Piogaster pilosator (Aubert, 1958)Figs 13, 14, 15Polysphincta pilosator Aubert, 1958: 79. Transferred to Piogaster by Aubert (1960 b).Piogaster rugosa Perkins, 1958: 266. Synonymized by Aubert (1965). Type lost.Piogaster punctulata Perkins, 1958, (syn. by Aubert, 1967 with doubt, indicated by “? ”; not adopted by subsequent authors).Piogaster pilosator caucasica Kasparyan, 1981: 70. syn. nov.Diagnosis.Piogaster pilosator can be distinguished from its congeners by possession of the combination of the following: 1) mesopleuron rugose (Fig. 14 B); 2) propodeum rugose to rugose punctate (Figs 13 B, 14 D); 3) in specimens with head primarily dark brown to black (Figs 15 C), metasoma is not pale yellow to white.This species is most similar to P. daisetsuzana, P. punctulata, and P. ussuriensis. Piogaster pilosator can be distinguished from P. daisetsuzana by the absence of medial transverse striations on the propodeum (present in P. daisetsuzana, Fig. 8 D) and the absence of the elongated fore wing vein ramellus (Fig. 14 F) or ramellus present only as a short stub (Fig. 13 A) (ramellus present and long in P. daisetsuzana, Fig. 8 F). Piogaster pilosator can be differentiated from P. ussuriensis by the colour of the head and metasoma and the lengths of the ovipositor sheaths (see couplet 7 of key). Piogaster ussuriensis has a primarily dark brown to black head and a contrasting pale yellow-white metasoma (Fig. 18 A) whereas P. pilosator specimens have the head and metasoma colour variable, from pale yellow to black (Figs 15 A – C), but never with contrasting dark brown to black head and pale metasoma as found in P. ussuriensis. Piogaster ussuriensis has ovipositor sheaths that range from 0.4–0.5 × as long as hind tibia (Fig. 18 A), while P. pilosator has ovipositor sheaths that range from 0.7–0.9 × as long as hind tibia (Figs 13 A, 15 A, 15 C). Piogaster pilosator can be differentiated from P. punctulata by the sculpture of the mesopleuron: rugose in P. pilosator (Fig. 14 B), and punctate in P. punctulata (Fig. 17 A).Redescription.Adult. Female [values in square brackets are from lectotype]. Body length 4.0–6.4 [6.4] mm. FW length 3.2–4.8 [4.6] mm. Head. Antenna with 20–24 [22] flagellomeres. Clypeus 1.9–2.4 [2.3] × as wide as high, matte to subpolished, weakly rugulose with fine, dense, setiferous punctation located mainly ventrally and dorsally, medially sparse to absent, with long white setae. Face, frons and vertex (Figs 13 C, 13 D) matte with short to long white to yellow setae, sculpture highly variable: all regions can be partly or completely rugulose, rugose, rugose punctate, granulate or punctate [rugose punctate]. Gena matte to subpolished, strigose ventrally, strigose to irregularly rugose medially and dorsally, with short to medium length white to yellow setae. Occipital carina complete (Figs 13 A, 14 A). MSL 0.8–1.0 [1.0] × as long as BWM. OOD 1.2–2.0 [1.6] × as long as LOD. Mesosoma. Pronotum with epomia absent; matte, rugulose to rugose to rugose punctate [rugose punctate], with sparse to dense short to medium length white setae. Mesoscutum matte, densely punctate to densely rugose punctate [densely rugose punctate], with dense short to medium length white to yellow setae. Scutellum matte, rugulose to rugose, with dense short to long white setae. Mesopleuron matte to subpolished, rugulose to rugose [rugose] with sparse to dense medium to long white setae (Figs 14 B, 15). Metapleuron