Biodiversity Data Journal :
Taxonomic Paper
|
Corresponding author: Christer Hansson (christerdennis@gmail.com), Stefan Schmidt (schmidt.s@snsb.de)
Academic editor: Simon van Noort
Received: 30 Sep 2020 | Accepted: 20 Nov 2020 | Published: 16 Dec 2020
© 2020 Christer Hansson, Stefan Schmidt
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hansson C, Schmidt S (2020) A revision of European species of the genus TetrastichusHaliday (Hymenoptera: Eulophidae) using integrative taxonomy. Biodiversity Data Journal 8: e59177. https://doi.org/10.3897/BDJ.8.e59177
|
|
The European species of the genus Tetrastichus (Insecta, Hymenoptera, Eulophidae, Tetrastichinae) are revised with 93 species, including 50 species described as new. The revision was conducted using an integrative taxonomic approach, based on DNA barcoding in combination with morphological characters. The Tetrastichinae are a biologically diverse and species-rich group of parasitoid wasps with numerous complexes of morphologically often very similar species that attack a wide range of hosts in over 100 insect families in 10 different orders. The genus Tetrastichus is, with almost 500 described species, the third largest genus of Tetrastichinae. Although biological information is lacking for most species, current data indicate that Tetrastichus species are gregarious koinobiont endoparasitoids developing on juvenile stages of mainly holometabolous insects. Due to their host specificity, several species of Tetrastichus are used as biological control agents.
The European species of Tetrastichus Haliday (Hymenoptera: Eulophidae) are revised using a combination of externo-morphological and DNA barcoding data. This is the first integrative approach for any of the large genera of the Tetrastichinae. A total of 93 species are included, of which 50 are described as new: T. agonus sp. n., T. antonjanssoni sp. n., T. argei sp. n., T. argutus sp. n., T. asilis sp. n., T. ballotus sp. n., T. bledius sp. n., T. broncus sp. n., T. calcarius sp. n., T. calmius sp. n., T. clisius sp. n., T. cosidis sp. n., T. cumulus sp. n., T. cyprus sp. n., T. delvarei sp. n., T. doczkali sp. n., T. elanus sp. n., T. elodius sp. n., T. ennis sp. n., T. enodis sp. n., T. erinus sp. n., T. evexus sp. n., T. fadus sp. n., T. fenrisi sp. n., T. flaccius sp. n., T. gredius sp. n., T. iasi sp. n., T. illydris sp. n., T. incanus sp. n., T. inscitus sp. n., T. intruitus sp. n., T. johnnoyesi sp. n., T. lacustrinus sp. n., T. ladrus sp. n., T. lanius sp. n., T. lazius sp. n., T. lixalius sp. n., T. lycus sp. n., T. marcusgrahami sp. n., T. minius sp. n., T. mixtus sp. n., T. nataliedaleskeyae sp. n., T. nymphae sp. n., T. pixius sp. n., T. scardiae sp. n., T. splendens sp. n., T. sti sp. n., T. suecus sp. n., T. tacitus sp. n. and T. tartus sp. n. Two keys for the identification of species are presented, one for females and one for males. Based on DNA barcode sequences for 70 of the species, a Maximum Likelihood tree to assess phylogenetic relationships within the genus is presented. These 70 species are also characterised by a combination of CO1 and morphological data. The remaining 23 species, without a DNA barcode, are characterised by morphological data. Using a combination of data from the morphology and CO1 or morphological data only, the species are separated into three species groups (clito-, hylotomarum-, murcia-groups) with 41 unplaced species outside these groups. Hosts are known for 27 of the species and they are gregarious, koinobiont endoparasitoids on a wide range of immature stages of holometabolous insects and appear to be very host specific. The first host record for Lepidoptera (Tineidae) in Europe is included.
integrative taxonomy, DNA barcoding, new species, parasitoids, Tetrastichinae, taxonomic revision, morphology, parasitoid, gregarious endoparasitoids, distribution.
The Tetrastichinae (Hymenoptera: Eulophidae) are one of the largest groups of parasitoid Hymenoptera. They are, with currently about 1650 species in 91 genera, the largest subfamily of the family Eulophidae, that include 297 genera and about 4500 species (
The subfamily is a group of particular importance, even within parasitoid wasps that are generally essential for maintaining biological diversity in terrestrial ecosystems (
Taxonomists have largely ignored tetrastichines because they are taxonomically extremely difficult due to a large number of species and their morphological similarity. Both factors have been a major obstacle for taxonomic revisions, based on morphological characteristics. For the present study, we chose an integrative taxonomic approach, based on DNA barcoding combined with features in the external morphology. Barcoding allows for sequencing large numbers of specimens at a reasonable cost and it provides a standardised set of molecular characters that, together and in combination with morphological data, can be used to assess species boundaries. Examination of morphology allows the inclusion of species for which DNA data could not be gathered, for example, museum specimens of rare species that are often very old and pose problems for DNA sequencing.
