Holotype, ♀ (NMS, Edinburgh), “[England], Otmoor N. R., Oxon., H[ost]: Abraxas grossulariata [on] Prunus spinosa, HLC [= host larva collected] 13.v.[19]79, PLE [parasitoid larva evident = mummification in the case of Aleiodes] 11.vi.[19]79, PIE [= parasitoid imago emerged] 4.vii.[19]79, M.R. Shaw”. Paratypes (74 ♀, 34 ♂): 52 ♀, 25 ♂ reared from larvae of the geometrid Abraxas grossulariata (Linnaeus) collected in v/vi, em (v)vi/vii as follows: 14 ♀, 4 ♂ (NMS, RMNH) England, Oxford, Otmoor, 1972, 1973, 1979, M.R. Shaw; 2 ♀, 3 ♂ (NMS, BMNH) England, West Sussex, Littlehampton, 1978, 1979, A.A. Allen; 1 ♂ (NMS) England, West Sussex, Hove, 1982, A.R. Cronin; 3 ♀, 1 ♂ (NMS, AAC) England, Surrey, Salfords, 1976, A.A. Allen; 1 ♀, 1 ♂ (NMS, RMNH) England, Berks, Maidenhead Thicket, 1979, M.R. Britton; 4 ♀, 4 ♂ (NMS, RMNH) England, Bucks, Butlers Hangings, 1979, M.R. Shaw; 2 ♀ (NMS) England, Bucks, Milton Keynes, 1984, J.P. Brock; 1 ♀ (BMNH) England, Cambridge, 1913, L. Doncaster; 2 ♀ (NMS) England, Westmorland, Beetham, 1991, M.R. Shaw; 11 ♀, 1 ♂ (NMS, BMNH) Scotland, Fife, St Andrews, 1935, 1936, 1938, D.J. Jackson; 1 ♀ (NMS) Scotland, Stirling, D.J. Jackson; 8 ♀, 7 ♂ (NMS, RMNH) Scotland, Orkney, Mainland, Waulkmill Bay, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Mainland, Caldale Bottom, 2009, K.P. Bland; 2 ♀ (NMS) Scotland, Orkney, Mainland, Redland, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Hoy, Nowt Bield, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Hoy, Enegars, 2004, S. Gauld; 2 ♀ (SDEI) Germany, Sachsen-Anhalt, Wolfen, 1957, B. Stehlik; additionally 1 ♀ (NMS) from the Otmoor locality, host larva collected 8.x.1978, mummified 4.vi.1979, emerged 3.vii.1979, M.R. Shaw. Non-reared specimens: 1 ♂ (NMS) England, Cambridge, Chippenham Fen, 9.vii.1983, M.R. Shaw; 1 ♀ (NMS) England, Hunts, Monks Wood, 31.viii.2005, G.R. Broad; 1 ♂ (NMS) England, East Gloucester, Eastleach, 8.viii.2006, M.R. Shaw; 1 ♀ (BMNH) East Cornwall, Botusfleming, Marshall collection; 4 ♀ (BMNH) England, Oxford, Stanton St. John, 19.viii. 1968 (1) and 4.ix.1968 (3), J.P. Brock; 1 ♂ (BMNH) England, Herts, Whetstone, 24.vii.1961, P.H. Ward; 1 ♀, 1 ♂ (BMNH) England, Northampton, Spratton, x.1975 and vii.1976 respectively, I. & P. Gauld; 2 ♀ (BMNH) British Isles, Harwood coll.; 1 ♀ (BMNH) presumed British, A. Matthews in Lyle coll.; 3 ♀ (BMNH) presumed British, Stephens coll.; 1 ♀ (CMIM) England, Dorset, Weymouth 24.vi.1899, Peachell; 1 ♀ (CMIM) England, West Suffolk, Old Newton; 1 ♂ (CMIM) England, East Suffolk, Monk’s Soham, 18.vii.1933; 2 ♀, 1 ♂ (AAC) England, South Devon, Shaldon, 7.viii.1978 (1 ♀) and 6.viii.1979 (1 ♀, 1 ♂), A.A. Allen; 2 ♀ (NMS, RMNH) Wales, Anglesey, Llangristiolus, 27.viii–25.ix.1982, S.A. & D.C. Wilkinson; 1 ♀ (NMS) Scotland, West Ross, Sheildaig, viii. 1991, I. MacGowan; 2 ♂ (NMS) Scotland, South Uist, Loch Eynort, vi.1988, D. Whiteley; 2 ♀ (ALC, RMNH) Czech Republic, South Bohemia, Šumava Boubinský Pralés, virgin forest, 1000–1300 m, light trap, 22–24.vii.2003, I. Jaroš & K. Spitzer; 1 ♀ (ALC), Czech Republic, South Bohemia, Šumava Mts, 740 m, peat bogs near Mrtvý, light trap, 21–24.viii.2001, I. Jaroš & K. Spitzer; 1 ♂ (NMS) Finland, Satakunta, Pori, 1991, K. Ruohomäki; 1 ♀ (NMS) Sweden, Gotland, Sundre, Barrshage, vii/viii.2004, N. Ryrholm.

MRS391 (Sweden JF962827, CO1), MRS636 (Scotland HQ551278/HQ551264, CO1), MRS637 (Scotland HQ551262, CO1 + KU682263, 28S), MRS694 (Scotland HQ551277, CO1).

Univoltine and presumed monophagous parasitoid of Abraxas grossulariata, overwintering in the living host larva and killing it in early summer in its penultimate instar. Mummy (Fig. 8) black and moderately swollen. In Britain very widespread and present in most well-established colonies of the host, on its various foodplants (rearing records from Prunus spinosa, Ribes uva-crispa, Ribes nigrum, Calluna vulgaris and Euonymus japonicus). It has not been reared from collections of the congeneric Abraxas sylvata (Scopoli) which hibernates as a pupa and would not provide a means for the parasitoid to overwinter. The essentially univoltine host overwinters as a small caterpillar, within which the parasitoid overwinters as an early (probably first) instar larva. The mummies of penultimate instar hosts are made in exposed situations on very narrow stems etc. in about early June and, no doubt aided by their almost black and hence heat-absorbing colour, adult emergence follows quickly; the mummies otherwise being subject to high levels of pseudohyperparasitism. The adult females are unwilling to mate soon after their emergence in the morning but become highly receptive in early evening, offering themselves to males at the first contact which sometimes happens before the males are fully aware of them, and repeated copulation bouts with the same or another male frequently ensue over the next tens of minutes. The new generation of hosts is not available for several weeks, but the adult females are long-lived (confirmed in captivity), and have been collected from June well into the autumn (as late as October). The host has been in severe decline in Britain in recent years, apart from its strong presence on Calluna in parts of Scotland, which remain also a stronghold for the otherwise declining parasitoid. When surveying for the presence of the parasitoid, undersized hosts collected in mid to late May are the most likely to yield results.

Figures 7–8. 

Aleiodes abraxanae sp. n., ♀, holotype. 7 habitus lateral 8 mummy of Abraxas grossulariata (Linnaeus).

Pterostigma of both sexes blackish or dark brown medially, border between dark and pale part sharp, contrasting with each other (Figs 7, 9); temples directly strongly narrowed (Fig. 18) and comparatively wide in lateral view (Fig. 17); OOL about equal to diameter of ocellus; propodeum and first tergite yellowish or medially brown (Fig. 11); vein 2-SR of fore wing yellowish as vein 1-R1 (Fig. 9); head moderately transverse (Fig. 18); antennal segments of ♀ 41–45, of ♂ 40–44; subapical antennal segments slender (Fig. 15); vertex may be distinctly rug(ul)ose and mesopleuron only coriaceous medially (Fig. 10); body entirely brownish yellow, at most propodeum and first tergite medially brown. Similar to A. hellenicus Papp, 1985, but A. hellenicus has pterostigma of both sexes completely pale yellowish and precoxal area usually with rugae medially.

Figures 9–19. 

Aleiodes abraxanae sp. n., ♀, holotype. 9 wings 10 mesosoma lateral 11 propodeum and anterior half of metasoma dorsal 12 hind leg lateral 13 fore femur lateral 14 antenna 15 apical segments of antenna 16 head anterior 17 head lateral 18 head dorsal 19 basal segments of antenna.

Holotype, ♀, length of fore wing 5.1 mm, of body 6.2 mm.

Head. Antennal segments of ♀ 43, length of antenna 1.3 × fore wing, its subapical segments about 2.3 × as long as wide; frons only coriaceous, matt; OOL 0.9 × diameter of posterior ocellus and coriaceous; vertex coriaceous, matt; clypeus convex, coriaceous; ventral margin of clypeus thick and depressed (Fig. 16); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 16) and face coriaceous; length of eye 3.8 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum coriaceous and occipital carina interrupted by somewhat less than width of ocellus (Fig. 18); clypeus partly above lower level of eyes (Fig. 16); length of malar space 0.3 × height of eye in lateral view; eyes distinctly protruding (Figs 16, 18).

Mesosoma. Mesoscutal lobes largely coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli narrow and smooth, posteriorly lost in rugose area; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron and metapleuron coriaceous, matt; mesopleuron above precoxal area (except smooth and shiny speculum) coriaceous, but dorsally rugose; mesosternal sulcus narrow and shallow, impressed and without carina posteriorly; mesosternum angulate posteriorly; scutellum slightly convex, coriaceous, and carinate laterally; propodeum evenly convex and rugose but anteriorly weakly so, median carina complete, without tubercles.

Wings. Fore wing: r 0.4 × 3-SR (Fig. 9); 1-CU1 horizontal, 0.35 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Figs 7, 9); cu-a vertical, not parallel with CU1b, straight (Fig. 9); 1-M slightly curved posteriorly. Hind wing: apical half of marginal cell parallel-sided; 2-SC+R short and longitudinal; m-cu present but unpigmented.

Legs. Tarsal claws setose; hind coxa coriaceous, largely matt; hind trochantellus 2.6 × longer than wide; length of fore and hind femora 6.1 and 4.9 × their width, respectively (Figs 12–13); inner apex of hind tibia without comb; length of inner hind spur 0.25 × hind basitarsus.

Metasoma. First tergite nearly as long as wide posteriorly, moderately convex and latero-posteriorly lamelliform; first and second tergites densely and finely longitudinally rugose, robust (Fig. 11), with distinct median carina; medio-basal area of second tergite obsolescent; second suture shallow and crenulate; basal half of third tergite finely rugose, remainder of metasoma largely superficially coriaceous; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath densely setose.

Colour. Brownish yellow; apical fifth of antenna and dorsally propodeum dark brown; ovipositor sheath black; palpi, tegulae, apical 0.4 of first tergite and more or less second tergite pale yellowish; veins (but distally from 2-SR yellowish) and pterostigma (except yellow base and apex) dark brown; border between dark and pale part of pterostigma sharp, contrasting with each other (Figs 7, 9); wing membrane subhyaline.

Variation. Length of fore wing 4.4–5.3 mm; antennal segments of ♀ 41(10), 42(18), 43(30), 44(6), and 45(1), of ♂ 40(3), 41(7), 42(8), 43(4), 44(4); stemmaticum of male black and of female brownish yellow; basal 0.2–0.5 of pterostigma pale yellow, rarely largely yellow and only medially darkened; first tergite yellowish or infuscate medially.

Named after the generic name of its host: Abraxas Leach.

*British Isles (England, Scotland, Wales), *Czech Republic, *Finland, *Germany, *Sweden.

The males of this species have on average about one antennal segment less than females.

Redescribed ♀ and holotype of A. heterogaster (KBIN), “[Belgium], Campine, 1833”, “A. heterogaster mihi, det. C. Wesmael”, “Belgique, Charleroi/teste Papp, J., 1983”, “Holotypus Aleiodes heterogaster Wesm., 1838 / Papp, 1983”. The type series of R. albitibia is lost.

*Austria, British Isles (England: V.C.s 3, 11, 15, 22, 58, 59, 61, 69; Wales: V.C. 49; Scotland: V.C.s 72, 77, 80, 88, 92, 96, 97, 98, 99; Ireland: (V.C.s H1, H19, H20), *Czech Republic, Finland, France, Germany, Hungary, Netherlands (DR: Wijster, LI: Asselt; Castelre, GE: Heerde; Putten; Tongeren, NB: Tilburg (Kaaistoep), NH: Muiderberg, OV: Buurse (Schipbeek)), Poland, *Spain, Sweden, Switzerland. Specimens in NMS, BMNH, BZL, CNC, OUM, RMNH, SDEI, USNM, ZSSM, I. Kakko collection, and WUIM.

MRS210 (Scotland EU979574, CO1), MRS383 (Sweden JF962835/ KU682238, CO1), MRS753 (Sweden KU682248, CO1).

Univoltine parasitoid of arboreal notodontids, overwintering in a highly distinctive mummy (Fig. 21). Specimens (in NMS unless indicated) reared from wild collected arboreal Notodontidae identified as Notodonta dromedarius (Linnaeus) (12 [5 are OUM, 2 are USNM]; T.H. Ford, M.J. Morgan, M.R. Shaw, A.W. Stelfox, W.A. Watson), Eligmodonta ziz-zac (Linneaus) (2:1; M.R. Shaw), Pheosia tremula (Clerck) (8 [1 is AAC]; A.A. Allen, B.T. Parsons, M.R. Shaw), Pheosia gnoma (Fabricius) (2 [BMNH]; G. Graham-Smith). Host range experiments had the following outcomes: E. ziz-zac 2:32\10\\8+2; Pterostoma palpina (Clerck) 2:10\0\\-; Ptilodon capucina (Linnaeus) 1:5\0\\-; Clostera pigra (Hufnagel) 2:9\0\\-; Phalera bucephala (Linnaeus) 2:2\0\\-. The developmental biology of this species is rather unusual in several respects. On approaching the host (E. ziczac in all the following observations, which are based on two female A. albitibia) the female seems to depend on its fore and middle tarsi more than its antennae for host assessment, although antennation does occur (possibly as much to desensitise the host as to investigate it). The host is not paralysed during attack: the female more or less pounces on the host once accepted and aligns herself along the host’s body, which she grasps with her front and middle legs during oviposition, and fans her wings for short bursts repeatedly during the oviposition process (this may have indicated that venom was being injected, but if so it did not appear to have a significantly paralysing action), which in some cases lasted as long as 10–15 minutes – in these cases usually with more than one insertion. In fact, probably because the host was not temporarily parasitised and so was not sluggish, self-superparasitism happened very easily. The metasoma is only weakly curled downwards during oviposition, after which the female left the host without the usual post-oviposition period of association. When young, the females host-fed non-destructively on occasion. Development of the parasitoid larva was extremely rapid at ca 22–25 °C, with mummification ensuing after as few as 7–10 days from oviposition. The unusual and highly distinctive mummy (Fig. 21) is very shiny and dark mahogany brown in colour. It consists of the caudal portion of the host (usually from abdominal segment 3 onwards), strongly delimited from the anterior portion which usually shrivels up and becomes detached, leaving a sharp rim. The parasitoid pupates in a capsule which is formed in the most caudal half of this structure (usually in abdominal segments 5–8), dorsally raised, thinly lined with silk, and sealed from a more ventral and anterior inflation of the host’s cuticle that is of approximately equal volume but fully open at its anterior end. During the mummification process this area fills with liquid and bubbles, but fluids do not spread to the substrate and the mummy does not become stuck down. Once dried and hardened, the whole structure looks as though it contains two pupating parasitoids, and indeed some authors have been misled by this (Hammond and Smith 1957). It is possible that the empty chamber may serve to decoy pseudo-hyperparasitoids, though this seems unlikely to be its main function. The mummy forms on the host’s food plant, usually on a leaf surface from which it is easily dislodged, and overwinters in the leaf litter. Aleiodes albitibia occurs particularly in wet, bushy places: bearing in mind that its hosts (evidently rather restricted within Notodontidae) feed on trees and shrubs (Salix, Populus, Alnus and Betula) that often overhang water, the form of the mummy may also be an adaptation to prevent submersion and perhaps also results in dispersion by water, as it floats easily and is not wetted. The winter is passed in the mummy, and A. albitibia is univoltine, with a flight period in Britain of roughly JuneAugust.

Figures 20–21. 

Aleiodes albitibia (Herrich-Schäffer), ♀, England. 20 habitus lateral 21 mummy of Notodonta dromedarius (Linnaeus).

Third antennal segment robust (Fig. 34); area in front of anterior ocellus without tubercle; OOL 0.4–0.5 × diameter of posterior ocellus (Fig. 32); mesopleuron strongly shiny and precoxal area not impressed (Fig. 24); pterostigma dark brown; vein 1r-m of hind wing about as long as vein 1-M (Fig. 23); inner hind tibial spur 0.4–0.5 × hind basitarsus; inner side of basal half of hind tibia whitish, rarely largely dark brown; inner side of hind tibia with bristly setae and no comb apically; third tergite curved medio-posteriorly in dorsal view (Fig. 25) and longer than second; metasomal tergites largely black (also laterally so), sometimes with a large yellow or ivory central patch on second tergite.

Figures 22–34. 

Aleiodes albitibia (Herrich-Schäffer), ♀, Switzerland. 22 fore wing 23 hind wing 24 mesosoma lateral 25 propodeum and anterior half of metasoma dorsal 26 hind leg lateral 27 apical segments of antenna 28 fore femur lateral 29 head anterior 30 head lateral 31 ovipositor sheath lateral 32 head dorsal 33 antenna 34 basal segments of antenna.

Holotype of A. heterogaster, length of fore wing 5.4 mm, of body 5.6 mm.

Head. Antennal segments of ♀ 46, long setose, length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide; frons weakly depressed, finely rugose medially, remainder superficially micro-granulate; OOL 0.4 × diameter of posterior ocellus, and finely granulate; vertex finely granulate, with some rugulae posteriorly, rather dull; clypeus normal, micro-granulate; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.5 × minimum width of face (Fig. 29); length of eye 2.8 × temple in dorsal view (Fig. 32); occiput behind stemmaticum granulate with some rugulae, narrow; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view; occipital carina widely interrupted medio-dorsally and ventrally (Fig. 30).

Mesosoma. Mesoscutal lobes largely granulate and with punctulation, matt; prepectal carina complete, rather weak; precoxal area of mesopleuron largely smooth (but in other specimens usually micro-granulate with some punctulation); mesopleuron above precoxal area strongly shiny and smooth; metapleuron largely coriaceous; scutellum granulate and finely punctate, no distinct carina; propodeum evenly convex, short, anteriorly granulate, medially and posteriorly rugose, median carina complete, without tubercles.

Wings. Fore wing: r 0.5 × 3-SR (Fig. 22); 1-CU1 horizontal, 0.35 × 2-CU1; r-m 0.5 × 3-SR; second submarginal cell rather long (Fig. 22); cu-a inclivous, curved posteriorly; 1-M straight posteriorly. Hind wing: marginal cell subparallel-sided, its apical width 1.0 × width at level of hamuli (Fig. 23); 2-SC+R shortly longitudinal (but in other specimens subquadrate); m-cu absent; M+CU:1-M = 5:3; 1r-m about as long as 1-M.

Legs. Tarsal claws yellowish setose; hind coxa sparsely punctulate, and granulate; hind trochantellus robust; length of fore femur, hind femur and basitarsus 5.4, 3.7 and 6.6 × their width, respectively (Figs 26, 28); length of inner hind spur 0.5 × hind basitarsus, as long as outer spur.

Metasoma. First tergite robust (Fig. 25); first and second tergites rather coarsely longitudinally (reticulate-)rugose, robust, with distinct median carina; medio-basal area of second tergite minute, triangular; second suture shallow; third tergite as long as second tergite and largely granulate, anteriorly with some rugulae; remainder of metasoma smooth, compressed; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath slender and rather shiny.

Colour. Black; malar area, narrow stripe along eyes dorsally, fore and middle legs (but telotarsi infuscate), hind coxa, trochanter trochantellus and femur, palpi and tegulae yellowish; pterostigma and most veins dark brown; basal 0.6 of hind tibia ivory; remainder of hind tibia and tarsus blackish.

