Aleiodes pseudoseriatus van Achterberg & Shaw, 2024

van Achterberg, Cornelis, Shaw, Mark R., Fernandez-Triana, Jose & Quicke, Donald L. J., 2024, Resolution of the Aleiodes seriatus (Herrich-Schäffer, 1838) - aggregate in the western Palaearctic (Hymenoptera, Braconidae, Rogadinae), with description of a new species, ZooKeys 1208, pp. 241-258 : 241-258

Aleiodes pseudoseriatus van Achterberg & Shaw sp. nov.

Figs 3–4 , 5–17 , 18–19 , 20

Type material.

Holotype, ♀ ( NMS), “ Italy, Veneto, Vittoria Veneto (VT), Frazione di fais , 46.017 N 12.274 E (WGS 48), 450 m, 18. vii. 2016, [at] UV light, D. Dal Pos ”, “ MRS Aleiodes DNA 1064 ”, “ DNA COI worked ” . Paratypes: 1 ♂ ( BZL), Austria: Bad Ischl, OÖ [= Oberösterreich], Höherstein, 820 m, lux SW-wand, Forststrasse, N. 47.686 ° E 13.689 °, 3. vii. 2010, N. Pöll ”; 1 ♂ ( MSC), A [ustria]: Oberösterreich, 13 km SSW Reichraming, Krahlalm, 47 ° 46 ' N, 14 ° 23 ' E 22. vi. 2011, 680– 850 m, M. Schwarz ”; 1 ♂ ( BZL) “ A-OÖ [= Austria: Oberösterreich], Linz-Urfahr, Pragerstrasse, N 48.19. 16 E 14.17. 36 18–19. vii. 2013, Trefenthaler ”; 1 ♂ ( BZL), id., but “ KGA Riesenhof, Parz 60 E 14.16. 15 N 48.19. 06, 5–8. ix. 2013 ”; 1 ♂ ( RMNH), “ Belgium: Liège, Mt. Rigi, 650 m, 1–2. viii. 1986, at light, C. Bank, RMNH ”; 2 ♂ ( RMNH), id., but 2. viii. 1986, C. v. Achterberg; 1 ♀ ( NMS), “ Bulgaria: W Stara Planina Mts, confluence of Penkova and Berkovska rivers, 558 m, N 43.2233 E 023.07596, 10. ix. 2021, S. Beshkov & A. Nahirnić-Beshkova ”; 1 ♂ ( BZL), “ CZ [= Czech Republic]: Bohemia, C. Budéjovice, D. Voda, N 48 ° 58 ’ E 14 ° 32 ’ 470 m, M. Halada 10. vii. 2001 ”; 1 ♀ ( NMS), “ [ England:], Cumbria, Howe, Whitbarrow, [at] MV light, 24. viii. [19] 95, M. R. Shaw ”; 1 ♂ ( NMS), “ Cum [bria], Roudsea Wood, at light, 15. vii. [20] 06, M. R. Shaw ”, “ MRS Aleiodes DNA 455 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ MRS Aleiodes DNA 616 ”, “ DNA CO 1 worked ”; 1 ♂ ( NHMUK), “ England: E. Kent, West Wood, TR 1426143868, MV light, 29. viii. 2011 ”, “ MRS Aleiodes DNA 819 ”, “ DNA CO 1 worked ”; 1 ♀ ( NHMUK), “ England, Cornwall, Ding Dong, Tredinnick Stack, SW 444348 MV light trap, J. Herbert, BMNH (E) 2012-41 ”; 1 ♂ ( NHMUK), “ [England]: Hen Wood, SU 6522 Hants VC 11, 23. vii. 2013 MV ”; 1 ♀ ( NMS), “ Estonia: Piargu, Raplamaa Farmland, [N] 59.122167, [E] 24.831745, 9. ix. 2019 MV light, Kaido Kärner ”, “ MRS Aleiodes DNA 1103 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ MRS Aleiodes DNA 1104 ”, “ DNA CO 1 worked ”; 1 ♀ ( NMS), “ France: Côte d’Or, Abbaye de la Bussière, La Bussière-sur-Ouche, at light, 19. vii. 2003, M. R. Shaw ”, “ MRS Aleiodes DNA 262 ”, “ DNA COI worked ”; 1 ♀ ( NMS), id., but “ MRS Aleiodes DNA 254 ”, “ DNA CO 1 worked ”; 1 ♀ ( NMS), id., but “ MRS Aleiodes DNA 252 ”, “ DNA CO 1 worked ”; 1 ♂ ( RMNH), “ France: Finistère, Forêt du Cranou, 7 km E [of] le Faou, on