The genus Tetrastichus has been attributed to
Tetrastichus species have been found on all continents (
Biological information is known for about one-third of the species (29 out of 93) treated here. The available information from literature and from our own experience strongly indicate that Tetrastichus species are gregarious koinobiont endoparasitoids developing on juvenile stages of mainly holometabolous insects from all the four major orders: Coleoptera, Diptera, Hymenoptera and Lepidoptera (Table
Tetrastichus species |
Host |
T. agrilocidus Graham |
Agrilus sp. (Coleoptera: Buprestidae), Xylotrechus pantherinus Savenius (Coleoptera: Cerambycidae) ( |
T. argei sp. n. |
Gregarious endoparasitoid on Arge ustulata (L.) (Hymenoptera: Argidae) |
T. atratulus (Nees) |
Ptecticus tenebrifer (Walker) (Diptera: Stratiomyiidae) (in Japan) ( |
T. brachyopae Graham |
Gregarious koinobiont endoparasitoid on Brachyopa spp. (Diptera: Syrphidae), clutch size varying from 7–18 specimens, with a strong female bias ( |
T. clito (Walker) |
Cassida murraea L., C. rubiginosa Müller, C. deflorata Suffrian ( |
T. coeruleus (Nees) |
Crioceris asparagi (Coleoptera: Chrysomelidae), emerges from host cocoons, but oviposits already in the host eggs ( |
T. crioceridis Graham |
Crioceris duodecimpunctata L. (Coleoptera: Chrysomelidae), egg-larval parasitoid ( |
T. cyprus sp. n. |
Reared from an unidentified Coleoptera and from an unidentified leaf miner on broad bean (Vicia faba L.) |
T. epilachnae (Giard) |
Epilachna argus (Geoffroy in Fourcroy), E. chrysomelina F., Subcoccinella vigintiquattropunctata L. (Coleoptera: Coccinellidae) ( |
T. halidayi (Graham) |
Agropus ahrensi Germar (Coleoptera: Chrysomelidae) ( |
T. heeringi Delucchi |
Agrilus aurichalceus Redtenbacher, A. integerrimus Ratzeburg, A. viridis (L.) (Coleoptera: Buprestidae) ( |
T. hylotomarum (Bouché) |
Arge ochropus (Gmelin), A. pagana (Panzer) (Hymenoptera: Argidae), Athalia cordata Audinet-Serville, Cladius pectinicornis (Geoffroy) (Hymenoptera: Tenthredinidae), parasitising host larvae and pupae ( |
T. julis (Walker) |
Lema (Oulema) spp. (Coleoptera: Chrysomelidae), gregarious endoparasitoid of host larva ( |
T. legionarius Giraud |
Gregarious endoparasitoid of larvae and pupae of Lipara lucens Meigen (Diptera: Chloropidae) ( |
T. leocrates (Walker) |
Rhynchaenus alni (L.) ( |
T. lyridice (Walker) |
Plagiodera versicolora (Laich.)? (Coleoptera: Chrysomelidae). This record is doubtful, needs checking ( |
T. macrops (Graham) |
Probably Cis spp. (Coleoptera: Cisiidae) in Polyporaceae ( |
T. melosomae Graham |
Chrysomela vigintipunctata (Scopoli) (Coleoptera: Chrysomelidae) ( |
T. miser (Nees) |
Rhynchaenus alni (L.), R. fagi (L.), R. pilosus (F.), R. quercus (F.), R. salicis (L.), Ramphus oxyacanthae (Marsham) (Coleoptera: Curculionidae) ( |
T. murcia (Walker) |
Geosargus sp. (Diptera: Stratiomyiidae), endoparasitoid emerging from pupa ( |
T. nymphae sp. n. |
Gregarious endoparasitoid on Galerucella nymphaeae (L.) (Coleoptera: Chrysomelidae) |
T. pilemostomae Graham |
Pilemostoma fastuosa (Schaller) (Coleoptera: Chrysomelidae) ( |
T. polyporinus Askew |
Possibly Dacne sp. ( |
T. scardiae sp. n. |
Scardia boletella (Fabricius) (Lepidoptera: Tineidae) |
T. setifer Thomson |
Lilioceris lilii (Scopoli) ( |
T. solvae Graham |
Xylomyia cabrerae (Becker) (Diptera: Xylomyiidae) in twigs of Euphorbia canariensis ( |
T. telon (Graham) |
Agrilus viridis L. (Coleoptera: Buprestidae) ( |
T. temporalis (Graham) |
Associated with Phalaris arundinacea (Poaceae) ( |
T. ulmi Erdös |
Scolytus rugulosus (Müller) and Leperisinus orni Fuchs (Coleoptera: Scolytidae), Agrilus sp. and possibly Anthaxia sp. (Coleoptera: Buprestidae) ( |
The sex ratio of clutches is female biased (e.g.
