Taeger et al. (2010) also listed Abia nitens var. vernetensis Pic, 1928 (type locality: France, Allier) as a junior synonym of Abia nitens auct. However, no type specimen has been examined subsequent to its description, as far as we are aware. Pic wrote [translated from French] “differs from the typical form in the black-marked base of the posterior femora”. This suggests that it possibly does not belong to A. brevicornis.

Abia nitens was successively mentioned as occurring in Sweden by Fallén (1807), Dahlbom (1836), and Thomson (1871), who all placed Tenthredo sericea as its synonym. Thomson’s description of leg color indicates that he had before him specimens of the species called by Taeger (1998) A. sericea, or perhaps A. candens [not distinguished from the former in Thomson’s time]. Influenced by the continued, widespread use of the name Abia nitens in Scandinavian literature, Taeger et al. (2006) and ArtDatabanken (2015) included Sweden within the range of A. brevicornis [as A. nitens auct.]. In fact, no Swedish specimens of A. brevicornis have been located in the MZLU or NHRS collections, and there are no published records from other Fennoscandian countries. Abia brevicornis is restricted to extremely dry, summer-warm sites, where its larval hosts occur (Scabiosa spp.) (Liston and Späth 2006). Probably it has a strongly continental distribution: its most north-westerly known localities in Europe are in Central Germany, whereas it is not definitely known in France (Noblecourt 2020), and a single old record from Spain needs confirmation. Although a presence of A. brevicornis in southern Sweden cannot be ruled out, we consider it likely that all references to A. nitens auct. in Sweden relate to either A. nitens [= sericea] or A. candens.

To help resolve the taxonomic disagreements in the interpretation of these nominal taxa, and promote the future stability of nomenclature, a neotype is designated for Astatus punctatus:

Astatus punctatus Klug, 1803. Neotype ♀ (DEI-GISHym21255, Fig. 4A–D), hereby designated. Germany, Brandenburg, Landkreis Märkisch-Oderland, Müncheberg, Trebnitz, 52.535°N, 14.204°E, damp meadow, swept from Alopecurus pratensis, 16.05.2015, leg. A. Liston (deposited in the SDEI). Labelling [printed on pale paper if not stated otherwise]: “Germany: Brandenburg; Landkreis Märkisch-Oderland, Müncheberg Trebnitz 16.05.2015 leg. A. D. Liston”, “21255” [handwritten] with part of a leg gummed to card], “DEI-GISHym21255”, “♀ Calameuta punctata (Klug) [handwritten] det. A. Liston 2018”, “NEOTYPE ♀ Astatus punctatus Klug, 1803 designated A. Liston 2022” [red]. Klug’s description states that abdominal segment 4 has obscure, paired dorsal spots; segment 5 four separate yellow spots, one pair laterally, the other dorsally; segments 6 and 7 with spots on their lower posterior margins; segment 8 immaculate; segment 9 completely yellow. Accordingly, we selected as neotype a specimen with small pale markings on terga 4–7 as well as 8–10 (Fig. 4A–D). The abdomen of a second female (DEI-GISHym21260) collected at the same place and time has fewer and less extensive pale markings and is thus intermediate in this respect to other female C. punctata specimens collected in Germany and all known Finnish and Estonian specimens, which have a completely black abdomen apart from terga 8–10 (Fig. 3A, C).

Figure 3. 

Calameuta punctata (Klug, 1803) ♀ (DEI-GISHym12236). A. Dorsal habitus; B. Head and thorax, dorsal. Mesoscutum (arrow); C. Lateral habitus; D. Maxillary palp. Palpomeres 5 and 6 (arrows); E. Head and thorax, ventral. Scale bars: 1 mm.

Figure 4. 

Calameuta punctata (Klug, 1803): neotype ♀ Astatus punctatus Klug, 1803 (DEI-GISHym21255). A. Dorsal habitus; B. Lateral habitus; C. Abdomen, lateral; D. Abdomen tip, lateral. Scale bar: 1 mm.

Astatus floralis and A. analis have in the past generally been treated as synonyms of Calameuta haemorrhoidalis auct. [our variabilis], e.g. by Konow (1905a). The opinion that A. floralis is a synonym of Cephus pygmeus (Linnaeus, 1767), as in de Dalla Torre (1894), cannot be accepted: Klug’s description of leg colour does not fit C. pygmeus. Our reason for placing A. floralis and A. analis as synonyms of Calameuta punctata rather than of C. variabilis is based primarily on one of the main characters which distinguishes C. punctata from C. variabilis: the structure of the maxillary palps. In the description of Astatus which precedes the descriptions of A. floralis, A. analis and A. punctatus, Klug characterized the genus thus: “Palpi[...]anteriores[...]sexarticulati, articulis duobus baseos cylindricis, aequalibus, tertio crassiori, longiori, subcylindrico, quarto longissimo, graciliori, quinto brevissimo, ultimo longitudine fere tertii subulato[...]”. The described proportions of maxillary palpomeres 5 and 6 therefore fit C. punctata (Fig. 3D), not C. variabilis (Fig. 5B). Calameuta variabilis is unique in Calameuta in having maxillary palpomeres 5 and 6 of almost equal length (Gussakovskij 1935; Benson 1968; Zombori 1978). Zombori (1978) correctly identified Calameuta variabilis [which he called C. haemorrhoidalis] as a taxon distinct from C. punctata, and summarized the characters that distinguish them, but interpreted some of the names wrongly. Notably, Zombori (1978) did not mention the major contradiction in the morphology of the maxillary palps, as described by Klug, when he tentatively suggested that A. floralis and A. analis might be synonyms of haemorrhoidalis auct. [“the description of the latter two [floralis, analis] rather corresponds to the one given by Fabricius for C. haemorrhoidalis, accordingly, they are considered as synonyms of the latter name.”]. Zombori’s main reason for doubting that analis was synonymous with punctata, seems to have been the wording of Klug’s descriptions, which suggested that the thorax of analis is shinier than that of punctata. Apart from this, Klug’s description of A. analis fits the darker forms within the rather wide range of variability in the female sex of C. punctata. Strangely, in his discussion of these names, Zombori (1978) does not mention Calameuta filum at all.

Figure 5. 

Calameuta variabilis (Mocsáry, 1886). A. Lateral habitus and labels, lectotype ♀ Cephus variabilis Mocsáry, 1886; B. Maxillary palp ♀ (DEI-GISHym14072). Palpomeres 5 and 6 (arrows); C. Thorax, dorsal ♀(DEI-GISHym14072). Mesoscutum (arrow).

The explicit collection data given by Klug (1803) for Astatus floralis (“Locus in editioribus argillosis; in floribus”), A. analis (“Locus in editioribus; in floribus”), and A. punctatus (“Locus in floribus”) are, in part, not easy to interpret. Clearly, “in floribus” means that the specimens were collected from flowers. We think that “in editioribus argillosis” refers simply to the type of locality, i.e. an elevated place on clayey ground. This fits well with the type of sites at which C. punctata has recently been collected in Germany (see below).

The synonymy of Calameuta filum with C. punctata can be proposed with a high degree of confidence. The characters described by Gussakovskij (1935) for the former are precisely those used by Zombori (1978) to characterize the latter. The same characters are also given by Viitasaari (1975) in his description of Finnish specimens identified as Calameuta filum, and which he compared with a syntype of that species. Viitasaari (1984) subsequently noted that Calameuta punctata sensu Zombori (1978) and C. filum are probably conspecific.

The only recorded host plant of Calameuta punctata is Alopecurus pratensis L. (Vikberg 1978; Liston 2015), on which it is possibly monophagous. Accordingly, C. punctata occurs mostly in rather moist places. Its wider geographic range is not entirely clear, particularly because the identity of Calameuta pravei (Dovnar-Zapolskij, 1926) remains unresolved. This has been considered to be a valid species (e.g. Llorente and Gayubo 1990), or a synonym of C. punctata (e.g. Taeger et al. 2010). Calameuta pravei was recorded by Gussakovskij (1935) from Transcaucasia, Crimea, and the western Kopet-Dagh (Turkmenia). Llorente and Gayubo (1990) added records from Spain. Excluding these records, C. punctata is known from south-east Russia, Transcaucasia and south Siberia to Irkutsk (Gussakovskij 1935, as Cephus filum), and central and northern Europe (specimens examined by us). The distribution of C. punctata is therefore rather different from that of C. variabilis (see below), but their ranges overlap at least in parts of south-central Europe.

Calameuta variabilis [as C. haemorrhoidalis auct.] has been stated to occur in Germany based on the mention of Germany as the type locality in the original descriptions by Fabricius (1781) of Tenthredo haemorrhoidalis and by Klug (1803) of Astatus punctatus, A. floralis and A. analis, coupled to an apparently faulty understanding of which taxa are represented by these names. Blank et al. (1998, 2001) listed C. haemorrhoidalis from Germany, only for Berlin-Brandenburg, dating the record respectively as “vor 1803” and “1802”. This refers to the type material of Klug’s species. Later, Liston et al. (2012) treated C. haemorrhoidalis auct. [C. variabilis] as extinct in Germany, and added C. punctata to the German list, based on recently collected specimens. Although it cannot be ruled out that C. variabilis once occurred in Germany, but has since disappeared, we think it more likely that in historical times only C. punctata ever occurred there, and propose in future to include only it in the list of German Symphyta. Fennoscandian and Estonian specimens which were previously identified as C. filum also belong to C. punctata.

Based on COI sequences, C. punctata is split into two barcode clusters. Three specimens from Finland and one from Estonia are identical (BOLD:ACQ7596), but differ from two German specimens by 5.0–5.5% (no BIN assigned yet, GenBank accessions MW353981 and MW353982). The BOLD:ACQ7596 is closer to C. pallipes, differing by a minimum of 4.1%.

Lectotype of Cephus variabilis Mocsáry, 1886, hereby designated: ♀, id nr.017651 HNHM Hym.coll. (Fig. 5A); labels (Fig. 5A). Type locality: Romania, Herkulesfürdõ (HNHM). Paralectotypes (all HNHM): 5♂ and 2♀ also belong to the type series and have been labelled as paralectotypes: details of their sexes and localities were given by Zombori (1978). Note that the type series is heterogeneous, and contains specimens of Calameuta punctata as well as C. variabilis. Zombori noted the heterogeneity of the type series, but did not publish a lectotype designation, although the specimens were labelled by him as lectotype and paralectotypes. The types were found by Z. Vas in the HNHM, grouped as stated by Zombori. We do not follow Zombori’s intention according to his labels, but designate the female from Herculesfürdö (= Baile Herculane, Romania) (id nr. 017651 HNHM Hym. coll., labelled by Z. Vas) as lectotype (Fig. 5A). Thus, Calameuta variabilis (Mocsáry, 1886) comb. nov. can be used as a valid name for Calameuta haemorrhoidalis sensu Gussakovskij et auct.

