Aleiodes fortipes (Reinhard, 1863)

van Achterberg, Cornelis, Shaw, Mark R. & Quicke, Donald L. J., 2020, Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis group, ZooKeys 919, pp. 1-259 : 1

Aleiodes fortipes (Reinhard, 1863) Figs 322-324 , 325-338 , 339-342

Rogas fortipes Reinhard, 1863: 272; Shenefelt 1975: 1229 [examined].

Aleiodes (Neorhogas) fortipes ; Papp 1985a: 158, 1987a: 333, 1987b: 35, 1991a: 75.

Aleiodes fortipes ; Papp 2005: 176; Lozan et al. 2010: 17; Butcher et al. 2012: 14.

Rhogas freyi Hellén, 1927: 25-26; Papp 1985a: 158 (unnecessary lectotype designation and as synonym of A. fortipes ), 2005: 176 [examined].

Rogas freyi ; Shenefelt 1975: 1229-1230; Tobias 1986: 75 (transl.: 121).

Type material.

Holotype of A. fortipes , ♂ (ZMB), "Gallia [France]", “Type”, "Coll. H. Rhd.", “26723”, " fortipes Rhd.", “Holotypus”, " Rogas fortipes Reinh., 1863, ♂, Papp, 1983.". Holotype of A. freyi , ♂ (ZMH), "[Finland], Nagu", "R. Frey", " Freyi n. sp., Hellén det.", "Mus. Zool. H:fors, sp. typ. No. 5363, Rhogas Freyi Hellén”, "Lectotypus Rogas freyi Hellén, design. Tobias", " Aleiodes % ♂ fortipes Rh., det. Papp J., 1983/ compared with ♂ holotype of A. fortipes ". The lectotype designation is superfluous because it is evident from the description that the author had only one male.

Additional material.

Austria, British Isles (England: V.C.s 16, 26, 28), Bulgaria, Czech Republic, Finland, France, Germany, Hungary, Netherlands (GE: 't Harde, Nunspeet), Poland, Spain, Sweden, Turkey. Specimens in ZJUH, BZL, CMIM, FC, MTMA, NMS, RMNH, SDEI. It has been collected in open or understory habitats, including (but not exclusively) growths dominated by Vaccinium and/or Calluna below sparse conifers. Generally, found on sandy well-drained soils in England (Breck heaths of East Anglia) and the Netherlands (Veluwe).

Molecular data.

MRS650 (France), MRS807 (Poland).

Biology.

The flight time of this univoltine species is (April)May-June, and ca 10 months of the year is spent as an exposed mummy. The only mummy seen (Fig. 324 ) formed in captivity firmly attached beneath a thin stem and would have been positioned low down in the vegetation, but probably aerially. It is light brown, moderately slender, and the parasitoid occupied approximately abdominal segments 3-8. The host was Idaea sp. ( Geometridae ), either I. aversata (Linnaeus) or I. straminata (Borkhausen), and the rearing arose when a few larvae of the foregoing were collected (MRS) along with an adult female of A. fortipes at the same site in Poland (22.v.2016) and offered to the parasitoid, which had been fed honey water, on 24.v.2016. Although two of the caterpillars were well-grown, in their final instars and at least twice as long as the parasitoid, one was accepted avidly. This host was first pricked several times, at intervals. Paralysis was rather slow to take effect and not complete until after the host was revisited for oviposition: a single insertion of ca 30 seconds duration, with no post-oviposition association (the parasitoid simply walked away after oviposition). The host mummified on 9.vi.2016 and an adult female emerged on 22.v.2017. The other host was rejected after being pricked just once, and later died. A penultimate instar caterpillar of the same host aggregate was also parasitised but died after an ecdysis. Subsequent barcoding (through the kindness of Axel Hausmann, ZSSM) of the dead caterpillars revealed one specimen each of I. aversata and I. straminata , leaving the precise determination of the successful host unclear. It is possible that the parasitised host had already been attacked before it was collected, but the rather long time before mummification occurred suggests not. In any case, at least one Idaea species in the aversata / straminata group clearly serves as host. Some individuals of the long and slender, morphologically very different, larvae of Idaea muricata (Hufnagel) were also offered. Although possibly of less interest to the parasitoid, one penultimate instar larva (1.7 times the length of the female parasitoid) was immediately parasitised (a single prick for eventual paralysis, followed after an interval by a single insertion for oviposition lasting just more than a minute), but this larva later produced a moth. Final instars of this very elongate species of caterpillar were generally ignored, but one did elicit a downwards curl of the metasoma without, however, being stung.