Variation. Antennal segments of ♀: 43(8), 44(14), 45(14), 46(3), 47(3), 49(1); of ♂: 39(1), 40(7), 41(7), 42(4), 43(3), 44(2). Second metasomal tergite may be largely yellowish or ivory (except laterally), or only with pale basal patch; hind tibia may be largely dark brown; base of pterostigma and first tergite medio-apically completely black (typical A. heterogaster) or yellowish (typical A. albitibia); width of hypoclypeal depression 0.4–0.5 × minimum width of face; mesopleuron usually with faint brownish longitudinal streak ventrally.

Males average about four fewer antennal segments than females. As is the case for the vast majority of Aleiodes species, only one parasitoid develops in each host (paceHammond and Smith 1957).

Holotype, ♀ (RMNH, Leiden), “Nederland (Dr.), Wijster, opposite Biol. Station, 12–19.viii.1977, C. v. Achterberg”. Paratypes (15 ♀): 1 ♀ (NMS), England, Cumbria, Whitbarrow, Howe, MV light, 24.viii.1995, M.R. Shaw; 1 ♀ (NMS), England, Norfolk, Scarning, TF981120, 6.vii–1.ix.1988, A.P. Foster/NCC; 1 ♀ (NMS), England, Norfolk, Sutton, TQ373235, water trap, 21.viii–4.ix.1986, A.P. Foster/NCC; 2 ♀ (NMS), Wales, Anglesey, Fedw Fawr, SH6081, MV light, 11.viii.2003, M.R. Shaw; 1 ♀ (NMS), Wales, Gwent, Magor Marsh, ST425865, water trap 8–21.vii.1988, P. Holmes/NCC; 1 ♀ (Tullie House Museum, Carlisle), Scotland, Dumfriesshire, Gretna, Springfield, 17.viii.1939, J. Murray; 1 ♀ (BMNH), Jersey, Trinity, Howard Davis Farm, Rothamsted trap 18.iv–3.xii.2004, A. Vautier/P. Gould; 1 ♀ (FMNH), Finland, U. Vantaa. 6690:384. ex larva Hypenodes humidalis 27.v.1974, cocoon [in which the mummy formed?] 19.vi.1974, em. 3.vii.1974, E.O. Peltonen; 1 ♀ (BZL), Greece, Thráki NW, Mt. Menikio, 12.viii.2010, J. Halada; 1 ♀ (NMS), Lithuania, Cerkelia peat bog, 3.ix.2006, A. Lozan; 1 ♀ (M. Riedel Collection), Russia, E. Siberia, 10 km E Irkutsk, 8.viii.2005, Berlov; 1 ♀ (NMS), Portugal, Azores, ca 2008 [per D.L.J. Quicke, no further data]; 1 ♀ (MRS), China, Yangtze River near Fengdu, 15.vii.2002, M.R. Shaw; 1 ♀ (RMNH), Japan, Kusakai, Kawai V., Iwate, 3–4.viii.1981, A. Takasu.

MRS172 (China KU682231, CO1), MRS279 (Wales KU682232, CO1), MRS280 (Wales KU682233, CO1), MRS822 (Azores KU682246, CO1).

No males have been seen, suggesting that this species might be thelytokous. Only a single reared specimen examined, from Hypenodes humidalis Doubleday (Erebidae: Hypenodinae). From the specimen labelling, the mummy appears to be formed in the host cocoon (but this has not been examined) and the adult emerged the same year. From this, and the flight data (vii-viii), it is surmised that it is a univoltine species, overwintering in the partly fed host larva. Hypenodes humidalis occurs in both acidic and alkali marshy areas, and the larva feeds on plant debris certainly including dead or dying Molinia caerulea (G.M. Haggett, personal communication). Indeed, when known the collecting sites of A. angustipterus have mostly been wet grasslands, including fens and bogs, but at least one specimen was collected in woodland on a limestone hill (Whitbarrow) which may suggest a wider host range.

Head subglobular (Fig. 46) and body slender; antenna of ♀ without a pale submedial band; antennal segments of ♀ 36–40; eye rather small (Fig. 45); OOL 1.2 × posteior ocellus; speculum of mesopleuron rugose or reticulate and dull as is remainder of mesopleuron (Fig. 38); propodeum slightly elongate (Fig. 38); fore wing narrow (Fig. 36); pterostigma brown; hind coxa distinctly shorter than first tergite; hind femur 6–7 × as long as its maximum width; hind trochantellus slender (Fig. 40); dorsal carinae of first metasomal tergite lamelliform protruding basally; second tergite with small smooth triangular area medio-basally and tergite rather short (Fig. 39); third tergite weakly sculptured; fourth tergite partly or entirely without sharp lateral crease, fourth and following tergites partly retracted and largely smooth. Morphologically similar to A. jakowlewi from Finland, Sweden and N. Russia, but A. jakowlewi has the hind coxa about as long as first tergite and second tergite comparatively long (hind coxa distinctly shorter than first tergite in A. angustipterus (Fig. 35) and second tergite comparatively short (Figs 35, 39)); fourth tergite with distinct sharp lateral crease and basally rugulose (fourth tergite partly without distinct sharp lateral crease, partly retracted and largely smooth); third tergite strongly sculptured (third tergite weakly sculptured); pterostigma dark brown with basal third pale (pterostigma dark brown); eye comparatively large (eye comparatively small); antenna of ♀ sometimes with a narrow white or pale yellowish submedial band (antenna of ♀ without a pale submedial band); antennal segments of female 49–52 (36–40).

Figure 35. 

Aleiodes angustipterus sp. n., ♀, holotype, habitus lateral.

Figures 36–47. 

Aleiodes angustipterus sp. n., ♀, holotype. 36 fore wing 37 hind wing 38 mesosoma lateral 39 propodeum and anterior half of metasoma dorsal 40 hind leg lateral 41 fore femur lateral 42 antenna 43 basal segments of antenna 44 head anterior 45 head lateral 46 head dorsal 47 apical segments of antenna.

Holotype, ♀, length of fore wing 3.2 mm, of body 4.1 mm.

Head. Antennal segments of ♀ 36, length of antenna 1.1 × fore wing, its subapical segments about 1.4 × as long as wide; frons granulate, rather shiny; OOL and POL 1.2 and 1.5 × width of posterior ocellus, respectively; vertex superficially granulate-coriaceous, rather shiny; clypeus convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 44); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 44) and face coriaceous with some rugulae; length of eye 2.4 × temple in dorsal view and temple directly narrowed behind eye; head subglobular (Fig. 46); occiput behind stemmaticum coriaceous with satin sheen; occipital carina complete and dorsally arched (Fig. 46); clypeus partly above lower level of eyes (Fig. 44); length of malar space 0.4 × height of eye in lateral view; eyes somewhat protruding (Figs 44, 46).

Mesosoma. Mesoscutal lobes coriaceous-rugulose, matt, but medio-posteriorly longitudinally rugose and anteriorly low; notauli narrow and crenulate, but sculpture largely lost; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron, area below it and mesosternum largely reticulate-rugose; remainder of mesopleuron (including speculum) rugose or rugulose and matt (Fig. 38); metapleuron rugose, matt; mesosternal sulcus deep and sparsely crenulate; mesosternum rounded posteriorly; scutellum slightly convex, rugulose, and laterally with irregular carina; propodeum flattened, without tubercles and coarsely rugose, median carina incomplete, posterior 0.3 absent.

Wings. Fore wing: r 0.4 × 3-SR (Fig. 36); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Fig. 36); cu-a vertical, not parallel with CU1b, straight; 1-M straight and 1-SR angled with 1-M. Hind wing: apical half of marginal cell slightly widened; 2-SC+R short; m-cu obsolescent.

Legs. Tarsal claws with yellow bristles; hind coxa rugulose and with spaced oblique rugae, with satin sheen and 0.7 × as long as first tergite; hind trochantellus 2.4 × longer ventrally than wide; length of fore and hind femora 5.8 and 6.1 × their width, respectively (Figs 40–41); inner apex of hind tibia without distinct comb; length of inner hind spur 0.25 × hind basitarsus.

Metasoma. First tergite 1.1 × as long as wide posteriorly, convex anteriorly and dorsal carinae lamelliform protruding basally; first and second tergites longitudinally rugose, robust (Fig. 39), with distinct median carina; medio-basal area of second tergite minute; second suture narrow and crenulate; basal half of third tergite largely superficially coriaceous, with some fine longitudinal elements; third tergite with complete sharp lateral crease, absent on following tergites; ovipositor sheath densely setose and apically acute.

Colour. Dark brown; head (except stemmaticum), mesoscutum and scutellum medially, tegulum, legs (but femora largely infuscate), patch on posterior third of first tergite, large triangular patch on second tergite (Fig. 39) and anterior patch of third tergite brownish yellow; fourth–seventh tergites yellow; mouthparts, humeral plate and metasoma ventrally pale yellow; ovipositor sheath black; veins and pterostigma dark brown; wing membrane infuscate.

Variation. Antennal segments of ♀: 36(2), 37(4), 38(2), 39(3), 40(2). The male is unknown. Pale patches of first and third tergites sometimes absent; hind femur 6.1–7.0 × as long as wide and hind trochantellus 2.4–3.0 × longer ventrally than wide. Central antennal segments vary from 1.2–1.5 × as long as wide, but in one specimen about 2.2 times – although its metasoma (at least posteriorly) is female, it seems possible that this individual is an intersex.

From “angustus” (Latin for “narrow”) and “pteron” (Greek for “wing”), because of the narrow wings.

*British Isles (England, Scotland, Wales, Jersey), *Finland, *Greece, *Lithuania, *Netherlands, *Portugal (Azores), *Russia (Siberia), *China (Chongqing), *Japan (Honshu).

CO1 sequences obtained from the paratypes from Azores and China group closely with those from Britain, and this seldom-collected species appears to have a very wide distribution.

Holotype of A. apiculatus, ♀ (ZMB), “Type”, “Coll. H. Rhd”, “Germania, [surroundings of] Bautzen”, “Holotypus Rogas apiculatus Reinh., 1863, ♀, Papp, 1983”, “Aleiodes pallidator Thunb., ♀, det. Papp, J., 1984/ var. apiculatus (Fahr.)”. Paratype of A. negativus, ♀ (BMNH) from Russia (Siberia: Tuvinskaya ASSR).

*England (V.C.s 9, 17, 20, 23, 24, 29, 31), *Poland. Specimens in NMS, BMNH, RMNH, AAC.

MRS028 (England EF115455, CO1 + EF115440, 28S), MRS064 (England KU682218, CO1), MRS079 (England KU682222, CO1), MRS407 (England KU682239, CO1).

This rather poorly-known species is a probably monophagous parasitoid of Euproctis similis (Fuessly) (Erebidae: Lymantriinae), from which we have seen 5 rearings (England, Poland; A.A. Allen, S.D. Beavan, M.R. Shaw, L. Sukovata) in addition to a reared paratype of A. negativus from the same host. Although evidently not obligatorily so (see below), it is probably largely univoltine, and the winter is passed inside the diapausing host larva. The host is arboreal, and when parasitised shows strong climbing behaviour just before being mummified, such that mummies are formed in exposed positions. It was readily reared from E. similis in culture, but quantitative data are not available owing to high overwintering mortality. The notes that follow relate to a single, virgin, female. This female showed great interest in an egg mass of its host, antennating the dense covering of setae left by the female moth, and probing also with the ovipositor but probably without attempting to oviposit except into fully eclosed larvae as they exited from the felted covering. Neither legs nor antennae were used to manipulate such hosts, and the process was achieved with a single insertion of the ovipositor. In subsequent trials, second instar hosts were offered naked, and it was clear that there was an injection of a temporarily paralysing venom (detected by a clear jerk of the wings) before actual oviposition took place, although the ovipositor was usually not removed in the interim. As with the emerging first instars, the use of antennae or legs to hold the host was minimal so usually the ovipositor was all that was in contact with the host, pinning it against the substrate, and oviposition usually took about (often just over) 2 minutes, without a period of post-oviposition assessment or at most with only a minimal one. The long setae of third instar hosts were a good deal harder for the female to penetrate, but some ovipositions into this instar also occurred.

In culture A. apiculatus proved to be, like its host, partly plurivoltine. The host invariably overwinters as a partly grown larva in a densely spun hibernaculum, and the adult moths appear in the following vi/vii. In captivity, a small proportion of host larvae (available from about vii onwards) from the resulting eggs fed up rapidly and produced a second generation of the moth, while the majority developed only slowly and entered hibernation in the autumn (often not until the end of ix) while still relatively small, joined at that time by offspring of the second generation. The parasitoid invariably overwinters as a small larva within these diapausing hosts. From overwintering hosts mummification takes place in about (v–)vi the following year, and emergence of the adult parasitoids in about (vi–)vii, to oviposit into the young hosts that appear soon after. In host individuals with the accelerated growth pattern the host was mummified in about viii and the adult parasitoids emerged in ix (N = 4). At this time host larvae, from both generations, are still available prior to constructing their hibernacula. The cohort of hosts with accelerated growth that produced a second generation during the culture experiments arose in control groups as well as among the parasitised hosts, so this behaviour was not the result of having been parasitised: rather, it seems likely that only the growth of host individuals independently destined for a second generation would have provoked similar early development by the parasitoid. Hosts bearing the parasitoid entered winter diapause on average an estimated 8–10 days sooner than unparasitised ones. For the hibernaculum, the parasitised hosts constructed a weak outer web, moulted, and then made a much denser inner chamber isolated from the exuvium, while unparasitised controls usually moulted before commencing construction of a single chamber. Parasitised hosts (N = 10) broke diapause in spring over a period of 22 days, on average 8.0 days later than controls (N = 9) which emerged from their hibernacula over a period of 11 days (see also A. pallidator which exhibits similar behaviour).

Despite the possibility of plurivoltinism revealed in culture experiments the capture dates, in Britain (vi–)vii–ix(–x), suggest that a single generation of rather long-lived individuals is the norm. It appears to have colonised Britain only recently; the first specimens known to us were collected in 1999 in Berkshire, since when it has been taken in MV traps in the SE corner of England fairly regularly. It is unlikely to have been long-overlooked in Britain, as its rather common and attractive host larva is conspicuous, readily identified, often reared and, when mummified by this parasitoid, often easily seen in a sun-exposed position.

Head transverse in dorsal view and directly narrowed ventrally in anterior view; eye rather large; OOL 0.5 × width of posterior ocellus; scapus and pedicellus of ♀ at least partly blackish, contrasting with yellowish middle of antenna and antenna of ♀ in dorsal view bicoloured, first–fifth[–eighth] and few apical segments more or less dark brown, remainder of antenna yellowish, antenna of ♂ entire yellowish; antennal segments of ♀ 46–49; length of malar space of ♀ 0.25–0.30 × height of eye in lateral view (Fig. 60; of ♂ 0.30 times); speculum of mesopleuron granulate and with satin sheen; fore wing rather narrow (Fig. 50); vein m-cu of fore wing straight and angled to vein 2-CU1 (Fig. 50); pterostigma dark brown with its basal half largely pale yellow; hind femur about 4.5 × as long as its maximum width; hind trochantellus about 2.6 × as long ventrally as wide (Fig. 53); hind tibia completely brownish yellow (Fig. 53); tarsal claws with distinct fine pecten (Fig. 54); dorsal face of propodeum medium-sized and rounded posteriorly (Figs 51, 52), first metasomal tergite lamelliform protruding latero-anteriorly (Fig. 62); second metasomal tergite rather stout and with minute smooth triangular area medio-basally (Fig. 52); third tergite finely sculptured; apical half of third and fourth tergite without sharp lateral crease and superficially granulate and with satin sheen; fourth metasomal tergite of ♀ black latero-posteriorly (Figs 52, 53; of ♂ brownish yellow).

Figures 48–49. 

Aleiodes apiculatus (Fahringer), ♀, England. 48 habitus lateral 49 mummy of Euproctis similis (Fuessly).

Figures 50–62. 

Aleiodes apiculatus (Fahringer), ♀, England. 50 wings 51 mesosoma lateral 52 propodeum and anterior half of metasoma dorsal 53 hind leg lateral 54 outer hind claw lateral 55 fore femur lateral 56 antenna 57 apical segments of antenna 58 basal segments of antenna 59 head anterior 60 head lateral 61 head dorsal 62 base of first tergite dorsal.

Redescribed ♀ (NMS) from England, length of fore wing 5.9 mm, of body 6.8 mm.

Head. Antennal segments of ♀ 47, length of antenna 1.2 × fore wing, its subapical segments 1.6 × as long as wide; frons granulate, with satin sheen and some rugae; OOL and POL 0.5 and 0.6 × width of posterior ocellus, respectively; stemmaticum strongly protruding; vertex rugulose-granulate, with satin sheen; clypeus convex and punctulate-coriaceous; ventral margin of clypeus thick and convex (Fig. 59); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 59) and face mainly granulate with transverse rugulae; length of eye 3.4 × temple in dorsal view and temple moderately narrowed behind eye; occiput behind stemmaticum rugulose-granulate; occipital carina reduced medio-dorsally and complete ventrally, without crenulae and dorsally curved (Fig. 61); clypeus above lower level of eyes (Fig. 59); length of malar space 0.30 × height of eye in lateral view; eyes rather protruding (Figs 59–61).

Mesosoma. Length of mesosoma 1.7 × its height; mesoscutal lobes finely granulate, matt, but medio-posteriorly irregularly rugose and anteriorly high; notauli medium-sized and crenulate; prepectal carina medium-sized, remaining separate far from anterior border; precoxal area of mesopleuron and area above it distinctly rugose; remainder of mesopleuron (including speculum) granulate and with satin sheen (Fig. 51); metapleuron distinctly granulate and with satin sheen; mesosternal sulcus shallow and sparsely crenulate; mesosternum rather angulate latero-posteriorly but rounded medially; scutellum slightly convex, finely granulate, and antero-laterally with carina; propodeum weakly convex, without tubercles, anteriorly granulate, medially coarsely rugose and posteriorly with longitudinal carinae, median carina complete.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 50); 1-CU1 slightly oblique, 0.5 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.45 × 3-SR; second submarginal cell rather long (Fig. 50); cu-a slightly inclivous, not parallel with CU1b, straight (Fig. 50); 1-M straight posteriorly and 1-SR angled to 1-M. Hind wing: marginal cell parallel-sided, but slightly narrowed submedially; 2-SC+R medium-sized, slender; m-cu short and only slightly pigmented; M+CU:1-M = 4:3; 1r-m 0.7 × 1-M.

Legs. Tarsal claws rather small and with distinct fine pecten (Fig. 54); hind coxa finely granulate, with satin sheen and 0.8 × as long as first tergite; hind trochantellus 2.6 × longer ventrally than wide; length of fore and hind femora 6.0 and 4.6 × their width, respectively; inner apex of hind tibia without distinct comb; length of inner hind spur 0.35 × hind basitarsus.

Metasoma. First tergite 1.1 × as long as wide posteriorly, stout, convex anteriorly and latero-anteriorly distinctly lamelliform; first and second tergites densely coarsely longitudinally rugose (Fig. 52), third tergite basally rugulose and with distinct median carina up to middle of third tergite; medio-basal area of second tergite minute; second suture rather wide and crenulate; remainder of third tergite granulate and following tergites shiny and superficially granulate; apical half of third and fourth tergites without sharp lateral crease; ovipositor sheath densely setose and apically truncate.

Colour. Brownish yellow; scapus and pedicellus of ♀ at least partly blackish, contrasting with yellowish middle of antenna and antenna of ♀ in dorsal view bicoloured, first–fifth[–eighth] and 2–3 apical segments more or less dark brown, remainder of antenna yellowish; malar space, mandible, palpi, tegulae, pronotum anteriorly, basal half of pterostigma, trochanters and trochantelli, fore and middle coxae, and ventral half of metasoma ivory or pale yellow; face medially, frons and vertex medially, stemmaticum, occiput dorsally, mesoscutum laterally narrowly, scutellar sulcus, axilla, scutellum posteriorly, metanotum, third (except antero-lateral corner)–sixth tergites, fourth–sixth sternites and ovipositor sheath black or dark brown; telotarsi slightly infuscate; veins and apical half of pterostigma dark brown; wing membrane slightly infuscate.

