward the mesopleural pit (Figures 5, 153).

Posterodorsal metapleural sulcus

Directly ventral to the metapleural carina there is often a line of cells in Scelionidae (Figs 5, 139, 189). We here coin the name posterodorsal metapleural sulcus for this character. In many species this sulcus is a clearly defined line of cells, whereas in others its definition is challenged by coarse sculpture or irregular forms of the cells that comprise the sulcus.

Generic diagnosis of Holarctic Trissolcus

The characters presented in Talamas et al. (2015a) will separate most, but not all, species of Palearctic Trissolcus from Telenomus. A four-merous clava is found in T. hyalinipennis and T. oobius, compounded by as many as 6 clavomeres that can be found in Telenomus. Trissolcus perepelovi and T. plautiae have a frons that can be largely smooth, reducing the reliability of a sculptured frons as a diagnostic character for Trissolcus. The ratio of width to length of T2: wider than long in Trissolcus and longer than wide in Telenomus, remains unequivocal in the Palearctic region as far as we know.

In the Nearctic, we encountered a species of Telenomus (USNMENT01197254), reared from lepidopteran eggs, with a 5-merous clava and T2 wider than long. This species can easily be separated from Trissolcus by an occipital carina that extends to the anterior articulation of the mandible. In all Holarctic Trissolcus, the occipital carina extends to the posterior articulation of the mandible.

The subacropleural sulcus is present in all species of Holarctic Trissolcus with the sole exception of T. exerrandus. We have yet to thoroughly examine Telenomus from the Palearctic region, and thus cannot give a complete analysis of this character’s diagnostic power, but nonetheless we consider it to be a useful generic character for Trissolcus.

Species groups

The creation of species-groups can be very useful for identifying subgeneric lineages that represent monophyletic groups or morphologically similar clusters of species. However, as more species are examined, many species cannot be placed in species-groups without drastically altering their limits. Trissolcus stoicus presents a challenge to our concepts of species-groups because it exhibits characters suggesting relation to both the flavipes and thyantae groups. The complete hyperoccipital carina aligns it with the flavipes group, but it has 6 clypeal setae. The posteroventral metapleuron has 2 large setae, perhaps an intermediate form between the glabrous metapleuron of the flavipes group and the dense setation typical of the thyantae group. This species also lacks notauli and well-defined episternal foveae of the thyantae group and we here do not treat it as part of any species group. As the taxonomic treatment of Trissolcus expands geographically, the concepts of species groups will likely require modification, as has occurred following our revision of the Palearctic species.

basalis species group

The basalis species-group in the context of the Nearctic fauna was defined by a glabrous metapleuron and the absence of a hyperoccipital carina. We here continue to approach this cluster of species as the basalis group, although where the basalis group ends become increasingly blurry as more species are examined, particularly in Southeast Asia.

flavipes species group

Multiple species in the flavipes group from the Palearctic region do not comply with the characters in the key to species of Nearctic Trissolcus by Talamas et al. (2015a). The notaulus may be very weakly indicated and present only at the posterior margin of the mesoscutum, as in T. gonopsidis, and the hyperoccipital carina in T. eriventus may be unrecognizable. Use of these characters is further complicated by species with a hyperoccipital carina that are well outside of the flavipes group (T. carinifrons and T. stoicus). Regarding the Holarctic fauna and what we have seen from Southeast Asia, the number of clypeal setae (4 or fewer) is the most reliable character for identifying the flavipes group and is the primary character that we use for its delimitation.

Kononova (2014) presented a new concept of the flavipes species group, one based on a postmarginal vein longer than the stigmal vein. The vast majority of species in Trissolcus have this form of wing venation, and thus this concept offers very little for subgeneric classification or identification. We prefer more restrictive species group concepts because they enable us to produce more efficient identification tools. However, there is presently no objective basis for selection of one species-group concept over another because there is no phylogeny to demonstrate which, if any, of the concepts correspond to monophyletic groups.

thyantae species group

The thyantae species-group in the Nearctic is characterized by setae in the posteroventral portion of the metapleuron, presence of notauli, episternal foveae present as large, distinct cells that are often elongate horizontally and form a continuous line from the dorsal limit of the postacetabular sulcus (or acetabular carina) to the mesopleural pit, a transversely strigose speculum, and six clypeal setae. These characters are valid for the Palearctic species, but there are some notable species (T. larides, T. levicaudus, T. tersus) with a densely setose metapleuron that lack the other defining characters of the thyantae group. Some of these species are also morphologically close to species that lack setation of the metapleuron. We here choose not to expand the concept of the thyantae group to include all species with a setose metapleuron until additional phylogenetic analysis determines whether or not this character delimits a monophyletic lineage.

The relationships between species of the thyantae group across the Northern Hemisphere warrant further investigation. We here present a concept of T. scutellaris that contains a great deal of variation in the sculpture of the frons, mesoscutum and mesoscutellum, and in the degree of development of the mesopleural carina. Among the specimens at hand there is a continuous gradient of sculptural development, and clear lines cannot be drawn between T. scutellaris and the species that we treat as junior synonyms. In the Nearctic, these characters appear to be fixed and are thus used in species diagnosis. We were unable to find morphological characters to separate the more rugose specimens of T. scutellaris (Palearctic) from T. ruidis Johnson (Nearctic), and the smoother specimens from T. valkyria Johnson & Talamas (Nearctic). We refrain from treating the Nearctic species as junior synonyms at this time because of their disjunct distributions and the possibility that these are cryptic species. Molecular data will undoubtedly be useful in determining how well our morphological species concepts match concepts generated from DNA data; the combination of these data will help determine with greater confidence if T. scutellaris is a Holarctic species.

Illumination

High quality optics and bright, diffuse lighting are necessary for observing the characters in this key. The authors recommend fluorescent desk lamps, or fiber optic lamps with a mylar shield between the tip of the light pipes and the specimen. The mylar should be placed close to the specimen, with the light source approximately 10 cm from the mylar (Figures 1–2) for the optimal light-scattering effect. The characters used in this key are evaluated with brightfield microscopy.

Figures 1–2. 

Configuration of mylar shield for reducing glare.

Key to Trissolcus of Europe (females)

1 Vertex between lateral ocelli with hyperoccipital carina (Figures 43, 72); carina sometimes weakened medially (Figures 47, 200, 206–207); clypeus with 4 or fewer setae (Figures 69–70, 85, 92, 221) 2 (flavipes group)
Vertex between lateral ocelli without hyperoccipital carina (Figures 29, 104, 108); clypeus with 6 or 8 setae (Figures 97, 123, 172–177) 8
2 Laterotergite 1 with line of setae dorsally (Figures 5, 146, 149) Trissolcus saakowi (Mayr), p. 115
Laterotergite 1 without setae (Figures 28, 35) 3
3 At intersection with malar sulcus, orbital furrow expanded with medial margin well-defined (Figures 37–38, 48, 84, 86–87, 93) 4
At intersection with malar sulcus, orbital furrow absent or without a defined medial margin (Figures 42, 44, 65, 69–70, 73–76) 7
4 Mesoscutum with median mesoscutal carina (Figure 51) Trissolcus edessae Fouts, p. 49
Mesoscutum without median mesoscutal carina (Figures 200, 206–208) 5
5 Mesopleuron with episternal foveae distant from postacetabular sulcus and mesopleural pit, often weakly indicated (Figure 202); vertex with hyperoccipital carina weakened medially (Figure 200) Trissolcus tumidus (Mayr), p. 148
Mesopleuron with episternal foveae forming a continuous line of cells from postacetabular sulcus to mesopleural pit (Figures 37, 88, 133, 197, 219); vertex with hyperoccipital carina uniform and robust (Figures 72, 89) 6
6 Frons above antennal scrobe with irregular raised lines (Figures 92–93); mesoscutum with microsculpture effaced posteriorly (Figure 91); median lobe of mesoscutum with oblique rugulae (Figure 91); mesoscutellum entirely smooth (Figure 91) Trissolcus kozlovi Rjachovskij, p. 79
Frons above antennal scrobe with transverse rugae or microsculpture alone, but not irregular lines (Figures 84–87); mesoscutum usually with microsculpture present throughout (Figure 89), sometimes with sculpture effaced posteriorly; rugulae on mesoscutum, if present, oriented longitudinally between notauli; mesoscutellum smooth or with microsculpture Trissolcus japonicus (Ashmead), p. 74
7 Frons above antennal scrobe with coarse arcuate rugae (Figures 44, 45) Trissolcus cultratus (Mayr), p. 45
Frons above antennal scrobe with shallow rugulae forming a circle (Figures 69–70) Trissolcus flavipes (Thomson), p. 63
8 Metapleuron with posteroventral portion densely setose (Figures 154, 213) 9 (thyantae group)
Metapleuron without setae below metapleural sulcus (Figures 20, 56–58, 78) 10 (basalis group)
9 Gena distinctly broad and bulging in lateral view (Figure 213); compound eye with posterior and dorsal margins separated from hyperoccipital carina by a broad furrow (Figure 213) Trissolcus viktorovi Kozlov, p. 157
Gena narrow to moderately broad in lateral view (Figures 154, 209); compound eye with posterior and dorsal margins narrowly separated from hyperoccipital carina (Figures 156–157, 212) Trissolcus scutellaris (Thomson), p. 120
10 Antenna with 4 clavomeres (A8A11 with basiconic sensilla) (Figures 80, 82) Trissolcus hyalinipennis Rajmohana & Narendran, p. 70
Antenna with 5 clavomeres (A7A11 with basiconic sensilla) (Figures 6, 141–142) 11
11 Mesopleuron without episternal foveae (Figures 16, 132, 140); metapleuron with anteroventral extension short, not reaching base of mesocoxa (Figures 118, 128) Trissolcus rufiventris (Mayr), p. 111
Mesopleuron with at least one episternal fovea (Figures 74, 105, 171) indicated or metapleuron with anteroventral extension extending to base of mesocoxa (Figures 52, 57–58, 171) 12
12 Metapleuron with paracoxal sulcus and metapleural epicoxal sulcus indicated by distinct foveae (Figures 52, 56–58) Trissolcus elasmuchae (Watanabe), p. 53
Metapleuron with paracoxal sulcus or metapleural epicoxal sulcus poorly defined or not indicated externally (Figures 20, 171) 13
13 Lateral pronotum with netrion sulcus complete dorsally, netrion distinct (Figures 170–171); lateral mesoscutum with mesoscutal humeral sulcus usually comprised of cells (Figure 169), occasionally present as a smooth furrow Trissolcus semistriatus (Mayr), p. 129
Lateral pronotum with netrion sulcus incomplete dorsally, netrion often poorly defined (Figure 22); lateral mesoscutum with mesoscutal humeral sulcus present as a smooth furrow (Figures 20–21) Trissolcus basalis (Wollaston), p. 30
Figure 3. 

Line drawing of dorsal Trissolcus.

Figures 4–5. 

4 Line drawing of lateral Trissolcus 5 Line drawing of lateral Trissolcus.

Figure 6. 

Trissolcus japonicus, female (USNMENT00896000), head, mesosoma, metasoma, ventral view. Scale bar in millimeters.

Figure 7–12. 

7 T. mitsukurii, female (USNMENT00675722), axillar crescent, lateral view 8 T. tumidus, female (USNMENT00979425), axillar crescent, lateral view 9 T. flavipes, holotype of T. circus, female (ZMAS 0143), axillar crescent, lateral view 10 T. japonicus, female (USNMENT00675989), axillar crescent, lateral view. 11 T. saakowi, female (USNMENT00977544) axillar crescent, lateral view 12 T. nycteridaner, female (USNMENT00896178), axillar crescent, lateral view. Scale bars in millimeters.

Key to Trissolcus of the Palearctic region (females)

1 Inner margin of eye with orbital furrow broad and transversely rugulose at midpoint of eye (Figure 61); clypeus with 2 setae (Figures 69–70) Trissolcus eriventris (flavipes group in part), p. 57
Inner margin of eye with orbital furrow smooth (Figure 48), narrow (Figures 54, 97), or poorly defined at midpoint of eye; clypeus with 2–8 setae 2
2 Vertex between lateral ocelli with hyperoccipital carina (Figures 24, 43, 72), carina sometimes weakened medially (Figures 47, 200, 206–207) 3
Vertex between lateral ocelli without hyperoccipital carina (Figures 29, 104, 108) 18
3 Clypeus with 6 setae (Figure 26); mesopleural carina complete, extending to ventral mesopleural carina (Figure 25); mesoscutum with notauli weakly developed or absent (Figures 24, 184) 4
Clypeus with 4 or fewer setae (Figures 39, 66); mesopleural carina variable; mesoscutum often with distinct notauli (Figures 63, 68, 89) 5 (flavipes group)
4 Gena smooth and shining posterior to genal carina (Figure 185); mesopleuron with mesepimeral sulcus comprised of transversely elongate cells (Figure 185); mesopleuron with speculum divided by dorsoventral furrow (Figure 185); radicle shorter than width across clypeus (Figure 188) Trissolcus stoicus (Nixon), p. 139
Gena with microsculpture and rugae posterior to genal carina (Figure 25); mesopleuron with mesepimeral sulcus comprised of more of less circular foveae (Figure 25); mesopleuron with speculum undivided (Figure 27); radicle as long as width across clypeus (Figure 26) Trissolcus carinifrons (Cameron), p. 35
5 Laterotergite 1 with line of setae dorsally (Figures 5, 146, 149) Trissolcus saakowi (Mayr), p. 115
Laterotergite 1 without setae (Figures 28, 35) 6
6 At intersection with malar sulcus, orbital furrow expanded with medial margin well-defined (Figures 37–38, 48, 84, 86–87, 93) 7
At intersection with malar sulcus, orbital furrow absent or without a defined medial margin (Figures 42, 44, 65, 69–70, 73–76) 14
7 Mesoscutum with median mesoscutal carina (Figure 51) Trissolcus edessae Fouts, p. 49
Mesoscutum without median mesoscutal carina (Figures 89, 195, 200) 8
8 Mesopleuron with episternal foveae distant from postacetabular sulcus and mesopleural pit, often weakly indicated (Figure 202); vertex with hyperoccipital carina weakened medially (Figure 200) Trissolcus tumidus (Mayr), p. 148
Mesopleuron with episternal foveae forming a continuous line of cells from postacetabular sulcus to mesopleural pit (Figures 37, 88, 133, 197, 219); vertex with hyperoccipital carina uniform and robust (Figures 72, 89) 9
9 Frons between median ocellus and antennal scrobe with a dorsoventral furrow surrounded by area of effaced microsculpture (Figures 196–197); T2 with longitudinal striae present only in medial third of tergite (Figure 195) Trissolcus trophonius (Nixon), p. 145
Frons without dorsoventral furrow (Figures 84, 87, 92, 135) or if furrow present, then microsculpture not effaced (Figures 85–86); T2 with striae present throughout anterior half of tergite (Figures 36, 89, 134) 10
10 Frons above antennal scrobe with irregular raised lines (Figures 92–93); mesoscutum with microsculpture effaced posteriorly (Figure 91); median lobe of mesoscutum with oblique rugulae (Figure 91); mesoscutellum entirely smooth (Figure 91) Trissolcus kozlovi Rjachovskij, p. 79
Frons above antennal scrobe with transverse rugae (Figures 37–38) or rugae absent (Figures 84, 135–136, 221) but not irregular lines; mesoscutum sometimes with sculpture effaced posteriorly; rugulae on mesoscutum, if present, oriented longitudinally between notauli; mesoscutellum smooth or with microsculpture 11
11 Frons with parallel arched rugae within and above antennal scrobe, rugae often extending onto lateral frons (Figures 37–38); genal carina usually present (Figures 37–38); mesoscutum often with longitudinal rugae posteriorly; ventral frons with facial striae usually pronounced (Figure 37); clypeus with 4 setae, lateral clypeal setae about half as long as medial setae (Figure 39) Trissolcus corai Talamas, sp. n., p. 41
Frons sometimes with transverse striation directly above antennal scrobe, but not extending to median ocellus or extending onto lateral frons; genal carina absent (Figure 88); mesoscutum without rugae posteriorly; ventral frons without facial striae (Figures 84–86); clypeus with 2 or 4 setae, clypeal setae approximately equal in length (Figure 85) 12
12 Clypeus with 2 setae (Figure 221) Trissolcus vindicius (Nixon), p. 160
Clypeus with 4 setae (Figures 85, 135) 13
13 Frons directly below median ocellus with broad smooth area and without a dorsoventral furrow (Figures 135–136) Trissolcus plautiae (Watanabe), p. 108
Frons directly below median ocellus with microsculpture and usually a dorsoventral furrow (Figures 84–87) Trissolcus japonicus (Ashmead), p. 74
14 Eyes densely setose (Figure 63–66); mesopleuron without subacropleural sulcus (Figure 64) Trissolcus exerrandus Kozlov & Lê, p. 71
Eyes bare or with short scattered setae (Figures 69–70, 42–45); mesopleuron with subacropleural sulcus indicated by line of foveae (Figures 42, 71) 15
15 Frons above antennal scrobe with coarse arcuate rugae (Figures 44–45) Trissolcus cultratus (Mayr), p. 45
Frons between antennal scrobe and median ocellus without coarse rugae (Figures 69–70, 73, 116) 16
16 Mesoscutum with notaulus well developed, at least 1/3 length of mesoscutum (Figure 68); frons with shallow rugulae forming a circle directly above antennal scrobe (Figures 69–70); mesopleuron with episternal foveae forming a continuous line between mesopleural pit and postacetabular sulcus (Figure 67) Trissolcus flavipes (Thomson), p. 63
Mesoscutum with notaulus abbreviated, present only at posterior margin of mesoscutum (Figures 72, 113, 115); frons above antennal scrobe with faint transverse striation (Figure 116) or with microsculpture alone (Figure 73); mesopleuron with episternal foveae distant from postacetabular sulcus (Figures 74, 114) 17
17 Mesoscutellum with microsculpture, otherwise smooth (Figure 72); setal bases on mesoscutellum simple; mandible broad (Figures 74, 76); ventral frons with facial striae pronounced (Figures 73, 75–76) Trissolcus gonopsidis (Watanabe), p. 66
Mesoscutellum rugose, medial rugae oriented longitudinally (Figure 115); setal bases on mesoscutellum distinctly pustulate (Figures 115); mandible slender (Figure 117); ventral frons without facial striae (Figures 116–117) Trissolcus nycteridaner Talamas, sp. n. , p. 96
18 Metapleuron with posteroventral portion densely setose (Figures 94, 103, 154, 189, 213), rarely sparsely setose only in ventral portion 19
Metapleuron without setae below metapleural sulcus (Figures 20, 2728, 56–58, 78, 99, 110) 24 (basalis group)
19 Mesopleural carina with row of foveae along its dorsal margin (Figure 189) Trissolcus tersus Lê, p. 142
Mesopleural carina without foveae along its dorsal margin (Figures 154, 209), or mesopleural carina absent (Figure 94) 20
20 Mesoscutum without notauli (Figures 95–96, 104); mesopleuron with a single episternal fovea (Figures 103, 105) or foveae entirely absent (Figure 94); T1 without longitudinal striae posterior to basal costae (Figures 95, 104) 21
Mesoscutum with notauli (Figures 155–156, 164–167, 210, 214); mesopleuron with episternal foveae extending from postacetabular sulcus to mesopleural pit (Figures 154, 209, 213); T1 with longitudinal striae posterior to basal costae (Figures 155, 210, 214) 22 (thyantae group)
21 T2 longitudinally striate anteromedially (Figures 95–96); mesopleuron without episternal foveae (Figure 94); mesoscutum longitudinally striate posteriorly (Figure 96); crenulate furrow present along anterior margin of metascutellum (Figures 95–96) Trissolcus larides Nixon, p. 83
T2 entirely smooth (Figure 104); mesopleuron with one episternal fovea (Figures 103, 105); mesoscutum areolate rugulose throughout (Figure 104); line of deep punctures present along anterior margin of metascutellum (Figure 104) Trissolcus levicaudus Talamas sp. n., p. 89
22 Gena distinctly broad and bulging in lateral view (Figure 213); compound eye with posterior and dorsal margins separated from hyperoccipital carina by a broad furrow (Figure 213); mesoscutum with median mesoscutal carina (Figure 215) Trissolcus viktorovi Kozlov, p. 157
Gena narrow to moderately broad in lateral view (Figures 154, 209); compound eye with posterior and dorsal margins narrowly separated from hyperoccipital carina (Figures 156–157, 212); mesoscutum with or without median mesoscutal carina 23
23 Lateral mesoscutum with mesoscutal suprahumeral sulcus and mesoscutal humeral sulcus comprised of conspicuous cells (Figure 156); posterior vertex without medial depression (Figures 155–156, 164–167); occipital carina smoothly arched medially (Fig. 164); metapleuron below metapleural sulcus with setation ventrally and posteriorly (Figure 154) Trissolcus scutellaris (Thomson), p. 120
Lateral mesoscutum with mesoscutal suprahumeral sulcus and mesoscutal humeral sulcus comprised of smooth furrows or very small and poorly defined cells (Figures 210, 212); posterior vertex depressed medially (Figure 211); occipital carina prominent and flat medially; metapleuron below metapleural sulcus with sparse setation ventrally, without setation posteriorly (Figure 209) Trissolcus vesta Kozlov & Lê, p. 154
24 Antenna with 4 clavomeres (A8A11 with basiconic sensilla) (Figures 80, 82) 25
Antenna with 5 clavomeres (A7A11 with basiconic sensilla) (Figures 6, 141–142) 26
25 Fore wing with length of postmarginal vein equal to or less than length of stigmal vein (Figures 124–127) Trissolcus oobius (Kozlov), p. 100
Fore wing with length of postmarginal vein at least twice length of stigmal vein (Figure 79) Trissolcus hyalinipennis Rajmohana & Narendran, p. 70
26 Antennae with distinctly large clava (Figure 107, 109); orbital furrow expanded at intersection with malar sulcus (Figures 109–111); laterotergite 1 with a line of setae along dorsal margin (Figure 110) Trissolcus mitsukurii (Ashmead), p. 92
Antenna with clava moderately large to slender (Figures 16, 30, 132, 178–183); orbital furrow uniform in width (Figures 30, 54, 99, 105); laterotergite 1 with or without setae 27
27 Mesopleuron without episternal foveae (Figures 16, 132, 140); metapleuron with anteroventral extension short, not reaching ventral mesopleural carina (Figures 118, 128) 28
Mesopleuron with at least one episternal fovea (Figures 105, 114, 171) indicated or metapleuron with anteroventral extension extending to ventral mesopleural carina (Figures 57–58, 99, 171) 30
28 Mesopleural carina extending from mesopleural pit to ventral mesopleural carina (Figure 15); mesoscutellum rugose throughout Trissolcus atys (Nixon), p. 26
Mesopleural carina incomplete, at most extending half the distance from the mesopleural pit to ventral mesopleural carina; mesoscutellum smooth or with shallow microsculpture 29
29 Ventral mesopleuron distinctly bulging (Figure 132); mesocoxa oriented parallel to long axis of body (Figure 128); dorsal frons with sculpture effaced, sometimes entirely smooth and shining (Figures 131–132); A7 with two basiconic sensilla; metasoma dark brown to black (Figures 128, 130) Trissolcus perepolovi (Kozlov), p. 105
Ventral mesopleuron not distinctly bulging (Figure 140); mesocoxa oriented at an angle of ~45º relative to long axis of body; dorsal frons evenly and densely covered in microsculpture (Figure 144); A7 with one basiconic sensillum (Figures 141–142); metasoma dark brown to bright yellow (137–139) Trissolcus rufiventris (Mayr), p. 111
30 Metapleuron with paracoxal sulcus and metapleural epicoxal sulcus indicated by distinct foveae (Figures 52, 56–58) Trissolcus elasmuchae (Watanabe), p. 53
Metapleuron with paracoxal sulcus or metapleural epicoxal sulcus poorly defined or not indicated externally (Figures 20, 99, 171, 225) 31
31 Mesoscutellum smooth or with coriaecious microsculpture (Figures 20–21, 168–169) 32
Mesoscutellum coarsely rugose (Figures 29, 34, 98, 100, 226) 33
32 Lateral pronotum with netrion sulcus complete dorsally, netrion distinct (Figures 170–171); lateral mesoscutum with mesoscutal humeral sulcus usually comprised of cells (Figure 169), occasionally present as a smooth furrow Trissolcus semistriatus (Mayr), p. 129
Lateral pronotum with netrion sulcus incomplete dorsally, netrion often poorly defined (Figure 22); lateral mesoscutum with mesoscutal humeral sulcus present as a smooth furrow (Figures 20–21) Trissolcus basalis (Wollaston), p. 30
33 Mesopleuron with femoral depression coarsely rugose throughout (Figure 99); radicle yellow (Figures 99, 102) Trissolcus latisulcus (Crawford), p. 86
Mesopleuron with femoral depression smooth or with arched rugae ventrally (Figures 28, 225); radicle dark brown to black (Figures 30–32, 222–224) 34
34 Frons directly below anterior ocellus with one or more dorsoventral rugae, sculpture effaced lateral of rugae (Figures 222, 224) Trissolcus yamagishii Ryu, p. 164
Frons with arched rugae around median ocellus or with microsculpture alone (Figures 31–33) Trissolcus comperei (Crawford), p. 37

Species descriptions

Trissolcus atys (Nixon)

Figures 13–16; Morphbank 1

Telenomus atys Nixon, 1935: 76, 86, 96 (original description, keyed); Risbec, 1950: 558, 560 (description, keyed).

Trissolcus atys (Nixon): Masner, 1965: 124 (type information, generic transfer); Johnson, 1992: 623 (cataloged, type information).

Description

Female body length: 0.88–0.71 mm (n=3). Male body length: 0.74 mm (n=1). Body color: head and mesosoma black, metasoma brown.

Figures 13–16. 