Taxus , 27. vi. 1988, M. J. Gijswijt ”; 1 ♀ ( RMNH), “ France: Doubs, RN Lac de Remoray, 16. viii. 2009, Mal. tr [ap] 3, 948242 / 6634536, H. Gens, RMNH ’ 23 ”; 1 ♀ ( NMS), “ Finland: Oulu, Ketolanoja, Muhos, Mal. tr. 5–19. viii. [20] 05, N. Laurenne ”; 1 ♀ ( NMS), “ Germany: Bayerswald, 2001, M. Kuhlmann ”, “ MRS Aleiodes DNA 222 ”, “ DNA CO 1 worked ”; 1 ♂ ( RMNH), “ Germany: Thüringen, NP Hainich, nr Eisenach, [reared] from Fagus sylvatica stems, 12. vi- 3. vii. 2008, M. Gossner, RMNH ’ 08 ”; 1 ♀ ( ZSM), “ [Germany]: Ober Bayern, Garmisch, 12–1300 m, 10. viii. 1936, E. Bauer ”; 2 ♀ ( ZSM), “ [Germany]: Ebenhausen, Isart, viii. [19] 40, K. V. Rosen ”; 7 ♂ ( MTMA), “ Hungary, Nógrád m., Bátonyterenye (Kistererenye), Csente, Kertvárosi kert ”, “ 48.0074992 ° / 19.8180737 ° [= 20. viii – 9. ix. 2016], P. G. Sulyán, lámpázás (6) ”; 1 ♂ ( MTMA), id., but “ [= 24. ix – 30. ix. 2016] ... lámpázás (8) ”; 1 ♂ ( MTMA), id., but [= 15. x. 2016] ... lámpázás (9) ”; 1 ♀ ( NMS), “ [ Ireland]: Wexford, 5. vii. [19] 02, J. J. F. X. King ”; 1 ♀ ( NHMUK), “ [Ireland]: Kilkea Deerpark, Co, W [e] x [ford], 4. vi. 1937, A. W. Stelfox ”; 3 ♀ + 2 ♂ ( NMS), “ Italy: Veneto, Riserva Naturale Integrale Bosco Nordio, Chioggia, 45.122 N 12.260 E, 28. vii. 2016, D. Dal Pos ”; 2 ♂ ( NMS), id., but “ 3. vi. 2016 ”; 1 ♂ ( NMS), “ Netherlands: Noord Holland duinreservaat, Egmond aan Zee, MV 8. vii. 2016, M. R. Shaw ”, “ MRS Aleiodes DNA 838 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ MRS Aleiodes DNA 839 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ MRS Aleiodes DNA 840 ”, “ DNA CO 1 worked ”; 2 ♂ ( NMS), id., but no DNA labels 1 ♂ + 1 ♀ ( RMNH), “ Netherlands: Gld, Tongeren, 3. ix. 1991, B. v. Aartsen ”; 1 ♀ ( RMNH), id., but 9. vii. 1989, C. J. Zwakhals; 1 ♀ ( RMNH), “ Netherlands: LI, Brunssum-Treebeek, c. 100 m, 50 ° 56 ' 17 " N, 5 ° 56 ' 58 " E, garden, at light, 25–31. vii. 2018, G. Lommen, RMNH ”; 1 ♂ ( RMNH), id., but 3–10. vi. 2018; 1 ♀ ( RMNH), “ [Netherlands: UT,] 3 bergen [= Driebergen], Six [c. 1860] ”; 1 ♀ ( RMNH), “ [? Netherlands, Hilvarenbeek], H. B., 3. vii ”; 1 ♂ ( RMNH), “ Nederland: Gld, ‘ t Harde, 16. viii. 1993, B. v. Aartsen ”; 1 ♀ ( RMNH), “ Netherlands [: FR], Fochtelo, 4. ix. 2001, B. v. Aartsen; 1 ♀ ( RMNH), “ Netherlands: DR, Borger, Boswachterij Borger, UTM LD, 495693, SBB-vak 26, 25–28. vii. 1993, Mal. tr [ap], L. Witmond ”; 1 ♀ ( RMNH), “ Netherlands: NB, Tilburg, Kaaistoep, at light, 18. vii. 2017, 128.8–394.6, T. Peeters, RMNH ’ 18 ”; 1 ♀ ( RMNH), “ [Netherlands:] Gld, Epe, de Dellen, 19. vii. 1994, B. v. Aartsen ”; 1 ♂ ( RMNH), “ [Netherlands:] OV, Hasselt, Stadsgaten, 24. vii. 1994, B. v. Aartsen ”; 3 ♀ ( RMNH), “ Netherlands: NB, Achtmaal, O. Bluisse Heide, MT, R. D. 97–386, 5. viii. 2015, E. Brosens ”; 1 ♂ ( RMNH), id., but 15. viii. 2015; 1 ♀ ( NMS), “ Norway: RY Hølland, 58.52445 N 5.83518 E, 17. vii – 2. vii. 