Cephus atripes Stephens, 1835 has sometimes been listed as a synonym of C. variabilis, e,g. by Taeger et al. (2010, under Calameuta haemorrhoidalis). The type specimen, or specimens, is probably lost. The description is short, and does not state the sex of the described specimens(s). de Dalla Torre (1894) treated C. atripes as a synonym of C. pygmeus (Linnaeus, 1767) and Konow (1905a) as a synonym of C. haemorrhoidalis. The color pattern described by Stephens does not fit very well with either of these, in either sex, nor with any other known north-west European cephid species. Stephens’ name is best treated as a species inquirendae.

According to Macek et al. (2020, under Calameuta haemorrhoidalis) the host plants of C. variabilis are various Poaceae, including cereals such as rye (Secale cereale) and wheat (Triticum). According to our personal experiences, adults occur in dry places, mostly on or near wild grasses. The taxon to which the name C. variabilis is now applied has so far been found only in the West Palaearctic, and has an essentially Mediterranean distribution, summarized by Gussakovskij (1935) as comprising southern Europe, Crimea, Caucasus, western Turkmenia (Kopet-Dagh) and Syria. In Central Europe, it reaches at least as far north as Hungary (Zombori 1978), but specimens recorded from Austria and the Czech Republic (see Taeger et al. 2006) should be checked.

Cephalcia intermedia has in recent years mostly been understood as a distinct species close to C. arvensis (e.g. Viitasaari 2002b, Taeger et al. 2006), but with more extensively dark-patterned adults, occurring in northern Europe and the central European mountains. Because of its dark coloration, specimens of C. intermedia can also be mixed up with C. lariciphila (Wachtl, 1898), but the host plant of C. intermedia is Picea, as in C. arvensis, not Larix as in C. lariciphila. The status of C. intermedia and the supposed differences to C. arvensis were discussed by Vikberg (1982), Shinohara (1985), Midtgaard (1987) and Viitasaari (2002b). The only differences between C. arvensis and C. intermedia are in coloration: mainly of the abdomen, and less so of the antennae and legs. Shinohara (1985) pointed out that an unbroken range of color variability occurs between pale C. arvensis and the dark specimens identified as C. intermedia. This is in accordance with our observations, as the amount of dark color on the male abdomen varies considerably even in a single locality (Fig. 6). Small differences in colouration of larvae are possibly not constant, as the larvae of C. intermedia used for comparison were offspring of a single female. At present, there seems to be no convincing evidence for treating C. intermedia as a species distinct from C. arvensis, and we therefore synonymize them.

Figure 6. 

Cephalcia arvensis Panzer, 1802. Variability in colour pattern, dorsal habitus. A–D. Males (ZMUO.032092, ZMUO.033004, ZMUO.030747, GP.108460). E, F. Females (ZMUO.040720, GP.108471).

Substantiating the statement about the host plant by Benson (1952), the ZMUO specimens, collected in the Helsinki area, were reared from larvae feeding on cultivated Fragaria.

Figure 7. 

A. Claremontia confusa (Konow, 1886). Dorsal habitus ♀ (ZMUO.044470); B, C. Claremontia brevicornis (Brischke, 1883), dorsal habitus; B. ♀ (ZMUO.039550); C. ♂ (ZMUO.031990).

ZMUO specimens, from Finland, are not reared; Sanguisorba, recorded as a host plant by Benson (1952), can be excluded as a possible host plant at these localities, but Potentilla erecta is present in abundance at one of the localities in Eastern Finland.

Konow (1886a), in a key, described the tibiae of Claremontia confusa (as Blennocampa confusa) as mainly black with only the knees very narrowly whitish and the protibia only whitish on the anterior face, and wrote that the tibiae of C. puncticeps were mainly pale. Benson (1952) and Lacourt (2020), among others, distinguished Claremontia brevicornis (as confusa) from C. puncticeps using several characters, also including the color of the metatibia of females. However, Brischke (1883) described the metatibia of Claremontia brevicornis, a nominal species not known to Konow (1886a), as extensively yellow-white. Numerous barcoded females (SDEI, ZMUO) belong to two separate COI sequence clusters (distance 3.1–5.8%), which correlate fully with their leg color. Accordingly, we think that the original description of C. brevicornis refers to what has more recently come to be known as C. puncticeps, and that the correct name for the other species is C. confusa. Unlike some previous authors, we did not detect a clear difference in the sculpture of the head of the two forms. Distinction of the males is problematic, because of the lack of reliably identified male specimens of C. confusa. Benson (1952) stated that C. confusa “is entirely parthenogenetic, at least in Britain”. On the other hand, several male specimens of C. brevicornis (Fig. 7C) have been barcoded, and can thus definitely be associated with that species. Based mainly on barcode-sequenced specimens in ZMUO, females of Claremontia confusa and C. brevicornis may be separated as follows.

Dolerus aericeps Thomson, 1871. Lectotype ♀ hereby designated, labelled: “Båst[ad]” [Type locality: Sweden, Skåne, Båstad], “aericeps” [blue line along upper margin], “MZLU 2013 416” [pale green, printed, loan record], “Lectotypus 2014 Dolerus aericeps ♀ Thomson, 1871 M.Heidemaa design.” [printed, red], “Dolerus aericeps Thomson, 1871 M.Heidemaa det.”. In excellent condition. MZLU. Paralectotypes: 2♀, 2♂. MZLU.

Tenthredo eglanteriae Fabricius, 1793. Lectotype ♀ hereby designated, labelled: “eglan[...]riae”, “LECTOTYPE Tenthredo eglanteriae Fabricius, 1793 des. SMBlank 2008” [red, printed], “Dolerus eglanteriae (F.) = bajulus Serv. [handwritten] det. S.M.Blank 2008 ✓” [printed], “ZMUC-GISHym 1014”, “Dolerus aericeps Thomson, 1871 M.Heidemaa det.” [printed]. Left hind tarsus and some other tarsomeres missing. ZMUC.

Among 2♀ and 1♂ specimens preserved in the ZMUC collection under the name T. eglanteriae, only the above female can be regarded as a syntype. The scutellum of the other female is red, and the tip of the abdomen of the male black. Both characters conflict with Fabricius’ original description. Compared with the other two specimens, the pale parts of the legs of the lectotype are darkened. The lectotype agrees with the species which has for a long time been known as Dolerus aericeps Thomson, 1871 (Zhelochovtsev 1994), characterized for example by the laterally directed and distally curved setae of the ovipositor sheath. Lacourt (2000) used the name D. bajulus Serville, 1823 for this species, but Blank et al. (2009) presented arguments for the use of the name D. aericeps (nomen protectum) versus D. bajulus (nomen oblitum). Tenthredo eglanteriae was synonymized with T. germanica by Klug (1819) and with Athalia glabricollis Thomson, 1870 by Konow (1897a). Since Brullé (1846), T. eglanteriae (nomen oblitum) has never again been used as valid, while between 1971–2021 the name Dolerus aericeps (nomen protectum) was used as valid by more than 120 authors in over 150 publications (Article 23.9.1, ICZN 1999). One paralectotype male of D. cothurnatus Serville belongs to this species (see under D. germanicus).

Tenthredo (Dolerus) palustris Klug, 1818. Lectotype ♂ hereby designated, labelled: “14190” [collection catalog no.], “cotypus von Dol. palustris Klg. = aericeps Thoms. O.Conde det 1941” [handwritten by Otto Conde], “GBIF-GISHym 2342” [printed], “Lectotypus [printed] ♂ Tenthredo (Dolerus) palustris Klug, 1818 [handwritten in block letters] M.Heidemaa des. [printed]” [red], “Dolerus aericeps Thomson, 1871 M.Heidemaa det.”, “Zool. Mus. Berlin” [printed]. Left forewing and metatarsi missing, half of the genital capsule glued on a paper card and pinned with the lectotype. Paralectotype ♂ (“GBIF-GISHym 2341”) with same labelling (except “Paralectotypus”). All in ZMHB.

Formerly placed as a synonym of D. cothurnatus auct. (= D. junci (Stephens, 1835)), e.g. by Taeger et al. (2010).

Dolerus bajulus Serville, 1823. Lectotype ♀ hereby designated, labelled: “Dolerus bajulus, Lep.”, “Lectotype [red, printed in block letters]”, “Lectotype D. bajulus LEP. det.: H.Haris 1996” [white], “D. aericeps THOMS. ♀ det.: A.Haris”, “Lectotypus [printed] ♀[handwritten] Dolerus bajulus Serville, 1823 [handwritten in block letters] M.Heidemaa design.” [printed, red]. “Dolerus aericeps Thomson, 1871 M.Heidemaa det” [printed]. MNHN.

Holotype ♀. Figs 8A, B, 9B, C.

Colour. Fig. 8A, B. Body black with terga 2–7 and sterna 2–7 orange. Terga 8–9 brownish black, 10 brownish basally but yellowish brown apically, apical third of protibiae brownish. Ventral margin of valvula 3 and cerci brownish. Wings clear.

Figure 8. 

Dolerus aericepsellus sp. nov. A, B. Holotype ♀ (ZMUO.044566); A. Dorsal habitus; B. Lateral habitus and labels (below); C, D. Paratype ♂ (ZMUO.035740); C. Dorsal habitus; D. Lateral habitus.

Measurements (mm). Body length: 5.0, distance from tegula to base of pterostigma: 3.1, head breadths: 1.46 (max. at eyes), 1.28 (max. behind eyes), 1.07 (min. behind eyes), head length (behind eyes): 0.21, max diameter of eye: 0.62, breadth of postocellar field: 0.36, OC: 0.96, OOCL: 0.35, OOL: 0.20, POL: 0.19. Length of: metafemur: 1.18, metatibia: 1.76, ovipositor sheath (from the base of valvifer 2 to the apex of valvula 3): 1.01. Length of flagellomeres: 1: 0.48, 2: 0.41, 3: 0.37, 4: 0.31, 5: 0.29, 6: 0.26, 7: 0.25.

Large structures. Clypeal emargination almost half as deep as clypeal median length. Clypeus asymmetrical, its antero-lateral lobes round at apex, the left lobe longer than the right. Distance between antennal sockets 1.6× as long as the malar space. Length of antennomere 3 about 1.24× that of 4. Oblique furrow not outlined. Distance between cenchri about 1.1× as long as width of cenchrus. Metatarsomere 1 about 1.75× as long as 2. Mesoscutellar appendage with weakly outlined ridge medially.

Setae. Metascutellum with few setae (about 5). Abdomen dorsally mostly pubescent from segment 7 to last one (laterally and ventrally from 2 to last one). Longest setae of valvula 3 in dorsal view markedly curved in apical third, and forming an angle of about 80–90° (Fig. 9B).