Species of Tetrastichus are, like most other Tetrastichinae, taxonomically exceptional amongst Eulophidae because of their high morphological similarity, hampering a taxonomic revision, based on morphology alone. Unless collected through rearing when the host is known, species of Tetrastichus are very difficult from which to get a series of specimens. Sweep netting and Malaise trapping only yield one or a few specimens at best per collecting event. The rarity of many species, with only very few specimens of each species available for study, prevents the assessment of intraspecific variation and further complicates the definition of species borders. To alleviate the situation, for the present study, an integrative approach was employed, with DNA barcoding as a first step to obtain operational taxonomic units that were subsequently examined morphologically, using criteria that were traditionally used for delimiting species, i.e. morphologically diagnostic characters. In the absence of sufficient morphological differences between OTUs, a conservative approach was followed in that the OTUs were regarded as conspecific until further evidence is available to assess their species status. In these cases, additional genetic markers are needed to disentangle complexes of closely-related species. In several species, the taxonomy has to rely on very few specimens, often with even fewer DNA barcodes and their species status will need to be evaluated as further specimens (and sequence data) become available. Furthermore, the number of species in Europe is probably much higher than currently known and there is, thus, a high probability that species of Tetrastichus that are not included in the present revision will be encountered in the future, in particular with material that is collected in areas other than the main study area in this article and in
For DNA extraction, whole specimens were sent to the Canadian Centre for DNA Barcoding (CCDB) in Guelph, Canada, for DNA extraction and barcode sequencing and subsequent recovery of vouchers for preparation and morphological study. A complete list of voucher specimens included in the revision is given in Suppl. material S1. DNA extraction, PCR amplification and sequencing were conducted at CCDB, using standardised high-throughput protocols (
The data include collecting locality, geographic coordinates, elevation, collector, one or more digital images, identifier and voucher depository. Sequence data can be obtained through BOLD and include a detailed LIMS report, primer information and access to trace files. The sequences are also available on GenBank (for accession numbers, see Suppl. material S1).
Due to the expected presence of complexes of cryptic species, i.e. species that are morphologically virtually indistinguishable, all suitable specimens were subjected to DNA barcoding, even at the risk of obtaining a long series of the same species. The final BOLD dataset (DS-TTSEUR) contains 1,128 specimen records, 1,126 of which are associated with a DNA sequence. Amongst the specimens with a barcode sequence, 860 were assigned a Barcode Index Number (BIN) by the BOLD system, resulting in 73 species with one or more BINs. A total of 60 species were represented by a single BIN, 24 species with two BINs and one species (T. sinope) with three BINs (with a single specimen each).
The initial concerns with obtaining multiple sequences of the same species turned out to be unwarranted. A total of 22% of the species were represented by singletons, almost two thirds (63%) by 1-5 specimens and only two species with over 100 specimens each (Fig.
Sequence divergence statistics were calculated using the Kimura two parameter model of sequence evolution (
For phylogenetic analyses, the COI sequences were aligned using Muscle (
ETHZ Swiss Federal Institute of Technology, Entomological Collection, Zürich, Switzerland; GD Private collection of Gérard Delvare, Montpellier, France; HNHM Hungarian Natural History Museum, Budapest, Hungary; MZLU Biological Museum, Entomology, Lund University, Sweden; NHM the Natural History Museum, London, United Kingdom; NHRS Swedish Museum of Natural History, Stockholm, Sweden; NMW Naturhistorisches Museum, Wien, Austria; OUMNH Oxford University Museum of Natural History, United Kingdom; SMTP Swedish Malaise Trap Project, Station Linné, Ölands Skogsby, Sweden; UCRC University of California, Riverside, Entomology, USA; USNM United States National Museum of Natural History, Washington, D.C., USA; ZSM SNSB-Zoologische Staatssammlung München, Munich, Germany.
With some exceptions (below), the morphological terms used here are explained and illustrated (figs. 6–12) in
To avoid the problems associated with the requirement by the International Code of Zoological Nomenclature (
The colour images of the type specimens of previously-described species were made using Canon camera equipment including an EOS 5D Mark IV body, MP E-65 macro lens and macro twin lite MT-24 EX. The camera was attached to a Cognisys stackshot macro rail system. The images of type specimens for the newly-described species were done with the same equipment, except that the macro lens was substituted with a Canon tele-zoom lens, 70–300 mm (using only 135 & 200 mm), with a 10× Mitutoyo microscope lens attached. The picture stacking was done with Helicon Focus version 6 software and Adobe Photoshop was used for image processing.
The SEM micrographs are from uncoated specimens and were done with a Hitachi SU 3500 scanning electron microscope, in low vacuum.
Tetrastichus
For a complete list of synonyms, see
European species of Tetrastichus can be diagnosed by a combination of three features: submarginal vein in fore wing with one dorsal seta, mid-lobe of mesoscutum with at least three adnotaular setae on each side and propodeum with a ± complete lateral longitudinal carina that splits into two in posterior part (like an inverted "Y", Fig.
Tetrastichus spp.
Figs
Eyes with long setae (e.g. Fig.
FEMALE holotype (Fig.
Colour. Body with weak metallic greenish-blue tinges, scape yellowish-brown with dorsal edge brown, pedicel and flagellum dark brown, tegulae dark brown, wings hyaline with veins pale yellowish-brown, coxae and femora concolorous with body, trochanters dark brown, tibiae and tarsi yellowish-brown.
MALE. Unknown.
Antenna very short with F1 1.6×, F2 1.4×, F3 1.1× and clava 2.2× as long as wide; POL/OOL 2.2; distance between SMG 1.7× distance SMG to SLG; ovipositor sheaths protruding slightly beyond apex of Gt7.
Named after the collector of the holotype, the Swedish entomologist Anton Jansson, who collected insects mainly in the vicinities of the city Örebro (Närke Province), where he also lived and worked as a journalist.