2020 Mal. tr. A. T. Mjøs ”; 1 ♂ ( NMS), “ Serbia: Kasan, N of Prepollent, 1256 m, 43 ° 19 ' 35 " N, 19 ° 96 ' 44 " E, 3. vii. 2019, C. W. Plant ”, “ MRS Aleiodes DNA 1057 ”, “ DNA COI worked ”; 1 ♀ ( NMS), “ Serbia: Tzaribrod (Dimitrovgrad) distr., Vištni Kamen above Bačevo Village, 763 m, N 43.0271, E 022.8239 11. viii. 2021, S. Beshkov & A. Nahirnić-Beshkova ”, “ MRS Aleiodes DNA 1129 ”, “ DNA CO 1 worked ”; 1 ♀ + 3 ♂ ( NMS), “ Serbia: Suva Planina, Preslap, 1186 m, N 43.19473 E 022.24400, 30. vi. 2021, S. Beshkov & A. Nahirnić-Beshkova ”; 1 ♀ ( NHMUK), “ Yugoslavia, Slovenia, Postojne, 24. vii, R. L. Coe ”; 1 ♀ ( RMNH), “ Espana [= Spain:] Huesca, Torla, 1035 m, 8–26. vii. 1974, J. Wolschrijn ”; 1 ♀ ( NMS), “ Sweden: Bohuslän, Tossene, Åby, MV, 9. vii – 13. viii. 2013, N. Ryrholm ”, “ MRS Aleiodes DNA 864 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ 14. viii – 21. xi. 2013 ” and no DNA labels; 1 ♂ ( NMS), “ Sweden: Bohuslän, Tossene, Stora Hultet MV, 8. viii – 21. xi. 2013 N. Ryrholm, “ MRS Aleiodes DNA 867 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id., but “ MRS Aleiodes DNA 868 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), id, but “ 28. v – 5. viii. 2013 ” and no DNA labels; 1 ♂ ( NMS), “ Sweden: Gästrikland, Staffen, Grinduga, MV, 23. vii – 9. 9. 2013 N. Ryrholm ”, “ MRS Aleiodes DNA 870 ”, “ DNA CO 1 worked ”; 1 ♂ ( NMS), “ Sweden: Ha [lland], Ysby Perstorp, 1–8. viii. 2004, N. Ryrholm, NMS Z 2004.167 ”, “ MRS Aleiodes DNA 385 ”, “ DNA CO 1 worked ”; 2 ♂ ( NMS), id., but no DNA labels; 1 ♂ ( NMS), “ Sweden: Skåne, Ö Hoby, Spraggehusen, MV, 1. ix – 30. x. 2013 N. Ryrholm ”, “ MRS Aleiodes DNA 854 ”, “ DNA CO 1 worked ”; 2 ♂ ( NMS), id., but no DNA labels: 1 ♂ ( NMS), “ Sweden: Skåne, Spraggehusen, MV, 20. v – 16. vii. 2017 N. Ryrholm / C. Källender ”, “ MRS Aleiodes DNA 983 ”, “ DNA CO 1 worked ”; 2 ♀ + 4 ♂ ( NMS), “ Sweden: Skåne, Käseberga, Käseberga, 17. vii – 14. ix. 2013, N. Ryrholm ”; 1 ♀ ( NHMUK), “ Sweden: Sk [åne], Degaberga, 8. vii. 1938, D. M. S. P [erkins] & J. F. P [erkins], B. M. 1938-414 ”; 1 ♀ + 1 ♂ ( NHMUK), id., but “ 10. vii. 1038 ”; 1 ♀ ( NHMUK), id., but “ 14. vii. 1938 ”; 3 ♀ ( NHMUK), “ Sweden: Skåne, Löderup, 27. vii. 1938, D. M. S. P [erkins] & J. F. P [erkins], B. M. 1938-414 ” 1 ♀ ( NMS), Switzerland: BE, Lenk, Brandegg, 1540 m, 29. vi – 3. vii. 2019, M. R. Shaw ”, “ MRS Aleiodes DNA 1033 ”, “ DNA CO 1 worked ”; 1 ♀ ( NMS), id., but “ MRS Aleiodes DNA 1034 ”, “ DNA CO 1 worked ”; 1 ♀ ( NHMUK), “ Switzerland: Grindelwald, viii. 1937, G. Nixon ”; 1 ♀ ( RMNH), “ CH [= Switzerland]: Lauerz, SZ, Schuttwald, 480 m, 8. viii. 1990, Lf, L. Rezbanyai-Reser ”; 1 ♂ ( RMNH), id., but 26. vi. 1990; 1 ♂ ( RMNH), id., but 11. ix. 1991; 1 ♂ ( RMNH), id., but Sägel (Ried), 455 m, 24. vii. 1990. Most unassociated males are considered too doubtfully determined to be treated as paratypes.