T. atys, female (USNMENT00896187) 13 female (USNMENT00896187) head, mesosoma, metasoma, dorsal view 14 female (USNMENT00896187) head, anterior view 15 female holotype (B.M. TYPE HYM. 9.245a), head and mesosoma, lateral view 16 female (USNMENT00896187) head and mesosoma, anterolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: present. Sculpture of malar sulcus: faintly rugulose. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: transversely strigose ventrally, absent dorsally. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: areolate. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: transversely strigose; with granular microsculpture. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: irregularly rugulose. Postacetabular sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by smooth furrow with a small number of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: areolate, longitudinally striate along posterior margin. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: pointed. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, trochanters and proximal femora pale brown, elsewhere yellow.

Metasoma. Longitudinal striae on T1 posterior to basal costae: absent. Number of sublateral setae (on one side): 2. Setation of laterotergite 1: absent. Longitudinal striation of T2: faintly present anteriorly. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present.

Diagnosis

Trissolcus atys can be separated from other Palearctic species in the basalis group by the absence of episternal foveae (Figure 16), a rugose mesoscutellum (Figures 13, 15), fine transverse striation on the frons (Figure 14) and a mesopleural carina that extends from the mesopleural pit to the ventral mesopleural carina (Figure 15).

Material examined

Holotype, female, T. atys: UGANDA: Kampala, XI-1930, reared from egg, H. Hargreaves, B.M. TYPE HYM. 9.245a (deposited in BMNH). Other material: (2 females, 1 male) KENYA: 1 female, 1 male, USNMENT01109872–01109873 (USNM). UNITED ARAB EMIRATES: 1 female, USNMENT00896187 (CNCI).

Trissolcus barrowi (Dodd) nomen dubium

Figures 17–19; Morphbank 2

Telenomus barrowi Dodd, 1920: 356 (original description).

Microphanurus barrowi (Dodd): Nixon, 1938: 124, 137 (description, generic transfer, keyed); Nixon, 1943: 138 (keyed).

Microphanurus borrowi (Dodd): Risbec, 1950: 569 (keyed, error).

Trissolcus barrowi (Dodd): Masner, 1965: 124 (type information, generic transfer); Johnson, 1992: 623 (cataloged, type information); Rajmohana K. & Narendran, 2007: 102 (keyed).

Description

Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Figures 17–19. 

T. barrowi, female holotype (B.M. TYPE HYM. 9.323) 17 metasoma, lateral view 18 metasoma, dorsal view 19 T1–T2, dorsolateral view. Scale bars in millimeters.

Material examined

Holotype, female, T. barrowi: INDIA: Himachal Pradesh St., Dalhousie, 11.VIII.1906, reared, H. J. W. Barrow, B.M. TYPE HYM. 9.323 (deposited in BMNH).

Comments

The holotype specimen consists of a metasoma and a wing, which are insufficient to separate this species from other species of Trissolcus, nor can it be confidently distinguished with the descriptions by Dodd (1920) and Nixon (1938). The condition of the type is so poor that designation of a neotype is warranted. However, this is confounded by the host association on the holotype labels that indicate that it emerged from the egg of a hawk moth (Sphingidae)- which is most likely an error. In the absence of material from vicinity of the type locality required for designation of a neotype, we presently treat T. barrowi as nomen dubium.

Trissolcus basalis (Wollaston)

Figures 20–23; Morphbank 3

Trissolcus africanus (Fouts) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3175; Morphbank4; Lectotype designation. We here designate syntype specimen MCSN 0015 (deposited in MCSN) as the lectotype of Trissolcus africanus (Fouts).

Trissolcus lodosi (Szabó) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3259; Morphbank5

Trissolcus sulmo (Nixon) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3315; Morphbank6; Lectotype designation. We here designate syntype specimen B.M. TYPE HYM. 9.318 (deposited in BMNH) as the lectotype of Trissolcus sulmo (Nixon).

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)); Talamas, Johnson & Buffington, 2015: 64 (type information).

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 africanus Fouts, 1934: 105, 106 (original description, keyed).

Microphanurus basalis (Wollaston): Nixon, 1935: 96, 100 (description, generic transfer, synonymy, keyed); Nixon, 1943: 138 (keyed); Risbec, 1950: 570, 571 (variation, keyed).

Microphanurus sulmo Nixon, 1938: 123, 126 (original description, keyed); Nixon, 1943: 138 (keyed); Risbec, 1950: 569 (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 & Lê, 1977: 516 (keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 121 (description); Graham, 1984: 100 (variation); Johnson, 1985b: 432, 434 (description, keyed); Johnson, 1991: 212, 213, 214, 216 (diagnosis, keyed); Ghahari, Buhl & Kocak, 2011: 594 (listed); Mao, Valerio, Austin, Dowton & Johnson, 2012: 194 (presentation of mitochondrial genome, phylogenetic position); Fusu, Bin & Popovici, 2013: 263 (description of chromosomes); Kononova, 2014: 1425 (keyed); Talamas, Johnson & Buffington, 2015: 60, 61 (diagnosis, keyed, lectotype designation); Kononova, 2015: 263 (keyed).

Trissolcus maderensis (Wollaston): Masner, 1965: 126 (type information, generic transfer).

Trissolcus piceipes (Dodd): Masner, 1965: 127 (type information, generic transfer).

Trissolcus sulmo (Nixon) syn. n.: Masner, 1965: 128 (type information, generic transfer); Johnson, 1992: 638 (cataloged, type information).

Asolcus sulmo (Nixon): Voegelé, 1969: 151 (keyed).

Trissolcus megacephalus (Ashmead): Johnson, 1983: 448 (type information).

Trissolcus africanus (Fouts) syn. n.: Bin, 1974: 463 (generic transfer, type information); Masner, 1976: 76 (systematic position).

Asolcus lodosi Szabó, 1981: 197 (original description).

Trissolcus lodosi (Szabó) syn. n.: Kononova, 2014: 1425 (keyed); Kononova, 2015: 262 (keyed).

Description

Female body length: 1.07–1.28 mm (n=20). Male body length: 0.91–1.24 mm (n=11). Body color: head, mesosoma, and metasoma black.

Figures 20–23. 

T. basalis, female lectotype (B.M. TYPE HYM. 9.304) 20 head, mesosoma, metasoma, lateral view 21 head, mesosoma, metasoma, dorsal view 22 head and mesosoma, anterolateral view 23 head, anterior view. Scale bars in millimeters.

Head. Color of radicle: black; brown; dark brown. Length of radicle: equal to or greater than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally. Color of A7A11 in female: dark brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: bulging. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: smooth. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent; weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: faintly striate. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 2; 1. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: absent; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by small punctures; present as a smooth furrow. Mesopleural epicoxal sulcus: present as a smooth furrow. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: mostly smooth with faint rugulae radiating from metapleural pit. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: multiple rows of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present.

Diagnosis

Among Palearctic species of Trissolcus, T. basalis is most similar to T. semistriatus with which it shares episternal foveae that are distant from both the postacetabular sulcus and the mesopleural pit and an absence of coarse macrosculpture on the mesoscutum and mesoscutellum. The best feature for separating these species is the netrion sulcus, which is dorsally complete in T. semistriatus (Figures 170, 171) and dorsally undefined in T. basalis (Figure 22). The postacetabular sulcus in T. basalis is typically indicated by small and poorly defined cells, sometimes appearing as a smooth furrow. In T. semistriatus the cells of the postacetabular sulcus are distinct and regularly spaced. The mesoscutal humeral sulcus in T. basalis is always present as a smooth furrow whereas in T. semistriatus it is nearly always comprised of distinct cells. Trissolcus basalis is also similar to T. elasmuchae, from which it can be separated by the absence of a well-defined paracoxal sulcus in the ventral half of the metapleuron, and by the form of the episternal foveae, which in T. elasmuchae from a more or less continuous line from the postacetabular sulcus to the mesopleural pit.

Material examined

Lectotype, female, T. basalis: PORTUGAL: Madeira Reg. Autó., Madeira Island, VII-1855, Wollaston, B.M. TYPE HYM. 9.304 (deposited in BMNH). Lectotype, female, Microphanurus africanus: SOMALIA: Shabeellaha Dhexe Reg., Giohar (Villaggio Duca degli Abruzzi), 1.III.1926, Paoli, MCSN 0015 (deposited in MCSN). Holotype, female, T. megacephalus: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, no date, H. H. Smith, USNM Type No. 2525 (deposited in USNM). Lectotype, female, M. sulmo: SRI LANKA: Central Prov., Nuwara Eliya Dist., Talawakele (Talawakelle), 1932, C. B. R. King, B.M. TYPE HYM. 9.318 (deposited in BMNH). Paratype, female, Asolcus lodosi : TURKEY: 1 female, Hym.Typ.No. 1886, Mus.Budapest (HNHM). Other material: (189 females, 53 males, 54 unknowns) AUSTRALIA: 22 females, 4 males, USNMENT0087208800872090, 00903007, 0095454200954563 (USNM). BERMUDA: 10 unsexed, USNMENT0098942200989423, 00989425, 0098942700989431, 0098945700989458 (USNM). CUBA: 6 unsexed, USNMENT00989407–00989412 (USNM). CYPRUS: 3 females, 1 male, USNMENT00916553, 00916555, 00916580, 00916582 (BMNH). DOMINICA: 1 unsexed, USNMENT00989459 (USNM). DOMINICAN REPUBLIC: 9 unsexed, USNMENT00989413–00989421 (USNM). EGYPT: 2 females, 2 unsexed, USNMENT00872006–00872009 (USNM). FRANCE: 9 females, USNMENT0089605500896060, 0089607000896071, 00896296 (CNCI). HUNGARY: 1 male, USNMENT00916996 (BMNH). ISRAEL: 1 female, UCRC ENT 296979 (UCRC). ITALY: 1 female, USNMENT00916293 (BMNH). JORDAN: 4 females, USNMENT00916495–00916498 (BMNH). MALAYSIA: 1 female, 1 male, USNMENT00916371, 00916372 (BMNH). MONTENEGRO: 1 female, USNMENT00896249 (CNCI). MONTSERRAT: 16 females, 2 males, USNMENT00954513–00954530 (USNM). MOROCCO: 1 female, 1 male, USNMENT00896088, 00896109 (CNCI). PORTUGAL: 21 females, 6 males, USNMENT00916186, 00916188, 0091619600916200, 00916204, 0091620700916209, 00916214, 00916219, 00916221, 0091622400916225, 0091622800916233, 00916239, 0091624200916245 (BMNH). SAINT VINCENT AND THE GRENADINES: 13 unsexed, USNMENT0098943200989434, 0098944000989449 (USNM). SPAIN: 2 males, USNMENT00916183–00916184 (BMNH). TRINIDAD AND TOBAGO: 10 unsexed, USNMENT0076495000764951, 0098945000989456, 00989461 (USNM). TURKEY: 2 females, 1 male, OSUC 17739, 17742, USNMENT00916024 (BMNH). UNITED STATES: 105 females, 35 males, 2 unsexed, OSUC 131149131186, 154353, 7339 (OSUC); USNMENT0087210300872109, 0095402200954023, 00954393, 0095443600954437, 0095444300954452, 0095445400954469, 0095448000954512, 0095453100954541, 0095456400954570, 0110906501109069, 0110907101109074, 01109077, 01109079, 0110908801109089 (USNM). VENEZUELA: 1 unsexed, USNMENT00989460 (USNM).

Trissolcus carinifrons (Cameron)

Figures 24–27; Morphbank 7

Immsia carinifrons Cameron, 1912: 105 (original description); Kieffer, 1926: 393 (description).

Telenomus carinifrons (Cameron): Dodd, 1920: 355 (description, generic transfer).

Microphanurus carinifrons (Cameron): Nixon, 1938: 124, 138 (description, generic transfer, keyed); Nixon, 1943: 137 (keyed); Risbec, 1950: 569 (keyed).

Trissolcus carinifrons (Cameron): Masner, 1965: 125 (type information, generic transfer); Mani & Sharma, 1982: 143 (description); Johnson, 1992: 624 (cataloged, type information); Rajmohana K. & Narendran, 2007: 102 (keyed).

Description

Female body length: 1.82 mm (n=1). Body color: head, mesosoma, and metasoma black.

Figures 24–27. 

T. carinifrons, female holotype (B.M. TYPE HYM. 9.312) 24 head, mesosoma, metasoma, dorsolateral view 25 head, mesosoma, metasoma, ventrolateral view 26 head, anterior view 27 mesosoma, lateral view. Scale bars in millimeters.

Head. Color of radicle: dark brown. Length of radicle: equal to or greater than width of clypeus. Color of A1A6 in female: yellow. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: bulging. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: coarsely rugose. Preocellar pit: present. Setation of lateral frons: moderately dense. Sculpture directly ventral to preocellar pit: absent. Macrosculpture of lateral frons: rugose. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: rugulose. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 3. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: concentrically strigose. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: coarsely rugose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as clearly defined line of cells. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: coarsely rugose, lines oriented more longitudinally along posterior margin. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: absent. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: two rows of deep, well-defined cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae and fifth tarsomeres brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in most of tergite, extending posteriorly to transverse line of setae. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Trissolcus carinifrons can be separated from all the species treated here by the combination of 6 setae on the clypeus, the presence of a complete hyperoccipital carina, and a mesepimeral sulcus comprised of circular foveae.

Material examined

Holotype, female, I. carinifrons: INDIA: Uttarakhand St., Forest Research Institute (FRI / IFRI), Dehra Dun, 3.VIII.1910, at light, V. S. Iyer, B.M. TYPE HYM. 9.312 (deposited in BMNH).

Trissolcus comperei (Crawford)

Figures 28–29, 30–34; Morphbank 8

Trissolcus itoi Ryu syn. n.; http://bioguid.osu.edu/xbiod_concepts/3248; Morphbank9

Telenomus comperei Crawford, 1912: 1 (original description); Baltazar, 1966: 172 (cataloged, type information, distribution).

Trissolcus comperei (Crawford): Masner & Muesebeck, 1968: 72 (type information, generic transfer); Johnson, 1992: 625 (cataloged, type information).

Trissolcus itoi Ryu syn. n., 1984: 37, 52 (original description, keyed).

Description

0.93–1.51 mm (n=19). Body color: head, mesosoma, and metasoma black.

Figures 28–29. 

T. comperei 28 female (USNMENT00872396), head, mesosoma, metasoma, lateral view 29 female (USNMENT00872397), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 30–34. 

T. comperei 30 female holotype of T. itoi syn. n. (Type No. 2220 Kyushu Univ.) 31 female holotype (USNMENT00989064), head, anterolateral view 32 female (USNMENT00872399), head, anterior view 33 female (USNMENT00916352), head, anterior view 34 female (USNMENT00872397), mesosoma, T1–T2, posterodorsal view. Scale bars in millimeters.

Head. Color of radicle: black; dark brown. Length of radicle: equal to or greater than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: pale brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; strigose, roughly concentric around median ocellus. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: horizontally striate, striae of antennal scrobe extending to lateral frons; absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: rugulose; absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: antero-posteriorly striate; finely rugulose. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2; 1. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose; with granular microsculpture. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: concentrically strigose. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: irregularly rugulose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly; rugose. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: indicated by smooth furrow with a small number of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose; rugulose, becoming longitudinally striate posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent; indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: multiple rows of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, femora and tibiaie yellow to pale brown, elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in most of tergite, extending posteriorly to transverse line of setae. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present.

Diagnosis

Trissolcus comperei is similar to T. yamagishii, T. latisulcus, and T. carinifrons with which it shares a long dark radicle and coarse sculpture of the mesoscutum and mesoscutellum. Trissolcus comperei can be separated from them by the characters presented in the key: sculpture of the frons below the median ocellus, absence of a hyperoccipital carina, and a femoral depression without coarse rugae throughout. The rugae that are present in the femoral depression are located ventrally and are arched and parallel. This is a very useful diagnostic character for the species, although the rugae are fainter in smaller specimens. Trissolcus aloysiisabaudiae from East Africa shares with T. comperei this sculptural pattern of the femoral depression and is a very similar species, excluding the sculpture of the mesonotum and the presence of a robust longitudinal mesoscutellar carina. These two species should certainly be compared in a greater context of African species.

Comments

Specimens from China have bright yellow legs and non-claval flagellomeres. Specimens from Japan have darker legs and flagellomeres. The pattern of macrosculpture on the frons and the coloration of the legs and antennae are essentially identical to the pattern found in T. semistriatus: specimens from higher latitudes tend to be darker with more robust facial sculpture.

Material examined

Holotype, female, T. comperei: CHINA: Guangdong Prov., Guangzhou (Canton), no date, reared from egg, G. Compere, USNMENT00989064 (deposited in USNM). Holotype, female, T. itoi: JAPAN: Niigata Pref., Mt. Kanegura Yama, 19.VIII.1970, K. Yamagishi, Type No. 2220 Kyushu Univ. (deposited in KUEC). Other material: (16 females) CHINA: 5 females, USNMENT00916347, 0091634900916352 (BMNH). INDIA: 1 female, USNMENT00916361 (BMNH). JAPAN: 8 females, OSUC 144487144488, 542357, 542365, 542371, 542424 (CNCI); OSUC 75840–75841 (OSUC). SOUTH KOREA: 1 female, USNMENT00896147 (CNCI). UNITED ARAB EMIRATES: 1 female, USNMENT00896183 (CNCI).

Trissolcus corai Talamas, sp. n.

Figures 35–36, 37–39; Morphbank 10

Description

Female body length: 1.11–1.41 mm (n=20). Male body length: 1.10–1.23 mm (n=2). Body color: head, mesosoma, and metasoma black.

Figures 35–36. 

Trissolcus corai female paratype (USNMENT00977542) 35 head, mesosoma, metasoma, lateral view 36 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 37–39. 

Trissolcus corai 37 female paratype (OSUC 542370), head, mesosoma, metasoma, ventrolateral view 38 female paratype (USNMENT00896028), head, mesosoma, metasoma, ventrolateral view 39 female paratype (USNMENT01223664), ventral head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow; yellow, becoming brown distally. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate; faintly rugulose. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: moderately dense. Sculpture directly ventral to preocellar pit: absent; microsculptured; weakly transversely striate. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present; absent medially, present laterally. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly; rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; weakly rugulose posteriorly; rugulose, becoming longitudinally striate posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Trissolcus corai belongs to the cluster of species of the flavipes group with 4 clypeal setae, which includes T. japonicus, T. kozlovi, and T. plautiae. It can be separated from all of these by the transverse sculpture on the frons and by the distinctly smaller size of the lateral clypeal setae relative to the median pair. The number of clypeal setae is the best character for separating T. corai from T. vindicius, which can be very similar, particularly because the rugae on the frons of some T. corai may be weakly developed. In most T. corai, the mesoscutum between the notauli has longitudinal rugae, which will separate T. corai from T. japonicus, T. kozlovi, T. plautiae, and T. vindicius.

Etymology

This species is named for Joe Cora, former database manager at The Ohio State University, for his multitudinous contributions to the development of cybertaxonomic tools and the bioinformatics of Platygastroidea.

Associations

emerged from egg of Cappaea tibialis Hsiano & Cheng: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female: CHINA: Beijing Prov., Haidian, Baiwangshan Mountain N40º02’05.31 E116º15’21.86, 15.XII.2015, reared from egg, T. Haye, USNMENT01223979 (deposited in USNM). Paratypes: (94 females) CHINA: 8 females, USNMENT0122397301223974, 0122397701223978 (OSUC); USNMENT01223965, 0122397001223972 (USNM). JAPAN: 79 females, OSUC 144392144408, 144411144435, 144437, 144439144444, 144446144454, 144456144467, 542356, 542366542367, 542369542370, 542372, 542379, 542381, USNMENT00896310 (CNCI). SOUTH KOREA: 5 females, USNMENT0089602700896028, 00896030, 00896037, 00896047 (CNCI). TAIWAN: 2 females, OSUC 63888, 76838 (OSUC). Other material: (2 females, 4 males) CHINA: 2 females, 2 males, USNMENT01223664, 01223966, 01223969, 01223976 (USNM). JAPAN: 2 males, OSUC 144409144410 (CNCI).

Trissolcus cultratus (Mayr)

Figures 40–41, 42–45; Morphbank 11

Trissolcus striatellus Kononova syn. n.; http://bioguid.osu.edu/xbiod_concepts/354844

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); Rjachovskij, 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); Talamas, Johnson & Buffington, 2015: 54, 71 (removed from synonymy with Trissolcus flavipes (Thomson), diagnosis, keyed, type information).

Trissolcus striatellus Kononova syn. n., 2014: 744 (original description, diagnosis).

Description

Female body length: 1.19–1.98 mm (n=20). Male body length: 1.03–1.68 mm (n=5). Body color: head, mesosoma, and metasoma black.

Figures 40–41. 

Trissolcus cultratus 40 female lectotype (NHMW 0008A), head, mesosoma, metasoma, lateral view 41 female (USNMENT00916251), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 42–45. 

Trissolcus cultratus 42 female (USNMENT00916251), head and mesosoma, ventrolateral view 43 female (USNMENT00764849), head and mesosoma, dorsolateral view 44 female lectotype (NHMW 0008A), head, anterolateral view 45 female (USNMENT00916251), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: dark brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye; present only at base of mandible. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Macrosculpture of frons between antennal scrobe and anterior ocellus: parallel arcuate rugae. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: sparse. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: obliquely striate, striae of upper frons extending ventrolaterally. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma: Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: irregularly rugulose; absent. Postacetabular sulcus: formed by large cells; formed by small punctures. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly. Posterodorsal metapleural sulcus: present as line of foveae; poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent; present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent; present. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose.

Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae dark brown to black, femora and tibia yellow to dark brown, trochanters and tarsi yellow to pale brown. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in most of tergite, extending posteriorly to transverse line of setae. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

The arched rugae on the frons of Trissolcus cultratus separate it from most species of the flavipes group, but it is shared with T. exerrandus and T. corai. Trissolcus exerrandus has densely setose eyes whereas those of T. cultratus are essentially bare. The orbital furrow in T. corai is well defined ventrally and that of T. cultratus is not (compare Figures 37, 38 to Figures 44, 45).

Material examined

Lectotype, female, T. cultratus: Palearctic region: no date, G. Mayr, NHMW 0008A (deposited in NHMW). Paralectotype: Palearctic region: 1 male, NHMW 0008B (NHMW). Other material: (198 females, 26 males, 4 unsexed) AUSTRIA: 5 females, 2 unsexed, USNMENT00979612–00979613 (CUIC); OSUC 75765–75767 (OSUC); USNMENT0067594300675944 (USNM). BELGIUM: 1 female, USNMENT00896165 (CNCI). CHINA: 8 females, UCRC ENT 142635, 143817 (UCRC); USNMENT0076494500764947, 0076496200764963, 00764983 (USNM). CZECH REPUBLIC: 1 female, 3 males, USNMENT0089631100896314 (CNCI). FRANCE: 11 females, 1 male, USNMENT00916035, 0091603700916038, 00916041, 00916083, 00916125, 00916131, 00916147 (BMNH); OSUC 75753–75756 (OSUC). GERMANY: 1 female, USNMENT00872117 (USNM). HUNGARY: 5 females, 1 unsexed, USNMENT00896072, 00896073 (CNCI); OSUC 75771–75773, 75783 (OSUC). JAPAN: 32 females, 6 males, OSUC 144472144480, 542363, 542374, 542412, 542415, USNMENT00896136, 00896138, 00896140, 00896305, 0089630700896309, 00896315, 00896339, 00896341 (CNCI); OSUC 75784, 75786–75788 (OSUC); UCRC ENT 297012 (UCRC); USNMENT0067573000675737, 00675761, 00764849 (USNM). POLAND: 6 females, USNMENT0091660400916609 (BMNH). RUSSIA: 34 females, USNMENT0089604800896054, 0089607400896075, 0097928200979286, 00979289 (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, 0089601500896016, 00896018, 00896019, 00896029, 00896032, 0089604400896046, 0089611200896116, 0089611800896119, 0089612100896122, 0089613400896135, 00896157, 00979237, 0097924600979250, 00979253, 00979280 (CNCI). SWEDEN: 10 females, 4 males, USNMENT0091610500916110, 0091611200916114, 00916310, 0091631500916317, 00916319 (BMNH). SWITZERLAND: 26 females, 2 males, USNMENT00979222–00979226 (CNCI); USNMENT00916973–00916989, 00954000– 00954005 (USNM). TAIWAN: 2 females, OSUC 542386 (CNCI); UCRC ENT 112210 (UCRC). UNITED KINGDOM: 21 females, 6 males, 1 unsexed, USNMENT0091624700916254, 0091639300916408, 0091641100916412, 00916416, 00916418 (BMNH).

Trissolcus edessae Fouts

Figures 46–47, 48–51; Morphbank 12

Trissolcus edessae Fouts, 1920: 65 (original description); Masner & Muesebeck, 1968: 72 (type information); Johnson, 1984a: 799, 801 (description, keyed); Johnson, 1987: 289, 300 (diagnosis, keyed); Johnson, 1992: 626 (cataloged, type information); Talamas, Johnson & Buffington, 2015: 56, 74 (diagnosis, keyed, type information).

Description

Female body length: 1.27–1.67 mm (n=31). Male body length: 1.18–1.70 mm (n=8). Body color: head, mesosoma, and metasoma black.

Figures 46–47. 

Trissolcus edessae, female (USNMENT00916178) 46 head, mesosoma, metasoma, lateral view 47 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 48–51. 

Trissolcus edessae, female (USNMENT00916178) 48 head and mesosoma, anterolateral view 49 head, anterior view 50 head and mesosoma, ventrolateral view 51 mesosoma, dorsal view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: faintly rugulose. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: transversely strigose ventrally, absent dorsally. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: sparse. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: rugose. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: weakly differentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: dorsoventrally strigose in dorsal half. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth area small because cells of surrounding sulci are large. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: present. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: absent. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally, absent medially. Setation of S2: present posteromedially.