Variation. Antennal segments of ♀ 46(1), 47(10), 48(4), 49(1); ♂ 41(1), 42(2), 43(8), 44(3), 45(6), 46(2). Length of fore wing 5–6 mm, of body 5–7 mm. Males are brownish yellow, but stemmaticum black and antenna apically, occiput dorsally, mesoscutum laterally, scutellum posteriorly, metanotum, propodeum medially, first tergite except posteriorly and second tergite laterally somewhat infuscate; malar space, palpi, tegulae, pronotum, fore and middle coxae, trochanters and trochantelli, first tergite medio-apically and middle of second tergite pale yellowish.

Similar to A. pallidator (Thunberg), but the latter differs by having the tarsal claws only bristly setose, the hind trochantellus ventrally 2.2 × as long as wide, the antennal segments of ♀ with 51–57 segments; the stemmaticum less protuberant, the pterostigma yellow and the body of ♀ entirely brownish yellow. The extent of dark colouration is highly variable, and is often poorly developed in the British population. In males especially, the colour (including scape and even stemmaticum) can be rather uniform orange to light honey-brown. Because it can lack the colour characters usually plain in females, the male of this species can superficially resemble some of the relatively large orange species with big ocelli and antennal segments in the range 41–48 that fall into the residual circumscriptus-group not dealt with in this paper. Good recognition characters for male A. apiculatus include its somewhat bristly antenna and legs, its enlarged fifth tarsal segment (especially in the fore leg), its relatively strongly sculptured second metasomal tergite with weak mediolateral depressions, its weakly pectinate claws, and the stronger (though weak) development of a comb at the apex of the hind tibia. The synonymy with Aleiodes negativus (Tobias) is accepted; the examined females of A. apiculatus have the antenna with 46–49 segments (the holotype has 49 segments). The examined paratype of A. negativus (BMNH) was reared from E. similis and has 47 antennal segments. According to Tobias (1961)A. negativus female types (including the holotype) should have 35–38 antennal segments and the male types 42–43 segments; most likely the antennal counts for the female types given by Tobias result from a lapsus or typographical error and should be 45–48.

Lectotype, ♀ (ZIL), “Lpl” [= Lapland, North Sweden]).

3 ♀, 2 ♂ (G. Várkonyi personal coll., NMS) Finland, Ks. Salla 752.61, Värriö H, ex Pygmaena fusca, collection dates (of host larva) between 15.vi and 28.vi.1995, G. Várkonyi; 1 ♂ (G. Várkonyi personal coll.) Finland: Ks. Salla 752.61 Värriö H 21.vi.1995, G. Várkonyi; 1 ♀ (BMNH) Finland, Kuusano, Mäntytunturi, on snow, 29.vi.1935, G.J. Kerrich; 1 ♂ (NMS) S. Norway, Jotunheimen, Giendersheim, 1000–1500 m, 7.vii.[1966], J.E. & R.B. Benson; 2 ♂ (BMNH) S. Norway, Buskerud Fylke Geilo, 1000–1100 m, 16–24.vi.1965, J.E. & R.B. Benson; 1 ♂ (RMNH) Norway, Oppdal, Kongsvoll, Vestbekken, 28.vi.1978, J.O. Solem; 1 ♀ (BMNH) Sweden, Torne Lappmark, Tornehamn, 4.vii.1954, J.E. & R. B. Benson; 1 ♀, 1 ♂ (BMNH, NMS) Switzerland, Grisons, Müstairtal, Funt da S. Charl, 2400 m, 9.vi 1960 and 3.vii.1960, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Grisons, Engadine National Park, 2380 m, 1.vii.1960, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Valais, Arolla, 8000 ft, 9.vii.1935, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Valais, near Verbier, 8000–8500 ft, 27.vi.1959, J.E. & R.B. Benson.

This boreoalpine species is univoltine, passing the winter as a mummy. The only known host is the ennomine (Macariini) geometrid Pygmaena fusca (Thunberg) (5:1; G. Várkonyi/Finland), which feeds on Empetrum and Vaccinium (G. Várkonyi personal communication) and probably occurs throughout the range of the parasitoid. The small mummy (Fig. 64) is short, broad and dorsally elevated.

Figures 63–64. 

Aleiodes arcticus (Thomson), ♀, Switzerland. 63 habitus lateral 64 mummy of Pygmaena fusca (Thunberg) from Finland.

Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 70); OOL 1.9 × diameter of posterior ocellus; mesoscutum, orbita and malar space black; precoxal sulcus largely granulate; trochanters, trochantelli and pterostigma largely black(ish); mesoscutum without a longitudinal carina on mesoscutum medio-posteriorly; apical half of marginal cell of hind wing parallel-sided or slightly widened; vein M+CU1 of fore wing apically at about same level as vein 2-CU1 (Fig. 65); vein r of fore wing 0.6–0.9 × vein 3-SR (Figs 63, 65); vein 1-SR of fore wing linear with vein 1-M (Fig. 65); all femora and tibiae reddish or yellowish brown; fore and hind femora moderately stout (Figs 68–69); fourth metasomal tergite curved posteriorly in dorsal view (Fig. 67), lateral crease distinct and following tergites more or less retracted; length of fore wing 3.4–3.9 mm.

Figures 65–75. 

Aleiodes arcticus (Thomson), ♀, Switzerland. 65 wings 66 mesosoma lateral 67 metasoma dorsal 68 hind leg lateral 69 fore femur lateral 70 head anterior 71 head lateral 72 head dorsal 73 antenna 74 apical segments of antenna 75 basal segments of antenna.

Redescribed ♀ (BMNH) from Müstairtal (Switzerland), length of fore wing 3.7 mm, of body 4.6 mm.

Head. Antennal segments 40, length of antenna as long as fore wing, its subapical segments about 1.7 × as long as wide; frons mainly superficially granulate and with some rugulae anteriorly, weakly shiny; OOL 1.9 × diameter of posterior ocellus and granulate as vertex, with satin sheen; clypeus moderately convex, narrow and coriaceous; ventral margin of clypeus thick and depressed (Fig. 70); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 70) and face mainly coriaceous with some rugulae dorsally and long setae; length of eye 1.1 × temple in dorsal view and temple subparallel-sided behind eye; occiput behind stemmaticum granulate and occipital carina absent ventrally and narrowly interrupted dorsally; clypeus below lower level of eyes (Fig. 70); length of malar space 0.5 × height of eye in lateral view; eyes somewhat protruding (Figs 70–72).

Mesosoma. Mesoscutal lobes largely granulate-coriaceous, matt and medio-posteriorly rugose, middle lobe without a longitudinal carina; notauli narrow, shallow and very finely crenulate; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) granulate, but dorsally finely rugose; medially metapleuron granulate and with some rugae, rather shiny; mesosternal sulcus narrow and rather deep, with longitudinal carina posteriorly; mesosternum rounded posteriorly; scutellum moderately convex, mainly granulate and largely non-carinate laterally; propodeum rather directly lowered posteriorly and granulate-rugose, median carina complete, without tubercles.

Wings. Fore wing: r 0.7 × 3-SR (Fig. 65); 1-CU1 subhorizontal, 0.1 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.8 × 3-SR; second submarginal cell rather small (Fig. 65); vein M+CU1 of fore wing apically at about same level as vein 2-CU1 (Fig. 65); vein 1-SR of fore wing linear with vein 1-M; cu-a subvertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 65); 2-SC+R short and longitudinal; m-cu present, pigmented.

Legs. Tarsal claws setose; hind coxa granulate-coriaceous, with satin sheen; hind trochantellus twice longer ventrally than wide; length of fore and hind femora 5.5 and 4.0 × their width, respectively (Figs 68–69); inner apex of hind tibia without comb; length of inner hind spur 0.4 × hind basitarsus.

Metasoma. First tergite 0.7 × as long as wide posteriorly, convex and latero-posteriorly non-lamelliform; first–second tergites finely and densely irregularly rugulose and with median carina (Fig. 67); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite with median carina (except posteriorly), third–fourth tergites very finely rugulose-coriaceous; fourth tergite convex medially and apically; fourth tergite with sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose.

Colour. Black (including coxae); palpi basally, tegulae, pterostigma, veins, trochanters and trochantelli dark brown; remainder of palpi and legs yellowish brown; wing membrane slightly infuscate.

Variation. Antennal segments of ♀ 39(1), 40(2), 41(0), 42(1), of ♂ 38(1), 39(2), 40(2); length of fore wing 3.4–3.7 mm; maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face; vein r of fore wing 0.6–0.8 × vein 3-SR; median carina of middle mesoscutal lobe absent or weakly indicated; legs (except basally) vary from largely yellowish brown to largely dark brown with base of hind femur and tibiae paler than remainder of legs; second submarginal cell of fore wing rather variable in shape, but some are as trapezoidal as in A. reticulatus, with which this species is closely related.

Recorded as British by Morley (1916), but in error as the three specimens (CMIM) on which the record was based have been examined and prove to belong to A. similis (Curtis). A series from Austria (Hohe Tauern, various altitudes ca 2300 m (MSC)) and also specimens from similar elevations in Switzerland (BMNH) have a habitus similar to A. arcticus but differ considerably from our concept of A. arcticus in being more coarsely rugose (including mesopleuron), the head being longer and behind the eyes narrower, outer orbits brownish (i. e. lighter in colour than the rest of the temple), antennal segments longer, second cubital cell usually longer. It is considered to be a different species placeable in the bicolor-group, but with apical tergites retracted and hind coxa short.

Holotype, ♀ (NMS, Edinburgh), “France, Olonne, ex Macaria artesiaria, mummy coll[ected] on Salix repens, vi.1984, N. Hall”.

Apart from the host (determined as a result of adults of Macaria artesiaria (Denis & Schiffermüller) (Geometridae) being reared from caterpillars morphologically corresponding to the mummy and collected at the same time), nothing is known of the biology of this species. The holotype was excavated (fully formed but dead) from the mummy more than a year after it had been collected in apparently freshly made condition on a twig of its foodplant.

Maximum width of hypoclypeal depression 0.3 × minimum width of face (Fig. 84); OOL 2.4 × diameter of posterior ocellus; mesoscutum, orbita and malar space yellowish brown; precoxal sulcus largely granulate; trochanters, trochantelli and pterostigma largely dark brown or black(ish); mesoscutum without a longitudinal carina medio-posteriorly; apical half of marginal cell of hind wing parallel-sided or slightly widened; vein M+CU1 of fore wing apically above level of vein 2-CU1 (Fig. 78); vein r of fore wing about 0.7 × vein 3-SR (Figs 76, 78); vein 1-SR of fore wing angled with vein 1-M (Fig. 78); all femora and tibiae dark brown; fore and hind femora moderately stout (Figs 87–88); fourth metasomal tergite curved posteriorly in dorsal view (Fig. 81), lateral crease distinct and following tergites more or less retracted; length of fore wing about 3 mm. Very similar to A. reticulatus (Noskiewicz) but the latter differs by the black mesoscutum, the reddish or yellowish brown femora and tibiae, the less slender fore and hind femora (Figs 313–314 vs 87–88) and by having vein 1-SR of fore wing linear with vein 1-M (Fig. 305 vs 78).

Figures 76–77. 

Aleiodes artesiariae sp. n., ♀, holotype. 76 habitus lateral 77 mummy of Macaria artesiaria (Denis & Schiffermüller) after extraction of adult.

Figures 78–88. 

Aleiodes artesiariae sp. n., ♀, holotype. 78 wings 79 mesosoma dorsal 80 mesosoma lateral 81 metasoma dorsal 82 antenna (incomplete) 83 basal segments of antenna 84 head anterior 85 head lateral 86 head dorsal 87 hind leg lateral 88 fore femur lateral.

Holotype, ♀, length of fore wing 2.9 mm, of body 3.6 mm.

Head. Antennal segments 33+ (incomplete), length of antenna at least as long as fore wing, its subapical segments somewhat longer than wide; frons mainly superficially granulate, moderately shiny; OOL 2.4 × diameter of posterior ocellus and granulate as vertex, with satin sheen; clypeus moderately convex, narrow and coriaceous, 0.4 × width of face; ventral margin of clypeus thick and depressed (Fig. 84); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 84) and face mainly coriaceous with some rugulae dorsally and long setae; length of eye 1.8 × temple in dorsal view and temple sub-parallel-sided behind eye and narrowed posteriorly; occiput behind stemmaticum granulate and occipital carina absent ventrally and complete (but irregular) dorsally; clypeus partly up to lower level of eyes (Fig. 84); length of malar space 0.4 × height of eye in lateral view; eyes slightly protruding (Figs 84–86).

Mesosoma. Mesoscutal lobes largely granulate-coriaceous, matt and medio-posteriorly rugose, middle lobe without a longitudinal carina; notauli narrow, shallow and very finely crenulate; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) granulate, but dorsally rugose; medially metapleuron granulate, rather shiny; mesosternal sulcus narrow and rather deep, without longitudinal carina posteriorly; mesosternum rather angulate posteriorly; scutellum moderately convex medially and depressed laterally, mainly granulate and largely non-carinate laterally; lunula moderately wide; propodeum rather directly lowered posteriorly and granulate-rugose, median carina complete, without tubercles.

Wings. Fore wing: r 0.7 × 3-SR and linear with 3-SR (Fig. 78); 1-CU1 oblique and widened, 0.25 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.7 × 3-SR; second submarginal cell rather small and square (Fig. 78); vein M+CU1 of fore wing apically above level of vein 2-CU1 (Fig. 78); vein 1-SR of fore wing angled with vein 1-M; cu-a subvertical, not parallel with CU1b, straight; 1-M straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 78); 2-SC+R medium-sized and longitudinal; m-cu present and slightly pigmented.

Legs. Tarsal claws setose; hind coxa granulate-coriaceous, with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 5.4 and 5.2 × their width, respectively (Figs 87–88); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus.

Metasoma. First tergite 0.7 × as long as wide posteriorly, convex and non-lamelliform latero-posteriorly and basally; first–second tergites finely and densely irregularly rugulose and with median carina (Fig. 81); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite superficially coriaceous and with median carina (except posteriorly), third–fourth tergites very finely rugulose-coriaceous; fourth tergite convex medially and apically, shiny and with sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose.

Colour. Black (including coxae, middle and hind trochanters); palpi, tegulae, pterostigma, veins, first and second tergites and remainder of legs dark brown; malar space, orbita, mesoscutum (but middle lobe somewhat infuscate medio-anteriorly), scutellum laterally, pronotum, mesopleuron (except postero-ventrally) yellowish brown; wing membrane slightly infuscate.

From the specific epithet of its host.

*France.

Lectotype here designated, ♀ (NRS), “Färöar [= Faroe Isl., Denmark], Klinck”, “Triangisvaag”, “♀ Aleiodes circumscriptus var. bistrigatus Roman, C. van Achterberg, 1984. Lectotype”, “178, 84”, “Riksmuseum Stockholm”, “NHRS-HEVA 000003802”. Paralectotypes: 3 ♂ (NRS) with same locality labels as lectotype.

None.

Unknown.

Apical half of hind femur (partly) dark brown, darker than hind trochanter and trochantellus; face with distinct rugae; antenna of female with 39 segments and third segment stout, 4^th–7^th segments moderately stout (but less than in A. diarsianae; Fig. 164), of male with 38 segments; OOL 1.7 × diameter of posterior ocellus (Fig. 97); clypeus 0.4 × as wide as face (Fig. 95); mesopleuron black dorso-posteriorly (Fig. 90); malar space and inner orbita dark brown and temple near eye (= external orbita) reddish-brown; first tergite strongly widened apically; eye (of male) elongate in lateral view; middle third of hind femur yellowish brown or dark brown; vertex moderately setose, rather shiny and blackish posteriorly; mesosternum dark brown. Close to A. borealis (Thomson, 1892), but this species has less antennal segments (♀: 32–34 segments), palpi and legs more or less infuscate and the clypeus wider (about 0.5 × width of the face). The shape of the subbasal antennal segments is similar to that of series from Scandinavia mentioned under A. diarsianae, but this series has the temples directly narrowed behind the eyes and the males have 42–47 antennal segments and females 44–45. Similar to the A. pictus-aggregate (e.g. A. pictus (Herrich-Schäffer, 1838) and A. nigriceps Wesmael, 1838), but these are usually smaller and less robust species, having the face without distinct rugae or only a few rugae dorsally, the mesosternum usually widely orange-brown (and the mesopleuron usually without rugae in A. nigriceps), the malar space partly or completely yellowish brown, the first tergite less widened apically, the fore and hind tarsi comparatively slender, the eye normal in lateral view and the antenna less robust.

Figures 89–98. 

Aleiodes bistrigatus Roman, ♀, lectotype. 89 wings 90 mesosoma lateral 91 basal half of metasoma dorsal 92 hind leg lateral 93 hind femur lateral 94 fore femur lateral 95 head anterior 96 head lateral 97 head dorsal 98 basal segments of antenna.

Lectotype, ♀, length of fore wing 4.0 mm, of body 5.5 mm.

Head. Antennal segments of ♀ 39, length of antenna 1.3 × fore wing, its subapical segments about 1.9 × as long as wide and third segment stout (Fig. 98); frons coriaceous and posteriorly rugulose, with satin sheen; OOL 1.7 × diameter of posterior ocellus and coriaceous; vertex rugulose-coriaceous, rather shiny; clypeus slightly convex, indistinctly sculptured; ventral margin of clypeus rounded and depressed (Fig. 95); width of hypoclypeal depression 0.38 × minimum width of face and face distinctly transversely rugose (Fig. 95); length of eye 1.6 × temple in dorsal view and temple gradually narrowed behind eye (Fig. 97); occiput behind stemmaticum coriaceous and with some rugulae, occipital carina interrupted dorsally; clypeus partly above lower level of eyes and 0.4 × as wide as face (Fig. 95); length of malar space 0.6 × length of eye in lateral view; eyes slightly protruding (Figs 95–97).

Mesosoma. Mesosoma 1.7 × as long as high; mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli complete and moderately wide, weakly crenulate and posteriorly widened and rugose; prepectal carina medium-sized and lamelliform, reaching anterior border; precoxal area of mesopleuron very coarsely rugose, connected to rugosity of dorso-anterior part of mesopleuron; speculum nearly smooth and shiny (Fig. 90); metapleuron granulate, matt and posteriorly rather tuberculate; mesosternal sulcus narrow and deep; mesosternum rounded posteriorly; scutellum elongate, slightly convex, coriaceous and laterally largely carinate; propodeum rather flat dorsally, laterally and apically rather rugose, anteriorly only weakly so, median carina complete, but posteriorly irregular.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 89); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.65 × 2-SR, and 0.40 × 3-SR; second submarginal cell medium-sized (Fig. 89); cu-a vertical, not parallel with CU1b, straight; 1-M slightly curved posteriorly and not continuous with 1-SR. Hind wing: apical half of marginal cell parallel-sided or nearly so; 2-SC+R short and longitudinal; m-cu present; 1r-m distinctly oblique and 0.7 × 1-M.

Legs. Tarsal claws setose; hind coxa coriaceous but partly superficially rugulose, largely matt; hind trochantellus twice longer ventrally than wide; length of fore and hind femora 5.4 and 5.0 × their width, respectively (Figs 93–94); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus.

Metasoma. First tergite 0.9 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope comparatively wide (Fig. 91); first–third tergites densely and distinctly longitudinally rugose, robust (Fig. 91), with distinct median carina; medio-basal area of second tergite absent; second tergite 1.5 × as long as third tergite; second suture moderately impressed and crenulate; remainder of metasoma largely superficially coriaceous; fourth and apical fifth of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) densely setose.

Colour. Black or brownish black; antenna yellowish brown, but scapus dorsally and apical seventh of antenna dark brown; palpi, temple near eyes, legs (except infuscate subapical part of hind femur), tegulae, longitudinal stripe on mesopleuron, mesoscutum posteriorly, metasoma baso-ventrally, first tergite medio-apically, middle of second tergite and third tergite medio-basally largely yellowish; veins and pterostigma (except yellowish basal third and centrally) dark brown; border between dark and pale part of pterostigma diffuse (Fig. 89); wing membrane subhyaline.