Sweden.
Unknown.
Holotype deposited in MZLU.
Eulophus atratulus
Tetrastichus puncticoxae
See
Female antenna with length of pedicel+flagellum 1.15–1.35× width of mesoscutum, F1 2.0–2.5×, F2 1.6–2.0× and clava (incl. spicule) 2.4–3.0× as long as wide; submedian grooves of mesoscutellum nearer to sublateral grooves than to each other; apical part of ovipositor sheaths does not reach apex of Gt7. Similar to T. dasyops, but with longer flagellum and shorter ovipositor sheaths. Male with relatively short hairs on eyes, about 0.3× OD.
(Former) Czechoslovakia, France, Germany, Italy, Japan, (former) Yugoslavia, Russia, United Kingdom (
Ptecticus tenebrifer (Walker) (Diptera: Stratiomyiidae) (in Japan) (
Type material: neotype ♀ (NHM, type no. 5.3627). Additional material (292♀ 55♂): Austria 2♀ (MZLU, UCRC), France 51♀ 2♂ (CIRAD, NHM, ZSM), Germany 10♀ (MZLU, ZSM, UCRC), Italy 10♀2♂ (NHM, UCRC), Romania 132♀ 50♂ (MZLU, NHM, ZSM), Russia 3♀ (UCRC), Sweden 64♀ 1♂ (MZLU, NHM, NHRS, SMTP, ZSM), United Kingdom 20♀ (NHM).
Tetrastichus brachyopae
See
Submedian grooves of mesoscutellum equidistant from each other and from sublateral grooves; female gaster with ovipositor sheaths projecting beyond apex of Gt7 ; female antennal funicle rather stout, its parts relatively short, F1 1.5–1.7×, F2 1.4–1.5× as long as broad; male with relatively short setae on eyes, about 0.3× OD.
(Former) Czechoslovakia (
Gregarious koinobiont endoparasitoid on Brachyopa spp. (Diptera: Syrphidae), clutch size varying from 7–18 specimens, with a strong female bias (
Type material: holotype ♀ (NHM, type no. 5.3628). Additional material (39♀ 3♂): the Netherlands 13♀ 3♂ (NHM), Sweden 26♀ (MZLU, NHM, ZSM).
Tetrastichus dasyops
See
Female with length of pedicel+flagellum 1.0–1.05× width of mesoscutum, F1 1.7–2.0×, F2 1.2–1.6× and antennal clava (incl. spicule) 2.1–2.35× as long as wide; submedian grooves of mesoscutellum nearer to sublateral grooves than to each other; ovipositor sheaths reach apex of Gt7or slightly beyond. Similar to T. atratulus, but with shorter antennal flagellum and longer ovipositor sheaths.
United Kingdom (
Unknown.
Type material: holotype ♀ (NHM, type no. 5.3626). Additional material (158♀ 14♂): France 7♀ 1♂ (GD, NHM), Italy 5♀ (NHM), Romania 3♀ (NHM), Russia 16♀ 1♂ (MZLU, UCRC), Spain 1♀ (NHM), Sweden 122♀ 12♂ (MZLU, NHM, NHRS), Switzerland 1♀ (NHM), United Kingdom 3♀ (NHM).
FEMALE holotype (Fig.
Colour. Body with weak coppery tinges, entire antenna dark brown, tegulae dark brown, wings hyaline with veins yellowish-white, coxae and femora black, trochanters dark brown, tibiae dark brown to black with apex yellowish-brown, tarsi yellowish-brown or dark brown.
Variation. Scape yellowish-brown (one specimen) or pale brown (one specimen).
MALE. Body length 1.3 mm. Head. Width/length (dorsal view) 2.5, width/length (frontal view) 2.1, mouth width/malar space 1.4, widths head/mesosoma 1.2. Antenna. F1–F4 without basal whorls of setae, scape length/eye height 0.9, scape length/width 2.5, ventral plaque placed in central part of scape, lengths ventral plaque/scape 0.6, pedicel+flagellum length/mesosoma width 2.0, length/width F1, F2, F3, F4 3.0, 2.7, 2.7, 2.7, clava length/width 6.7, lengths pedicel/F1 0.7, lengths F1/F2 1.1, F1/F3 1.1, F1/F4 1.1, lengths F1, F2, F3, F4/clava 0.5, 0.4, 0.4, 0.4.
Colour. As in female.
Female with antennal flagellum long, for example, F1 and F2 both 2.5× and F3 1.9× as long as wide and pedicel+flagellum 1.32× as long as width of mesoscutum; ovipositor sheaths not protruding beyond apex of Gt7. Male with scattered setae and without externo-dorsal, sub-basal compact whorls of long dark setae on funiculars. Through the erect setae on vertex and erect adnotaular setae, similar to species in the murcia -group.
Hungary, Romania.
Unknown.
Holotype deposited in NHM, paratypes in MZLU, NHM.
FEMALE holotype (Fig.
Colour. Body with weak coppery tinges, scape, flagellum and pedicel dark brown, tegulae dark brown, wings hyaline with veins yellowish-brown to brown, coxae black with metallic green tinges, trochanters dark brown, femora black, tibiae yellowish-brown, tarsi dark yellowish-brown to infuscate.