Diagnosis

Trissolcus edessae is unique among the species of the flavipes group treated here in that it has a medial mesoscutal carina. It bears no particular affinity to other Palearctic species in the flavipes group, but is morphologically similar to two species in the Nearctic: T. brochymenae and T. euschisti, from which it may be separated by the distinctly bicolored antennae, with A1A6 yellow, and A7A11 brown (Figures 46, 47). The labrum of T. edessae has a pair of distinct pits that are easily visible with light microscopy (Figure 50), whereas these pits in other species typically require scanning electron microscopy for observation. This character has yet to be fully explored throughout the genus because the labrum is often occluded, but we suspect that it will be informative.

Material examined

Holotype, female: UNITED STATES: LA, Orleans Parish, New Orleans, 23.VII.1919, C. E. Smith, USNMENT00872412 (deposited in USNM). Other material: SPAIN: 1 female, USNMENT00916178 (BMNH). (37 females, 3 males, 6 unsexed) EL SALVADOR: 5 unsexed, USNMENT00989220–00989221 (CNCI); USNMENT0076498000764981, 00764993 (USNM). UNITED STATES: 36 females, 3 males, USNMENT0095434600954377, 0095438400954390 (USNM).

Comments

The specimen illustrated in Figures 4651 is the only known representative of T. edessae outside of the Nearctic. We compared this specimen directly to the type of T. edessae and found nothing to suggest that it is a different species. The collecting locality of Valencia, Spain, is noteworthy given the large volume of shipping traffic that passes through this port city and we consider it likely that T. edessae was delivered to Valencia on cargo. Determination of whether or not it became established in Europe will require focused sampling, as even in the Nearctic it is not a very common species. This phenonenon of species spreading around the world, as is the case with T. edessae and T. japonicus (Talamas et al. 2015), emphasizes the importance of a cosmopolitan perspective when conducting alpha taxonomy.

Trissolcus elasmuchae (Watanabe)

Figures 52–53, 54–58; Morphbank 13

Trissolcus davatchii (Javahery) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3210; Morphbank14

Trissolcus monirus Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3268; Morphbank15

Asolcus elasmuchae Watanabe, 1954: 21, 22 (original description).

Asolcus davatchii Javahery, 1968: 419, 422 (original description, keyed).

Trissolcus polarica Rjachovskij, 1972: 74 (original description, synonymized by Kononova (1974)); Kononova, 1974: 72 (junior synonym of Trissolcus elasmuchae (Watanabe)).

Trissolcus elasmuchae (Watanabe): Kononova, 1974: 72 (diagnosis, synonymy); Kozlov & Lê, 1977: 510 (keyed); Kozlov, 1978: 636 (description); Kozlov & Kononova, 1983: 109 (description); Ryu & Hirashima, 1984: 37, 55 (description, keyed); Kononova, 1995: 96 (keyed); Petrov, 2013: 326 (keyed).

Trissolcus davatchii (Javahery) syn. n.: Kozlov & Lê, 1977: 516 (keyed, generic transfer); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 120 (description); Fergusson, 1984: 230 (type information).

Trissolcus monirus Lê syn. n., 1985: 165 (original description); Johnson, 1992: 632 (cataloged, type information); Lê, 1997: 24 (keyed); Lê, 2000: 312, 318 (description, keyed, type information).

Description

Female body length: 1.10–1.63 mm (n=20). Male body length: 1.06–1.07 mm (n=2). Body color: head, mesosoma, and metasoma dark brown to black.

Figures 52–53. 

T. elasmuchae, female (USNMENT00896150) 52 head, mesosoma, metasoma, lateral view 53 head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 54–58. 

T. elasmuchae 54 female (OSUC 144486), head, anterior view 55 female paratype (USNMENT00746982), head and mesosoma, anterolateral view 56 female paratype (USNMENT00746982), mesopleuron and metapleuron, lateral view 57 female (USNMENT00896150), mesopleuron and metapleuron, lateral view 58 female (USNMENT00916427), mesopleuron and metapleuron, lateral view. Scale bars in millimeters.

Head. Color of radicle: pale brown to dark brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: pale to dark brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow; moderately bulging. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: smooth; antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus; uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; strigose, roughly concentric around median ocellus; transversely strigose. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: dorsoventrally fluted. Macrosculpture of lateral frons: absent; rugose; horizontally striate ventrally, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation; separated by less than one ocellar diameter. Hyperoccipital carina: present only posterior to lateral ocellus; absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose; striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2; 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: smooth; transversely strigose; weakly transversely wrinkled. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: absent; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae parallel to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose; smooth area small because cells of surrounding sulci are large. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of distinct foveae. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as clearly defined line of cells. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose; reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout; effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present; absent. Notaulus: absent; indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells; multiple rows of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: between 2 and 3 times as long as stigmal vein. Color of legs: coxae dark brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite; present in anteromedial portion of the tergite. Setation of T2: present in a transverse line posteriorly; present in a transverse line and along lateral margin. Setation of laterotergite 2: absent. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Among Palearctic species, Trissolcus elasmuchae is most similar to T. semistriatus with which it shares a great deal of variability in the sculpture of the frons (compare Figures 54–55 to figures 172–177). Trissolcus elasmuchae can readily be separated from T. semistriatus by numerous characters: the episternal foveae in T. semistriatus are distinctly separate from both the mesopleural pit and the dorsal limit of the posacetabular sulcus whereas in T. elasmuchae the episternal foveae are more numerous and form a continuous line from the postacetabular sulcus to the mesopleural pit; the paracoxal sulcus in the ventral half of the metapleuron is indicated by deep cells in T. elasmuchae, and indicated in T. semistriatus at most as spaces between rugae that radiate from the anterior margin of the metapleuron to the metapleural pit. This form of the paracoxal sulcus is found in a few species of Trissolcus in the New World, T. zakotos, T. radix, and T. solocis, but is not known to us from any other Palearctic species. Trissolcus elasmuchae also has the metapleural epicoxal sulcus indicated by deep cells, which is atypical for the Palearctic fauna.

Material examined

Allotype of T. elasmuchae: JAPAN: 1 male, USNMENT00764939 (EIHU). Paratypes of T. elasmuchae: JAPAN: 2 females, USNMENT00764982 (EIHU); USNMENT00872005 (USNM). Holotype, female, A. davatchii: UNITED KINGDOM: England, Windsor and Maidenhead Unit. Auth., Silwood Park, 1966, reared, B.M. TYPE HYM. 9.796 (deposited in BMNH). Paratypes of T. davatchii: UNITED KINGDOM: 1 male, OSUC 17732 (BMNH). Holotype, female, T. monirus: VIETNAM: Dac Nong Prov., rice seed / rice, Dao Nghia, 25.V.1979, IEBR 0047 (deposited in IEBR). Other material: (45 females, 3 males) CHINA: 2 females, UCRC ENT 296991–296992 (UCRC). FRANCE: 1 female, USNMENT00916119 (BMNH). INDIA: 11 females, UCRC ENT 296980–296990 (UCRC). JAPAN: 13 females, OSUC 144391, 144484144486, 542358, 542364, 542380, 542417, 542419542421, USNMENT00896306, 00896328 (CNCI). SOUTH KOREA: 8 females, 2 males, OSUC 144483, USNMENT00896012, 00896020, 00896021, 00896043, 00896150, 00896151, 00896158 (CNCI); OSUC 542388542389 (OSUC). SWEDEN: 4 females, 1 male, USNMENT00916045, 00916048, 00916111, 00916300, 00916312 (BMNH). TAIWAN: 1 female, OSUC 75842 (OSUC). UNITED KINGDOM: 5 females, USNMENT00916420, 0091642600916428, 00916430 (BMNH).

Comments

With T. davatchii and T. monirus treated as junior synonyms, T. elasmuchae takes on a curious distribution, at least as far as we have documented it, with specimens from Japan, South Korea, India, Vietnam and Europe, reaching as far West as England. However, this distribution is consistent with a pattern that we see in the distributions of other species of Trissolcus. Trissolcus flavipes, for example, is known from Sweden, England, the Asian Far East, and SE Asia.

Trissolcus eriventus

Figures 59, 60–62; Morphbank 16

Trissolcus eriventus Lê, 1997: 23, 25 (original description, keyed); Lê, 2000: 312, 315 (description, keyed, type information).

Description

Female body length: 1.15–1.17 mm (n=5). Body color: head, mesosoma, and metasoma black.

Figure 59. 

Trissolcus eriventus female (USNMENT00916459), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 60–62. 

Trissolcus eriventus 60 female (USNMENT00916458), head, mesosoma, metasoma, dorsal view 61 female (USNMENT00916457) 62 female (USNMENT00916458). Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: dense. Punctation of lateral frons: present along medial margin of orbital furrow. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: uncertain, present only posterior to lateral ocellus; uncertain, absent; uncertain, effaced medially. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum; weakly differentiated from sculpture of dorsal pronotum. Location of pronotal suprahumeral sulcus: posterior half of pronotum.

Mesosoma. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: rugulose and pustulate along ventral half of mesopleural carina. Patch of striae at posteroventral end of femoral depression: uncertain, absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: irregularly rugulose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae.

Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: uncertain, absent; uncertain, present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent; present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: punctures visible in at least ventral half, not quite striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Trissolcus eriventus is a straightforward species to identify; the rugosity of the orbital furrow (Figure 61) is known to us only from this species.

Material examined

Holotype, female: VIETNAM: Hanoi Prov., Nghia Dô, Hanoi, 19.VIII.1982, IEBR 0046 (deposited in IEBR). Other material: CHINA: 5 females, USNMENT00916344, 0091645700916460 (BMNH).

Comments

The hyperoccipital carina in this species is highly variable, ranging from complete to essentially absent between the posterior ocelli. This level of variability is unusual given the stability of this character in other species of Trissolcus.

Trissolcus exerrandus Kozlov & Lê

Figures 63–66; Morphbank 17

Trissolcus exerrandus Kozlov & Lê, 1976: 657, 661 (original description, keyed); Kozlov & Lê, 1977: 502 (keyed); Kozlov, 1978: 629 (description); Kozlov & Kononova, 1983: 80 (description); Kononova, 1995: 92 (keyed).

Description

Female body length: 1.15–1.17 mm (n=5). Body color: head, mesosoma, and metasoma black.

Figures 63–66. 

Trissolcus exerrandus 63 female holotype (ZMAS 0141), head, mesosoma, metasoma, dorsal view 64 female (USNMENT00896159), head and mesosoma, lateral view; 65 female holotype (ZMAS 0141), head, anterior view 66 female (USNMENT00896159), head and mesosoma, ventral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: dense. Punctation of lateral frons: present along medial margin of orbital furrow. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: uncertain, present only posterior to lateral ocellus; uncertain, absent; uncertain, effaced medially. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum; weakly differentiated from sculpture of dorsal pronotum. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: rugulose and pustulate along ventral half of mesopleural carina. Patch of striae at posteroventral end of femoral depression: uncertain, absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: irregularly rugulose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: uncertain, absent; uncertain, present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent; present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: punctures visible in at least ventral half, not quite striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

The transversely arched rugae on the frons ally this species with T. cultratus, with which it shares a poorly defined orbital furrow at its intersection with the malar sulcus. However, it is unlikely to be confused with this species because the densely setose eyes of T. exerrandus separate it from other species of Palearctic Trissolcus. We interpret the mesopleuron to be devoid of episternal foveae: cells at the dorsal limit of the postacetabular sulcus are sometimes slightly larger, but predominantly are the same size as the cells of ventral portion of the sulcus. Faint rugulae are present where the episternal foveae are typically present, leading us to believe that they are simply not externally developed in T. exerrandus. The subacropleural sulcus, present in all other Palearctic specis of Trissolcus, is conspicuously absent in T. exerrandus.

Material examined

Holotype, female: RUSSIA: Primor’ye Terr., Shkotovo, 21.VIII.1961, M. Kozlov, ZMAS 0141 (deposited in ZIN). Paratypes: (2 females, 1 male) MONGOLIA: 1 female, USNMENT00916627 (ZIN). RUSSIA: 1 female, 1 male, USNMENT00916626, 00916628 (ZIN). Other material: (2 females) SOUTH KOREA: 1 female, USNMENT00896159 (CNCI). TAIWAN: 1 female, OSUC 542428 (CNCI).

Trissolcus flavipes (Thomson)

Figures 9, 67–70; Morphbank 18

Trissolcus circus Kozlov & Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3201; Morphbank19

Trissolcus crassus Kononova syn. n.; http://bioguid.osu.edu/xbiod_concepts/354845

Telenomus flavipes Thomson, 1860: 170 (original description).

Aphanurus Flavipes (Thomson): Kieffer, 1912: 72 (description, generic transfer).

Microphanurus flavipes (Thomson): Kieffer, 1926: 91, 96 (description, generic transfer, keyed).

Trissolcus flavipes (Thomson): Kozlov, 1968: 198, 200 (description, synonymy, lectotype designation, keyed); Voegelé, 1969: 149 (keyed);

Trissolcus circus Kozlov & Lê syn. n., 1976: 659, 666 (original description, keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 629 (description); Kozlov & Kononova, 1983: 86 (description); Kononova, 1995: 92 (keyed); Samin, Shojai, Ghahari & Kocak, 2010: 8 (new distribution record for Iran); Petrov, 2013: 325 (keyed).

Trissolcus crassus Kononova syn. n., 2014: 745 (original description, diagnosis).

Description

Female body length: 1.32–1.62 mm (n=8). Body color: head and mesosoma black, metasoma black to reddish brown.

Figures 67–70. 

Trissolcus flavipes 67 female (USNMENT00916409), head, mesosoma, metasoma, lateral view 68 female (USNMENT00916409), head, mesosoma, metasoma, dorsal view 69 female lectotype (NHRS-HEVA 000002617), head, anterior view 70 female (holotype of T. circus) (ZMAS 0143), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: pale brown; yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: A3A4 yellow, elsewhere yellow to brown. Color of A7A11 in female: pale brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: with circular ring, ring margined dorsally by arched rugae. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: moderately dense. Sculpture directly ventral to preocellar pit: absent. Macrosculpture of lateral frons: absent. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent; present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout; effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent; present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent; present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: punctures visible in at least ventral half, not quite striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout; absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): uncertain, 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: uncertain, sparsely present in posterolateral corner. Setation of laterotergite 2: absent; present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: present posteromedially.

Diagnosis

Trissolcus flavipes can be easily identified by the circular impression on the frons directly above the antennal scrobe. Transverse striation may exist surrounding this impression to varying degrees but even in these cases the circular marking remains distinct. The lack of a well-defined orbital furrow near the malar sulcus, the presence of only 2 clypeal setae, and the visible dorsal limit to the cells inside of the axillar crescent (Figure 9) also help to identify this species from other members of the flavipes species group.

Material examined

Lectotype, female, T. flavipes: SWEDEN: Västra Götaland Co., Bohuslän Prov., no date, Boheman, NHRS-HEVA 000002617 (deposited in NHRS). Holotype, female, T. circus: RUSSIA: Primor’ye Terr., Troitsy Bay, 11.VII–15.VII.1972, M. Kozlov, ZMAS 0143 (deposited in ZIN). Paratypes of T. circus: RUSSIA: 2 females, 1 male, USNMENT00916631–00916633 (ZIN). Other material: (7 females, male) JAPAN: 4 females, OSUC 542416, 542418, 542422542423 (CNCI). RUSSIA: 1 male, USNMENT00916634 (ZIN). THAILAND: 1 female, OSUC 523952 (OSUC). UNITED KINGDOM: 2 females, USNMENT00916409, 00916415 (BMNH).

Comments

Trissolcus flavipes has a peculiar distribution, the specimens that we examined directly are from Europe, Eastern Asia, and Thailand and among them we found no characters that suggested a European species and an Asian species. Kononova (2014) published a photograph of the frons of T. crassus that illustrates the circular marking that we have found to be diagnostic for the species. Her description of T. crassus is entirely consistent with our concept of the species, and it is on this basis that we treat T. crassus as a junior synonym of T. flavipes. The two Ukrainian specimens from which she described T. crassus serve to help bridge the distributional gap between the opposite ends of the Palearctic. Notable intraspecific variation occurs in the degree to which microsculpture is expressed in the femoral depression, ranging from distinctly microsculptured throughout to entirely smooth.

Trissolcus gonopsidis (Watanabe)

Figures 71–72, 73–76; Morphbank 20

Trissolcus conidioles Kozlov & Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/179841; Morphbank21

Asolcus gonopsidis Watanabe, 1951: 23, 25 (original description, keyed); Watanabe, 1954: 22 (keyed).

Trissolcus gonopsidis (Watanabe): Kozlov, 1968: 199 (keyed); Kozlov & Lê, 1977: 510 (keyed); Kozlov, 1978: 635 (description); Kozlov & Kononova, 1983: 108 (description); Ryu & Hirashima, 1984: 37, 51 (description, keyed); Johnson, 1992: 629 (cataloged, type information); Kononova, 1995: 95 (keyed); Kononova, 2014: 1422 (keyed); Kononova, 2015: 259 (keyed).

Trissolcus conidioles Kozlov & Lê syn. n., 2000: 314, 362 (original description).

Trissolcus conodioles Kozlov & Lê: Lê, 2000: 312 (keyed, misspelling).

Description

Female body length: 1.40 mm (n=1). Male body length: 1.19 mm (n=1). Body color: head, mesosoma, and metasoma black.

Figures 71–72. 

Trissolcus gonopsidis 71 male allotype (USNMENT00764938), head, mesosoma, metasoma, lateral view 72 female (OSUC 542413), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 73–76. 

Trissolcus gonopsidis 73 female (OSUC 542413) 74 male allotype (USNMENT00764938), head and mesosoma, ventrolateral view 75 male allotype (USNMENT00764938), head, anterior view 76 female (OSUC 542413), head, ventral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: uncertain, bulging; uncertain, narrow. Genal carina: absent; present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: moderately dense. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: anterior half or pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of elongate cells. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: weakly rugulose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: punctures visible in at least ventral half, not quite striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Setation of posterior scutellar sulcus: present. Form of metascutellum: multiple rows of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anteromedial portion of the tergite. Setation of T2: present throughout area posterior to striae. Setation of laterotergite 2: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Trissolcus gonopsidis is a distinctive species: the facial striae of T. gonopsidis are robust (Figures 73–76) and, in combination with episternal foveae distant from the postacetabular sulcus (Figure 74), separate it from other species of the flavipes group.

Associations

emerged from egg of Gonopsis affinis (Uhler): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female of T. conidioles: VIETNAM: Thai Nguyen Prov., Thái Nguyên, no date, IEBR 0039 (deposited in IEBR). Paratypes of T. gonopsidis: JAPAN: 2 females, USNMENT00764937 (EIHU), USNMENT01197395 (ZIN). Allotype: JAPAN: 1 male, USNMENT00764938 (EIHU). Other material: JAPAN: 1 female, OSUC 542413 (CNCI).

Trissolcus hyalinipennis Rajmohana & Narendran

Figures 77–78, 79–83; Morphbank 22

Trissolcus indicus (Subba Rao & Chacko): Fergusson, 1983: 209 (generic transfer, type information).

Trissolcus hyalinipennis Rajmohana & Narendran: Rajmohana K. & Narendran, 2007: 101, 102 (replacement name, keyed).

Description

Female body length: 0.95–0.98 mm (n=3). Male body length: 0.83 mm (n=1). Body color: head and mesosoma black, metasoma black to reddish brown.

Figures 77–78. 

T. hyalinipennis 77 female (USNMENT01109062), head, mesosoma, metasoma, dorsal view 78 female (USNMENT01109060), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 79–83. 

T. hyalinipennis 79 female (USNMENT01109060), venation of fore wing, dorsal view 80 female (USNMENT01109061), antennal clava, ventral view 81 female (USNMENT01109061), head and mesosoma, dorsolateral view 82 female (USNMENT01109060) head and antenna, anterior view 83 female (USNMENT01109060) head, mesosoma, metasoma, dorsolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: pale brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 0. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: faintly striate. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: smooth. Mesopleural pit: simple. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by small punctures. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: pointed. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: absent.

Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum.

Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: faintly present anteriorly. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Setation of S2: present only in medial third.

Diagnosis

Trissolcus hyalinipennis is most similar to T. oobius, with which it shares a 4-merous clava. They may be separated by wing venation: the postmarginal vein is shorter than the stigmal vein in T. oobius and approximately twice as along as the stigmal vein in T. hyalinipennis (compare Figure 79 to Figures 124–127). Apart from this character, we found no other means of separating these two species.

Material examined

Other material: (6 females, 3 males) INDIA: 1 male, UCRC ENT 296999 (UCRC). PAKISTAN: 5 females, USNMENT01109060–01109064 (USNM). PALESTINE: 1 female, USNMENT00916601 (BMNH). SPAIN: 1 male, USNMENT00896154 (CNCI). TURKEY: 1 male, USNMENT00896194 (CNCI).

Comments

Our determination of these specimens is based on their consistency with the description of Subba Rao and Chacko (1961). Many of our specimens were reared from the eggs of Bagrada hilaris (=Bagrada cruciferarum) (subsequently reared on eggs of Podisus maculiventris Say), which is also consistent with the biology presented in the original description. The holotype of this species is now considered to be lost (Rajmohana Keloth, personal communication), and thus designation of a neotype is needed to fully stabilize the name. We refrain from doing so because the reared specimens that we have on hand were collected in Pakistan, sufficiently far from New Delhi to not be considered from the same area, and the specimen from New Delhi that we do have is a male.

Trissolcus japonicus (Ashmead)

Figures 6, 10, 84–87, 88–89; Morphbank 23

Trissolcus dobashii Buhl syn. n.; http://bioguid.osu.edu/xbiod_concepts/223373; Morphbank24

Trissolcus cercus Kozlov & Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3199; Morphbank25

Trissolcus mirus Kononova syn. n.; http://bioguid.osu.edu/xbiod_concepts/354848

Trissolcus pontus Kozlov & Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3292; Morphbank26

Dissolcus japonicus Ashmead, 1904: 73 (original description); Kieffer, 1926: 124, 125 (description, keyed).

Trissolcus japonicus (Ashmead): Masner & Muesebeck, 1968: 72 (type information, generic transfer); Hirashima & Yamagishi, 1981: 153 (description, synonymy); Ryu & Hirashima, 1984: 37, 43 (description, keyed); Talamas, Buffington & Hoelmer, 2013: 114 (description, synonymy, type information).

Trissolcus cercus Kozlov & Lê syn. n., 1976: 659, 666 (original description, keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 89 (description).

Trissolcus pontus Kozlov & Lê syn. n., 1976: 659, 664 (original description, keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 88 (description); Kononova, 1995: 93 (keyed).

Trissolcus dobashii Buhl syn. n., 1996: 128 (original description).

Trissolcus halyomorphae Yang: Qiu, Yang & Tao, 2007: 62 (unavailable: nomen nudum); Yang, Yao, Qiu & Li, 2009: 40 (original description); Talamas, Buffington & Hoelmer, 2013: 114 (junior synonym of Trissolcus japonicus (Ashmead)).

Trissolcus mirus Kononova syn. n., 2014: 749 (original description, diagnosis).

Description

Female body length: 1.16–1.85 mm (n=21). Male body length: 1.15–1.51 mm (n=20). Body color: head, mesosoma, and metasoma black.

Figures 84–87. 

84 Trissolcus cercus (=T. japonicus), female holotype (ZMAS 0145), head, anterior view 85 Trissolcus japonicus, female holotype (USNMENT00831865), head, anterior view 86 Trissolcus pontus (=T. japonicus), female holotype (ZMAS 0144), head, anterior view 87 Trissolcus dobashii (=T. japonicus), female holotype (zmuc00021257), head, anterior view. Scale bars in millimeters.

Figures 88–89. 

Trissolcus japonicus 88 female (USNMENT00979297), head, mesosoma, metasoma, lateral view 89 female (USNMENT00675989), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Head. Color of radicle: orange; yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: dark brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate; faintly rugulose. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: absent. Setation of lateral frons: moderately dense. Punctation of lateral frons: sparse. Sculpture directly ventral to preocellar pit: dorsoventrally fluted. Macrosculpture of lateral frons: absent. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose; weakly transversely wrinkled. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly; rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; weakly rugulose anteriorly, otherwise absent. Pattern of mesoscutal microsculpture: uniform throughout; effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent; present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout; present laterally, absent medially. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: multiple rows of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Color of legs: coxae dark brown to black, femora and tibia yellow to dark brown, trochanters and tarsi yellow to pale brown. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite; present in anterior two-thirds of tergite. Setation of T2: present throughout posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

Trissolcus japonicus is very similar to T. plautiae and T. kozlovi. Trissolcus japonicus can be separated from T. plautiae by the sculpture directly below the median ocellus: in T. japonicus this area is covered with microsculpture and often a dorsoventral furrow is present. In T. plautiae this area is entirely smooth and without microsculpture. Matsou et al. (2013) used the presence of a sublateral setae to separate T. japonicus from T. plautiae. However, in our experience with specimens reared in quarantine, sublateral setae are not uncommon in T. japonicus. Our preliminary analysis of molecular data is based on DNA extracted non-destructively from specimens of T. japonicus collected in China, Japan, and South Korea, and specimens of T. plautiae from China and Japan. In the specimens used for this analysis, and in the specimens used for monographic work, we found that the pattern of sculpture below the anterior ocellus is stable in both species. It is on this basis that we use this character to separate T. plautiae and T. japonicus, which is consistent with the findings of Matsuo et al. (2013).

Trissolcus japonicus and T. kozlovi can be separated by the sculpture of the frons directly above the antennal scrobe and sculpture of the mesoscutum between the notauli. The sculpture of the frons above the antennal scrobe in T. kozlovi is irregular whereas in T. japonicus, this area of the frons is either smooth or with weakly developed transverse lines of sculpture. The posteromedial mesoscutum in T. kozlovi is obliquely striate, with the lines of sculpture extending anterolaterally from the midline. In T. japonicus there is no macrosculpture in the posterior half of the mesoscutum.