There are two particularly significant aspects to the successful rearing. The first is that these Idaea species overwinter as quite well-grown larvae, so during the flight period of the parasitoid they are in late instars, and attacking hosts at this stage is an unusual strategy for Aleiodes (but see A. aterrimus and A. sibiricus ). The second is that we know of no other Aleiodes species apart from A. sibiricus (q. v.) among those whose host overwinters as a larva that fails to take advantage of that to overwinter as an early instar larva within it. The apparently riskier strategy taken by A. fortipes , in both respects, may be plesiomorphic.

Aleiodes fortipes is the only known West Palaearctic species in which males have small, subapical setose pore (probably associated with pheromone release) situated mid-dorsally on each of the 4th-6th metasomal tergites (Fig. 340 ). We also expect these pores to be present in A. caucasicus , which is only doubtfully distinct from A. fortipes , but we have not seen the male of A. caucasicus . Similar, probably homologous, pores are also a feature of males of Aleiodes (Hemigyroneuron) species which are found in the near East, Oriental and Afrotropical regions (Butcher & Quicke, 2015). Outside of Hemigyroneuron , metasomal pores are also found the New World Aleiodes cameronii (Dalla Torre) species complex and in a number of undescribed Madagascan Aleiodes . In Hemigyroneuron the pores have been shown to connect with large sub-tergal glands (Butcher & Quicke, 2011). Collectively these taxa form a basal clade in our molecular phylogeny (Fig. 1 ).

Aleiodes fortipes is the only species among those treated in this part of our revision with known hosts outside the Noctuidae and, although no host is known for rather a lot of these species, the apparently basal position of A. fortipes in the group is noteworthy and using geometrid hosts may also be plesiomorphic. The rather slender ovipositor sheath (Fig. 322 ) is another indication for its basal position. It is interesting that the known hosts of both A. fortipes and of the subgenus Hemigyroneuron are all Geometridae (two species of Hemigyroneuron with examined mummies, India and S. Africa, cited by Butcher & Quicke, 2011 [a label record indicating a pierid host of a 3rd species is also cited in that paper but is discounted here because no mummy was present]). An Australian species described under Hemigyroneuron with examined mummy reported to be that of a geometrid by Butcher & Quicke (2016) is probably (a) actually not a member of A. (Hemigyroneuron) and (b) may be from a lasiocampid (W. Moore in litt.). The hosts of members of the Aleiodes cameronii complex, based on multiple rearings in both North America and Costa Rica include both Geometridae and Erebidae (Eiseman & Charney, 2010; http://v4.boldsystems.org).

Diagnosis.

Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 332 ); 2nd-10th antennal segments yellowish, contrasting with remaining darker segments; clypeus obtuse apically and not protruding in lateral view (Fig. 334 ); precoxal area largely smooth, at most with some aciculae or punctures medially (Fig. 327 ); tegulae brown; lobes of mesoscutum finely coriaceous-granulate and rather dull, with satin sheen; vein 1-CU1 of fore wing 0.4-0.6 × vein 2-CU1 (Fig. 332 ); length of hind femur 3.5-3.8 × its maximum width (Fig. 331 ); hind tarsal claws small and only yellowish or brownish setose (Fig. 338 ); body of ♂ completely black, antenna completely blackish, dark brown or with some segments yellowish subbasally and 4th-6th tergites with a setose medio-dorsal depression; length of fore wing 3.7-5.0 mm.

Description.

Holotype of A. freyi , ♂, length of fore wing 4.5 mm, of body 5.3 mm.

Head. Antenna incomplete, (length of antenna of ♀ from Santon Downham 1.2 × fore wing, its subapical segments rather robust: Fig. 336 ); frons smooth anteriorly and with coarse curved striae posteriorly; OOL 2.7 × diameter of posterior ocellus, and rather regularly and rather coarsely striate; vertex transversely striate, rather shiny; clypeus narrow, strongly curved dorsal margin, rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 334 ); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 332 ); length of eye 1.4 × temple in dorsal view (Fig. 333 ); vertex behind stemmaticum transversely rugose; clypeus near lower level of eyes; face coarsely transversely rugose; length of malar space 0.4 × length of eye in lateral view.

Mesosoma. Mesoscutal lobes very densely coriaceous-granulate, with vague micro-reticulate sculpture, matt; precoxal area of mesopleuron largely smooth (except some micro-sculpture) medially, rather depressed; remainder of mesopleuron largely smooth, except some punctures and antero-dorsally coarsely rugose; scutellum superficially granulate and with some punctures; propodeum coarsely rugose-reticulate and medio-longitudinal carina nearly complete.

