Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 1: Introduction, key to species groups, outlying distinctive species, and revisionary notes on some further species

Abstract Seven new species of the genus Aleiodes Wesmael, 1838 (Braconidae: Rogadinae) are described and illustrated: Aleiodes abraxanae sp. n., Aleiodes angustipterus sp. n., Aleiodes artesiariae sp. n., Aleiodes carminatus sp. n., Aleiodes diarsianae sp. n., Aleiodes leptofemur sp. n., and Aleiodes ryrholmi sp. n. A neotype is designated for each of Aleiodes circumscriptus (Nees, 1834) and Aleiodes pictus (Herrich-Schäffer, 1838), and both species are redescribed and illustrated. Aleiodes ochraceus Hellén, 1927 (not Aleiodes ochraceus (Curtis, 1834)) is renamed as Aleiodes curticornis nom. n. & stat. rev., and redescribed and illustrated. Aleiodes bistrigatus Roman, 1917, Aleiodes nigriceps Wesmael, 1838, and Aleiodes reticulatus (Noskiewicz, 1956), are re-instated as valid species. A lectotype is designated for Aleiodes bistrigatus Roman. An illustrated key is given to some distinctive species and the residual species groups along which further parts of an entire revision of western Palaearctic species of Aleiodes and Heterogamus will be organised. Biology, host associations and phenology are discussed for the keyed species (in addition to the above, Aleiodes albitibia (Herrich-Schäffer, 1838), Aleiodes apiculatus (Fahringer, 1932), Aleiodes arcticus (Thomson, 1892), Aleiodes cantherius (Lyle, 1919), Aleiodes esenbeckii (Hartig, 1834), Aleiodes jakowlewi (Kokujev, 1898), Aleiodes modestus (Reinhard, 1863), Aleiodes nigricornis Wesmael, 1838, Aleiodes pallidator (Thunberg, 1822), Aleiodes praetor (Reinhard, 1863), Aleiodes seriatus (Herrich- Schäffer, 1838) sensu lato, Aleiodes testaceus (Telenga, 1941), Aleiodes ungularis (Thomson, 1892), and Aleiodes varius (Herrich-Schäffer, 1838)) which are dealt with in full here (with the exception of Aleiodes seriatus s.l. which is, however, included in the key). The experimental methodology covering the revision as a whole, which involves some behavioural investigation, is outlined.


Introduction
Over the past 45 years, during a programme aimed at investigating host relations of Ichneumonoidea in which Rogadinae and some other cyclostome braconid groups have been strong foci, the second author has reared many western European species of Aleiodes from their Lepidoptera hosts, and also received donations of specimens reared by a large number of lepidopterists. Investigation of Aleiodes host ranges has also involved some experimentation using short-term cultures by the second author, and in some cases this has been motivated by, and crucial for, elucidating species-level taxonomy. The large amount of collected data will be used for a revision of the western Palaearctic species of the genera Aleiodes and Heterogamus, also covering the host range, phenology and other aspects of biology of as many species as our data permit.
In this first paper we give a key to the species groups that will be dealt with in further parts, and to some of the more distinctive species which are then dealt with in full here. Some species that urgently need valid names are newly described and/or redescribed to clarify the confused nomenclature and status of some nominal species. One valid species (A. ochraceus Hellén, 1927) is renamed as A. curticornis (nom. n. & stat. rev.), because Hellén's name is a junior homonym; and moreover not a synonym (as proposed by Papp 1985) of A. gastritor (Thunberg, 1822). Some parts of the genus appear to have radiated relatively recently and even in the well-studied British fauna it is probable that further biological research will reveal that some of the taxa currently recognised as single species are in fact aggregates of biologically distinct entities resistant to morphological separation. An example is given in this paper with the A. pictus-aggregate, and a relevant speciation hypothesis has been suggested by M.R. Shaw (1994Shaw ( , 2002, M.R. Shaw and Horstmann (1997) and Stigenberg and M.R. Shaw (2013). We outline the experimental methods involved in the short-term culture experiments involved in elucidating that complex (which will apply also to other parts of the revision), and the ways in which both natural rearings and experimental results are and will be presented. Overview information on the general developmental biology of Aleiodes will be given elsewhere (M.R. Shaw, in prep.), though particular points of interest will be mentioned under the species concerned and, to facilitate that, a brief description of the "normal" oviposition sequence is given in this paper.