The characters used to separate T. japonicus and T. kozlovi are based on examination of a small number of specimens of the latter species, and they are so similar that we considered that they may actually represent variation within a single species. However, because we are still able to consistently separate them based on morphology, we consider it best to continue to treat them as separate species. Analysis of the DNA sequence and biology of T. kozlovi should be a priority in the future, particularly because as BSMB spreads eastward in Europe its distribution is likely to overlap with that of T. kozlovi.

Material examined

Holotype, female, T. dobashii: JAPAN: Fukuoka Pref., woodland / vegetation consisting mainly of bamboo, Mount Aburayama, 10.I.1996, sweeping, P. N. Buhl, zmuc00021257 (deposited in ZMUC). Holotype, female, T. cercus: RUSSIA: Astrakhan’ Reg., Astrakhan Nature Reserve, 20-VII, M. Y. Asse, ZMAS 0145 (deposited in ZIN). Holotype, female, T. pontus: RUSSIA: Primor’ye Terr., Ussuriyskiy (Suputinskiy) Nature Reserve, 2.VIII.1961, Kovalev, ZMAS 0144 (deposited in ZIN). Holotype, female, Dissolcus japonicus: JAPAN: Kanagawa Pref., Ashigarashimo Dist., Hakone Town, no date, Koebele, USNMENT00831865 (deposited in USNM). Paratypes: CHINA: 2 females, USNMENT00872401, 00872402 (USNM). Other material: (185 females, 97 males) CHINA: 109 females, 73 males, USNMENT0091634000916346, 0091646200916464 (BMNH); USNMENT0097919000979198, 0097920000979221 (CNCI); OSUC 75616 (OSUC); UCRC ENT 142613, 142653, 142678, 142682, 142724, 142738, 142750, 143837, 143886, 143900 (UCRC); USNMENT00675704, 0067573800675739, 0067574100675743, 0067574600675749, 00675986, 0067598800675989, 00764964, 00764984, 00916255, 0091671000916787, 00916796, 0091680100916813, 00916815, 0091689500916899, 0091690000916903, 0091691800916930 (USNM). JAPAN: 16 females, 9 males, OSUC 542354, 542373, 542549, USNMENT00896137, 0089613900896340 (CNCI); UCRC ENT 158378 (UCRC); USNMENT0067570900675713, 0067571500675716, 00675755, 00675770, 0087212500872133 (USNM). RUSSIA: 3 females, USNMENT00979287 (CNCI); UCRC ENT 297007, 297010 (UCRC). SOUTH KOREA: 18 females, 11 males, USNMENT00896014, 00896026, 0089603300896036, 0089603800896042, 00896117, 00896120, 00979251, 00979254 (CNCI); USNMENT0067570500675708, 0067571800675720, 0067572300675729 (USNM). TAIWAN: 2 females, UCRC ENT 112211, 296942 (UCRC). UNITED STATES: 37 females, 4 males, USNMENT01109017–01109019 (CNCI); USNMENT01059357, 0105935901059402, 0105940401059407, 0105940901059412, 0105941401059417, 0105942001059422, 0105942401059427, 01059430, 01081080, 01109016, 0110902001109026, 01109132, 01109134, 01109419, 01109562 (USNM).

Comments

Trissolcus cercus, which we here treat as a junior synonym, was collected in western Russia, far from the distribution of what we otherwise know to be the native range of T. japonicus. In the late 1960s, Podisus maculiventris, which is a known host of Trissolcus japonicus, was introduced into Eastern Europe in the late 1960’s as a biological control agent (Schaefer & Panizzi 2000). The eggs associated with the holotype of T. cercus are included on the pin, and their morphology is consistent with that of eggs of P. maculiventris. The presence of T. japonicus in Eastern Europe may be the result of an adventive population of T. japonicus that followed the introduction of P. maculiventris, and then declined along with the stink bug population.

Trissolcus kozlovi Rjachovskij

Figures 90, 91–93; Morphbank 27

Trissolcus amplus Kononova syn. n.; http://bioguid.osu.edu/xbiod_concepts/354847

Trissolcus Kozlovi Rjachovskij, 1975: 4 (original description).

Trissolcus kozlovi Rjachovskij: Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 629 (description); Kozlov & Kononova, 1983: 85 (description); Johnson, 1992: 630 (cataloged, type information); Kononova, 2014: 1421 (keyed); Kononova, 2015: 258 (keyed).

Trissolcus amplus Kononova syn. n., 2014: 747 (original description, diagnosis).

Description

Female body length: 1.08–1.33 mm (n=7). Body color: head, mesosoma, and metasoma black.

Figure 90. 

Trissolcus kozlovi, female holotype (ZMAS 0147), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 91–93. 

Trissolcus kozlovi 91 female (USNMENT00916624), head, mesosoma, metasoma, dorsal view 92 female holotype (ZMAS 0147), head, anterior view 93 female holotype (ZMAS 0147), head, anterolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow; brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: basal A1 and distal A2 yellow, elsewhere brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 4; 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: irregularly rugose, rugae effaced. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: present along medial margin of orbital furrow. Sculpture directly ventral to preocellar pit: absent. Macrosculpture of lateral frons: absent. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose, with oblique symmetryl in relation to longitudinal midline. Pattern of mesoscutal microsculpture: effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate; punctures visible in at least ventral half, not quite striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown, femora yellow to dark brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: sparsely present throughout area not covered by laterotergite.

Material examined

Holotype, female, T. Kozlovi: RUSSIA: Voronezh Reg., Ramon’, 13.VIII.1972, V. Rjachovskij, ZMAS 0147 (deposited in ZIN). Paratypes: RUSSIA: 3 females, USNMENT00916623–00916625 (ZIN). Other material: MOLDOVA: 3 females, USNMENT00916620–00916622 (ZIN).

Diagnosis

Trissolus kozlovi is a difficult species to treat taxonomically because it is represented by a small number of specimens that are extremely similar to T. japonicus and T. plautiae. It remains diagnosable among the specimens at hand by subtle, but seeming reliable characters: from both T. japonicus and T. plautiae it may be separated by the obliquely oriented sculpture on the posterior mesoscutum between the notauli. The sculpture on the frons above the interantennal process is also useful for separating T. kozlovi from these species: in T. kozlovi the frons has irregular lines of sculpture; in T. plautiae this part of the frons is entirely smooth and in T. japonicus it is covered with microsculpture, sometimes with weak transvere striation or a dorsoventral furrow below the median ocellus. Assessment of this character’s utility for species-level identification requires additional specimens and, if possible, corroboration with molecular data.

Comments

In most specimens, including the holotype, there are 4 clypeal setae present. The lateral setae are about half the length of the median pair, and in some specimens the lateral setae appear to be absent. In some cases a seta is absent from one side only, in some, the lateral setae are appressed to the labrum or clypeus, probably as a preservation artifact, in others neither are evident. We suspect that these setae are broken off, but because we cannot be certain, the description for this character is coded as having both 2 and 4 setae.

Trissolcus larides (Nixon)

Figures 94, 95–97; Morphbank 28

Microphanurus larides Nixon, 1943: 138, 141 (original description, keyed); Risbec, 1950: 569 (keyed).

Trissolcus larides (Nixon): Masner, 1965: 126 (type information, generic transfer); Johnson, 1992: 630 (cataloged, type information).

Description

Female body length: 1.00–1.19 mm (n=4). Body color: head and mesosoma black, metasoma black to reddish brown.

Figures 94. 

Trissolcus larides female (USNMENT00896166), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 95–97. 

Trissolcus larides 95 female (USNMENT00896174), head, mesosoma, metasoma, dorsal view 96 female (USNMENT00896174), mesosoma, posterolateral view 97 female (USNMENT00896216), head, anterior view. Scale bars in millimeters

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: pale brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent; present as short grooves. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: absent. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus; absent. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: smooth. Mesopleural pit: simple. Mesopleural carina: complete; absent; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by small punctures. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: longitudinally striate posteriorly, otherwise absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: absent. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: absent. Form of metascutellum: furrow with broad interspaces between crenulae. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: undifferentiated from remainder of propodeum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: weakly striate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: absent. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anteromedial portion of the tergite. Setation of T2: absent. Setation of laterotergite 2: present. Striation of S2: absent. Setation of S2: present only in medial third.

Diagnosis

Trissolcus larides is unique among species with a setose metapleuron in that the episternal foveae are entirely absent. It is most similar to T. levicaudus with which it shares an antero-posteriorly compact habitus, the absence of notauli, a smooth speculum, and T1 smooth posterior to the basal costae. The two may be separated by a number of characters: the anteromedial portion of T2 is striate in T. larides and smooth in T. levicaudus; the mesoscutal humeral and mesoscutal suprahumeral sulci are smooth furrows in T. larides and comprised of cells in T. levicaudus; T. levicaudus has a single episternal fovea and T. larides has none; the prespecular sulcus extends ventrally to the mesopleural pit in T. levicaudus but not in T. larides; the cells along the anterior margin of the metascutellum are deep punctures in T. levicaudus and in T. larides this sulcus is a furrow with broad interspaces between crenulae; the wing membrane at the junction of the marginal, postmarginal, and stigmal veins is pigmented in T. larides and hyaline in T. levicaudus.

Material examined

Holotype, female, Microphanurus larides: SENEGAL: Diourbel Rég., Bambey, 31.V.1939, reared from egg, J. Risbec, B.M. TYPE HYM. 9.310 (deposited in BMNH). Other material: UNITED ARAB EMIRATES: 4 females, USNMENT00896166, 00896174, 00896184, 00896216 (CNCI).

Trissolcus latisulcus (Crawford)

Figures 98–102; Morphbank 29

Trissolcus felis Kozlov & Lê, syn. n.; http://bioguid.osu.edu/xbiod_concepts/179842; Morphbank30

Trissolcus legatarius Lê, syn. n.; http://bioguid.osu.edu/xbiod_concepts/3254; Morphbank31

Telenomus latisulcus Crawford, 1913: 244 (original description); Baltazar, 1966: 172 (cataloged, type information, distribution).

Telenomus oecleus Dodd, 1913: 163, 168 (original description, keyed, synonymized by Johnson (1991)); Johnson, 1991: 223 (junior synonym of Trissolcus latisulcus (Crawford)).

Telenomus darwinensis Dodd, 1915: 7 (original description, synonymized by Johnson (1991)); Johnson, 1991: 223 (junior synonym of Trissolcus latisulcus (Crawford)).

Telenomus erigone Dodd, 1915: 8 (original description, synonymized by Johnson (1991)).

Aphanurus banksi Gahan, 1921: 349 (original description, synonymized by Johnson (1991)); Johnson, 1991: 223 (junior synonym of Trissolcus latisulcus (Crawford)).

Microphanurus oecleus (Dodd): Kieffer, 1926: 93, 95, 111 (description, generic transfer, keyed).

Microphanurus priapus Nixon, 1938: 125, 133 (original description, keyed, synonymized by Johnson (1991)); Nixon, 1943: 139 (keyed); Risbec, 1950: 570 (keyed); Johnson, 1991: 223 (junior synonym of Trissolcus latisulcus (Crawford)).

Trissolcus priapus (Nixon): Masner, 1965: 127 (type information, generic transfer).

Microphanurus banksi (Gahan): Baltazar, 1966: 173 (cataloged, type information, distribution).

Trissolcus banksi (Gahan): Masner & Muesebeck, 1968: 71 (type information).

Trissolcus latisulcus (Crawford): Masner & Muesebeck, 1968: 73 (type information); Johnson, 1991: 213, 223 (description, synonymy, keyed); Rajmohana K. & Narendran, 2007: 102 (keyed).

Asolcus priapus (Nixon): Voegelé, 1969: 151 (keyed).

Trissolcus legatarius Lê syn. n., 1982: 223 (original description); Johnson, 1992: 631 (cataloged, type information); Lê, 2000: 312, 317 (description, keyed, type information).

Trissolcus darwinensis (Dodd): Johnson, 1988: 239 (lectotype designation, generic transfer).

Trissolcus erigone (Dodd): Johnson, 1988: 239 (type information, generic transfer).

Trissolcus oecleus (Dodd): Johnson, 1988: 240 (lectotype designation, generic transfer).

Trissolcus felis Kozlov & Lê syn. n., 2000: 312, 316, 363 (original description, keyed).

Description

Female body length: 1.13–2.17 mm (n=21). Male body length: 1.16–1.97 mm (n=7). Body color: head, mesosoma, and metasoma black.

Figures 98–102. 

T. latisulcus 98 female holotype (USNMENT00989065), head, mesosoma, metasoma, dorsal view 99 female (OSUC 523953), head and mesosoma, ventrolateral view 100 female holotype (USNMENT00989065), mesosoma, posterodorsal view 101 female holotype (USNMENT00989065), dorsal mesosoma, lateral view 102 female holotype (USNMENT00989065), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow; yellow, becoming brown distally. Color of A7A11 in female: pale brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: moderately bulging. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: coarsely rugose. Preocellar pit: present. Setation of lateral frons: sparse; moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: absent; microsculptured; dorsoventrally carinate. Macrosculpture of lateral frons: rugose. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: present only posterior to lateral ocellus. Macrosculpture of posterior vertex: rugulose. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: rugose. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: weakly differentiated from sculpture of dorsal pronotum. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2; 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: smooth; weakly transversely wrinkled; with granular microsculpture. Mesopleural pit: uncertain, simple. Mesopleural carina: complete. Sculpture of femoral depression: rugose perpendicular to long axis of femoral depression. Patch of striae at posteroventral end of femoral depression: uncertain, present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: coarsely rugose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae.

Mesoscutal humeral sulcus: indicated by a line of cells; indicated by smooth furrow with a small number of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose, becoming longitudinally striate posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent; present. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in most of tergite, extending posteriorly to transverse line of setae. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

The coarse sculpture found throughout the femoral depression (Figure 99) distinguishes T. latisulcus from other Palearctic species in the genus. The femoral depression in T. comperei and T. carinifrons has parallel arched rugae (Figures 27, 28) but these are limited to the ventral portion of this area. The anterior margin of the mesoscutellum often is distinctly raised, creating the appearance of an enlarged scutoscutellar sulcus (Figure 101), which is also useful for identifying this species.

Material examined

Holotype, female, T. latisulcus: TAIWAN: Taipei Special Muni., Taipei (Taihoku), VI-1911, T. Shiraki, USNMENT00989065 (deposited in USNM). Paratype of T. latisulcus: TAIWAN: 1 female, USNMENT00872011 (USNM). Holotype, female, A. banksi: PHILIPPINES: Laguna Prov., Los Baños, 15.VII.1918, C. S. Banks, USNMENT00989095 (deposited in USNM). Holotype, female: VIETNAM: Hoa Binh Prov., Van Mai, 30.V.1982, IEBR 0040 (deposited in IEBR). Holotype, female, T. legatarius: VIETNAM: Gia Lai Prov., forest, Ha Nung, 8.XI.1979, X. H. Lê, IEBR 0045 (deposited in IEBR). Other material: (18 females, 9 males, 1 unsexed) CHINA: 4 females, 1 male, USNMENT0091646500916468, 00916470 (BMNH). INDIA: 4 females, 1 male, 1 unsexed, USNMENT00916358, 00916360 (BMNH); OSUC 144507 (CNCI); USNMENT00872013–00872015 (USNM). LAOS: 2 females, USNMENT00764922, 00764924 (BPBM). MALAYSIA: 4 females, 4 males, OSUC 144516, 542429542435 (CNCI). PHILIPPINES: 1 male, USNMENT00872016 (USNM). THAILAND: 4 females, 2 males, USNMENT00764926 (BPBM); OSUC 144510, 144513144514 (CNCI); OSUC 523953 (OSUC); UCRC ENT 134998 (UCRC).

Comments

We observe significant size variation in this species, suggesting that it is a generalist attacking eggs of a variety of sizes, and perhaps this contributes to the expansive distribution of Trissolcus latisulcus from eastern Australia to India.

Trissolcus levicaudus Talamas, sp. n.

Figures 103, 104–106; Morphbank 32

Description

Female body length: 0.74–0.80 mm (n=7). Body color: head and mesosoma black, metasoma black to reddish brown.

Figure 103. 

Trissolcus levicaudus female holotype (OSUC 144511), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 104–106. 

Trissolcus levicaudus 104 female holotype (OSUC 144511), head, mesosoma, metasoma, dorsal view 105 female holotype (OSUC 144511), head and mesosoma, ventrolateral view 106 female paratype (OSUC 144508), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: brown; pale brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; transversely strigose ventrally, absent dorsally. Preocellar pit: absent; present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: absent. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 1. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: smooth. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout; absent; present laterally, absent medially. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: mostly smooth, with furrow along ventral margin. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: absent. Number of sublateral setae (on one side): 0; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: absent. Setation of T2: present in a transverse line medially. Setation of laterotergite 2: absent; present. Striation of S2: absent. Setation of S2: present posteromedially.

Diagnosis

Trissolcus levicaudus is closest morphologically to T. larides, with which it shares a setose posterior metapleuron, absence of notauli, a smooth speculum, and T1 without striae posterior to the cells along the anterior margin. It may be separated from T. larides by the absence of striation on T2, a metapostnotum extending to the lateral margin of the metascutellum in dorsal view, and the entirely hyaline fore wing.

Etymology

The species epithet “levicaudus”, meaning “without knots” or “smooth” refers to the smoothness of the metasomal tergites.

Material examined

Holotype, female: THAILAND: Uthai Thani Prov., Khao Nang Rum Wildlife Research Station, 400m, V-1986, malaise trap, M. Allen, OSUC 144511 (deposited in CNCI). Paratypes: (6 females) INDIA: 3 females, OSUC 144505, 144508144509 (CNCI). THAILAND: 1 female, OSUC 144512 (CNCI). UNITED ARAB EMIRATES: 2 females, USNMENT00896167, 00896173 (CNCI).

Trissolcus mitsukurii (Ashmead)

Figures 7, 107–108, 109–111; Morphbank 34

Trissolcus rudus Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3297; Morphbank35

Telenomus mitsukurii Ashmead, 1904: 72 (original description, keyed); Esaki, Hori, Hozawa, Ishii, Issiki, Kawada, Kawamura, Kinoshita, Kishida, Koidzumi, Kojima, Kuwana, Kuwayama, Marumo, Niijima, Oguma, Okamoto, Shinji, Shiraki, Takahashi, Uchida, Ueno, Yamada, Yano, Yokoyama & Yuasa, 1932: 345 (description, distribution).

Liophanurus Mitsukurii (Ashmead): Kieffer, 1912: 61 (generic transfer).

Telenomus oecleoides Dodd, 1914: 122 (original description, synonymized by Johnson (1991)).

Microphanurus oecleoides (Dodd): Kieffer, 1926: 95, 116 (description, generic transfer, keyed).

Phanurus mitsukurii (Ashmead): Kieffer, 1926: 50, 56 (description, generic transfer, keyed).

Asolcus mitsukurii (Ashmead): Watanabe, 1951: 22, 25 (generic transfer, description, keyed); Watanabe, 1954: 22 (keyed); Tachikawa, 1965: 284 (description).

Trissolcus mitsukurii (Ashmead): Masner & Muesebeck, 1968: 73 (lectotype designation, generic transfer); Safavi, 1968: 415 (keyed); Voegelé, 1969: 149 (keyed); Kozlov & Lê, 1976: 659 (keyed); Kozlov & Lê, 1977: 506 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 92 (description); Ryu & Hirashima, 1984: 36, 39 (description, keyed); Johnson, 1991: 213, 224 (description, synonymy, keyed); Kononova, 1995: 95 (keyed); He et al., 2004: 318 (description).

Trissolcus rudus Lê syn. n., 1983: 24, 25 (original description, keyed); Johnson, 1992: 635 (cataloged, type information); Lê, 1997: 23 (keyed); Lê, 2000: 311, 320 (description, keyed, type information).

Trissolcus oecleoides (Dodd): Johnson, 1988: 240 (type information, generic transfer); Johnson, 1991: 224 (junior synonym of Trissolcus mitsukurii (Ashmead)).

Description

Female body length: 1.04–1.74 mm (n=20). Male body length: 0.90–1.53 mm (n=19). Body color: head, mesosoma, and metasoma black.

Figures 107–108. 

T. mitsukurii 107 female holotype (USNMENT00989031), head, mesosoma, metasoma, ventral view 108 female (USNMENT00675722). Scale bars in millimeters.

Figures 109–111. 

T. mitsukurii 109 female holotype (USNMENT00989031), head, mesosoma, metasoma, lateral view 110 female (USNMENT00675722), head and mesosoma, lateral view 111 female (USNMENT00675722), head and mesosoma, ventrolateral view. Scale bars in millimeters.

Head. Color of radicle: orange. Length of radicle: less than width of clypeus. Color of A1A6 in female: apical A1 and dorsal A2 brown, elsewhere orange. Color of A7A11 in female: black. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: moderately bulging. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: rugose. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: dorsoventrally fluted. Macrosculpture of lateral frons: weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: present only posterior to lateral ocellus. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: antero-posteriorly striate. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: weakly differentiated from sculpture of dorsal pronotum. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: irregularly rugulose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: rugulose. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: absent. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: undifferentiated from remainder of propodeum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae and femora dark brown to black, elsewhere yellow to pale brown and becoming darker at distal tarsomeres.

Metasoma. Anteromedial portion of metasomal depression: antero-posteriorly striate. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1. Setation of laterotergite 1: present. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present.

Diagnosis

Among Palearctic species in the basalis group T. mitsukurii can be identified by the combination of the greatly enlarged antennae clava (A7A11) in females (Figures 107, 109), the size of which is emphasized in comparison to the compressed A4A6, an orbital furrow that is expanded at its intersection with the malar sulcus (Figures 109–111), and the line of setae along the dorsal margin of the first laterotergite (Figure 110). During the course of this revision we examined a number of species from southeast Asia that share the above-mentioned diagnostic characters, but to our knowledge the distributions of these species do not extend into temperate Asia.

Material examined

Holotype, female, T. mitsukurii: JAPAN: Shizuoka Pref., Atami City, no date, Koebele, USNMENT00989031 (deposited in USNM). Paratype of T. mitsukurii: JAPAN: 1 female, USNMENT00675967 (USNM). Holotype, female, T. rudus: VIETNAM: Hanoi Prov., Nghia Dô, Hanoi, 7-VII-1978, X. H. Lê, IEBR 0042 (deposited in IEBR). Other material: (39 females, 29 males, 5 unsexed) CHINA: 1 female, USNMENT00916469 (BMNH). JAPAN: 28 females, 27 males, 5 unsexed, OSUC 542353, 542355, 542362, 542376542378, 542425, USNMENT00896316 (CNCI); USNMENT0067572100675722, 0067575000675754, 0067575600675760, 0067576200675769, 00675771, 0067584700675859, 0067596800675979, 00764853, 0087200000872004 (USNM). SOUTH KOREA: 1 female, USNMENT00896149 (CNCI). THAILAND: 5 females, OSUC 542391542395 (OSUC). UNITED STATES (quarantine): 4 females, 2 males, USNMENT00872119–00872124 (USNM).

Trissolcus nycteridaner Talamas, sp. n.

Figures 12, 112–113, 114–117; Morphbank 35

Description

Female body length: 1.50–1.79 mm (n=10). Male body length: 1.60 mm (n=1). Body color: head, mesosoma, and metasoma black.

Figures 112–113. 

Trissolcus nycteridaner, female holotype (USNMENT00896182) 112 head, mesosoma, metasoma, lateral view 113 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 114–117. 

Trissolcus nycteridaner 114 female holotype (USNMENT00896182), mesosoma, lateral view 115 female holotype (USNMENT00896182), head and mesosoma, posterodorsal view 116 female paratype (USNMENT00896175), head, anterior view 117 female holotype (USNMENT00896182), head, anteroventral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally. Color of A7A11 in female: pale brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: faintly rugulose. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: moderately dense. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: absent. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: rugose. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 3; 4 or more. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by open crenulae; formed by small punctures. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells; absent. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: absent. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: longitudinally strigose medially. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: absent. Form of metascutellum: coarsely rugose. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: present throughout posterolateral corner and lateral portions of tergite. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally, absent medially. Setation of S2: present only in medial third.

Diagnosis

Trissolcus nycteridaner can be identified by the presence of a hyperoccipital carina, the undefined orbital furrow at the malar sulcus, the extremely slender mandibles, and the longitudinal sculpture on the mesoscutellum and posterior mesoscutum. It is similar to T. gonopsidis, but can be easily distinguished by the slender mandibles and absence of facial striae. Among species of the flavipes group, it is unusual in that the mesoscutal humeral sulcus is a mostly smooth furrow.

Etymology

The name of this species was inspired by young Luke Meladossi of Scarsdale, New York, who inquired if a species could be named for the superhero Batman. Like Batman, species of Trissolcus are dark in color, rarely seen by the public, kill bad guys (stink bugs) for the benefit of humankind, and are at times unfairly vilified on the basis of their name (parasitic wasp). The epithet derives from the Greek words “nycteris,” meaning “bat”, and “aner,” meaning “man”, and is treated as a noun in apposition.

Material examined

Holotype, female: UNITED ARAB EMIRATES: #8173, 25°24’N 56°17’E, Wadi Wurayah, 25.III.2007, sweeping, F. Menzel & A. Stark, USNMENT00896182 (deposited in CNCI). Paratypes: UNITED ARAB EMIRATES: 9 females, USNMENT0089617500896178, 0089618000896181, 0089618500896186, 00896218 (CNCI). Other material: UNITED ARAB EMIRATES: 1 male, USNMENT00896179 (CNCI).

Trissolcus oobius (Kozlov)

Figures 118–119, 120–127; Morphbank 36

Trissolcus aglaope (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3176; Morphbank37

Trissolcus dirrhope (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3216; Morphbank38

Trissolcus dryope (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3220; Morphbank39

Trissolcus lampe (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3251

Trissolcus merope (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3266; Morphbank40

Trissolcus niceppe (Kozlov & Lê) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3272; Morphbank41

Aporophlebus oobius Kozlov, 1972: 670 (original description); Kononova, 1973: 439, 440 (description, keyed); Kozlov & Lê, 1976: 350 (keyed).