Figure 9. 

Dolerus aericepsellus sp. nov. A. Head frontal, paratype ♂ (ZMUO.035740); B. Sawsheath dorsal, holotype ♀; C. Thorax lateral, holotype ♀, metepimeron (arrow); D–G. Paratype ♂ (ZMUO.035740); D. Thorax dorsal; E. Thorax lateral; F. Head dorsal; G. Penis valve.

Macrosculpture. Pits on head rather irregular in size and distribution, more distinct and separated on postocular and postocellar area. Density and size of pits on thorax vary, the largest (partly fused) on lateral sides of median mesoscutal lobes and on mesoscutellum. Pits on lateral lobes of mesoscutellum and near the median mesoscutal groove sparse and small. Pits on mesepisternum largest, partly isolated and round, partly polygonal and fused (Fig. 9C). Distinct pits on pectus nearly absent (Fig. 9C).

Microsculpture. Meshes on mesoscutellar appendage extensive, on tergum 1 absent, on metepimeron (Fig. 9C) and on ventral surface of metacoxa absent. Sculpticells on anterior half of katepimeron of mesopleuron rib-like, irregular and of uneven height, and absent on metepimeron, pectus, and terga (all glossy).

Male. Figs 8C, D, 9A, D–G. Closely resembles the female, but the anterior margin of tergum 2 blackish, basal 2/3 of hind tibia (with apical spurs), and base of metatarsomere 1 reddish yellow (Fig. 8C, D). Penis valve valviceps long and narrow (Fig. 9G).

Measurements (mm). Body length (paratype male ZMUO.035740): 5.3, distance from tegula to base of pterostigma: 3.2, max. diameter of eye: 0.60, head breadth: 1.47 (max. at eyes), breath of postocellar field: 0.38, head length (behind eyes): 0.24, OC: 1.01, OOCL: 0.18, OOL: 0.29, POL: 0.21, metafemur: 1.25, metatibia: 1.64, metatarsus: 0.50. Length of flagellomeres: 1: 0.59, 2: 0.51, 3: 0.50, 4: 0.44, 5: 0.41, 6: 0.39, 7: 0.41.

Three specimens with DNA barcode sequence data are available. They comprise a distinct cluster BIN BOLD:ABV8002, with 0.3% intraspecific variability. Based on sequences longer than 600 bp, the minimum distance to the closest relative D. aericeps (n=26) is 3.9%, but a few specimens identified as D. yukonensis from North America show a minimum distance of even less, 3.6%.

Adults of D. aericepsellus are most similar to D. aericeps Thomson, but are distinguished by the following characters: smaller body size (similar to D. elderi Kincaid, 1900, or smaller); very short, strongly narrowing postocular area; glossy metascutellum with few distinct pits and setae (3–6); pectus without distinct pits; and thorax black, including tegulae (based on the barcoded specimens: 1♀ 2♂). Lateral postocellar furrows very distinct in both sexes (Fig. 9F), like in D. aericeps and D. incisus. The thorax of female D. aericeps is usually largely red, whereas in the D. aericepsellus paratype female it is almost entirely black. Dolerus aericeps ab. nigricollis was described by Lindqvist (1943) from northern Finland close to the localities where D. aericepsellus was collected. We examined the type specimen of Dolerus aericeps ab. nigricollis deposited in the MZH and found that it is not conspecific with D. aericepsellus but with D. aericeps, although it has a nearly completely black thorax, with two obscure reddish flecks. Some other examined specimens of D. aericeps from northern Finland show reduced red markings on the thorax, suggesting that this character varies in D. aericeps.

Holotype ♀. Type locality: Finland, Enontekiö, Pousu. Labelled [white, printed if not given otherwise]: “FIN Le Enontekiö Pousu 7651:3266 [68.846°N, 21.197°E] 27.6.2020 Marko Mutanen leg.”, “Sawfly tissue 2020-1079 M.Mutanen” [greenish], “http://id.zmuo.oulu/fi/ZMUO.044566”, “Holotypus [block letters] ♀ Dolerus aericepsellus Heidemaa & Mutanen, 2022 [handwritten in block letters] des. M. Heidemaa” [red]. Good condition: wings spread, left fore and mid legs removed as tissue samples. ZMUO.

Paratypes. Total: 1 ♀, 3 ♂: Finland: Lapponia inarensis, Utsjoki Pulmankijärvi, YKJ grid coordinates: 7761:3539 [69.928°N, 28.031°E], 1♂ (ZMUO.033245) 07.07.2017, leg. M. Mutanen, N. Mutanen, A. Mutanen, SDEI. 1♂ (ZMUO.035740) (Fig. 9A, D–G), same collection data as preceding, but 05.07.2018, ZMUO. Lapin Lääni, Nuorgam 17 km SSE, 69.944°N, 28.041°E, 1♀ (BC ZSM HYM 11450), leg. M. Kraus, ZSM. 1♂ (BC ZSM HYM 11451), same collection data as preceding, ZSM.

The name is an adjective derived from the species name of its closely similar sister species, D. aericeps Thomson, using the Latin diminutive suffix -ellus.

Tenthredo (Dolerus) anthracina Klug, 1818: 302. ♀, ♂. Syntypes. Type locality: Germany, Gartz in Pommern. Lectotype ♂ hereby designated, labelled: “14205” [collection catalog no., printed], “Anthracina Kl.” [Klug’s handwriting], “Kl.” [Klug’s handwriting], “GBIF-GISHym 2308” [printed], “Zool. Mus. Berlin” [printed], “Lectotypus ♀ Tenthredo (Dolerus) anthracina Klug, 1818 M.Heidemaa des. ‘22 [handwritten in block letters]” [red], “Dolerus anthracinus (Klug, 1818) M.Heidemaa det. [printed] not nitens Zadd.” [handwritten in block letters]. ZMHB.

Tenthredo (Dolerus) coracina Klug, 1818. Lectotype ♀ hereby designated, labelled: “14207” [collection catalog no., printed], “Carinth. Kl.” [type locality: Austria, Carinthia, probably Klagenfurt], “Coracina Kl.” [Klug’s handwriting], “GBIF-GISHym 2311” [printed], “Zool. Mus. Berlin” [printed], “Lectotypus ♀ Tenthredo (Dolerus) coracina Klug, 1818 M.Heidemaa des. ‘22” [block letters, red], “Dolerus coracinus (Klug, 1818) M.Heidemaa det. [printed] = D. anthracinus auct.” [handwritten]. ZMHB.

Similarly to the melanic color form of D. yukonensis (see below), which was described as a distinct species, D. scoticus, by Cameron (1881a), the melanic form of D. junci was also described as a distinct species, D. thargitai, by Zombori (regarded as a synonym of D. yukonensis / D. scoticus until now). The melanic color form of D. junci has also been recorded from central Europe (Germany) and southern Europe (Italy, Switzerland: see, e.g. Pesarini 2012), but is not known from Fennoscandia. The males of the melanic forms of D. junci (01.06.1984) and D. yukonensis (24.05.1984) have been collected at the same locality at nearly the same time (Germany: Federsee, Brackenhof, leg. E. Jansen, CEJ). In addition to their different penis valve structure and other characters used in the keys by Benson (1952) and Zhelochovtsev (1994), D. junci and D. yukonensis differ in the shape and sculpture of the mesoscutellar appendage: almost unsculptured and without a distinct longitudinal keel in the former, and clearly sculptured and with a distinct keel in the latter. Most likely, the melanic color form of D. junci does not occur within the geographic range of D. incisus, but they can also be distinguished by the shape and the sculpture of the mesoscutellar appendage: meshed (nearly matt) and long, with a distinct median longitudinal keel (D. incisus) versus short and almost unsculptured (more or less glossy) and without a distinct median keel (D. junci).

Dosytheus junci Stephens, 1835. Lectotype ♀ hereby designated, labelled: “Type H.T. [holotype]” [round with wide red margin, printed; not a holotype but a syntype!], “B.M. TYPE HYM. [printed] 1.575.”, “B.M. TYPE HYM. [printed] Dosytheus Junci Stephens 1835.”, “Junci.”, “Stephens Coll. 53–46” [printed], “Dolerus palustris, Klug.” [printed], “Lectotypus 2012 Dosytheus junci [printed]♀ [symbol handwritten] Stephens, 1835 M.Heidemaa des. [printed]”, “Dolerus junci Stephens, 1835 M.Heidemaa det.” [printed]. Condition: legs partly destroyed, left midleg missing, the apical flagellomeres missing, apex of left forewing broken. BMNH.

The original description of D. junci by Stephens (1835) refers to syntypes (“Devonshire and near Windsor”). Although Kirby (1882) mentioned a “Type of D. junci”, this cannot be regarded as a lectotype designation, even though he was apparently only able to locate a single specimen.

Dolerus busaei Snellen van Vollenhoven, 1858. Photos of the lectotype and of the penis valve were checked. Left midleg and two segments of the right antenna missing; the genital capsule is glued on a card. RMNH. This nominal taxon was previously treated as a synonym of D. cothurnatus auct.

Dolerus cothurnatus auct. nec Serville, 1823. Haris and later Lacourt (2000) studied the syntypes (3 ♂) of D. cothurnatus Serville, but not their genitalia. Based on their penis valve structure, none of the syntypes fits D. cothurnatus auct.: the two syntypes (among them the one labelled as lectotype) represent D. germanicus and the third one is a male of D. aericeps. The name D. cothurnatus by Serville is placed here as a junior synonym of D. germanicus, by designating its lectotype (see also under D. germanicus), and the name Dolerus junci (Stephens, 1835) has to be used for D. cothurnatus auct. nec Serville.

Dolerus thargitai Zombori, 1994. The holotype and some paratypes (2♀, 4♂) were studied. HNHM.

Dolerus fumosus Stephens, 1835. Lectotype ♀ hereby designated, labelled: “Type H.T.” [round with wide red margin, printed], “B.M. TYPE HYM. [printed] 1.572.”, “B.M. TYPE HYM. [printed] Dolerus fumosus (Stephens 1835)”, “fumosus.”, “Stephens Coll. 53–46” [printed], “Lectotypus 2012 Dolerus fumosus ♀ [the symbol handwritten] Stephens, 1835 M.Heidemaa des.” [red, printed], “Dolerus sanguinicollis Klug M.Heidemaa det.” [printed] [Identified before it became clear that two species were mixed up under this name.] Condition poor: mid- and hind tarsi incomplete, 2 apical flagellomeres of the right antenna missing, abdomen partly damaged. BMNH.

Kirby (1882) referred to the “Types (♂, ♀)”. The number of syntypes of D. fumosus was not given by Stephens, but only one syntype female is now present in the collection (BMNH).