Variation. Paratypes with scape pale brown to yellowish-brown with dorsal edge brown, wing veins yellowish-white.
MALE. Body length 1.2 mm. Head. Width/length (dorsal view) 2.1, width/length (frontal view) 1.3, mouth width/malar space 1.1, eye height/malar space 1.3, widths head/mesosoma 1.1. Antenna. F1–F4 with basal whorls of setae, these setae reaching beyond apex of flagellomere attached to, scape length/eye height 1.0, scape length/width 2.6, ventral plaque placed in central part of scape, lengths ventral plaque/scape 0.8, length/width F1, F2, F3, F4 1.7, 1.8, 1.5, 2.0, lengths pedicel/F1 1.0, lengths F1/F2 0.9, F1/F3 1.1, F1/F4 0.8.
Female with antennal clava (incl. spicule) 2.8× as long as wide; POL/OOL 2.0; distance between SMG 1.7× distance SMG to SLG; ovipositor sheaths not protruding beyond apex of Gt7.
Sweden.
Unknown.
Holotype deposited in MZLU, paratypes in MZLU.
FEMALE holotype (Fig.
Colour. Body with weak metallic green and golden tinges, scape dark yellowish-brown, pedicel and flagellum dark brown, tegulae dark brown, wings hyaline with veins yellowish-white, coxae concolorous with body, trochanters dark brown, femora dark brown with very weak metallic tinges, tibiae and tarsi yellowish-brown.
MALE. Body length 1.1–1.4 mm. Head. Width/length (dorsal view) 2.0, width/length (frontal view) 1.2, mouth width/malar space 1.3, eye height/malar space 1.4, widths head/mesosoma 1.2. Antenna. F1–F4 with basal whorls of setae, these setae subequal in length to flagellomere attached to, scape length/eye height 1.1, scape length/width 3.2, ventral plaque placed in central part of scape, lengths ventral plaque/scape 0.7, pedicel+flagellum length/mesosoma width 1.8, length/width F1, F2, F3, F4 2.0, 2.7, 2.7, 2.3, clava length/width 5.3, lengths pedicel/F1 0.8, lengths F1/F2 0.8, F1/F3 0.8, F1/F4 0.9, lengths F1, F2, F3, F4/clava 0.4, 0.5, 0.5, 0.4.
Colour. Similar to female, but antennal scape dark brown.
Female with distance between posterior ocelli relatively long, POL/OOL= 2.4; F1 1.7× as long as wide; distance between SMG 1.5× distance SMG to SLG; ovipositor sheaths reaching to, but not protruding beyond, apex of Gt7. Male with antennal clava 5.3×, F4 2.3× and scape 3.2× as long as wide.
Hungary, Romania, Sweden.
Unknown.
Holotype deposited in MZLU, paratypes in MZLU, NHM, ZSM.
1♀ “HUNGARY, Balaton-Zamárdi, 1.VIII.1961, A. Sundholm” (NHM, ex coll. Hedqvist).
Cirrospilus Murcia
Tetrastichus trichops
See
Female gaster with ovipositor sheaths projecting distinctly beyond apex of Gt7 , length of projecting part about equal to length of hind basitarsus; setae on eyes very long, 0.8× OD; distance between posterior ocelli relatively short, POL/OOL= 1.5–1.6.
France, Germany, Sweden, United Kingdom (
Sargus (Geosargus) sp. (Diptera: Stratiomyiidae), endoparasitoid emerging from pupa (
Type material: lectotypes ♂ of C. murcia (type no. 5.1944) and ♀ of T. trichops (type no. 4869:1). Additional material (4♀): France 2♀ (GD), Russia 1♀ (MZLU), Sweden 1♀ (SMTP).
See
Female with antennal funicle stout, F1 and F2 both 1.1–1.25× as long as wide, pedicel 1.35–1.5× as long as F1; mid-lobe of mesoscutum without a median groove or with the groove indicated only near mesoscutellum.
Spain (Canary Islands, Tenerife) (
Xylomyia cabrerae (Becker) (Diptera: Xylomyiidae) in twigs of Euphorbia canariensis (
Type material: holotype ♀ (NHM, type no. 5.3629).
FEMALE holotype (Fig.
Colour. Body with weak metallic green tinges, scape, pedicel and flagellum dark brown, tegulae dark brown, wings hyaline with veins yellowish-brown to brown, coxae and femora concolorous with body, trochanters dark brown, tibiae yellowish-brown, fore tarsus pale brown, mid and hind tarsi yellowish-brown with T4 dark brown.
Variation. Scape pale brown in some paratypes.
MALE. Body length 1.2–1.5 mm. Head. Width/length (dorsal view) 1.8, width/length (frontal view) 1.1, mouth width/malar space 1.1, eye height/malar space 1.4, widths head/mesosoma 0.9. Antenna. F1–F4 with basal whorls of setae, these setae reaching beyond apex of flagellomere attached to, scape length/eye height 0.9, scape length/width 2.9, ventral plaque placed in central part of scape, lengths ventral plaque/scape 0.6, pedicel+flagellum length/mesosoma width 1.6, length/width F1, F2, F3, F4 2.0, 2.8, 2.8, 2.7, clava length/width 6.0, lengths pedicel/F1 1.0, lengths F1/F2 0.7, F1/F3 0.7, F1/F4 0.8, lengths F1, F2, F3, F4/clava 0.3, 0.4, 0.4, 0.4.