Variation. The male paralectotypes are very similar to the lectotype; one has a complete antenna with 38 segments and most of the hind femur darkened. One paralectotype has the hind coxa completely yellowish and the mesopleuron less coarsely rugose, but other paralectotypes have the mesopleuron coarsely sculptured and the hind coxa largely infuscate.

Possibly a Faroe Islands endemic.

Lectotype here designated, ♀ (BMNH), “2504” [on card], “cotype”, “[England,] New Forest, 4.v.1914, ex Semiothisa liturata, G.T. Lyle”, “G.T. Lyle Coll., B.M. 1930-579”, “Rhogas cantherius Lyle”. Paralectotypes: 4 ♀ + 3 ♂ (BMNH, CMIM), topotypic and from same host, but one non-reared paralectotype from Harwood collection.

*Austria, British Isles (England: V.C.s 11, 17, 19, 22, 24, 25, 28, 56), *Germany, *Netherlands (Breda; Melissant; Wageningen), *Russia, Slovakia, *Sweden. . Specimens in NMS, BMNH, RMNH, ZISP, FRAH, CC, CMIM, SDEI.

MRS777 (Sweden KU682249, CO1)), MRS787 (Sweden KU682253, CO1).

A parasitoid of conifer-feeding Macaria species (Geometridae), overwintering as a mummy. Specimens (in NMS unless specified) reared from Macaria liturata (Clerck) (22 [6 are BMNH, 3 CMIM, 2 SDEI, 1 FRAH]; P.E. Hatcher, G.T. Lyle, M.R. Shaw/England); Macaria signaria (Hübner) (2:2 [CC]; M. Čapek/Slovakia). Additionally 5 reared specimens, fortunately accompanied by the host mummy, had been labelled as reared from other conifer-feeding geometrids (Bupalus piniaria (Linnaeus) (2, ZISP), Eupithecia indigata (Hübner) (2, FRAH), Hylaea fasciaria (Linnaeus) (1, ZISP)) but in all cases examination of the host remains established that the host was in fact more consistent with a species of Macaria. Plurivoltine, overwintering in a mummy constructed on a conifer needle. The adult occurs in the field from May until well into September, and it is clear from the rearing data that its conifer-feeding Macaria hosts are likely to be sought across all of their foodplants (Abies cephalonica, Larix decidua, Pinus strobus, Pinus sylvestris and Pseudotsuga menzieseii are indicated on the data labels overall). No experimentation has been undertaken.

Antennal segments of ♀ 39–43, of ♂ 40–43; head strongly narrowed behind eyes (Fig. 110), yellowish anteriorly and mainly dark brown dorsally; OOL 0.6 × diameter of posterior ocellus; length of malar space of ♀ 0.2–0.3 × height of eye in lateral view (Fig. 109); scapus in lateral view rather oblique apically; occipital carina interrupted dorsally and complete ventrally (Fig. 109); eye 3.5–6.5 × as long as temple in dorsal view (Fig. 110); mesosternum and precoxal sulcus superficially granulate and with satin sheen; vein 2-CU1 of fore wing about 3 × vein 1-CU1 (Fig. 101); vein 1-SR narrow and linearly connected to vein 1-M and vein 1-M straight (Fig. 101); hind femur stout (Fig. 104); fourth metasomal tergite largely (superficially) coriaceous and shiny; length of fore wing 4–5 mm.

Figures 99–100. 

Aleiodes cantherius (Lyle), ♀, England. 99 habitus lateral 100 mummy of Macaria sp.

Figures 101–112. 

Aleiodes cantherius (Lyle), ♀, England. 101 wings 102 mesosoma lateral 103 propodeum and metasoma dorsal 104 hind leg lateral 105 antenna 106 basal segments of antenna 107 mesosoma dorsal 108 head anterior 109 head lateral 110 head dorsal 111 fore femur lateral 112 apical segments of antenna.

Redescribed ♀ (NMS) from Santon Downham (England), length of fore wing 4.6 mm, of body 5.0 mm.

Head. Antennal segments 43, length of antenna 1.3 × fore wing, its subapical segments about 2.3 × as long as wide (Fig. 112) and scapus in lateral view rather oblique apically; frons granulate and rather shiny; OOL and POL 0.6 and 0.8 × diameter of posterior ocellus, respectively and granulate; vertex granulate, dull; clypeus rather high, convex, coriaceous; ventral margin of clypeus thick (Fig. 108); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 108) and face mainly transversely rugose and granulate; length of eye 6.5 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum mainly granulate and occipital carina interrupted medio-dorsally and complete ventrally (Fig. 109); clypeus above lower level of eyes (Fig. 108); length of malar space 0.3 × height of eye in lateral view; eyes protruding (Fig. 109).

Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen, but medio-posteriorly with some rugae; notauli narrow, shallow and largely smooth; prepectal carina rather lamelliform medio-ventrally, nearly reaching anterior border of mesopleuron and latero-ventrally curved; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) superficially granulate, but dorsally rugulose; medially metapleuron superficially granulate, rather shiny; mesosternal sulcus narrow and rather deep, micro-crenulate, without carina posteriorly; mesosternum rather angulate posteriorly; scutellum finely coriaceous and non-carinate laterally; dorsal face of propodeum medium-sized, convex and rugulose, but posteriorly with some carinae and smooth in between and anteriorly mainly granulate, median carina complete, without tubercles.

Wings. Fore wing: r 0.5 × 3-SR (Fig. 101); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell elongate (Fig. 101); 1-SR slightly angled to 1-M and slender; cu-a rather inclivous, not parallel with CU1b, straight; 1-M slightly curved. Hind wing: apical half of marginal cell parallel-sided or nearly so (Fig. 101); 2-SC+R longitudinal; m-cu present as fold, unpigmented; M+CU:1-M = 3:2; 1r-m 0.7 × 1-M.

Legs. Tarsal claws yellowish setose; hind coxa superficially finely coriaceous, rather shiny; hind trochantellus 2.4 × longer ventrally than wide; length of fore and hind femora 6.2 and 4.0 × their width, respectively (Figs 104, 111); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus.

Metasoma. First tergite 1.2 × as long as wide posteriorly, flattened and latero-anteriorly narrowly lamelliform; first–second tergites and base of third tergite densely finely longitudinally rugose and with median carina; second tergite stout, 0.8 × longer than wide basally and 1.2 × as long as third tergite (Fig. 103); medio-basal area of second tergite minute; second suture deep and distinctly crenulate; remainder of metasoma largely superficially coriaceous and rather shiny; apical half of third and fourth tergite without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate.

Colour. Black or dark brown; palpi, pronotum postero-dorsally and tegulae pale yellowish; scapus and pedicellus ventrally (but dorsally more or less darkened), orbita, two stripes on mesoscutum, legs (but hind coxa more or less dark brown), first tergite medio-apically, second tergite (except postero-lateral corners), third and following tergites mainly yellowish brown (Figs 99, 103); pterostigma and veins dark brown, but base of pterostigma and vein 1-R1 of fore wing brownish yellow; wing membrane slightly infuscate.

Variation. Length of fore wing 3.6–4.5 mm, of body 3.8–4.6 mm; antennal segments of ♀ 39(1), 40(2; one is lectotype), 41(6), 42(2), 43(4), of ♂ 39(1), 40(6), 41(8), 42(3), 43(3); specimens have a characteristic pair of more or less obscure dorsal orange brown marks on the otherwise dark mesoscutum. Males examined have the metasoma dark brown apically, hind tibia (except ivory base) and tarsus more or less infuscated.

The two sexes have about the same number of antennal segments.

Holotype, ♀ (NMS, Edinburgh), “[France:] Corsica: Corte, Val de Restonica (Hôtel Colonna), 500 m, [at] light, 29.vii–3.viii.[20]01, M.R. Shaw”, “MRS Aleiodes DNA 102”. Paratypes (11 ♀ + 23 ♂): 1 ♀ + 1 ♂ (NMS, RMNH), same data as holotype; 1 ♀ (BMNH), “[Spain:] Mallorca, Sa Roca, P.N. de s’Albufera, MV light, 2–27.ix.2013, M.R. Honey BMNH(E) 2013-158”; 1 ♀ (NMS), “Spain: Zaragoza Prov., Los Monegros, Retuerta de Pina, 30TYL 27.94, J. Blasco-Zumeta, 5104, 8.viii.[19]92, NMSZ1997.026, swept from Suaeda vermiculata”; 2 ♀ + 1 ♂ (NMS, RMNH), id., but 28.vi.1992 and swept from Suaeda vera, ♂ swept from Osyris alba; 3 ♀ (NMS, RMNH), id., but 12.ix.1991 (1) or 10.ix.1993 (2) and collected at light; 2 ♂ (NMS, RMNH), id., but 10.vii.1993, at light; 9 ♂ (NMS, RMNH), id., but 10.ix.1993; 3 ♂ (NMS, RMNH), id., but 20.ix.1993; 2 ♂ (NMS, RMNH),, id., but ?1991; 2 ♂ (NMS), id., but 20.vii.1993; 1 ♂ (NMS), id., but 20.viii.1993; 2 ♀ (FC), “Esp.: Valencia, El Saler (Casal d’Esplai), T.M., 20–27.vii.1992 & 17–24.viii.1992, J.V. Falcó y F. Luna”; 2 ♂ (FC), “Esp.: Valencia, Moncada-TM blanca, 6–13.vii.1992 & 13–20.vii.1992, M.J. Verdú”; 1 ♀, (NMS), “[Spain:] Canary Islands, Tenerife, Anco Viejo, La Sabinita, 20.iii.1999, R.R. Askew”.

MRS055 (Corsica JF962818, CO1), MRS098 (Corsica KU682224, CO1), MRS102 (Corsica KU682225, CO1).

Unknown. This species is active at night and occurs in open habitats suggesting that its hosts live in low vegetation, but its voltinism is unclear.

Head weakly transverse (Fig. 122); body slender and entirely brownish yellow; antenna of ♀ (except scapus) dark brown; antennal segments of ♀ 34–37, of ♂ 35–40; eye rather small (Figs 121–123)); OOL equal to width of posterior ocellus; length of malar space of ♀ 0.5 × height of eye in lateral view (Fig. 123); speculum of mesopleuron smooth and shiny or superficially granulate; propodeum slightly elongate (Fig. 115); fore wing rather narrow (Fig. 114); vein m-cu of fore wing straight and angled to vein 2-CU1 (Fig. 114); pterostigma pale yellowish basally; hind coxa distinctly shorter than first tergite; fore wing subhyaline; apex of hind tibia with comb at inner side (Fig. 119); hind femur 5 × as long as its maximum width; hind trochantellus slender (Fig. 117); dorsal carinae of first metasomal tergite lamelliform protruding basally; second tergite without triangular area medio-basally (Fig. 116); third tergite weakly sculptured; fourth tergite without sharp lateral crease, fourth and following tergites partly retracted and largely smooth. Resembles A. testaceus (Telenga, 1941), but A. testaceus has no apical comb of the hind tibia (present in A. carminatus); surrroundings of precoxal sulcus largely smooth and shiny (mainly granulate and moderately shiny); veins 1-SR and r of fore wing longer (shorter); antenna yellowish brown basally (dark brown) and length of malar space 0.3–0.4 × height of eye in lateral view (0.5 ×). Resembles superficially A. curticornis nom. n., but A. curticornis has no apical comb of the hind tibia (present in A. carminatus); fore femur, third and penultimate antennal segments robust (slender) and antenna yellowish brown basally (dark brown). The presence of a hind tibial comb distinguished it from all the pale members of the A. circumscriptus group not treated in this paper.

Figure 113. 

Aleiodes carminatus sp. n., ♀, holotype, habitus lateral.

Figures 114–124. 

Aleiodes carminatus sp. n., ♀, holotype. 114 wings 115 mesosoma lateral 116 propodeum and metasoma dorsal 117 hind leg lateral 118 fore femur lateral 119 inner side of hind tibial apex lateral 120 basal segments of antenna 121 head anterior 122 head dorsal 123 head lateral 124 apical segments of antenna.

Holotype, ♀, length of fore wing 3.4 mm, of body 3.9 mm.

Head. Antennal segments of ♀ 35, length of antenna 1.2 × fore wing, its subapical segments about 1.7 × as long as wide; frons rugulose-granulate, with satin sheen; OOL and POL 1.0 and 0.8 × width of posterior ocellus, respectively; vertex granulate, rather dull and distinctly depressed near ocelli; clypeus convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 121); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 121) and face coriaceous with superficial rugulae; length of eye 2.4 × temple in dorsal view and temple roundly narrowed behind eye; occiput behind stemmaticum coriaceous with satin sheen; occipital carina widely interrupted medio-dorsally and ventrally weak and irregular (Figs 122–123); clypeus partly above lower level of eyes (Fig. 121); length of malar space 0.5 × height of eye in lateral view; eyes protruding (Fig. 122).

Mesosoma. Mesoscutal lobes coriaceous-granulate, with satin sheen, but medio-posteriorly longitudinally rugose and anteriorly steep; notauli obsolescent; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron (except posteriorly) and mesopleuron antero-dorsally distinctly rugose; remainder of mesopleuron (but speculum partly smooth and shiny) granulate and dull; metapleuron largely granulate, matt; mesosternal sulcus shallow and largely smooth; mesosternum rounded posteriorly; scutellum flat, granulate, and laterally with distinct carina, lunula narrow and parallel-sided; propodeum convex, without tubercles, rugulose anteriorly and remainder rugose, median carina complete.

Wings. Fore wing: r 0.7 × 3-SR (Fig. 114); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Fig. 114); cu-a slightly inclivous, nearly parallel with CU1b, straight; 1-M nearly straight and 1-SR distinctly angled with 1-M. Hind wing: apical half of marginal cell slightly widened; 2-SC+R short; m-cu absent.

Legs. Tarsal claws with yellow setae; hind coxa rugulose and with spaced oblique rugae, with satin sheen and 0.8 × as long as first tergite; hind trochantellus 2.8 × longer ventrally than wide (Fig. 117); length of fore and hind femora 6.6 and 4.8 × their width, respectively (Figs 117–118); inner apex of hind tibia with distinct comb (Fig. 119); length of inner hind spur 0.2 × hind basitarsus.

Metasoma. First tergite as long as wide posteriorly, convex anteriorly and dorsal carinae lamelliform protruding basally; first and second tergites longitudinally striate, robust (Fig. 116), with distinct median carina; medio-basal area of second tergite absent; second suture narrow and crenulate; third tergite largely longitudinally rugulose, but smooth posteriorly; third tergite with complete sharp lateral crease but this absent from following tergites; ovipositor sheath ventrally densely setose and remainder smooth, shiny and apically acute.

Colour. Yellowish brown; antenna (except scapus and pedicellus ventrally), ovipositor sheath and most of ventral part of metasoma dark brown; stemmaticum black; tegulae, pronotum partly and legs brownish yellow; veins brown; pterostigma pale yellowish, but slightly darkened laterally; wing membrane subhyaline.

Variation. Antennal segments of ♀: 34(5), 35(3), 36(2), 37(2), of ♂: 35(1), 36(1), 37(8), 38(3), 39(1), 40(2). In many specimens fore wing 2-SR is strikingly longer than r-m, but in others this is less distinctive. Hind femur sometimes brown, 4.5–4.9 × as long as wide and hind trochantellus 2.6–2.9 × longer ventrally than wide; occipital carina ventrally sinuate and reduced or complete; colour of body varies from nearly completely yellowish brown to largely brown. The female from Canary Islands is the darkest specimen examined with metasoma (except medial pale patch) and hind leg largely brown.

From “carmino” (Latin for “comb”), because of the comb on the hind tibia.

*France (Corsica), *Spain (mainland, Balearic and Canary Islands).

Males have on average about 2–3 more antennal segments than females.

Neotype here designated, ♀ (NMS, Edinburgh), “[Scotland], Rowardennan, Stirlings., Hypena proboscidalis [on] Urtica, 2.ix.[19]89, mum. 17.iv.[19]90, em. 10.vi.[19]90, M.R. Shaw (♀ 2 in 1990 Expts)”.

Widespread in western Europe: *Austria, Belgium, British Isles (England: V.C.s 1, 3, 4, 11, 12, 14, 17, 20, 22, 23, 25, 26, 27, 28, 30, 31, 32, 33, 58, 61, 62, 63, 64; Wales: V.C. 52; Scotland: V.C.s 72, 77, 84, 86, 87, 89, 99, 111; Ireland: Co. Cork), Bulgaria, Czech Republic, *Finland, Germany, Hungary, Italy, *Lichtenstein, Netherlands (FL: Lelystad (Oostvaardersplassen), FR: Ried, GE: Heerde; Tongeren; Brummen (Voorstonden), LI: Kerkrade; St. Pietersberg; Tegelen; Wrakelberg, NB: Bergen op Zoom, ZH: Asperen; Waarder; Lexmond), Norway, Spain, Slovakia, *Sweden. Specimens in NMS, BMNH, OUM, BZL, RMNH, MTMA, ZSSM, ZISP, World Museum Liverpool, CNC, USNM, UWIM, M. Riedel collection, H. Schnee collection, MSC, JLCWAE, and I. Kakko collection.

MRS062 (England EU979579, CO1 + KU682264, 28S), MRS073 (England KU682256, CO1), MRS074 (England KU682220, CO1).

Plurivoltine parasitoid of larvae of Hypena proboscidalis (Linnaeus, 1758) (Erebidae: Hypeninae), overwintering in the host larva. Mummy (Fig. 15) brown and moderately slender. Specimens (in NMS unless indicated) reared from Hypena proboscidalis (Linnaeus) (15 [1 BMNH, 1 OUM, 1 AAC, 3 H. Schnee collection]; A.A. Allen, G.M. Haggett, A. Hawkins, R.J. Heckford, S. Ratering, M.R. Shaw). It may be an absolute specialist on Hypena proboscidalis; related species that similarly feed on Urtica (H. obsitalis (Hübner) and H. obesalis (Treitschke)) overwinter as adults and would not (by themselves) be capable of supporting the parasitoid’s annual life cycle: indeed, it has been absent from several large collections of H. obesalis made in various localities in the Alps (M.R. Shaw). The oviposition sequence (observations from two females, and H. proboscidalis) is abnormal in that there is no separate pre-oviposition sting inducing temporary paralysis, nor is there a post-oviposition period of association. The host is scarcely antennated, but quickly recognised and pounced upon or snatched with the front two pairs of legs, and held aligned with the parasitoids’s body while the ovipositor is inserted for a prolonged period – sometimes there are several insertions, with self-superparasitism then often occurring. The egg is strongly attached to internal organs (gut and malpighian tubules both observed) at its narrow end. Because of subsequent disease in the stock, it is not possible to give quantitative results, but it appeared that enthusiastic attack on fourth instar hosts resulted in oviposition but no development, and that only earlier instar hosts were suitable. Mummification takes place on the host’s food plant, usually in a semi-concealed position.

Antennal segments of female 42–47, of male 42–46; fore femur of ♀ 5.4–5.7 × as long as wide (Fig. 131) and hardly sculptured, but of ♂ slenderer; scapus and pedicellus (yellowish) brown ventrally; temples directly narrowed behind eyes; precoxal area frequently with some rugae or rugulae; propodeum distinctly transversally rugose medially and median carina largely absent on posterior half of propodeum or irregular; posterior half of pterostigma of female largely dark brown; ivory part of malar space usually reaching clypeus (Fig. 134); mesosternum more or less blackish or dark brown, rarely completely reddish; hind femur of ♀ rather reddish brown, but may be largely infuscate in ♂; OOL about equal to diameter of posterior ocellus (Fig. 136); vein 1-CU1 of fore wing horizontal and vein cu-a short (Fig. 127); antenna dark brown or black (but scape below usually paler than face; Fig. 134), rarely completely yellowish brown; mesosoma black(ish) dorsally, especially mesoscutum and scutellum (but notaulic area may be brownish posteriorly); metasoma largely blackish with (pale) yellowish elliptical patch medially (Fig. 129). Similar to A. nigricornis Wesmael, 1838, which (like many A. circumscriptus) has the mesoscutum usually without light markings, but A. nigricornis has the fore femur more slender (6.7–7.4 × as long as wide) and very finely sculptured, the scapus and the pedicellus more or less infuscate or black ventrally, the precoxal area usually without rugae, the propodeum largely coriaceous medially and the median carina at least anteriorly present on posterior half of propodeum and regular, the posterior half of the pterostigma of female more or less yellowish, but usually apical third laterally darkened, the pale yellowish part of the malar space usually not reaching the clypeus and the mesosternum usually reddish or brownish.