Dolerus lucens André, 1880. Lectotype ♂ hereby designated, labelled: “Jászó” [Type locality: Hungary, Jászó (Jasov)], “272”, “Hongrie” [printed in block letters], “D. lucens, André.” [printed], “DEI GISHym 89371” [printed], “coll. Desbrochers [handwritten] k. [kollektsija = coll.] A. Jakovleva [nomin. A. Jakovlev]” [printed], “SYNTYPUS [printed in red] Dolerus lucens ANDRÉ, 1880 [handwritten] teste A. Taeger 20[printed]18 [handwritten]” [white label with a red frame], “Lectotypus [printed] Dolerus lucens André, 1880 [handwritten] M.Heidemaa des. [printed]” [red label], “Dolerus fumosus Stephens, 1835 M.Heidemaa det.” [white, printed], ZIN. Paralectotype: 1♂ with same original collecting labels as the lectotype, but the no. is “271” and “Dolerus lucens André (typ.)” [handwritten, probably by Ed. André]. Left forewing missing. HNHM.

Figure 11. 

Dolerus fumosus Stephens, 1835 and D. sanguinicollis (Klug, 1818) habitus. A, B. Dolerus fumosus ♀ (ZMUO.039144); C, D. D. fumosus ♂ (ZMUO.045650); E, F. D. sanguinicollis ♀ (DEI-GISHym14001).

The paralectotype of D. lucens mentioned above bears a lectotype label by L. Zombori (“Lectotypus ♂ Dolerus lucens […] des. Zombori 1980”), but Zombori never published a lectotype designation. Although the paralectotype male has a handwritten label, most probably by André (“Dolerus lucens André (typ.)”, Fig. 12I) and its penis valve is dissected, another syntype male (the right forewing and flagellum missing) is selected as the lectotype because it is deposited in André’s collection (in ZIN), with many other type specimens of species described by him.

Figure 12. 

Dolerus fumosus Stephens, 1835 and D. sanguinicollis (Klug, 1818). A. D. fumosus head dorsal; B. D. sanguinicollis head dorsal; C. D. fumosus ovipositor; D. D. sanguinicollis ovipositor; E. D. fumosus penis valve; F. D. sanguinicollis penis valve. G. D. fumosus katepimeron (arrow); H. D. sanguinicollis katepimeron (arrow); I. Dolerus lucens André, paralectotype ♂, label probably written by André.

The holotype female of D. sanguinicollis var. reicherti has reddish lateral mesoscutal lobes and median mesoscutal lobes only slightly reddish on the anterior part, but according to the other diagnostic characters mentioned above, it belongs to D. fumosus.

Tenthredo (Dolerus) sanguinicollis Klug, 1818: 305. ♀ [not explicitly stated, but indicated by colour characters]. Syntypes. Published type locality: Austria. Lectotype ♀ hereby designated, labelled: “14222” [printed], “Austr. Kl.” [Type locality: Austria, Carinthia, Klagenfurt], “sanguinicollis Kl.”, “Lectotypus” [printed in block letters] ♀ 2014 [handwritten] Tenthredo (Dolerus) sanguinicollis Klug, 1818 [handwritten], “Des. M.Heidemaa” [red label], “GBIF-GISHym 2343”, “Zool. Mus. Berlin” [printed], “Dolerus sangunicollis (Klug, 1818) M.Heidemaa det.”. Minor damage. 2♀ (same collecting data) labelled as paralectotypes (GBIF-GISHym: 2344, 2345). All in ZMHB.

Tenthredo pratensis Linnaeus, 1758. Lectotype. LSUK. Images of the lectotype (LINN 2501) in dorsal and lateral view and the labels are available (The Linnean Society of London 2022), but we did not examine the specimen itself.

Figure 13. 

Dolerus pratensis (Linnaeus, 1758) ♀ (ZMUO.045279). A. Dorsal habitus; B. Lateral habitus. Dolerus timidus (Klug, 1818) ♀ (ZMUO.032881); C. Dorsal habitus; D. Lateral habitus.

Tenthredo (Dolerus) deserta Klug, 1818. Holotype (GBIF-GISHym 2316; Fig. 15). Good condition, genitalia dissected and pinned with the specimen. ZMHB.

Figure 14. 

Dolerus timidus (Klug, 1818), D. variator Enslin, 1927 and D. pratensis (Linnaeus, 1758). A. Penis valve, lectotype Tenthredo timida; B. Penis valve, D. variator; C. Metepimeron (arrow), D. pratensis (ZMUO.032881); D. Metepimeron (arrow), D. timidus (ZMUO.029409), mirrored; E. Lancet, D. timidus; F. Lancet, D. pratensis.

Figure 15. 

Dolerus pratensis (Linnaeus, 1758), holotype ♂ Tenthredo deserta Klug, 1818. A. Penis valve; B. Dorsal habitus.

Tenthredo (Dolerus) timida Klug, 1818. Lectotype ♂ hereby designated, labelled: “14199” [printed catalog no.], “Var. T. timida Kl. [Klug’s handwriting]”, “GBIF-GISHym 2317”, “Zool. Mus. Berlin” [printed], “Lectotypus [printed] ♂ Tenthredo (Dolerus) timida Klug, 1818 [handwritten in block letters] M.Heidemaa design. [printed]” [red], “Dolerus timidus (Klug, 1818) M.Heidemaa det.” [printed]. Good condition, penis valve dissected, pinned with the specimen (Fig. 14A). Paralectotypes: 4♂ 5♀ (GBIF-GISHym: 2312, 2318, 2320–2326). All in ZMHB.

Tenthredo (Dolerus) dubia Klug, 1818. Lectotype ♂ hereby designated, labelled: “14198” [catalog no.], “Dolerus pratensis (L.) ♂ [handwritten] det. F. Koch [printed]”, “GBIF-GISHym 2315” [printed], “Zool. Mus. Berlin” [printed], “Lectotypus [printed] ♂ Tenthredo (Dolerus) dubia Klug, 1818 [handwritten in block letters] M.Heidemaa design. [printed]” [red], “Dolerus timidus (Klug, 1818) M.Heidemaa det.” [printed], Condition satisfactory, genitalia dissected, penis valves on a slide pinned with the specimen (they fit D. timidus). Paralectotypes: 1♂, 1♀ (“GBIF-GISHym 2313”, “… 2314”). All in ZMHB.

Dolerus dubius var. atratus Enslin, 1913. Lectotype ♂ hereby designated, labelled: “Helvet.[ia]”, “Type” [pink, printed], “Dolerus ♂ dubius var atratus [handwritten] Dr. Enslin det. [printed]”, “Sammlung Dr. Enslin” [printed], “GBIF-GISHym 3102”, “LECTOTYPE ♂ Dolerus dubius var. atratus Enslin, 1913 designated M. Heidemaa 2022” [red, printed]. Some damage: left flagellum missing, abdomen glued to a card together with the dissected penis valves and pinned with the specimen. ZSM.

This melanic color form was identified as D. timidus based on its penis valve structure and the sculpture of the metepimeron.

Dolerus liogaster Thomson, 1871. Lectotype ♂ hereby designated, labelled: “Lp. in.” [?Lapland], “LECTOTYPUS [printed] Dolerus liogaster ♂ Thomson, 1871 [handwritten in block letters] design. 1999 M. Heidemaa and M. Viitasaari [printed]” [red], “Dolerus liogaster ♂ [handwritten] M.Heidemaa det. [printed]” [white]. Penis valves dissected, pinned with the specimen. Paralectotypes: 2♀, 1♂. All in MZLU.

Figure 16. 

Dolerus varispinus complex. A. Head dorsal D. liogaster ♀; B. Head dorsal D. schmidti ♀; C. D. liogaster ♀ metepimeron; D. D. schmidti ♀ metepimeron (red arrow), katepimeron (blue arrow); E. D. liogaster fore wing SC₁ (arrow); F. D. schmidti fore wing SC₁; G. D. varispinus fore wing SC₁; H. D. liogaster ♀ katepimeron (arrow); I. D. schmidti ♀ katepimeron; J. D. varispinus ♀ katepimeron; K. D. liogaster serrulae; L. D. schmidti serrulae; M. D. varispinus serrulae; N. D. liogaster penis valve; O. D. schmidti penis valve; P. D. varispinus penis valve; Q. D. liogaster ♀ clypeus; R. D. schmidti ♀ clypeus; S. D. varispinus ♂ clypeus.

Dolerus schneideri Kiaer, 1898. Lectotype ♀ hereby designated, labelled: “Tromsö 22.6.77”, “lget expl” [spelling doubtful], “Ex coll. H. Kiaer” [printed], “261” [yellow], “LECTOTYPUS [printed] Dolerus schneideri ♀ Kiaer, 1898 [handwritten in block letters] design. 1999 M. Heidemaa and Viitasaari” [printed (excl. year), red], “Dolerus liogaster ♀M. Heidemaa det.” [printed label]. Ovipositor dissected and pinned with the specimen, right antenna missing. Paralectotypes: 2♀. All in ZMUN.

The synonymy of D. schneideri with D. liogaster was first proposed by Lindqvist (1943). Treated here as a color form of D. liogaster, but still separated in the key, because the melanic color form of D. liogaster also has black legs, and D. schmidti has a color form with a red-marked thorax. Specimens resembling D. schneideri are also known from the mountain areas of Central Europe, e.g. from Switzerland at altitudes of 1500–1700 m (Benson 1961).

Dolerus truncatus Lacourt, 1988. The holotype (CTN) was studied. Its clypeus emargination looks as if it is abnormally developed. The penis valve (Fig. 18) appears to be somewhat distorted, perhaps during preparation, but resembles the valve of D. liogaster. Tentatively, we treat D. truncatus as a junior synonym of D. liogaster.

Dolerus schmidti Konow, 1884. Lectotype ♀ hereby designated, labelled: “Dolerus Schmidti m. Hagen i/W.”, “Typus” [red, printed], “Holotypus” [red, printed], “GBIF-GISHym4231” [printed], “Dolerus 199[printed]7 schmidti KONOW M. Heidemaa” [handwritten], “Eberswalde coll. DEI”, “Coll. Konow”, “LECTOTYPE ♀ Dolerus schmidti Konow, 1884 designated M. Heidemaa 2022” [red, printed]. Ovipositor dissected, gummed to a card and pinned with the specimen. SDEI.

Konow described D. schmidti as a new species only because D. liogaster Thomson was not known to him at the time. Later D. schmidti was treated as a variety of D. liogaster by Konow (1890b, 1898, 1905b) and synonymized with it by Enslin (1913). The lectotype female was wrongly interpreted as a holotype by Oehlke and Wudowenz (1984) and followed by Heidemaa (2004).