Aporophlebus dirrhope Kozlov & Lê, 1976: 351, 368 (original description, keyed).

Aporophlebus dryope Kozlov & Lê, 1976: 351, 365 (original description, keyed).

Aporophlebus lampe Kozlov & Lê, 1976: 350, 360 (original description, keyed).

Aporophlebus merope Kozlov & Lê, 1976: 350, 359 (original description, keyed).

Aporophlebus niceppe Kozlov & Lê, 1976: 351, 369 (original description, keyed).

Trissolcus oobius (Kozlov): Kozlov & Lê, 1977: 518 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 126 (description); Kononova, 1995: 98 (keyed); Petrov, 2013: 326 (keyed).

Aporophlebus aglaope Kozlov & Lê, 1976: 351, 363 (original description, keyed).

Trissolcus aglaope (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 519 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 133 (description).

Trissolcus dirrhope (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 519 (generic transfer, keyed); Kozlov, 1978: 638 (description); Kozlov & Kononova, 1983: 134 (description).

Trissolcus dryope (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 519 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 132 (description); Kononova, 1995: 98 (keyed); Ghahari, Buhl & Kocak, 2011: 595 (listed).

Trissolcus lampe (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 518 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 129 (description).

Trissolcus merope (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 518 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 124 (description); Kononova, 1995: 98 (keyed).

Trissolcus niceppe (Kozlov & Lê) syn. n.: Kozlov & Lê, 1977: 519 (generic transfer, keyed); Kozlov, 1978: 638 (description); Kozlov & Kononova, 1983: 135 (description).

Description

Female body length: 0.68–1.04 mm (n=8). Body color: head and mesosoma black, metasoma black to reddish brown.

Figures 118–119. 

T. oobius, female paratype (USNMENT00916617) 118 head, mesosoma, metasoma, lateral view 119 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 120–127. 

T. oobius 120 female paratype of T. niceppe (USNMENT00916279), head and mesosoma, ventrolateral view 121 female paratype of T. oobius (USNMENT00916617), head, mesosoma, T1–T2, dorsolateral view 122 female paratype of T. merope (USNMENT00916614), head and anterior mesosoma, lateral view 123 female paratype of T. dirrhope (USNMENT00916275), head, anterior view 124 female paratype of T. merope (USNMENT00916614), venation of fore wing, dorsal view 125 female paratype of T. dirrhope (USNMENT00916275), venation of fore wing, dorsal view 126 female paratype of T. niceppe (USNMENT00916279), venation of fore wing, dorsal view 127 female paratype of T. oobius (USNMENT00916619), venation of fore wing, dorsal view. Scale bars in millimeters.

Head. Color of radicle: pale brown; yellow; brown. Length of radicle: less than width of clypeus. Color of A7A11 in female: pale brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 0. Color of A1A7 in female: yellow; pale brown. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent; present. Shape of ventral gena in lateral view: narrow. Genal carina: absent; present and extending dorsally to vicinity of lower margin eye; present only at base of mandible. Malar striae: absent. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: absent; present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent; weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation; separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: weakly transversely wrinkled. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent; present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout; present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by small punctures. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; weakly rugulose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent; indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout; absent; present laterally, absent medially. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: absent. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: shorter than length of stigmal vein. Color of legs: coxae brown, femora and tibiaie yellow to pale brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate; smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present; absent; weakly present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: absent; faintly present anteriorly. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present.

Diagnosis

Trissolcus oobius shares with T. hyalinipennis a 4-merous antennal clava in females. This character, in combination with a postmarginal vein that is shorter than the stigmal vein, unambiguously seperates it from all Palearctic species. Kozlov described numerous species based on small differences in the length of the postmarginal vein, and we here treat most of them as as junior synonyms. The length of the postmarginal vein varies from nearly absent to approaching the length of the stigmal vein in a continuous gradient such that separation based on arbitrary lengths is not useful for species identification (see Figures 124–127).

Material examined

Holotype, female, T. oobius: RUSSIA: Altay Terr., Kosh-Agach, 17.VI.1964, Kozlov, ZMAS 0130 (deposited in ZIN). Paratypes of T. oobius: RUSSIA: 4 unsexed, USNMENT00916616–00916619 (ZIN). Holotype, female, T. aglaope: TURKMENISTAN: Lebap Prov., Repetek, 6.VI.1968, Sugonyaev, ZMAS 0117 (deposited in ZIN). Paratype of T. oobius: TURKMENISTAN: 1 male, USNMENT00916326 (ZIN). Holotype, female, T. dirrhope: TURKMENISTAN: Lebap Prov., Halaç (Khalach), 25.VI.1967, ZMAS 0123 (deposited in ZIN). Paratype of T. dirrhope: TURKMENISTAN: 1 female, USNMENT00916275 (ZIN). Holotype, female, T. dryope: KAZAKHSTAN: Atyrau (Gurev) Reg., shore, Lake Inder, 11.VII.1974, V. V. Kostjukov, ZMAS 0124 (deposited in ZIN). Paratype of T. dryope: MONGOLIA: 1 female, USNMENT00916277 (ZIN). Holotype, female, T. lampe: KAZAKHSTAN: Atyrau (Gurev) Reg., Ural River Floodplain, Inderbor (Inderborskiy), 8.VII.1974, V. V. Kostjukov, ZMAS 0126 (deposited in ZIN). Holotype, female, T. merope: RUSSIA: Altay Terr., Kosh-Agach, 11.VII.1964, Kozlov, ZMAS 0127 (deposited in ZIN). Paratype of T. merope: RUSSIA: 1 female, USNMENT00916614 (ZIN).Paratypes of T. niceppe: (2 females) ARMENIA: 1 female, USNMENT00916279 (ZIN). TURKMENISTAN: 1 female, ZMAS 0129 (ZIN). Other material: (4 females) IRAN: 1 female, USNMENT00896237 (CNCI). ISRAEL: 2 females, USNMENT00896142, 00896145 (CNCI). TURKMENISTAN: 1 female, UCRC ENT 296993 (UCRC).

Comments

The degree to which longitudinal rugae extend from the basal costae of T1 is variable, ranging from essentially absent to distinctly present. Similarly, anterior T1 may be faintly striate or entirely smooth. Microsculpture on the mesoscutellum varies from present throughout to absent, and in some specimens the microsculpture is only present laterally. Specimens in the type series of T. oobius are larger than the specimens in the type series of species here treated as junior synonyms. These larger specimens have coarser sculpture on the frons and a distinct preocellar pit. In the smallest specimens the preocellar pit is absent and in specimens of intermediary size the pit is absent or very small.

Trissolcus perepelovi (Kozlov)

Figures 128–132; Morphbank 42

Trissolcus deserticola (Kozlov) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3214; Morphbank43

Trissolcus dircus Kozlov & Lê syn. n.; http://bioguid.osu.edu/xbiod_concepts/3215; Morphbank44

Aporophlebus deserticola Kozlov, 1972: 668 (original description); Kozlov & Lê, 1976: 350 (keyed).

Aporophlebus perepelovi Kozlov, 1972: 669 (original description); Kozlov & Lê, 1976: 350 (keyed).

Trissolcus deserticola (Kozlov) syn. n.: Kozlov & Lê, 1977: 518 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 127 (description); Kononova, 1995: 98 (keyed).

Trissolcus perepelovi (Kozlov): Kozlov & Lê, 1977: 518 (generic transfer, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 129 (description); Kononova, 1995: 98 (keyed).

Trissolcus dircus Kozlov & Lê syn. n., 1977: 516, 521 (original description, keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 120 (description); Radjabi, 2001: 111 (keyed).

Description

Female body length: 1.00–1.54 mm (n=5). Body color: head, mesosoma, and metasoma black.

Figures 128–132. 

T. perepelovi 128 female paratype (USNMENT00916339), head, mesosoma, metasoma, lateral view 129 female paratype of T. dircus syn. n. (USNMENT00916278), head and mesosoma, dorsal view 130 female paratype (USNMENT00916338), mesosoma, dorsal view 131 female paratype of T. dircus syn. n. (USNMENT00916278), head, anterior view 132 female paratype (USNMENT00916338), head, anterolateral view. Scale bars in millimeters.

Head. Color of radicle: brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent; present and extending dorsally to vicinity of lower margin eye; present only at base of mandible. Malar striae: absent. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: absent. Macrosculpture of lateral frons: absent; horizontally striate ventrally, striae of antennal scrobe extending to lateral frons. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. .

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: smooth. Mesopleural pit: simple; extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: absent; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent; present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: absent; present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by small punctures; present as a smooth furrow. Mesopleural epicoxal sulcus: formed by open crenulae; present as a smooth furrow. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture; present as coarse rugae. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: absent. Posterodorsal margin of axillular carina: round. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about equal to length of stigmal vein. Color of legs: coxae dark brown, femora yellow to dark brown, elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: absent. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anteromedial portion of the tergite. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present.

Diagnosis

Trissolcus perepelovi can be distinguished by the distinctly bulging ventral mesopleuron, the absence of episternal foveae, striation on T2 that is limited to the anteromedial portion of the tergite, and the smooth area on the frons between the antennal scrobe and the median ocellus. Variability occurs in the extent to which the microsculpture on the frons is effaced. In the paratypes of T. deserticola and T. dircus that we examined, the area does not extend to the inner margins of the compound eyes, whereas in the type series of T. perepelovi it may either extend to the margin of the compound eye or be separated by a patch of microsculpture. The expanded smooth area in the type series of T. perepelovi corresponds to a reduction in surface sculpture of the gena and reduction in the size of the genal carina, which are much more pronounced in the paratypes of T. deserticola and T. dircus. As in many other species of Trissolcus, the color of the antennal segments preceding the clava is not stable and A3A6 vary from yellow to brown. Similar to T. semistriatus and T. comperei, the extent to which the striae of the antennal scrobe extend onto the ventral frons is variable within this species. The paratype of T. dircus shows definitive striation on the ventral frons, whereas this striation is very weak in the type series of T. perepelovi and absent in the paratype of T. deserticola.

Material examined

Holotype, female, Aporophlebus perepelovi: MONGOLIA: Bayanhongor Prov., buckwheat / saxaul, 24–35km E Dzamiin Bilgihi (Talin-Bilgekh) Spring, 17.VIII.1969, M. Kozlov, ZMAS 0131 (deposited in ZIN). Paratypes: MONGOLIA: 3 females, USNMENT00916338–00916339, 00916615 (ZIN). Holotype, female, Trissolcus dircus TURKMENISTAN: Lebap Prov., 70km NW Türkmenabat (Chardzhou), 31.V.1965, M. Kozlov, ZMAS 0122 (deposited in ZIN). Paratype of T. dircus: TURKMENISTAN: 1 female, USNMENT00916278 (ZIN). Holotype, female, Aporophlebus deserticola: TURKMENISTAN: Lebap Prov., 70km NW Türkmenabat (Chardzhou), 30.V.1965, M. Kozlov, ZMAS 0121 (deposited in ZIN). Paratype of A. deserticola: TURKMENISTAN: 1 female, USNMENT00916325 (ZIN).

Trissolcus plautiae (Watanabe)

Figures 133, 134–136; Morphbank 45

Asolcus plautiae Watanabe, 1954: 18, 22 (original description, keyed, synonymized by Hirashima & Yamagishi (1981)); Johnson, 1992: 630 (type information).

Trissolcus plautiae (Watanabe): Kozlov, 1968: 198 (keyed); Kozlov & Lê, 1976: 658 (keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 629 (description); Hirashima & Yamagishi, 1981: 153 (junior synonym of Trissolcus japonicus (Ashmead)); Kozlov & Kononova, 1983: 86 (description); Kononova, 1995: 92 (keyed); Kononova, 2014: 1421 (keyed); Kononova, 2015: 258 (keyed).

Description

Female body length: 1.03–1.39 mm (n=26). Male body length: 0.92–1.26 mm (n=20). Body color: head, mesosoma, and metasoma black.

Figure 133. 

Trissolcus plautiae female (USNMENT00916880), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 134–136. 

Trissolcus plautiae 134 female paratype (USNMENT00903180), head, mesosoma, metasoma, dorsal view 135 female (USNMENT00872403), head, anterior view 136 female paratype (USNMENT00872403), head and mesosoma, anterolateral view. Scale bars in millimeters.

Head. Color of radicle: orange; pale brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: dark brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: sparse. Sculpture directly ventral to preocellar pit: absent. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: smooth; weakly transversely wrinkled. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells; formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly; rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: extending to antero-admedian line. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent; present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown, femora yellow to dark brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0; 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present throughout posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

See diagnosis of T. japonicus.

Material examined

Paratypes: JAPAN: 2 females, USNMENT00903180–00903181 (USNM). Other material: (122 females, 25 males, 1 unsexed) CHINA: 90 females, 23 males, UCRC ENT 142662, 143855, 143907 (UCRC); USNMENT00872403, 0091669000916709, 00916795, 0091684000916859, 0091686100916894, 0091690400916917, 0091693100916951 (USNM). JAPAN: 8 females, 1 male, 1 unknown, OSUC 144489, 398858, 542359, 542414 (CNCI); OSUC 75843–75848 (OSUC). RUSSIA: 6 females, UCRC ENT 297000, 297006, 297008, 297011, 329910, 329920 (UCRC). SOUTH KOREA: 18 females, USNMENT0097922700979236, 0097923800979240, 0097924200979245, 00979252 (CNCI). TAIWAN: 1 male, OSUC 144515 (CNCI).

Trissolcus rufiventris (Mayr)

Figures 137–138, 139–144; Morphbank 46

Trissolcus protogyne (Voegelé) syn. n.; http://bioguid.osu.edu/xbiod_concepts/309567; Morphbank47

Telenomus rufiventris Mayr, 1907: 158 (original description).

Prophanurus Rufiventris (Mayr): Kieffer, 1912: 41, 59 (description, generic transfer).

Dissolcus rufiventris (Mayr): Kieffer, 1926: 124 (description, generic transfer, keyed); Meier, 1940: 79, 80 (description, keyed); Rjachovskij, 1959: 82 (keyed).

Microphanurus anitus Nixon, 1939: 131, 132 (original description, keyed, synonymized by Delucchi (1961)); Delucchi, 1961: 54 (junior synonym of Asolcus rufiventris (Mayr)).

Asolcus rufiventris (Mayr): Masner, 1959: 380 (diagnosis, variation); Delucchi, 1961: 45, 54 (description, synonymy, keyed); Viktorov, 1964: 1014, 1023 (description, keyed); Voegelé, 1964: 28 (keyed); Szabó, 1976: 176 (keyed).

Telenomus rubriventris Szabó, 1959: 169 (original description, synonymized by Kozlov (1968)); Kozlov, 1968: 211 (junior synonym of Trissolcus rufiventris (Mayr)).

Trissolcus rufiventris (Mayr): Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 415 (keyed); Kozlov, 1968: 199, 211 (description, lectotype designation, keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 514 (keyed); Kozlov, 1978: 636 (description); Kozlov & Kononova, 1983: 117 (description); Kononova, 1995: 96 (keyed); Koçak & Kilinçer, 2003: 303, 313 (keyed, description); Ghahari, Buhl & Kocak, 2011: 596 (listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 326 (keyed).

Asolcus rufiventris rufiventris (Mayr): Voegelé, 1969: 151 (keyed).

Asolcus rufiventris protogyne Voegelé syn. n., 1969: 151 (original description, keyed, considered a valid species but described as a subspecies).

Description

Female body length: 0.99–1.24 mm (n=18). Male body length: 1.08–1.24 mm (n=6). Body color: head, mesosoma and T1 reddish brown to black, remainder of metasoma yellow to reddish brown.

Figures 137–138. 

T. rufiventris female lectotype (NHMW 0005) 137 head, mesosoma, metasoma, lateral view 138 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 139–144. 

T. rufiventris 139 female (USNMENT01059199), head, mesosoma, metasoma, dorsolateral view 140 female (USNMENT01059197), head, mesosoma, ventrolateral view 141 female (USNMENT00675886), antennal clava, ventral view 142 female (USNMENT00675886), antennal clava, ventral view 143 female lectotype (NHMW 0005), venation of fore wing, dorsal view 144 female (USNMENT00989158), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: pale to dark brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 1. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent; present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 0. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple; extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by small punctures. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: uncertain, dorsoventrally strigose. Posterodorsal margin of axillular carina: pointed. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent. Longitudinal striation of T2: absent; faintly present anteriorly. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present. Striation of S2: absent. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

Trissolcus rufiventris is usually easily identifiable by the color of the metasoma, which in some cases is bright yellowish-orange in color. This color pattern is not known to us from other species and thus can be used effectively to confirm the identity of this species. However, the metasoma may be dark brown, approaching black, and thus color should not be used to exclude specimens from this species. Although it is less accessible, the most reliable character for T. rufiventris is the presence of a single basiconic sensillum on A7. All other species of Palearctic Trissolcus have either 2 basiconic sensilla on A7 or none.

Material examined

Lectotype, female, T. rufiventris: RUSSIA: Rostov Reg., Rostov-na-Donu (Rostow am Don) City, V-1905, reared, I. Vassiliev, NHMW 0005 (deposited in NHMW). Paralectotypes: RUSSIA: 3 females, 2 males, 1 unsexed, USNMENT0098915800989159, 0105903501059037 (NHMW). Other material: (30 females, 15 males) ARMENIA: 1 female, USNMENT00979996 (ZIN). FRANCE: 2 females, USNMENT00896293, 00896330 (CNCI). ISRAEL: 1 female, USNMENT00896143 (CNCI). ITALY: 1 female, USNMENT00896204 (CNCI). MOROCCO: 4 females, 5 males, USNMENT00896091 (CNCI); USNMENT01059197, 0105920001059207 (MNHN). TURKEY: 20 females, 10 males, USNMENT00675860–00675889 (USNM). TURKMENISTAN: 1 female, UCRC ENT 296994 (UCRC).

Trissolcus saakowi (Mayr)

Figures 11, 145, 146–147, 148–153; Morphbank 48

Trissolcus mentha Kozlov & Lê, syn. n.; http://bioguid.osu.edu/xbiod_concepts/3265; Morphbank49

Trissolcus radjabii Iranipour, syn. n.м http://bioguid.osu.edu/xbiod_concepts/244989; Morphbank50

Telenomus Saakowi Mayr, 1903: 397 (original description).

Microphanurus saakowi (Mayr): Kieffer, 1926: 91, 95 (description, generic transfer, keyed).

Trissolcus saakovi (Mayr): Kozlov, 1968: 198, 202 (description, emendation, lectotype designation, keyed); Fabritius, 1972: 30 (keyed); Szabó, 1975: 266 (keyed); Kozlov & Lê, 1976: 658 (keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 629 (description); Kozlov & Kononova, 1983: 84 (description).

Trissolcus mentha Kozlov & Lê syn. n.: Kozlov & Lê, 1977: 510 (keyed); Kozlov & Lê, 1977: 1253 (original description); Kozlov, 1978: 635 (description); Kozlov & Kononova, 1983: 107 (description); Ghahari, Buhl & Kocak, 2011: 596 (listed).

Trissolcus radjabii Iranipour syn. n., 2010: 66 (original description, diagnosis); Ghahari, Buhl & Kocak, 2011: 596 (listed).

Trissolcus saakowi (Mayr): Ghahari, Buhl & Kocak, 2011: 596 (listed).

Description

Female body length: 1.06–1.94 mm (n=9). Male body length: 1.69 mm (n=1). Body color: head, mesosoma, and metasoma black.

Figure 145. 

Trissolcus saakowi female lectotype (NHMW 0003), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 146–147. 

Trissolcus saakowi 146 female lectotype (NHMW 0003), head, mesosoma, metasoma, dorsolateral view 147 female (USNMENT00977544), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 148–153. 

Trissolcus saakowi 148 female lectotype (NHMW 0003), head, anterolateral view 149 female (USNMENT00977544), T1– T2, posterolateral view 150 female (OSUC 259830, paratype of T. radjabii syn. n.) 151 female (USNMENT00977544), mesosoma, posterodorsal view 152 female (USNMENT00977544), head, anterior view 153 female lectotype (NHMW 0003), mesosoma, lateral view. Scale bars in millimeters.

Head. Color of radicle: orange; pale brown; yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: pale brown; yellow; yellow, becoming brown distally. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent; present as short grooves. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: sparse; dense. Sculpture directly ventral to preocellar pit: dorsoventrally fluted; microsculptured. Macrosculpture of lateral frons: absent; horizontally striate ventrally, striae of antennal scrobe extending to lateral frons. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus; rugose. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple; extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; reticulate anteriorly, becoming longitudinally strigose posteriorly; longitudinally striate posteromedially, otherwise absent; longitudinally striate posteriorly, otherwise absent; longitudinally striate. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: absent; indicated only at posterior margin of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: uncertain, absent; uncertain, present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: rugulose punctate. Macrosculpture of mesoscutellum: rugose throughout; rugose laterally. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae dark brown to black, trochanters and femora yellow to brown, elsewhere yellow to orange. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: present. Longitudinal striation of T2: present in most of tergite, extending posteriorly to transverse line of setae. Setation of T2: present throughout posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally, absent medially; absent. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Setation on the first laterotergite unambiguously separates T. saakowi from all Palearctic species in the flavipes group. This character occurs in T. stoicus, T. mitsukurii, and rarely in T. semistriatus, all of which are quite different species in terms of their diagnostic characters and general appearance. Most similar to T. saakowi in the Palearctic is T. tumidus, with which it shares the punctate-rugulose interior of the axillar crescent (Figures 8, 11). Trissolcus saakowi may be separated from T. tumidus by setation of the first laterotergite and the absence of a well defined orbital furrow at its intersection with the malar sulcus. These two species have similar distributions, stretching from the the Middle East to Far Eastern Asia.

Material examined

Lectotype, female, T. saakowi: TURKMENISTAN: Askhabad (Askabad) Dist., VII-1902, reared, J. Vassillief, NHMW 0003 (deposited in NHMW). Paralectotype of T. saakowi: TURKMENISTAN: 1 female, USNMENT01059011 (NHMW). Holotype, female, T. mentha: UZBEKISTAN: Samarkand Reg., Samarqand (Samarkand), 13.VII.1957, E. Popova, ZMAS 0146 (deposited in ZIN). Paratypes of T. radjabii: IRAN: 5 females, OSUC 259828259832 (OSUC). Other material: (23 females, 14 males) AFGHANISTAN: 3 females, 1 male, USNMENT00916471-00916474 (BMNH). CHINA: 1 female, USNMENT00977544 (USNM). GREECE: 1 female, 1 male, USNMENT00916066 (BMNH); USNMENT00896234 (CNCI). INDIA: 1 female, 1male, USNMENT00916359 (BMNH); OSUC 144506 (CNCI). IRAN: 2 females, USNMENT00916481, 00916484 (BMNH). MOROCCO: 10 females, 9 males, USNMENT00896068, 00896077-00896078, 00896081-00896082, 00896092, 00896095-00896102, 00896104-00896105, 00896108, 00896327, 00896332 (CNCI). PAKISTAN: 2 females, 2 males, USNMENT00916354-00916357 (BMNH). SPAIN: 1 female, USNMENT00896334 (CNCI). TURKEY: 2 females, CNC424591, CNC424592T (CNCI).

Comments

Trissolcus saakowi exhibits variability in the sculpture of the mesonotum and the punctation and sculpture of the frons lateral to the antennal scrobe. The mesoscutum may be longitudinally striate throughout, reticulate anteriorly with longitudinal lines medially or posteriorly, or there may be no pronounced macrosculpture at all. The notaulus is typically very short, but may be indistinguishable from mesoscutal striation or even entirely absent. The holotype of T. mentha is smaller than most specimens of T. saakowi, which we attribute to development in a smaller host egg. It also exhibits less pronounced surface sculpture than the larger specimens of T. saakowi and the notauli are absent. Female specimens from Pakistan and China have dense punctation on the lateral frons and striation that extends laterally from the antennal scrobe onto the ventrolateral frons. Two males from Pakistan have label data identical to that of the females, and have the sparse punctation and absence of macrosculpture on the frons as do males from other regions.

Trissolcus scutellaris (Thomson)

Figures 154, 155–157, 158–163, 164–167; Morphbank 51

Trissolcus choaspes (Nixon) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3200; Morphbank52

Trissolcus evanescens Kieffer syn. n.; http://bioguid.osu.edu/xbiod_concepts/3233; Morphbank53

Trissolcus festivae (Viktorov) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3235; Morphbank54

Trissolcus ghorfii (Delucchi & Voegele) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3240; Morphbank55

Trissolcus histani (Voegele) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3243; Morphbank56

Trissolcus simoni (Mayr) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3308; Morphbank57

Trissolcus vassilliewi (Mayr) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3328; Morphbank58

Trissolcus volgensis (Viktorov) syn. n.; http://bioguid.osu.edu/xbiod_concepts/3332; Morphbank59

Telenomus scutellaris Thomson, 1860: 171 (original description); Mayr, 1879: 699, 706 (description, keyed).

Telenomus Simoni Mayr, 1879: 699, 705 (original description, keyed); Mayr, 1903: 398 (description of male, emendation of Mayr (1879)).

Telenomus simonii Mayr: Dalla Torre, 1898: 519 (emendation).

Telenomus Vassilliewi Mayr, 1903: 399 (original description).

Trissolcus evanescens Kieffer syn. n., 1904: 47 (original description); Kieffer, 1926: 126, 127 (description, keyed); Dessart, 1975: 9 (type information).

Aphanurus Scutellaris (Thomson): Kieffer, 1912: 76 (description, generic transfer).

Trissolcus Evanescens Kieffer: Kieffer, 1912: 20 (description).

Trissolcus Simoni (Mayr): Kieffer, 1912: 21 (description, generic transfer).