Figures 125–126. 

Aleiodes circumscriptus (Nees), ♀, neotype. 125 habitus lateral 126 mummy of Hypena proboscidalis (Linnaeus).

Figures 127–137. 

Aleiodes circumscriptus (Nees), ♀, neotype. 127 wings 128 mesosoma lateral 129 propodeum and metasoma dorsal 130 hind leg lateral 131 fore femur lateral 132 antenna 133 apical segments of antenna 134 head anterior 135 head lateral 136 head dorsal 137 basal segments of antenna.

Neotype, ♀, length of fore wing 4.9 mm, of body 5.3 mm.

Head. Antennal segments 46, length of antenna 1.3 × fore wing, its subapical segments about 1.6 × as long as wide; frons coriaceous and posteriorly rugulose, weakly shiny; OOL equal to diameter of posterior ocellus and coriaceous; vertex coriaceous, with satin sheen; clypeus moderately convex, coriaceous; ventral margin of clypeus thick and depressed (Fig. 134); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 134) and face mainly coriaceous with some rugae dorsally; length of eye 3.2 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum coriaceous and occipital carina nearly complete, interrupted dorsally by somewhat less than width of ocellus (Fig. 136); clypeus partly above lower level of eyes (Fig. 134); length of malar space 0.4 × height of eye in lateral view; eyes moderately protruding (Fig. 136).

Mesosoma. Mesoscutal lobes largely coriaceous, matt, but medio-posteriorly with a few longitudinal rugae; notauli narrow, shallow and crenulate, but posterior half absent; prepectal carina lamelliform medio-ventrally, reaching anterior border; precoxal area of mesopleuron coriaceous and with some rugae medially; mesopleuron above precoxal area (except large smooth and shiny speculum) coriaceous, but dorsally rugose; medially metapleuron coriaceous, matt; mesosternal sulcus narrow and rather deep, with carina posteriorly; mesosternum rather angulate posteriorly; scutellum nearly flat, coriaceous and largely non-carinate laterally; propodeum rather flat and coriaceous but posteriorly with some rugae, median carina present but absent on posterior half, without tubercles.

Wings. Fore wing: r 0.2 × 3-SR (Fig. 127); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 127); cu-a vertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell parallel-sided or nearly so (Fig. 127); 2-SC+R short and longitudinal; short stub of m-cu present, unpigmented.

Legs. Tarsal claws setose; hind coxa superficially coriaceous, with satin sheen; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 5.7 and 4.3 × their width, respectively (Figs 130–131); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus.

Metasoma. First tergite as long as wide posteriorly, flattened and latero-posteriorly lamelliform; first tergite coriaceous and finely irregularly longitudinally rugose; second tergite robust (Fig. 129), without distinct median carina, with satin sheen and superficially rugulose; medio-basal area of second tergite absent; second suture shallow and largely crenulate; basal half of third tergite indistinctly rugulose, remainder of metasoma largely superficially coriaceous and rather shiny; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath largely densely setose.

Colour. Black or brownish black; antenna brown, but scapus dorsally and laterally dark brown; palpi, malar space up to eyes, mandible, tegulae, fore and middle coxae, trochanters and trochantelli, bases of fore and middle femora, medio-apical fifth of first tergite, medially second tergite and medio-basal patch of third tergite pale yellowish (Fig. 129); orbita (except latero-ventrally) brownish yellow (Figs 134–136) and remainder of head dark brown; mesopleuron ventrally yellowish brown with darker mesosternum; hind femur (except basally) fuzzy brown (Fig. 130), remainder of legs brownish yellow; veins and pterostigma (except yellow basal 0.4 and apex) dark brown; border between dark and pale part of pterostigma fairly sharp, contrasting with each other (Fig. 127); wing membrane subhyaline.

Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀: 42(1), 43(1), 44(17), 45(46), 46(28), 47(4); of ♂: 42(13), 43(30), 44(38), 45(25), 46(1); notauli absent posteriorly or shallowly impressed; mesoscutum sometimes with weak diffuse reddish colouration posteriorly, along notaulic courses; orbita sometimes completely yellowish; mesosternum varying from (frequently) almost black, and then strongly contrasting with the reddish lower third of the mesopleuron, to reddish brown; median carina of propodeum sometimes traceable to posterior margin.

Males have on average about one fewer antennal segment than females.

Lectotype of A. ochraceus Hellén here designated (FMNH), ♀, “[Finland,] Jomala”, “Hellén”, “829”, “Coll. Hellén: Aleiodes ochraceus Hellén”, “http://id.luomus.fi./GL3421”; one ♀ paralectotype (topotypic, GL3420) and one ♂ paralectotype (Nystad, GL3419).

*Austria, Finland, France (*mainland and *Corsica), *Hungary, Italy (*mainland and *Sicily), *Romania, *Spain, *Slovakia, *Slovenia, *Turkey. Specimens in NMS, BMNH, RMNH, MTMA, FMNH, ZSSM, FC, JLC.

MRS056 (Corsica JF962825, CO1), MRS336 (Italy JF973341, CO1), MRS338 (Italy KU682235, CO1]), MRS342 (Italy KU682236, CO1), MRS343 (Italy JF962826/KU682237, CO1).

Unknown. Most specimens have been collected at night in July and August in open situations. It may be univoltine and have hosts in low vegetation.

Head subglobose (Fig. 150); antennal segments of ♀ 28–35, and stout (Fig. 147), of ♂ 39–42; antenna of ♀ 0.8–1.1 × as long as fore wing, longer in ♂; OOL of ♀ 1.2 × diameter of posterior ocellus; occiput coriaceous-rugose or -rugulose (Fig. 150); second submarginal cell of fore wing rather narrow (Fig. 139); dorsal face of propodeum long and (slightly) angularly protruding postero-laterally (Fig. 142); fore femur stout (Fig. 145); hind femur rather wide (Fig. 143); sexes strongly dimorphic, the ♂ having larger ocelli, and slender and more numerous antennal segments; body completely yellowish brown, except black stemmaticum.

Figure 138. 

Aleiodes curticornis nom. n., ♀, Italy, Tyrol, near Laudes, habitus lateral.

Figures 139–151. 

Aleiodes curticornis nom. n., ♀, Turkey, Agri, but 143 of ♀ from Tyrol. 139 fore wing 140 hind wing 141 mesosoma lateral 142 propodeum and metasoma dorsal 143 hind leg lateral 144 basal segments of antenna 145 fore femur lateral 146 antenna 147 apical segments of antenna 148 head anterior 149 head lateral 150 head dorsal 151 propodeum and first tergite lateral.

Redescribed ♀ (RMNH) from Agri (Turkey), length of fore wing 3.2 mm, of body 3.7 mm.

Head. Antennal segments 28, length of antenna 0.8 × as long as fore wing, its subapical segments slightly longer than wide (Fig. 147); head subglobose in dorsal view (Fig. 150); frons granulate-rugulose and slightly shiny; OOL 1.2 × diameter of posterior ocellus and very finely rugulose as is vertex, with satin sheen; clypeus moderately convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 148); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 148) and face mainly transversely rugulose and with medium-sized setae; length of eye 2.2 × temple in dorsal view and temple subparallel-sided behind eye; occiput behind stemmaticum rugulose and occipital carina present ventrally and widely interrupted dorsally; clypeus near lower level of eyes (Fig. 148); length of malar space 0.5 × height of eye in lateral view (Fig. 149); eyes somewhat protruding (Figs 148–150).

Mesosoma. Pronotum medio-anteriorly distinctly convex; mesoscutal lobes largely coriaceous, matt and medio-posteriorly rugulose, notauli narrow and sparsely finely crenulate and posteriorly absent; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron finely rugose; mesopleuron above precoxal area (except partly smooth and shiny speculum) superficially granulate, but dorsally rugose; medially metapleuron granulate and matt; mesosternal sulcus narrow and rather shallow, without carina posteriorly; mesosternum rounded posteriorly; scutellum slender, moderately convex, mainly granulate and non-carinate laterally; dorsal face of propodeum largely rugose, long and (slightly) angularly crest-like or tuberculate protruding postero-laterally, median carina present but irregular and similar to surrounding sculpture.

Wings. Fore wing: r 0.5 × 3-SR (Fig. 139); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell rather slender (Figs 138–139); vein M+CU1 of fore wing apically at same level as vein 2-CU1 (Fig. 139); vein 1-SR of fore wing short and linear with vein 1-M (Fig. 139); cu-a subvertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 140); 2-SC+R narrow and longitudinal; m-cu present, but unpigmented.

Legs. Tarsal claws setose; hind coxa coriaceous, with some oblique striae and satin sheen and about reaching apex of first tergite; hind trochantellus 2.4 × longer ventrally than wide; length of fore and hind femora 4.5 and 4.2 × their width, respectively (Figs 143, 145); inner apex of hind tibia without comb; length of inner hind spur 0.4 × hind basitarsus.

Metasoma. First tergite 0.9 × as long as wide posteriorly, rather flattened medially and latero-posteriorly non-lamelliform; first–second tergites longitudinally rugose and with median carina (Fig. 142); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite without median carina; third–fourth tergites finely coriaceous; fourth tergite flat medially and apically truncate; fourth tergite without sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose.

Colour. Yellowish brown; palpi, tegulae, pterostigma, veins (but parastigma and part of basal veins dark brown) and legs yellow; stemmaticum and ovipositor sheath black; wing membrane subhyaline.

Variation. Sexual dimorphism is unusually pronounced in this species, in respect of the large ocelli and the slenderer and much higher number of antennal segments of the male. Antennal segments of ♀ 28(2), 30(6), 31(2), 32(1), 33(1), 34(3), 35(1) and of ♂ 39(2), 40(1), 41(5), 42(1); antenna of ♀ 0.8–1.1 × as long as fore wing; stemmaticum black or brown; hind femur of ♀ moderately robust (Fig. 143) to rather swollen.

Holotype, ♀ (NMS), “[U.K.], Wales: Anglesey, Fedw Fawr, ex indet. Noctuid swept at night ex Calluna etc., 23.v.[19]97, mum. 29.v.[19]97, em. 16.vi.[19]97, died 1.ii.[19]98, M.R. Shaw, ♀ A. diarsianae in 1997 expts.”, “Host remains compatible with Diarsia sp., possibly brunnea or mendica det M.R. Shaw, 2013”. Paratypes (3 ♀ + 55 ♂): 42 ♂ (NMS, RMNH, BMNH) progeny of the holotype, 29 cultured in the noctuid Diarsia mendica (Fabricius), oviposition in range 26.vii–3.viii.1997, mummification 18–27.v.1998, emergence 25.vi–4.vii.1998 and 13 in Diarsia rubi (Vieweg), oviposition 23–30.viii.1997, mummification 4.iv–9.v.1998, emergence 7–25.vi.1998; 1 ♂ (NMS) England: Westmorland, Arnside Knott, ex Diarsia ? brunnea (Denis & Schiffermüller) on Calluna, coll. 5.v.1984, mum. 16.v.1984, em. 19.vi.1984, M.R. Shaw; 1 ♀ + 1♂ (NMS) Scotland: Orkney, ex Diarsia brunnea on Calluna, coll. v.1977, em. vi.1977, R.I. Lorimer; 1 ♀ (NMS) Scotland, East Perth, Drumderg, NO2055, ix.2012, A. Huff; 1 ♂ (NMS) Scotland, South Aberdeen, Glen Tanar 16.vii–4.viii.1986, I. MacGowan; 1 ♂ (NMS) Scotland, South Aberdeen, Braemar, Morrone Birkwood, 12.vii–6.viii.1984, B.D. Batty; 1 ♂ (NMS) Scotland, Easterness, Loch Garten, vi.1984, J.A. Owen; 3 ♂ (NMS) Scotland, Easterness, river Nethy shingle bank, NJ0214, 19.vi–5.vii.1999 (1 ♂) and 5–19.vii.1999 (2 ♂), M. Edwards); 2 ♂ (NMS) Scotland, Elgin, Bognacruie, NJ0415 19.vii–3.viii.1999 (1 ♂) and 3–23.viii.1999 (1 ♂), M. Edwards; 1 ♂ (NMS) Scotland, Elgin, Elchies, NJ2146, 27.vii–9.viii.1999, B. Hicks; 2 ♂ (NMS) Scotland, Shetland, HU335730, 15–19.vii.2004, C. Sullivan; 1 ♀ (ZSSM), Netherlands: Nijmegen, ex Diarsia rubi, Bauer; 1 ♀ (RMNH), France: Besse en Chande SSE, Puy de Dôme, 13.vi.1976, H. Teunissen.

MRS030 (Wales JF962600, CO1), MRS135 (Scotland KU682257, CO1 + EU854345, 28S).

Univoltine and possibly partly plurivoltine parasitoid of low feeding noctuid larvae (especially, perhaps exclusively, Diarsia spp.) on moorland vegetation such as Calluna, overwintering in the host larva. Mummy (Fig. 153) probably made in concealment, blackish and swollen. The above list of paratypes includes specimens reared in culture, with the following experimental outcomes: Diarsia mendica (Fabricius) 1:47\45\\30+6; D. rubi (Vieweg) 1:47\45\\33+5. In both cases mortality was rather heavy in the overwintering young larvae, as also in control groups. In Britain a northern insect, apparently restricted to broadly moorland habitats where it is a parasitoid of low-feeding noctuid larvae, possibly exclusively in the genus Diarsia. When using univoltine hosts it is certainly capable of being univoltine, overwintering as a small larva within the overwintering young host, which is killed in its penultimate instar to form a characteristic swollen mummy more or less concealed near ground level (in culture, all those from D. rubi were made on tissues lining the base of the container; in the case of D. mendica a small proportion formed on food plant, but again in low situations). At least one suitable host (D. rubi) is at least partly plurivoltine; it is not entirely clear how the parasitoid responds to this, but D. rubi larvae parasitised in culture in late viii all overwintered, while about 10% of the control cohort fed up to become autumn moths, which may suggest a tendency towards univoltine constraint by the parasitoid. The adults are long-lived and females can probably persist in the field from midsummer right through the latter half of the summer; on the other hand, some of the male collection dates recorded above are late enough to suggest plurivoltinism.

Figures 152–153. 

Aleiodes diarsianae sp. n., ♀, holotype. 152 habitus lateral 153 mummy of Diarsia sp.

Apical half of hind femur (partly) dark brown, darker than hind trochanter and trochantellus (Fig. 157); face with distinct rugae; antenna of ♀ with 36–40 segments and third segment stout, of ♂ with 40–46 segments, 4^th–7^th antennal segments of both sexes stout (Fig. 164); OOL 1.4 × diameter of posterior ocellus (Fig. 163); clypeus 0.4 × as wide as face; mesosternum usually at least narrowly black posteriorly (Fig. 155); malar space and temple near eye dark reddish brown or dark brown; first tergite strongly widened apically (Fig. 156); eye elongate (of ♂) in lateral view; middle third of hind femur yellowish brown or dark brown; vertex moderately setose, rather shiny and more or less blackish or infuscate posteriorly; mesosternum variable, but dark brown in all British specimens; POL 1.2–1.7 × diameter of posterior ocellus; eye 2.2–2.5 × as long as temple in dorsal view (Fig. 163). Very similar to A. bistrigatus (Roman), but the latter has the temples less directly narrowed behind the eyes, 4^th–7^th antennal segments less robust and fewer antennal segments in the males. Similar to the A. pictus-aggregate (i.e. A. pictus (Herrich-Schäffer, 1838) and A. nigriceps Wesmael, 1838) but these usually are smaller and less robust species, having the face without distinct rugae or only a few rugae dorsally, the mesosternum usually widely orange brown (and the mesopleuron usually without rugae in A. nigriceps), the malar space partly or completely and the temple near the eye (= external orbita) yellowish brown, the first tergite less widened apically, the fore and hind tarsi comparatively slender, the mummy slender and usually brownish (Figs 231, 273), the eye normal in lateral view and the antenna less robust. The holotype is much darker than at the start of its life (it lived in a humid tube for over 7 months); this happens to most experimental females.

Figures 154–164. 

Aleiodes diarsianae sp. n., ♀, holotype. 154 wings 155 mesosoma lateral 156 propodeum and metasoma dorsal 157 hind leg lateral 158 fore femur lateral 159 apical segments of antenna 160 antenna 161 head anterior 162 head lateral 163 head dorsal 164 basal segments of antenna.

Holotype, ♀, length of fore wing 4.0 mm, of body 5.5 mm.

Head. Antennal segments of ♀ 40, length of antenna 1.3 × fore wing, its subapical segments about 1.5 × as long as wide and third segment stout (Figs 159–160, 164); frons coriaceous and posteriorly rugulose, with satin sheen; OOL 1.4 × diameter of posterior ocellus and coriaceous; vertex coriaceous, matt; clypeus slightly convex, indistinctly sculptured; ventral margin of clypeus rounded and depressed; width of hypoclypeal depression 0.36 × minimum width of face and face distinctly transversely rugose (Fig. 161); length of eye 2.5 × temple in dorsal view and temple rather directly narrowed behind eye (Fig. 163); occiput behind stemmaticum coriaceous and with some rugulae, occipital carina interrupted dorsally by somewhat less than width of ocellus (Fig. 163); clypeus partly above lower level of eyes and 0.4 × as wide as face (Fig. 161); length of malar space 0.5 × length of eye in lateral view; eyes moderately protruding (Figs 161–163).

Mesosoma. Mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli complete and moderately wide, weakly crenulate and posteriorly widened and rugose; prepectal carina medium-sized and lamelliform, reaching anterior border; precoxal area of mesopleuron largely widely rugose, mesopleuron above precoxal area (except nearly smooth and shiny speculum) largely rugose (Fig. 155); metapleuron coriaceous, matt and posteriorly rather tuberculate; mesosternal sulcus narrow and deep, absent and replaced by carina medio-posteriorly; mesosternum rounded posteriorly; scutellum elongate, slightly convex, coriaceous and laterally largely carinate; propodeum rather flat dorsally (depressed laterally and posteriorly, rather tuberculate latero-posteriorly) and strongly rugose but anteriorly weakly so, median carina complete, but posteriorly irregular.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 154); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.65 × 2-SR, and 0.55 × 3-SR; second submarginal cell comparatively large (Figs 152, 154); cu-a weakly oblique, not parallel with CU1b, straight; 1-M slightly curved posteriorly. Hind wing: apical half of marginal cell parallel-sided or nearly so; 2-SC+R short and longitudinal; m-cu present and weakly pigmented (Fig. 154).

Legs. Tarsal claws setose; hind coxa coriaceous but partly superficially rugulose, largely matt; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 5.1 and 4.3 × their width, respectively (Figs 157–158); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus.

Metasoma. First tergite 0.8 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope comparatively wide (Fig. 156); first–third tergites densely and distinctly longitudinally rugose, robust (Fig. 156), with distinct median carina; medio-basal area of second tergite absent; second suture moderately impressed and crenulate; remainder of metasoma largely superficially coriaceous; fourth and apical fifth of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) densely setose.

Colour. Black or brownish black; antenna pale brown, but scapus dorsally and apical seventh of antenna dark brown; palpi, and tegulae pale yellowish (Fig. 152); orbita posteriorly and dorsally brownish yellow (Figs 162–163); mesosoma orange brown, but propleuron, mesoscutal lobes medially, metanotum laterally, anterior half of propodeum and metapleuron, mesopleuron dorso-posteriorly and mesosternum black; metasoma largely dark orange brown (Figs 152, 156); hind femur (except basally) fuzzy dark brown (Fig. 157) and remainder of legs yellowish brown; veins and pterostigma (except yellow basal 0.2 and apex) dark brown; border between dark and pale part of pterostigma sharp, contrasting with each other (Fig. 154); wing membrane subhyaline. This specimen had lived in a humid tube for 7 months, and its colour had deepened considerably over this time.

Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀ 36(1), 38(3), 39(1), 40(1), of ♂ 40(2), 41(2), 42(4), 43(13), 44(16), 45(14), 46(3); mesosoma largely black to largely orange-brown; OOL of male slightly longer than diameter of posterior ocellus and apical half of antenna dark brown; fifth maxillary palp segment slender to moderately widened and rather long; first tergite (except medio-posteriorly) black (♂) or entirely dark reddish-brown (♀) and second tergite black or reddish laterally; in British females only posterior segments somewhat darkened; in British males first tergite more or less blackish in anterior half as well, but second and third tergites usually (almost) fully orange, sometimes with infuscation sublaterally on second tergite (especially anteriorly).; mesopleuron medially and propodeum rugose or superficially rugulose; few females seen, but in one very extensively orange specimen the legs are almost completely orange, with only slight infuscation in the apical half of hind femur. May be confused with A. borealis (Thomson, 1892), but this species has less antennal segments (♀: 32–34 segments), palpi and legs more or less infuscate and the clypeus wider (about 0.5 × width of the face).

We have seen 3 ♀ + 11 ♂ (NMS) from Sweden (Bohuslän and Västerbotten) and Finland (Kuusamo and Saarijärvi) that come close to A. diarsianae, but differ in being less robust (T1 less expanded apically; antennal segments longer in relation to width, especially basally), less strongly sculptured (fewer rugae on face; mesopleuron with only weak rugae), and having somewhat larger eyes. They also have slightly more antennal segments, at least in the female sex (the two females with intact antennae have 44 and 45 segments, the males have 42(1), 43(1), 44(1), 45(6), 47(1)), and the females have T1 more or less extensively blackish in anterior half, unlike the British A. diarsianae females seen, in which it is uniformly orange. CO1 sequences have been obtained for two localities (Västerbotten and Kuusamo; respectively MRS304 GenBank KU682234, and MRS692 GenBank KU682247): they form a well-isolated clade with A. diarsianae but differ from it by 8 fixed base-pairs. One of the Finnish males was reared from a noctuid mummy collected on a twig in a bog (N.R. Fritzén) later kindly identified from its CO1 sequence as Coenophila subrosea (Stephens) by Dr Katja Kramp (SDEI). Another male (in NMS) from Norway (Turtagrö, Sogn og Fjordane) has 42 antennal segments and probably belongs to the same species; it was reared from an unidentified dark noctuid mummy on Betula nana (K.P. Bland), which, like the Finnish one, is somewhat swollen but not as extensively so as in the considerable number of British A. diarsianae mummies we have seen. Both of these specimens, in common with some (but not all) males from Sweden, have the maxillary palp more swollen than seen in the British material. More material is needed to settle the status of these Fennoscandian populations, but we provisionally regard them as probably a different species near A. diarsianae.

Named after the generic name of its host: Diarsia Hübner.

*British Isles (England, Wales, Scotland), *France, *Netherlands.

Males have on average about 3–4 more antennal segments than females.

Holotype of R. esenbeckii, ♂ (ZSSM), “715, [Germany, Charlottenburg]”, “Esenbeckii n.”, together with mummy of Dendrolimus pini (L.); holotype of R. corsicus, ♀ (MTMA), “[France, Corsica,] Ajaccio”, “praetor Reinh.? (Corsica)”.

f. esenbeckii: Austria, *Croatia, Czech Republic, France (*mainland and Corsica) *Netherlands (Muiderberg), Germany, Spain (*Mallorca); f. dendrolimi: China, *Finland, Russia, Switzerland. Specimens in NMS, ZSSM, RMNH, BZL, MTMA, BMNH, SDEI, ZISP

MRS180 (Finland EU979581,CO1 + EU854329, short 28S), MRS500 (Mallorca JF962845/KU682240, CO1).

Apart from the examined holotype (see above) all the reared specimens we have seen (of form dendrolimi) were from Siberian populations of Dendrolimus superans sibericus (Rozhkov) (Lepidoptera: Lasiocampidae) (7). Aleiodes esenbeckii is a well-known parasitoid of Dendrolimus species. In the central part of its range the host species D. pini (Linnaeus) is normally univoltine, but in southern European (Mediterranean) populations it is at least bivoltine (Vadim V. Zolotuhin, pers. comm.), while in northern Siberia the host D. superans sibericus Tchetverikov usually has a 2-year life cycle. The parasitoid overwinters inside the diapausing host larva, and adapts its seasonality according to that of the host: Boldaruev (1958) reports that in Siberia A. dendrolimi spends over 20 months inside its host larva, which is mummified in late May to June following the second winter of the 2-year cycle, but in captivity (and also ocassionally in the wild) the host, and similarly the parasitoid, can be induced to develop within a year or even less. He also found that the adult parasitoid is long-lived and that females develop eggs only after a prolonged period of feeding on honeydew. Specimens seen from Europe have been collected from late May–October, no doubt reflecting local seasonality of the host.

Antennal segments of ♀ 56–62; head entirely brownish yellow apart from stemmaticum (Figs 173–175): f. esenbeckii) or blackish (Figs 185–187: part of f. dendrolimi); scapus in lateral view ratherly oblique apically; OOL 0.3 × diameter of posterior ocellus; occipital carina complete ventrally (Fig. 174); length of malar space 0.25 × height of eye in lateral view; vein 2-CU1 of fore wing 0.7–1.2 × vein 1-CU1 (Fig. 167); vein 1-SR angled to vein 1-M and vein 1-M distinctly curved (Fig. 167); vein r of fore wing gradually merging into vein 3-SR; tarsal claws small (Fig. 170); fourth metasomal tergite superficially coriaceous; length of fore wing 6–10 mm. Often confused with A. varius (as A. procerus), but differs by the relative lengths of veins 1- and 2-CU1 and by the number of antennal segments.

Figures 165–166. 

Aleiodes esenbeckii (Hartig), ♀, Spain, Mallorca. 165 habitus lateral 166 mummy of Dendrolimus pini (Linnaeus) of holotype of A. esenbeckii from Germany.

Figures 167–177. 

Aleiodes esenbeckii (Hartig), ♀, Spain, Mallorca. 167 wings 168 mesosoma lateral 169 propodeum and metasoma dorsal 170 hind leg lateral 171 fore femur lateral 172 antenna 173 head anterior 174 head lateral 175 head dorsal 176 apical segments of antenna 177 basal segments of antenna.

Figures 178–179. 

Aleiodes esenbeckii (Hartig) f. dendrolimi (Matsumura), ♀, Russia, Sakhalin. 178 habitus lateral 179 mummy of Dendrolimus spectabilis (Butler) from China.

Figures 180–189. 

Aleiodes esenbeckii (Hartig) f. dendrolimi (Matsumura), ♀, Russia, Sakhalin. 180 wings 181 mesosoma lateral 182 propodeum and metasoma dorsal 183 hind leg lateral 184 antenna 185 head anterior 186 head lateral 187 head dorsal 188 fore femur lateral 189 basal segments of antenna.

Redescribed ♀ (NMS) from Mallorca (Spain), length of fore wing 7.6 mm, of body 9.5 mm.

Head. Antennal segments 59, length of antenna 1.3 × fore wing, its subapical segments about 1.9 × as long as wide and scapus in lateral view rather oblique apically; frons superficially granulate, rather shiny; OOL 0.3 × diameter of posterior ocellus and granulate; vertex superficially coriaceous, with satin sheen; clypeus rather high, convex dorsally and flattened ventrally, coriaceous and with long setae; ventral margin of clypeus thick and gradually depressed (Figs 173, 185); width of hypoclypeal depression 0.4 × minimum width of face (Figs 173, 185) and face mainly rugose with interspaces coriaceous; length of eye 4.7 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum finely rugose and occipital carina curved and interrupted medio-dorsally and complete ventrally (Figs 174, 186); clypeus entirely above lower level of eyes (Figs 173, 185); length of malar space 0.25 × height of eye in lateral view; eyes strongly protruding (Figs 173–175, 185–187).

Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen; notauli narrow, shallow and mainly coriaceous; prepectal carina rather lamelliform medio-ventrally, almost reaching anterior border of mesopleuron and latero-ventrally curved; precoxal area of mesopleuron coriaceous, without fine rugae medially; mesopleuron above precoxal area (including shiny and granulate speculum) coriaceous, but dorsally finely rugose (cf. Fig. 181); medially metapleuron superficially granulate and shiny; mesosternal sulcus narrow and deep, without carina posteriorly; mesosternum angulate posteriorly; scutellum superficially coriaceous and carinate antero-laterally; dorsal face of propodeum rather long and coriaceous, posterior face short, hardly differentiated, with some short carinae and smooth in between, median carina complete and with weak tubercles postero-laterally.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 167); 1-CU1 horizontal, 1.1 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.45 × 3-SR; second submarginal cell stout (Fig. 167); 1-SR rather angled to 1-M; cu-a somewhat reclivous and curved; 1-M slightly curved. Hind wing: marginal cell parallel-sided submedially and slightly widened apically (Fig. 167); 2-SC+R short and subquadrate; m-cu present as slightly pigmented vein; M+CU:1-M = 6:5; 1r-m 0.7 × 1-M.

Legs. Tarsal claws with fine brownish pecten basally; hind coxa finely coriaceous, with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 5.7 and 4.5 × their width, respectively; inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus; hind basitarsus wider than following segments.

Metasoma. First tergite 1.3 × as long as wide posteriorly, flattened and latero-anteriorly widely lamelliform; first–second tergites densely finely irregularly rugulose and with fine median carina; second tergite as long as wide basally and 1.4 × as long as third tergite (Fig. 169); minute medio-basal area of second tergite present; second suture rather deep, widened medially and distinctly crenulate; third and following tergites superficially coriaceous and shiny; fourth tergite largely without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate.

Colour. Yellowish brown; antenna (except yellow scapus and pedicellus) dark brown; stemmaticum black; hypopygium, middle and hind tarsi more or less infuscate; pterostigma and veins of middle third of wings dark brown (Fig. 167); other veins brownish yellow; wing membrane subhyaline.

Variation. Length of fore wing 6–10 mm, of body 7.5–11.5 mm; antennal segments of ♀ 58(1), 59(2), 60(3), (and of f. dendrolimi: 60(2), 61(2), 62(1), 63(1)), of ♂ 55(1), 56(2), 57(1), 58(2), 59(2), 61(1) (of f. dendrolimi: 54(1), 61(1)); latero-anterior lamella of first tergite rather wide or narrow; marginal cell of hind wing parallel-sided or slightly narrowed submedially; f. dendrolimi has head partly, palpi, mesosoma ventrally and posteriorly, metasoma and legs more or less dark brown or blackish; rarely nearly entire head black.

A. esenbeckii f. dendrolimi differs morphologically only in colouration and occurs in the East Palaearctic region and in boreal Europe, perhaps reflecting a 2-year life cycle. The CO1 sequences (between Mallorcan f. esenbeckii and Finnish f. dendrolimi) are, however, divergent, differing by at least 32 base pairs in the barcode region (around 5%) (D.L.J. Quicke, pers. comm.), suggesting effective genetic isolation of at least these populations. For mediterranean specimens the name A. corsicus Szépligeti, 1906, is available. From limited data males appear to average about 3 fewer antennal segments than females, in both forms.

*Finland: 2 ♀ (NMS), Pohjois-Savo, Kangaslampi, Malaise trap, 26.vi–17.vii.2004 and 19.vii–1.viii.2005, N.M. Laurenne; 1 ♀ (I. Kakko collection) Finland, Loppi Topeno, 67410xx: 33468xx, 24.viii.2006, I. Kakko; *Sweden: 2 ♀, 1 ♂ (BMNH), Skåne, Röstanga 6.vii.1938, D.M.S & J.F. Perkins; *Slovakia: 1 ♀ (MTMA), Smolnieká Huta, 2.vii.1956, M. Čapek.

MRS355 (Finland JF962849, CO1).

Nothing is known of the biology of this predominantly boreal species.

Head transverse in dorsal view and directly narrowed ventrally; eye rather large; antenna of ♀ sometimes with a narrow white or pale yellowish submedial band, scapus and pedicellus of ♀ similarly coloured as medial fifth of antenna; antennal segments of ♀ 49–52; OOL equal to width of posterior ocellus; length of malar space of ♀ 0.30–0.40 × (of ♂ 0.25 times) height of eye in lateral view (Fig. 200); speculum of mesopleuron rugose or reticulate and dull as remainder of mesopleuron; propodeum distinctly elongate (Fig. 190); fore wing narrow (Fig. 191); vein m-cu of fore wing straight and angled to vein 2-CU1 (Fig. 191); pterostigma dark brown with its basal third pale yellow; hind femur 4.9–5.5 × as long as its maximum width; hind trochantellus 2.2–2.7 × as long ventrally as wide (Fig. 194); hind tibia infuscate subapically, contrasting with yellowish apex of tibia (Fig. 194); tarsal claws only bristly setose, without distinct pecten; first metasomal tergite with lamella latero-anteriorly (Fig. 202), second metasomal tergite rather long; second tergite with small smooth triangular area medio-basally (Fig. 193); third tergite coarsely sculptured; fourth tergite with distinct sharp lateral crease and basally rugulose. Very similar to the East Palaearctic A. parentalis Belokobylskij, 2000; this species differs mainly by the wider hind femur, the subbasally yellowish first tergite (except two dark brown patches) and the more robust apical antennal segments.

Figure 190. 

Aleiodes jakowlewi (Kokujev), ♀, Finland, Kangaslampi, habitus lateral.

Figures 191–202. 

Aleiodes jakowlewi (Kokujev), ♀, Finland, Kangaslampi. 191 wings 192 mesosoma lateral 193 propodeum and metasoma dorsal 194 hind leg lateral 195 fore femur lateral 196 apical segments of antenna 197 basal segments of antenna 198 antenna 199 head anterior 200 head lateral 201 head dorsal 202 base of first metasomal tergite dorsal.

Redescribed ♀ (NMS) from Kangaslampi (Finland), length of fore wing 4.5 mm, of body 5.5 mm.

Head. Antennal segments of ♀ 49, length of antenna 1.2 × fore wing, its subapical segments 1.7–1.8 × as long as wide; frons granulate, with satin sheen; OOL and POL 1.0 and 0.8 × width of posterior ocellus, respectively; vertex distinctly rugulose-granulate, with satin sheen; clypeus convex and coriaceous; ventral margin of clypeus thick and convex (Fig. 199); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 199) and face mainly coarsely granulate with some rugulae; length of eye 3 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum rugulose-coriaceous; occipital carina complete, with short crenulae and dorsally curved (Figs 200–201); clypeus partly above lower level of eyes (Fig. 199); length of malar space 0.35 × height of eye in lateral view; eyes rather protruding (Figs 199–201).

Mesosoma. Length of mesosoma 1.8 × its height; mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose and anteriorly low; notauli narrow and crenulate; prepectal carina medium-sized, remaining separate from anterior border; precoxal area of mesopleuron and area above it largely rugose (Fig. 192); remainder of mesopleuron (including speculum) granulate and with satin sheen; metapleuron distinctly granulate and with satin sheen; mesosternal sulcus deep and sparsely crenulate; mesosternum rather angulate posteriorly; scutellum slightly convex, coriaceous, and laterally with carina; propodeum flattened, without tubercles and largely coarsely rugose, median carina complete.

Wings. Fore wing: r 0.5 × 3-SR (Fig. 191); 1-CU1 horizontal, 0.3 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 191); cu-a inclivous, parallel with CU1b, straight (Fig. 191); 1-M straight and 1-SR angled to 1-M. Hind wing: marginal cell parallel-sided; 2-SC+R short; m-cu short and only weakly pigmented; M+CU:1-M = 15:10; 1r-m 0.7 × 1-M.

Legs. Tarsal claws with yellow bristles and small; hind coxa rugose-granulate, with satin sheen and 0.9 × as long as first tergite; hind trochantellus 2.7 × longer ventrally than wide; length of fore and hind femora 7.0 and 4.9 × their width, respectively (Figs 194–195); inner apex of hind tibia without distinct comb; length of inner hind spur 0.25 × hind basitarsus.

Metasoma. First tergite 1.2 × as long as wide posteriorly, stout, convex anteriorly and latero-anteriorly distinctly lamelliform; first–fourth tergites densely finely rugose (Fig. 193), with distinct median carina up to middle of third tergite; medio-basal area of second tergite minute; second suture medium-sized and crenulate; third and fourth tergites with complete sharp lateral crease; fifth and following tergites retracted; ovipositor sheath mainly densely setose and apically acute.

Colour. Dark brown; palpi, humeral plate, trochanters and trochantelli, fore and middle coxae, and ventral half of metasoma ivory or pale yellow; orbita posteriorly and tegula brown; legs (but hind femur (except basally) dark brown and fore and middle femora and hind tibia subapically infuscate), first tergite posteriorly, second tergite (except laterally) brownish yellow; ovipositor sheath black; veins and pterostigma (but basal third pale yellow) dark brown; wing membrane rather infuscate.

Variation. Antennal segments of ♀ 49(3), 50(1); of ♂ 48(1). In some females the antenna is distinctly white-banded (over about 23^rd–27^th segments) but in others, even from the same locality, the antenna is completely brownish. The anterior ocellus is sometimes enlarged, but this too seems to be variable and is not always noticeable.

From limited data males appear to have fewer antennal segments than females.

Holotype, ♀ (NMS, Edinburgh), “[England:] Norfolk, Santon, ex Stilbia anomala on Deschampsia flexuosa, 18.xii.[20]01, mum. ?ii.[20]02, em. 12.v.[20]02, G. M. Haggett”, “MRS Aleiodes DNA 154”. Paratypes (475 ♀, 412 ♂) from England (West Cornwall, East Cornwall, South Devon, North Devon, Isle of Wight, North Somerset, North Wilts, South Wilts, Dorset, North Hants, South Hants, West Sussex, East Sussex, East Kent, West Kent, Surrey, South Essex, Herts, Middlesex, Berks, Oxford, Bucks, West Suffolk, East Norfolk, West Norfolk, Cambridge, Bedford, Hunts, Northampton, East Gloucester, West Gloucester, Warwick, Stafford, Leicester, Derby, Chester, South Lancaster, Mid-west York, North-west York, Westmorland), Scotland (Dumfries, Ayr, Lanark, Berwick, Haddington, Edinburgh, Linlithgow, Fife, Stirling, West Perth, East Perth, Elgin, Easterness, Westerness, Dunbarton, Kintyre, West Ross), Wales (Glamorgan, Merioneth, Caernarvon, Monmouth, Pembroke, Anglesey), Isle of Man, Guernsey, Jersey, Andorra (St. Julia), Austria (Niederösterreich: Raglitz; Poysdorf; Hainburg; Oberösterreich: Linz, Kirchschlag; Sensengebirge Bärenriedlau), Belgium (Mt. St. Pierre), Bulgaria (Rodopi: Galabovo), Cyprus (Yermasoyia Riv.; Yeroskipon; Cherkes; Limasol; Paramytha), Czech Republic (Sumava, 1000–1300m; Moravia: Mikulov), France (Orsay; Hautes-Alpes: Briançon), Finland (Kainuu: Kuhmo), Germany (Bonn: Rheinhöhenweg im Kottenforst; Schleswig-Holstein: Lübeck; Niedersachsen: Berkhof; Gottingen: Hann-Munden; Bonn; Rheinland, Köln-Flittard; Bavaria: Wiesen/Spessart; Lower Saxony: Harzburg; Mullingen; Alfeld; Oberhaverbeck; Hürth-Fischenich; Ahlem; Saxony: Zöbigker; Baden-Württemberg: Heidel/Jagdl; Goslar, Astfeld), Gibraltar (Botanical Garden), Greece (Lakonia, Parnon Oros, 1700 m; id., Taygetos, 1000–1200 m; Thessalia: Mt Olympos; Pisadia; Kozani). Hungary (Budapest: Budaliget; Budakeszi, Hársbokorhegy; Pécs, Tettye), Ireland (WI: Ballinclea; SK: Caragh Lake; DU: Phoenix Park; Carlow, Antrim), Italy (Tuscany: Gorgona Isl.; Sicily: Catania; Mt. Etna, Milo Fornazzo, 800 m; Bolzano, Sarntal; 1250 m; Südtirol: Ahrntal; Trentino: Riva s. Garda; Trentino: Tremalzo), Netherlands (DR: Wijster; Borger, FL: Lelystad (Oostvaardersplassen & Jagersbos), GE: Gortel; Ede; Epe; Heerde; Nunspeet; Tongeren, NB: Baarle-Nassau; Eindhoven; Geertruidenberg; Hoogerheide; Nijmegen, Berg en Dal; Vierlingsbeek; Etten-Leur; Hilvarenbeek; Rijen; Bergen op Zoom; Oss; Raamdonksveer; Tilburg (Kaaistoep), LI: Arcen; Geulle; Lemelerberg; Neercanne; Wrakelberg; Grubbenvorst; St. Pietersberg; Vilt; Wolder, NH: Weesp; Texel, Oudeschild, UT: Linschoten, ZH: Meijendel; Oostvoorne; Ouddorp; Lexmond; Melissant; Nieuwkoopse Plassen; Noordwijk; Rotterdam; Voorschoten; Waarder, FR: Ried; Terschelling, ZE: Haamstede; Westenschouwen), Norway (Oppland, Lom-Lia), Slovakia (B. Karpaty-Jaktar, Drietoma), Spain (Teruel, Tramacastilla; Navarra, Alsasua, 600 m; Mallorca, Porto Cristo; Zaragoza: Juslibol), Sweden (Skåne: Järahusen; Ystad; Böste; Spraggehusen; Ö. Väringe; Spukke; Härjedalen: Tänndalen; Duvberget; Halland: Åsa Närsbokrok), and Switzerland (GR: Sent Surains, Val Gronda, 1500 m). Paratypes in NMS, AAC, BMNH, BZL, M. Riedel collection, RMNH, H. Schnee collection, SYKE, MSC, ZJUH, MCZ and ZSSM.

