The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps.

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the "Angiosperm Terrestrial Revolution". Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3-180.5 Ma) and a crown age of 162.2 Ma (153.9-169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history

Spaniopus belokobylskiji Tselikh, 2015, sp. nov.

<i>Spaniopus belokobylskiji</i> sp. nov. <p>(Figs 7–12)</p> <p> <b>Description.</b> <i>Female</i>. Body length 2.7–3.0 mm; fore wing length 2.2–2.4 mm. Colour: head, mesosoma and metasoma dark metallic green with diffuse coppery lustre; antenna with scape yellowish-brown, pedicel basally brown and apically yellowish-brown, flagellum brown; coxae brown, femora and tibiae yellowish-brown, and tarsi yellow except last segment brown; fore wing with dark broad spot under M and S, venation yellowish-brown; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.1–2.2× as broad as long and 1.1–1.2× as broad as mesoscutum; in frontal view 1.25× broader than high. POL 1.1–1.2× OOL. Eye height 1.4× eye length and 1.6× as long as malar space. Frons 1.5× broader than eye height. Distance between antennal toruli and lower margin of clypeus 0.8× distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape as long as eye height and 1.4× as long as eye length, reaching to anterior edge of median ocellus; pedicel 1.6–1.8× as long as broad and as long as F1; combined length of pedicel and flagellum 0.8–0.9× breadth of head; anelli equal in length; flagellum almost filiform; F1 1.5× as long as broad and hardly constricted basally; F2–F5 longer than broad and F6 subquadrate; each funicular segment with one row of dense sensilla; clava 2.0–2.2× as long as broad.</p> <p>Mesosoma 1.50–1.55× as long as broad. Pronotal collar medially 0.2× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with distinctly frenal line. Propodeum medially 0.70–0.75× as long as scutellum; with median carina complete and irregular, costula absent; nucha subglobose and strongly reticulate, occupying 0.5× length of propodeum. Fore wing 2.3× as long as maximum width; basal cell bare; speculum open; M as long as P and 1.2–1.25× as long as S; P 1.2–1.25× as long as S.</p> <p>Metasoma ovate-acuminate, 1.5–1.6× as long as broad and 1.1× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Unknown.</p> <p> <b>Material examined.</b> Holotype: ♀, <b>RUSSIA,</b> “Primorskiy Territory, near Vladivostok, 21.VII 1961, E. Shuvakhina” (ZISP). Paratype: 1 ♀, <b>RUSSIA,</b> “Primorskiy Territory, 30 km SE Ussuriysk, 12–17.VII.2001, S. Belokobylskij” (ZISP).</p> <p> <b>Distribution.</b> Russia (South of Far East).</p> <p> <b>Biology.</b> Unknown.</p> <p> <b>Etymology.</b> Named in honour of the Russian hymenopterist, Dr. Sergey A. Belokobylskij.</p> <p> <b>Comparative diagnosis</b>. <i>Spaniopus belokobylskiji</i> belongs to a group that includes most known species, in which females have as relative shorter F1 that is hardly constricted basally and the second anellus is distinctly transverse; differences from other species are shown in the key. Females are similar to those of <i>S. fulvicornis</i> and <i>S. monospilus</i> by the fore wing having a broad dark spot under M and S, but differ from females of both in having a shorter F1 and M as long as P.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 46-48, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East

Tselikh, Ekaterina (2015): Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East. Zootaxa 4058 (1): 41-65, DOI: http://dx.doi.org/10.11646/zootaxa.4058.1.