Molecular data.

We have DNA barcoded material from England, Estonia, France, Germany, Italy, Netherlands, Serbia, Sweden and Switzerland (see Fig. 1 ).

Biology.

The record ( Fahringer 1934) of A. “ vittiger ” from Atolmis (as Gnophria ) rubricollis (Linnaeus) ( Lepidoptera : Erebidae , Arctiinae, Lithosiini ) is presumed to relate to this species, but we have not seen a reared specimen ourselves except for one partially formed adult extracted from a mummy of this host from Austria that is, unfortunately, not in good enough condition to be determined unequivocally as A. pseudoseriatus . However, we have barcoded the dead parasitoid prepupa (MRS 935) from a failed mummy of this moth from the Netherlands and it clusters in the tree unequivocally with A. pseudoseriatus . Also, at a site in S. Cumbria, England where A. pseudoseriatus is the only one of the two relevant Aleiodes we have found (and barcoded), we have on several occasions obtained mummies of A. rubricollis that must undoubtedly have harboured A. pseudoseriatus , though unfortunately, none survived to produce adults of the parasitoid. The host is increasingly widely distributed and abundant in Europe, and its larva feeds on algae on (often dead) twigs of trees, perhaps with a special liking for conifers, from about July into October. It overwinters as a pupa (unlike Eilema griseola ), so in this case the parasitoid overwinters in the host mummy and has proved to be difficult to rear. To judge from their behaviour in captivity, parasitised Atolmis rubricollis larvae probably descend from trees to mummify in the litter rather than the mummy forming on twigs. Aleiodes pseudoseriatus is univoltine with a flight time from the very end of June to September.