Colour. Similar to female but scape always dark brown and tibiae brown in some specimens.
Female with antennal clava (incl. spicule) 2.7× as long as wide; ovipositor sheaths protruding beyond apex of Gt7. Male with antennal clava 6.0×, F4 2.3× and scape 2.9× as long as wide.
Germany, Hungary, Romania.
Unknown.
Holotype deposited in MZLU, paratypes in MZLU, NHM, ZSM.
FEMALE holotype (Fig.
Colour. Body with weak metallic green and coppery tinges, scape yellowish-brown with dorsal edge brown, pedicel and flagellum dark brown, tegulae blackish, wings hyaline with veins yellowish-brown, coxae and femora concolorous with body, trochanters dark brown, tibiae yellowish-brown, tarsi yellowish-brown with T4 brown.
MALE. Unknown.
Distance between submedian grooves 2.0× distance between submedian and sublateral grooves; antennal clava (incl. spicule) long, 3.3× as long as wide; ovipositor sheaths not protruding beyond apex of Gt7.
Romania.
Unknown.
Holotype deposited in NHM.
Body bright metallic and shiny; ovipositor sheaths retracted in dry specimens and do not reach apex of Gt7 (as in Fig.
FEMALE holotype (Fig.
Colour. Body metallic blue-green, entire antenna dark brown, tegulae black with metallic tinge, wings hyaline with veins yellowish-brown, coxae and femora concolorous with body, trochanters dark brown, tibiae yellowish-brown, fore tarsus brown, mid and hind tarsi yellowish-brown with T3–4 brown.
MALE. Body length 1.7–1.8 mm. Head. Width/length in dorsal view 2.3, width/length in frontal view 1.2, eye height/malar space 1.2, mouth width/malar space 1.1, widths head/mesosoma 1.0. Antenna. F1–F4 without basal whorls of setae, scape length/eye height 1.1, scape length/width 2.4, ventral plaque placed in central part of scape, lengths ventral plaque/scape 0.7, pedicel+flagellum length/mesosoma width 1.5, length/width F1, F2, F3, F4 1.7, 2.0, 2.0, 2.2, clava length/width 4.4, lengths pedicel/F1 1.0, lengths F1/F2 0.8, F1/F3 0.8, F1/F4 0.7, lengths F1, F2, F3, F4/clava 0.3, 0.5, 0.5, 0.5.
Colour. As in female.
Tibiae yellowish-brown; female with malar space 0.8× eye height, scape 3.5× as long as wide, antennal clava 3.4× as long as wide; mesoscutellum with ratio distance between submedian grooves to distance between submedian and sublateral grooves 1.4.
Sweden.
Holotype deposited in MZLU, paratypes in MZLU, NHM, ZSM.
Gregarious endoparasitoid on Arge ustulata (L.) (Hymenoptera: Argidae), 14♀ and 3♂ have been reared from the same host specimen. The label information does not specify which stage of the sawfly that was parasitised. Another Tetrastichus species, T. hylotomarum, has been recorded from larvae and pupae of Arge spp. (Graham 1991), but not specifically A. ustulata.
1♀ 1♂ ”SWEDEN, Skåne, Kullaberg, 28.ii.1966, P. Benander, ex Arge ustulata”.
FEMALE holotype (Fig.
Colour. Body metallic bluish-green, scape yellowish-brown, pedicel and flagellum pale brown, tegulae dark brown with metallic tinges, wing venation yellowish-brown, coxae and femora concolorous with body, trochanters dark brown, tibiae and tarsi yellowish-brown, T4 pale brown.
MALE. Unknown.
Tibiae yellowish-brown; female with malar space 0.9× eye height, POL/OOL 2.4, F1 1.6×, F2 1.9×, F3 1.7× as long as wide.
Germany.
Unknown.
Holotype deposited in ZSM.
See
Female flagellum long and slender, for example, F1 2.4–2.7×, F3 2.2–2.5× and clava 4.5–5.2× as long as wide.
United Kingdom (
Unknown.
24♀: Czech Republic 1♀ (NHM), Sweden 20♀ (MZLU, NHM, SMTP), United Kingdom 3♀ (NHM).
FEMALE holotype (Fig.
Colour. Body golden-green, scape yellowish-brown with dorsal edge dark brown, pedicel and flagellum dark brown, tegulae black with metallic tinge, wings hyaline with venation yellowish-white, coxae and femora concolorous with body, trochanters black, tibiae yellowish-brown, fore tarsus brown, mid and hind tarsi with T1–3 yellowish-brown, T4 brown.
MALE. Unknown.
Antennal clava 3.8×, F1 1.0× and F2 2.1× as long as wide, F1 1.3× as long as pedicel; ratio POL/OOL = 1.9; length/width of enclosed space between submedian grooves 2.7.
Sweden.
Holotype deposited in MZLU.
Unknown.
See
Antennal clava 3.4–3.6× as long as wide; malar space 0.8–0.9× height of eye; mesoscutellum with enclosed space between submedian grooves 2.7× as long as wide.
