Received April 15, 2009

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. 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. Kasparyan a and J.-P. Kopelke b a Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia b Forschungsinstitut 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 Pontopristia. A key to species and subspecies is given and illustrated by 55 figures. For each species the literature, synonymy, 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 ichneumon-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 foodplants (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 gallmaking sawflies, two new species of the subfamily Ctenopelmatinae (Anoncus gallicola Kasparyan et Kopelke and Rhinotorus brachycerus Kasparyan et Kopelke) were described. A key to the species of this complex of Ichneumonidae was elaborated. The species 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. Species 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 deposited 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), collections 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 Museum of Lund University, Sweden (Mus. Lund, curator R. Danielsson), Naturhistoriska Riksmuseet, Stock- 71

72 KASPARYAN, KOPELKE holm, Sweden (Mus. Stockholm, curator T. Kronestedt), 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 Natural History, Washington, USA (Mus. Washington, curator R. Carlson), the American Entomological Institute, Gainesville, Florida, USA (AEI, curator D. Wahl). The authors cordially thank the curators of these collections and also A. Reschikov who furthered obtaining the material from the USA, Canada, and Finland (including photographs of some type specimens). 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 Forschungsgemeinschaft: 1990, 1993, 1999, 2000) and the research foundations (Ungerer-Stiftung, Rüppell Expeditions Fond) of the Senckenbergische Naturforschende Gesellschaft (SNG: 1985, 1988, 1997, 2001). Genus SAOTIS Förster, 1869 Type species Mesoleius (Saotus) brevispina Thomson, designated by Viereck, 1914. Dalla Torre, 1902 : 272 (catalogue, bibliogr.); Schmiedeknecht, 1914 : 2807 2820 (key, descriptions of species); Yu, Horstmann, 1997 : 436 437 (catalogue, bibliogr.); Kasparyan, Shaw, 2003 : 351 355 (key; = Iskarus); Kasparyan, 2007 : 547 551 (in: Kasparyan and Khalaim, 2007) (key). Iskarus Kolarov, 1987 : 70; type species Iskarus seleuciformis Kolarov, 1987 (= Saotis mirabilis Schmiedeknecht, 1914). This relatively small Holarctic genus comprises about 22 species, among which 20 have been described from Europe. The species of the genus can be readily distinguished by the strongly compressed abdomen 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 gallmaking nematine sawflies of the genera Pontania and Phyllocolpa (Tenthredinidae, Nematinae). One species, Saotis albionis Kasp., was reared in Finland from the larva of a nematine, Pontopristia amentorum developing in catkins of Salix. The evolutionary links of Saotis with the genus Mesoleius s. l. (including Campodorus) are rather clear. The peculiarities of the females behavior during the process of the host infestation have not been described and the morphofunctional significance of the compressed apical segments of the abdomen is unknown. The following 12 species-groups can be distinguished in the genus (the most important apomorphies are given in italics). 1 1. The brevispina species-group includes S. brevispina 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 convex 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 markings; 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 malar space ( gena ) to basal width of mandibles; i. ar. ap, index propodealis, ratio of area apicalis length to total length of propodeum along midline; i. terg, index tergalis, ratio of length of first metasomal tergite to its maximum width; i. nrv, index nervellaris, 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.

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 73 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 Phyllocolpa (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 (Greenland) 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 (immovably 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 (including 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 unknown. 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 unknown. 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 having basal flagellomeres shorter (i. fl 0.9). Small insects with fore wing 3.5 mm long; Antenna with 22 flagellomeres; i. gen about 0.5 (Fig. 28); i. nrv 0.28; i. fem 4.6; i. terg 1.7. Scutellum black; coxae and trochanters I predominantly black. Pterostigma pale with brownish margins. Host: Pontania. 7. The nigriscuta species-group; includes S. heteropus 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: Phyllocolpa. 8. The pygidiator species-group; includes S. pygidiator 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); nervulus usually interstitial, i. nrv 0.1 0.25, and pterostigma 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 basitarsus; areola of propodeum is narrow (Fig. 40); i. terg 1.7 2.0; tergite 2 with very fine and smoothed granulation 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.