A female paratype (not barcoded but confidently determined and from the area in England (S. Cumbria) where only A. pseudoseriatus has been found (and barcoded )), was offered cultured larvae of the lithosiin arctiine Eilema griseola (Hübner) at various stages of growth in viii. 1995, by day and at dusk when she was more active, but apart from very brief antennation on a minority of occasions she showed no interest in them.

Diagnosis.

Subbasal cell of fore wing with small glabrous patch apically (a in Fig. 20 ); pterostigma variable, often with less-developed pale yellowish patch or entirely brown antero-basally (Figs 3 , 5 ); hind femur of ♀ usually 4.7–5.5 × longer than wide; pterostigma usually dark brown or brown antero-basally, rarely yellowish (Figs 5 , 18 , 19 ); if ♀ hind femur partly dark brown, then usually paler ventrally than laterally; fourth antennal segment dark brown (Figs 3 , 15 ), if brown then darker than scapus ventrally (Fig. 18 ); vein 1 - M of fore wing of ♂ and surrounding area often darker than in A. seriatus . On average with about 3 more antennal segments than A. seriatus in both sexes. We have also seen the holotype of Rogas kuslitzkyi Tobias, 1976 , from Azerbaijan and believe it can be ruled out to belong to A. pseudoseriatus (see also notes on barcoded specimens from Primorsky Krai below).

Description.

Holotype, ♀, length of fore wing 5.2 mm, of body 5.9 mm.

Head. Antenna incomplete, but according to label originally with 47 segments, length of antenna in ♀ paratype from England 1.3 × fore wing and its subapical segments medium-sized (Fig. 17 ); frons granulate and distinctly depressed laterally; OOL 1.5 × diameter of posterior ocellus, granulate and matt; depression near posterior ocellus granulate; vertex largely granulate-coriaceous, rather dull; clypeus coriaceous; ventral margin of clypeus depressed (Fig. 12 ); face granulate but dorsally rugulose; width of hypoclypeal depression 0.4 × minimum width of face (Fig. 12 ); length of eye 3.6 × temple in dorsal view (Fig. 13 ); vertex behind stemmaticum rugulose-granulate; clypeus largely above lower level of eyes; length of malar space 0.3 × length of eye in lateral view.

Mesosoma. Mesoscutal lobes finely granulate-coriaceous, matt; precoxal area of mesopleuron rugulose but posteriorly absent, and area above it finely granulate; metapleuron densely granulate and ventrally rugose; metanotum with short median carina anteriorly and distinct depression posteriorly; scutellum finely granulate; propodeum rather long and flat, granulate anteriorly and densely rugose posteriorly, medio-longitudinal carina complete, and without protruding carinae laterally.

Wings. Fore wing: r 0.4 × 3 - SR (Fig. 5 ); 1 - CU 1 horizontal, 0.7 × 2 - CU 1; r-m 0.3 × 3 - SR; second submarginal cell medium-sized (Fig. 5 ); cu-a inclivous, straight; 1 - M straight posteriorly; 1 - SR as wide as 1 - M; surroundings of M + CU 1, 1 - M and 1 - CU 1 setose, but subbasal cell with small glabrous patch apically (a in Fig. 20 ). Hind wing: marginal cell parallel-sided, its apical width 1.1 × width at level of hamuli (Fig. 5 ); 2 - SC + R as long as wide; short m-cu present anteriorly; vein 2-1 A absent (Fig. 5 ); M + CU: 1 - M: 1 r-m = 30: 18: 18.

Legs. Tarsal claws rather robust, bristly setose and very finely yellowish pectinate; hind coxa rather shiny and only very superficially micro-sculptured, dorsally granulate; hind trochantellus rather slender (Fig. 11 ); length of hind femur and basitarsus 5.1 and 8.3 × their width, respectively; length of inner hind spur 0.2 × hind basitarsus; apex of hind tibia with distinct comb at inner side (Fig. 10 ).