Dolerus soniensis Dubois, 1920. Lectotype ♀ hereby designated, labelled: “La Cambre Mai”, “Collection Dubois”, “D. sonienis ♀ det [handwritten] Ed. Dubois 19 [printed]”, “Dolerus soniensis Dub ♀ [handwritten] Don et dét. [a gift and the determination by] Ed. Dubois 1920 [printed]”, “Type [printed with red ink]” [a pale label with a red frame], “Dolerus 2000 [printed] schmidti KONOW [handwritten] M.Heidemaa det. [printed]”, “Lectotypus ♀Dolerus soniensis Dubois, 1920 M.Heidemaa des.” [printed white label with a red frame]. Ovipositor dissected and glued on a piece of card, pinned with the specimen. RBINS.

Dolerus varispinus Hartig, 1837. Lectotype examined. ZSM.

Dolerus rugosus Konow, 1884. Lectotype ♂ hereby designated, labelled: “Dolerus rugosus m [mihi]. fbg. 4.84” (Fürstenberg / Mecklenburg, IV.1884) [with a double black frame], “rugosus D. T. [Dalla Torre]”, “Syntypus” [red, printed]; “Lectotypus [printed] ♂ Dolerus rugosus Konow, 1884, design. M. Heidemaa and Viitasaari [printed] 1999 [handwritten]” [red label], “GBIF-GISHym 3770” [printed], “Coll. Konow” [printed], “Eberswalde coll. DEI” [printed], “Dolerus 1997 [printed] varispinus Hartig [handwritten] M. Heidemaa det. [printed]”. SDEI. The taxonomic identity of the lectotype is certain despite its poor condition, because the katepimeron of the mesopleuron is preserved.

Tenthredo tristis Fabricius, 1804. Lectotype ♂ hereby designated, labelled: “tristis”, “LECTOTYPE Tenthredo tristis Fabricius, 1775 [sic!] des. SMBlank 2008” [red, printed], “Dolerus bimaculatus (Geoff.) [handwritten] det. S.M.Blank 2008 ✓” [printed], “Dolerus bimaculatus (Geoffroy) M.Heidemaa det.” [printed], “ZMUC-GISHym 1067”. Left flagellum, right flagellomeres 6–7, left fore tarsomeres 4–5 missing. ZMUC.

Contrary to Fabricius’ original description, the posterior half of tergum 2 is red in addition to terga 3–4.

The application of the name Tenthredo bimaculata to this particular Dolerus species remains uncertain until the type material (syntype(s) in coll. E. L. Geoffroy, MNHN or Museum d’Historié Naturelle d’Autun, France?) should be located, or a neotype designated. The diagnosis by Geoffroy (1785) for Tenthredo bimaculata is uninformative: “deux taches blanches au corcelet”, but based on the description by Geoffroy (1762) it was regarded as a possible senior synonym of D. tristis (F.) by de Dalla Torre (1894). Konow (1897a, 1905b) subsequently affirmed this synonymy.

Dolerus coruscans Konow, 1890. Fürstenberg/Mecklenburg [Germany]. Lectotype ♂ hereby designated, labelled: “Dolerus varispinus Htg fbg. [Fürstenberg] 4.84.” [Type locality: Germany, Brandenburg, Fürstenberg], “Dolerus coruscans Knw.”, “Type” [red, printed], “Syntypus” [red, printed], “Coll. Konow” [printed], “GBIF-GISHym 3762” [printed], “Lectotypus [printed] ♂ Dolerus coruscans Konow, 1884 M.Heidemaa design. ‘22”, “Dolerus [printed] ♂ coruscans Konow, 1890 [handwritten in block letters] M.Heidemaa det. [printed]”. Antennomeres 5–9 missing, genitalia dissected, pinned with the specimen. SDEI. Paralectotype: 1♀ (GBIF-GISHym 3761), Germany, Schönberg [probably Brandenburg]. Here determined as D. anthracinus. SDEI.

Oehlke and Wudowenz (1984) correctly linked D. coruscans Konow, 1890 to the description of D. varispinus Hartig sensu Konow, 1884, but following the definitions used in the Code (ICZN 1999), they were wrong in categorizing it as a nomen novum proposed as a replacement for a previously existing name. Dolerus coruscans was, in fact, described as a new species, by indication on the description by Konow (1884). Konow (1890a) altered his opinion on his 1884 identification of D. varispinus, and stated that this taxon represented D. anthracinus sensu Thomson. Goulet (1986) followed Konow’s 1890 opinion, but this cannot be accepted, because Konow clearly based his 1884 description on material in his own collection, not in Thomson’s collection. Although D. coruscans was later synonymized (Enslin 1909) with D. nitens Zaddach, it has been used in published works more than once as a valid name since 1899 (e.g. Dittrich 1905; Fedchenko 1905; Dalglish 1914) and cannot be treated as a nomen oblitum. However, it was listed as a synonym of D. anthracinus by Liston (1981), while Konow (1890b, 1905b) and Cameron (1893) regarded it as possibly conspecific with D. possilensis Cameron. A recent study of the syntypes of D. coruscans (♀, ♂) revealed that the penis valve of the lectotype male (Fig. 17A) is identical to D. nigrominutus Haris, 1998 (the holotype and two paratype males studied) but the penis valve drawing by Haris (1998) was apparently based on a deformed valve because the other valves examined (including those of the holotype) have no such distinct depression on the ventroapical margin near the valvispina. The paralectotype female of D. coruscans (see above) is probably a female (the only one known at present) of D. anthracinus Klug (nec auct.).

Figure 17. 

A–H Dolerus coruscans Konow, 1890; A. Lectotype ♂, penis valve; B. ♀ head frontal (Austria, Burgenland, SDEI); C. ♀ head dorsal (Hungary, HNHM); D. ♀ thorax lateral (Hungary, HNHM); E. ♀ thorax and base of abdomen (Hungary, HNHM); F. ♀ apex of abdomen ventral (Austria, Burgenland, SDEI); G. ♀ valvula 3 dorsal (Hungary, HNHM); H. ♀ valvula 3 lateral (Hungary, HNHM); I. Dolerus picipes (Klug, 1818). ♀ valvula 3 dorsal (ZMUO.045856).

Figure 18. 

Dolerus truncatus Lacourt, 1988, holotype (? = D. liogaster). Penis valve.

The female of D. coruscans, based on two specimens, resembles D. picipes in many characters (the males are even more similar) and the melanic form of D. liogaster with black legs, but differs by its mostly smaller body size of 6–7 mm and the rather inconspicuous setation of the ovipositor sheath (valvula 3) (Fig. 17G). Due to their similarity, the diagnostic characters of coruscans are compared to their states in picipes.

Head. (Fig. 17B, C) Closely resembles D. picipes, but the minimal distance between the antennal sockets is about 2× the length of the malar space (clearly shorter in picipes). Pits on face and vertex more unevenly distributed and vary more in their size. Postocellar field less distinctly defined than in picipes and lateral postocellar furrows less distinct: short and pit-like, rather than long and line-like in picipes. Glossy patches beside the lateral postocellar furrows less distinct (almost no difference between the males).

Thorax. (Fig. 17D, E) The katepimeron of the mesopleuron (Fig. 17D) with more numerous and distinct pits than in picipes. The lateral mesoscutal lobes (Fig. 17E) less distinctly and rather sparsely pitted.

Abdomen. (Fig. 17F–H) The ovipositor sheath in dorsal view broadens slightly towards the apex (Fig. 17G). Cerci (Fig. 17H) yellowish (black in picipes, Fig. 17I). Setae on valvula 3 rather short and delicate compared to most other Poodolerus species (Fig. 17G, H). Setae on the abdominal terga shorter and sparser, except on the apical terga. The species can be easily differentiated from D. picipes as follows.

Dolerus coruscans is recorded from Austria (N-Burgenland, Ungerberg, Weiden am See, 1♀, 24.03.1957, leg. F. Kasy, SDEI), Germany (lectotype: see above), Hungary (“Budapest”, 1♀, SDEI. Haris 1998, 2003: “Budakeszi, Csepel, Látrány, Rákospalota”; Roller and Haris 2008: “Simontornya, Csévharaszt, Szécsény”, in total 14♂), and Greece (Nom. Grevená, Kónitsa E 28 km Vasilítsa Ski Resort, 40.050°N, 21.050°E, 1780 m, 2♂, 11.05.2007, leg. S. M. Blank et al. / leg. M. Wei, SDEI. The records of D. coruscans from Austria and Greece are the first for these countries. The species apparently has a wide distribution, but is probably seldom collected because of its rarity and early flight period. Probably it has also sometimes remained unidentified, or has been misidentified. Possibly, the males of this species in collections could have been placed under D. picipes and its rather small females have perhaps been mistaken for dwarf females of D. liogaster / D. puncticollis (both species also have melanic color forms).

Haris (2003) speculated that the female of this species might be wingless, or have reduced wings. Present data exclude this possibility. Nor does the female remain unknown, as Haris (1998) suggested it would, because of the destruction of the very small type locality of D. nigrominutus. In Central Europe D. coruscans has been collected early in the season (March–April).

Dolerus ferrugatus Serville, 1823. Lectotype ♀ hereby designated, labelled: “Saint-Fargeau” [round], “Lectotype” [red, printed in block letters], “Lectotype D. ferrugatus LEP. ♀ det. A.Haris 1996”, “D. ferrugatus LEP. ♀ det.: A.Haris”, “Lectotypus [printed]♀ Dolerus ferrugatus Serville, 1823 [handwritten in block letters] M.Heidemaa design. [printed]” [red], “Dolerus ferrugatus Serville, 1823 M.Heidemaa det.” [printed]. In good condition (the right flagellum missing). MNHN.

Dolerus brevicornis Thomson, 1871. Lectotype ♀ hereby designated (type locality: Sweden, Öland), labelled: “Ö.” [Öland], “MZLU 2013 414” [green, printed (a loan record label)], “Lectotypus [printed] ♀ 2014 Dolerus brevicornis Thomson, 1871 [handwritten in block letters] M.Heidemaa des. [printed]” [red], “Dolerus ferrugatus Serville, 1823 M.Heidemaa det.” [printed]. 2♀ (from Öland and Scania), 1♂ (the melanic form, from Lund) labelled as paralectotypes. All in MZLU.

Dolerus thomsoni var. miricolor Konow, 1887. Lectotype ♂ hereby designated, labelled: “f. [Fürstenberg] 12/4. [18]84.”, “Dolerus v. [var.] miricolor Knw. [Konow] fbg. 12/4. 84. Typus” [with double black frame], “Dolerus ferrugatus Lep. f. miricolor Knw. Typus. ♂ det. OConde [Otto Conde] 1943”, “Typus” [red, printed], “Holotypus” [red, printed], “Coll. Konow” [printed], “GBIF-GISHym 4215” [printed], “Lectotypus [printed] ♂ Dolerus thomsoni var. miricolor [handwritten in block letters] M.Heidemaa des. [printed]” [red], “Dolerus ferrugatus Serville, 1823 M.Heidemaa det.” [printed]. Right flagellum missing, penis valves dissected (a microscope slide: Symphyta coll. Nr: 159). All in SDEI.