matte to subpolished, rugose, with dense long white to yellow setae, sometimes setae absent medially (Fig. 14 C). Propodeum matte to subpolished, rugose to rugose punctate [rugose punctate], with dense long white setae (Fig. 14 D). Propodeum with pleural carina variable, absent to complete (Fig. 14 C), sometimes partially complete anteriorly or posteriorly [complete but weak]; lateral longitudinal carina present in posterior 0.2–0.5 [0.2], all other carinae absent. Wings. Fore wing. Vein Rs + M without ramellus extending significantly into cell 1 M + 1 R 1 (Fig. 14 F), at most, a stub of ramellus present (Fig. 13 B). Vein 2 rs-m 0.5–0.9 [0.8] × as long as M between 2 rs-m and 2 m-cu (Figs 14 E, 14 F). Vein 2 m-cu not thickened or angulate between bullae (Fig. 14 F). Hind wing. Vein 1 / Cu & cu-a inclivous and strongly angled apically where 2 / Cu intercepts, 2 / Cu intercepting in lower 0.3–0.4 [0.3], 2 / Cu long, tubular to spectral (Fig. 14 E). Metasoma. T 1 matte to subpolished, densely punctate to punctate reticulate [punctate reticulate] with dense long white setae anteriorly, dense medium to long white setae posteriorly (Figs 13 B, 14 G). T 1 median dorsal carina absent (lateral margin of dorsomedial sulcus rounded), or bordering sulcus at extreme base or entire length of sulcus, 0.1–0.4 × length of T 1; dorsolateral carinae present anterior to spiracle, or complete to almost complete but medially and / or posteriorly weak [complete to 0.9 length of T 1] (Fig. 14 C). T 2 – T 5 matte to subpolished, densely punctate to punctate reticulate, with dense medium length white setae (Fig. 14 H). T 6 subpolished, densely shallowly punctate, with dense medium length white setae (Fig. 14 H). T 7 and T 8 subpolished, densely finely punctate, with dense medium length white setae (Fig. 14 H). Tergites either without grooves and tubercles or with weak paired anterior tubercles and a weak medial transverse groove on T 2 – T 3 (Fig. 14 H). Ovipositor sheath 0.7–0.9 [0.8] × as long as hind tibia (Figs 13 A, 15 A, 15 C). Colour. Head, mesosoma and metasoma highly variable, yellow-white (Fig. 15 A) to yellow / orange-brown (Fig. 13 A) to red-brown (Fig. 15 B) to dark brown-black (Fig. 15 C), paler specimens generally darker dorsally and ventrally than laterally. Clypeus yellow-white (Fig. 15 A), orange-yellow, orange-brown or dark brown-black (Fig. 15 C), [yellow white (Fig. 13 C)]. Face pale yellow (Fig. 15 A), pale to dark orange, completely brown-orange, dark brown medially and brown-orange laterally, or dark brown-black (Fig. 15 C); ventrolaterally paler (light brown, yellow or white) in some of the predominantly orange specimens [orange, paler ventrolaterally (Fig. 13 C)]. Frons pale yellow, pale to dark brown-orange (Fig. 13 D), medium brown or dark brown-black; ventral orbit may be same colour as medially or paler, dorsal orbit is paler in all specimens (pale region contiguous with pale marking laterally on vertex) [pale brown-orange (Figs 13 B, 13 C)]. Vertex medially pale yellow, pale to dark brown-orange, medium to dark brown (Fig. 13 D) or brown-black; anterolaterally with white-yellow to yellow longitudinal ovoid, crescentic or subrectangular markings (Figs 13 B, 13 D), darker markings may also be present, especially posteriorly and in ocellar triangle [pale brown orange medially with crescentic yellow-white markings laterally (Fig. 13 B)]. Gena except ventral to eye yellow white (Fig. 15 A), light