MRS154 (England KU682229, CO1), MRS156 (England JF962813, CO1), MRS157 (England KU682230, CO1), MRS515 (Netherlands KU682260, CO1).

A parasitoid of a wide range of low feeding noctuid larvae, as listed below. Overwinters as a small larva in the host, which is killed before it is in its final instar. Mummy (Fig. 204) largely dark brown (summer generations paler) and slender, usually formed in a prominent position at least in spring. Specimens (in NMS unless indicated) reared from wild-collected Noctuidae identified as Abrostola triplasia (Linnaeus) (1; J.L. Gregory), Ammoconia caecimacula (Denis & Schiffermüller) (4:1; J. Connell/Austria), Autographa gamma (Linneaus) (2; G.E. King/Spain, E. Haeselbarth/Germany; 1 (RMNH), G. Peters/Germany), ?Cerastis rubricosa (Denis & Schiffermüller) (1; J.L. Gregory), Cucullia chamomillae (Denis & Schiffermüller) (2; A.A. Allen [1 is AAC]), ?Diarsia rubi (Vieweg) (1; T.H. Ford), Dicestra trifolii (Hufnagel) (1; G.M. Haggett), Euplexia lucipara (Linnaeus) (6:1; P. Baker), Lacanobia oleracea (Linnaeus) (2:1; P. Baker), Melanchra pisi (Linnaeus) (5; P. Baker, A. Lord, M.R. Shaw), Mythimna ferrago (Fabricius) (1; J. L. Gregory), ?Mythimna impura (Hübner) (3:1; M. R. Hall), ?Mythimna littoralis (Curtis) (1; F.D. Bennett), Noctua comes Hübner (14; J. Connell[?], D. Hackett[?], G.M. Haggett, E. Haeselbarth, R. Hinz, R.A. Softly), Noctua fimbriata (Schreber) (2; G.M. Haggett, R.A. Softly), Noctua interjecta Hübner (2; M.R. Hall, G.M. Haggett), Noctua janthina (Denis & Schiffermüller) (1; G.M. Haggett), Noctua orbona (Hufnagel) (3; G.M. Haggett), Noctua pronuba (Linnaeus) (5 [2 are BMNH]; R.A. Softly), Orthosia gracilis (Denis & Schiffermüller) (1; M.R. Shaw), Paradiarsia glareosa (Esper) (1; G.M. Haggett), Phlogophora meticulosa (Linnaeus) (10; M.R. Shaw, R.A. Softly, J. Voogd), Shargacucullia verbasci (Linnaeus) (24 [5 are OUM, 1 is AAC]; A.A. Allen, F.C. Woodforde, M.R. Shaw), Standfussiana lucernea (Linnaeus) (1; R.F. Logan), Stilbia anomala (Haworth) (4; G.M. Haggett), Xestia agathina (Duponchel) (12 [6 are ZSSM, 2 are AAC]; A.A. Allen, E. Bauer, A. Dobson, M.R. Shaw), Xestia baja (Denis & Schiffermüller) (1; J.L. Gregory), Xestia castanea (Esper) (8 [5 are ZSSM]; E. Bauer, K.P. Bland[?], M.R. Shaw), Xestia xanthographa (Denis & Schiffermüller) (31 [1 is ZSSM, 1 is RMNH]; E. Bauer, M.R. Britten, J. Connell, G.M. Haggett, M.R. Hall, N. Hall, R. Hinz, M.R. Shaw, R.A. Softly), and unidentified noctuids (81 [14 are BMNH], mostly collected as mummies). Specimens in NMS reared in culture experiments are included in the type material (but not in the above host list), and these experimental results (using females from the overwinter generation to parasitize summer hosts) are as follows: Diarsia rubi 1:16\16\\7+4 [several others were retarded but died]; Dicestra trifolii 5:28\28\\24+4; Lacanobia oleracea 6:28\22\\0+16 [several others were retarded but died]; Melanchra persicariae (Linnaeus) 4:22\13[several others pricked and paralysed without oviposition]\\0+13; Orthosia cerasi (Fabricius) 7:55\6\\5+1; Orthosia gothica (Linneaus) 6:40\1\\0+1; Orthosia gracilis 8:27\16\\14+2; Orthosia incerta (Hufnagel) 6:23\4\\2+1; Phlogophora meticulosa 9:49\44\\29+14: Shargacucullia verbasci 1:1\1\\1+0. In Britain adult flight times peak around May, July–August and October. However, in common with most Aleiodes, the adults are very long-lived and can be found in most months (but we have no British records from January to March). Both sexes come feely to light, but it is also active by day. The males tend to court non-conspecific females (at least of some species) with as much – though unsuccessful – vigour as with conspecifics. All host records are from Noctuidae feeding on low plants, but within that group this species has an unusually broad host range. Many (but not all) of the overwintering hosts, such as Noctua and Xestia species (parasitized from September to November), feed on Poaceae and grow slowly during mild periods in the winter, with mummification by the parasitoid in the host’s 3^rd or 4^th instar sometimes as early as February, but more often during March or April. The resulting adults again parasitize low-feeding noctuids, with a similarly broad range of hosts, but mostly on plants other than Poaceae. Adults of both sexes are often swept from trees and bushes (especially in late summer and autumn) but they are probably merely feeding on honeydew rather than seeking hosts: although some Noctua and Xestia species that feed through the winter on grasses do sometimes oviposit on tree leaves, with the resulting larvae feeding thereon until (easily!) displaced, their falling to the field layer usually happens in the first instar (MRS, personal observation) when they are probably still too small to be easily parasitized by A. leptofemur. It is rather remarkable that the host larvae, especially of the overwintering generation, regularly (perhaps invariably) climb out of their normal living space, to be mummified fully exposed high on stems of various kinds (very often on dead grass seed heads), on tree trunks, fence posts etc. Related common species such as A. nigriceps, A. pictus and A. similis (Curtis, 1834) [the latter to be treated in a subsequent part of this revision], which (at least overwinter) parasitize ecologically similar and closely related – or in many cases the same – noctuid species, do not cause their hosts to do this, but instead the hosts parasitized by these species seek concealment before mummification. Consequently, mummies of this group found exposed in nature are almost invariably those of A. leptofemur. It is an obvious suggestion that this helps the parasitoid to avoid idiobiont parasitism (pseudohyperparasitism) in the dangerous field layer, including by virtue of faster development to the relative safety of the adult stage (the spring-forming mummies are very dark and presumably absorb insolation energy well), but it does leave open the question why the other (related, and similarly plurivoltine) species mentioned above have not adopted the same habit.

Figures 203–204. 

Aleiodes leptofemur sp. n., ♀, holotype. 203 habitus lateral 204 mummy of Stilbia anomala (Haworth).

Length of fore femur 6.4–8.0 × its maximum width (Fig. 209) and hind femur parallel-sided (Fig. 208); mesosternum usually black(ish); face with some weak transverse rugae dorsally; OOL 1.5 × diameter of posterior ocellus; temple roundly narrowed (Fig. 214); scapus ventrally and usually basal half of antenna (dark) brown, rarely yellowish; hind femur slender, basally largely yellowish and frequently infuscate subapically, but remaining nearly always paler than ventral side of scapus; if rarely hind femur is distinctly infuscate (Fig. 208) then often also hind coxa (at least basally) and base of hind tibia infuscate (Fig. 203); face usually black or dark brown medially and near eyes yellowish brown; antennal segments of ♂ 35–40, usually 36–38, less than of ♀, which has usually 37–39 segments; pterostigma tending to be dark brown medially (Fig. 205). Similar to A. borealis (Thomson, 1892) and to species of the A. pictus aggregate. They differ by having the length of the fore femur 5.4–6.4(–7.3) × its maximum width and the hind femur more or less weakly swollen; if more than 6.4 × then the face without transverse rugae dorsally, the hind femur comparatively wide basally, the mesosternum yellowish or the temple comparatively wide, or the scapus ventrally and the basal half of the antenna yellowish brown; if the scapus is dark brown or blackish then the scapus is similarly coloured as the hind femur subapically; colour of the hind femur variable, usually dark brown or blackish subapically; face usually completely black or rarely yellowish; antennal segments of male 37–45, averages about one segment more than of female, which has 36–45 segments (32–34 in A. borealis, of which we have not seen a male with complete antenna); pterostigma is variable, but often yellowish medially; clypeus distinctly transverse and less depressed ventrally, and hind femur rather micro-sculptured. Some dark males of A. similis are very like pale males of A. leptofemur and in extreme cases scarcely separable. The new species shares with the East Palaearctic A. angustatus (Papp, 1971) the elongate and paralle-sided fore and hind femora. Aleiodes angustatus has the body entirely yellowish brown, the ocelli larger (POL slightly less than the diameter of the posterior ocellus and OOL about 1.2 times diameter of ocellus), the antenna of ♀ with 47–49 (♂: 46–49) segments, the second metasomal tergite nearly parallel-sided, the precoxal sulcus witrh distinct rugae and the pterostigma yellow.

Figures 205–214. 

Aleiodes leptofemur sp. n., ♀, holotype. 205 wings 206 mesosoma lateral 207 propodeum and metasoma dorsal 208 hind leg lateral 209 fore femur lateral 210 basal segments of antenna 211 apical segments of antenna 212 head anterior 213 head lateral 214 head dorsal.

Holotype, ♀, length of fore wing 4.5 mm, of body 4.4 mm.

Head. Antennal segments 39, length of antenna 1.1 × fore wing, its subapical segments about 1.8 × as long as wide (Fig. 211) and basal segments comparatively wide (Fig. 210); frons granulate-coriaceous, with satin sheen; OOL 1.5 × diameter of posterior ocellus and coriaceous; vertex granulate-coriaceous, with satin sheen; clypeus distinctly convex (Fig. 213), coriaceous; ventral margin of clypeus rounded and depressed (Fig. 212); width of hypoclypeal depression 0.4 × minimum width of face and face coriaceous, dorsally somewhat rugulose (Fig. 212); length of eye 2.5 × temple in dorsal view and temple gradually roundly narrowed behind eye (Fig. 214); occiput behind stemmaticum granulate-coriaceous, occipital carina interrupted by somewhat more than width of ocellus (Fig. 214); clypeus partly above lower level of eyes (Fig. 212); length of malar space 0.4 × length of eye in lateral view; eyes moderately protruding (Figs 212–214).

Mesosoma. Mesoscutal lobes finely coriaceous, with satin sheen, medio-posteriorly with a few rugulae; notauli complete and narrow, largely smooth and posteriorly reduced; prepectal carina narrow lamelliform, reaching anterior border; precoxal area of mesopleuron granulate-coriaceous, mesopleuron with superficially granulate and shiny speculum and rugose dorso-anteriorly (Fig. 206); metapleuron granulate-coriaceous, matt and posteriorly not tuberculate; mesosternal sulcus narrow and rather deep, shallow medio-posteriorly and no carina; mesosternum angulate posteriorly; scutellum elongate, slightly convex, granulate-coriaceous and laterally largely without carina; propodeum rather flat dorsally, not tuberculate latero-posteriorly, and coriaceous with median carina complete.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 205); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.35 × 3-SR; second submarginal cell comparatively large (Fig. 205); cu-a weakly oblique, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apically marginal cell nearly twice as wide as its minimum width; 2-SC+R short and longitudinal; m-cu short and not pigmented (Fig. 205).

Legs. Tarsal claws setose; hind coxa finely coriaceous, largely matt; hind trochantellus 2.4 × longer than wide; length of fore and hind femora 6.9 and 6.9 × their width, respectively (Figs 208–209); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus.

Metasoma. First tergite 1.1 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope medium-sized (Fig. 207); first and second tergites densely and longitudinally rugulose (Fig. 207), with distinct median carina; medio-basal area of second tergite absent; second suture narrow and finely crenulate; third tergite coriaceous and remainder of metasoma largely smooth and shiny; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) moderately setose.

Colour. Black; antenna dark brown; palpi largely brown; tegulae, malar space ventrally and triangular patch on second tergite pale yellowish (Fig. 207); inner orbita as dark as face centrally; outer orbita posteriorly and dorsally, malar space dorsally and mesoscutum medio-posteriorly brownish yellow (Fig. 213); mesopleuron with ventral brownish yellow stripe (Fig. 206); hind coxa largely, apical half of hind femur, base of hind tibia and tarsi rather fuzzy dark brown (Figs 203, 208) and remainder of legs yellowish brown; veins and pterostigma (except yellow basal 0.3 and slightly apex) dark brown; border between dark and pale part of pterostigma rather sharp, contrasting with each other (Fig. 205); wing membrane subhyaline.

Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀ 35(3), 36(16), 37(79), 38(98), 39(70), 40(20), 41(1), 42(1), of ♂ 35(22), 36(64), 37(89), 38(60), 39(17), 40(4), 41(1)); mesosoma largely black to largely orange brown; medial length of second tergite 0.8–0.9 × its basal width; OOL of male slightly longer than diameter of posterior ocellus and apical half of antenna dark brown; mesopleuron medially and propodeum rugose or superficially rugulose. Specimens of the summer generation(s) are usually overall paler than those from the overwinter generation. The face usually dark centrally with the inner orbits paler but sometimes face completely black (as in the type, from the overwinter generation), less often completely orange or darkened only near clypeus (males more likely than females to exhibit these extremes). Extent of orange markings on mesoscutum extremely variable, but almost always distinct; metasoma only rarely wholly black or dark brown. Colour of pterostigma very variable, sometimes pale greyish and only faintly darker near posterior margin.

The broad host range, which has (at least in part) been experimentally verified, may contribute to the variability of this species. We have seen a large number of summer-generation female specimens from S. Europe (Portugal, Greece, Turkey and most notably a long series from South Bulgaria from Rodopi in BZL) that consistently differ in colour from summer specimens from Britain in the combination of a slightly darker pterostigma, uniformly pale legs, and the metasomal tergites posterior to the central pale area tending to be reddish brown rather than blackish, and they are also slightly smaller. Because of its relative uniformity in contrast to the variability of what we otherwise regard as A. leptofemur, it seem possible that this material represents a different species and we have not included it in the type series.

This common and widely distributed species is named after its slender femora (“leptos” = Greek for “thin”).

*Andorra, *Austria, *Belgium, *British Isles (England, Wales, Isle of Man, Scotland, Ireland, Guernsey, Jersey), *Bulgaria, *Cyprus, *Czech Republic, *France, *Finland, *Germany, *Gibraltar (British territory), *Greece, *Hungary, *Italy, *Netherlands, *Norway, *Slovakia, *Spain, *Sweden, *Switzerland. The southern European countries are included provisionally (see above under variation).

Males have on average about one fewer antennal segments than females. Both authors have left determination labels for this species incorrectly as A. borealis (Thomson) on a large number of specimens in many collections (up until about 2006 for CvA; until 2007 for MRS), which are now impossible to correct.

Lectotype ♀ (ZMB) from Germany examined.

Widespread in western Europe: *Austria, British Isles (England: V.C.s 11, 17, 22, 28, 29, 31, 32, 38, 39, 52, 57, 58, 61, 62, 63, 64; Wales: V.C.s 48, 52: Scotland: V.C.s 88, 96, 97), Bulgaria, *Czech Republic, *Denmark, Finland, *France, Germany, Italy, Netherlands (DR: Borger; Wijster, GE: Heerde; Brummen (Voorstonden), FR: Terschelling (Midsland-Noord, dunes), LI: Epen, ZH: Asperen; Meijendel (dunes)), Poland, *Romania, Russia, Slovakia, Sweden, Switzerland. Specimens in NMS, MNHN, BMNH, RMNH, NRSM, BZL, MTMA, USNM, CNC, CMIM, M. Riedel coll., JLC, Delémont, ZSM, MCZ, SDEI, FRAH, WAE, UWIM, ZMC, ZMUO.

MRS282 (Wales JF962850, CO1 + KU682267, 28S).

A univoltine parasitoid of a wide range of Eupithecia species (Geometridae: Larentiinae), mummifying the host in its pupation chamber and overwintering in the mummy. Specimens reared from wild collected hosts determined as follows: Eupithecia absinthiata (Clerck) (2 NRS), Eupithecia exiguata (Hübner) (1 ZMUH), Eupithecia gelidata hyperboreata Staudinger (2 SDEI), Eupithecia goossensiata Mabille (2 NMS, 1 OUM; T.H. Ford), Eupithecia innotata (Hufnagel) (2 ZMUH, 1 ZMUO); I. Itämes), Eupithecia lariciata (Freyer) (1 NMS, 2 Copenhagen, 2 Delémont, 10 FRAH), Eupithecia nanata (Hübner) (22 NMS, 5 BMNH, 5 OUM]; T.H. Ford, G.T. Lyle), Eupithecia satyrata (Hübner) (1 NRS), Eupithecia subfuscata (Haworth) (3 OUM; T.H. Ford), Eupithecia succenturiata (Linnaeus) (1 NMS, 1 OUM; T.H. Ford), Eupithecia vulgata (Haworth) (6 NMS, 2 OUM; T.H. Ford, M.R. Shaw) and 13 (NMS) from undetermined Eupithecia spp on various low plants (including Artemisia, Lotus and Pimpinella). Experimental culture result in E. vulgata 2:16\16\\7+1. While it has been reared particularly from hosts feeding on field layer plants (as is the habit of the majority of European Eupithecia species), the rearing records also include a substantial number from E. lariciata on Larix; however, it seems probable that many of the specimens seen were reared in a nursery context, in which young Larix would have presented as part of the field layer, and it is perhaps significant that A. modestus was not found to be a prominent parasitoid of E. lariciata on mature Larix sampled in the Alps (although it was indeed reared from it in small numbers at most sampling sites: Kenis et al. 2005). The host larva is usually fully grown and cocooned in the soil as a prepupa by the time mummification occurs, and the wide and sharp-rimmed clypeus of the adult probably reflects the need to chew its way through the host’s cocoon and soil. The winter is passed in the mummy, which is rather tough and dark, slightly dorso-ventrally flattened but not keeled (Fig. 216), and sometimes weakly upcurled. Typically the thoracic and first two abdominal segments are contracted, with the parasitoid occupying approximately abdominal segments 3–8 in a thin silken lining. The mummy is not stuck down, though probably the usual ventral opening for the expulsion of fluid occurs none the less (however, this is unconfirmed). The adult flight time in Britain is approximately from late June through August.