Microphanurus scutellaris (Thomson): Kieffer, 1926: 91, 99 (description, generic transfer, keyed).

Microphanurus vassilliewi (Mayr): Kieffer, 1926: 91, 97 (description, generic transfer, keyed).

Trissolcus simoni (Mayr): Kieffer, 1926: 126, 127 (description, keyed); Meier, 1940: 79, 80 (description, keyed); Rjachovskij, 1959: 82 (keyed); Safavi, 1968: 414 (keyed); Kozlov, 1968: 198, 204, 207, 210 (description, lectotype designation, synonymy, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 30 (keyed); Szabó, 1975: 266 (description, lectotype designation, keyed); Kozlov & Lê, 1977: 506 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 96 (description); Koçak & Kilinçer, 2003: 302, 303 (keyed, description); Ghahari, Buhl & Kocak, 2011: 597 (listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 325 (keyed).

Microphanurus choaspes Nixon, 1939: 130, 131 (original description, keyed).

Microphanurus vassilievi (Mayr): Meier, 1940: 80 (description, emendation, keyed).

Trissoscelio evanescens (Kieffer): Kelner-Pillault, 1958: 152 (type information, spelling error).

Asolcus scutellaris (Thomson): Masner, 1959: 379 (diagnosis, variation); Kozlov, 1963: 295 (diagnosis); Viktorov, 1964: 1013, 1018 (description, keyed); Szabó, 1976: 176 (keyed).

Trissolcus (Microphanurus) vassilievi (Mayr): Rjachovskij, 1959: 83 (keyed).

Asolcus ghorfii Delucchi & Voegele: Delucchi, 1961: 44, 48 (description, keyed); Delucchi & Voegelé, 1961: 37 (original description); Voegelé, 1964: 28 (keyed); Voegelé, 1965: 111 (keyed); Baur, 2001: 67 (type information).

Asolcus simoni reticulatus Delucchi, 1961: 44, 49 (original description, keyed, synonymized by Kozlov (1968)); Kozlov, 1968: 210 (junior synonym of Trissolcus simoni (Mayr)); Baur, 2001: 67 (type information).

Asolcus vassilievi (Mayr): Delucchi, 1961: 44, 45 (description, keyed); Viktorov, 1964: 1013 (keyed); Voegelé, 1964: 28 (keyed); Voegelé, 1965: 111 (keyed).

Asolcus simoni (Mayr): Kozlov, 1963: 295 (diagnosis); Delucchi, 1963: 13 (diagnosis); Viktorov, 1964: 1013, 1016 (diagnosis, keyed); Voegelé, 1964: 28 (keyed); Voegelé, 1965: 111 (keyed).

Asolcus reticulatus Delucchi: Delucchi, 1963: 13 (diagnosis, changed to species status); Viktorov, 1964: 1015 (variation); Voegelé, 1964: 28 (keyed); Voegelé, 1965: 111, 112 (keyed).

Asolcus festivae Viktorov, 1964: 1013, 1020 (original description, keyed).

Asolcus reticulatus reticulatus (Delucchi): Viktorov, 1964: 1013 (keyed).

Asolcus reticulatus volgensis Viktorov, 1964: 1013, 1015 (original description, keyed).

Asolcus histani Voegele, 1965: 109, 111, 112 (original description, keyed).

Trissolcus scutellaris (Thomson): Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 414 (keyed); Kozlov, 1968: 199, 210 (diagnosis, lectotype designation, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 510 (keyed); Kozlov, 1978: 634 (description); Kozlov & Kononova, 1983: 105 (description); Kononova, 1995: 95 (keyed); Petrov, 2013: 325 (keyed).

Trissolcus reticulatus reticulatus (Delucchi): Viktorov, 1967: 91 (generic transfer, keyed).

Trissolcus reticulatus volgensis (Viktorov): Viktorov, 1967: 91 (generic transfer, keyed).

Trissolcus reticulatus (Delucchi): Safavi, 1968: 414 (keyed).

Trissolcus festivae (Viktorov) syn. n.: Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 414 (keyed); Kozlov, 1968: 199, 211 (description, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 510 (keyed); Kozlov, 1981: 187 (keyed); Kozlov & Kononova, 1983: 635 (description); Kozlov & Kononova, 1983: 107 (description); Kononova, 1995: 95 (keyed); Ghahari, Buhl & Kocak, 2011: 595 (listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 325 (keyed).

Trissolcus vassilievi (Mayr) syn. n.: Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 414 (keyed); Kozlov, 1968: 199, 208 (description, lectotype designation, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 508 (keyed); Kozlov, 1978: 634 (description); Kozlov & Kononova, 1983: 103 (description); Kononova, 1995: 95 (keyed); Koçak & Kilinçer, 2003: 302, 303 (keyed, description); Ghahari, Buhl & Kocak, 2011: 598 (listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 325 (keyed).

Trissolcus choaspes (Nixon) syn. n.: Kozlov, 1968: 199, 210 (diagnosis, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 507 (keyed); Kozlov, 1978: 634 (description); Kozlov & Kononova, 1983: 102 (description); Johnson, 1992: 625 (cataloged, type information); Petrov, 2013: 325 (keyed); Kononova, 2014: 1423 (keyed); Kononova, 2015: 260 (keyed).

Trissolcus ghorfii (Delucchi & Voegele) syn. n.: Safavi, 1968: 414 (keyed); Voegelé, 1969: 149 (keyed); Kozlov & Lê, 1977: 507 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 99 (description).

Trissolcus histani (Voegele) syn. n.: Safavi, 1968: 414 (keyed); Voegelé, 1969: 149 (keyed); Kozlov & Lê, 1977: 507 (keyed); Kozlov, 1978: 632 (description); Kozlov, 1981: 187 (keyed); Kozlov & Kononova, 1983: 99 (description).

Trissolcus volgensis (Viktorov) syn. n.: Safavi, 1968: 414 (keyed); Kozlov, 1968: 198 (change to species status, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 30 (keyed); Kozlov & Lê, 1977: 506 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 94 (description); Kononova, 1995: 95 (keyed); Petrov, 2013: 325 (keyed).

Lectotype designation

The specimen selected by Kozlov as the lectotype of T. simoni (NHMW 0004) was sent to Mayr by A. Saakow in 1901 according to Mayr (1903). Because this specimen was not part of the original type series it is ineligible to be designated as a lectotype. Szabó (1975) designated a lectotype from Mayr’s syntype series, but did not indicate which of the three specimens on the pin was meant to be the lectotype. To stabilize the nomenclature, we here select one of these specimens (USNMENT01029022b, deposited in NHMW) as the lectotype.

Figures 154. 

Trissolcus scutellaris female lectotype of T. simoni (USNMENT01059022b), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 155–157. 

Trissolcus scutellaris 155 female lectotype of T. simoni (USNMENT01059022b), head, mesosoma, metasoma, dorsal view 156 female (NHMW 0004), head and mesosoma, dorsolateral view 157 female (NHMW 0004), head and mesosoma, ventrolateral view. Scale bars in millimeters.

Figures 158–163. 

Trissolcus scutellaris 158 male, paralectotype of T. vassilliewi (USNMENT01029319), head, anterior view 159 female (USNMENT00896094), head, anterior view 160 female (USNMENT00896103), head, anterolateral view 161 female paralectotype of T. simoni (USNMENT01059022a), head, anterior view 162 female lectotype of T. simoni (USNMENT01059022b), head, anterodorsal view 163 female (USNMENT01059024), head, anterodorsal view. Scale bars in millimeters

Figures 164–167. 

Trissolcus scutellaris 164 female (USNMENT00896335), head and mesosoma, dorsal view 165 female (USNMENT00872172), head and mesosoma, dorsal view 166 female (USNMENT00896206), head and mesosoma, dorsal view 167 female (USNMENT01059030), head and mesosoma, dorsal view. Scale bars in millimeters.

Description

Female body length: 0.89–1.47 mm (n=11). Male body length: 0.91–1.06 mm (n=5). Body color: head, mesosoma, and metasoma black.

Head. Color of radicle: pale brown to dark brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: variably yellow to black. Color of A7A11 in female: pale brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; rugose; areolate. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent; sparse; dense. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: areolate; absent; rugose. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: rugulose; absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate; finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete; well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by open crenulae; formed by small punctures. Mesopleural epicoxal sulcus: formed by open crenulae; formed by small punctures; present as a smooth furrow. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: absent; smooth posteriorly, cells of metapleural sulcus extending posteriorly into horizontal striae. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent; present. Macrosculpture of mesoscutum: rugulose; absent; longitudinally striate posteriorly, otherwise absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout; absent; faintly present posteriorly. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: absent; present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: between 2 and 3 times as long as stigmal vein. Color of legs: coxae dark brown to black, femora yellow to brown, trochanters and distal tarsomeres sometimes brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: absent; present in anterior half of sternite not covered by laterotergites. Setation of S2: present posteromedially.

Diagnosis

Among Palearctic species in the thyantae group, T. scutellaris is closest to T. vesta from which it may be separated by composition of the mesoscutal suprahumeral sulcus, the shape of the vertex and setation of the metapleuron (see couplet 23 of the key).

Associations

emerged from egg of Eurygaster integriceps Puton: [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; emerged from egg of Eurygaster integriceps Puton: [Hemiptera: Heteroptera: Pentatomoidea: Scutelleridae]; egg parasite of sunn pest : [Hemiptera: Heteroptera: Pentatomoidea]

Material examined

Paratypes of T. choaspes: GERMANY: 1 female, 1 male, OSUC 17713, USNMENT0119670 (BMNH). Holotype, female, T. evanescens: AUSTRIA: Wien (Vienna), Lainz, VII-1892, J. J. Kieffer, MNHN 0079 (deposited in MNHN). Paratypes of T. festivae: RUSSIA: 1 female, 1 male, USNMENT00916329, 00916335 (ZIN). Lectotype, female, T. Simoni: PALEARCTIC: no date, USNMENT01059022b (deposited in NHMW). Paralectotypes of T. Simoni: PALEARCTIC: 2 females, USNMENT01059022a, 01059022c (NHMW). Lectotype, female, T. Vassilliewi: PALEARCTIC: no date, G. Mayr, NHMW 0006 (deposited in NHMW). Paralectotypes of T. Vassilliewi: TURKMENISTAN: 3 females, 11 males, 7 unsexed, USNMENT00979609 (CUIC); USNMENT0091666700916675, 01029319, 0105901201059021 (NHMW). Paratypes of T. volgensis: RUSSIA: 1 female, 2 males, USNMENT00916328, 00916336, 00916337 (ZIN). Lectotype, female, T. scutellaris: SWEDEN: Smaland, no date, C. H. Boheman, NHRS-HEVA 000003871 (deposited in NHRS). Other material: (80 females, 15 males, 15 unsexed) BULGARIA: 1 female, USNMENT00872172 (BMNH). CROATIA: 3 females, USNMENT00872407, 00916590, 00916594 (BMNH). FRANCE: 10 females, 4 males, USNMENT00916039, 0091607600916077, 00916086, 00916128, 00916133 (BMNH); USNMENT00896125, 00896230, 0089625200896253, 00896289, 00896329, 0089633500896336 (CNCI). GEORGIA: 8 females, 1 male, 10 unsexed, NHMW 0004, USNMENT0091665600916666, 01059025, 0105902901059034 (NHMW). GREECE: 3 females, USNMENT00916055, 00916065 (BMNH); USNMENT00896233 (CNCI). HUNGARY: 1 female, OSUC 523954 (OSUC). ITALY: 3 females, 1 male, USNMENT00916091 (BMNH); USNMENT0089620600896207, 00896209 (CNCI). MACEDONIA: 1 male, USNMENT00916595 (BMNH). MOROCCO: 13 females, 2 males, USNMENT00916999 (BMNH); USNMENT0089607900896080, 0089608300896086, 0089608900896090, 00896094, 00896103, 0089610600896107, 00896326, 00896337 (CNCI). PORTUGAL: 11 females, 1 male, USNMENT00916187, 00916189, 0091619200916193, 00916195, 00916205, 00916218, 00916220, 00916227, 00916235, 00916240 (BMNH); USNMENT00896304 (CNCI). RUSSIA: 2 females, USNMENT00896069, 00896110 (CNCI). SOUTH KOREA: 7 females, OSUC 542399542402, 542404542406 (OSUC). SPAIN: 4 females, USNMENT00916162, 00916164 (BMNH); USNMENT00896323–00896324 (CNCI). SWEDEN: 3 females, 2 males, NHRS-HEVA 000003872 (NHRS); USNMENT00916069, 00916072, 00916301, 00916318 (BMNH). SYRIA: 1 female, OSUC 17708 (BMNH). TURKEY: 8 females, 3 males, USNMENT00675772–00675780, 00675965–00675966 (USNM). UNKNOWN: 2 females, 5 unsexed, USNMENT00979610 (CUIC); 01059010, 0105902301059024, 0105902601059028 (NHMW).

Comments

Examination of specimens across a wide geographical range revealed that the characters used by previous authors to separate T. volgensis and T. scutellaris, (and T. ghorfii from T. histani) are variable. The microsculpture of the mesoscutellum varies from distinctly present throughout to entirely absent (Figures 164–167). Mesoscutal characters are similarly variable: posteromedial striation, a median mesoscutal carina, and raised rugulose sculpture vary along a continuous gradient from absent to moderately pronounced (Figures 164–167). Kozlov’s 1968 key to Palearctic Trissolcus used the distance between the notauli (referred to as parapsidal grooves) to separate T. ghorfii and T. histani, specifically that the distance between the notauli (measured posteriorly) is more than twice the length of the notauli in the latter species. We examined a series of T. histani that was compared to the holotype and determined by Voegele, and in none of these specimens is this the case. The sculpture of the dorsal frons was not mentioned in the most recent keys by Kononova (2013), Petrov (2013) and Kozlov & Kononova (1983). Were it not for a continuous spectrum of variation, the sculptural development of the frons might separate T. vassilliewi from T. scutellaris. We present a plate of images (Figs 158–163) that illustrate the variation that occurs within this species, even within Mayr’s type series of T. simoni. In the lectotype of T. vassilliewi the frons outside of the antennal scrobe has microsculpture but is otherwise smooth (as in Figure 158). In other specimens, smooth shallow punctures are present along the inner orbits (Fig. 160) or across the dorsal frons (Fig. 161). The punctures may be surrounded by slightly raised rugulae, or, as in the lectotypes of T. simoni and T. volgensis, the rugae are coarse (Figures 162–163). Similarly, the degree of transverse striation directly above the antennal scrobe varies from absent to pronounced. Other structures that vary within this species are the epomial carina, which is always present, but may be robustly or weakly indicated, and the median mesoscutal carina, which is either absent or weakly indicated.

Trissolcus semistriatus (Nees von Esenbeck)

Figures 168–171, 172–177, 178–183; Morphbank 60

Trissolcus artus Kozlov & Lê syn. n.м http://bioguid.osu.edu/xbiod_concepts/3185; Morphbank61

Trissolcus djadetshko (Rjachovskij) syn. n.м http://bioguid.osu.edu/xbiod_concepts/3218; Morphbank62

Trissolcus grandis (Thomson) syn. n.м http://bioguid.osu.edu/xbiod_concepts/3242; Morphbank63

Trissolcus manteroi (Kieffer) syn. n.м http://bioguid.osu.edu/xbiod_concepts/3260; Morphbank64

Trissolcus nigripedius (Nakagawa) syn. n.м http://bioguid.osu.edu/xbiod_concepts/3274; Morphbank65; Neotype designation. Watanabe (1951) determined specimen EIHU 0003 to be T. nigripedius based on the original description of Nakagawa. It is on this basis that we designate EIHU 0003 as the neotype for this species.

Trissolcus pentatomae (Rondani) syn. n.м http://bioguid.osu.edu/xbiod_concepts/3285; Morphbank66

Teleas semistriatus Nees von Esenbeck, 1834: 290 (original description); Ratzeburg, 1852: 182 (description).

Telenomus frontalis Thomson, 1860: 170 (original description, synonymized by Kozlov (1968)); Kozlov, 1968: 214 (junior synonym of Trissolcus grandis (Thomson)).

Telenomus grandis Thomson, 1860: 169 (original description).

Telenomus nigripes Thomson, 1860: 170 (original description, synonymized by Kozlov (1968)); Kozlov, 1968: 214 (junior synonym of Trissolcus grandis (Thomson)); Fergusson, 1984: 230 (lectotype designation).

Telenomus Nigrita Thomson, 1860: 172 (original description, synonymized by Kozlov (1968)); Kozlov, 1968: 214 (junior synonym of Trissolcus grandis (Thomson)).

Telenomus ovulorum Thomson, 1860: 171 (original description, synonymized by Mayr (1879)); Mayr, 1879: 704 (junior synonym of Telenomus semistriatus (Nees von Esenbeck)).

Telenomus semistriatus (Nees von Esenbeck): Thomson, 1860: 171 (description, generic transfer); Mayr, 1879: 699, 701, 704 (description, synonymy, keyed).

Teleas (?) PentatomaeRondani: Rondani, 1874: 135 (nomen nudum).

Teleas pentatomae Rondani, 1877: 199 (original description).

Telenomus pentatomae (Rondani): Dalla Torre, 1898: 518 (generic transfer).

Telenomus nigritus Thomson: Dalla Torre, 1898: 517 (emendation).

Asolcus nigripedius Nakagawa, 1900: 17 (original description); Watanabe, 1951: 21, 26 (description, type information, keyed); Watanabe, 1954: 22 (keyed).

Telenomus Manteroi Kieffer, 1909: 268 (original description).

Aphanurus Frontalis (Thomson): Kieffer, 1912: 81 (description, generic transfer).

Aphanurus Grandis (Thomson): Kieffer, 1912: 76 (description, generic transfer).

Aphanurus Nigrita (Thomson): Kieffer, 1912: 79 (description, generic transfer).

Aphanurus nigripes (Thomson): Kieffer, 1912: 75 (description, generic transfer).

Aphanurus Semistriatus (Nees von Esenbeck): Kieffer, 1912: 74 (description, generic transfer).

Liophanurus Pentatomae (Rondani): Kieffer, 1912: 69 (description, generic transfer).

Aphanurus Manteroi (Kieffer): Kieffer, 1912: 84 (description, generic transfer).

Liophanurus pentatomae (Rondani): Kieffer, 1926: 71 (description).

Microphanurus frontalis (Thomson): Kieffer, 1926: 91, 103 (description, generic transfer, keyed).

Microphanurus grandis (Thomson): Kieffer, 1926: 91, 99 (description, generic transfer, keyed); Debauche, 1947: 256 (diagnosis).

Microphanurus nigripes (Thomson): Kieffer, 1926: 91, 98 (description, generic transfer, keyed).

Microphanurus nigritus (Thomson): Kieffer, 1926: 91, 100 (description, generic transfer, keyed).

Microphanurus manteroi (Kieffer): Kieffer, 1926: 91, 102 (description, generic transfer, keyed); Boldaruyev, 1969: 163, 170 (description, keyed).

Microphanurus semistriatus (Nees von Esenbeck): Kieffer, 1926: 91, 97 (description, generic transfer, keyed); Nixon, 1939: 131, 134 (description, keyed); Meier, 1940: 80 (description, keyed); Rjachovskij, 1959: 84 (keyed).

Microphanurus alexeevi Meier, 1949: 114 (original description, not seen: reference from Kozlov (1963), synonymized with Asolcus semistriatus (Nees von Esenbeck) by Kozlov (1963)); Rjachovskij, 1959: 83 (keyed); Kozlov, 1963: 295 (junior synonym of Asolcus semistriatus (Nees von Esenbeck)).

Microphanurus schtepetelnikovae Meier, 1949: 114 (original description, not seen: reference from Kozlov (1963), synonymized with Asolcus semistriatus (Nees von Esenbeck) by Kozlov (1963)); Kozlov, 1963: 295 (junior synonym of Asolcus semistriatus (Nees von Esenbeck)).

Asolcus semistriatus (Nees von Esenbeck): Masner, 1959: 376 (diagnosis, variation); Delucchi, 1961: 44, 59 (diagnosis, taxonomic status, keyed); Kozlov, 1963: 295 (synonymy); Viktorov, 1964: 1013, 1020 (variation, keyed); Kochetova, 1966: 558 (description of immature stages); Javahery, 1968: 419 (keyed); Voegelé, 1969: 150 (keyed); Szabó, 1976: 176, 178 (description, neotype designation, keyed).

Microphanurus djadetshko Rjachovskij, 1959: 84, 87 (original description, keyed).

Microphanurus stschepetilnicovae Meier: Rjachovskij, 1959: 83 (keyed, spelling error).

Asolcus grandis (Thomson): Delucchi, 1961: 44, 60 (description, keyed); Voegelé, 1964: 28 (keyed); Javahery, 1968: 419 (keyed); Voegelé, 1969: 150 (keyed).

Asolcus nigribasalis Voegelé, 1962: 155 (original description); Voegelé, 1964: 28 (keyed); Voegelé, 1965: 96, 108 (variation, diagnosis, keyed); Voegelé, 1969: 151 (junior synonym of Asolcus djadetshko (Rjachovskij)).

Asolcus djadetshko (Rjachovskij): Viktorov, 1964: 1015, 1021 (description, generic transfer, removed from synonymy with Telenomus scutellaris Thomson, keyed).

Trissolcus nigripedius (Nakagawa) syn. n.: Masner, 1964: 146 (generic transfer); Ryu & Hirashima, 1984: 37, 56 (description, keyed); He et al., 2004: 321 (description).

Trissolcus djadetshko (Rjachovskij): Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 415 (keyed); Kozlov, 1968: 200 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 512 (keyed); Kozlov, 1978: 636 (description); Kozlov, 1981: 187 (keyed); Kozlov & Kononova, 1983: 115 (description); Kononova, 1995: 96 (keyed); Koçak & Kilinçer, 2000: 171 (description, diagnosis, new distribution record for Turkey); Koçak & Kilinçer, 2003: 303, 313 (keyed, description); Ghahari, Buhl & Kocak, 2011: 595 (listed); Petrov, 2013: 326 (keyed).

Trissolcus grandis (Thomson): Viktorov, 1967: 91 (generic transfer, keyed); Safavi, 1968: 416 (keyed); Kozlov, 1968: 200, 214 (description, lectotype designation, synonymy, keyed); Viggiani & Mineo, 1974: 156, 160, 161 (description, keyed); Kozlov & Lê, 1977: 512 (synonymy, keyed); Kozlov, 1978: 636 (description); Kozlov, 1981: 187 (keyed); Kozlov & Kononova, 1983: 110 (description); Kononova, 1995: 96 (keyed); Doganlar, 2001: 112 (description); Koçak & Kilinçer, 2003: 302, 307 (keyed, description); Buhl & O’Connor, 2010: 154 (distribution); Ali, 2011: 10 (keyed); Ghahari, Buhl & Kocak, 2011: 596 (host association, listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 326 (keyed).

Trissolcus manteroi (Kieffer) syn. n.: Kozlov, 1968: 199 (keyed); Fabritius, 1972: 31 (keyed); Bin, 1974: 462 (type information); Kozlov & Lê, 1977: 514 (keyed); Kozlov, 1978: 636 (description); Kozlov & Kononova, 1983: 117 (description); Koçak & Kilinçer, 2000: 174 (description, diagnosis, new distribution record for Turkey); Koçak & Kilinçer, 2003: 302, 310 (keyed, description of female); Koçak & Kodan, 2006: 41 (description of male); Ghahari, Buhl & Kocak, 2011: 596 (listed); Kononova, 2014: 1424 (keyed); Kononova, 2015: 262 (keyed).

Asolcus nixomartini Javahery, 1968: 419, 429 (original description, keyed, synonymized by Kozlov & Lê (1977)); Kozlov & Lê, 1977: 512 (junior synonym of Trissolcus grandis (Thomson)).

Trissolcus semistriatus (Nees von Esenbeck): Safavi, 1968: 416 (keyed); Kozlov, 1968: 200 (keyed); Fabritius, 1972: 31 (keyed); Kozlov & Lê, 1977: 512 (keyed); Kozlov, 1978: 636 (description); Kozlov & Kononova, 1983: 113 (description); Graham, 1984: 92 (variation); Kononova, 1995: 96 (keyed); Koçak & Kilinçer, 2003: 302, 305 (keyed, description); Ali, 2011: 10 (keyed); Ghahari, Buhl & Kocak, 2011: 597 (listed); Guz, Kocak & Kilincer, 2013: 87 (description, phylogenetic relationships); Petrov, 2013: 326 (keyed).

Asolcus silwoodensis Javahery, 1968: 419, 425 (original description, keyed, synonymized by Kozlov & Lê (1977)); Kozlov & Lê, 1977: 512 (junior synonym of Trissolcus grandis (Thomson)).

Asolcus djadestshko (Rjachovskij): Voegelé, 1969: 151 (synonymy, keyed, spelling error).

Trissolcus pentatomae (Rondani): Bin, 1974: 463 (generic transfer, lectotype designation).

Trissolcus artus Kozlov & Lê syn. n., 1977: 512, 519 (original description, keyed); Kozlov, 1978: 636 (description); Kozlov & Kononova, 1983: 112 (description); Kononova, 1995: 96 (keyed).

Trissolcus nigribasalis (Voegelé): Kozlov & Lê, 1977: 518 (keyed); Kozlov, 1978: 637 (description); Kozlov & Kononova, 1983: 124 (description).

Trissolcus nigripes (Thomson): Fergusson, 1978: 120 (generic transfer).

Trissolcus nixomartini (Javahery): Fergusson, 1978: 120 (generic transfer); Fergusson, 1984: 230 (type information).

Trissolcus silwoodensis (Javahery): Fergusson, 1978: 120 (generic transfer); Fergusson, 1984: 230 (type information).

Description

Female body length: 0.99–1.33 mm (n=20). Male body length: 0.94–1.20 mm (n=20). Body color: head, mesosoma, and metasoma black.

Figures 168–171. 