to dark orange brown (Fig. 15 B) to brown-black (Fig. 15 C), posteriorly and ventrally darker in some specimens; area ventral to eye (posterior of malar space) with white to yellow marking in form of vertical stripe (Fig. 15 B) or complete area paler (Fig. 13 A), this marking apparently absent in holotype of Piogaster pilosator caucasica Kasparyan, 1981 (Fig. 15 A) and Dagestan specimen (Fig. 15 C) [light orange-brown except yellow-white ventral to eye (Fig. 13 A)]. Occiput generally concolourous with posterior of vertex and gena or a bit darker. Mandibles except teeth yellow, light brown, orange-brown, medium to dark brown or black, teeth brown / black [dark brown (Fig. 13 C)]. Maxillary and labial palps yellow (Fig. 15 A), medium brown (Fig. 15 B) to brown / black (Fig. 15 C), darker basally than apically [medium brown (Fig. 13 A)]. Antenna light yellow to light brown basally, light to medium brown apically (Fig. 13 B). Mesosoma including pronotum, mesoscutum, scutellum, mesopleuron, mesosternum, post-scutellum, metapleuron and propodeum highly variable (see start of colour description and Figs 13, 14, 15). Palest (Fig. 15 A) and darkest (Fig. 15 C) specimens uniformly coloured; specimens with intermediate darkness (yellow-brown including holotype, orange or red-brown) with lateral areas (pronotum, mesopleuron, metapleuron, lateral propodeum) generally paler than dorsal and ventral areas (mesoscutum, scutellum, post-scutellum, anterior, dorsal propodeum and mesosternum) (Figs 13 A, 13 B, 15 B); yellow-brown and orange-brown specimens may have mesoscutum with paler, longitudinal markings where notauli would be and / or anteriorly and medioposteriorly in some specimens. Tegula white to pale brown [pale brown (Fig. 13 A)] in specimens with mesosoma predominantly yellow, yellow-brown or orange-brown; tegula brown to black in specimens with mesosoma predominantly red-brown to brown-black (Fig. 13 E). Wings hyaline, veins including stigma pale brown to transparent, Sc + R medium to dark brown. Legs highly variable, coxae yellow-white (Fig. 15 A), yellow-brown, orange, light brown to dark brown-black (Fig. 15 C) [orange (Fig. 13 A)], trochanters similar to coxae (yellow-white to dark brown), trochanters can be unicolorous with coxae in palest (Fig. 15 A) or darkest (Fig. 15 C) specimens, or have some dark and light colour in specimens of intermediate darkness [as in holotype, Fig. 13 B]; femora of holotype of P. p. caucasica completely yellow (Fig. 15 A), all other specimens have femora predominantly white to yellow with light to dark brown longitudinal stripe on dorsal and / or ventral surfaces and laterally at base in most specimens (especially darker ones) (Figs 13 A, 13 B, 15 B, 15 C), tibiae with dorsal and in most specimens, ventral surface dark with light to dark brown longitudinal stripe (holotype of P. p. caucasica with light brown dorsally near apex only); tarsi from pale brown to dark brown, generally paler ventrally than dorsally and apically compared to basally [pale brown (Fig. 13 A)]. Metasomal tergites yellow-white, pale yellow (Fig. 15 A), orange-yellow (Fig. 13 B) or dark brown-black (Figs 15 B, 15 C); specimens with dark mesosoma (red-brown to brown-black) have dark metasoma (brown to brown-black) (Figs 15 B, 15 C); whereas specimens with pale mesosoma (yellow to orange-brown) can have metasomas ranging from yellow-white to orange-brown. Most specimens have the metasoma uniformly