Wings. Fore wing: r 0.6 × 3-SR (Fig. 325 ); 1-CU1 horizontal and somewhat widened, 0.45 × 2-CU1; r-m 0.65 × 3-SR; 2nd submarginal cell rather short (Fig. 325 ); cu-a vertical, straight; 1-M rather curved posteriorly; 1-SR short and narrow; surroundings of M+CU1, 1-M and 1-CU1 evenly setose; m-cu subvertical, slightly diverging from 1-M posteriorly. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 326 ); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 80:57; 1r-m 0.6 × 1-M; 1-M straight.

Legs. Tarsal claws small but robust and only yellowish setose (Fig. 338 ); hind coxa largely rugulose-granulate; hind trochantellus medium-sized; length of hind femur and basitarsus 3.8 and 7.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus; fore femur 4.8 × as long as wide.

Metasoma. First tergite evenly convex, 1.3 × as long as wide apically; 1st and 2nd tergites with weak medio-longitudinal carina and together with basal half of 3rd tergite densely and finely longitudinally rugose; medio-basal area of 2nd tergite narrow but rather distinct (Fig. 329 ); 2nd suture rather shallow; remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; 4th-6th tergites with a setose medio-dorsal depression.

Colour. Dark brown or blackish; palpi dark brown; mesopleuron with reddish brown streak; legs yellowish brown but tarsi, apex of hind femur (and indistinctly apices of fore and middle femora, and of tibiae) and base of hind coxa infuscate; tegulae and pterostigma brown; wing membrane slightly infuscate.

Variation. Maximum width of marginal cell of hind wing 2.0-2.6 × its width near hamuli (Fig. 326 ); vein 1-CU1 of fore wing 0.4-0.6 × vein 2-CU1; length of hind femur 3.5-3.8 × its maximum width; length of 1st tergite 1.0 (♀)-1.3 (♂) × its apical width. Propodeum and metapleuron posteriorly, 1st and 2nd tergites and base of 3rd tergite of ♀ more or less brown, basal third of antenna (except scapus and base of pedicellus) brownish yellow or yellowish brown and ovipositor sheath rather slender, with long setae and apically narrowed (Fig. 323 ). Antennal segments: ♀ 38(1), 39(1), 41(1), 43(1), 44(1), 45(1); ♂ 36(1), 39(3), 40(6), 41(3), 42(7), 43(2), 44(2), 45(1). The number of antennal segments appear to be comparable between the sexes. Males have 1st-3rd metasomal tergites completely black, and basal third of antenna completely blackish, dark brown or with some subbasal segments yellowish. The male apical tergites (besides pores, see above) are type 1, setae rather dense, no fringe observed and probably absent (Fig. 341 ).

Distribution.

*Austria, *British Isles (England), Bulgaria, Czech Republic, Finland, France, *Germany, Hungary, *Netherlands, *Poland, *Spain, *Sweden, *Turkey.

Figures

Figures 322 - 324. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 322 habitus lateral 323 ovipositor sheath lateral 324 mummy of Idaea sp. (either I. aversata (Linnaeus) or I. straminata (Borkhausen )), Poland, Dybki.

Figures 325 - 338. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 325 fore wing 326 hind wing 327 mesosoma lateral 328 mesosoma dorsal 329 1 st - 3 rd metasomal tergites dorsal 330 fore femur lateral 331 hind femur lateral 332 head anterior 333 head dorsal 334 head lateral 335 base of antenna 336 apex of antenna 337 antenna 338 inner hind tarsal claw.

Figures 339 - 342. Aleiodes fortipes (Reinhard), ♂, England, Santon Downham 339 habitus lateral 340 3 rd- 7 th metasomal tergites dorsal 341 id. lateral 342 metasoma dorsal. Arrows indicating setose depressions or pores of 4 th- 6 th tergites.

Figures 1 - 6. Terminology and measurements used in this paper 1 wing venation: pa = parastigma, pt = pterostigma, 1 = marginal cell, 2 a, b, c = 1 st, 2 nd and 3 rd submarginal cell, respectively, 3 a, b = 1 st and 2 nd discal cell, respectively, 4 a = 1 st subdiscal cell, 5 = basal cell, 6 = subbasal cell 2 head, dorsal aspect: a = length of eye, b = length of temple 3 head, lateral aspect: c = width of temple, d = width of eye, e = height of eye, f = width of malar space (measured as actual true distance in its own plane) 4 head, anterior aspect: g = width of face, h = width of hypoclypeal depression 5 fore femur, lateral aspect: i = length, j = width 6 1 st metasomal tergite, dorsal aspect: k = length of tergite (measured from adductor), l = apical width of tergite.