The melanic colour form of D. ferrugatus, described by Konow as D. thomsoni var. miricolor, has sometimes been mixed up with D. pachycerus Hartig, 1837. Apart from their different penis valves, they can be separated by the shape and the structure of the mesoscutellar appendage: long, concave at the sides, and with a distinct longitudinal keel in D. pachycerus compared with D. ferrugatus.

Tenthredo germanica Fabricius, 1775. Lectotype ♀ hereby designated, labelled: “germanica”, “LECTOTYPE Tenthredo germanica Fabricius, 1775 des. SMBlank 2008”, [red, printed], “ZMUC-GISHym 1050”, “Dolerus germanicus (F.) [handwritten] det. S.M.Blank 2008 ✓” [printed], “Dolerus germanicus (Fabricius) M.Heidemaa det.” [printed], “ZMUC-GISHym 1043”. Left flagellum, right fore tarsus, some additional distal tarsomeres missing. ZMUC.

Two specimens are present in the ZMUC collection under the name Tenthredo germanica. Klug (1819) referred to one of them, which has black legs and an infuscated tip of the abdomen (“Das vorhandene Exemplar hatte schwarze Beine und einen an der Spitze schwärzlichen Hinterleib, [...]”). This specimen is selected as the lectotype. It corresponds with the current concept of Dolerus germanicus, e.g. in the shape of the sawsheath (valvula 3) setation. The species needs a thorough revision, because DNA barcoding divides the specimens into two BINs (BOLD:AAI9736 and BOLD:ABV8027) separated by a minimum of over 5% divergence, indicating the possible presence of two distinct species. The two BINs are associated with color differences, of which some appear stable within the cluster and others are variable. Based on about 80 barcoded specimens from Finland, specimens of BOLD:ABV8027 consistently have entirely black hind legs, while those of BOLD:AAI9736 usually have largely orange hind femora and tibiae, although sometimes they are completely black. Additionally, females with a red mesoscutellum always fall in BOLD:AAI9736 and always have red tegulae, pronotum and mesoscutum, whereas in BOLD:ABV8027 the mesoscutellum is always entirely or mostly black, but the tegulae, pronotum and mesoscutum vary from black to red. It is also worth mentioning that the division of specimens into two groups was not reflected in two nuclear genes (POL2 and NaK). Likewise, no useful structural characters have yet been found, but a more detailed systematic study of the male genitalia might be helpful.

Dolerus arcticus Thomson, 1871. Lectotype ♀ hereby designated, labelled: “Lapp” [Lapland] [printed], “arcticus m[mihi]” [Thomson’s handwriting], “MZLU 2013 421” [green, printed loan label], “Lectotypus [printed] ♀ 2014 Dolerus arcticus Thomson, 1871 [handwritten in block letters] M.Heidemaa des. [printed]” [red], “Dolerus germanicus (Fabricius) M.Heidemaa det.” [printed]. Ovipositor dissected, glued on a piece of paper pinned below the specimen. Paralectotypes: 3♀ [MZLU 422, 424, 425], and 1♂ [MZLU 423]. All in MZLU.

Dolerus cothurnatus Serville, 1823. Lectotype ♂ hereby designated, labelled: [green, round, without text], “Lectotype” [red, printed in block letters], “D. cothurnatus LEP. Det.: A.Haris”, “Lectotype D. cothurnatus LEP. Det.: A.Haris”, “Lectotypus [printed] ♀[handwritten], 2014 Dolerus cothurnatus Serville, 1823 [handwritten in block letters] M.Heidemaa des. [printed]” [red label], “Dolerus germanicus (Fabricius, 1775) M.Heidemaa det.” [printed]. Two paralectotype males were also examined: one of them is D. germanicus, the other is D. aericeps. All in MNHN.

A study of the penis valves of the three syntypes of D. cothurnatus revealed that two of them belong to D. germanicus, including the specimen here designated as the lectotype, and one to D. aericeps. These old specimens all have rather pale wings, but probably they have faded: D. germanicus and D. junci [=cothurnatus auct.] typically have darker wings. The specimen labelled by Haris as lectotype is designated here as a lectotype for D. cothurnatus Serville and D. cothurnatus is regarded as a synonym of D. germanicus (see above, under Dolerus cothurnatus auct. nec Serville, 1823).

Dosytheus fuscipennis Stephens, 1835. Lectotype ♀. Labelled: “Type H.T. [holotype]” [round with a red margin, printed], “B.M. TYPE HYM. [printed] 1.578.”, “B.M. TYPE HYM. [printed] Dosytheus fuscipennis Stephens 1835.”, “fuscipennis.”, “Dolerus pratensis, L.” [printed], “Stephens Coll. 53–46” [printed], “Dolerus germanicus (Fabricius) M.Heidemaa det.” [printed]. Condition good, legs partly destroyed. BMNH.

The number of specimens was not given by Stephens. The species description should therefore be assumed to have been based on syntypes. However, Kirby (1882) referred to the “♀. (Type of D. fuscipennis.) S. Scotland. J. F. Stephens”, and according to the labels and the collection catalogue this was the only specimen from the Stephens collection in the BMNH. The specimen fits the original description, and Kirby’s statement qualifies as a valid lectotype designation, because there is no evidence in the original description that more than one type specimen existed (ICZN 1999 Article 74.6.).

Dosytheus hyalinalis Stephens, 1835. Lectotype ♀ hereby designated, labelled: “B.M. TYPE HYM [printed] 1.577.”, “B.M. TYPE HYM [printed] Dosytheus hyalinalis. (Stephens 1835)”, “hyalinalis Steph”, “42 1b” [round], “Dolerus pratensis, L.” [printed], “Dolerus germanicus (F.) M.Heidemaa det.” [printed]. BMNH.

In the original description Stephens indicates that he had more than one specimen, because he gives color characters for both sexes. Kirby’s statement “♀ Type of D. hyalinalis” (Kirby 1882) refers to the only syntype from the Stephens collection that was present in the BMNH, but cannot be accepted as a lectotype designation, because it is clear that Stephens based his description on more than one specimen (ICZN 1999 Article 74.5.). The synonymy with D. pratensis, based on a misidentification, was adopted for example by de Dalla Torre (1894) and most other subsequent works.

Dosytheus xanthopus Stephens, 1835. The only specimen found in the collection (BMNH) labelled as “xanthopus” and “B.M. TYPE 1.257.” belongs to D. germanicus. It cannot be a syntype of D. xanthopus, because the specimen disagrees with the original description: its mesoscutellum is orange, not black as described, and the locality label (“Kent, Darenth, J. F. Stephens BM 1853 – 42”, probably added by Benson) does not match the locality given by Stephens. Interestingly, it was labelled by Benson (handwritten): “Dolerus etruscus?”.

Dolerus Gessneri [sic!] André, 1880: 273. Sex not given. Syntypes. Type locality: Switzerland. Lectotype ♀ hereby designated, labelled: “Suisse”, “DEI GISHym 89367” [printed], “coll. Desbrochers [handwritten] k. [kollektsija = coll.] A. Jakovleva [nomin. A. Jakovlev]” [printed], “SYNTYPUS [printed in red] Dolerus gessneri ANDRÉ, 1880 [handwritten] teste A. Taeger 20[printed]18 [handwritten]” [white with a red frame]. “Lectotype Dolerus gessneri André 1880 vide A. Taeger 2022 des.” [red, printed]. Good condition, but slightly dusty. Paralectotypes: 2♂ (one without flagella) with the same data (except “♂” [printed], “DEI-GISHym 89365” and “D… 89366” and “Paralectotypus” instead of “Lectotypus”). All in ZIN.

The color form of Dolerus gessneri with a red-banded abdomen was described as a distinct species, D. labiosus Konow, 1897 (Konow 1897b), but was later mostly treated as a synonym (sometimes also as a subspecies) of D. gessneri, because intermediate color forms with variable red markings on the abdomen occur. The specimens from the Russian Far East with completely black forelegs, which have sometimes been determined as D. gessneri, probably belong to some other Dolerus (subgen. Equidolerus) species. Dolerus gessneri, a Holarctic species, needs a revision, because COI barcoding data suggest that it could include more than one species, and some morphological characters correlate with the genetic data. There are three BIN clusters (BOLD:AAL2317, BOLD:ACE7304, BOLD:ACE3617) diverging by 1.8–2.3%. BOLD:AAL2317 is closest to D. timidus and BOLD:ACE7304 is closest to D. pratensis (minimum divergence in both cases 1.2%). All three BIN clusters are represented in Fennoscandia, two of which (Fig. 19) have been found in Finland.

Figure 19. 

Dolerus gessneri (André, 1880) (ZMUO.045118). A. Dorsal habitus; B. Lateral habitus. Dolerus sp. near gessneri (ZMUO.045279); C. Dorsal habitus; D. Lateral habitus.

Tenthredo gonagra Fabricius, 1781. Lectotype ♀ hereby designated, labelled: “gonagra”; “LECTOTYPE Tenthredo gonagra Fabricius, 1781 des. SMBlank 2008” [red, printed: paralectotypes labelled in corresponding way], “ZMUC-GISHym 1050”, “Dolerus gonager (F.) [handwritten] det. S.M.Blank 2008 ✓” [printed], “Dolerus gonager (Fabricius, 1781) M.Heidemaa det.” [printed]. Right flagellomeres 6–7, right hind tarsus missing. Paralectotype 1♀: “gonagra”, “ZMUC-GISHym 1051”. ZMUC.

The type specimens correspond with Zhelochovtsev’s definition of Dolerus (Poodolerus) gonager, which has red knees, superficially pitted lateral mesonotal lobes and backwards directed setae of the parallel-sided ovipositor sheath (similar to Zhelochovtsev 1994: fig. 165.2, but with some of the longest hairs curved distally).

Dolerus femoratus Eversmann, 1847. Lectotype ♀ hereby designated, labelled: “Kas[an] 16.V.”, “femoratus. Kl.” [with black frames], “DEI-GISHym 30068” [white, printed], “Lectotype Dolerus femoratus Eversmann 1847 vide A. Taeger 2014 des.” [red, printed]. In good condition. Belongs to D. gonager. Paralectotypes: 2♀ (DEI-GISHym 30069, 30070), are D. puncticollis Thomson, 1871. All in ZIN.