74 KASPARYAN, KOPELKE 9. The granulator species-group; includes only S. granulator sp. n. The species may easily be distinguished 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); hypopygium truncate on hind margin and slightly concave 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 narrowed (Fig. 50); i. gen 0.33; i. fl 0.9; nervulus interstitial; ovipositor sheath as wide as hind femur; tergites completely black, without thin pale band on hind margin. 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. Mesoscutum 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 depressed, ovipositor sheath small. Mesothorax red. Host unknown. In the World catalogue of Ichneumonidae, Hypamblys salicis Teunissen, 1953 (holotypum examinavi; = Synodites) was erroneously placed (Yu and Horstmann, 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.

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 75 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 species (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 triangular 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. Flagellum 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 rufous, 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 yellow, usually truncate at apex (Fig. 1). Mesopleuron with distinct and rather dense punctures in lower 0.6. Antenna usually with 24 26 flagellomeres. 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 yellow; fore and middle coxae and trochanters yellow; 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; ovipositor 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

76 KASPARYAN, KOPELKE Figs. 1 10. Genus Saotis, female (Ichneumonidae, Ctenopelmatinae): (1 8) apical part of abdomen [(1) S. brevispina, (2) S. compressiuscula, (3) S. varicoxa, (4) S. bilineata, (5) S. nigriscuta (holotype), (6) S. morleyi, (7) S. dorsata, (8) S. nigriventris]; (9) S. varicoxa, scutellum, dorsal view; (10) S. bilineata, apical part of hind tibia with spurs. 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 separated from black mesopleuron by yellow longitudinal 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 antennal sockets, and with yellow markings above clypeus, on malar space and on cheeks; mesoscutum 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

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 77 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]. 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 usually with 27 30 segments...... 20. S. tricolor Thoms. (var. liopleuris Thoms.) Tergite 1 about 2.0 2.3 times as long as wide. Tergites 7 and 8 white laterally, tergite 8 almost entirely white (except for dorsal spot at base) (Figs. 6, 33). i. fl 0.9 0.95; segment 5 of hind tarsus not longer than segment 3. i. terg 2.2; i. gen 0.28. Epipleura of tergites 3 8 and all sternites 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 extreme base....... 14. Saotis morleyi Fitton (= albiventris Kasp.) [this species was considered to be a junior synonym 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; thorax 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-

78 KASPARYAN, KOPELKE ellar segments; clypeus impressed along lower margin; the body black to brown, with the following 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 almost 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 polished; hind tibia darkened dorsally, dirty whitish to dirty rufous in middle 0.5 ventrally. Lectotype: 24 flagellar segments; mesoscutum with anterolateral 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 yellowish brown (in type) to yellowish, truncate apically (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 tergite 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 often with basal blackish spot. All coxae and trochanters predominantly black...... 19. S. subarctor sp. n. Epipleura of tergite 4 and 5 not separated by crease (occasionally separated in S. dorsata). Mesoscutum 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, tergite 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 yellow, 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) separated by crease; ovipositor sheath brownish...... 8. S. dorsata Thoms. Metasomal tergites 2 4 dorsally black, without reddish brown markings. Ovipositor sheath wider and shorter. Mesoscutum black or often with anterolateral 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 granulation. Malar space very short (i. gen about 0.25). Sternites blackish, without distinct longitudinal 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 hypopygium (Fig. 39). Mesoscutum, scutellum and abdominal tergites black; coxae and hind trochanter I black or reddish; other trochanters predominantly yellow... 9. S. granulator sp. n. Thorax and tergites 1 4(5) predominantly or partly polished, or without distinct granulation. Malar