(Former) Czechoslovakia, Ireland, Sweden, United Kingdom and (former) Yugoslavia (
Holotype deposited in MZLU, paratypes in MZLU.
Unknown.
9♀, Sweden (NHM).
Eulophus coeruleus
Tetrastichus asparagi
See
Mouth opening very wide, 1.8–2.0 × malar space; body bright metallic blue to bluish-purple; ovipositor does not reach apex of Gt7 ; male funiculars with short whorled setae, 0.5–0.7 × as long as funicular attached to.
France, Germany, Hungary, Italy, The Netherlands, United Kingdom (
Crioceris asparagi (L.) (Coleoptera: Chrysomelidae), emerges from host cocoons, but oviposits in host eggs or larvae (
Type material: lectotype ♀ of E. coeruleus (OUMNH). Additional material (72♀ 18♂): China 1♀ (UCRC), France 2♀ (GD, NHM), Norway 3♀ 1♂ (NHM), Sweden 63♀ 17♂ (MZLU, NHM, ZSM), United Kingdom 3♀ (NHM).
FEMALE holotype (Fig.
Colour. Body golden-green, antenna dark brown, tegulae black with metallic tinges, wings hyaline, wing venation yellowish-white, coxae and femora concolorous with body, trochanters blackish, tibiae and tarsi yellowish-brown.
MALE. Unknown.
Hind coxa with a strong, sharp and complete carina along posterior margin; malar space 0.9× eye height; antennal clava 3.9× as long as wide; mesoscutellum with ratio distance between SMG/distance between SMG and SLG 1.4, length/width of enclosed space between submedian grooves 2.6.
Sweden.
Holotype deposited in MZLU.
Unknown.
FEMALE holotype (Fig.
Colour. Body metallic blue-green, scape dark brown with base dark yellowish-brown, pedicel and flagellum dark brown, tegulae black with metallic tinge, wings hyaline with veins yellowish-brown, coxae and femora concolorous with body, trochanters black, tibiae yellowish-brown, fore tarsus dark brown, mid and hind tarsi yellowish-brown with T4 brown.
Variation. Colour of scape varies from completely dark brown to yellowish-brown with apicodorsal ⅓ dark brown, but most specimens with colour as holotype. Paratypes with body metallic blue or blue-green.
MALE (Fig.
Colour. As in female, but scape completely dark brown. Body metallic blue-green or purple.
Mid and hind tibiae yellowish-brown; mesoscutellum with distance between submedian grooves short, distance between SMG/distance between SMG and SLG 1.2; female with malar space 0.9× height of eye, antenna with F2 2.0×, F3 1.8× and clava 3.8× as long as wide; male antenna with scape 2.2× and clava 5.0× as long as wide and scape length 1.0× height of eye.
Russia and Sweden.
Holotype deposited in MZLU, paratypes in MZLU, NHM, SMTP and ZSM.
Unknown.
1♀ ”SWEDEN, Skåne, Dalby, Ö. Mölla, 21-27.viii.1998, yellow pan trap, R. Danielsson (MZLU)
FEMALE holotype (Fig.
Colour. Body with metallic greenish-blue, entire antenna dark brown, tegulae black, wings hyaline with veins yellowish-brown to brown, coxae concolorous with body, trochanters dark brown, fore and mid femora dark brown, hind femur concolorous with body, tibiae and tarsi yellowish-brown.
MALE. Unknown.
Similar to T. halidayi with the wide mouth and large and oddly-shaped mandibles (as in Figs. 2a and 65d). Differs from T. halidayi in having female gaster short ovate, 1.3× as long as wide, ovipositor sheaths short and not visible in dorsal view, cerci on gaster placed ventrally and not visible in dorsal view, setae on vertex shorter, 0.45× OD and body strongly metallic.
Cyprus.
The specimens in the type series are not in pristine condition, all specimens being more or less broken and dirty. The holotype is missing right wing pair and right pedicel+flagellum is broken off and glued separately to the card.
Reared from an unidentified Coleoptera and from an unidentified leaf miner on broad bean (Vicia faba L.).
Holotype ♀ ”CYPRUS, Phlasson, 30.XII.1967, G.P. Georghiou. Parasite of Coleoptera #521” (UCRC). Paratypes (3♀): 2♀ with same label data as holotype (UCRC), 1♀ from same locality and date as holotype, but ex leaf miner on broad bean (UCRC).
FEMALE holotype (Fig.
Colour. Body with golden-green, scape yellowish-brown with apical ⅓ brown, pedicel and flagellum dark brown, tegulae black with metallic tinge, wings hyaline with veins brown to fuscous, coxae and femora concolorous with body, trochanters dark brown, tibiae yellowish-brown, fore tarsus brown, mid and hind tarsi yellowish-white with T4 brownish.
Variation. Body blue-green in paratype.
MALE. Unknown.
Mesoscutellum with distance between submedian grooves long, distances between submedian grooves/submedian and sublateral grooves 1.9, ratio length/width of enclosed space between submedian grooves 2.2; tibiae yellowish-brown; female with malar space 0.8× eye height; antennal clava 4× as long as wide; POL/OOL 2.0.