Dolerus magnicornis Eversmann, 1847. Lectotype ♂ hereby designated, labelled: “Orb” [Orenburg], “magnicornis mihi” [with black frames], “DEI-GISHym 30067” [white, printed], “Syntype Dolerus magnicornis Eversmann 1847 vide A. Taeger 2014” [red, printed], Lectotype Dolerus magnicornis Eversmann 1847 M.Heidemaa des.” [red, printed]. In good condition. Belongs to D. gonager. ZIN.

Finland: Regio kuusamoensis, Kuusamo Siikauopaja, YKJ grid coordinates: 7362:3612 [66.334°N, 29.511°E], 2♀ (ZMUO.035155, ZMUO.035156), 12.06.2018. 1♀ (ZMUO.045265), 20.06.2020. All specimens leg. M. Mutanen. ZMUO.

First record in the Palaearctic. Recorded previously only in boreal North America: Alaska, Alberta, Yukon, and North West Territories (Goulet 1986). The genetic and morphological affinity of D. incisus to D. junci and D. yukonensis suggests that the host plant of D. incisus is also a species of Equisetum (Goulet 1986). The habitat in Finland is a shallow oxbow lake (meander) of the Oulanka river with plenty of Equisetum fluviatile, which is also the host of D. junci. This habitat matches perfectly with that given for D. incisus in North America by Goulet (1986). Several other Dolerus species occur syntopically, including the closely related D. junci Stephens (= D. cothurnatus auct., see above). The melanic form of D. incisus is recorded from Finland (all three are females, see above, Fig. 20A) and Norway (Heimdalsmunnen, alt 1150 m, leg. E. Heibo, CEH). Melanic females of D. subarcticus (recorded from Norway and Kamtschatka, see under D. subarcticus) can be distinguished by their postocellar furrows, the sawsheath (valvula 3) shape, and its apical setation (Fig. 20B). Melanic males can be separated most reliably by their penis valves (Fig. 20C, 20D). Only the melanic colour form of D. incisus has so far been recognized from Fennoscandian countries, whereas in Canada and the USA both the black color form and a form with red-banded abdomen and partly reddish legs are known. We studied paratypes of D. incisus: 2♀, 2♂, some of them dissected, CNC.

Figure 20. 

Dolerus incisus Goulet, 1986. A. Dorsal habitus ♀; B. Valvula 3 dorsal; C. Penis valve; D. Dolerus subarcticus Hellén, 1956, penis valve.

Dolerus possilensis Cameron, 1882a: 178–179. ♀. Syntypes (assumed). Type locality: Possil Marsh near Glasgow, Scotland. Lectotype ♀ hereby designated, labelled: “B.M. TYPE HYM. 1.573” [printed], “Dolerus possilensis Cam.”, “Cameron 96–76. [printed] Possil” [handwritten], “B.M. TYPE HYM. [printed] Dolerus possilensis (Cameron 1882)”, “Lectotypus [printed] Dolerus possilensis Cameron, 1882 [handwritten in block letters] M.Heidemaa design. [printed]” [red]. Most of the abdomen missing. Ovipositor preparation apparently lost, but probably figured in Cameron (1882a: plate XIX, fig. 8). BMNH.

Probably a parthenogenetic species: no male has ever been recorded. The only known Fennoscandian record is from Sweden: 1♀ [labelled as Dolerus sp.] “Resarö Waxholm / 6 1915”, leg. R. Malaise, NHRS. New to the Swedish fauna.

Dolerus puncticollis Thomson, 1871: 286. ♀, ♂. Syntypes. Published type locality: [Sweden]. Lectotype ♂ hereby designated, labelled (printed on pale paper if not stated otherwise): “Lund” [type locality Sweden, Lund], “Lectotypus [printed] 1♂ Dolerus puncticollis Thomson, 1871 [handwritten in block letters] M.Heidemaa design. [printed]” [red]. Paralectotypes (3♀, 2♂, Öland, Lund, Scania; the locality labels of 2♀ are illegible). All in MZLU.

Penis valve of the lectotype as in Fig. 21. The taxonomic status of a melanic colour form closely resembling D. puncticollis (recorded from Portugal and Greece) is still under study.

Figure 21. 

Dolerus puncticollis Thomson, 1871. Penis valve, lectotype.

Dolerus subarcticus Hellén, 1956. Lectotype ♀ hereby designated, labelled (printed on pale paper if not stated otherwise): “Fennia, Li. Utsjoki, Outakoski, 29.6.1947 [type locality], leg. Hellén” [printed], “Lectotypus 2014, Dolerus subarcticus, ♀ Hellén, 1956, M.Heidemaa design.” [red], “Dolerus subarcticus Hellén, 1956, M.Heidemaa det.” [white]. In good condition. 10♀, 8♂ paralectotypes also examined: Ivalo, Kantalaks, Kilpisjärvi, Lutto, Muonio, Paanajärvi, Petsamo, Umba, Utsjoki (Outakoski). All in MZH.

Dolerus willoughbyi Benson, 1956. 2♀, 2♂ paratypes studied. BMNH.

The melanic (black) form (♂) and a nearly black form, with obscure reddish patches on abdominal terga 2–4 (♀), are known from Norway (near Hovet and near Geitryggtunnelen, leg. E. Heibo, CEH), and the melanic form also from Kamtschatka, Russia (a syntype female of D. pratensis var. totus determined by R. Malaise, in NHRS, see also under D. yukonensis).

Dolerus vulneratus Mocsáry, 1878: 199. ♀. Syntypes (assumed). Type locality: Siberia. Lectotype ♀ hereby designated, labelled: “Siber[ia]”, “Dolerus vulneratus ♀ Mocs.”, “Holotypus [printed with red] ♀ Dolerus vulneratus Mocsáry, 1878 Zombori, 1977 [handwritten]”, “Lectotypus [printed] ♀ Dolerus vulneratus Mocsáry, 1878 M.Heidemaa des. ‘22” [handwritten, red]. “Dolerus vulneratus Mocsáry, 1878 M.Heidemaa det.” [printed]. HNHM.

The “holotype” label attached by Zombori has no nomenclatural significance, because this interpretation was never published. Moreover, the number of specimens was not given in the original description and the ♀ symbol does not necessarily indicate that Mocsáry had only one specimen, even if only one specimen was found in the author’s collection. Such assumptions about the status of specimens as holotypes should be avoided according to the Code (ICZN 1999: Articles 72.4.7, 73F). The larva is known through rearing and observations by Ponomarev (2022). Host plant: unidentified Poaceae spec.

Dolerus scoticus Cameron, 1881. Lectotype ♀. BMNH.

Dolerus pratensis var. totus Malaise, 1931. Lectotype ♀ hereby designated, labelled: “KAMTSCHATKA Malaise” [printed], “586” [pink, printed], “Typus” [red, printed], “D. pratensis var totus n. sp. Type [handwritten] Malaise det. [printed]”, “Dolerus scoticus ♀ Cam [handwritten] A.Zhelochovtsev det 19[printed] 33 [handwritten]”, “Dolerus germanicus totus Mal.” [printed and folded], “Lectotypus [printed] Dolerus pratensis var. totus Malaise, 1931 M.Heidemaa des. [handwritten in block letters]” [red label], “Dolerus 2009 [printed] yukonensis [handwritten] M.Heidemaa det. [printed]”. NHRS. Paralectotypes: 1♀, “280” [printed], “Dolerus 20 [printed] 11 subarcticus [handwritten] M.Heidemaa det. [printed]” (a melanic female). NHRS. 1♀, “Dolerus v. toto [sic!] n. var. Paratypus ♀ Malaise det. [written by R. Forsius?]”, “Dolerus yukonensis Norton M.Heidemaa det.”, MZAT (coll. Forsius).

Dolerus yukonensis has a Holarctic distribution, and both red-banded/-marked as well as melanic color forms occur. According to Goulet (1986) the color forms developed independently in the coastal areas of both continents. This speaks against the subspecific status of the melanic color forms suggested by Benson (1959). According to Benson (1934a) D. scoticus Cameron can be distinguished from melanic forms of D. yukonensis by the strongly developed temporal furrows, like in D. junci and D. incisus, which are absent in D. yukonensis. He also wrote that “Dolerus totus Malaise, 1931, from Kamtchatka may be the same species as D. scoticus Cam.” Benson (1934b) synonymized them (and D. arcticola Kiaer, 1898) with D. scoticus and later all of them with D. yukonensis (Benson 1958: corrigenda). Latterly, he regarded D. scoticus as a subspecies of D. yukonensis (Benson 1959) and reported it from Switzerland (Benson 1961) and southern Norway, Hallingskarvet District (Benson 1966). However, Benson was not aware of the presence of melanic forms of D. incisus and D. subarcticus in Norway, the males of which can most reliably be identified by their penis valves (Fig. 20C, D). The sculpture of the abdominal terga in D. yukonensis, as in D. subarcticus, is not as stable as is often assumed. The depth of the clypeus emargination and the lateral postocellar furrows also vary. However, no potential male of D. scoticus, which could support its status as a distinct species as proposed by Haris (2000), has so far been recognized. The lectotype female of D. scoticus probably belongs to D. yukonensis, and certainly not to D. junci or D. incisus. Here we retain the synonymy of D. scoticus with D. yukonensis proposed by Benson (1958) and concurred with by Goulet (1986), but admit that more DNA sequence data are needed to test the taxonomic status of the melanic forms, which show at least some coinciding structural differences. We have studied melanic and red-marked color forms from the Palaearctic, but only red-marked forms from the Nearctic.

The species boundaries between willow-feeding taxa of the Empria immersa group (E. immersa, E. camtschatica, E. plana, and E. improba) have proved to be difficult to elucidate (Prous et al. 2014, 2020). In Fennoscandia, two forms can commonly be found at the same time and place (Prous et al. 2014): E. immersa with a dark pterostigma and short antenna, and E. camtschatica with a pale pterostigma and long antenna. These two forms can also be distinguished by larval morphology (Fig. 22). Based on ex ovo rearings by M. Prous (two females from Sweden and Estonia) and ex larva rearings by Ponomarev (2022) of E. immersa, and ex ovo rearings by M. Prous of E. camtschatica (using two females from Sweden), the main difference seems to be in head coloration: E. immersa with an occipital fleck or stripe (Fig. 22F–M) and E. camtschatica with occipital and parietal stripes (Fig. 22A–E). An additional difference may be that glandubae (white conical warts) are more prominent in E. immersa than in E. camtschatica. Although based on limited specimen sampling, genome scale data (Prous et al. 2020) support E. immersa as a distinct species most consistently compared to the other species in the E. immersa group. In Fennoscandia, taxonomy is complicated by the presence of occasional specimens identifiable as E. plana, somewhat intermediate in morphology between E. immersa and E. camtschatica (pterostigma like E. camtschatica, saw intermediate). Genome scale data of one E. plana female from Sweden do not indicate affinity with E. immersa, but do show at least some affinity with E. camtschatica (Sweden), E. improba (Canada), and one other E. plana (Hokkaido, Japan) (see fig. 5 in Prous et al. 2020). Given the above, we synonymize E. plana and E. camtschatica with E. improba, because clear boundaries between these taxa cannot at present be drawn. Thus, in Europe, the specimens with dark pterostigma, short antennae and more prominent serrulae of the saw can be identified as E. immersa, and those with pale pterostigma, usually longer antennae, and less prominent serrulae as E. improba (see Prous et al. 2014). In North America, however, at least some E. improba specimens look externally more like E. immersa (dark pterostigma and metafemur), while the serrulae of the saw resemble E. camtschatica. Lacourt (2020) suggested that E. camtschatica could be a synonym of E. improba, but genetically these taxa are not necessarily closer to each other than they are to E. plana (Prous et al. 2020). If the circumscription of E. improba as proposed here is considered incorrect, then it remains unclear how many additional species should be recognized, and how these should be delimited. For example, in Europe the morphological distinction between E. camtschatica and E. plana is not clear, although these forms can be more reliably distinguished from E. immersa.