Figures 215–216. 

Aleiodes modestus (Reinhard), ♀, England, Santon Downham. 215 habitus lateral 216 mummy of Euphithecia vulgata (Haworth).

Antennal segments of both sexes 37–(40–45)–47 and third segment rather slender (Fig. 223); ventral margin of clypeus thin and protruding (Fig. 226); maximum width of hypoclypeal depression about 0.5 × minimum width of face (Fig. 225); length of malar space of female 0.2–0.3 × height of eye in lateral view (Fig. 226); OOL about equal to diameter of posterior ocellus (Fig. 227); area in front of anterior ocellus with a minute smooth tubercle (Fig. 227); vein r of fore wing 0.5–0.6 × vein 3-SR; vein 1-SR of fore wing short (Fig. 217); vein 1r-m of hind wing distinctly shorter than vein 1-M (Fig. 217); mesopleuron above precoxal area strongly shiny and superficially sculptured; third tergite without distinct striae; pterostigma largely yellow or yellowish brown; inner side of basal half of hind tibia yellowish; head and mesosoma largely blackish.

Figures 217–229. 

Aleiodes modestus (Reinhard), ♀, Netherlands, Wijster. 217 wings 218 mesosoma lateral 219 propodeum and metasoma dorsal 220 hind leg lateral 221 fore femur lateral 222 antenna 223 basal segments of antenna 224 apical segments of antenna 225 head anterior 226 head lateral 227 head dorsal 228 base of hind tibia 229 inner apex of hind tibia.

Figured ♀ (RMNH) from Wijster (Netherlands), length of fore wing 5.4 mm, of body 5.6 mm.

Head. Antennal segments of ♀ 43, moderately setose, length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide (Fig. 224); frons rather flat and granulate; OOL equal to diameter of posterior ocellus and finely granulate; vertex finely granulate, with satin sheen; clypeus rather large, micro-granulate; ventral margin of clypeus thin and rather protruding forwards (Fig. 226); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 225); length of eye 1.9 × temple in dorsal view (Fig. 227); occiput behind stemmaticum mainly granulate with some rugulae; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view; occipital carina complete; with a minute smooth tubercle in front of anterior ocellus.

Mesosoma. Mesoscutal lobes densely and finely granulate and with punctulation, matt; prepectal carina complete, distinct; precoxal area of mesopleuron with some rugulae medially; mesopleuron above precoxal area strongly shiny, sparsely punctate and with some superficial micro-granulation (Fig. 218); metapleuron largely granulate; scutellum flat, micro-granulate and without lateral carina; propodeum convex, finely rugose and interspaces micro-granulate (Fig. 219), without tubercles, median carina complete, weak and rather irregular.

Wings. Fore wing: r 0.6 × 3-SR (Fig. 217); 1-CU1 horizontal, slender, 0.35 × 2-CU1; r-m 0.5 × 3-SR; second submarginal cell rather long (Fig. 217); cu-a vertical, straight posteriorly; 1-M slightly curved posteriorly. Hind wing: marginal cell subparallel-sided but slightly constricted medially, its apical width equal to width at level of hamuli (Fig. 217); 2-SC+R subquadrate; m-cu faintly indicated; M+CU:1-M = 5:4; 1r-m 0.6 × as long as 1-M.

Legs. Tarsal claws yellowish setose; hind coxa granulate; hind trochantellus rather robust; length of fore femur, hind femur and basitarsus 5.7, 5.2 and 8.6 × their width, respectively (Figs 220–221); length of inner hind spur 0.25 × hind basitarsus.

Metasoma. First tergite rather robust (Fig. 219); first and second tergites densely and rather finely rugose, with rather weak median carina, reduced posteriorly; medio-basal area of second tergite absent; length of second tergite 0.9 × its basal width; second suture rather deep and distinctly crenulate; third tergite 0.8 × as long as second tergite, anterior two-thirds rugose and remainder of metasoma superficially granulate and punctate, somewhat compressed; fourth and apex of third tergite without sharp lateral crease; ovipositor sheath widened and setose.

Colour. Black; palpi, tegulae and pterostigma yellow; veins (except basally) dark brown; malar area ventrally, orbita dorsally and posteriorly, pronotum anteriorly and ventrally, mesopleuron antero-dorsally and postero-ventrally, hind coxa (but basally black), medio-posterior part of mesoscutum, first tergite latero-posteriorly, second and third tergites dark reddish brown; apical half of hind femur largely dark brown, except apically; antenna, telotarsi and clypeus dark brown; remainder of legs yellowish brown or blackish; wing membrane subhyaline.

Variation. Antennal segments of ♀ 38(1), 39(0), 40(0), 41(1), 42(18), 43(27), 44(12), 45(4) and of ♂ 40(1), 41(1), 42(2), 43(37), 44(18), 45(2); length of antenna of ♂ 1.3 × fore wing; length of eye of ♀ 1.2–1.9 × temple in dorsal view, of ♂ 1.6–2.5 times; dark specimens have mesosoma, metasoma and hind coxa nearly completely black; pale specimens have scapus, pedicellus, clypeus, entire orbita, notaulic area of mesoscutum, metapleuron posteriorly, first tergite apically and most of second–fourth tergites dark reddish brown; pterostigma sometimes slightly infuscate laterally, but remaining largely yellow; the infuscation of the hind femur is sometimes diffuse; vein cu-a of fore wing vertical or oblique; rugae of precoxal area may be entirely absent.

The number of antennal segments does not differ appreciably between the sexes. The specimens reported from Hungary by Papp (1983, 1985a, 2005) are misidentified A. fortipes (Reinhard).

Lectotype, ♀ (KBIN) from Belgium examined.

*Austria, Belgium, British Isles (*England: V.C.s 3, 8, 12, 19, 20, 22, 23, 24, 25, 27, 28, 29, 30, 31, 33, 40, 58, 62, 65; *Wales: V.C.s 41, 45, 52; *Isle of Man: V.C. 71; *Scotland: V.C.s 75, 77, 83, 84, 89, 97, 99, 101, 105; *Ireland: V.C. H5), *Jersey, *Bulgaria, *Czech Republic, *Finland, France, Germany, Hungary, *Italy, *Lithuania, *Netherlands (FR: Terschelling (Formerum), GE: Putten, NB: Geertruidenberg; Oploo; Udenhout (de Brand), NH: Overveen; Muiderberg, LI: Tegelen; St. Pietersberg; Venlo; Maastricht, UT: Harmelen, ZE: Cadzand; Oostkapelle; ZH: Delft; Den Haag; Wassenaar; Leidschendam), *Norway, *Poland, *Portugal, Russia, *Slovakia, Spain, *Sweden and Switzerland. Specimens in NMS, BMNH, RMNH, BZL, SDEI, ZSSM, FMNH, MTMA, CNC, P.-N. Libert collection, JLC, WAE, H. Haraldseide collection, and MSC.

MRS075 (England KU682221, CO1), MRS144 (Scotland KU682228, CO1), MRS613 (Wales KU682243, CO1), MRS783 (Austria KU682250, CO1).

At least partly a plurivoltine parasitoid of larvae of low-feeding noctuine Noctuidae inhabiting grassy sites; most frequent in humid situations. Mummy (Fig. 231) dark brown or brown and slender, only slightly or not swollen, forming low down in concealment. Specimens (in NMS unless stated otherwise) reared from wild-collected Noctuidae identified as Diarsia rubi (Vieweg) (10 [2 are ZSSM]; G.M. Haggett, M.R. Shaw), Lycophotia porphyyrea (Dennis and Schiffermüller) (1 [ZSSM/Netherlands]), Noctua fimbriata (Schreber) (1; J. Connell/Austria), Xestia c-nigrum (Linnaeus) (1; J. Connell/Austria), Xestia sexstrigata (Haworth) (3; G.M. Haggett), and indet. green noctuid (1). A further specimen was reared from a noctuid mummy collected in a Carex tussock (D.G. Notton). Specimens in NMS reared in culture experiments as follows: Xestia sexstrigata 6:20\20\\15+2, Xestia xanthographa (Denis & Schiffermüller) 4:47\47\\3+30 [several others were retarded but died], Noctua fimbriata 6:45\40\\24+0 [several others were retarded but died], Diarsia rubi 6:89\83\\74+9, Phlogophora meticulosa (Linnaeus) 7:43\19\\0+12 [several others were retarded but died]. No ovipositions resulted from the following culture experiments: Diarsia mendica (Fabricius) 1:5\0\\-, Xanthorhoe fluctuata (Linnaeus) 3:19\0\\-, Xanthorhoe montanata (Denis & Schiffermüller) 3:15\0\\-, Camptogramma bilineata (Linnaeus) 6:13\0\\-. While many of its hosts are univoltine with overwintering larvae, in which the parasitoid overwinters as an early instar larva, A. nigriceps has the capacity to be plurivoltine (e.g. in D. rubi), and its long summer flight period, including abundant males in late summer, indicates that it regularly is so. Experimental rearings suggest that even within the genera it uses as hosts it specialises narrowly, and these experiments also clearly distinguish between it and both A. pictus and A. leptofemur. The mummy forms low down, more or less in concealment, and is very seldom collected (unlike that of A. leptofemur). The adult emerges soon after. In captivity adult females of this species are long-lived, but have always died off during the winter, and the flight period is ca May–September. Males are sluggish in their courtship, even towards conspecific females which, however, typically offer themselves for mating immediately.

Figures 230–231. 

Aleiodes nigriceps Wesmael, ♀, Wales, Anglesey. 230 habitus lateral 231 mummy of Xestia sexstrigata (Haworth).

Antennal segments of female (38–)39–43, of male (38–)39–44; pale area of facial (= inner) orbita of female usually narrow or absent (Fig. 239); OOL 1.6 × diameter of posterior ocellus; face usually wholly black or dark brown, rarely wholly yellow brown; mesopleuron without distinct rugulae; mesosternum yellowish or reddish, if darkened then dark area usually not sharply defined; width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 239); mesoscutum antero-laterally and pronotum medio-anteriorly often black or dark brown; face without distinct rugae or only a few rugae medio-dorsally; ventrally mesopleuron usually widely orange brown; malar space partly or completely and temple near eye orange brown; length of first tergite of ♀ 1.1–1.2 × its apical width; second tergite comparatively wide (Fig. 234); fore and hind femora comparatively slender (Figs 235–236); pterostigma partly darkened anteriorly; mummy slightly or not swollen and dark brown or brown; middle third of hind femur partly or entirely dark brown. Very similar to A. pictus (Herrich-Schäffer, 1838), but A. pictus has the antennal segments of ♀ (35–)36–40, of ♂ (36–)37–41, the mesopleuron with some rugulae (at most very weakly evident in A. nigriceps), the mesosternum almost always strongly darkened or black and dark area usually sharply defined, malar space somewhat longer than in A. nigriceps, and the mesoscutum antero-laterally and pronotum medio-anteriorly more often yellowish (but variable in both species). The yellowish colouration is frequently more brownish in A. pictus (tending to orange in A. nigriceps), the legs are on the whole less slender, and the paler area at the extreme apex of the hind femur tends to be more extensive. In practice, the majority of specimens of A. nigriceps have both the mesosternum and third tergite completely yellowish orange, and such specimens are easy to recognise as this combination is rarely approached in A. pictus. In extreme examples of A. nigriceps the mesoscutum and (less often) even the scutellum may be wholly black, and when the tergites and scape are also predominantly dark they may resemble A. nigricornis, but in that species the hind femur is only very rarely extensively darkened, and the number of antennal segments is greater (though with a small overlap).

Figures 232–242. 

Aleiodes nigriceps Wesmael, ♀, Wales, Anglesey. 232 wings 233 mesosoma lateral 234 propodeum and metasoma dorsal 235 hind leg lateral 236 fore femur lateral 237 apical segments of antenna 238 antenna 239 head anterior 240 head lateral 241 head dorsal 242 basal segments of antenna.

Redescribed ♀ (NMS), “Culture: [Wales], Anglesey, Llangristiolus, from ♀ 3:2 [ex] Diarsia rubi in Xestia sexstrigata, ovip. 28.v.[20]08, mum. 4.iv.[20]09, em. 22.v.2009, died viii.2009, ♀ 2/09, M.R. Shaw”. Length of fore wing 4.4 mm, of body 5.5 mm.

Head. Antennal segments 43, length of antenna 1.3 × fore wing, its subapical segments about 1.6 × as long as wide; frons granulate-coriaceous, with satin sheen; OOL 1.6 × diameter of posterior ocellus (POL 1.4 times); vertex granulate-coriaceous, with satin sheen; clypeus distinctly convex and largely smooth; ventral margin of clypeus thick and depressed (Fig. 239); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 239) and face mainly coriaceous with some rugulae medio-dorsally; length of eye 2.3 × temple in dorsal view and temple roundly narrowed behind eye (Fig. 241); occiput behind stemmaticum coriaceous and occipital carina absent medio-dorsally, its interruption slightly less than width of stemmaticum (Fig. 241); clypeus partly above lower level of eyes (Fig. 239); length of malar space 0.4 × height of eye in lateral view; eyes moderately protruding (Figs 239–241).

Mesosoma. Mesoscutal lobes finely coriaceous, with satin sheen, but medio-posteriorly longitudinally rugose; notauli narrow, moderately impressed and finely crenulate, but posteriorly lost in rugose area; prepectal carina narrow lamelliform and reaching anterior border; precoxal area of mesopleuron coriaceous and without rugae medially; mesopleuron above precoxal area (except small smooth and shiny speculum) coriaceous, but antero-dorsally rugose; medially metapleuron coriaceous, matt; mesosternal sulcus narrow and moderately deep, but posteriorly shallow and with a carina; mesosternum rather angulate posteriorly; scutellum nearly flat, coriaceous and largely non-carinate laterally; propodeum rather flat and coriaceous but posteriorly with some rugae, median carina complete, without tubercles.

Wings. Fore wing: r 0.2 × 3-SR (Fig. 232); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 232); cu-a oblique, but not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell somewhat widened apically (Fig. 232); 2-SC+R short and longitudinal; short stub of m-cu present, pigmented.

Legs. Tarsal claws setose; hind coxa superficially coriaceous, with satin sheen; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 6.3 and 5.6 × their width, respectively (Figs 235–236); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus.

Metasoma. First tergite 1.2 × as long as wide posteriorly, convex, but posteriorly flattened; first and second tergites and base of third tergite densely longitudinally rugose; second tergite robust (Fig. 234), with distinct median carina, matt; medio-basal area of second tergite obsolescent; second suture shallow and crenulate; third tergite (except basally) largely coriaceous, remainder of metasoma largely smooth and rather shiny; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath largely densely setose.

Colour. Black or brownish black; antenna brown, but scapus and pedicellus dorsally and laterally and apical third of antenna dark brown; palpi and tegulae pale yellowish (Fig. 230); outer orbita posteriorly and dorsally brownish yellow (Figs 240–241) and remainder of head black; mesopleuron (except dorsally), mesosternum and metapleuron medially orange brown; hind femur (except basally) dark brown (Fig. 235); scutellum, first tergite medio-apically, second and third tergites, part of fourth tergite and remainder of legs brownish yellow; veins (except dark brown veins 1-SR, 1-M, r and CU1) brown; pterostigma yellowish, but anteriorly and posteriorly darkened (Fig. 232); wing membrane subhyaline.

Variation. Length of fore wing 4.3–4.4 mm; antennal segments of ♀ 38(1), 39(9), 40(44), 41(44), 42(27), 43(13), 44(2), 45(1); of ♂ 37(1), 38(4), 39(31), 40(82), 41(113), 42(33), 43(12), 44(7), 45(3); vein r of fore wing 0.2–0.5 × vein 3-SR; length of first tergite of ♀ 1.1–1.2 × its apical width; pterostigma medially yellow or dark brown; medio-posteriorly mesoscutum black or brownish yellow; second and third tergites yellowish, infuscate or dark brown laterally, mesosternum orange brown or infuscate.

The two sexes have about the same number of antennal segments. The lectotype of A. nigriceps has the antenna mutilated; according to the original description it had 40 or 41 segments. Aleiodes nigriceps is often considered to be a synonym of A. circumscriptus, e.g. Papp (1985) but the selection of a neotype for A. circumscriptus in the present work resolves that issue (and the two species differ in, among other things, the number of antennal segments).

Lectotype, ♀ (KBIN) from Belgium examined.

*Austria, Belgium, British Isles (England: V.C.s 2, 3, 7, 11, 16, 17, 20, 22, 23, 24, 25, 29, 30, 31, 34, 38, 57, 58, 59, 60, 63, 64, 65, 66, 67, 69; Wales: V.C.s 35, 44, 48, 52; Scotland: V.C.s 72, 75, 78, 79, 83, 85, 86, 87, 88, 89, 90, 95, 96, 97, 98, 99, 101, 102, 105, 107, 108, 111; Ireland: V.C.s H21, H28, H29, “Westport”), Czech Republic, *Denmark, France, Finland, Germany, Italy, Netherlands (GE: Ede; Nunspeet; Heerde; Otterlo, Velp, LI: Grubbenvorst, NB: Geertruidenberg; Helvoirt; Bergen op Zoom; Etten-Leur, NH: Overveen; Muiderberg, ZH: Den Haag; Meijendel; Oegstgeest; Voorschoten; Asperen; Waarder), *Norway, Poland, Russia, Slovakia, Sweden, Switzerland. Specimens in NMS, BMNH, RMNH, BZL, MTMA, NRS, SDEI, CC, FMNH, ZSSM, OUM, NMI, CMIM, Sheffield Museum, SMNS, ZISP, SYKE, ZMUO, USNM, CNC, UWIM, M. Riedel collection, JLC, MSC, AAC, WAE, I. Kakko collection, H. Haraldseide collection.

MRS216 (Scotland EU979585, CO1 + AJ784934, 28S), MRS373 (Sweden KU682258, CO1), MRS790 (Scotland KU682254, CO1), MRS794 (England KU682255, CO1).

A plurivoltine parasitoid of Noctuidae, using Apamea species in which to overwinter, and (possibly exclusively) Orthosia gothica (Linnaeus) in early summer. The mummy is moderately dark brown, rather elongate (Fig. 244) and often found in exposed positions. Specimens of the overwintering generation reared from noctuids identified as Apamea crenata (Hufnagel) (2 ZSSM, 1 NMS, 1 NRS; E. Haeselbarth, R.I. Lorimer), Apamea epomidion (Haworth) (2 ZSSM, 1 OUM; E. Haeselbarth), Apamea ? monoglypha (Hufnagel) (1 NMS; M.R. Britton), Apamea ? remissa (Hübner) (1 H. Haraldseide/Norway) and ?Apamea sp. (11, hosts unidentified or misidentified but with preserved mummies that are consistent with Apamea species (det. M.R. Shaw), having at least moderately large shining warts and a well-developed prothoracic plate: 7 (2 as Xestia xanthographa (Denis & Schiffermüller)) NMS, 4 (as Polia spp.) OUM, 1 (as Noctua fimbriata (Schreber)) RMNH, 1 AAC). Specimens of the summer generation reared from Orthosia gothica (3 NMS; R.I. Lorimer, M.R. Shaw), and unidentified noctuid mummies compatible with O. gothica (8 NMS, 1 ZMUO, 1 ZSSM, 2 H. Haraldseide collection). Other specimens (both generations) with a host recorded (Mythimna ferrago (Fabricius) in ZSSM, Epirrita autumnata (Borkhausen) in CC) have not been accompanied by mummies and we regard the records as dubious. Experimental results from female ex Apamea ? monoglypha; with 2^nd instar hosts: Orthosia gothica 1:9\9\\6\3, Orthosia cerasi (Fabricius) 1:10\ 0\\-, Orthosia incerta (Hufnagel) 1:4\0\\-; with 3^rd instar hosts O. gothica 1:8\8\\0\8. The single female trialed with hosts undertook considerable non-destructive concurrent host feeding on O. gothica. Adult flight time approximately May to August, with females persisting until October.