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 79 space longer, i. gen 0.3 0.5 (except for S. caucasica). Sternites usually yellowish, with distinct longitudinal fold; apex of hypopygium usually projecting backward... 13. 13. Abdomen strongly elongate, i. terg 2.8 (in holotype), black, tergites 2 and 3 without white apical band, tergites 7, 8 and ovipositor sheath brown (Fig. 15); sternites fuscous, hypopygium brownish yellow; ovipositor sheath as wide as hind femur; 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; nervulus almost interstitial (weakly postfurcal). Mesopleuron with smoothened granulation and sparse hairs; metapleuron with superficial granulation; 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 tegulae 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 tergites (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 perpendicular 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 anterolateral spots; scutellum black. Nervulus interstitial to slightly postfurcal; i. nrv 0.1 0.2; i. terg 1.7 2.0. Ovipositor sheath (Fig. 42) usually reddish brown, about 0.8 times as wide as hind femur (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 entirely 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 yellow spots near clypeal fovea. Flagellum almost uniformly yellowish rufous. Pterostigma pale yellowish brown... 14c. 14b. Coxae and trochanters I black. 23 26 flagellomeres. Pterostigma pale brown with dark margins. 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 flagellomere about 8 times as long as wide at midlength. Clypeus immediately above lower margin with distinct median convexity (Fig. 34). Ovipositor sheath black, very wide, about 1.2 times as wide as hind femur (Fig. 5). Nervulus interstitial or weakly postfurcal; spurs of hind tibia brownish. All coxae, trochanters and femora usually light reddish. Holotype: fore wing 4.9 mm; flagellum with 26 segments, slightly shorter (0.97) than fore wing; propodeal carinae vestigial; 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.

80 KASPARYAN, KOPELKE 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]. 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 yellow 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 propodeum very narrow and separated from apical area (Fig. 23). Segment 3 of hind tarsus 1.2 1.3

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 81 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]. 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 dorsoapical 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; trochanters 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; scutellum black... 4. S. boreator sp. n. 19. First tergite 2.2 2.9 times as long as wide, its lateral margins beyond spiracles subparallel (Fig. 19); flagellum with 21 24 segments; i. gen about 0.4. Tergite 2 polished in apical 0.3; apical 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

82 KASPARYAN, KOPELKE 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. Flagellum 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; epipleura 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 segments 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-segmented; i. fl 1.06; i. fem 4.2; i. nrv 0.28; tergite 1 weakly narrowed toward base (Fig. 48); ovipositor 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 reddish (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 margin. 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 punctures; 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 posterior 0.3 of second tergite and almost hairless; tergites 3 7 on hind edges moderately concave and with narrow pale band. Hind tibia white in middle 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 narrowed 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

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 83 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. longiventris, 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]. 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 rufous, at apex and dorsally brownish...... 19. S. subarctor sp. n. Antenna with 24 29 flagellomeres. Hind coxae and hind trochanters usually uniformly pale reddish or yellow. Hind tibia in middle part (0.2 0.4) whitish. i. fl 1.0 1.1... 27.

84 KASPARYAN, KOPELKE 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 segment of flagellum 5 times as long as wide at middle... 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 Roman, 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, usually 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 predominantly 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 nigriscuta), and the malar space is longer. Structurally the new species resembles S. compressiventris; both species 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 complete; 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 extending to apical 0.7 of tergite. Tergites 2 and 3 finely granulate, smooth and shiny in apical 0.2, their

A TAXONOMIC REVIEW OF ICHNEUMON-FLIES 85 epipleura separated by crease; subsequent tergites almost smooth; epipleura of tergite 4 weakly separated. 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, mandible, 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 trochanters 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 anterolateral 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 pronotum, 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 resembles 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 flagellum consists of 19 21 segments (22 24-segmented in S. heteropus and about 26 27-segmented in S. nigriscuta). S. boreator differs from S. alpinator (another small species with broad ovipositor sheath) in