Spaniopus dissimilis Walker 1833

<i>Spaniopus dissimilis</i> Walker, 1833 <p>(Figs 13–18, 40–42)</p> <p> <i>Spaniopus dissimilis</i> Walker, 1833: 466; lectotype ♂ (BMNH, examined). Type locality: [Great Britain] "near London". <i>Spaniopus elegans</i> Foerster, 1856: 56; holotype ♂, perhaps lost (Bouček 1972: 312). Type locality: [Germany]. <i>Polyscelis modestus</i> Gahan, 1922: 11; holotype ♀ (USNM, not examined). Type locality: [USA] "Pennsylvania, Hanover".</p> <p>Synonymy by Graham (1969: 705).</p> <p> <b>Description.</b> <i>Female</i>. Body length 1.9–2.4 mm; fore wing length 1.4–2.0 mm. Colour: head, mesosoma and metasoma dark metallic green with diffuse coppery lustre; antenna with scape yellowish-brown, pedicel and flagellum brown or sometimes hardly paler basally; legs yellowish-brown; fore with or without infuscate spot under M and S; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.0× as broad as long and 1.15–1.20× as broad as mesoscutum; in frontal view 1.2–1.3× broader than high. POL 1.0–1.1× OOL. Eye height 1.40–1.45× eye length and 1.2–1.3× as long as malar space. Frons 1.4–1.5× broader than eye height. Distance between antennal toruli and lower margin of clypeus 0.8–0.9× distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape 0.9–1.0× as long as eye height and 1.3× as long as eye length; pedicel 1.7× as long as broad and 1.3× as long as F1; combined length of pedicel and flagellum 0.9× breadth of head; second anellus distinctly transverse, about 2.0× as long as first anellus; F1 1.3–1.4× as long as broad, hardly constricted basally; F2 and F3 longer than broad, F4 and F6 subquadrate; each funicular segment with one row of dense sensilla; clava 1.9–2.0× as long as broad.</p> <p>Mesosoma 1.5–1.6× as long as broad. Pronotal collar medially 0.2× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.65–0.70× as long as scutellum; usually with median carina complete, costula rarely complete; nucha subglobose and strongly reticulate, occupying 0.5× length of propodeum. Fore wing 2.3× as long as maximum width; basal cell bare; speculum open; M 1.35–1.56× as long as P and 1.66–1.8× as long as S; P 1.3–1.4× as long as S.</p> <p>Metasoma ovate-acuminate, 1.36–1.53× as long as broad and 1.2–1.37× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 1.4–1.5 mm. Colour: antenna with pedicel, F1–F6 and clava yellowish-brown; fore wing hyaline; mid tibia brown. POL 1.45–1.5× OOL. Antenna with pedicel 0.9–1.1× as long as F1; F1–F6 longer than broad; combined length of pedicel and flagellum 1.3× breadth of head; clava 2.6–2.8× as long as broad. Mesosoma 1.8× as long as broad. Fore wing with M 1.0–1.1× as long as P. Inner margin of mid tibia more broadly flattened than its outer margin. Metasoma 0.7–0.8× as long as mesosoma. Otherwise similar to female.</p> <p> <b>Material examined. GREAT BRITAIN:</b> 1 ♂ (lectotype), “ LECTOTYPE ”, “grass in fields near London August”, “ <i>Spaniopus dissimile</i> Walker ”, “Stood under this name in old B. M. Coll C. Waterhouse”, “ <i>Spaniopus dissimilis</i> (Walk.) ♂ det. M.W.R. de V. Graham, 1956 ”, “B.M. TYPE HYM. 5 1810”, “ BMNH (E) #953702”; 1 ♀, “Ross-shire, Black Isle vii.1951 N.W. Hussey”,“Ex <i>Megastigmus infested</i> seed.”,“Pres by Com Inst Ent B M 1953- 623”,“ <i>Trichomalus near sunides</i> MWG. 1952”, “ BMNH (E) #953704”. <b>SWEDEN:</b> 2 ♀, 2 ♂, “ SWEDEN. SK. Fjellfota sjö. 31.vii.1938 J.F.P. B.M. 1938-414” (all from BMNH). <b>RUSSIA:</b> 1 ♀, “Primorskiy Territory, Spassk- Dal’niy, 30 km W, 3–4.VII.1984, Belokobylskij”; 1 ♀, “Primorskiy Territory, Natural Reserve “Kedrovaya pad’”, 18.VIII.1976, Storozheva” (both from ZISP).</p> <p> <b>Distribution.</b> Belgium, Canada, Czech Republic, Germany, Great Britain, Netherland, Russia, Slovakia, Sweden, United States of America (Noyes 2015).</p> <p> <b>Biology.</b> Primary parasitoid of <i>Mayetiola destructor</i> (Say, 1817) (Diptera, Cecidomyiidae) (Thompson 1958; Peck 1963; Graham 1969; Bouček 1972; Dzhanokmen 1978; Herting 1978; Burks 1979; Noyes 2015); <i>Diplolepis rosae</i> (Linnaeus, 1758) (Herting 1977; Jennings 2004; Askew <i>et al.</i> 2006; Lotfalizadeh 2007; Noyes 2015) and <i>Phanacis caulicola</i> (Hedicke, 1939) (Askew <i>et al.</i> 2006; Noyes 2015) (Hymenoptera: Cynipidae); lepidopteran pupa of <i>Coleophora</i> sp. (Coleophoridae) (Askew 1970; Noyes 2015) and <i>Elachista poae</i> Stainton, 1855 (Elachistidae) (Vidal 1993; Noyes 2015). Secondary parasitoid of hymenopteran <i>Apanteles</i> sp. (Braconidae), <i>Platygaster vernalis</i> (Myers, 1917) and <i>P. zosine</i> Walker, 1835 (Platygastridae) (Peck 1963; Noyes 2015).</p> <p> The records as egg parasitoids of <i>Erigone atra</i> and <i>E. dentipalpis</i> (Araneae: Liniphyidae) (Vidal 1997; Noyes 2015) should be verified very carefully.</p> <p> <b>Comparative diagnosis</b>. <i>Spaniopus dissimilis</i> is very similar to <i>S. amoenus</i>, the differences between these species being given in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 48-50, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus peisonis Erdos 1957

<i>Spaniopus peisonis</i> (Erdös, 1957) <p>(Figs 46–52)</p> <p> <i>Gyrinophagus peisonis</i> Erdös, 1957: 64; holotype ♀, not available for study (Dr. Z. Vas, personal communication) (HNHM, not examined). Type locality: [Hungary] “Vörs”.</p> <p> <i>Spaniopus peisonis</i>: Bouček, 1972: 307 –308.</p> <p> <b>Description.</b> <i>Female</i>. Body length 2.6–2.8 mm; fore wing length 1.25–2.1 mm. Colour: head, mesosoma and metasoma dark metallic green or blush-green with diffuse coppery lustre; antenna and legs brown; fore wing usually with one lunate spot below S, sometimes with additional small subapical spot; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.0× as broad as length and 1.24–1.30× as broad as mesoscutum; in frontal view 1.3× broader than high. POL as long as OOL. Eye height 1.3× eye length and 1.45–1.55× as long as malar space. Frons 1.45–1.60× broader than eye height. Distance between antennal toruli and lower margin of clypeus 0.8–0.9× as long as distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape 1.1–1.2× as long as eye height and 1.4–1.5× as long as eye length; pedicel 1.8–2.0× as long as broad and 1.4–1.5× as long as F1; combined length of pedicel and flagellum 0.9–1.0× breadth of head; second anellus of antenna transverse and 1.9–2.0× as long as first one; flagellum filiform; F1 1.15–1.20× as long as broad, weakly constricted basally; F2–F3 subquadrate, F4–F6 subtransverse; each funicular segment with one row of dense sensilla; clava 2.0× as long as broad.</p> <p>Mesosoma 1.65–1.08× as long as broad. Pronotal collar medially 1.16–1.18× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.9× as long as scutellum; with median carina complete, costula irregular; nucha subglobose and reticulate, occupying at least 0.5× length of propodeum. Fore wing 2.27–2.35× as long as maximum width; basal cell of fore wing bare; speculum open; M 1.0–1.1× as long as P and 1.5–1.7× as long as S; P 1.5–1.67× as long as S.</p> <p>Metasoma sublanceolate, 1.7–2.1× as long as broad and 1.40–1.55× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 1.6–2.0 mm. Colour: head, mesosoma and metasoma metallic green with diffuse coppery lustre; scape and pedicel yellowish-brown, flagellum brown; legs yellowish-brown; fore wing hyaline. Eye height 1.7× as long as malar space. Combined length of pedicel and flagellum 1.10–1.15× breadth of head; F1–F6 longer than broad; clava 2.2× as long as broad. Fore wing with M 1.3× as long as S; P 1.25–1.3× as long as S. Otherwise similar to female.</p> <p> <b>Material examined. CZECH REPUBLIC:</b> 1 ♀, “Boh. or., Týniště nad Orl., obora, 8.8.59, Bouček”; 1 ♀, “Týniště n. Orl., BOHEMIA, Bouček, 10.8.1959 ”; 1 ♀, “ ČSR., Bohemia, Sobeslav, 22/7.55, A. Hoffer”; 1 ♀, “Břehyně u Doks, BOHEMIA, Bouček, 17.7.63 ” (all from BMNH). <b>FRANCE</b> *: 1 ♀, “ FRANCE. Hérault Reg. Montpellier, H. Lavagne leg.”. <b>GREAT BRITAIN:</b> 2 ♀, 1 ♂, “BRANDON Suffolk 10.vi.73 ex. <i>G. i n c l u s a</i> in Phragmites.”, “♀ <i>Spaniopus peisonis</i> (Erd.) det. Z. Bouček, 1973”; 1 ♀, “ ENGLAND: Cambs. Wicken Fen 2.vi.1976 ”, “♀ <i>Spaniopus peisonis</i> (Erd.) det. Z. Bouček, 1980”, “ BMNH #953708”. <b>HUNGARY:</b> 1 ♂ (allotype), “Vörs, 1955. VI.16 dr. Erdös”, “De Phragmite vulgari Ltm.”, “Allotypus ♂ <i>Gyrinophagus peisonis</i> Erd., 1957 ”, “ <i>Gyrinophagus peisonis</i> Erd. det Erdös”, “♂”, “Hym. Typ. No. 6725 Mus. Budapest”, “Hungarian Natural History Museum Hymmenoptera Coll. Budapest” (HNHM).</p> <p> <b>Distribution.</b> Belgium, Czech Republic, France *, Germany, Great Britain, Hungary, Netherland, Slovakia, Sweden (Noyes 2015).</p> <p> <b>Biology.</b> Primary parasitoid of <i>Giraudiella inclusa</i> (Frauenfeld, 1862) (Diptera, Cecidomyiidae) (Bouček 1972; Dzhanokmen 1978; Tscharntke <i>et al.</i> 1991; Tscharntke 1992; Noyes 2015).</p> <p> <b>Comparative diagnosis</b>. Females of <i>S. peisonis</i> along with those of most known species of <i>Spaniopus</i> have F1 comparatively shorter and hardly constricted basally, and the second anellus distinctly transverse. Differences between these species are given in the key to females. Males of <i>S. peisonis</i> along with those of <i>S. fulvicornis</i>, <i>S. japonicus</i>, <i>S. nigriceps</i> and <i>S. sasacolae</i> have the mid tibia neither broadened or flattened, but <i>S. peisonis</i> males distinctly differ from those of the other four species by its bare basal cell and shorter pedicel.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 58-60, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus amoenus Foerster 1856

<i>Spaniopus amoenus</i> Foerster, 1856 <p>(Figs 1–6)</p> <p> <i>Spaniopus amoenus</i> Foerster, 1856: 56; ♂ type specimens perhaps lost (Bouček 1972: 312). Type locality: [Germany].</p> <p> <b>Description.</b> <i>Female</i>. Body length 1.8–2.4 mm; fore wing length 1.8–1.9 mm. Colour: head, mesosoma and metasoma dark metallic green with diffuse coppery lustre; antenna with scape and pedicel, and F1–F3 yellowishbrown; legs yellowish-brown; fore wing with infuscate spot under M and S; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.0–2.1× as broad as long and 1.1–1.2× as broad as mesoscutum; in frontal view 1.2–1.3× broader than high. POL 1.3× OOL. Eye height 1.4× eye length. Frons 1.35–1.4× broader than eye height. Distance between antennal toruli and lower margin of clypeus 1.08–1.10× distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape 0.90–0.95× as long as eye height and 1.33× as long as eye length; pedicel 2.0× as long as broad and 1.5–1.6× as long as F1; combined length of pedicel and flagellum 0.9× breadth of head; second anellus distinctly transverse, about 2.0× as long as first anellus; F1 1.3–1.4× as long as broad and hardly constricted basally; F2–F3 longer than broad and F4–F6 subquadrate; each funicular segment with one row of dense sensilla; clava 1.8× as long as broad.</p> <p>Mesosoma 1.10–1.15× as long as broad. Pronotal collar medially 0.2× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.70–0.75× as long as scutellum; usually with median carina and costula complete; nucha subglobose and strongly reticulate, occupying 0.50× length of propodeum. Fore wing 2.3–2.4× as long as maximum width; basal cell bare; speculum open; M 1.2× as long as P and 1.48–1.53× as long as S; P 1.2–1.3× as long as S.</p> <p>Metasoma ovate-acuminate, 1.70–1.75× as long as broad and 1.4× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 1.6–1.8 mm. Fore wing length 1.3 mm. Colour: antenna with pedicel and F1–F4 yellow, and F5, F6 and clava brown; fore wing hyaline. Head in dorsal view 1.2–1.3× as broad as mesoscutum and in frontal view 1.7× as broad as high. Antenna with pedicel 1.3× as long as broad and 1.3× as long as F1; F1–F6 longer than broad; clava 1.8× as long as broad; combined length of pedicel and flagellum 1.2× breadth of head. Fore wing with M 0.9× as long as P and 1.1–1.2× as long as S; basal cell with several hairs in apical half. Mid tibia with only outer edge flattened. Metasoma 1.3× as long as broad and 1.0–1.1× as long as mesosoma. Otherwise similar to the female.</p> <p> <b>Material examined. FRANCE:</b> 2 ♀, “ FRANCE. FI. Huelgoat. 31.v.1954. J.F. Perkins. B.M. 1954-396”. <b>GREAT BRITAIN:</b> 1 ♂, “Burnham Beeches Bucks. ENGLAND 17.8.75. Bouček”. <b>HUNGARY:</b> 1 ♀, 1 ♂, “Mecsek, Misana 1953. X. 24. leg. dr. Kaszab”, “avar rostalva”; 1 ♂, “Budapest. Biró. 1927. IX.21 ”. <b>SLOVAKIA:</b> 1 ♀, “Helmec. údoli, Dargov. vrchy, SLOVAKIA or. 5. 8. 54. Bouček” (all from BMNH).</p> <p> <b>Distribution.</b> Belgium, Czech Republic, France, Germany, Great Britain, Hungary, Slovakia, Sweden (Noyes 2015).</p> <p> <b>Biology.</b> Unknown.</p> <p> <b>Comparative diagnosis</b>. <i>Spaniopus amoenus</i> is very similar to <i>S. dissimilis</i>; the differences between these species are given in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 44-46, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus sasacolae Kamijo 1981

<i>Spaniopus sasacolae</i> Kamijo, 1981 <p>(Figs 53–58)</p> <p> <i>Spaniopus nigriceps</i> Kamijo, 1981: 3 –5; holotype ♀ (EIHU, examined). Type locality: [Japan] “Sapporo, Hokkaido”.</p> <p> <b>Description.</b> <i>Female</i>. Body length 2.8–3.4 mm; fore wing length 2.4–2.6 mm. Colour: head, mesosoma and metasoma coppery colouration, with some bluish-green reflections; scape and pedicel yellowish-brown, flagellum brown; legs yellowish-brown; fore wing with light brownish tint; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.1–2.2× as broad as length and 1.25× as broad as mesoscutum; in frontal view 1.3–1.4× broader than high. POL 1.0–1.2× OOL. Eye height 1.25–1.35× eye length and 1.6–1.7× as long as malar space. Frons 1.45–1.6× broader than eye height. Distance between antennal toruli and lower margin of clypeus as long as distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4 or 4:4. Antenna with scape as long as eye height and 1.3× as long as eye length; pedicel of antenna 2.0× as long as broad and as long as F1; combined length of pedicel and flagellum 0.8–0.9× breadth of head; second anellus of antenna transverse and 2.0× as long as first one; flagellum clavate; F1 2.0× as long as broad, constricted basally; F2 and F3 longer than broad, F4 subquadrate, F5 and F6 transverse; each funicular segment with one row of dense sensilla; clava 1.5× as long as broad.</p> <p>Mesosoma 1.6× as long as broad. Pronotal collar medially 1.15× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.6× as long as scutellum; with median carina complete, costula absent; nucha subglobose and reticulate, occupying at least 0.5× length of propodeum. Fore wing 2.2–2.4× as long as maximum width; basal cell of fore wing pilose; speculum open; M as long as P and 1.4–1.6× as long as S; P 1.4× as long as S.</p> <p>Metasoma 1.9–2.0× as long as broad and 1.23× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 2.5 mm. Colour: head, mesosoma and metasoma metallic green or metallic blue with diffuse coppery lustre; scape, pedicel, both anelli and F1 yellow, F2–F4 yellowish-brown, F5, F6 and clava brown; fore wing hyaline. POL 1.4× OOL. Combined length of pedicel and flagellum 1.1× breadth of head; flagellum slender, F1–F6 longer than broad. Metasoma 1.3× as long as broad and as long as mesosoma. Otherwise similar to the female.</p> <p> <b>Material examined. JAPAN:</b> 1 ♀ (holotype), “Sapporo, Hokkaido, 24.V 1.1960, TAKAGI-S.”, “ Holotype <i>Spaniopus sasacolae</i> ♀ Kamijo”; 1 ♂ (paratype), “Iwamizawa, Hokkaido, 15.VII.1980, K. KAMIJO”, “ Paratype <i>Spaniopus sasacolae</i> ♂ Kamijo” (both from EIHU). <b>RUSSIA</b> *: 1 ♀, “Kuril Islands, Kunashir, Alekhino, 11–13.V 1973, Kerzhner” (ZISP).</p> <p> <b>Distribution.</b> Japan, Russia (South of Far East)*, (Kamijo 1981; Noyes 2015).</p> <p> <b>Biology.</b> Primary parasitoid of <i>Hasegawaia sasacola</i> Monzen, 1937 (Diptera, Cecidomyiidae) (Kamijo 1981; Noyes 2015).</p> <p> <b>Comparative diagnosis</b>. This species is similar to <i>S. monospilus</i> and <i>S. fulvicornis</i>; the differences between the species are given in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on page 61, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus japonicus Kamijo 1981

<i>Spaniopus japonicus</i> Kamijo, 1981 <p>(Figs 32–35)</p> <p> <i>Spaniopus japonicus</i> Kamijo, 1981: 5 –8; holotype ♀ (EIHU, examined). Type locality: [Japan] “Tsubetsu, Hokkaido”.</p> <p> <b>Description.</b> <i>Female</i>. Body length 1.5–2.6 mm; fore wing length 1.4–1.9 mm. Colour: head, mesosoma and metasoma dark metallic green with diffuse coppery lustre; scape and pedicel yellowish-brown, flagellum brown; legs yellowish-brown; fore wing hyaline; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.1–2.2× as broad as length and 1.3× as broad as mesoscutum; in frontal view 1.2–1.3× broader than high. POL 1.2–1.3× OOL. Eye height 1.5× eye length and 1.9–2.1× as long as malar space. Frons 1.4– 1.5× broader than eye height. Distance between antennal toruli and lower margin of clypeus 0.8× as long as distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape as long as eye height and 1.20–1.25× as long as eye length; pedicel 2.00–2.25× as long as broad and 1.3–1.4× as long as F1; combined length of pedicel and flagellum 0.9–1.1× breadth of head; second anellus of antenna transverse, 1.9–2.0× as long as first one; flagellum weakly clavate; F1 1.45–1.60× as long as broad, weakly constricted basally; F2–F4 longer than broad, F5 subquadrate, F6 distinctly transverse; each funicular segment with one row of dense sensilla; clava 1.8–2.0× as long as broad.</p> <p>Mesosoma 1.40–1.55× as long as broad. Pronotal collar medially 1.17–0.20× as long as mesoscutum. Scutellum finely reticulate, 1.0–1.1× as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.65× as long as scutellum; with median carina complete, costula absent; nucha subglobose and reticulate, occupying at most 0.4× length of propodeum. Fore wing 2.30–2.35× as long as maximum width; basal cell of fore wing only with several hairs apically; speculum open; M 1.2–1.3× as long as P and 1.5–1.6× as long as S; P 1.4× as long as S.</p> <p>Metasoma ovate-acuminate, 1.50–1.75× as long as broad and 1.2–1.3× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 1.5–2.0 mm. Colour: head and mesosoma metallic green or metallic bluish-green with diffuse coppery lustre; metasoma blackish or dark metallic green, basally with yellowish-brown or brown spot; scape yellow, pedicel and funicle yellowish-brown, clava brown. Pedicel 1.1–1.2× as long as F1; F1 1.45–1.6× as long as broad, F2–F6 longer than broad; F1–F6 with long hairs; clava 2.5× as long as broad. Mesosoma 1.65–1.70× as long as broad. Metasoma 1.10–1.45× as long as broad. Otherwise similar to female.</p> <p> <b>Material examined. JAPAN:</b> 1 ♀ (holotype), “Tsubetsu Hokkaido 14.VIII.1976 K. KAMIJO”, “ Holotype <i>Spaniopus japonicus</i> ♀ Kamijo”, “ HOST <i>Apanteles liparidis</i> Bouche ” (EIHU); 1 ♀, 1 ♂ (paratypes), with the same label as in holotype (all from EIHU). <b>RUSSIA</b> *: 1 ♀, “Kamchatka Territory, Natural Park “Nalychevo”, 5.VII 2013, Tselikh, Rachin”; 1 ♀, “Magadan Province, 12 km N Klepka, 50 km N Magadan, 13.VIII 1975, Barkalov”; 1 ♀, “Primorskiy Territory, Novokachalinsk, coast of Chanka Lake, forest, 12– 16.08.2003, S. Belokobylskij” (all from ZISP).</p> <p> <b>Distribution.</b> Japan, Russian Far East*, South Korea (Kamijo 1981; Noyes 2015).</p> <p> <b>Biology.</b> Primary parasitoid of the dipteran <i>Masakimyia pustulae</i> Yukawa et Sunose, 1976 (Cecidomyiidae) (Kamijo 1981; Sunose 1984; Noyes 2015); lepidopterans <i>Caloptilia theivora</i> (Walsingham, 1891) (Lee <i>et al.</i> 1995; Noyes 2015), <i>Spulerina corticola</i> Kumata, 1964 (Kamijo 1981; Sunose 1984; Noyes 2015), <i>Metriochroa</i> sp. (Gracillariidae) (Sunose 1984; Noyes 2015) and <i>Cymolomia hartigiana</i> (Saxesen, 1840) (Tortricidae) (Kamijo 1981; Sunose 1984; Noyes 2015). Secondary parasitoid of <i>Protapanteles liparidis</i> (Bouche, 1834) (Hymenoptera, Braconidae) (Kamijo 1981; Sunose 1984; Noyes 2015).</p> <p> <b>Comparative diagnosis</b>. This species is very similar to <i>S. nigriceps</i>; the differences between these species are showed in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 52-55, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus fulvicornis Boucek 1972

<i>Spaniopus fulvicornis</i> Bouček, 1972 <p>(Figs 19–24)</p> <p> <i>Spaniopus fulvicornis</i> Bouček, 1972: 309 –310; holotype ♀ (BMNH, examined). Type locality: [Czech Republic] “ Bohemia ”.</p> <p> <b>Description.</b> <i>Female</i>. Body length 1.9–2.6 mm; fore wing length 1.4–2.0 mm. Colour: head, mesosoma and metasoma dark metallic green with diffuse coppery lustre; antenna yellowish-brown; coxae dark metallic green, femora and tibiae yellowish-brown, tarsi yellow, last segment of tarsus brown; fore wing with dark broad spot under M and S, venation yellowish-brown; ovipositor sheath dark brown.</p> <p>Head in dorsal view 2.0–2.1× as broad as long and 1.24–1.25× as broad as mesoscutum; in frontal view 1.23× broader than high. POL 1.25–1.35× OOL. Eye height 1.4× eye length and 1.8–2.0× as long as malar space. Frons 1.4–1.45× broader than eye height. Distance between antennal toruli and lower margin of clypeus as long as distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly emarginate. Mandible formula 3:4. Antenna with scape as long as eye height and 1.25–1.4× as long as eye length, reaching to anterior edge of median ocellus; pedicel 2.0–2.1× as long as broad and 1.1–1.23× as long as F1; combined length of pedicel and flagellum 0.9× breadth of head; second anellus of antenna subquadrate, 2.0–2.1× as long as first one; flagellum almost filiform; F1 1.65–1.90× as long as broad, distinctly constricted basally; F2 longer than broad, F3 and F4 subquadrate, F5 and F6 distinctly transverse; each funicular segment with one row of dense sensilla; clava 1.55–1.70× as long as broad.</p> <p>Mesosoma 1.53–1.70× as long as broad. Pronotal collar medially 0.2× as long as mesoscutum. Scutellum finely reticulate, about as long as broad, with noticeable but shallow frenal line. Propodeum medially 0.66–0.7× as long as scutellum; with median carina weak and costula absent; nucha subglobose and strongly reticulate, occupying 0.4–0.5× length of propodeum. Fore wing 2.3–2.4× as long as maximum width; basal cell bare, basal fold with several hairs; speculum open; M 1.6–1.7× as long as P and 1.96–2.0× as long as S; P 1.2–1.3× as long as S.</p> <p>Metasoma ovate-acuminate, 1.7–1.85× as long as broad and 1.0–1.15× as long as mesosoma. Ovipositor sheath projecting slightly beyond apex of metasoma.</p> <p> <i>Male</i>. Body length 1.5–1.6 mm. Antenna with pedicel 1.8× as long as broad; combined length of pedicel and flagellum 1.1× breadth of head; F1 1.5× as long as broad; F2–F4 longer than broad, F5 and F6 quadrate; clava 2.5× as long as broad. Basal cell of fore wing with several hairs; M 1.8× as long as S (Bouček 1972).</p> <p> <b>Material examined. CZECH REPUBLIC:</b> 1 ♀ (holotype) “ Holotype ”, “Sedlou Litoměřic, BOHEMIA Bouček”, “ HOLOTYPUS ♀ <i>Spaniopus fulvicornis</i> Bouček ”, “Pres by Com Inst Ent B M 1972-2”, “B.M. TYPE HYM. 5. 2177”, “ BMNH (E) #953700” (BMNH). <b>RUSSIA *:</b> 1 ♀, “Primorskiy Territory, Natural Reserve “Ussuriisky”, River Sokolovka, 30.VII.1975, Storozheva”; 1 ♀, “Primorskiy Territory, Vladivostok, Okeanskaya, forest, 30.VII.2001, S. Belokobylskij” (both from ZISP); 1 ♀, “Primorskiy Territory, Lazo, 43 31 16 N 134 3 52 E, 1547 m. Mountain top, MT. 04.viii.2001 ”, “M. Quest coll. BMNH (E) 2009-59 ” (BMNH); 1 ♀, “Yevreyskaya Autonomic Province, 20 km NW Amurzet, 17 VI 1985, Kasparyan” (ZISP).</p> <p> <b>Distribution.</b> Czech Republic, Russia (South of Far East)*, Slovakia, Sweden (Noyes 2015).</p> <p> <b>Biology.</b> Unknown.</p> <p> <b>Comparative diagnosis</b>. Females of <i>Spaniopus fulvicornis</i> are very similar to those of <i>S. monospilus</i>, but males are similar to those of <i>S. nigriceps</i> and <i>S. japonicus</i>. The differences between these species are given in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on page 50, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt

Spaniopus varicornis Boucek 1972

<i>Spaniopus varicornis</i> Bouček, 1972 <p>(Figs 63–66)</p> <p> <i>Spaniopus varicornis</i> Bouček, 1972: 310 –312; holotype ♂ (BMNH, examined). Type locality: [Slovakia] “Smokovec”.</p> <p> <b>Description.</b> <i>Male</i>. Body length 1.35–1.60 mm; fore wing length 1.2–1.3 mm. Colour: head, mesosoma and metasoma metallic green with diffuse coppery lustre; antenna with scape, F1, F2, F5 and F6 yellow, pedicel, F3, F4 and clava brown; legs yellowish-brown, mid tibia apically brown; fore wing with dark broad spot under M and S.</p> <p>Head in dorsal view 1.8–1.9× as broad as length and 1.35–1.40× as broad as mesoscutum; in frontal view 1.25× broader than high. POL 1.4–1.5× OOL. Eye height 1.3× eye length and 2.2–2.45× as long as malar space. Distance between antennal toruli and lower margin of clypeus 0.90–0.95× as long as distance between antennal toruli and median ocellus. Clypeus radially striate, its lower margin weakly subemarginate. Mandible formula 3:4. Antenna with scape 0.9× as long as eye height and 1.2× as long as eye length; pedicel 1.7–1.8× as long as broad and 1.4–1.6× as long as F1; combined length of pedicel and flagellum 1.36× width of head; F1–F6 subquadrate; each funicular segment with one row of dense sensilla; clava 2.9–3.0× as long as broad.</p> <p>Mesosoma 1.95–2.0× as long as broad. Pronotal collar medially 1.13× as long as mesoscutum. Scutellum finely reticulate, with noticeable but shallow frenal line. Propodeum medially 0.7× as long as scutellum; with median carina complete and irregular; nucha reticulate, occupying 0.45× length of propodeum. Inner margin of mid tibia weakly concave submedially. Basal cell of fore wing pilose; speculum open; M 1.25–1.28× as long as P and 1.56–1.60× as long as S; P 1.25–1.30× as long as S.</p> <p>Metasoma 1.20–1.45× as long as broad and 0.7× as long as mesosoma.</p> <p> <i>Female</i>. Body length 2.0 mm. Colour: head, mesosoma and metasoma metallic green or blush-purple often with diffuse coppery lustre; fore wing hyaline. Head in dorsal view 2.0× as broad as length and 1.26× as broad as mesoscutum. Combined length of pedicel and flagellum 0.9× width of head; clava 2.0× as long as broad. Fore wing M 1.04× as long as P and 1.13× as long as S; P 0.9× as long as S. Metasoma ovate-acuminate 1.6× as long as broad (Bouček 1972).</p> <p> <b>Material examined. SLOVAKIA:</b> 1 ♂ (holotype), “ Slovakia, Vys. Tatry, Starý Smokovec, 29.8.58. Bouček”, “ Holotype ”, “ HOLOTYPUS ♂ <i>Spaniopus varicornis</i> Bouček ”, “Presented to BMNH 1976 Z. Bouček”, “B.M. TYPE HYM. 5. 2400”, “ BMNH (E) #953701” (BMNH). <b>RUSSIA</b> *: 1 ♂, “Kamchatka Territory, 80 km S Lazo, Natural Reserve “Kronotsky”, Ipuin; N 55° 06.969 E 159°57.925, 18.VIII 2013, Tselikh, Rachin” (ZISP).</p> <p> <b>Distribution.</b> Russia (Far East)*, Slovakia (Noyes 2015).</p> <p> <b>Biology.</b> Unknown.</p> <p> <b>Comparative diagnosis</b>. <i>Spaniopus varicornis</i> is very similar to <i>S. hedqvisti</i>; the differences between the species are given in the respective keys to the sexes.</p>Published as part of <i>Tselikh, Ekaterina, 2015, Review of the world species of Spaniopus Walker, 1833 (Hymenoptera: Pteromalidae), with description of a new species from the Russian Far East, pp. 41-65 in Zootaxa 4058 (1)</i> on pages 61-64, DOI: 10.11646/zootaxa.4058.1.2, <a href="http://zenodo.org/record/239870">http://zenodo.org/record/239870</a&gt