Metasoma. First tergite distinctly convex medially, as long as wide apically; first and second tergites with medio-longitudinal carina, weakly indicated on third tergite; first tergite densely longitudinally rugose; second and third tergites more or less obliquely rugulose (Fig. 8 ); medio-basal area of second tergite triangular and minute (Fig. 8 ); second suture deep and distinctly crenulate; remainder of metasoma superficially micro-sculptured or smooth; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath widened, with medium-sized slanted setae and apically subtruncate (Fig. 10 ).

Colour. Dark brown; palpi, legs (but base of hind tibia dark brown), mandible (except dark brown teeth), malar space, clypeus and tegulae pale yellowish; orbita, propleuron, side of pronotum, mesosternum anteriorly, scutellum largely, first tergite medio-apically, second tergite medially (area widened posteriorly) and third tergite antero-medially yellowish brown; antenna, veins and pterostigma (but slightly paler basally than medially) mainly dark brown; third-sixth tergites posteriorly and laterally ivory (Figs 3 , 8 ); wing membrane subhyaline, but surroundings of veins 1 - M, 1 - SR, 1 - CU 1 and r of fore wing more or less infuscate (Figs 5 , 18 ).

Distribution

(from type material involved in this study): Austria, Belgium, Bulgaria, Czech Republic, England, Estonia, France, Finland, Germany, Hungary, Ireland, Italy, Netherlands, Norway, Serbia, Slovenia, Spain, Sweden, Switzerland.

Etymology.

The species is named “ pseudoseriatus ”, because of its similarity to A. seriatus .

Variation.

Pterostigma colour is rather variable, often with indistinct pale yellowish patch or entirely brown antero-basally, but sometimes with distinct yellowish basal patch; hind femur of ♀ usually 4.7–5.5 times longer than wide; ♀ with 46 (1), 47 (3), 48 (8), 49 (15), 50 (13), 51 (1) antennal segments and ♂ with 48 (1), 50 (2), 51 (5), 52 (14), 53 (14), 54 (11), 55 (5), 56 (4), 58 (1) antennal segments; fourth antennal segment dark brown (Fig. 15 ), if brown then darker than scapus ventrally, rarely both are yellow; hind femur entirely yellowish brown or with faint brown small patch to large dark brownish part; metasoma with typical black pattern. Specimens with almost unmarked metasoma seem to occur very rarely or possibly not at all. Males have, on average, about three or four more antennal segments than females.

Figures

Figure 1. Maximum likelihood tree based on all DNA barcode sequence data for taxa included in this paper, rooted using sequences from three related Aleiodes species, with full bootstrap support values for selected branches.

Figures 3, 4. Aleiodes pseudoseriatus sp. nov., paratype, ♀ (3), and mummified caterpillar of Atolmis rubricollis (Linnaeus) with A. pseudoseriatus larva within (4), both from England, S. Cumbria, Whitbarrow. 3 habitus, lateral view 4 mummy, dorsal view.

Figures 5–17. Aleiodes pseudoseriatus sp. nov., holotype, ♀, Italy, Vittoria Veneto, but 17 of paratype ♀ from England, Whitbarrow. 5 wings 6 mesosoma, lateral view 7 mesosoma, dorsal view 8 propodeum and metasoma, dorsal view 9 fore femur, lateral view 10 ovipositor sheath 11 hind femur, lateral view 12 head anterior 13 head, dorsal view 14 head, lateral view 15 base of antenna 16 antenna 17 apex of antenna.

Figures 18–19. Aleiodes pseudoseriatus sp. nov., paratype, ♂, England, S. Cumbria, Roudsea Wood. 18 habitus, lateral view 19 wings.

Figures 20–25. Aleiodes seriatus aggregate: holotype of A. pseudoseriatus sp. nov. (20), Aleiodes seriatus (Herrich-Schäffer), lectotype of A. vittiger Wesmael, ♀, Belgium (21–25) 20, 21 detail of distal half of subbasal cell of fore wing (“ a ” indicating comparatively large glabrous patch and “ aa ” a minute glabrous patch) 22 wings 23 fore femur 24 antennae 25 base of antenna. Photographs by Julian Lalanne except 20.