T. semistriatus 168 female (USNMENT01109059), head, mesosoma, metasoma, dorsal view 169 female (USNMENT01109059), head and mesosoma, dorsolateral view 170 female neotype (NHMW 0007A), head and mesosoma, anteroventral view 171 female (USNMENT01109059), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Figures 172–177. 

T. semistriatus 172 female (USNMENT00896318), head, anterior view 173 female (USNMENT00896062), head, anterior view 174 female (USNMENT00896262), head, anterior view 175 female (USNMENT00896226), head, anterior view 176 female (USNMENT00896205), head, anterior view 177 female (USNMENT00896254), head, anterior view. Scale bars in millimeters.

Figures 178–183. 

T. semistriatus 178 female (USNMENT00896318), head, mesosoma, metasoma, lateral view 179 female (USNMENT00896062), head, mesosoma, metasoma, lateral view 180 female (USNMENT00896262), head, mesosoma, metasoma, lateral view 181 female (USNMENT00896226), head, mesosoma, metasoma, ventrolateral view 182 female (USNMENT00896254), head, mesosoma, metasoma, ventrolateral view 183 female (USNMENT00896205), head, mesosoma, metasoma, ventrolateral view. Scale bars in millimeters.

Head. Color of radicle: dark brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: variably yellow to black. Color of A7A11 in female: pale brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent; present. Shape of ventral gena in lateral view: narrow. Genal carina: present only at base of mandible. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse; moderately dense. Punctation of lateral frons: absent; sparse. Sculpture directly ventral to preocellar pit: dorsoventrally fluted. Macrosculpture of lateral frons: horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half ofpronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: absent. Shape of dorsal margin of anterior lobe of axillar crescent: acute. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, femora and tibia yellow to dark brown, trochanters and tarsi yellow to pale brown. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 1. Setation of laterotergite 1: absent; present. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present in a transverse line and along lateral margin. Setation of laterotergite 2: present.

Diagnosis

Trissolcus semistriatus is most similar to T. basalis from which it can be separated by the complete netrion sulcus, distinct and regular cells of the postacetabular sulcus and, in most cases, by the distinct cells of the mesoscutal humeral sulcus (always indicated by a smooth furrow in T. basalis and very rarely so in T. semistriatus). It is also similar to T. elasmuchae from which it can be separated by the absence of a well defined paracoxal sulcus in the ventral portion of the metapleuron.

Comments

The immense geographical distribution of Trissolcus semistriatus, in combination with its capacity for variation in size, color, and surface sculpture (Figures 172183) have resulted in numerous redescriptions of this species. It is by examining specimens across the Palearctic land mass that we have been able to document intermediate forms that connect phenotypes previously treated as discreet. The general trend is that specimens from higher latitudes tend to be larger, darker, and have more robust sculpture on the frons. The sculpture on the frons in Trissolcus semistriatus, T. basalis and T. elasmuchae have similar patterns when rugae are present and they share the same spectrum of sculptural development, varying from coarsely rugose to absent. Preliminary results on a study of phenotypic plasticity in Trissolcus indicate that color and size are directly influenced by temperature and host species. Species with distributions as large as T. semistriatus, T. basalis, and T. elasmuchae are likely to be oligophagous or polyphagous, increasing the degree of phenotypic variation that can occur.

Material examined

Neotype, female, T. semistriatus: PALEARCTIC: no date, NHMW 0007A (deposited in NHMW). Neoparatype: PALEARCTIC: 1 female, NHMW 0007B (NHMW). Holotype, female, T. artus: RUSSIA: Altay Terr., Kosh-Agach, 6.VII.1964, Kozlov, ZMAS 0119 (deposited in ZIN). Paratype of T. artus: RUSSIA: 1 female, USNMENT00916276 (ZIN). Paratypes of T. djadetshko: UKRAINE: 5 females, USNMENT00954012–00954016 (ZIN). Lectotype, female, T. grandis: SWEDEN: Dalarna (Dalecarlia), no date, C. H. Boheman, NHRS-HEVA 000003869 (deposited in NHRS). Holotype, female, T. manteroi: ITALY: Liguria, Genoa, 9.VIII.1997, G. Mantero, MCSN 0013 (deposited in MCSN). Lectotype, male, T. pentatomae: ITALY: no date, MZUF 0001 (deposited in MZUF). Other material: (417 females, 86 males, 7 unsexed) AUSTRIA: 18 females, USNMENT0091610000916104, 0091629000916292, 0091629500916299, 0091632000916324 (BMNH). BULGARIA: 2 females, USNMENT00872173, 00872174 (BMNH). CHINA: 19 females, UCRC ENT 142621, 142628, 142631, 142641, 142649, 142657, 142666, 142672, 142677, 142707, 142737, 142744, 143823, 143841, 143857, 143859, 143882, 143909 (UCRC); USNMENT00979424 (USNM). CROATIA: 11 females, 1 male, USNMENT0087240500872406, 0087240800872409, 0091658800916589, 00916598 (BMNH); USNMENT00896155–00896156 (CNCI); USNMENT00675788–00675790 (USNM). CYPRUS: 69 females, 6 males, USNMENT0091650000916513, 0091651500916523, 00916525, 0091652700916528, 0091653000916549, 0091655100916552, 00916554, 0091655700916579, 00916581, 0091658300916584 (BMNH). CZECH REPUBLIC: 2 females, USNMENT00896164, 00896217 (CNCI). FRANCE: 66 females, 14 males, 1 unsexed, USNMENT00916074, 00916078, 00916080, 00916082, 0091608700916089, 00916115, 00916117, 00916123, 00916126, 00916130, 00916136, 00916139, 00916141, 00916148, 00916150 (BMNH); USNMENT00896066, 0089612300896124, 0089612600896130, 0089613200896133, 0089623100896232, 0089623500896236, 0089624100896246, 00896251, 0089625400896259, 0089626100896288, 0089629000896292, 00896294, 0089629800896299, 0089630200896303, 00896338 (CNCI). GREECE: 16 females, 2 males, USNMENT00916054, 0091605600916058, 00916062, 00916064, 00916499, 0091658500916586 (BMNH); USNMENT0089606100896065, 00896131, 0089624700896248, 00896250 (CNCI). HUNGARY: 7 females, USNMENT00896211–00896214, 00896219 (CNCI); OSUC 523955523956 (OSUC). IRAN: 11 females, 6 males, 2 unsexed, USNMENT0091647500916476, 00916480, 00916482, 00916485, 00916487 (BMNH); OSUC 145630, USNMENT0089622500896229, 0089623800896240 (CNCI); OSUC 556692556693 (HMIM); OSUC 556691, 76299 (OSUC). IRAQ: 4 females, USNMENT00916488–00916491 (BMNH). IRELAND: 1 female, 1 unsexed, USNMENT00916138, 00916425 (BMNH). ISRAEL: 1 female, USNMENT00896141 (CNCI). ITALY: 17 females, USNMENT00916042, 0091609200916099, 00916285, 00916597 (BMNH); USNMENT00896201–00896203, 00896205, 00896208, 00896220 (CNCI). JAPAN: 3 females, OSUC 542382 (CNCI); OSUC 70461–70462 (OSUC). KAZAKHSTAN: 1 female, USNMENT00916599 (BMNH). LEBANON: 1 female, USNMENT00916602 (BMNH). MACEDONIA: 1 female, USNMENT00916596 (BMNH). MOROCCO: 18 females, 10 males, 1 unsexed, USNMENT0091600000916009, 0091699000916994, 00916998 (BMNH); USNMENT00896087, 00896093, 00896333 (CNCI); USNMENT01059170–01059172, 01059174–01059177, 01059179–01059181 (MNHN). PORTUGAL: 21 females, 2 males, 1 unsexed, USNMENT00916170, 0091617200916175, 00916177, 00916185, 0091619000916191, 0091620100916203, 00916206, 0091621000916213, 0091621500916217, 0091622300916234, 0091623600916237 (BMNH). RUSSIA: 2 females, USNMENT00896111 (CNCI); UCRC ENT 133647 (UCRC). SERBIA: 4 females, 3 males, USNMENT00916592 (BMNH); USNMENT00675782–00675787 (USNM). SLOVAKIA: 1 female, USNMENT00916603 (BMNH). SOUTH KOREA: 2 males, OSUC 542387, 542390 (OSUC). SPAIN: 16 females, 1 male, USNMENT00916151, 0091615400916157, 0091615900916161, 00916166, 0091616800916169, 00916179 (BMNH); USNMENT00896153, 00896320–00896322, 00896325 (CNCI). SWEDEN: 10 females, 5 males, USNMENT0091604600916047, 0091605100916053, 0091606700916068, 00916070, 0091630500916311 (BMNH); USNMENT00896067 (CNCI). SWITZERLAND: 20 females, 4 males, USNMENT00896297 (CNCI); 0091695200916972, 0110905801109059 (USNM). TURKEY: 49 females, 27 males, OSUC 17730, USNMENT0091603200916034, 00916071 (BMNH); USNMENT0089619000896193, 0089619500896199, 00896223, 00896319 (CNCI); USNMENT0067579100675810, 0067581200675846, 0067598000675984 (USNM). UNITED KINGDOM: 26 females, 3 males, 1 unsexed, USNMENT00916246, 00916410, 00916419, 0091642200916424, 00916429, 0091643100916437, 0091643900916441, 0091644400916454, 00916456 (BMNH); USNMENT00896318 (CNCI).

Trissolcus stoicus (Nixon)

Figures 184, 185–188; Morphbank 67

Microphanurus stoicus Nixon, 1938: 124, 135 (original description, keyed); Nixon, 1943: 138 (keyed); Risbec, 1950: 569 (keyed).

Trissolcus stoicus (Nixon): Masner, 1965: 128 (type information, generic transfer).

Telenomus stoicus (Nixon): Lê, 1981: 13 (generic transfer, keyed).

Description

Female body length: 0.93–1.11 mm (n=2). Body color: head, mesosoma, and metasoma black.

Figure 184. 

Trissolcus stoicus, female (OSUC 285262), head, mesosoma, metasoma, dorsal view. Scale bar in millimeters.

Figures 185–188. 

Trissolcus stoicus 185 female holotype (B.M. TYPE HYM. 9.314), head and mesosoma, lateral view 186 female holotype (B.M. TYPE HYM. 9.314), head and mesosoma, dorsolateral view 187 female (OSUC 285262), mesosoma, posterolateral view 188 female (OSUC 285262), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow. Color of A7A11 in female: pale brown; yellow, becoming darker distally. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: weakly transversely strigose. Preocellar pit: absent. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Macrosculpture of lateral frons: weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present medially, absent laterally. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: dorsoventrally bisected by deep furrow. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete. Sculpture of femoral depression: rugulose and pustulate along ventral half of mesopleural carina. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: absent. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: coarsely rugose. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: cells of metapleural sulcus extending posteriorly into rugae. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of elongate cells. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: present. Macrosculpture of mesoscutum: coarsely areolate. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Notaulus: absent; indicated only at posterior margin of mesoscutum. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: rugose throughout. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent; present. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: mostly smooth, with furrow along ventral margin. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae dark brown to black, trochanters and femora yellow to brown, elsewhere yellow to orange. Anteromedial portion of metasomal depression: antero-posteriorly striate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: present. Longitudinal striation of T2: present laterally, absent medially. Setation of T2: present at posterior termination of striae. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present laterally and in anterior half of median third. Setation of S2: present posteromedially.

Diagnosis

Trissolcus stoicus can be distinguished from most species treated here by having 6 clypeal setae and a complete hyperoccipital carina, a combination which is otherwise known to us only from T. carinifrons. These are otherwise dissimilar species and can be separated by the sculpture of the gena (coarse in T. carinifrons and smooth in T. stoicus) and the shape of the foveae that form the mesepimeral sulcus (circular in T. carinifrons and elongate in T. stoicus). The form of the metanotal trough, in which the cells are present at the bottom of a narrow furrow is shared with T. exerrandus, from which it may separated by the presence of dense setation on the eyes in the latter. The distribution of striae on T2 in T. stoicus, in which striae are present laterally and absent medially, is unusual for Trissolcus, at least in our experience. However, this character is not fully visible in the holotype specimen and we are thus hesitant to use it for identification with a sample size of only three specimens, particularly when a wealth of other characters exists to identify this species.

Associations

emerged from egg of Coffea Linnaeus: [Rubiales: Rubiaceae]; emerged from egg of Hemiptera : [Hemiptera]; emerged from egg of Lepidoptera : [Lepidoptera]

Material examined

Lectotype, female, M. stoicus: MALAYSIA: Pahang St., Malay Peninsula, Pahang, 16.VI.1932, G. H. Corbett, B.M. TYPE HYM. 9.314 (deposited in BMNH). MALAYSIA: 1 female, UCRC ENT 297004 (UCRC). THAILAND: 1 female, OSUC 285262 (OSUC).

Lectotype designation

We here designate specimen B.M. TYPE HYM. 9.314 (deposited in BMNH) to be the lectotype of this species.

Comments

Trissolcus stoicus exhibits two characters that suggest in may be an intermediate form between the flavipes and thyantae species groups. The hyperoccipital carina and setation of the first laterotergite ally it with the flavipes group. However, the presence of 6 clypeal setae and 2 large setae in the posteroventral metapleuron suggest relation to the thyantae group.

Trissolcus tersus

Figures 189–190, 191–193; Morphbank 68

Trissolcus tersus Lê, 1985: 166 (original description); Johnson, 1992: 638 (cataloged, type information); Lê, 2000: 312, 320 (description, keyed, type information).

Description

Female body length: 0.99–0.81 mm (n=2). Body color: head, mesosoma, and metasoma black.

Figures 189–190. 

Trissolcus tersus female (EMBT ENT 0001839) 189 head, mesosoma, metasoma, lateral view 190 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 191–193. 

Trissolcus tersus female (EMBT ENT 0001839) 191 head, mesosoma, anterodorsal view 192 head and mesosoma, ventrolateral view 193 head, anterior view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: basal A1 and distal A2 yellow, elsewhere pale brown. Color of A7A11 in female: dark brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as elongate striation. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: present only at base of mandible. Malar striae: present. Sculpture of malar sulcus: smooth. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: absent. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: lateral ocellus and eye without continuous scleritic separation. Hyperoccipital carina: complete. Anterior margin of occipital carina: simple.

Mesosoma. Epomial carina: absent. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Posterior pronotal sulcus: absent. Number of episternal foveae: 3. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: weakly transversely wrinkled. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: smooth area large, surrounded by well defined sulci. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae dark brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: smooth.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: absent. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: absent.

Diagnosis

Trissolcus tersus can be identified by the line of foveae along the dorsal margin of the mesopleural carina (Figures 189, 192), a character known to us so only from this species. The setation of the posteroventral metapleuron (Figure 189) and the presence of facial and malar striae (Figure 193) are also useful for its identification.

Link to distribution map

Material examined

Holotype, female: VIETNAM: Hoa Binh Prov., rice field, Thuong Tien, 11.XI.1978, IEBR 0043 (deposited in IEBR). Other material: (9 females) INDONESIA: 1 female, CNC424618 (CNCI). THAILAND: 7 females, CNC424502–424505, CNC424607, CNC424619 (CNCI); EMBT ENT 0001839 (USNM).

Trissolcus trophonius (Nixon)

Figures 194–195, 196–198; Morphbank 69

Microphanurus trophonius Nixon, 1938: 123, 127 (original description, keyed); Nixon, 1943: 136, 140 (diagnosis, keyed); Risbec, 1950: 568 (keyed).

Trissolcus trophonius (Nixon): Masner, 1965: 128 (type information, generic transfer).

Description

Female body length: 0.94–1.08 mm (n=2). Body color: head, mesosoma, and metasoma black.

Figures 194–195. 

Trissolcus trophonius, female holotype (B.M. TYPE HYM. 9.315) 194 head, mesosoma, metasoma, lateral view 195 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 196–198. 

Trissolcus trophonius, female holotype (B.M. TYPE HYM. 9.315) 196 head, anterior view 197 head and mesosoma, anterolateral view 198 head and mesosoma, dorsolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: equal to or greater than width of clypeus; less than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally; basal A1 and distal A2 yellow, elsewhere pale brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 4. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: smooth. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: dorsoventrally fluted. Macrosculpture of lateral frons: weakly horizontally striate, striae of antennal scrobe extending to lateral frons. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: absent. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma: Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: uncertain, percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by open crenulae. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: rugose. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indicated by a line of elongate cells. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; weakly rugulose anteriorly, otherwise absent. Pattern of mesoscutal microsculpture: effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: absent. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: absent. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about 1.5 times as long as stigmal vein. Color of legs: coxae brown, legs elsewhere yellow; coxae and femora brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anteromedial portion of the tergite. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: present posteromedially.

Diagnosis

The presence of 4 clypeal setae and posteriorly effaced sculpture of the mesoscutum are shared with T. plautiae, T. japonicus, and T. kozlovi. It may separated from them by the dorsoventral groove that extends from the median ocellus to the antennal scrobe, the effaced sculpture lateral to this groove, and the striation on T2 that is limited to the anteromedial portion of the tergite.

Lectotype designation

We here designate specimen B.M. TYPE HYM. 9.315 (deposited in BMNH) to be the lectotype of this species.

Material examined

Lectotype, female, M. trophonius: INDONESIA: Sumatera Utara Prov., Asahan Regency, Negaga Estate, 50–60m, 10.XII.1934, reared, Schneider, B.M. TYPE HYM. 9.315 (deposited in BMNH). Other material: THAILAND: 1 female, UCRC ENT 297005 (UCRC).

Trissolcus tumidus (Mayr)

Figures 8, 199–202, 203–205, 206–208; Morphbank 70

Trissolcus cephalotes Kozlov & Lê syn. n.м http://bioguid.osu.edu/xbiod_concepts/3198; Morphbank71

Trissolcus delucchii Kozlov syn. n.м http://bioguid.osu.edu/xbiod_concepts/3212; Morphbank72

Trissolcus pierrot Mineo, O’Connor & Ashe syn. n.м http://bioguid.osu.edu/xbiod_concepts/274651; Morphbank73

Telenomus tumidus Mayr, 1879: 699, 703 (original description, keyed).

Aphanurus Tumidus (Mayr): Kieffer, 1912: 74 (description, generic transfer).

Microphanurus tumidus (Mayr): Kieffer, 1926: 91, 96 (description, generic transfer, keyed).

Asolcus tumidus (Mayr): Delucchi, 1961: 44, 52 (description, keyed); Voegelé, 1964: 28 (keyed).

Trissolcus delucchii Kozlov syn. n., 1968: 198, 203 (original description, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 30 (keyed); Kozlov & Lê, 1976: 658 (keyed); Kozlov & Lê, 1977: 506 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 90 (description); Ryu & Hirashima, 1984: 36, 41 (description, keyed); Ghahari, Buhl & Kocak, 2011: 595 (listed, note on incorrect Lepidopteran association from Modarres Awal (1997)).

Trissolcus tumidus (Mayr): Safavi, 1968: 415 (keyed); Kozlov, 1968: 198, 204 (diagnosis, keyed); Voegelé, 1969: 149 (keyed); Fabritius, 1972: 30 (keyed); Kozlov & Lê, 1976: 658 (keyed); Kozlov & Lê, 1977: 504 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 89 (description); Ryu & Hirashima, 1984: 36, 42 (description, keyed); Kononova, 1995: 95 (keyed); Ghahari, Buhl & Kocak, 2011: 598 (listed); Petrov, 2013: 324 (keyed).

Trissolcus cephalotes Kozlov & Lê syn. n., 1976: 658, 661 (original description, keyed); Kozlov & Lê, 1977: 506 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 93 (description).

Trissolcus pierrot Mineo, O’Connor & Ashe syn. n., 2010: 27 (original description, placement within Trissolcus pierrot group); Kononova, 2014: 1421 (keyed); Kononova, 2015: 258 (keyed).

Description

Female body length: 0.94–1.53 mm (n=21). Male body length: 0.89–1.03 mm (n=3). Body color: head, mesosoma, and metasoma black.

Figures 199–202. 

Trissolcus tumidus 199 female holotype (NHMW 0002), head, mesosoma, metasoma, lateral view 200 female holotype of T. cephalotes syn. n. (ZMAS 0142), head, mesosoma, metasoma, dorsal view 201 female holotype of T. cephalotes syn. n. (ZMAS 0142), head, anterior view 202 female (USNMENT00979425), mesopleuron, lateral view. Scale bars in millimeters.

Figures 203–205. 

203 Trissolcus tumidus, female holotype (NHMW 0002), head and antennae, anterolateral view 204 Trissolcus cephalotes, female holotype (ZMAS 0142), head and antenna, anterolateral view 205 Trissolcus delucchii, female holotype (ZMAS 0148). Scale bars in millimeters.

Figures 206–208. 

Trissolcus tumidus 206 female paratype of T. delucchii syn. n. (USNMENT00916636), head and mesosoma, dorsal view 207 female (USNMENT00979425), mesosoma, dorsal view 208 female (OSUC 523869), mesosoma, dorsal view. Scale bars in millimeters.

Head. Color of radicle: yellow; brown; dark brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: variably yellow to black. Color of A7A11 in female: dark brown to black. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: bulging; narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: effaced medially. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: finely rugulose; striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum; weakly differentiated from sculpture of dorsal pronotum. Location of pronotal suprahumeral sulcus: posterior half of pronotum. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: distinctly separate from mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression; present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by open crenulae; formed by small punctures. Mesopleural epicoxal sulcus: formed by large cells; formed by small punctures. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: absent. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; longitudinally striate posteromedially, otherwise absent; longitudinally striate posteriorly, otherwise absent. Pattern of mesoscutal microsculpture: uniform throughout; effaced posteriorly lateral of notaulus. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: rugulose punctate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown to black, femora and tibia yellow to dark brown, trochanters and tarsi yellow to pale brown. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present throughout anterior half of tergite; present in anteromedial portion of the tergite. Setation of T2: uncertain, present throughout posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: absent. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

Trissolcus tumidus is unusual among Palearctic species in the flavipes group because the episternal foveae are clearly separated from the cells of the postacetabular sulcus and the mesopleural pit, a character shared with T. gonopsidis and T. nycteridaner. In most species of the flavipes group these foveae form a more or less continuous line between the postacetabular sulcus and the mesopleural pit. This character, in combination with a glabrous first laterotergite and a well-defined and ventrally expanded orbital furrow, separate it from other species of the flavipes group.

Material examined

Holotype, female, T. tumidus: AUSTRIA: Lower Austria St., Markt Piesting, 1872, Tschek, NHMW 0002 (deposited in NHMW). Holotype, female, T. cephalotes: TAJIKISTAN: Dushanbe Indep. City, Pavlovsky Institute of Zoology and Parasitology (IZIP), trees, Dushanbe, 18.X.1961, V. Triapitsyn, ZMAS 0142 (deposited in ZIN). Paratype of T. cephalotes: TAJIKISTAN: 1 female, USNMENT00916629 (ZIN). Holotype, female, T. delucchii: ARMENIA: Erevan City, Yerevan (Erevan), 23.VI.1959, A. Avetjan, ZMAS 0148 (deposited in ZIN). Paratypes of T. delucchii: (4 females) ARMENIA: 3 females, USNMENT00916635–00916637 (ZIN). Holotype, female, T. pierrot: ITALY: Genova, San Lorenzo di Cas., VIII-1936, F. Solari, MCSN 0014 (deposited in MCSN). Other material: (34 females, 5 males) AFGHANISTAN: 1 male, USNMENT00916640 (ZIN). CHINA: 29 females, 4 males, USNMENT00979423, 0097942500979429, 0097962000979646 (USNM). RUSSIA: 1 female, OSUC 523869 (OSUC). SOUTH KOREA: 1 female, USNMENT00896013 (CNCI). TAJIKISTAN: 1 female, USNMENT00916641 (ZIN). TURKEY: 2 females, USNMENT00916030–00916031 (BMNH). TURKMENISTAN: 2 females, USNMENT00916630, 00916639 (ZIN).

Comments

Petrov (2013) used the longitudinal striation of the mesoscutum between the notauli and the smooth and shining frons between the antennal scrobe and median ocellus as identifying characters. In the lectotype of T. tumidus, longitudinal striation is present between the notauli but the frons is not entirely smooth and shining. Our examination reveals that both of these characters are size dependent. Large specimens tend to have longitudinal striation between the notauli, pustulate microsculpture throughout the mesoscutum (Figure 208), microsculpture throughout the frons (Figures 203, 205), and a gena that does not sharply recede behind the compound eye. Smaller specimens have less development of mesoscutal sculpture (Figures 206–207), the frons is smooth and shining (Figure 204), there is no striation between the notauli (Figure 206), and the gena recedes sharply behind the compound eye.

Trissolcus vesta Kozlov & Lê

Figures 209–212; Morphbank 74

Trissolcus vesta Kozlov & Lê: Kozlov & Lê, 1977: 507 (keyed); Kozlov & Lê, 1977: 1253 (original description); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 97 (description); Kononova, 1995: 95 (keyed).

Description

Female body length: 0.85–0.88 mm (n=2). Body color: head and mesosoma black, metasoma black to reddish brown.

Figures 209–212. 

Trissolcus vesta female paratype (USNMENT00916613) 209 head, mesosoma, metasoma, lateral view 210 head, mesosoma, metasoma, dorsal view 211 head and mesosoma, anterodorsal view 212 mesosoma, dorsolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: absent. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: smooth. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: absent. Setation of lateral frons: sparse. Punctation of lateral frons: absent. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: simple. Mesopleural carina: complete. Sculpture of femoral depression: faintly rugulose. Patch of striae at posteroventral end of femoral depression: present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: present as a smooth furrow. Mesopleural epicoxal sulcus: formed by small punctures; present as a smooth furrow. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: absent; smooth posteriorly, cells of metapleural sulcus extending posteriorly into horizontal striae. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: absent. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: faintly present posteriorly. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present. Form of metascutellum: single row of cells. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about three times as long as stigmal vein. Color of legs: coxae and femora brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present in a transverse line posteriorly. Setation of laterotergite 2: present. Striation of S2: absent. Setation of S2: sparsely present throughout area not covered by laterotergite.