Figure 22. 

Larvae of Empria on Salix. A–E. E. improba (Cresson, 1880); F–M. E. immersa (Klug, 1818).

COI barcodes of a few specimens identified as E. nigrans are indistinguishable from those of a large number of E. ephippium (red-marked females). Vikberg et al. (2011) discussed the distribution of the two forms in Fennoscandia and more widely in Europe, noting that the patterns of occurrence are complex, and that they are regionally often sympatric, but that the dark form becomes more common towards the West, suggesting that climatic causes may play a role in maintaining this polymorphism. No other differences have been detected between the red-marked and black forms. We therefore revert to treating them as conspecific colour forms.

Although we have not studied the lectotype of Lophyrus catocalus, the interpretation of its taxonomic status is possible after referring to Gussakovskij’s description of G. verticalis and the other publications cited above. We agree with Pschorn-Walcher (1982) and Viitasaari and Varama (1987) that the two forms are an expression of individual variability, linked to population differences with a geographical component. Accordingly, we treat Gilpinia catocala as a junior synonym of G. pallida.

Female. Figs 23, 24A–C, 26A, B

Colour. Fig. 26A, B. Black. Pale are: palpi, tegula, legs (except for arioli and extreme inner tip of metatibia); in fore wing base and apex of costa, apex of subcostal, base of stigma (Fig. 23A), R1, 1A, 2A+3A [color differences between veins of hind wing not clearly definable]; margins of median excision of abdominal tergum 1, medio-distal part of terga 2–4 (–6) [pale area on tergum 3 is the largest of these], all sterna, more or less the downturned parts of terga 2–8, more or less hypopygium and ovipositor valvifer 2. Wing membranes hyaline.

Figure 23. 

Heptamelus viitasaarii sp. nov. ♀ (ZMUO.033969). A. Fore wing costa and pterostigma; B. Antenna; C. Head dorsal; D. Lower head frontal; E. Head lateral; F. Thorax dorsal. Mesoscutellar appendage (arrow); G. Abdomen lateral; H. Thorax lateral.

Head. Pedicel about as long as scape. Head in lateral view with widest point on temple about equal to length of pedicel (Fig. 23E). Malar space slightly shorter than diameter of anterior ocellus. Lowest part of gena densely sculptured, without shining interspaces between the ill-defined pits (Fig. 23E). Clypeus densely pitted, weakly shiny; anterior margin widely emarginate, to about 0.4 of its length (Fig. 23D). Setae on upper head as long, or longer than, diameter of anterior ocellus (Fig. 23E).

Thorax. Whole pronotum except for small antero-ventral area dull, with sculpture (Fig. 23H). Pits on upper mesepisternum large and well-defined (Fig. 23H). Mesoscutellum with large, scattered pits anteriorly and laterally, interspaces shiny; medially and posteriorly nearly without pits and entirely unsculptured (Fig. 23F). Anterior of mesoscutellar appendage densely pitted, with numerous setae, only small posterior area unpitted (Fig. 23F).

Abdomen. Anterior of tergum 1 densely setose. Terga 2 and 3 glabrous. Terga increasingly setose from tergum 4 to apex of abdomen. Weak surface sculpture on tergum 2 becoming stronger on more distal terga. Sawsheath in dorsal view broad, with blunt tip (Fig. 24C). In lateral view exposed length of cercus about 0.5× as long as exposed upper length of valvula 3 (Fig. 23G). Lancet (Fig. 24B, C): 13–14 serrulae; apical serrulae short and high.

Figure 24. 

A–D. Heptamelus viitasaarii sp. nov. A. Lancet (DEI-GISHym31999); B. Tip of lancet (DEI-GISHym31999); C. Valvula 3 dorsal (ZMUO.033969); D. Penis valve (ZMUO.058165); E. Heptamelus dahlbomi (Thomson, 1870) (DEI-GISHym83629) penis valve.

Body length : 4.5–8.0 mm

Male. Figs 24D, 26C

Similar to female, except for: red-brown antennal flagellum with basal two antennomeres more or less black; metatibiae completely pale; abdomen completely dark with small, obscure pale markings medially on terga 3–5. Penis valve: Fig. 24D.

Body length : 4.5–5.5 mm

Variability : 5–6 flagellomeres, depending on whether or not the distal one is subdivided; the proportions of this antennomere are thus highly variable. Number of pits on upper mesepisternum variable. The scape and pedicel of females may be more or less pale.

Holotype ♀ (ZMUO.044130). Type locality: Finland, Karelia borealis, Kitee Otravaara, YKJ grid coordinates: 6868:3663 [61.8877N 30.1096E]; 04.08.2019 [larva] in Athyrium filix-femina (L.), Rearing 380/2019, leg. M. Mutanen. Labels white, printed, except for red holotype label. Deposited in the ZMUO.

Paratypes. Total: 92♀, 6♂. Finland [leg. M. Mutanen and in ZMUO unless otherwise stated]:

Karelia borealis, Kitee: Otravaara [Type locality]; 17♀ (including ZMUO.044126, ZMUO.044127, ZMUO.044128, ZMUO.044129, ZMUO.044131, ZMUO.044132, ZMUO.044133, ZMUO.044134, ZMUO.044135, ZMUO.044136), 04.08.2019 [larvae] in Athyrium filix-femina (L.), Rearing 380/2019. 7♀ (including ZMUO.034957, ZMUO.034958, ZMUO.034959), Otravaara, YKJ grid coordinates: 6868:3663 [61.888°N, 30.110°E], 19.06.2018. 30♀ (including ZMUO.058509, ZMUO.058510, ZMUO.058511, ZMUO.058512, ZMUO.058513, ZMUO.058514, ZMUO.058515, ZMUO.058516, ZMUO.058517, ZMUO.058518, ZMUO.058519, ZMUO.058520), Otravaara, YKJ grid coordinates: 6868:3663 [61.888°N, 30.110°E], 24.08.2020, leg. M. Mutanen and M. Prous. Host: Athyrium filix-femina, rearing 174/2020. 1♀ (ZMUO.046037), 15.06.2020. ZMUO and SDEI. Otravaara, YKJ grid coordinates: 6868:3664 [61.887°N, 30.129°E]; 8♀ (including ZMUO.034859, ZMUO.034860), 19.06.2018. 1♀ (ZMUO.034995), 18.06.2018. Pajarinmäki, YKJ grid coordinates: 68899:36670 [62.078°N, 30.197°E]; 1♀ (ZMUO.032885), at light, 11.07.2017 [end date of about 5-day period]. 1♀ (ZMUO.033391), at light, 18.07.2017 [end date of about 7-day period]. 2♀ (including ZMUO.034899), 20.06.2018. 1♂ (ZMUO.061917), 14.06.2021, leg. A. Liston, M. Mutanen, M. Prous. 1♂ (ZMUO.061893), 15.06.2021, leg. N. Kiljunen, A. Liston, M. Mutanen, M. Prous. 4♂ 8 ♀♀ (including ZMUO.060946, ZMUO.060947, ZMUO.060948, ZMUO.060949, ZMUO.060950, ZMUO.060951, ZMUO.060952, ZMUO.060953, ZMUO.060954, ZMUO.060955, ZMUO.058165), 18.06.2021, leg. N. Kiljunen, A. Liston, M. Mutanen, M. Prous. 2 larvae from Matteuccia struthiopteris (L.) (ZMUO.060557, ZMUO.060558) and 2 larvae from Athyrium filix-femina (L.) (ZMUO.060561, ZMUO.060562), 26.07.2021, leg. M. Mutanen, M. Prous. Papinniemi, YKJ grid coordinates: 6883:3656 [62.025°N, 29.990°E]; 6♀ (including ZMUO.034761, ZMUO.034762), 18.06.2018, ZMUO and SDEI. 1♀ (ZMUO.040497), 12.06.2019. Puhos, YKJ grid coordinates: 6889:3653 [62.080°N, 29.938°E]; 2♀ (including ZMUO.034968), 20.06.2018. Hiidensaari, YKJ grid coordinates: 6891:3668 [62.092°N, 30.226°E]; 1♀ (ZMUO.034908), 20.06.2018. Potoskavaara, YKJ grid coordinates: 6893:3670 [62.109°N, 30.266°E]; 1♀ (ZMUO.046430), 28.06.2020, leg. Tupu Vuorinen. Satulavaara, YKJ grid coordinates: 6877:3672 [61.964°N, 30.289°E]; 3♀ (including ZMUO.061792, SDEI-GISHym14061), 14.06.2021, leg. A. Liston, M. Mutanen and M. Prous. ZMUO and SDEI.

Karelia australis: Imatra Kaikkallio, YKJ grid coordinates: 6778:3594 [61.104°N, 28.752°E]; 1♀ (ZMUO.058146), 16.06.2021, leg. A. Liston, M. Mutanen, N. Kiljunen, M. Prous. Joutseno Kuurmanpohja Sunninmäki [61.071°N, 28.731°E]; 1♀ (ZMUO.016815), 06.06.2015, leg. Jussi Vilen, coll. Matti Viitasaari.

Tavastia australis: Kangasala Keisarinharju, YKJ grid coordinates: 6818:3347 [61.444°N, 24.141°E]; 1♀ (ZMUO.061964), 08.06.2021.

Finland: Nylandia, Vantaa, YKJ grid coordinates: 669:37, 17.06.2021, 1♀, leg. Miikka Friman.

Named after Matti Viitasaari, who first recognised a very large female of this species as probably not belonging to Heptamelus ochroleucus or H. dahlbomi.