Sweden.
Holotype deposited in MZLU, paratypes in SMTP.
Unknown.
FEMALE holotype (Fig.
Colour. Body metallic blue-green, scape yellowish-brown with dorsal edge darker, pedicel and flagellum dark brown, tegulae black with metallic tinge, wings hyaline with veins yellowish-white, coxae concolorous with body, trochanters dark brown, fore and mid femora dark brown with metallic tinges and with apical ⅓ yellowish-brown, hind femur concolorous with body, tibiae yellowish-brown, fore tarsus dark brown, mid and hind tarsi yellowish-brown with T4 dark brown.
MALE. Unknown.
Tibiae yellowish-brown; female with malar space 1.0× eye height; scape 3.6× and antennal clava 3.9× as long as wide; mesoscutellum with ratio distance between submedian grooves to distance between submedian and sublateral grooves 1.4. Similar to T. helviscapus, but with longer antennal clava in female, 3.9× as long as wide.
Sweden.
Holotype deposited in MZLU, paratypes in MZLU and NHM.
Unknown.
FEMALE holotype (Fig.
Colour. Body metallic greenish-blue, antenna dark brown, tegulae dark brown, wing venation yellowish-brown, coxae and femora concolorous with body, trochanters dark brown, tibiae yellowish-brown, fore tarsus brown, mid and hind tarsi yellowish-brown with T4 brown.
MALE. Unknown.
Mesoscutellum 1.1× as long as wide with enclosed area between submedian grooves 3.1× as long as wide; length/width F1, F2, F3, clava 2.2, 2.3, 2.1, 4.0; fore wing with marginal vein 2.7× as long as stigmal vein.
Sweden.
Holotype deposited in SMTP.
Unknown.
Holotype ♀ “SWEDEN, Hälsingland, Hudiksvalls kommun, Stensjön, 62.054°N 16.172°E, 27.vii.2005, SMTP” (SMTP).
See
Tibiae yellowish-brown; female with malar space 0.7× eye height; scape 4.0× and antennal clava 3.0× as long as wide; mesoscutellum with ratio distance between submedian grooves to between submedian and sublateral grooves 1.4 and enclosed space between submedian grooves 2.7× as long as wide.
Moldova, Russia, United Kingdom, (former) Yugoslavia (
Unknown.
France, 2♀ (GD); Morocco, 2♀ (GD).
Eulophus hylotomarum
See
Female antenna: F3 1.0–1.5× as long as wide, clava 2.6–3.0× as long as wide; male antenna: funiculars without an externo-dorsal, sub-basal compact whorl of long setae, F1 1.1–1.4× as long as wide and about as long as pedicel, distinctly shorter than F2; both sexes with mid and hind tibiae broadly infuscate, sometimes mainly black; body bright metallic green.
Bulgaria, (former) Czechoslovakia, France, Germany, Hungary, The Netherlands, Russia, Sweden and United Kingdom (
Reared from Arge ochropus (Gmelin), A. pagana (Panzer) (Hymenoptera: Argidae), Athalia cordata Audinet-Serville, Cladius pectinicornis (Geoffroy) (Hymenoptera: Tenthredinidae), parasitising host larvae and pupae (
11♀ 2♂: France 3♀ (NHM), Italy 1♀ (ZSM), Romania 1♀ (NHM), Sweden 6♀ 2♂ (MZLU, NHM, SMTP).
FEMALE holotype (Fig.
Colour. Body metallic bluish-green, gaster with purple tinges, scape yellowish-brown, flagellum and pedicel dark brown, tegulae black with metallic tinge, wings hyaline and venation yellowish-white, coxae and femora concolorous with body, trochanters black, tibiae yellowish-brown, fore tarsus with T1–3 brown and T4 dark brown, mid and hind tarsi with T1–3 yellowish-brown and T4 dark brown.
MALE (Fig.
Colour. Scape dark brown. Otherwise similar to female.
Mesoscutellum with submedian grooves diverging towards posterior part; female antenna with clava 3.3× as long as wide.
Romania.
Holotype and paratype deposited in NHM.
Unknown.
FEMALE holotype (Fig.
Colour. Mesoscutum metallic blue-green and mesoscutellum golden-green, scape yellowish-brown, pedicel and flagellum dark brown, tegulae black with metallic tinges, wing venation yellowish-white, coxae concolorous with body, trochanters dark brown, fore and mid femora dark brown with apex yellowish-brown, hind femur black with metallic tinge and apex yellowish-brown, tibiae yellowish-brown, tarsi with T1–3 yellowish-brown and T4 brown.
MALE. Unknown.
Mesoscutellum 0.8× as long as wide, length/width of enclosed space between submedian grooves 2.6; mesoscutum bluish and mesoscutellum greenish; antennal clava 3.3× as long as wide.
Germany and Romania.
Holotype deposited in ZSM, paratype in MZLU.
Unknown.
See
Antenna with F1 0.8× as long as F2 and only very slightly longer than the pedicel, F3 1.9× as long as wide; tibiae yellowish-brown; body bright metallic green.
United Kingdom (
Unknown.
4♀: France 2♀ (NHM), Sweden 1♀ (MZLU), United Kingdom 1♀ (NHM).