Associations

emerged from egg of Capnoda nigroaenea Jakovlev: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female: RUSSIA: Altay Terr., Kurayskiy Mts., Kosh-Agach, 2400m, 22.VII.1964, Kozlov, ZMAS 0134 (deposited in ZIN). Paratypes: RUSSIA: 2 females, USNMENT00916613, 00916648 (ZIN).

Diagnosis

Trissolcus vesta is likely to be one of the more difficult species to identify due to the subtlety of some of its diagnostic characters, specifically the posterior vertex and the setation in the ventral metapleuron. In addition to metapleuron setation it has the suite of characters that delimit the thyantae group: notauli, 6 clypeal setae, and well defined episternal foveae that extend from the mesopleural pit to the dorsal limit of the acetabular carina. In T. vesta the posterior vertex is medially depressed, creating a triangular area in dorsal view that serves as an important diagnostic character that was recognized by Kozlov (1978), but also one that is best interpreted in the context of other species. Of additional use are the mesoscutal humeral and suprahumeral sulci, which are comprised of smooth furrows or very small and irregular cells, contrasting the large cells found in the most similar Palearctic species, T. scutellaris.

Comments

The key to species of Palearctic Trissolcus by Kozlov (1978) used microsculpture of the mesoscutellum as a diagnostic character for this species. Contrary to his key, we observed that the sculpture of the mesoscutellum is significantly different from that of the mesoscutum: the mesoscutum is distinctly coriacieous throughout and microsculpture can be seen only very faintly on the posterior portion of the mesoscutellum under high magnification.

Trissolcus viktorovi Kozlov

Figures 213–216; Morphbank 75

Trissolcus viktorovi Kozlov, 1968: 198, 206 (original description, keyed); Fabritius, 1972: 30 (keyed); Kozlov & Lê, 1977: 507 (keyed); Kozlov, 1978: 632 (description); Kozlov & Kononova, 1983: 101 (description); Petrov, 2013: 325 (keyed).

Trissolcus Viktorovi Kozlov: Voegelé, 1969: 149 (keyed, emendation).

Tissoleus victorovi Kozlov: Kozlov, 1981: 187 (keyed, spelling errors).

Description

Female body length: 1.03–1.16 mm (n=16). Male body length: 0.94–1.26 mm (n=6). Body color: head, mesosoma, and metasoma black.

Figures 213–216. 

Trissolcus viktorovi 213 female paratype (USNMENT00916021), head, mesosoma, metasoma, lateral view 214 female paratype (USNMENT00916647), head, mesosoma, metasoma, dorsal view 215 female paratype (USNMENT00916010), mesosoma, dorsal view 216 female paratype (USNMENT00916014), head, anterior view. Scale bars in millimeters.

Head. Color of radicle: brown. Length of radicle: less than width of clypeus. Color of A1A6 in female: brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. .Facial striae: absent. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: bulging. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: uniform in width between midpoint of eye and malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent. Preocellar pit: present. Setation of lateral frons: sparse. Punctation of lateral frons: absent; sparse. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: present only posterior to lateral ocellus. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate.

Mesosoma. Epomial carina: absent. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: rugose anteroventrally. Patch of striae at posteroventral end of femoral depression: present, striae weakly developed and perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: present as a smooth furrow. Mesopleural epicoxal sulcus: present as a smooth furrow. Setation of posteroventral metapleuron: present. Sculpture of dorsal metapleural area: cells of metapleural sulcus extending posteriorly into rugae. Posterodorsal metapleural sulcus: poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; absent. Anteroventral extension of metapleuron: not extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow; indicated by a line of cells. Median mesoscutal carina: present. Macrosculpture of mesoscutum: absent. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: flat, appearing fused with lateral margin of mesoscutum. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: absent. Setation of posterior scutellar sulcus: absent. Form of metascutellum: thick, with only a narrow band of small punctures anteriorly. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae and femora brown, elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anteromedial portion of the tergite. Setation of T2: sparsely present in posterolateral corner. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: sparsely present throughout area not covered by laterotergite.

Diagnosis

T. viktorovi is a distinctive species that can be identified from other species of the thyantae group by the broad gena, the smooth furrow between the hyperoccipital carina and the dorsal and posterior margins of the compound eye. The median mesoscutal carina is distinct in T. viktorovi and is useful for confirming its identity, but this feature may also be found in other Palearctic species of the thyantae group.

Associations

emerged from egg of Dolycoris baccarum (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Eurydema festivum (Linnaeus): [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from Eurydema ventralis Kolenati: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]; emerged from egg of Pentatominae Leach: [Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae]

Material examined

Holotype, female: ARMENIA: Tavus Prov., Noyemberyanskiy Dist., 4.VII.1960, G. Viktorov, ZMAS 0138 (deposited in ZIN). Paratypes: ARMENIA: 3 females, 3 males, USNMENT00916642–00916647 (ZIN). Other material: (13 females, 3 males) PORTUGAL: 3 females, USNMENT00916222, 00916226, 00916241. TURKEY: 10 females, 3 males, USNMENT0091601000916014, 0091601600916022, 00916029 (BMNH).

Trissolcus vindicius (Nixon)

Figures 217–218, 219–221; Morphbank 76

Microphanurus vindicius Nixon, 1938: 123, 128 (original description, keyed); Nixon, 1943: 137 (keyed); Risbec, 1950: 569 (keyed).

Trissolcus vindicius (Nixon): Masner, 1965: 128 (type information, generic transfer); Lê, 1983: 24 (keyed); Johnson, 1992: 640 (cataloged, type information); Lê, 2000: 312, 323 (description, keyed).

Description

Female body length: 0.99–1.63 mm (n=18). Male body length: 1.44 mm (n=1). Body color: head, mesosoma, and metasoma black.

Figures 217–218. 

Trissolcus vindicius 217 female holotype (B.M. TYPE HYM. 9.316), head, mesosoma, metasoma, dorsolateral view 218 female (USNMENT00896037), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 219–221. 

Trissolcus vindicius 219 female (USNMENT00896037), head, mesosoma, metasoma, lateral view 220 female (USNMENT00896024), head and mesosoma, ventrolateral view 221 female holotype (B.M. TYPE HYM. 9.316), head, anterolateral view. Scale bars in millimeters.

Head. Color of radicle: yellow. Length of radicle: less than width of clypeus. Color of A1A6 in female: yellow to brown. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: absent. Number of clypeal setae: 2. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: narrow. Genal carina: absent. Malar striae: absent. Sculpture of malar sulcus: antero-posteriorly striate. Orbital furrow: expanding in size ventrally, strongly so at intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: absent; weakly transversely strigose. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: moderately dense. Sculpture directly ventral to preocellar pit: microsculptured. Macrosculpture of lateral frons: absent. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: complete. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: coarsely crenulate; finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: complete. Pronotal suprahumeral sulcus in posterior half of pronotum: clearly indicated by cells. Location of pronotal suprahumeral sulcus: percurrent. Number of episternal foveae: 4 or more. Course of episternal foveae ventrally: abutting postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: well defined anteriorly, poorly defined to absent posteriorly. Sculpture of femoral depression: smooth. Patch of striae at posteroventral end of femoral depression: absent. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present dorsally. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by large cells. Mesopleural epicoxal sulcus: formed by large cells. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: smooth in anterodorsal corner, coarsely rugose posteriorly; cells of metapleural sulcus extending posteriorly into rugae. Posterodorsal metapleural sulcus: present as line of foveae; poorly defined to absent. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent; indicated by a line of elongate cells. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: present as clearly defined line of cells; present as coarse rugae. Mesoscutal humeral sulcus: indicated by a line of cells. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: absent; weakly rugulose anteriorly, otherwise absent. Pattern of mesoscutal microsculpture: uniform throughout; effaced posteriorly. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Parapsidal line: present. Notaulus: extending at least 1/3 length of mesoscutum. Median protuberance on anterior margin of mesoscutellum: absent; present. Protruberance on anterior margin of mesoscutellum directly posterior to notaulus: present. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Area bounded by axillar crescent: striate. Macrosculpture of mesoscutellum: absent. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent. Setation of posterior scutellar sulcus: present.

Form of metascutellum: coarsely rugose. Metanotal trough: foveate, foveae occupying more than half of metanotal height. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae dark brown, legs elsewhere yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 0. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present throughout posterolateral corner and lateral portions of tergite. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: present in anterior half of sternite not covered by laterotergites. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

Trissolcus vindicius is delimited more by a unique combination of characters than by a single distinctive feature. It can be separated from other species in the flavipes group by the presence of 2 clypeal setae, a well-defined and ventrally expanded orbital furrow, episternal foveae that extend from the postacetabular sulcus to the mesopleural pit, and a mesoscutum without a median mesoscutal carina. See also diagnosis of T. corai.

Lectotype designation

We here designate specimen B.M. TYPE HYM. 9.316 (deposited in BMNH) as the lectotype for T. vindicius.

Material examined

Lectotype, female, M. vindicius: INDONESIA: Jawa Tengah Prov., Java Isl., Wonogiri Regency, 2000ft, 1937, J. S. Phillips, B.M. TYPE HYM. 9.316 (deposited in BMNH). Other material: (18 females, 1 male) INDONESIA: 1 female, USNMENT00916385 (BMNH). JAPAN: 10 females, 1 male, OSUC 144436, 144438, 144445, 144455, 144468, 542360542361, 542368, 542375, 542426542427 (CNCI). NEPAL: 2 females, OSUC 76418–76419 (OSUC). SOUTH KOREA: 5 females, USNMENT00896022–00896025, 00896148 (CNCI).

Comments

The two specimens from Nepal fit within our concept of T. vindicius, but exhibit some notable variation. They are much smaller, have broader mandibles, the microsculpture of the mesoscutum is slightly effaced posteriorly, and the occipital carina is finely crenulate and nearly smooth along its anterior margin. In one specimen the episternal foveae are fewer in number and dorsoventrally elongate relative to the roughly circular shape that is typical for T. vindicius, possibly an allometric consequence of its smaller size.

Trissolcus yamagishii Ryu

Figures 222–224, 225–226; Morphbank 77

Trissolcus artatus Kozlov & Lê, syn. n.; http://bioguid.osu.edu/xbiod_concepts/179840; Morphbank78

Trissolcus yamagishii Ryu, 1984: 37, 53 (original description, keyed).

Trissolcus artatus Kozlov & Lê, syn. n., 2000: 312, 313, 361 (original description, keyed).

Description

Female body length: 1.33–1.76 mm (n=19). Male body length: 1.29–1.58 mm (n=2). Body color: head, mesosoma, and metasoma black.

Figures 222–224. 

T. yamagishii 222 female holotype (Type No. 2221 Kyushu Univ.), head, anterior view 223 female holotype (Type No. 2221 Kyushu Univ.), head and mesosoma, anteroventral view 224 female (USNMENT00896189), head and antennae, anterior view. Scale bars in millimeters.

Figures 225–226. 

T. yamagishii 225 female holotype (Type No. 2221 Kyushu Univ.), head, mesosoma, metasoma, lateral view 226 female (USNMENT00903336), head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Head. Color of radicle: black. Length of radicle: equal to or greater than width of clypeus. Color of A1A6 in female: yellow, becoming brown distally. Color of A7A11 in female: brown. Number of basiconic sensilla on A6: 0. Number of basiconic sensilla on A7: 2. Facial striae: present as short grooves. Number of clypeal setae: 6. Microsculpture on gena directly above mandibular condyle: present. Shape of ventral gena in lateral view: moderately bulging. Genal carina: present and extending dorsally to vicinity of lower margin eye. Malar striae: absent. Sculpture of malar sulcus: smooth; antero-posteriorly striate. Orbital furrow: medially delimited by ridge or carina at midpoint of eye, poorly defined or absent near intersection with malar sulcus. Macrosculpture of frons between antennal scrobe and anterior ocellus: small number of dorsoventral striae directly ventral to anterior ocellus. Preocellar pit: present. Setation of lateral frons: moderately dense. Punctation of lateral frons: absent. Macrosculpture of lateral frons: rugose. OOL: separated by less than one ocellar diameter. Hyperoccipital carina: absent. Macrosculpture of posterior vertex: absent. Microsculpture on posterior vertex along occipital carina: present. Anterior margin of occipital carina: finely crenulate to smooth.

Mesosoma. Epomial carina: present. Macrosculpture of lateral pronotum directly anterior to netrion: striate, striae formed by elongation of cells of netrion sulcus. Netrion sulcus: incomplete. Pronotal suprahumeral sulcus in posterior half of pronotum: undifferentiated from sculpture of dorsal pronotum. Number of episternal foveae: 3; 2. Course of episternal foveae ventrally: distinctly separate from postacetabular sulcus. Course of episternal foveae dorsally: extending to mesopleural pit. Subacropleural sulcus: present. Speculum: transversely strigose. Mesopleural pit: extending ventrally into dorsoventral furrow parallel to mesopleural carina. Mesopleural carina: complete. Sculpture of femoral depression: faintly rugulose. Patch of striae at posteroventral end of femoral depression: present, striae perpendicular to long axis of femoral depression. Setal patch at posteroventral end of femoral depression: present. Microsculpture of anteroventral mesopleuron: present throughout. Macrosculpture of anteroventral mesopleuron: absent. Postacetabular sulcus: formed by open crenulae. Mesopleural epicoxal sulcus: formed by open crenulae. Mesofurcal pit: absent. Setation of posteroventral metapleuron: absent. Sculpture of dorsal metapleural area: mostly smooth with faint rugulae radiating from metapleural pit. Posterodorsal metapleural sulcus: present as line of foveae. Paracoxal sulcus in ventral half of metapleuron: indistinguishable from sculpture to absent. Anteroventral extension of metapleuron: extending to base of mesocoxa. Metapleural epicoxal sulcus: absent or indistinguishable from sculpture. Mesoscutal humeral sulcus: present as a simple furrow. Median mesoscutal carina: absent. Macrosculpture of mesoscutum: reticulate anteriorly, becoming longitudinally strigose posteriorly. Pattern of mesoscutal microsculpture: uniform throughout. Mesoscutal suprahumeral sulcus: comprised of cells. Length of mesoscutal suprahumeral sulcus: about half the length of anterolateral edge of mesoscutum. Notaulus: absent. Median protuberance on anterior margin of mesoscutellum: absent. Shape of dorsal margin of anterior lobe of axillar crescent: round. Sculpture of anterior lobe of axillar crescent: dorsoventrally strigose. Posterodorsal margin of axillular carina: pointed. Area bounded by axillar crescent: smooth. Macrosculpture of mesoscutellum: longitudinally strigose medially, rugulose laterally. Microsculpture on mesoscutellum: present throughout. Median mesoscutellar carina: absent; present. Setation of posterior scutellar sulcus: present. Form of metascutellum: coarsely rugose. Metanotal trough: smooth in dorsal half, with line of foveae ventrally. Metapostnotum: invaginated near lateral edge of metascutellum. Length of postmarginal vein: about twice as long as stigmal vein. Color of legs: coxae black; trochanters and femora brown, with femora becoming much lighter distally, tibiae and tarsi pale brown to yellow. Anteromedial portion of metasomal depression: punctate or crenulate.

Metasoma. Longitudinal striae on T1 posterior to basal costae: present. Number of sublateral setae (on one side): 2; 1; 3. Setation of laterotergite 1: absent. Longitudinal striation of T2: present in anterior two-thirds of tergite. Setation of T2: present in a transverse line and along lateral margin. Setation of laterotergite 2: present. Posteriorly directed setae on medial S1: present. Striation of S2: absent. Setation of S2: present throughout area not covered by laterotergite.

Diagnosis

Trissolcus yamagishi can be distinguished from other Palearctic Trissolcus in the basalis group by the complete mesopleural carina and the characteristic sculpture of the frons in which a dorsoventral carina below the anterior ocellus is flanked by areas of effaced microsculpture (Figures 222, 224). The characters used to separate T. yamagishii and T. comperei (=T. itoi) in the key to species by Ryu (1984) are not reliable. Variation occurs in the degree of rugosity in the mesoscutellum in both T. yamagishii and T. comperei, and the antennae and legs are subject to color variability. Specifically, the legs in T. yamagishii have two color forms, one in which the legs are entirely yellow beyond the coxae and one in which the trochanters and basal femora are brown, becoming lighter distally. We examined a single male of this species which had bright yellow legs beyond the coxae.

Material examined

Holotype, female, T. yamagishii: SOUTH KOREA: Mt. Sudosan, 9.VII–12.VII.1971, K. Yamagishi, Type No. 2221 Kyushu Univ. (deposited in KUEC). Holotype, female, T. artatus: VIETNAM: Thai Nguyen Prov., Thái Nguyên, 14.IV.1986, A. Sharkov, IEBR 0038 (deposited in IEBR). Other material: (19 females, 2 males) INDIA: 1 female, 1 male, USNMENT00916366–00916367 (BMNH). LAOS: 1 female, USNMENT00903336 (BPBM). SOUTH KOREA: 1 female, USNMENT00896017 (CNCI). UNITED ARAB EMIRATES: 16 females, 1 male, USNMENT0089600500896010, 00896160, 0089616800896172, 0089618800896189, 00896215, 00896295, 00896331 (CNCI).

Comments

We here significantly expand the known distribution of this species to include India, Laos, Vietnam and the United Arab Emirates.

Phanuromyia biroi (Szabó), comb. n.

Figures 227–228; Morphbank 79

Asolcus biroi Szabó, 1965: 425 (original description).

Trissolcus biroi (Szabó): Johnson, 1992: 624 (cataloged, type information).

Link to distribution map

http://hol.osu.edu/map-large.html?id=3193

Figures 227–228. 

Phanuromyia biroi, female paratype (Hym.Typ.No. 4611, Mus.Budapest) 227 head, mesosoma, metasoma, lateral view 228 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Material examined

Paratype: SERBIA: 1 female, Hym.Typ.No. 4611, Mus.Budapest (HNHM).

Comments

This species clearly does not belong in Trissolcus and we place it in Phanuromyia Dodd sensu Taekul et al. (2013) based on the rugulose sculpture throughout T2 and the apparently 4-merous antennal clava. We were unable to assess the characters of the ventral frons, but given our confidence in transferring this species to Phanuromyia, we did not find it necessary to attempt dissolving the surrounding glue.

Telenomus fulmeki (Soyka), comb. n.

Figures 229–230; Morphbank 80

Microphanurus fulmeki Soyka, 1942: 175 (original description).

Trissolcus fulmeki (Soyka): Kononova, 2014: 1426 (possibly in Telenomus); Kononova, 2015: 264 (possibly in Telenomus).

Lectotype designation

For the stability of nomenclature, we here designate specimen SDEI 0001A (deposited in SDEI) as the lectotype for Telenomus fulmeki.

Figures 229–230. 

Telenomus fulmeki 229 female lectotype (SDEI 0001A), head, mesosoma, metasoma, lateral view 230 male paralectotype (SDEI 0001B), head, mesosoma, metasoma, lateral view. Scale bars in millimeters.

Material examined

Lectotype, female, M. fulmeki: AUSTRIA: Burgenland, Eisenberg, 17.VI.1941, reared, L. Fulmek, SDEI 0001A (deposited in SDEI). Paralectotype: AUSTRIA: 1 female, SDEI 0001B (SDEI).

Comments

We transfer this species to Telenomus based on the 3-merous antennal clava and T2 that is wider than long. The sculptural details of the head are difficult to assess because of the state of the lectotype and paralectotype specimens. The visible characters, including the transverse rugae of the dorsal frons, place this species in the laricis species group of Telenomus sensu Johnson (1984b) and Huggert (1983).

Telenomus testaceus , comb. n.

Figures 231–233; Morphbank 81

Trissolcus testaceus Lê, 1983: 24 (original description, keyed); Johnson, 1992: 638 (cataloged, type information); Lê, 2000: 312, 321 (description, keyed, type information).

Link to distribution map

http://hol.osu.edu/map-large.html?id=3319

Figures 231–233. 

Telenomus testaceus, female holotype (IEBR 0041) 231 head, mesosoma, metasoma, dorsal view 232 head and mesosoma, lateral view 233 head, anterolateral view. Scale bars in millimeters.

Material examined

Holotype, female: VIETNAM: Gia Lai Prov., K’Bang Dist., forest edge, Buôn Luói, 27.XI.1978, IEBR 0041 (deposited in IEBR).

Acknowledgments

Our ability to assess species identities of Trissolcus across the Palearctic would not have been possible without the generous contributions of specimens, particularly type material, from a large number of collections. For making this revision possible we thank Sergey Belokobilskij (ZIN) for hosting a visit of the first author to the Zoological Institute and an extensive loan Kozlov types; David Notton (BMNH) for sending a great many primary types across the Atlantic; and to Shepherd Myers (formerly of BPBM), Lubomír Masner and Sophie Cardinal (CNCI), Yoshizawa Kazunori and Masahiro Hara (EIHU), Charuwat Taekul (EMBT), Zoltán Vas (HNHM), George Melika, Phạm Thái and Mến Trần (Vietnamese National Museum of Nature), Satoshi Kamitani (KUEC), Roberto Poggi (MCSN), Agnièle Touret-Alby (MNHN), Luca Bartolozzi (MZUF), Vizek Manuela and Dominique Zimmerman (NHMW), Hege Vårdal (NHRS), Luciana Musetti (OSUC), Andrew Liston (SDEI), Serguei Triapitsyn (UCRC), and Lars Vilhelmsen (ZMUC). We also thank Tim Haye (Centre for Agricultural Bioscience International) and Walker Jones (USDA/ARS) for many valuable specimens and host associations, Joe Cora (formerly of OSUC) for database support, Istvan Miko for commentary on morphological characters, and Smithsonian interns Joe Thompson, Amy Cutler, and Ian Realo for contributing specimen images. Norman Johnson (OSUC) and Lubomír Masner provided valuable taxonomic input. Dr. Miguel Alonso-Zarazaga (MNCN) was extremely helpful in the construction of species names. This project was funded by the National Institute of Food and Agriculture, U.S. Department of Agriculture, Specialty Crop Research Initiative under award number 2011-51181-30937, and the Systematic Entomology Lab, USDA-ARS. We are extremely grateful to Tracy Leskey (USDA-ARS Appalachian Fruit Research Station) for funding that extended the postdoctoral fellowship of the first author. The USDA does not endorse any commercial product mentioned in this research. USDA is an equal opportunity provider and employer.

References

  • Ali WK (2011) The level of sunn pest oophagous parasitoids (Hymenoptera: Scelionidae) in infested wheat fields of northern governorate in Iraq with an identification key of Trissolcus specis. Bulletin of the Iraq Natural History Museum 11: 7–15.
  • Ashmead WH (1881) On a parasite bred from the eggs of the orange tree plant bug; being another insect friend of the orange-grower. Florida Agriculturist 4: 181–181. 82
  • Ashmead WH (1887) Studies on the North American Proctotrupidae, with descriptions of new species from Florida. Entomologica Americana 3: 73–119. 83
  • Ashmead WH (1888) Descriptions of some unknown parasitic Hymenoptera in the collection of the Kansas State Agricultural College, received from Prof. E. A. Popenoe. Bulletin of the Kansas Agricultural Experiment Station Appendix 3: i–viii. 84
  • Ashmead WH (1894) Report on the parasitic Cynipidae, part of the Braconidae, the Ichneumonidae, the Proctotrypidae, and part of the Chalcidinae. Part III. Zoological Journal of the Linnean Society of London 25: 188–254. 86
  • Ashmead WH (1896) Report on the parasitic Hymenoptera of the island of Grenada, comprising the families Cynipidae, Ichneumonidae, Braconidae, and Proctotrypidae. Proceedings of the Zoological Society of London 1895: 742–812. 87
  • Ashmead WH (1900) Report upon the aculeate Hymenoptera of the islands of St. Vincent and Grenada, with additions to the parasitic Hymenoptera and a list of the described Hymenoptera of the West Indies. Transactions of the Royal Entomological Society of London 1900: 207–367. https://doi.org/10.1111/j.1365-2311.1900.tb02379.x
  • Ashmead WH (1904) Descriptions of new Hymenoptera from Japan -- I. Journal of the New York Entomological Society 12: 65–84. 88
  • Baltazar CR (1966) A catalogue of Philippine Hymenoptera (with a bibliography, 1758-1963). Pacific Insects Monographs 8: 1–488.
  • Baur H (2001) The Hymenoptera (Chalcidoidea, Ichneumonoidea, Platygastroidea) described by Vittorio Delucchi: an annotated catalogue. Journal of Natural History 35: 55–125. 89https://doi.org/10.1080/002229301447899
  • Bin F (1974) The types of Scelionidae [Hymenoptera: Proctotrupoidea] in some Italian collections (Museums of Genoa and Florence, Institute of Portici). Entomophaga 19: 453–466. 90https://doi.org/10.1007/BF02372781
  • Boldaruyev VO (1969) [Egg parasites of the subfamily Telenominae (Hymenoptera, Scelionidae), reared from the eggs of harmful insects. ] Trudy Buryatskogo Instituta Estestvennykh Nauk Buryatskii Filial Siirskogo Otdeleniya Akademii Nauk SSSR 7: 156–171.
  • Brues CT (1908) Hymenoptera. Fam. Scelionidae. Genera Insectorum 80: 1–59. 91
  • Brues CT (1910) Notes and descriptions of North American parasitic Hymenoptera. VIII. Bulletin of the Wisconsin Natural History Society 8: 45–52. 92
  • Brues CT (1916) Serphoidea (Proctotrypoidea). Pages 529–577 in Viereck. The Hymenoptera or, wasp-like insects, of Connecticut. Guide to the Insects of Connecticut, Part III. Bull. State Geol. Nat. Hist. Surv. No. 22, 824 pp. 93
  • Buhl PN (1996) Two new species of Scelionoidea from East Asia (Hymenoptera: