A Taxonomic Review of Ichneumon-flies (Hymenoptera ...€¦ · KASPARYAN, KOPELKE ENTOMOLOGICAL...

ISSN 0013–8738, Entomological Review, 2010, Vol. 90, No. 1, pp. 71–98. © Pleiades Publishing, Inc., 2010. Original Russian Text © D.R. Kasparyan, J.-P. Kopelke, 2010, published in Entomologicheskoe Obozrenie, 2010, Vol. 89, No. 1, pp. 235–266.

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A Taxonomic Review of Ichneumon-flies (Hymenoptera, Ichneumonidae), Parasitoids of Gall-forming Sawflies

(Hymenoptera, Tenthredinidae) on Salix. Part II. Review of the Palaearctic Species of the Genus Saotis Förster

with Description of Four New Species D. R. Kasparyana and J.-P. Kopelkeb

aZoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia bForschungsinstitut Senckenberg, Frankfurt am Main, Germany

Received April 15, 2009

Abstract—This is the second communication on Ichneumonidae, parasitoids of gall-making sawflies on Salix. Twenty-one Palaearctic species of the specialized genus Saotis are revised and classified into 12 species groups. Original data on 29 species of Saotis hosts are given: 18 spp. of Phyllocolpa, 10 Pontania and 1 species of Ponto-pristia. A key to species and subspecies is given and illustrated by 55 figures. For each species the literature, syn-onymy, diagnosis, material and hosts are given. Four new species are described: Saotis alpinator sp. n., S. boreator sp. n., S. granulator sp. n. and S. subarctor sp. n. DOI: 10.1134/S0013873810010069

In the preceding paper (Kasparyan and Kopelke, 2009), the authors considered a complex of ichneu-mon-flies (Ichneumonidae), parasitoids of gall-making sawflies on Salix. This complex includes about 45 species belonging to 15 genera of 6 Ichneumonidae subfamilies. The general list of hosts and their food-plants (33 species of Salix) was given, with reared species of ichneumonids; a total of 75 species of hosts belonging to the genera Pontania Costa, 1859 (40 spp.), Phyllocolpa Benson, 1960 (21 spp.), and Euura Newman, 1837 (14 spp.) were recorded. An annotated list was given for about 30 species of ichneumonids (except for Saotis) reared from gall-making sawflies, two new species of the subfamily Ctenopelmatinae (Anoncus gallicola Kasparyan et Ko-pelke and Rhinotorus brachycerus Kasparyan et Ko-pelke) were described. A key to the species of this complex of Ichneumonidae was elaborated. The spe-cies of the specialized genus Saotis were not included in the key and are treated here, in the second paper, with a review of all the recorded Palaearctic species. Four new species of Saotis, reared from Pontania and Phyllocolpa, are described.

MATERIALS Nearly the whole material reared (except for that

specially noted) was collected by J.-P. Kopelke. Spe-

cies of sawflies (hosts) and their food-plants were also determined by J.-P. Kopelke. Names of gall-forming sawflies adopted in Kopelke’s (1994, 1999, 2003) publications are used leaving aside the most recent nomenclature changes. The material reared is depos-ited mostly in the Forschungsinstitut Senckenberg, Frankfurt am Main (SMF). The names of countries in “Material” are given in bold, and those of provinces and Latin names of insects, in italics.

In addition to the extensive material from SMF, we also used here insects examined by D.R. Kasparyan (types and comparative material) from the following collections: the Zoological Institute of the Russian Academy of Sciences in St. Petersburg (ZIN), the Natural History Museum, London, Great Britain (NHM, curator G. Broad), the National Museums of Scotland, Edinburgh, Great Britain (NMS, curator M. Shaw), the collection of R. Hinz in Zoologische Sammlung des Bayerischen Staates, Munich, Germany (Mus. Munich, curators E. Diller, S. Schmidt), collec-tions of R. Jussila in Turku University, Finland (Mus. Turku), and W. Hellén, Zoological Museum, Helsinki, Finland (Mus. Helsinki, curator A. Albrecht), types of Saotis, Lathrostizus and Mesoleius s. l. from the Mu-seum of Lund University, Sweden (Mus. Lund, curator R. Danielsson), Naturhistoriska Riksmuseet, Stock-

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holm, Sweden (Mus. Stockholm, curator T. Kronest-edt), Zoologisches Museum, Humboldt-Universität, Berlin, Deutschland (Mus. Berlin, curator F. Koch), the Zoological Department of the Hungarian Natural History Museum, Budapest, Hungary (Mus. Budapest, curator Csősz Sandor), the National Museum of Natu-ral History, Washington, USA (Mus. Washington, curator R. Carlson), the American Entomological In-stitute, Gainesville, Florida, USA (AEI, curator D. Wahl).

The authors cordially thank the curators of these collections and also A. Reschikov who furthered ob-taining the material from the USA, Canada, and Finland (including photographs of some type spe-cimens).

The participation of D.R. Kasparyan in this work was supported by a grant from the Russian Foundation for Basic Research (no. 07-04-00454). The financial support of J.-P. Kopelke was provided by the German Research Foundation (Deutsche Forschungsge-meinschaft: 1990, 1993, 1999, 2000) and the research foundations (Ungerer-Stiftung, Rüppell Expeditions Fond) of the “Senckenbergische Naturforschende Ge-sellschaft” (SNG: 1985, 1988, 1997, 2001).

Genus SAOTIS Förster, 1869

Type species Mesoleius (Saotus) brevispina Thom-son, designated by Viereck, 1914.

Dalla Torre, 1902 : 272 (catalogue, bibliogr.); Schmiedeknecht, 1914 : 2807–2820 (key, descriptions of species); Yu, Horstmann, 1997 : 436–437 (ca-talogue, bibliogr.); Kasparyan, Shaw, 2003 : 351–355 (key; = Iskarus); Kasparyan, 2007 : 547–551 (in: Kasparyan and Khalaim, 2007) (key).—Iskarus Kola-rov, 1987 : 70; type species Iskarus seleuciformis Ko-larov, 1987 (= Saotis mirabilis Schmiedeknecht, 1914).

This relatively small Holarctic genus comprises about 22 species, among which 20 have been de-scribed from Europe. The species of the genus can be readily distinguished by the strongly compressed ab-domen of the female; the ovipositor sheath is short and broad (Figs. 1–8, 11–15, 41–45). The body is about 4–6 mm long (up to 15 mm in S. mirabilis); the fore wing has no areolet. The insects are parasitoids of gall-making nematine sawflies of the genera Pontania and Phyllocolpa (Tenthredinidae, Nematinae). One spe-cies, Saotis albionis Kasp., was reared in Finland from the larva of a nematine, Pontopristia amentorum de-

veloping in catkins of Salix. The evolutionary links of Saotis with the genus Mesoleius s. l. (including Cam-podorus) are rather clear. The peculiarities of the fe-males’ behavior during the process of the host infes-tation have not been described and the morphofunc-tional significance of the compressed apical segments of the abdomen is unknown.

The following 12 species-groups can be distingui-shed in the genus (the most important apomorphies are given in italics).1

1. The brevispina species-group includes S. bre-vispina Thoms., S. varicoxa Thoms., S. subarctor sp. n. It differs in having epipleura of tergites 4 and 5 separated by crease (in S. brevispina the crease is not always distinct), ovipositor sheath truncate at apex, yellow or reddish yellow (Figs. 1, 3, 44, 45). Antenna with 21–26 flagellomeres; i. fl 1.1–1.2; i. gen 0.4–0.5; lower margin of clypeus broadly con-vex and laterally with small sharp flanges (Fig. 36); i. fem 4.3–5.0; i. nrv 0.05–0.22; i. terg 1.5–2.1; noticeable peculiarity of coloration (except for S. subarctor): yellow dorsolateral spots of scutellum (Fig. 9) or its yellow dorsolateral margins.

Hosts: no reliable data on the hosts available. 2. The tricolor species-group. 2a. The tricolor s. str.

subgroup includes S. bilineata Grav., S. morleyi Fitton (= albiventris Kasp.), S. tricolor Toms. (= liopleuris Thoms.). It may easily be distinguished by flagellum with (24)26–30 segments, by strongly curved in profile tergite 8 (Figs. 4, 6, 14), by short malar space (i. gen 0.25–0.3). First flagellar segment only 4–5 times as long as its width at middle; i. fl 0.9–1.07; hind femur rather thick (i. fem 3.9–4.4); hind tibia with distinct white ring in middle part and often tricolor (Fig. 16); mesoscutum and scutellum usually with yellow mark-ings; mesoscutum and tergites of abdomen shiny and almost smooth. _____________ 1 The basic morphometric characters and their abbreviations used

here, in the key and in descriptions of new species of Saotis are listed below: i. fl, index flagellaris, ratio of length of first and second flagellar segments combined to eye height (longitudinal diameter); i. gen, index genalis, ratio of minimum length of ma-lar space (“gena”) to basal width of mandibles; i. ar. ap, index propodealis, ratio of area apicalis length to total length of propo-deum along midline; i. terg, index tergalis, ratio of length of first metasomal tergite to its maximum width; i. nrv, index nervel-laris, ratio of length of lower (posterior) section of hind-wing nervellus to its entire length; i. fem, index femoralis, length to width ratio of hind femur (in lateral view); i. tars, ratio of lengths of first to fifth segments of hind tarsus.

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2b. The renovata subgroup: S. renovata Morley? (= morleyi sensu Kasparyan, 2007). It differs from species of subgroup tricolor in having tergite 8 not so strongly curvate in profile (Fig. 13), antenna only with 21–25 flagellomeres, malar space longer (i. gen about 0.4) and tergite 1 rather long (i. terg 2.1–2.9) (Fig. 18, 21); yellow pattern on thorax and coloration of legs as in other species of group. S. renovatus is possibly also links with species-group albionis. Hosts: 3 species of tricolor-group are reared from 9 species of Phyllo-colpa (for S. bilineata no the reliable data on the hosts).

3. The albionis–hoeli species-group; consists of S. albionis Kasp., and also S. hoeli Roman (Green-land) and S. clypeata Ashmead (Alaska). It differs in having long projecting hypopygium and usually strongly projecting and straight dorsally in profile tergites 7 and 8 (Figs. 31, 32); tergites 3–7 with hind margin medially notched; pygostyli merged (im-movably connected) with tergite 8. Antenna usually with 22–23 flagellomeres; i. fl 0.9–1.0; i. gen 0.33–0.45 ; i. fem about 4.7; i. terg 1.6–1.8; i. nrv 0.25–0.35; nervulus usually postfurcal; tarsal claws at base with 1–2 teeth. Thorax black or with weak yellow markings. Legs with varying coloration, hind tibiae may be brownish yellow or whitish with fuscous apex. Hosts: S. albionis is reared from Pontopristia (whose larva lives in catkins of Salix).

4. The nigriventris species-group; includes only S. nigriventris Thoms. (?= compressiventris Strobl). This species may easily be distinguished by presence of numerous small hairs on tergites of abdomen (in-cluding epipleura), and by tergite 8 which is strongly convex dorsally, with tooth on its hind margin, and with anal vertical slot upward this tooth (Fig. 8). Antenna with 22–24 flagellomeres; i. fl 0.9; malar space very short (i. gen 0.2–0.25); i. terg 4.4; areas of propodeum strongly obliterate; thorax completely black; ovipositor sheath from ochreous to brownish, with deep dorsal notch at base (Fig. 8). Hosts un-known.

5. The compressiuscula species-group; includes S. compressiuscula Thoms. and S. caucasica Kasp. The group differs in having very short cheeks (i. gen 0.2–0.25), antenna with 22–24 flagellomeres; i. fl 1.05–1.1; carinae of propodeum often very weak; nervulus interstitial or weakly postfurcal; i. nrv 0.1–0.28. Apical tergites of abdomen not modified, moderately compressed; ovipositor sheath rather broad

(Figs. 2, 49). Face and cheeks sometimes with yellow markings; mesoscutum with anterolateral yellow spots; epimeron yellow at apex; scutellum black; tergites 5 and 6 with epipleura entirely or predominantly pale (Fig. 2, 49). Legs reddish yellow; hind tibia without distinct white ring, usually light rufous with fuscous apex and lighter posteriorly at basal 0.6. Hosts un-known.

6. The alpinator species-group; includes only S. alpinator sp. n. This species by wide ovipositor sheath and rather weakly compressed abdomen (Fig. 43) similar to species-group nigriscuta and partly to compressiuscula. It differs from both groups in hav-ing basal flagellomeres shorter (i. fl 0.9). Small insects with fore wing 3.5 mm long; Antenna with 22 flagel-lomeres; i. gen about 0.5 (Fig. 28); i. nrv 0.28; i. fem 4.6; i. terg 1.7. Scutellum black; coxae and tro-chanters I predominantly black. Pterostigma pale with brownish margins. Host: Pontania.

7. The nigriscuta species-group; includes S. hete-ropus Thoms., S. nigriscuta Thoms., S. boreator sp. n. It differs in having ovipositor sheath wide and black (Figs. 5, 12, 41), which usually is wider than hind femur. Two basal flagellomeres always distinctly longer than maximum diameter of eye (i. fl 1.15–1.33). Antenna with 19–26 flagellomeres. Hosts: Phyllo-colpa.

8. The pygidiator species-group; includes S. py-gidiator Kasparyan et Kopelke, 2009, one Holarctic species with 4 subspecies. In most characters similar to nigriscuta-group; it differs from this group and other species-groups of Saotis in having hypopygium of female is covering with hairs erected and denser (Fig. 42), temples broader and swollen (Fig. 30); ner-vulus usually interstitial, i. nrv 0.1–0.25, and ptero-stigma usually pale (except for S. pygidiator ivan). Antenna with 23–28 flagellomeres; i. fl 1.25 ± 0.1; i. gen about 0.4 ± 0.1; i. fem 5.0 ± 0.5; the biggest spur of hind tibia usually shorter than half of hind basitar-sus; areola of propodeum is narrow (Fig. 40); i. terg 1.7–2.0; tergite 2 with very fine and smoothed granu-lation and with short and rather scarce hairs. Tergite 8 short and usually strongly retracted in abdomen; hypopygium truncated on hind margin and slightly concave medially; ovipositor sheath about 0.8 times as wide as hind femur laterally. Mesoscutum usually with yellow anterolateral spots, scutellum black. Hosts: Saotis p. pygidiator is reared from 3 species of Pontania.

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9. The granulator species-group; includes only S. granulator sp. n. The species may easily be distin-guished by distinct granulation that evenly covers the body (on tergites 4–8 the granulation is smoother). Sternites are more heavily sclerotized and usually without median longitudinal fold. Head not narrowed beyond eyes (Fig. 37); malar space short, i. gen about 0.25; i. fl 0.8–1.0; antenna with 21–24 flagellomeres; mesoscutum and scutellum black; i. nrv 0.33; i. terg 1.7–1.8; tergites 7 and 8 usually projecting beyond hypopygium along their entire length (Fig. 39); hy-popygium truncate on hind margin and slightly con-cave medially. Hosts: Saotis granulator is reared from 6 species of Pontania.

10. The longiventris species-group; includes only S. longiventris Thoms. It differs in strongly elongate abdomen (Fig. 15); i. terg 2.7. Head strongly nar-rowed (Fig. 50); i. gen 0.33; i. fl 0.9; nervulus intersti-tial; ovipositor sheath as wide as hind femur; tergites completely black, without thin pale band on hind mar-gin. Host unknown.

11. The dorsata species-group; includes only S. dorsata Thoms. It differs from the congeners in having ovipositor sheath narrower (Fig. 7) and tergites 2 and 3 with distinct reddish brown pattern. Mesoscu-tum and scutellum black (scutellum sometimes brownish at apex), subtegular ridge with yellow stripe. Antenna with 22–26 flagellomeres; i. fl 0.9; i. gen 0.3; i. fem 3.9–4.2; i. terg 2.0–2.2; third segment of hind tarsus 1.1–1.2 times as long as fifth segment. Epipleura of tergites 4 and 5 sometimes separated by a weak crease. Host unknown.

12. The mirabilis species-group; includes only S. mirabilis Schmied. This species differs from all congeners in having large size (length of body about 15 mm) and very long abdomen (abdomen 3 times as long as head and thorax combined); tergites 3–8 on hind margin with a deep median notch; tergite 8 de-pressed, ovipositor sheath small. Mesothorax red. Host unknown.

In the World catalogue of Ichneumonidae, Hypam-blys salicis Teunissen, 1953 (holotypum examinavi; = Synodites) was erroneously placed (Yu and Horst-mann, 2004) in the genus Saotis (see Kasparyan, 2004 : 48).

A LIST OF SAOTIS AND THEIR HOSTS Saotis boreator sp. n.: Phyllocola nudipectus (Vikberg, 1965) / Salix

phylicifolia L.

Saotis morleyi Fitton:

Phyllocolpa ischnocera (Thomson, 1862) / Salix myrsinifolia Salisb.

Ph. leucapsis (Tischbein, 1846) / Salix cinerea L.

Ph. plicaglauca Kopelke, 2007 / Salix glaucosericea Flod.

Ph. plicalapponum Kopelke, 2007 / Salix lapponum L.

Ph. scotaspis (Förster, 1854) / Salix viminalis L.

Ph. spirhelvetica Kopelke, 2007 / Salix helvetica Vill.

Saotis nigriscuta Thoms.:

Phyllocolpa acutiserra (Lindquist, 1948) / Salix lapponum L.

Saotis ?renovata Morley:

Phyllocolpa anomalpotera (Förster, 1854) / Salix caprea L.

Ph. purpurea (Cameron, 1884) / Salix purpurea L.

Phyllocolpa sp. / Salix udensis Trautv. et Mey.

Saotis subarctor sp. n.:

Phyllocolpa acutiserra (Lindquist, 1948) / Salix lapponum L.

Ph. nudipectus (Vikberg, 1965) / Salix phylicifolia L.

Ph. plicaphylicifolia Kopelke, 2007 / Salix phylicifolia L.

Saotis tricolor Thoms. (= liopleuris Thoms.):

Phyllocolpa anomalpotera (Förster, 1854) / Salix caprea L.

Ph. ischnocera (Thomson, 1862) / Salix myrsinifolia Salisb.

Ph. leucosticta (Hartig, 1837) / Salix caprea L.

Ph. nudipectus (Vikberg, 1965) / Salix phylicifolia L.

Ph. oblita (Serville, 1823) / Salix fragilis L. × S. pentandra L.

Ph. plicadaphnoides Kopelke, 2007 / Salix daphnoides Vill.

Ph. polita (Zaddach, 1883) / Salix purpurea L.

Phyllocolpa sp. on Salix miyabeana Seemen.



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Saotis sp.:

Phyllocolpa alienata (Tischbein, 1846) / Salix aurita L.

Saotis albionis Kasp.:

Pontopristia amentorum Förster / catkins of Salix caprea L., S. repens L.

Saotis alpinator sp. n.:

Pontania obscura Kopelke, 2005 / Salix foetida Schleich.

Saotis granulator sp. n.:

Pontania bella (Zaddach, 1876) / Salix aurita L.

P. glabrifrons Benson, 1960 / Salix lanata L.

P. gallarum (Hartig, 1837) / Salix caprea L.

P. hastatae Vikberg, 1970 / Salix hastata L.

P. nivalis Vikberg, 1970 / Salix glauca L.

P. norvegica Kopelke, 1991 / Salix borealis Fries.

P. varia Kopelke, 1991 / Salix myrsinifolia Salisb.

Saotis pygidiator Kasp. et Kopelke:

Pontania breviserratae Kopelke, 1989 / Salix breviserrata Flod.

P. foetidae Kopelke, 1989 / Salix waldsteiniana Willd.

P. hastatae Vikberg, 1970 / Salix hastata L.

P. varia Kopelke, 1991 / Salix myrsinifolia Salisb.

A KEY TO THE EUROPEAN SPECIES OF SAOTIS 1. Females ................................................................. 2. —Males ................................................................... 23. [Males of Saotis are morphologically uniform and very

difficult for identification. In males of most spe-cies (in contrast to females), face yellow, fore and middle coxae and trochanters usually also yellow, often numerous yellow spots present on sides of thorax. Tergite 3 often with reddish tri-angular spots at base and at apex. Males of many species unknown. A short key below (couplets 23–28) is based on the reared material, using such biological character as specialization of species on Pontania or Phyllocolpa].

2. Mesothorax predominantly red; metasoma much longer than head and thorax combined.—Fla-

gellum with 27–33 segments ................................ ........................ 13. S. mirabilis Schmiedeknecht.

—Mesothorax predominantly black; metasoma shorter ........................................................................... 3.

3. Epipleura (lateral part of tergite ventrad of spiracle) of tergites 5 and 6 white ................................... 4.

—Epipleura of tergites 5 and 6 black, brown, or ru-fous, or whitish only at distal edge .................... 8.

4. Scutellum black; flagellum with 22–24 segments.—Tergites 6–8 predominantly black dorsally (with white band on hind margin); ovipositor sheath broad, light brown (Fig. 2); metasoma moderately compressed. Syntypes: fore wing 3–5 mm long; head wide; i. fl 1.1, 1st flagellar segment about 5 times as long as wide at apex (in other species, usually shorter); malar space short, i. gen 0.25; clypeus widely impressed along lower margin, convex above impression; propodeal areas very indistinct; i. terg 1.8; fore and middle coxae and all trochanters yellow (hind trochanter I darkened dorsally); hind coxa reddish or brownish; hind tibia yellowish white, weakly darkened at base and in apical 0.3 ............................................ ............................. 7. S. compressiuscula Thoms.

—Scutellum with yellow spot; flagellum with 25–29 segments ........................................................... 5.

5. Tergites 6–8 predominantly light yellowish (with black dorsal spot at base); ovipositor sheath yel-low, usually truncate at apex (Fig. 1). Meso-pleuron with distinct and rather dense punctures in lower 0.6. Antenna usually with 24–26 flagel-lomeres. Epipleura of tergites 4 and 5 sometimes separated by crease. Hind tibia reddish brown, but dirty white on posterior side in basal 0.6. —Syntypes: malar space rather long, i. gen 0.4; i. terg 1.5; mesoscutum with yellow central and anterolateral spots; scutellum dorsally (except for black median spot at base) and postscutellum yel-low; fore and middle coxae and trochanters yel-low; propodeum with distinct areola and apical area ................................ 5. S. brevispina Thoms.

—Tergites 6–8 predominantly black dorsally, with rather narrow white band on hind margin; ovi-positor sheath not as above (Figs. 4, 6, 14). Mesopleuron in lower 0.6 without distinct and dense punctures, almost smooth. Antenna usually with 26–29 flagellomeres. Epipleura of tergites 4

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and 5 not separated by crease. Hind tibia usually tricolorous: with broad white ring at middle, and with black and reddish pattern in basal 0.3 and at apex (Figs. 16, 17). Apical tergite of abdomen (tergite 8) very strongly convex in lateral view (Figs. 6, 14) ...................................................... 6.

6. Mesosternum red or yellowish brown, usually sepa-rated from black mesopleuron by yellow longitu-dinal band. Apical margin of hind tibia shining and projecting; hind spurs brownish (Fig. 10); segment 5 of hind tarsus 1.33 as long as

segment 3.—Syntypes (S. emarginata): flagellum of female with 29 segments, i. gen about 0.33 (in male, 0.1); face with two yellow spots below an-tennal sockets, and with yellow markings above clypeus, on malar space and on cheeks; mesoscu-tum with two yellow anterolateral spots; i. terg 1.75; hind tibia tricolorous; tergites black with white epipleura (Fig. 4) ........................................ ...... 3. S. bilineata Grav. (= emarginata Thoms.).

—Mesosternum and mesopleuron black. Hind tibia at apical margin not shining and not projecting, hind

Figs. 1–10. Genus Saotis, female (Ichneumonidae, Ctenopelmatinae): (1–8) apical part of abdomen [(1) S. brevispina, (2) S. com-pressiuscula, (3) S. varicoxa, (4) S. bilineata, (5) S. nigriscuta (holotype), (6) S. morleyi, (7) S. dorsata, (8) S. nigriventris]; (9) S. va-ricoxa, scutellum, dorsal view; (10) S. bilineata, apical part of hind tibia with spurs.



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spurs white; fifth segment of hind tarsus hardly longer than segment 3. Flagellum with 26–29 segments ........................................................... 7.

7. Tergite 1 about 1.6 times as long as wide (Fig. 20). Tergites 7 and 8 almost entirely brownish; tergite 8 strongly swollen in profile and with superficial transverse impression beyond middle (Fig. 14). i. fl 1.03–1.07; flagellar segment 2 about 3.8 times as long as wide; clypeus convex before lower margin. Fore wing 4.5–5.8; flagellum usu-ally with 27–30 segments ..................................... .... 20. S. tricolor Thoms. (var. liopleuris Thoms.)

—Tergite 1 about 2.0–2.3 times as long as wide. Ter-gites 7 and 8 white laterally, tergite 8 almost en-tirely white (except for dorsal spot at base) (Figs. 6, 33). i. fl 0.9–0.95; segment 5 of hind tar-sus not longer than segment 3.—i. terg 2.2; i. gen 0.28. Epipleura of tergites 3–8 and all ster-nites whitish, ovipositor sheath yellow (Fig. 6).

Legs light reddish; fore and middle coxae (except for base), trochanters (except for dark dorsal spots on hind trochanter I) yellow; hind tibia with white band at midlength (0.25), with fuscous base (0.3) and apex (0.45), spurs white (Fig. 16); hind tarsus black, with segments 1–4 whitish at ex-treme base . ............................................................ ... 14. Saotis morleyi Fitton (= albiventris Kasp.)

[this species was considered to be a junior syno-nym of S. liopleuris Thoms., but differs from the latter in having tergite I slender and coloration of apex of abdomen paler].

8. Tergites very densely covered with short hairs. i. gen about 0.2. All coxae brownish black; tho-rax entirely black.—Tergites of metasoma black to brown, without white pattern; apical tergites (except dorsally), hypopygium and ovipositor usually ochraceous; hind margin of tergite 8 with median projection (Fig. 8). Holotype: 23/24 flag-

Figs. 11–26. Genus Saotis, female (except for Figs. 21, 22) (Ichneumonidae, Ctenopelmatinae): (11–14) apical part of abdomen [(11) S. hoeli; (12) S. heteropus, lectotype; (13) S. ?renovatus; (14) S. tricolor (lectotype of L. liopleuris)]; (15) abdomen of S. longiventris, holotype; (16, 17) hind tibia [(16) S. morleyi; (17) S. ?renovata]; (18–20) tergite 1, dorsal view [(18) S. ?renovata, (19) S. albionis, (20) S. tricolor (lectotype of S. liopleuris)]; (21, 22) S. renovata (♂), lectotype [(21) tergite 1, dorsal view; (22) clypeus]; (23–26) dorsal areas of propodeum [(23) S. heteropus, (24) S. hoeli, (25) S. ?renovata, (26) S. albionis].

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ellar segments; clypeus impressed along lower margin; the body black to brown, with the follow-ing parts white: clypeus, mandibles, hind corners of pronotum, tegulae, fore and middle trochanters II; all coxae and hind trochanter black; hind tibia whitish, weakly fuscous at extreme base and brownish in apical 0.35; mesopleuron granulate; epipleura of tergite 3 (except at base) and tergites 4–8 not separated by crease ................................. .................................... 16. S. nigriventris Thoms.

—Tergites with hairs much sparser and longer; malar space usually longer; coxae red or black; thorax entirely black or with yellow markings ............ 9.

9. Scutellum with a pair of yellow basolateral spots. (Fig. 9). Epipleura of tergite 4 (sometimes also of tergite 5) separated by crease. Mesopleuron al-most smooth. Antenna with 22–26 flagellomeres; i. gen 0.45; i. nrv 0.08–0.15. Metasoma beyond apical part of tergite 1 smooth and almost pol-ished; hind tibia darkened dorsally, dirty whitish to dirty rufous in middle 0.5 ventrally. Lectotype: 24 flagellar segments; mesoscutum with anter-olateral yellow spots; fore and middle coxae and trochanters white to yellow; hind coxa and hind trochanter blackish brown (in other specimens sometimes red); sternites white; epipleura rusty rufous; fore and middle tibiae rufous, darkened dorsally at middle. Metasomal tergites (3)4–7 with white spot on hind margin dorsally (the spot angulate anteriorly); ovipositor sheath from yel-lowish brown (in type) to yellowish, truncate api-cally (Fig. 3) ......................................................... ........ 21. S. varicoxa Thoms. (compressa Szépl.).

—Scutellum with yellow apical spot or entirely black. Epipleura of tergites 4 and 5 not separated by crease (except for S. subarctor and sometimes in S. dorsata). Other characters not entirely as above .............................................................. 10.

10. Epipleura of tergites 4 and 5 separated by crease (Figs. 44, 45). Thorax entirely black (except for white tegulae). Tergites 7 and 8, epipleura of ter-gite 6 and ovipositor sheath ochreaceous-yellow, sternites light yellow; sheath truncate at apex. 20–23 flagellomeres; i. fl about 1.1; i. gen 0.4–0.55. Nervellus intercepted close to its lower end (i. nrv 0.1–0.23). Nervulus postfurcal; i. terg 1.75–2.1. Areola and basal area of propodeum narrow. Clypeus convex on lower margin, with small sharp lateral lobes (Fig. 36). Mandible of-

ten with basal blackish spot. All coxae and tro-chanters predominantly black ............................... ......................................... 19. S. subarctor sp. n.

—Epipleura of tergite 4 and 5 not separated by crease (occasionally separated in S. dorsata). Meso-scutum and scutellum sometimes with yellow markings. Tergites 7 and 8, epipleura of tergite 6 and ovipositor sheath darker.—Other characters not entirely as above ....................................... 11.

11. Apical part of metasomal tergite 2 and median longitudinal band on tergite 3 reddish brown, ter-gite 4 usually reddish mediodorsally. Ovipositor sheath rather narrow (Fig. 7), about as long as segment 2 of hind tarsus. Mesoscutum black; scutellum black, occasionally reddish brown at apex; i. terg about 2.2. Sternites 1–4 reddish yel-low, sternites 5 and 6 rufous.—Legs light red, hind tibia whitish, slightly reddish at base and reddish brown in apical 0.27.—Holotype: fore wing 4.2 mm, flagellum with 26 segments (in other examined specimens with 22–24 segments); i. fl 0.9; i. gen 0.3; clypeus widely convex on lower margin; epipleura of tergites 2–3 (–5) sepa-rated by crease; ovipositor sheath brownish ......... ............................................ 8. S. dorsata Thoms.

—Metasomal tergites 2–4 dorsally black, without red-dish brown markings. Ovipositor sheath wider and shorter. Mesoscutum black or often with an-terolateral yellow spots; scutellum black or with yellow dorsal spot ........................................... 12.

12. Thorax (except for speculum) and tergites 1–4(5) matte, evenly covered with distinct dense gra-nulation. Malar space very short (i. gen about 0.25). Sternites blackish, without distinct longi-tudinal fold; hind edge of hypopygium truncate. Tergites 2–4 subequal and almost not compressed (Fig. 39).—Head not narrowed, temples beyond eyes parallel (Fig. 37). 21–24 flagellomeres; i. fl about 1.0; nervulus postfurcal; i. nrv about 0.33; i. terg 1.7–1.8. Propodeum with basal area and areola obliterated (Fig. 38). Tergites 2–4 about 1.5 times as long as wide at base, tergites 7 and 8 usually projecting beyond apex of hypopy-gium (Fig. 39). Mesoscutum, scutellum and ab-dominal tergites black; coxae and hind trochanter I black or reddish; other trochanters predomi-nantly yellow ................... 9. S. granulator sp. n.

—Thorax and tergites 1–4(5) predominantly or partly polished, or without distinct granulation. Malar



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space longer, i. gen 0.3–0.5 (except for S. cau-casica). Sternites usually yellowish, with distinct longitudinal fold; apex of hypopygium usually projecting backward ....................................... 13.

13. Abdomen strongly elongate, i. terg 2.8 (in holo-type), black, tergites 2 and 3 without white apical band, tergites 7, 8 and ovipositor sheath brown (Fig. 15); sternites fuscous, hypopygium brown-ish yellow; ovipositor sheath as wide as hind fe-mur; tergites 2 and 3 combined 1.4 times as long as hind femur. Head strongly narrowed backward (Fig. 50). Pterostigma yellowish rufous. Face black with a pair of small yellow spots distal to clypeal foveae.—Holotype: fore wing 3.8 mm long; antennae missing; i. fl 0.9, i. gen 0.33, i. fem 4.7, i. tars 3.9 : 1.9 : 1.45 : 0.95 : 1.3; ner-vulus almost interstitial (weakly postfurcal). Mesopleuron with smoothened granulation and sparse hairs; metapleuron with superficial granu-lation; propodeal areas indistinct, pleural carina absent; antenna reddish brown; clypeus, small spots on malar space immediately above upper angle of mandible (Fig. 51), mandible, palpi, hind corners of pronotum, apex of epimeron, all tegu-lae yellow. Legs pale rufous; hind tibia whitish with basal 0.1 and apical 0.27 reddish brown ...... .................................... 12. S. longiventris Thoms.

—Abdomen less elongate i. terg 1.5–2.1 (except for S. morleyi) and usually with hind margin of ter-gites (2)3–7 whitish (Figs. 42, 43); sternites often yellow; tergites 2 and 3 combined usually 1.0–1.2 as long as hind femur. Pterostigma usually brownish or blackish (except for S. pygidiator). Face completely black (except for S. alpinator and occasionally in S. pygidiator) .................. 14.

14. Hypopygium densely covered with erect hairs per-pendicular to its surface or slightly slanting to sternite base (Fig. 42). Head not narrowed behind eyes, temples swollen (Fig. 30). i. fl 1.25 ± 0.1; first flagellar segment 1.4–1.6 times as long as segment 2. Flagellum 23–28-segmented, about as long as fore wing (± 0.1). Pterostigma pale brown to yellowish. Mesoscutum with yellow anter-olateral spots; scutellum black. Nervulus intersti-tial to slightly postfurcal; i. nrv 0.1–0.2; i. terg 1.7–2.0. Ovipositor sheath (Fig. 42) usually red-dish brown, about 0.8 times as wide as hind fe-mur (17. S. pygidiator sp. n.) ........................ 14a.

—Hairs on hypopygium sparse and reclinate (slanting toward sternite apex). Head usually slightly or

distinctly narrowed.—Other characters not en-tirely as above ................................................. 15.

14a. Flagellum slightly longer (1.02–1.1) than fore wing. Face entirely black. Flagellum darkened dorsally ......................................................... 14b.

—Flagellum slightly shorter than fore wing (about 0.95 times as long as latter). Face often with yel-low spots near clypeal fovea. Flagellum almost uniformly yellowish rufous. Pterostigma pale yel-lowish brown ................................................ 14c.

14b. Coxae and trochanters I black. 23–26 flagel-lomeres. Pterostigma pale brown with dark mar-gins.—West Palaearctic ........................................ .. S. pygidiator pygidiator Kasparyan et Kopelke.

—Coxae, trochanters and femora uniformly light red. 26–28 flagellomeres. Pterostigma blackish brown with small basal white marking.—Transbaikalia .............................. S. pygidiator ivan Kasparyan.

14c. i. gen 0.4– 0.5; second flagellar segment 4.5–4.8 times as long as wide at midlength. Hind tibia almost uniformly yellowish rufous.—Northern Siberia ............... S. pygidiator arctor Kasparyan.

—i. gen 0.27–0.33; second flagellar segment 5–6 times as long as wide at midlength. Hind tibia yellowish rufous with fuscous markings before base and before apex.—Nearctic Region ............. ....................... S. pygidiator nearctor Kasparyan.

15. Ovipositor sheath very wide (1.1–1.3 as wide as hind femur), black or dark brown (Figs. 5, 41, 43) ......................................................................... 16.

—Ovipositor sheath not wider than hind femur and usually paler .................................................... 19.

16. Antenna with (24)25–27 flagellar segments; basal flagellomeres slender—i. fl about 1.25, first flag-ellomere about 8 times as long as wide at mid-length. Clypeus immediately above lower margin with distinct median convexity (Fig. 34). Ovi-positor sheath black, very wide, about 1.2 times as wide as hind femur (Fig. 5). Nervulus intersti-tial or weakly postfurcal; spurs of hind tibia brownish. All coxae, trochanters and femora usu-ally light reddish.—Holotype: fore wing 4.9 mm; flagellum with 26 segments, slightly shorter (0.97) than fore wing; propodeal carinae vestig-ial; hind margin of tergites 3–7 white (only on dorsum); sternites 3–6 yellowish white (Fig. 5); hind tibia pale brownish with ventral side in basal 0.7 dirty-whitish ........... 15. S. nigriscuta Thoms.

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—Antenna with 19–24(?25) flagellomeres; i. fl 0.9–1.15; transverse basal convexity of clypeus not projecting medially toward sharp lower margin. Small species with fore wing 3.0–3.9 mm (except for S. heteropus); nrl postfurcal ..................... 17.

17. Flagellar segments 1 and 2 combined hardly as long as maximum diameter of eye (in type, i. fl 0.93); i. gen about 0.5; i. nrv 0.28–0.40. Face with yellow spots immediately above clypeus (Fig. 28). Tergites 3–7 on hind margin with rather wide yellow band and their epipleura broadly yel-low in hind corners (Fig. 43). Holotype: fore wing 3.1 mm; flagellum 22-segmented; areas of propodeum distinct, apical area of propodeum

rather high; mesoscutum and scutellum entirely black; all coxae and trochanters I black ................ ............................................. 2. S. alpinator sp. n.

—Flagellar segments 1 and 2 combined longer than maximum diameter of eye (i. fl 1.14–1.25); i. gen 0.33–0.5; i. nrv 0.15–0.3. Face entirely black. Tergites 3–7 on hind margin with narrow whitish band and their epipleura black, ovipositor sheath blackish (Figs. 12, 41) .................................... 18.

18. Hind femur black with brownish apex; all coxae and trochanters I black. Fore wing 4–5 mm long; antenna with 23–24 flagellomeres. Areola of pro-podeum very narrow and separated from apical area (Fig. 23). Segment 3 of hind tarsus 1.2–1.3

Figs. 27–36. Genus Saotis, female (Ichneumonidae, Ctenopelmatinae): (27–29) head, anterior view, and two basal flagellomeres [(27) S. albionis, holotype; (28) S. alpinator; (29) S. pygidiator arctor]; (30) S. pygidiator pygidiator, head, dorsal view; (31–33) apical part of abdomen, lateral view [(31) S. albionis, holotype; (32) S. albionis, ex Pontopristia; (33) S. morleyi, holotype]; (34–36) clypeus [(34) S. nigriscuta, (35) S. albionis, (36) S. subarctor].



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times as long as segment 5. Thorax black with only tegulae white. Lectotype: fore wing 4.8 mm; antennae missing; i. fl 1.12; mandible brownish (in other examined specimens yellow); nervulus weakly postfurcal; i. gen 0.35; i. nrv 0.25; tergite 1 beyond spiracles with parallel margins (Fig. 55); i. terg 2.1; trochanters II blackish at base; hind femur in lectotype dark brown with brownish apical part (0.25); hind tibia pale rufous with reddish apex (0.25) and fuscous at very base (0.05); tergites 3–7 black, with yellowish dor-soapical spot and white apical margin; sternites pale brown, sternites 5 and 6 yellowish with brownish basal part; ovipositor sheath wide, blackish (Fig. 12). Ovipositor sheath wide, black with brownish apex (Fig. 12) ............................... ....................................... 10. S. heteropus Thoms.

—Hind femur red; coxae red, fuscous at base; tro-chanters yellow, hind trochanters I red; hind spurs white (in type). Holotype: fore wing 3.7 mm; flagellum 21-segmented; i. gen about 0.4–0.5. Areola of propodeum absent or obliterated. Mesoscutum with yellow anterolateral spot; scu-tellum black ......................... 4. S. boreator sp. n.

19. First tergite 2.2–2.9 times as long as wide, its late-ral margins beyond spiracles subparallel (Fig. 19); flagellum with 21–24 segments; i. gen about 0.4.—Tergite 2 polished in apical 0.3; api-cal tergites brownish, strongly compressed and rather strongly convex; ovipositor sheath light brown to brown (Figs. 12, 13). Fore and middle coxae and trochanters yellow. Hind tibia whitish, weakly infuscate at base (0.08) and reddish

Figs. 37–45. Genus Saotis, female (Ichneumonidae, Ctenopelmatinae): (37–39) S. granulator [(37) head, dorsal view; (38) propodeum and tergite 1, dorsal view; (39) abdomen, lateral view]; (40) S. pygidiator var. helvetica, propodeum and tergite 1, dorsal view; (41–45) apical part of abdomen, lateral view [(41) S. boreator; (42) S. pygydiator; (43) S. alpinator; (44, 45) S. subarctor].

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brown in apical 0.45 (Fig. 17). i. fl 0.97, i. nrv 0.1–0.2 ............................................18. S. renovata Morley? (= morleyi sensu Kasparyan, 2007).

—First tergite about 1.6–1.8 times as long as wide, its lateral margins beyond spiracles divergent (Fig. 20); flagellum with 20–23 segments or with 26–30 segments (rarely with 24 segments); i. gen about 0.2–0.35; nervulus postfurcal ............... 20.

20. Tergite 8 in profile very strongly convex and immediately beyond middle with light transverse impression; apical tergites brownish, strongly compressed (Fig. 14). i. gen about 0.25. Flagel-lum usually with 26–30 segments (small specimens with fore wing 3.5–4.0 mm may have 24–25 flagellomeres). i. fl about 1.05; i. fem 4.0–4.4. Hind tibia in basal 0.3 with wide dark basal ring, usually tricolorous: white at middle, black with reddish at base and at apex (as in Fig. 16). Tergite 2 with very fine granulation, almost polished in apical 0.3; epi-pleura of tergite not 3 separated by a crease. Segment 3 of hind tarsus usually slightly shorter than segment 5. i. nrv about 0.33 ......................... ................................................. S. tricolor Thoms.

—Tergite 8 different (Figs. 32, 49). Flagellum usually with 20–23 segments (rarely with up to 26 se-gments in S. albionis), shorter or only slightly longer than fore wing ...................................... 21.

21. i. gen 0.22. Tergite 8 slightly projecting (Fig. 49). Holotype: fore wing 5 mm; flagellum 23-seg-mented; i. fl 1.06; i. fem 4.2; i. nrv 0.28; tergite 1 weakly narrowed toward base (Fig. 48); ovi-positor sheath about 0.8 times as wide as hind femur; face black with small yellow spot between mandibular condyle and eye (Fig. 47) .................. .......................................... 6. S. caucasicus Kasp.

—i. gen about 0.35. Tergite 8 strongly projecting, in apical half somewhat similar to hypopygium (Figs. 11, 31, 32). i. fl 0.9–1.0; i. fem about 4.5; tarsal claws with 2 teeth at base; i. nrv 0.25–0.33. Hosts: ?larvae of Pontopristia (Nematinae) in catkins of Salix ............................................... 22.

22. Coxae, femora and trochanters predominantly red-dish (trochantelli yellow); thorax usually with yellow spots (at least hind corners of pronotum yellow; in holotype, mesonotum anteriorly with a pair of yellow dots, and scutellum with small

yellowish spot). Sternites yellow; i. terg 1.5–1.7.—Holotype: fore wing 4.8 mm, flagellum 4.1 mm long, 23-segmented; i. fl about 0.9; i. fem 4.5. Clypeus with impression before sharp lower mar-gin. Tergite 1 broad posteriorly (Fig. 19). Tergite 2 entirely granulate; epipleura of tergites 2 and 3 separated by a crease. Tergite 8 dorsally evenly and rather weakly convex, ovipositor sheath dark (Fig. 31).—Mesopleuron with superficial granulation, almost smooth and with fine punc-tures; metapleuron granulate; propodeal areas distinct (Fig. 26). i. tars 2.8 : 1.5 : 1.15 : 0.75 : 1.15 (segment 3 about as long as segment 5); i. nrv about 0.25. Tergites smooth behind poste-rior 0.3 of second tergite and almost hairless; ter-gites 3–7 on hind edges moderately concave and with narrow pale band. Hind tibia white in mid-dle 0.35 (white coloration expanding ventrally almost to base of tibia), light grayish in basal 0.25 and brownish gray in apical 0.4; spurs white, hind tarsus grayish, segments 1–4 whitish at very base ...................................... 1. S. albionis Kasp.

—Coxae, femora and trochanters predominantly black; thorax black with only tegulae white. Sternites blackish brown (Fig. 11); i. terg about 1.8. —Greenland .......................... 11. S. hoeli Roman

Males

23. Parasites of Pontania ........................................ 24.

—Parasites of Phyllocolpa ..................................... 26.

24. Malar space very short (i. gen 0.2). Mesopleuron (except for speculum) and mesoscutum evenly granulate, black. Antenna with 20–22 flagellomeres (in female up to 24) ................................. .......................................... 9. S. granulator sp. n.

—Malar space longer (i. gen 0.33–0.5). Mesopleuron in upper half and mesoscutum almost smooth, without distinct granulation; mesoscutum with anterolateral yellow spots. Antenna with 22–26 flagellomeres ................................................... 25.

25. Antenna with 22 flagellomeres, i. fl 1.0. Head nar-rowed backward. Scape and pedicel entirely light yellow ventrally. Hind femora light reddish. Alps ............................................. 2. S. alpinator sp. n.

—Antenna with (23)24–26 flagellomeres, i. fl 1.1–1.2. Head not narrowed backward, temples beyond



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eyes parallel or swollen. Scape ventrally usually brownish with yellowish apical marking, pedicel may be completely brownish. Hind femora from reddish brown to blackish .................................... .............. 17. S. pygidiator Kasparyan et Kopelke.

26. Antenna with 20–22 flagellomeres.—i. fl 1.15–1.25; first segment of flagellum without distinct rhinaria, about 6.5 times as long as wide at middle and 1.6 times as long as second

flagellomere. i. gen about 0.33, i. terg 2.0, i. nrv about 0.1. Hind coxae and trochanters I reddish brown to blackish dorsally. Hind tibia pale ru-fous, at apex and dorsally brownish ..................... .......................................... 19. S. subarctor sp. n.

—Antenna with 24–29 flagellomeres.—Hind coxae and hind trochanters usually uniformly pale red-dish or yellow. Hind tibia in middle part (0.2–0.4) whitish. i. fl 1.0–1.1 ........................ 27.

Figs. 46–55. Genus Saotis, female (Ichneumonidae, Ctenopelmatinae): (46–49) S. caucasica, holotype [(46) head, dorsal view; (47) head, anterior view, and two basal flagellomeres; (48) tergite 1, dorsal view; (49) apex of abdomen, lateral view; (50, 51) S. lon-giventris, holotype [(50) head, dorsal view; (51) head, anterior view, and two basal flagellomeres]; (52–55) S. heteropus, lectotype [(52) clypeus; (53) head, dorsal view; (54) eye, lateral view, and two basal flagellomeres; (55) tergite 1, dorsal view].

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27. Antenna with 24–25 flagellomeres. i. gen about 0.44; i. terg about 2.5 (lectotype—Figs. 21, 22) .. 18. S. renovata Morley

—Antenna with 26–29 flagellomeres. i. gen about 0.25, i. terg 2.0–2.9. First segment of flagellum with distinct rhinaria, about 4–5.5 times as long as wide at middle and 1.3–1.4 times as long as second flagellomere ....................................... 28.

28. Tergite 1 2.0–2.4 times as long as wide. First seg-ment of flagellum 5 times as long as wide at mid-dle ..................................... 20. S. tricolor Thoms.

—Tergite 1 at least 2.6 times as long as wide. First segment of flagellum 4.0–4.5 times as long as wide at middle .................... 14. S. morleyi Fitton

A REVIEW OF PALAEARCTIC SPECIES OF SAOTIS

1. Saotis albionis Kasparyan, 2007 (in: Kasparyan and Khalaim, 2007)

(Figs. 19, 26, 27, 31, 32, 35).

Kasparyan, Shaw, 2003 : 353–355, ♀ (as Saotis sp. 2; key, fig.; England); Kasparyan, Khalaim, 2007 : 550, ♀ [holotypus: ♀, England, Oxfordshire (NMS); description in key; fig.].

Diagnosis. Saotis albionis can be distinguished from the congeners by a large hypopygium, by tergites 7 and 8 (Figs. 31, 32) strongly projecting and straight dorsally in profile, and by the shape of the pygostyli. The species is closely related to S. clypeata Ashmead, 1902 from North America (Alaska) and S. hoeli Ro-man, 1933 from Greenland, but differs from both in having red hind coxae and femora and scutellum with yellow apical spot.—Holotype: fore wing 4.8 mm; flagellum 4.1 mm long (0.83 times as long as fore wing), with 23 flagellomeres. i. fl about 0.9–1.0; clypeus impressed along sharp lower margin (Fig. 35); i. gen about 0.35; i. nrv 0.25–0.33; i. fem 4.5; i. tars in holotype 2.8 : 1.5 : 1.1 : 0.7 : 1.1 (third segment as long as fifth segment); tarsal claws with 2 rather high teeth at base; propodeum with distinct areas (Fig. 26). Mesopleuron superficially granulate, almost smooth with fine punctures; metapleuron granulate. Tergite 1 broad on posterior margin (Fig. 19); i. terg 1.5–1.7; tergite 2 almost entirely granulate; epipleura of tergites 2 and 3 separated by crease; tergites 3–7 smooth and almost bare, their hind margins moderately concave at middle and with narrow pale band. Thorax black, usu-ally with yellow markings, at least hind corners of

pronotum yellow; in holotype mesoscutum anteriorly with a pair of yellow dots and scutellum with small yellow marking. Coxae, femora and trochanters pre-dominantly red, trochanters II yellow; hind tibia of holotype white in middle third (on ventral side white coloration extending almost to base of tibia), light gray in basal 0.25 and brownish gray in apical 0.4; spurs white; hind tarsus grayish, with segments 1–4 whitish at extreme base. Sternites yellow.

Material. Great Britain: 1 ♀ (holotype). Finland: “Munksnäs, Pontopristia suavis,” 1 ♀—3.5.1951, 1 ♂—29.4.1951 (Lindqvist). Russia: 1 ♀ (paratype), Yaroslavl Province.

Host. Larvae of Pontopristia amentorum Först. in catkins of Salix caprea, S. repens.

2. Saotis alpinator Kasparyan et Kopelke, sp. n. (Figs. 28, 43)

Saotis alpinator resembles S. nigriscuta Thoms. in having a similar coloration and broad dark ovipositor sheath, but the new species is smaller, its clypeus is without median convexity apically, the flagellum is about 22-segmented (26–27-segmented in S nigris-cuta), and the malar space is longer. Structurally the new species resembles S. compressiventris; both spe-cies have a small body, 22-segmented flagellum, short abdomen, and wide ovipositor sheath. Saotis alpinator differs from S. compressiventris in having a malar space longer, two basal flagellomeres shorter, and epipleura of tergite 4 weakly separated and darkened.

Description. Female (holotype). Fore wing 3.1 mm. Antenna with 22 flagellar segments; i. fl 0.93. Head not narrowed beyond eyes, temples parallel. i. gen about 0.5. Clypeus convex in profile, its lower margin sharp and evenly concave. Mesoscutum centrally finely granulate and matte, with lateral parts smooth and shiny. Mesopleuron in upper part almost smooth, in lower half granulate. Metapleuron matte, entirely granulate. Propodeum with distinct dorsal areas, but lateral longitudinal carinae absent; pleural carina com-plete; i. ar. ap 0.36. Nervulus distinctly postfurcal, nervellus intercepted about at lower 0.28. i. fem 4.6, i. tars 5.2 : 2.9 :2.4 : 1.4 : 2.2.

Tergites 1–3 matte, more or less evenly covered with short hairs, finely granulate, with hind margin polished. First tergite with subparallel lateral margins behind spiracles, i. terg 1.7; its dorsal carinae extend-ing to apical 0.7 of tergite. Tergites 2 and 3 finely granulate, smooth and shiny in apical 0.2, their



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epipleura separated by crease; subsequent tergites almost smooth; epipleura of tergite 4 weakly sepa-rated. Hind edge of tergites 6 and 7 slightly concave. Ovipositor sheath broad (Fig. 43), 1.2 as broad as hind femur.

Antenna brown with scape at apical 0.4 yellowish ventrally. Head and thorax black with clypeus, small spot on face immediately above clypeal fovea, mandi-ble, palpi, hind corners of pronotum, very small lateral spot on central lobe of mesoscutum anteriorly, and tegulae yellow; apex of epimeron slightly reddish. Coxae black with apical half of fore coxa and apex of middle coxae light yellow; trochanters I black, apex of fore and middle trochanters yellow and fore trochant-ers reddish posteriorly; all trochanters II yellow; legs beyond trochanters reddish rufous with tarsi brownish rufous, hind femora anteriorly slightly infuscate; hind tibia pale rufous with brownish dorsolateral spot in basal 0.3, and entirely brownish in apical 0.35. Pterostigma pale with brownish margins. Abdominal tergites blackish with brown tinge; tergites 3–8 with yellowish hind margin (Fig. 43); epipleura of tergites 5 and 6 pale reddish brown on lateral margin. Sternites yellow.

Male similar to female, but antenna pale brown with scape, pedicel and flagellomeres 1–3 yellow ventrally; face, malar space, spot behind mandible, large anter-olateral triangle spot on mesoscutum, subtegular ridge, and apex of epimeron yellow. Tergites 3–8 reddish brown; fore coxae, middle coxae (except for base) and all trochanters light yellow; hind tibia pale rufous with apical 0.35 reddish.

Material. Holotype: ♀, Austria, Tirol, Zillertaler Alpen, Hintertux/Weitental, 29.8. 1991, ex Pontania sp. 1 / foetida (Kopelke); Zucht Nr 11-T1 / 1991, Sl, Schlüpfdatum 02.06 1992. Paratype: 1 ♂, with labels as in holotype.

Host. Pontania obscura Kopelke 2005.

3. Saotis bilineata (Gravenhorst, 1829) (Figs. 4, 10) Gravenhorst, 1829 : 175, ♀ [Tryphon bilineatus;

holotype (lost): ♀, Poland: ‘Warmbrunn”= Tseplice-Slenski-Zdruj]; Woldstedt, 1878 : 41 (Mesoleius; Russia: St. Petersburg); Pfankuch, 1906 : 218 (Saotis; = emarginatus); Schmiedeknecht, 1914 : 2808, 2811 (key, description); Kasparyan, Shaw, 2003 : 352, 353 (key, fig.); Kasparyan, Khalaim, 2007 : 548 (key, fig.).—emarginatus Thomson, 1883 : 933, ♀ ♂ [Mesoleius (Saotus); syntypi: 2 ♀ 1 ♂, examinavi,

Sweden, “Ört”= Skåne, Örtofta (Mus. Lund)]; Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Fitton, 1982 : 56 (type); Kasparyan, 1996 : 196 (= ulbrichti).—ulbrichti Habermehl, 1910 : 690, ♀ (Homotropus; typum examinavi: ♀, Germany: Crefeld (SMF)].

Diagnosis (is given on syntypes emarginatus Thoms.). Antenna with 29 flagellomeres; malar space short, i. gen less than 0.3 in female and about 0.1 in male; Mesopleuron shiny, almost smooth; female with yellow spots below antennal sockets, immediately above clypeus, on genae beyond the mandibles. Mesosternum red with longitudinal yellow band in lower part of mesopleuron; anterolateral large spots on mesoscutum and spot on scutellum yellow (in the original description of Tryphon bilineatus, such yellow spots on thorax noted only before and below tegulae and on scutellum; red coloration of mesosternum not mentioned). Legs red with yellow fore and middle coxae and trochanters; hind tarsi brownish, hind tibia tricolor—blackish, with red base and apex and with wide white ring at middle (as in S. morleyi, Fig. 16, but spurs brownish); apical margin of hind tibia shiny and projecting posteriorly (Fig. 10); color pattern of abdomen—Fig. 4. Male with face completely yellow; thorax yellow, with black transverse stripe on prono-tum, black spot on mesoscutum and on upper part mesopleuron; metapleuron and propodeum entirely black.

Material. Sweden: 2 ♀ 1 ♂ (syntypes of S. emarginatus) and 1 ♀ among syntypes of liopleuris (Mus. Lund).

Hosts. We did not rear specimens of this species. [There are records on parasitism in “Pontania proxima Lep.” (Brischke, 1878, Fulmek, 1968) and in Hoplocampa (2 species) (Brischke, 1878, Dalla Torre, 1902; Dominquez, 1950)].

4. Saotis boreator Kasparyan & Kopelke, sp. n. (Fig. 41)

Diagnosis. Saotis boreator structurally closely re-sembles S. heteropus Thoms. especially in having broad black ovipositor sheath, and rather slender basal flagellar segments; but the coxae, trochanters, and hind femora of S. boreator are red (predominantly black in S. heteropus), the clypeus is flat, and the fla-gellum consists of 19–21 segments (22–24-segmented in S. heteropus and about 26–27-segmented in S. ni-griscuta). S. boreator differs from S. alpinator (an-other small species with broad ovipositor sheath) in

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having slender basal flagellar segments and darker face and abdomen (cf. Figs. 41 and 43).

Description. Female (holotype). Fore wing 3.7 mm. Flagellum 3.3 mm long, with 21 flagellomeres; i. fl 1.1. Head weakly narrowed, temples subparallel be-yond the eyes; i. gen about 0.5. Clypeus rather flat (except for basal part), its lower margin sharp and broadly concave. Mesoscutum matte and finely granu-late on central lobe, smooth and shiny on lateral parts. Mesopleuron in upper part almost smooth with longi-tudinal striation under subtegular ridge, in lower half finely granulate. Metapleuron matte, entirely granu-late. Propodeum with distinct dorsal areas, but lateral longitudinal carinae absent; pleural carina complete, areola narrow; i. ar. ap 0.39. Nervulus distinctly post-furcal, nervellus intercepted about at lower 0.2. i. fem 4.6, i. tars 3.3 : 1.6 : 1.3 : 0.8 : 1.2. Abdominal tergites finely granulate, more or less evenly covered with short hairs. Lateral margins of first tergite slightly divergence behind spiracles, i. terg 1.8; its dorsal cari-nae extend to the middle of tergite. Epipleura of ter-gites 2–3 separated by crease; subsequent tergites shiny with fine granulation, their epipleura not sepa-rated. Hind edge of tergites 6 and 7 very weakly con-cave. Ovipositor sheath broad (Fig. 41), about 1.2 as broad as lateral width of hind femur.

Antenna brown, with small yellowish ventral spot at apical 0.3 of scape. Head and thorax black with clypeus, mandible, palpi, hind corner of pronotum, small anterolateral spot on central lobe of mesoscu-tum, and tegulae yellow; apex of epimeron slightly reddish. Fore and middle legs pale reddish with a base of fore coxae slightly infuscate, trochanters yellowish and middle tarsi brownish. Hind coxa reddish brown with base dorsally almost black; hind trochanters red with yellow spots; hind tibia dirty whitish, light gray-ish dorsally at basal 0.3, and entirely grayish at apical 0.37; hind spurs white; hind tarsus grayish. Pterostigma pale brown with brown margins. Abdomi-nal tergites blackish; tergites 3–8 with narrow yellow band on hind margin (Fig. 41). Sternites yellow; ster-nite 6 weakly darkened at base.

Variability. Paratype from Norway chromatically similar to holotype; flagellum 19-segmented. Paratype from Finland larger: fore wing 4.3 mm, flagellum 22-segmented, anterolateral yellow spots on meso-scutum very large. Specimens from Canada (British Columbia, Stone Mt. Pk., 3800 ft, 15–18.7.1973, leg. H. et M. Townes) (AEI) very similar to this species,

but all having flagellum 1.1 times as long as fore wing; maybe they must be consider as subspecies.

Material. Holotype: ♀, Norway, N/Finnmark, Lak-selv/Stabursnes, 02.08.2001 (Kopelke); Zucht: Ko-pelke Nr SZ8/2001, Schlüpfdatum 13.05.2002; ex Phyllocolpa nudipectus (labeled: Phyllocolpa sp.) / Salix phylicifolia / verdrillte Blattrolle. Paratypes. Norway: N/Finnmark, S. Varanger, Nyrud, 06.08.2001 (Kopelke); Zucht: Kopelke Nr SZ25 / 2001, Schlüpfdatum 22.05.2002; ex Phyllocolpa nudipectus (labeled: Phyllocolpa sp.) / Salix phylicifo-lia / verdrillte Blattrolle (Kopelke det.).” Finland: 1 ♀, Suomi, EH, Urjala, 13.7.1966 (T. Reilin) (Mus. Turku).

Hosts. Phyllocolpa nudipectus (Vikberg, 1965).

5. Saotis brevispina (Thomson, 1883) (Fig. 1)

Thomson, 1883 : 934, ♀, ♂ [Mesoleius (Saotus); syntypi examinavi: 8 ♀, Sweden, “Lund” (Mus. Lund)], 1895 : 2018 (Saotus); Dalla Torre, 1902 : 275 (Saotis; literature); Schmiedeknecht, 1914 : 2814 (key, description); Perkins, 1962 : 450; Fitton, 1982 : 55 (type); Kasparyan, Shaw, 2003 : 352, 353 (key, figs.); Kasparyan, Khalaim, 2007 : 548 (key, figs.).

Diagnosis. The species belongs to the “brevispina” species group. However, such a character as the epi-pleura of tergites 4 and 5 separated by a crease, which is typical of the group, is not always distinct in this species. S. brevispina differs from S. subarctor in yel-low coloration of the lateral margins of the scutellum and of the epimeron, in the presence of large antero-lateral yellow spots on the mesoscutum, in yellow fore and middle coxae and trochanters, and light reddish hind coxae and femora. In these characters of colora-tion, S. brevispina is similar to S. varicoxa, but differs from it in the presence of a yellow spot at the centre of the mesoscutum, yellowish-rufous coloration of ter-gites 6–8 (Fig. 1) and epipleura of tergites 5 and 6, in the long antennae (in S. brevispina, the flagellum is longer than the fore wing; in S. varicoxa, it is slightly shorter or as long as the fore wing). S. brevispina also differs from both these species in having the lower half of the mesopleuron covered with distinct punc-tures.

Syntypes. Fore wing 4–5 mm. Antenna with 23–26 flagellomeres; flagellum about 1.1 as long as fore wing; first flagellar segment about 1.6 times as long as second segment; i. fl 1.1–1.2; i. gen 0.4–0.5; i. fem 4.3;



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Mesopleuron shiny, with moderately fine and rather dense punctures on its lower half; areola of pro-podeum moderately wide, separated from apical area; i. terg 1.5; tergite 2 subquadrate; abdomen beyond the tergite 3 very strongly compressed, knife-like, with separate erected hairs; epipleura of tergites 4 and 5 often separated by crease; ovipositor sheath truncated at apex (Fig. 1). Body black; clypeus, mandibles, usu-ally a weak spot behind their lower corner on temple, collar and hind corners of pronotum, tegulae, large dorsolateral spots on mesoscutum and spot at its cen-tre, postscutellum, Scutellum dorsally almost entirely (except for black median spot at base) yellow. Legs red; fore and middle coxae and trochanters and tro-chanters II of hind legs yellow; hind tibia dorsally and in apical 0.4 brownish red or brownish, its ventral part in basal 0.6 yellowish or dirty-white; hind tarsi brownish. Hind margins of tergites (except for tergite 1) whitish, 3 last tergites and sternites almost entirely yellowish-rufous (Fig. 1).

Material. Sweden: 8 ♀ (syntypes) from collection of Thomson (including 4 ♀ on one pin) (Mus. Lund). Poland: 1 ♀, Krynki at Bialostok, 10.08.2002 (J. Sa-woniewicz) (ZIN).

Hosts. We did not rear specimens of this species. [There are records on parasitism in Pontania viminalis (L.) and Phyllocolpa leucapsis Tischb. (Fulmek, 1968)].

6. Saotis caucasica Kasparyan, 2009 (Figs. 46–49)

Kasparyan, 2009 : 118, ♀ [holotype: ♀, Russia, Ka-rachai-Cherkess, Arkhyz (ZIN)].

Diagnosis. Saotis caucasica closely resembles S. compressiuscula, but is considerably larger, its hind femora are wider (i. fem 4.2), i. nrv is equal to 0.28; tergite 1 is weakly narrowed to the base (Fig. 48); the coloration is similar to that in S. compressiuscula, but the tergites are darker (cf. Figs. 2 and 49). It differs from the other species in very short malar space (see also couplet 21 of the key).

Material. Holotype.

7. Saotis compressiuscula (Thomson, 1883) (Fig. 2)

Thomson, 1883 : 934, ♀ ♂ [Mesoleius (Saotus); syntypi examinavi, “Ringsjön, Skåne,” Sweden (Mus. Lund)], 1894 : 2019 (Saotus); Dalla Torre, 1902 : 273 (Saotis; bibliogr.); Morley, 1911 : 170 (description);

Schmiedeknecht, 1914 : 2818 (key, description); Fitton, 1982 : 55 (type); Kasparyan, Shaw, 2003 : 351, 352 (England; fig., key); Kasparyan, Khalaim, 2007 : 548 (key, fig.).

Description. Small insects with body about 4 mm long, length of fore wing 3.5 mm. Head large, about 1.5 times as wide as mesoscutum between tegulae; antenna with 22–24 flagellomeres; i. fl 1.1; malar space short, i. gen 0.25; carinae of propodeum very weak; i. fem 5.0; i. nrv about 0.15; i. terg 1.55–1.75; tergite 1 strongly tapered to base, (at apical margin 2.5 times as wide as its minimum width at base); abdomen moderately compressed, tergites 2 and 3 almost quad-rate. Ovipositor sheath wide, roundish and brownish (Fig. 2). Head and thorax of female black; clypeus, ventral spot on scape, mandibles, collar, all corners of pronotum, large anterolateral spots on mesoscutum, all tegulae and spots below tegulae, fore and middle coxae and all trochanters (hind trochanters with brownish dorsal spot at base) yellow. Hind coxae red-dish or reddish brown; femora and tibiae reddish, hind tibiae yellowish white with weak infuscation at base and at apical 0.3. Tergite 1 black; beyond tergite 1 abdomen varies from blackish brown on tergite 2 to pale brownish at apex, the last tergites with wide whit-ish-yellow apical band; sternites and epipleura whit-ish-yellow (Fig. 45). In male lateral parts of frons, face entirely, prothorax almost completely and mesothorax ventrally yellow.

Material. Great Britain: 2 ♀ (NHM). Sweden: 1 ♀, 3 ♂ (syntypes) and 5 ♀ (Mus. Lund). Germany: 1 ♀, Eschwege, 4.06.1960, “2942” (Hinz) (Mus. Munich). Poland: 1 ♀, Mielnik, Siemiatycze, 27.6.2005 (J. Sawoniewicz) (ZIN).

Saotis compressiventris (Strobl, 1902)]

[This species is considered here as supposedly jun-ior synonym of Saotis nigriventris Thomson, 1894].

8. Saotis dorsata (Thomson, 1888) (Fig. 7)

Thomson, 1888 : 1264, ♀ [Mesoleius (Saotus); holotypum examinavi, Sweden: “Pål” (= Skåne, Pålsjö) (Mus. Lund)], 1894 : 2019 (Mesoleius); Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Schmiedeknecht, 1914 : 2819 (key, description); Fitton, 1982 : 56 (type); Jussila, 1991 : 39 (fig., Finland); Kasparyan, Shaw, 2003 : 352, 353 (fig., key); Kasparyan, Khalaim, 2007 : 548, 550 (key, fig.).

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Diagnosis. Saotis dorsata differs from the other congeners in the following characters: ovipositor sheath rather narrow (Fig. 7) and tergites 2 and 3 red-dish in considerable part. Thorax entirely black; sub-tegular ridge with yellow stripe. Scutellum sometimes brownish at apex. Antenna with 22–26 flagellomeres; i. fl 0.9; i. gen 0.3; i. fem 3.9–4.2; i. terg 2.0–2.2; 3rd segment of hind tarsus 1.1–1.2 times as long as 5th segment. Sternites 1–4 reddish yellow, sternites 5 and 6 rufous. Legs light red, hind tibia whitish, slightly reddish at base and brownish at apical 0.27.—Holotype (♀): fore wing 4.2 mm, flagellum 26-segmented (22–24 segments in other specimens); epipleura of tergites 2 and 3, and occasionally also those of tergites 4 and 5 separated by crease; oviposi-tor sheath brownish.

Material. Sweden: 1 ♀, 3 ♂ (syntypes). ?Poland: 1 ♀, “VI.1960 (Glovacki)”(Mus. Munich). Ukraine: 1 ♀, Crimea, Karadag, 3.VI.1990 (Kasparyan). Russia. Leningrad Province: 1 ♀, Kobralovo, 42 km S Saint-Petersburg, 17.VII.1984 (Kasparyan). Yamalo-Nenetzkij nation. okrug: 1 ♀, Ob’ River, 8 km below Labytnangi, 12.VII.1994 (Kasparyan); 11 ♀, Taz River, 40–100 km upward Ratta, 21.VII–7.VIII.1992 (Kasparyan) (ZIN) .

9. Saotis granulator Kasparyan & Kopelke, sp. n. (Figs. 37–39)

Saotis granulator may be distinguished from the other congeners by an almost evenly granulate thorax, very short malar space, obliterated areas of the propo-deum, and predominantly rufous elongate abdomen of the female (Fig. 39); the body of the female is black, with the legs red beyond the trochanters.

Description. Holotype (female). Length of fore wing 5 mm, body length 6.5 mm. Antenna with 23 flagellomeres, i. fl 1.0. Head almost not narrowed be-yond eyes (Fig. 37), temple at middle 1.6 times as long as eye in profile. Clypeus convex at basal 0.6, with lower edge sharp and broadly concave. i. gen 0.25. Thorax matte and almost completely evenly and dis-tinctly granulate. Mesopleuron in lower half with moderately dense hairs; speculum rather small, partly with fine granulation. Metapleuron densely and coarsely granulate. Propodeum (Fig. 38) with basal area and areola partly obliterated, apical area weakly bordered; pleural carinae absent. i. fem 4.7, i. tars 5 : 2.5 : 2.0 : 1.2 : 1.7. i. nrv about 0.33. Tergites dis-tinctly elongate, i. terg 1.6, tergites 2–4 about 1.5 as

long as their width in base; tergites 7 and 8 projecting beyond hypopygium along their entire length (Fig. 39). All tergites completely granulate, tergites 1 and 2 matte, subsequent tergites smoother and shining. Epipleura of tergites 2 and 3 narrow and separated by crease. Tergites 3–5 with almost straight (not dis-tinctly emarginate) hind margin.

Body black. Antennae brownish, with blackish scape and pedicel. Clypeus and palpi reddish yellow; mandible, hind corners of pronotum, and fore tegulae yellow; hind tegulae reddish yellow. Coxae black, fore coxa yellowish at apex. Legs beyond coxae predomi-nantly reddish rufous; trochanters yellow, hind tibia light rufous in middle 0.35, its base and apex reddish rufous, spurs pale rufous; hind tarsus pale brownish with extreme base of segments 1–4 paler. Pterostigma rather pale, brownish yellow with brown margins. Abdominal tergites black with brownish tinge, not bordered with pale on hind margins. Sternites brown, the basal sternites yellowish posteriorly. Ovipositor sheath brownish black.

Variability in females. Fore wing 3.5–5 mm; fla-gellum 21–24-segmented, 0.84–0.92 as long as fore wing; i. fl 0.8–1.0; i. gen 0.23–0.27; pleural carinae of propodeum sometimes complete. Body always covered with dense distinct granulation, matte. In European population mesoscutum, scutellum, hind coxae almost always black; hind tibia light rufous, usually slightly grayish brown on dorsal side and at apex.

Male. Similar to female but slender, genae shorter (i. gen about 0.2); flagellar segments 1 and 2 rather slender, respectively 6 and 4 times as long as their width at the middle, segment 1 without discernible rhinaria. Face and clypeus entirely, base of antenna ventrally, malar space, small spot immediately beyond the mandible; thorax completely black or with yellow markings on collar, anterolateral spots on mesoscutum, sometimes on subtegular ridge, on prepectus and in lower part of mesopleuron. Fore and middle coxae, apical part of hind coxa and all trochanters yellow; hind coxae in basal 0.6–0.8 black or rarely brownish-red. Hind edge of tergite 2, extreme base of tergite 3 and sometimes median elongate stripe or spots on its dorsum reddish yellow. Sternites 1–4 yellowish white.

Geographic variability. Females from Canada dif-fer in having fore wing 4.3–5.3 mm long, flagellum 20–22-segmented, about 0.95 times as long as fore wing; i. gen 0.15–0.2, granulation of mesopleuron and tergites finer, propodeum with areola more distinct;



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i. fem 5.0, i. tars 5.2 : 2.7 : 2.1 : 1.3 : 2.0. i. nrv 0.2–0.33, i. terg 1.8; tergites 3–5 with distinctly emar-ginate hind margin; coxae reddish, trochantelli yellow, hind tibia dull whitish at extreme base (0.1) and at middle (0.35), infuscate before base (0.2) and at apex (0.35); clypeus reddish brown, both pairs of tegulae yellow; in one female, scutellum reddish yellow dor-sally.

Material. Holotype: Norway, ♀, Norge/Nordland, Polarkreis, M[oi] Rana, 5.8.90 (Kopelke); Zucht: 33/V 1990, Schlüpfdatum 17.5.91, ex Pontania glabrifrons Benson (Kopelke) (Mus. Senckenberg). Paratypes. Norway, Nordland: same locality and host as in holotype, galls are collected 5.8.1990 : ex Pontania glabrifrons, 11 ♀, 3 ♂, “Zucht 33/O, P, Q, R, S, T, V/1990” and 2 ♀, “Zucht 34/F1/1990“, Schlüpfdatum 22.4–17.5.1991; ex Pontania nivalis Vikb. (Kopelke): Zucht: 7 ♀, 34/D1, G1, H1/1990; Schlüpfdatum 26.4–2.5.1991; ex Pontania varia Kopelke (Kopelke): Zucht: 5 ♀, 10 ♂, 35/C, G, I, K/1990; Schlüpfdatum 24.4–6.5.1991.

Norway / Troms, Ramfjorden, 12.8.1985 (11 ♀, 10 ♂) ex Pontania norvegiea / Salix borealis [labeled Pontania sp. 20 / Salix borealis: 1 ♀, 1 ♂, 53/K1, Schlüpfdatum 22.5.1986 (2927); 1 ♀, 53/N1, Schlüpf-datum 23.5.1986 (2927); 3 ♀, 53/V1, Schlüpfdatum 25.5.1986 (2927); 2 ♀, 53/V1, Schlüpfdatum 25.5.1986 (2929); 1 ♀, 57/T1, Schlüpfdatum 22.5.1986 (“Saotis sp. 5”); 1 ♂, 54/C, Schlüpfdatum 15.5.1986 (2932); 1 ♂, 54/D, Schlüpfdatum 16.5.1986 (2932) (epimeron yellow); 2 ♂, 54/E, Schlüpfdatum 20.5.1986 (2927); 1 ♀, 54/G, Schlüpfdatum 21.5.1986 (“Saotis sp. 4,” 2929); 1 ♀, Z54/G, Schlüpfdatum 21.5.86 (2927); 1 ♀, 2 ♂, 54/H, Schlüpfdatum 20.5.1986 (2927); 2 ♂, 53/I1, Schlüpfdatum 20.5.1986 (“Saotis sp. 2,” 2927); 1 ♂, 54/J1, Schlüpfdatum 20.5.1986 (2927). Norway / Finnmark, 1 ♀, Nr. 42/I, S.-Varanger, Vaggatem, 23.7.1985 (Kopelke), ex Pontania nivalis / Salix glauca, Schlüpfdatum 20.5.1986 (2927). Norway / Süd-Trondelag, 1 ♂, 54, F, Oppdal, ex Pontania norvegica / Salix borealis, RK4, 19.08.1988 (Kopelke), Schlüpfdatum 18.5.1989. Norway / Buskerund: 8 ♀, 9 ♂, 55/S, O, W, K/ 1990, Geilo, 15.8.90, ex Pontania varia Kopelke / Salix myrsinifolia (Kopelke), Schlüpfdatum 26.4–6.5.1991. Norway / Nordland, 1 ♀, 32 P1/1997, Fauske, 16.8.1997 (Kopelke), ex Pontania hastatae / Salix hastata, Schlüpfdatum 14.5.1998.

Sweden: 1 ♀, Kvikkjokk, “2927,” 11.7.1964; 1 ♀, Abisko, “2927,” 16.7.1964; 1 ♀, Kiruna, “2927,”

2.7.1968 (Mus. Munich, coll. R. Hinz). Germany?: 1 ♀, “Jllasberg K 29.5.50,” “M[esoleius] juvenilis Hlgr” (Mus. Hamburg). Finland: 1 ♀, Suomi, PS Nilsia 70/55, 16.6.1988 (R. Jussila), "7769” (Coll. R. Jussila, Turku). Russia (ZIN). Leningrad Province: 1 ♀, Kobralovo–Semrino, 40–42 km S St. Petersburg, 21.6.1980 (Kasparyan); 1 ♀, Osel’ki, N St. Petersburg, ex Pontania pedunculi Htg. / Salix aurita, 11.05. 1995 (A. Zinoviev); Yakutia: 1 ♀, Zhigansk, ex “3147” 07.1990 (A. Zinoviev). Canada, British Columbia: 3 ♀, Stone Mt. Pk., 3800 ± ft, 19, 22 and 24.VII.1973 (H. et M. Townes) (AEI).

Hosts. Pontania glabrifrons Benson,1960, P. hasta-tae Vikberg, 1970, P. nivalis Vikberg, 1970, P. norve-gica Kopelke, 1991, P. varia Kopelke, 1991, Pontania [pedunculi (Hartig, 1837)] = bella (Zaddach, 1876).

10. Saotis heteropus (Thomson, 1883) (Figs. 52–55)

Thomson, 1883 : 934, ♀ [Mesoleius (Saotus); lectotypum examinavi, “Lpl” (Lappland) (Mus. Lund)]; Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Schmiedeknecht, 1914 : 2809, 2816 (key, description); Roman, 1931 : 4–54; Meyer, 1936 : 295 (Russia); Jussila, 1965 : 71 (Finland); 1984 : 91; Fitton, 1982 : 57 (lectotype); Kasparyan, Shaw, 2003 : 354 (fig., key); Kasparyan, Khalaim 2007 : 549, 550 (key, fig.).

Diagnosis. Saotis heteropus is similar S. boreator sp. n. and S. nigriscuta Thoms. in wide black oviposi-tor sheath and slender basal flagellomeres; it can be distinguished from both species by a black mesoscu-tum, coxae, and trochanters, and by a blackish hind femora with light brownish apices.—Lectotype. Fore wing 4.8 mm; flagella broken; i. fl 1.12; i. gen 0.35; clypeus before lower margin with a median swelling; Mesopleuron and metapleuron granulate; Mesopleuron on upper 0.4 finely striate; nervulus weakly postfurcal; i. nrv 0.3; i. fem 5.0; in lectotype hind tarsi broken (In other specimens from Lappland segment 3 1.2 times as long as segment 5); i. terg 2.15; lateral margins of tergite beyond spiracles subparallel (Fig. 55). Antenna with flagellum pale rufous, flagellomere 1, scape and pedicel blackish brown. Body black; clypeus reddish yellow, mandibles brownish, hind corners of pronotum slightly reddish. All coxae and trochanters black, hind femora blackish with brownish apex; hind tibia pale rufous, at extreme base brownish with small dorsal brownish marking before base and entirely reddish brown at apex (0.25). Tergites 3–8 with whitish hind

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margin at the middle (Fig. 12), sternites yellow with basal infuscation.

Material. Sweden, Lappland: 1 ♀ (lectotype) and 1 ♂ (Mus. Lund); 2 ♀, Kiruna, “2929,” 18 and 19.07.1969 (R. Hinz) (Mus. Munich); 1 ♀, Abisco, 13.07.1954 (J. E. and R. B. Benson) (NHM). Russia (ZIN): 1 ♀, Polar Ural, 50 km NW Labytnangi, 800 m, Salix, 18.07.1994 (Kasparyan); 2 ♀, Yakutia, Tit-Ary, delta of Lena river, ex Phyllocolpa sp. on Salix sp., “3197,” 24 VII 1990, emerged 16.05.1991 (A. Zino-viev).

11. Saotis hoeli Roman, 1933 (Figs. 11, 24) Roman, 1933 : 9 [holotype: ♀, East Greenland

(Zoologisk Museum, Sars gt. 1, Oslo, Norway]; 1934 : 609; Carlson, 1979 : 602 (Nearctic); Kasparyan, Shaw, 2003 : 354 (figs., key).

Diagnosis. Saotis hoeli differs from S. albionis and from the Nearctic S. clypealis Ashm. in the black body and legs (except for tibiae and tarsi). A circumpolar species.

Material. Greenland (Denmark): 1 ♀ (hoeli Roman det.), E. Greenland Jamesonland, VIII 1933 (D. Lack) (NHM); “Nedre Midsommer Sö Green-land,” 8 and 22.07.1966 (Mus. Ottawa). Canada: Hazen Camp, 81°49’ N, 71°18’ W, 11.07.1962 (4 ♂), 15 and 24.07.1962 (2 ♀) (R. B. Madge) (Mus. Ottawa). Russia: 1 ♀, Novaya Zemlya, Matochkin Shar, 31.07.1925 (Vakulenko); 1 ♀, Kamchatka, volcano Avacha, 1000 м, tundra, 26.07.1985 (Kasparyan) (ZIN).

12. Saotis longiventris (Thomson, 1888) (Figs. 15, 50, 51)

Thomson, 1888 : 1263, ♀ [Mesoleius (Saotus); holotypum examinavi: ♀, Sweden, “Ört 28/V”= Skåne, Örtofta (Mus. Lund)]; 1894 : 2019 (Saotus); Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Schmiedeknecht, 1914 : 2810, 2819 (key, description); Fitton, 1982 : 58 (type); Koponen, Jussila, Vikberg, 2000 : 73; Kasparyan, Shaw, 2003 : 352, 353 (key, fig.); Kasparyan, Khalaim, 2007 : 548, 550 (fig., key).

Diagnosis: see couplet 13 of the key.

Material. Holotype.

13. Saotis mirabilis Schmiedeknecht, 1914 Schmiedeknecht, 1914 : 2808, 2810, ♀ [key, de-

scription; typum examinavi: ♀, Germany (Mus. Ber-

lin)]; Teunissen, 1948 : 26; 1953 : 30 (♂); Kasparyan, Shaw, 2003 : 351 (key; = seleuciformis).—seleuciformis Kolarov, 1987 : 71, ♀ [Iskarus; holotype ♀, Bulgaria (coll. J. Kolarov)].

Diagnosis. Saotis mirabilis can easily be recognized by a combination of an exceptionally long abdomen (2–3 times as long as the head and thorax combined), an almost entirely red mesothorax, and large size (the body length is 9–15 mm). Tergites 3–8 form a deep angulate median notch on the hind margin; tergite 8 is depressed, the ovipositor sheath is smaller than that in the other species.

Material. Germany: 2 ♀, Blankenburg in Thüringen (including type) (Mus. Berlin). Swit-zerland: 2 ♀, Bern, Kirschenfeld (Steck) (Mus. Ber-lin). Slovakia: 5 ♀ 3 ♂, reserve Šur, 12 km NO Brati-slava, 30.06.1991 (Kasparyan) (ZIN). Russia: 2 ♀, Sosnovo, 75 km N St. Petersburg, Malaise trap, 1–7.08.2008 (Kasparyan) (ZIN).

14. Saotis morleyi Fitton, 1976 (Figs. 6, 16, 33)

Fitton, 1976 : 341, nom. nov. pro Homocidus emarginatus Morley, 1911, praeocc., non Thomson, 1883 [holotypum examinavi: ♀, Scotland, Kincar-dineshire, Banchory (NHM)].—Saotis albiventris Kasparyan, 2007 (in: Kasparyan & Khalaim, 2007 : 548–549), ♀, ♂ [holotypum: ♀, England, Oxfordshire, Dry Sandford Pit, 30.VI–19.VII.1990 (K. Porter) (NMS), key, description, figs., distribution—Great Britain, Sweden, Russia], syn. n.

Diagnosis. The female of this species can easily be recognized by the very pale and strongly compressed apex of the abdomen (Figs. 6, 33); tergite 8 is convex upwards in profile, the ovipositor sheath is light yel-low; the characters typical of the tricolor species group (short malar space and coloration of the mesos-cutum, scutellum, and hind tibia (Fig. 16)) are also typical of this species.

Material. Great Britain. Scotland, ♀ (holotype of Homocidus emarginatus Morley) (NHM); 1 ♀, Scotland (NMS); England, 2 ♀ (holotype and paratype of Saotis albiventris Kasparyan) (NMS). Switzerland: 1 ♀, 19 R1/1991, CH/Wallis, Grimselpass, 2000 m, ex Phyllocolpa spirhelvetica (labeled: Phyllocolpa sp.) / Salix helvetica, 4.09.1991, Schlüpfdatum 14.05.1992; 3 ♀ 1 ♂, SZ, 31/1998, same locality, 29.08.1998, ex Phyllocolpa plicaglauca (labeled: Phyllocolpa sp.) / Salix glaucosericea, einfache Blattrolle,



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Schlüpfdatum 27.04.1999 (2 ♀ 1 ♂) and 29.04.1999 (1 ♀). Germany. 1 ♀ 1 ♂, SZ, 02a, Schleswig-Holstein, Kiel, Dietrichsdorf, Phyllocolpa leucapsis / Salix cinerea, Blattrolle verdrillt, 21.06.2003, Schlüpfdatum 10.02.2004 (♀) and 15.03.2004 (♂); 1 ♀, SZ 7, Hessen, Wüstensachsen, NWR Stirn- berg, Phyllocolpa leucapsis / Salix cinerea, Blattrolle verdrillt, 26.06.2003, Schlüpfdatum 18.02.2004; 1 ♀, SZ 15, Baden-Würtemberg, Radolfzell, Mindelsee, Phyllocolpa leucapsis / Salix cinerea, Blattrolle verdrillt, 21.06.2003, Schlüpfdatum 10.02.2004; 1 ♂, SZ 77, Mecklenburg-Vorpommern, Rügen, Stedar, Phyllocolpa scotaspis / Salix viminalis, Blattrolle einfach, 17.07.2005, Schlüpfdatum 24.05.2006; 1 ♀, Hessen/Vogelsberg, 1984 (Kopelke), Phyllocolpa leucapsis (labeled: Phyllocolpa sp.2) / Salix cinerea / Doppelrollgallen. Norway: 1 ♀, SZ, 55, Süd-Trondelag, Berkak, Phyllocolpa ischnocera / Salix myrsinifolia, Blattrfalte einfach, 21.08.2004, Schlüpfdatum 29.04.2005; N/Finnmark, 3 ♀ 2 ♂, SZ 27, S. Varanger, Vaggatem, 07.08.2001, ex Phyl-locolpa sp. / Salix caprea / verdrillte Blattrolle, Schlüpfdatum 15.05 and 17.05 2002. Sweden (in coll. Thomson): 2 ♀ 2 ♂, Pålsjö (among paralectotypi of S. liopleuris), 1 ♀, Pålsjö and 1 ♂, Scorpe (among specimens of S. tricolor) (Mus. Lund). Finland: 1 ♀, Sakkola (Hellén) (Mus. Helsinki).

Hosts. Phyllocolpa ischnocera (Thomson, 1862), Phyllocolpa leucapsis (Tischbein, 1846) Phyllocolpa plicaglauca Kopelke, 2007, Phyllocolpa scotaspis (Förster, 1854) Phyllocolpa spirhelvetica Kopelke, 2007.

15. Saotis nigriscuta (Thomson, 1883) (Figs. 5, 34)

Thomson, 1888 : 1264, ♀ [Mesoleius (Saotus); holotypum examinavi, Sweden: “Pål” (= Skåne, Pålsjö) (Mus. Lund)]; 1894 : 2018 (Saotus); Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Schmiedeknecht, 1914 : 2813 (key, description); Meyer, 1936 : 294 (Russia); Teunissen, 1948 : 26; 1953 : 30 (Netherlands); Hellén, 1953 : 9; Fitton, 1982 : 58 (holotype); Shaw, Kasparyan, 2003 : 26 (England); Kasparyan, Shaw, 2003 : 352–355 (fig., key; England); Kasparyan, Khalaim, 2007 : 549, 550 (key, fig.).

Diagnosis. Saotis nigriscuta can be distinguished from the other species of the “nigriscuta” group by a greater number of the flagellomeres (usually 25–27),

more slender basal flagellomeres (i. fl 1.2–1.32), and by the presence of the median swelling above the lower margin of the clypeus (Fig. 34); the head is usu-ally slightly narrowed backwards; the coxae and tro-chanters are usually red.

Material. Great Britain, England: 2 ♀ (NMS) and 1 ♀ (NHM). Sweden: ♀, holotype (Mus. Lund). Nor-way: 1 ♀, SZ 57/2001, N/Nordland, Polarkreis I (Kopelke) 15.08.2001; ex Phyllocolpa acutiserra / S. lapponum / verdrilltle Blattrolle, Nr Schlüpfdatum 27.5.2002. 1 ♀, Norge, Dovre, Vålåsjö, 14.07.1953 (W. Hellén) (Mus. Helsinki). 1 ♀, Norway/Finnmark, S. Varanger (ZIN). Russia: 5 ♀, Murmansk and Len-ingrad Provinces, Polar Ural, Transbaikalia (ZIN). Mongolia: 1 ♀, Central aimak (ZIN).

Hosts: Phyllocolpa acutiserra (Lindquist, 1948).

16. Saotis nigriventris Thomson, 1894

Thomson, 1894 : 2019, ♀ [Saotus; holotypum examinavi, Germany: “Mdrk 20.VI.83” (Mus. Lund)]; Dalla Torre, 1902 : 276 (Saotis; bibliogr.); Schmie-deknecht, 1914 : 2816 (key, description); Teunissen, 1948 : 26 (Netherlands); Hellén, 1953 : 9 (Finland); Fitton, 1982 : 80 (holotype); Jussila, 1991 : 39; Silfverberg, 1996 :47. Horstmann, 2001 : 69–103 (Germany); Kasparyan, Shaw, 2003 : 352–353 (fig., key); Kasparyan, Khalaim, 2007 : 548, 550 (key, fig., Russia).—compressiventris Strobl, 1903 : 20, ♀ ♂ [Mesoleius; lectotype: ♀, Austria, nr Admont, “Natter-riegel 8/6 Styria Strobl” (coll. Strobl, Admont)], syn. n.; Schmiedeknecht, 1914 : 2809, 2817 (Saotis; key, description); Heinrich, 1953 : 209; Horstmann, 1999 : 51 (lectotype design., labeled by Hinz).

Diagnosis. The species is easily recognizable by two autapomorphies: short and very dense pubescence of the abdomen and modification of tergite 8 (with a tooth on its hind margin and with the anal vertical slot above the tooth, Fig. 8). The antenna consists of 22–24 flagellomeres; i. fl = 0.9; i. gen = 0.2–0.25; the areas of the propodeum are obliterated; the thorax is entirely black; the ovipositor sheath vary from pale rufous to reddish brown, with a deep dorsal notch at the base (Fig. 8).

Material. Germany (holotype). Sweden. 10 spe-cimens, majority from Stockholm (Mus. Stockholm); 3 ♀, (“compressiventris Strobl” Perkins det.), “Swe-den SK,” Höör distr. and Ring sjö 31.V–19.VI.1938 (D. M. S. P. & J. F. Perkins) (NHM). Switzerland:

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1 ♀, Kandersteg / CH, 15.7.1976, “2938,” (R. Hinz) (Mus. Munich). Finland.: 5 ♀, Suomi V Sauvo, Ka-runa, 1–27.6.1998 (3 ♀) and 8–23.6.2000 (R. Jussila) Mus. Turku). Russia: 1 ♀, Primorskii Territory, Khanikheza river, near Roshchino, 2.6.1966 (Kas-paryan).

Remarks. Mesoleius compressiventris Strobl, 1902 is considered here as junior synonym of S. nigriventris Thomson, 1894. This synonymy is based on the results of examination of Thomson’s type and the conspecific specimens from the collections of Munich and Lon-don, which were determined by R. Hinz and J. Perkins as S. compressiventris and compared with the type of Strobl. In the original description of S. comres-siventris, Strobl noted that the tergites with “ziemlich dichte und deutliche Punktierung;” this character per-haps fits such an autapomorphy of S. nigriventris as dense short hairs covering the abdominal tergites. Some peculiarities of the coloration, mentioned in the description (the thorax and abdomen are completely black with the last abdominal segments being “mehr rotbraun” and the ovipositor sheath “braunrote”), are also similar to those in S. nigriventris.

17. Saotis pygydiator Kasparyan & Kopelke, 2009 (Figs. 29, 30, 40, 42)

Kasparyan & Kopelke, 2009 : 120.

Saotis pygidiator structurally closely resembles S. nigriscuta Thoms., especially, in having rather broad ovipositor sheath, slender basal flagellar seg-ments, and also in coloration; it differs from S. nigris-cuta and other congeners in the broad and swollen temples (Fig. 6) and in the erected, usually very dense and short pubescence of the female’s hypopygium (Fig. 9); the nervulus is usually interstitial, or rarely weakly postfurcal; i. nrv 0.1–0.25; the pterostigma is often pale.

Description. Fore wing 4–5 mm long. Antenna with 23–28 flagellar segments, flagellum more or less as long as fore wing (±0.07). i. fl 1.25 ± 0.1; i. gen 0.4 ± 0.1; i. terg 5.0 ± 0.5; greater spurs of hind tibia usually shorter than half length of hind basitarsus. Clypeus convex at basal 0.3–0.5, with long setae be-low basal convexity, its lower margin sharp and broadly concave. Head finely granulate; granulation very uniform on frons, smoothened on temples; face with fine conspicuous punctures. Nervulus interstitial or weakly postfurcal, nervellus intercepted about at lower 0.2 ± 0.08. Dorsal areas of propodeum present,

areola narrow. First abdominal tergite finely granulate, with its apical margin polished; i terg 1.7–2.0; Second tergite with very fine and smoothened granulation and with short and rather sparse hairs. Tergite 8 short and usually is strongly drawn in abdomen. Hypopygium and sometimes preceding sternites are covered with very dense and short erected hairs (Fig. 42); hypopy-gium truncated on hind margin and usually weakly concave at the middle. Ovipositor sheath moderately wide, about 0.8 as broad as lateral width of hind fe-mur.

Body black; antenna from brownish to yellowish ru-fous; mesoscutum usually with anterolateral yellow spots, scutellum black; posterior margin of tergites (3)4–7 with whitish band dorsally. Legs (usually ex-cept for coxae and trochanters) reddish, hind tibia from uniformly pale rufous to dull whitish with dark-ened dorsal side and apex. Pterostigma usually pale yellowish brown (dark brown in S. pygidiator ivan).

There are four geographical forms (Kasparyan, 2009); the differences between these forms and their distribution are given in the key (couplets 14a–14c). Here, only a short description of the European subspe-cies is given.

Saotis pygidiator pygidiator Kasparyan & Kopelke, 2009

Kasparyan et Kopelke, 2009 : 122, ♀ ♂ [Holotype: ♀, Austria, Steiermark , ex Pontania varia Kopelke (SMF)].

Description. Female (holotype). Fore wing 4.5 mm, flagellum 4.7 mm long. Antenna with 24 flagellar segments; i. fl 1.25. Head not narrowed, temples al-most parallel beyond eyes (Fig. 30); i. gen about 0.4. Nervulus interstitial; i. nrv 0.15; i. fem 5.0, i. tars 4.5 : 2.3 : 1.8 : 1.1 : 1.7; longest tibial spur about 0.46 as long as hind basitarsus. i terg 2.0. Hypopygium with dense erected hairs [in specimens from Switzerland, hypopygium weakly pubescent (var. helvetica)].

Antenna brownish, flagellum at basal 0.2 paler ven-trally. Head and thorax black with clypeus, mandible, palpi, hind corner of pronotum, a pair of small anter-olateral spots on mesoscutum, and tegulae yellow; apex of epimeron slightly reddish. Fore and middle legs pale reddish; coxae blackish at base and dorsally, and reddish ventrally; all trochanters I blackish, tro-chanters II yellowish; middle tarsi brownish. Hind coxa black, ventrally at apex reddish brown; hind fe-mur red; hind tibia dirty whitish ventrally, light



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brownish dorsally, completely dark brown at apical 0.3 and at basal 0.15; hind spurs whitish; hind tarsus light brownish with extreme base of segments 1–4 pale. Pterostigma yellowish with brown margins. Ab-dominal tergites blackish; tergites 4–7 dorsally with narrow yellow band on hind margin (Fig. 10). Ster-nites yellow on longitudinal fold; sternite 1 darkened at apical 0.4; hypopygium entirely yellow.

Male similar to female, but slender; fore wing 3.5–4 mm; flagellum about 23–24 segments; i. fl 1.14–125; cheeks shorter, i. gen about 0.35; coloration paler than in female: face, cheeks from bottom of eye to lower corner of mandible, ventral spot on scape and pedicel, fore and middle coxae almost entirely, all trochanters (except for hind trochanter I) light yellow. Hind femur darker, from reddish brown to dark brown.

Material. Austria, ♀ (holotype), Steiermark, Don-nersbach, Planneralm B [II] 10.8.89 (Kopelke), ex Pontania varia Kopelke (Kopelke det.) [on Salix myrsinifolia], [type of galls: RK4], Zucht : Kopelke Nr 15/Q1 1989 [number of sample], Schlüpfdatum 21.5.1990 (SMF). Paratypes. Austria: Steiermark , same data as in holotype [1 ♀ 6 ♂] but: 1 ♀—15/U1 1989, Schlüpfdatum 27.5.90; 2 ♂—15/E1 1989, Schlüpfdatum 17.5.90; 2 ♂—15/R1 1989, Schlüpfda-tum 14.5.90; 2 ♂—15/S1 1989, Schlüpfdatum 23.5.90; Tirol, 1 ♂, Zillertal, Hintertux, Weitental, 1700 m, 13.8[1989] (Kopelke leg.), ex Pontania varia (Ko-pelke det.), [RK4], Zucht: Kopelke Nr 25/S1 1989, Schlüpfdatum 28.5.90; 1 ♀ 1 ♂, Tirol, Ötztal, Öber-gurgl, 1800 m, 16.8 89 (Kopelke), ex Pontania varia (Kopelke det) / Salix myrsinifolia, [RK4], Zucht: Ko-pelke Nr 37/M and 37P/ 1989, Schlüpfdatum 27.9.89 and 28.5.90; 1 ♀, 1 ♂, Salzburg, Obertauern, 26.8.96 (Kopelke), ex Pontania foetidae / Salix waldsteiniana, (Kopelke det.), [RK2], Zucht: Kopelke Nr 2Q1 and 2R1 1996, Schlüpfdatum ♀—15.5.97, ♂—6.5.97. Germany? [in: R. Hinz collection, Mus. Munich]: 1 ♀, without locality, “P 104 1970,” “2310,” ex Pon-tania sp. Switzerland: Helvetia 3 ♀, 37/D/1984, 37/E/1984 and 54/S/1984, Wallis, Sass Almagell, 1900 m, 14.8.1984 and 18.8.1984, ex Pontania brevis-erratae (labeled: Pontania sp. 13) / Salix breviserrata (Kopelke det.), Schlüpfdatum 18.5 1985 and 22.5.1985 and ex P. hastatae; Graubünden, 1 ♀, 45, A1-G 1, C1/ 2000, Juppa, Pontania varia, Salix myrsinifolia, RK4, 15.08.2000 (Kopelke), Schlüpfda-tum 12.03.2001. All paratypes from Austria (except for 1 ♀ and 1 ♂ in ZIN) and from Switzerland on deposition in SMF.

Distribution. Central Europe. Host. Pontania breviserratae Kopelke 1989

(= ”Pontania sp. 13”) on Salix breviserrata with pea-shaped galls transected horizontally by the leaf blade; Pontania foetidae Kopelke 1989 on Salix wald-steiniana; Pontania hastatae Vikberg, 1970; Pontania varia Kopelke 1991 on Salix myrsinifolia.

18. Saotis renovata (Morley, 1911) (Figs. 13, 17, 18, 21, 22)

Morley, 1911 : 169, ♂ [Mesoleius; lectotypum ex-aminavi: ♂, England, Worcester (NHM)]. Schmiedek-necht, 1914 : 2808, 2812 (Saotis; key, description).—brevispina: Brifgman, 1887 (ex Pontania purpureae / S. purpureae).Saotis morleyi: Kasparyan, Khalaim, 2007 : 550, misidentification (key, distribution—Britain, Sweden, Russia).

Diagnosis. Saotis renovata differs from the other species of the tricolor group (except for S. morleyi) in the following characters: tergite 1 strongly elongate (Figs. 18, 21), tergite 8 less convex dorsally in profile (compare Figs. 13 and 14), number of flagellomeres of antenna only 21–25, malar space longer (i. gen about 0.4). Saotis renovata like the other species of the tri-color group has the following features: i. fl 0.95–1.05, 1st flagellomere 4–5 times as long as wide at middle; hind femur rather thick (i. fem 3.9–4.4). Mesoscutum (usually) and scutellum with yellow spots; hind tibia with white ring at middle part, fuscous (often blackish with reddish) at base and at apex (Fig. 17). The spe-cies is also similar to S. albionis in many characters, but differs in slender tergite 1 (i. terg 2.1–2.9), and in the shape of tergite 8 with pygostyli (compare Figs. 13 and 32).

Material. Great Britain. 3 ♀, England, including a specimen labeled as “cotype” of Homocidus emarginatus Morley: “Leister” (T. A. M[arshall] (NHM). Norway. 1 ♀, SZ 48/2001, Nordland, Lofot, Austvagoya, ex Phyllocolpa anomalpotera (labeled: Phyllocolpa sp.) / Salix caprea / eingerolle Blatrolle (Kopelke det.), 13.08.2001, Schlüpfdatum 15.05.2002. [Perhaps to this species belong 1 ♂, SZ 25/2001, Norway/Finnmark, S. Varanger, 6.08.2001 (Kopelke), ex Phyllocolpa nudipectus (labeled: Phyllocolpa sp.) / Salix phylicifolia, Schlüpfdatum 27.05.2002; and 1 ♂, SZ 27/2001 from same locality, but galls were collected 7.08.2001, Phyllocolpa sp. / Salix phy-licifolia, Schlüpfdatum 15.05.2002]. Germany. 1 ♀, Einbeck, “2088“, 14.05.1974, R. Hinz (Mus. Munich). Sweden. 1 ♀, Röstanga, 5.07.1938, DMSP&J. F. P.).

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Poland. 1 ♀, Byalystok, 9.07.2004, J. Sawoniewicz (ZIN). Russia. 1 ♀, Primorskii Territory, Vladivostok, Sanatornaya, ex Phyllocolpa sp. / Salix udensis, 27.06.1979, parasitoid emerged 28.05.1980 (A. Zi-noviev) (ZIN).

Hosts. Phyllocolpa anomalpotera (Förster, 1854), [?2 ♂ ex Phyllocolpa nudipectus Vikberg], Phyl-locolpa sp. / Salix udensis. Type (as noted by Morley) was received from a series reared from Phyllocolpa purpurea (Cameron, 1884): “bred by Fletcher from a new leaf-rolling sawflies, Nematus purpurea Cam., discovered by him in June, 1886, on Salix purpurea at Worcester” Morley, 1911 : 169)

19. Saotis subarctor Kasparyan & Kopelke, sp. n. (Figs. 36, 44, 45)

Saotis subarctor can easily be distinguished from the other congeners by the epipleura of tergite 4 com-pletely, and those of tergite 5 partly separated by a crease (except for the two other species of the bre-vispina species-group and S. dorsata), by the entirely black thorax, coxae, all trochanters I, and most of ter-gites, and by yellow coloration of sternites and ovi-positor sheath (Figs. 44, 45).

Description. Female. Fore wing 2.7–4.5 mm. An-tenna with 20–23(?24) flagellomeres; i. fl about 1.1. Head weakly roundly narrowed behind eyes, temple in middle part about 1.2 times as long as eye in profile; i. gen 0.4–0.5. Clypeus moderately convex, with nar-row sharp flange along lower margin in lateral 0.4. Mesoscutum centrally with fine superficial granu-lation, with smooth and shiny lateral lobes. Mesopleu-ron with fine granulation; in holotype, almost smooth and shiny below subtegular ridge. Propodeum with distinct dorsal areas, areola long and narrow (more or less similar to that in S. heteropus—Fig. 23); i. ar. ap 0.35; pleural and lateral longitudinal carinae present. Nervulus strongly postfurcal, distad of basal vein about 0.6–0.8 of its length; nervellus intercepted on lower 0.1–0.25; i. fem 4.6–5.0; i. tars (in holotype) 3.5 : 1.9 : 1.4 : 0.8 : 1.4. Tergites 1 and 2 granulate; i. terg varying from 1.7 (in holotype) to 2.1; tergite 1 strongly tapered toward base basally of spiracles, partly scabrous, with deep basal impression sur-rounded with carina; dorsal carinae distinct to spira-cle’s level. Tergite 3 very finely and superficially granulate, shiny; subsequent tergites almost polished. Epipleura of tergites 2–4 completely separated by crease; epipleura of tergite 5 weakly separated. Ter-

gites 5–8 strongly compressed, tergite 8 not enlarged. Ovipositor sheath 0.85 times as wide as lateral width of hind femur and projecting beyond hypopygium by about 0.75 of length of hypopygium (Figs. 44, 45).

Antenna blackish, sometimes with flagellum slightly brownish ventrally, or pale brownish (in Ca-nadian specimens). Head black; clypeus yellowish; mandibles yellowish to reddish brown, often with dark brown spot in basal 0.6. Palpi whitish, with partly infuscate apical segments. Thorax black, with whitish yellow tegulae. Coxae and trochanters I black; all tro-chanters II yellow; legs beyond trochanters brownish rufous; femora often darkened dorsally and posteri-orly, hind femur sometimes entirely blackish. Hind tibia almost uniformly rufous (in holotype) or dirty-rufous with blackish base and apex. Pterostigma pale brownish, usually slightly paler in basal 0.4. Tergites 1–6 black with thin whitish apical band on tergites 2–6 and with whitish epipleura on tergites 2–4; lateral parts of tergites 5–6 and tergites 7–8 almost entirely pale reddish brown or yellowish (Figs. 44, 45). Ster-nites yellow. Ovipositor sheath yellowish.

Male. Fore wing 3.3–4.0 mm; flagellum 20–21-segmented; i. fl 1.15–1.25. Malar space shorter than that in female, i. gen 0.33; i. terg 2.0. Male clearly differing from female in coloration: face, malar space, mandibles, cheeks beyond mandibles, spot on lower and hind corners of pronotum, large anter-olateral spot on mesoscutum, subtegular ridge, apex of epimeron, transverse band on mesosternum along prepectal carina, sometimes lower part of thorax al-most entirely, fore and middle coxae entirely, hind coxa in lower half (reddish to brown dorsally), and trochanters, all yellow. Scutellum and postscutellum black, as those in female. Abdomen black with apical reddish band on tergite 2 and with reddish dorsome-dian stripe (0.3) running from base to apex of tergite 3. Tergites 3–6 with white apical margin. All epipleura and sternites yellow.

Variability. Nearctic specimens on the whole big-ger, but flagellum slightly shorter than fore wing, and number of flagellomeres in large specimens often equal to 20–21.

Material. Holotype: Russia: 1 ♀, Murmansk Province, Kirovsk, Yuksporrjokk stream, 31.07.1974 (Kasparyan) (ZIN). Paratypes. Russia, 1 ♀,Yakutia, Tit-Ary in delta Lena riv., tundra, 22.07.1990 (Kasparyan) (ZIN). Norway: Nordland, 1 ♀, SZ 57/2001, Polarkreis I, 15.08.2001 (Kopelke), ex Phyl-



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locolpa acutiserra / Salix lapponum / verdrillte Blat-trolle (Kopelke det.), Schlüpfdatum 22.05.2002. N.-Trondelag 1 ♂, SZ53/2001, Skogan, 16.08.2001 (Kopelke),ex Phyllocolpa nudipectus / Salix phy-licifolia / Blattrolle schraubig (Kopelke det.), Schlüpf-datum 13.05.2002. Finnmark, 1 ♀, SZ 6/2001, Lakselv Staburrnes, 02.08.2001 (Kopelke), ex Phyllocolpa plicaphylicifolia [labeled Phyllocolpa sp.] / Salix phylicifolia / einfache Blattrolle (Kopelke det.), Schlüpfdatum 22.05.2002; 1 ♂, same data, but SZ 8/2001, ex Phyllocolpa nudipectus / Salix phylicifolia / verdrillte Blattrolle, Schlüpfdatum 6.05.2002; Finnmark, 2 ♂, SZ 32/2001, Ifjordfjellet / Gilojokka, 08.08.2001 (Kopelke), ex Phyllocolpa nudipectus [labeled as Phyllocolpa sp.] / Salix phylicifolia / ver-drillte Blattrolle (Kopelke det.), Schlüpfdatum 08.05.2002; Finnmark, 1 ♀, SZ 25/2001, S. Varanger, Nyrud, 06.08.2001 (Kopelke), ex Phyllocolpa nudipectus [labeled as Phyllocolpa sp.] / Salix phylici-folia / verdrillte Blattrolle (Kopelke det.), Schlüpfda-tum 22.05.2002. S.-Trondelag, 1 ♀, SZ 1/2001, Dovrefjell, 26.07.2001 (Kopelke), ex Phyllocolpa nudipectus [labeled as Phyllocolpa sp.] / Salix phylici-folia / verdrillte Blattrolle (Kopelke det.), Schlüpfda-tum 22.05.2002. Finland: 1 ♂, SZ 37/2001, S/ F Kil-pisjarvi, 10.08.2001, (Kopelke), ex Phyllocolpa nudipectus [labeled as Phyllocolpa sp.] / Salix phylici-folia / verdrillteBlattrolle (Kopelke det.), Schlüpfda-tum 13.05.2002. Finland: 1 ♀ Ivalo, 6546 (Hellén) (Mus. Helsinki). Canada. Quebeck, Payne Bay: 5 ♀—8 and 21.07.1958 (E.E. MacDougall), and 1 ♀—7.07.1958 (W.R.M. Mason) (Mus. Ottava); British Columbia: 3 ♀, Stone Mt. Pk., 3800 ± ft,17 and 20.07.1973 (H.&M. Townes) (AEI).

Host. Phyllocolpa acutiserra (Lindquist, 1948), Ph. nudipectus Vikberg, 1965), Ph. plicaphylicifolia Kopelke, 2007.

20. Saotis tricolor (Thomson, 1883) (Figs. 14, 20)

Thomson, 1883 : 933, ♀ [Mesoleius (Saotus); syntypi examinavi: 3 ♀, Sweden, “L-d” (= Lund) (Mus. Lund)], 1894 : 2019 (Saotus); Dalla Torre, 1902 : 276 (Saotis; literature); Schmiedeknecht, 1914 : 2809, 2815 (key, description); Fitton, 1982 : 59 (syntypes); Kasparyan, Shaw, 2003 : 352, 353 (key, figs.); Kasparyan, Khalaim, 2007 : 548, 549 (= liopleuris; key, figs.).—liopleuris Thomson, 1888 : 1263, ♀ [Mesoleius (Saotus); lectotypum (labeled Hinz 1962) examinavi: ♀, Sweden, “Pål” (= Skåne, Pålsjö) (Mus. Lund)]; 1894 : 2018 (“liosternus,”

lapsus); Dalla Torre, 1902 : 276 (Saotis); Schmie-deknecht, 1914 : 2809, 2815 (key, description); Meyer, 1936 : 295 (Russia); Fitton, 1982 : 57 (designation of lectotype); Kasparyan, Shaw, 2003 : 353 (key).

Diagnosis. Saotis tricolor differs from closely re-lated S. bilineata and S. morleyi in brownish colora-tion of tergites 7 and 8 (including epipleura) (cf. Fig. 14 and Figs. 4 and 6). It can be distinguished from S. renovatus (Fig. 13) by tergite 8 strongly swollen in profile, by short malar space (i. gen about 0.25), by a large number of flagellomeres in antenna (27–30), and less slender tergite 1 (i. terg about 1.6).

Material. Austria. Salzburg, ex Phyllocolpa ischnocera [label: ex Phyllocolpa / nigricans (= myrsinifolia)]: 1 ♀, 4E, Radstädter Tauern, Obertauern 1793m NN, 27.8.1991 (Kopelke), Schlüpfdatum 19.05.1992; 1 ♂, SZ 18, same locality, 27.8.1996, Schlüpfdatum 28.04.1997; 3 ♀ 5 ♂, SZ 19, same locality, 27.08.1996, Schlüpfdatum 30.4.1997 (1 ♀), 2.05.1997 (2 ♀), 5.05.1997 (4 ♂), 6.05. 1997 (1 ♂); 4 ♀ 1 ♂, same locality, SZ 16, 1.08.1998 (Kopelke), Schlüpfdatum 30.04.1999 (1 ♂), 3.05.1999 (3 ♀), 7.05.1999 (1 ♀). Salzburg, 1 ♀, SZ Z, Obertauern, ex Phyllocolpa plicadaphnoides (labeled: ex Phyllocolpa sp. / Salix daphnoides), 14.09.1995, Schlüpfdatum 24.05.1996. Salzburg, 3 ♀, SZ 26, ex Phyllocolpa polita (labeled: ex Phyllocolpa sp. / Salix purpurea). Salzburg, Dachsteigebirge, 23.08.1998, Schlüpfdatum 7.05.1999. Tirol, 1 ♀, SZ 20, Ötztal, Vent Gample, 1800m, NN, 28.08.1996, ex Phyllocolpa ischnocera, Schlüpfdatum 28.04.1997. 1 ♀, Tamischbachturm, Gesäusealpen (H. Franz) (Mus. Helsinki). Germany: 1 ♀, SZ C, Hessen, Wetterau/Kilianstatten, ex Phyllocolpa polita (labeled: ex Phyllocolpa sp. / Salix purpurea) 12.07.95, Schlüpfdatum 29.05.1996; 1 ♀, SZ 68/2005, D./ Meck.-Vorpommern, Rügen, Glowitz, 19.07.2005, ex Phyllocolpa oblita / Salix fragilis [× pentandra] / einfache Blattfalte, Schlüpfdatum: 18.02.2004. Norway: 3 ♀ 1 ♂, SZ 07, ex Phyllocolpa ischnocera (labeled: Phyllocolpa sp. / Salix myrsinifolia): Finnmark, Alta, 2.08.2001, Schlüpfdatum 13.05.2002 (1 ♂), 15–22.05.2002 (3 ♀); 1 ♂, SZ 25, Finnmark, S.-Varanger, Nyrud, ex Phyllocolpa nudipectus (labeled: Phyllocolpa sp. / Salix phylicifolia), 6.08.2001, Schlüpfdatum 27.05.2002; 1 ♀, SZ 48, Nordland, Lofot, Austvagoya, Phyllocolpa anomalpotera (labeled: Phyllocolpa sp. / Salix carea): 13.08.2001, Schlüpfdatum 15.05.2002. Belarus: 1 ♀,

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Pripyat’, 40 km W of Petrikov, 16.08.1979 (Kas-paryan) (ZIN). Russia (ZIN). Leningrad Province: 1 ♀, Krasnitzy, 16.07.1980 (Kasparyan); Transbaikalia Territory: 2 ♀, Ivan-ozero NW Chita, 30.07.1975 (Kasparyan); Primorskii Territory: 1 ♀, oz. Khanka, Novokachalinsk, ex Phyllocolpa sp. on Salix miyabeana (Zinoviev).

Hosts. Phyllocolpa anomalpotera (Förster, 1854), Ph. ischnocera (Thomson, 1862), Ph. plicadaphnoides Kopelke, 2007, Ph. nudipectus (Vikberg, 1965), Ph. oblita (Serville, 1823), Ph. polita (Zaddach, 1883); in Russia reared from Phyllocolpa sp. on Salix miyabeana Seemen; in Germany reared (Hinz, 1961) as Saotis liopleuris Thoms. from Phyllocolpa leucosticta Hartig.

21. Saotis varicoxa Thomson, 1894 (Figs. 3, 9) Thomson, 1894 : 2019, ♀ [Saotus; lectotypum ex-

aminavi: ♀, Sweden, “Pål”= Skåne, Pålsjö (Mus. Lund)]; Dalla Torre, 1902 : 276 (Saotis; bibl.); Schmiedeknecht, 1914 : 2810, 2818 (key, description); Meyer, 1936 : 296 (Russia); Fitton, 1982 : 80 (type); Kasparyan, Shaw, 2003 : 352–355 (key, fig.; UK); Kasparyan, 2004 : 48 (= compressus; = flavo-punctata); Kasparyan, Khalaim, 2007 : 548, 550 (fig., key).—compressus Szépligeti, 1898 : 76 [Zootrephes; holotypum examinavi: ♀, “1897.VI.12” (Mus. Budapest)].—flavopunctata Pfankuch, 1906 : 218, ♀ (type: Mesoleius bilineatus var. 1 in coll Gravenhorst, lost).

Diagnosis. The species may easily be distinguished by coloration of the scutellum (a pair of yellow spots at the base dorsally—Fig. 9), by yellowish ovipositor sheath with the truncate apex (Fig. 3), by separated epipleura of tergite 4 and usually 5. By these charac-ters it is similar to Saotis brevispina, but differs in having almost impunctate mesopleura, antennae slightly shorter, tergite 1 slender, and coloration of the apical part of the abdomen darker (cf. Figs. 1 and 3). However, all these characters are rather variable; S. variabilis may be only a form of S. brevispina.

Remarks. Saotis flavopunctata is considered here as synonym of S. varicoxa on the base of such notice-able character as the presence of two yellow spots at the base of the scutellum that is mentioned in the de-scription of Pfankuch; the other peculiarities of colora-tion in the description of flavopunctata are also similar to those in varicoxa.

Material. Great Britain. 2 ♀, England and Scot-land (NMS); 3 ♀, England (NHM). Sweden: 8 ♀

(syntypes), Lund (Mus. Lund). Bulgaria: Vitosha/BG, 5.07.1966 (Hinz) (Mus. Munich). Ukraine. 1 ♀, Cri-mean reserve, 15.06.1978 (Kasparyan) (ZIN). Russia. 1 ♀, Transbaikalia Territory, Adrianovka, 3.08.1975 (Kasparyan) (ZIN).

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