coloured, although it may be somewhat paler laterally on some segments and paler in posterior segments; however, some specimens have the posterior of T 2–5 with pale transverse bands [orange-brown, slightly paler medioposteriorly on T 5, posteriorly and laterally on T 6–7 (Fig. 13 A, B)]. Ovipositor sheath white or yellow (Figs 13 B, 15 A) to medium / dark brown (Fig. 15 C), uniformly coloured, darker apically or paler apically [yellow, light brown apically (Fig. 13 A)].Male. Unknown. Male specimens have been collected at the same localities as females of P. albina and P. pilosator; however, European males are morphologically cryptic and therefore species identifications of male specimens of both these species cannot be confirmed.Distribution.Fig. 30. Western Palaearctic: Austria (Kazmierczak 1990), Belgium (Verheyde et al. 2021), Bulgaria (Kolarov 1989, 1997), Finland (Koponen et al. 1995), France (Aubert 1960 b; Aubert 1960 a), Germany (new record), Norway (new record), Poland (Kazmierczak 2004), Russia (Kasparyan 1981), Sweden (Perkins 1958), Switzerland (Klopfstein et al. 2019 b), Ukraine (Varga 2021). Aubert (1969 b) listed a record of P. pilosator from England. However, this record references Perkins (1958) and therefore it is most likely referring to the holotype of P. punctulata which was from England as Aubert (1969 b) synonymized P. punctulata under P. pilosator (see more about this synonymy in the Comments of the P. punctulata description).Biology.Piogaster sp. (cf. pilosator (Aubert)) oviposited on the cephalothorax of S. cingulatus (Salticidae) in a laboratory setting (Takasuka et al. 2018). We were unable to locate these two specimens for identification; however, given that this was a European species, it seems very likely it was P. pilosator considering the clear morphological distinction from females of P. albina and P. lucida and the fact that the author who observed these interactions (N. Fritzén) had previously examined the type of P. punctulata (G. Broad, pers. comm.).Material examined.Primary types.Polysphincta pilosator pilosator Aubert, 1958 lectotype (hereby designated):France • ♀; Alpes-Maritimes, Ile Ste. Marguerite; 9. viii. 1958; J. A. Aubert; [MZLS] (Figs 13 A – C).Condition of type: Intact, pinned through mesoscutum.Piogaster pilosator caucasicus Kasparyan, 1981 holotype:Russia • 1 ♀; Caucasus, Krasnodar Krai Lazarevskoye, Sochi; 1. vi. 1976; Kasparyan; [ZIN] (Fig. 15 A).Condition of type: Metasoma detached from body and glued to point. Left hind leg missing. Distal three right maxillary palpomeres missing.Other material.Austria • 1 ♀; Tirol, Zillertal Alps, Schlegeisgrund; 5. vii. 1967; NHMW -HYM 0022253; [NHMW]. Belgium • 1 ♀; West Flanders; Ruiselede, Gulke Putten; 51.07528°N, 3.34472°E; 22. v. 2020; A. De Ketelaere; Light trap; [Personal collection of Augustijn De Ketelaere] (photo only). France • 1 ♀; Orléans, Ardon, Institute National de la Recherche agronomique, 2163 Avenue de la pomme de pin, 45075 Ardon; 12. iii. 1991; D. Rougon; Emerged 6. v. 1991; host unknown; MNHN -EY-EY 51; [MNHN] • 1 ♀; St. Tropez; vi. 1980; Z. Boucek; [NHMUK] • 1 ♀; same data for proceeding; NHMUK 012850732 [NHMUK] • 1 ♀; Lot-et-Garonne, Bernac; 26. vi – 3. vii. 1991; R. R. Askew; Malaise trap; [NMS] • 1 ♀; St. Tropez; 13. viii. 1980; Boucek; [NHMUK]. Germany • 1 ♀; Bayern, Berchtesgaden-Schönau, Königsbachalm; 47.5693°N, 13.01144°E; 1222 m; 30. vii – 9. viii. 2018; D. Doczkal & V. Voith; [private collection of H. Haraldseide] (photo only). Norway • 1 ♀; Sør-Trøndelag, Malvik, Skjeltjønnbekken; 63.37499°N, 10.67403°E; 8. vii – 5. viii. 2019; Arnstein Staverløkk; Malaise trap; NOCER-678; [NINA] • 1 ♀; Innlandet, Vågå, Lye-Fellese, EIS 71, ON; 61.86734°N, 9.05867°E; 30. vi – 4. viii. 2018; Frode Ødegaard; Malaise trap; NOCER-587; [NINA] • 1 ♀; Telemark, Vinje, Vådalen, Lauvheim; 59.74674°N, 7.84290°E; 28. iii – 11. v. 2022 [date interpreted, date listed verbatim as 28. iv – 11. iiv. 2022]; NINA / Arnstein Staverløkk; seminatural habitat; Malaise trap sample 2; CNC 1754423; [NINA] • 1 ♀; Sør-Trøndelag, Trondheim, Sommerlystvegen 22; 63.40531°N, 10.38222°E; 29. vi – 6. vii. 2014; E. Stur and T. Ekrem; 75 moh, Malaise trap; NTNU -VM-181569 [NTNU]. Russia • 1 ♀; Dagestan, Rutul; 29. vi – 1. vii. 2018; 1400–1500 m; Astafurova; CNC 310411; [ZIN]. Sweden • 1 ♀; Västergötland, Hisingen, Säve s: n, Sävholmen; 57.84333°N, 11.90667°E; 11–20. vii. 1996, Michael Sporrong; Malaise trap; damp to marshy meadows with Salix; [EUMJ] • 1 ♀; Gothenburg; 57.67907°N, 11.92913°E; 12. vi. 2020; Sweep net; garden; [Private collection of Johan Ennerfelt] (photo only) • 1 ♀; Västergötland, Västra Götaland, Härryda, Råda portar 60, Råda portar, Råda, Vg; 57.66816°N, 12.09508°E; 25. v. 2023; Oscar Josefsson; [SLU Artdatabanken] (photo only). Switzerland • 1 ♀; Jura, Delémont, CABI; 47.37306°N, 7.32472°E; 26. v – 19. vi. 2014; J. Squire; Malaise trap; forest edge; CNC 842257; [CNC] • 1 ♀; Engiadina Bassa / Val Müstair; iv. 2015; Marc Neuman; Ichn-2144; [NMBE] • 1 ♀; Grisons / Graubünden, Tschierv; 18. vii. 1951; J. de Beaumont; [MZLS]. Ukraine • 1 ♀; Odesa Region; Prymorske; 5. vi. 1996; Anatoly Kotenko; [SIZK] (photo only). Unknown • 1 ♀; 1915; J. Perez; [MNHN].Comments.Aubert (1958) did not assign a holotype in the original description of P. pilosator, but listed two type specimens (syntypes) which were acquired by the Musée de Zoologie in Lausanne, Switzerland. These specimens were assumed loaned and lost (Klopfstein and Baur 2011), but one of the two syntypes was located and examined for this study. This specimen has been designated as the lectotype. The type specimen of P. rugosa (now synonymized with P. pilosator) was also lost. It should have been at the Lund University Museum. The archives at the Lund University Museum list one ichneumonid of a new genus, according to Dr. Henry Townes, loaned to J. F. Perkins in 1958. The material was not returned after the specimen was described as Piogaster rugosa in Perkins (1958). The Natural History Museum in London, where J. F. Perkins was employed, has been searched and this specimen was not located.Piogaster pilosator caucasica was briefly described only in a key and the only characters of note are colour related, based on a single pale specimen collected in southcentral Russia (Kasparyan 1981) (Fig. 15 A). Upon examination of 18 P. pilosator specimens (including the P. p. caucasica specimen) and seeing photos of an additional 10 specimens, it is clear P. pilosator has a wide range of colour variation (Figs 15 A – C). We have seen another similarly pale yellow specimen from Bernac, France, which is geographically isolated from the type of P. p. caucasica. As we could find no further morphological characters distinguishing the holotype of P. p. caucasica from other P. pilosator, the subspecies caucasica is here placed in synonymy with P. pilosator.