Specimens, methods and presentation of records
Unless stated otherwise the following protocols apply here and to ensuing parts of this work. Distributions are based only on material studied by us, unless otherwise stated. Literature records (e.g. as in Shenefelt 1975 andYu et al. 2012) indicate much wider distributions for many of the species but, while this may often be true, we have in most cases no means of verifying the records. Indeed, we have noted a high level of previous misidentification in the material we have been able to examine, as is usual during taxonomic revision of poorly known groups. Country records that are not (at the time of writing) given in Fauna Europaea are indicated with an asterisk (*). In cases in which we have seen large numbers of specimens the distributional data are given only in summary form. For Britain this is usually just as Watsonian Vice County numbers (Dandy 1969) in order within countries but, when very numerous paratypes are involved (e.g. as in A. leptofemur sp. n.), the British Isles data are given as the names of Watsonian Vice Counties. For The Netherlands and other countries the province and locality are mentioned when possible. Host records are similarly based only on material that we have examined personally, although the host-ranges given here are likely to be a suitable basis from which to evaluate the probability that other host records (e.g. as in Shenefelt 1975 andYu et al. 2012) are accurate. Host determinations taken from specimen labels are subject to the usual possibilities of host misidentification (including the possibility that overlooked "mummies" (= the structure within which the Aleiodes pupates, formed of the hardened host larval skin) of radically different hosts were undetected in mass-rearing programmes of particular host species), although host mummies have often been retained with specimens permitting a re-evaluation of host determination in many cases, quite often leading to a different conclusion. Rather than transcribing often obsolete or misspelt names from data labels, we have updated nomenclature when appropriate. Some of the records that we believe to be particularly anomalous or to require extra interpretation are discussed further. However, the majority of the reared specimens have resulted from special rearing efforts under carefully controlled conditions (M.R. Shaw 1997) undertaken by one of us (MRS), and much material supporting these records is deposited in the National Museums of Scotland, Edinburgh (NMS); in additional cases the relevant depository is given. The host records are given quantitatively for reasons outlined by M.R. Shaw (1994): the number immediately following each host name indicates the number of examined specimens reared from that host, and the number separated by a colon that sometimes follows plural records indicates the number of localities from which the records result, if it is fewer than three. The majority of our host records are of British rearings, and the names of hosts that are included only as a result of non-British rearings are followed by the name of their country of origin. All rearing records given in quantitative form are from wild-collected hosts. Experimental investigation of host range has been undertaken for several British species, and the results are given separately for each Aleoides/ host pair in the summary form N: x \ y \\ p \ q + r, where N = the number of female parasitoids involved, x = the number of that host species offered, y = the number of hosts accepted, p = the number of accepted hosts that survived (mortality includes cannibalised hosts and dead hosts that couldn't be dissected), q = hosts that produced mummies of the Aleiodes (in a few cases this includes hosts that were dissected to reveal healthy parasitoid larvae with gut content), r = produced healthy Lepidoptera pupae. A concise summary of host range, to suggest searching environment as well as host taxa, is given for all species for which information is sufficient.
Except for the limited time it took to perform experiments and/or service rearing containers indoors, all livestock (including wild collected caterpillars harbouring parasitoids; Lepidoptera cultures; Aleiodes mummies awaiting emergence; adult Aleiodes, from whatever source; experimentally parasitized and control caterpillars) was kept in an unheated, well-ventilated and fully shaded detached outdoor shed that held outdoor shade temperatures (Reading, UK and Edinburgh, UK) generally to within 0.5 °C of ambient (M.R. Shaw 1997), except that adult Aleiodes to be used for experiments were generally removed to cooler conditions when the shed temperature reached 16-20 °C. They were kept individually in corked 2.5 × 7.5 cm sterilised (household bleach) glass tubes (hereafter "tube(s)") and dilute (initially 1:3) honey:water droplets were touched onto the insides of the tubes, and replenished or diluted as required to maintain ad libitum access to honey dilute enough to be imbibed. As far as possible, honey was sourced from areas remote from arable agriculture, and, in cases when the adult parasitoid had overwintered, a paste of pollen (obtained from health food shops) in dilute honey was additionally offered. Fresh tubes were used as soon as any sign of mould arose, and under these conditions females of most species could routinely be kept alive for at least 2 months; often considerably longer (regularly 4-6 months) and usually for well over a year in the case of species that overwinter as diapausing adults. Lepidoptera cultures used in experiments were obtained as eggs from captive females. Trials involving known or putative hosts were conducted by introducing a single female parasitoid to a single active host caterpillar of appropriate size (generally 2 nd instar) in a clean tube without vegetation; putative hosts were judged rejected after 10 minutes (prolonged to 5 contacts if these had not occurred in the time). Occasionally more than one species, or a range of sizes, were offered simultaneously, but in these cases rejections were scored only if all hosts simultaneously offered were rejected. Sometimes runs of parasitized hosts could be extended by offering hosts in proecdysis, as these are often accepted, but rejections were not scored using hosts that were subactive in this way. Similarly, because Aleiodes species are synovigenic, for scored trials no female was offered hosts after four ovipositions on that day (although most species are probably capable of up to at least 8 ovipositions on a reasonably warm day), but runs of fully acceptable hosts were sometimes extended beyond that. In addition to egg depletion, it was often clear that the supply of venom used to induce pre-oviposition paralysis was subject to temporary exhaustion, sometimes more so than eggs. Parasitized and (separately) control hosts were reared on carefully searched, clean, wild food plants as counted cohorts in 14 × 8 × 6 or 18 × 12 × 6 cm plastic boxes bottom-lined with 4-6 sheets of absorbent tissue (white lavatory roll); food and tissue was changed at least weekly, when each caterpillar was rigorously accounted for. All mummies were removed when found, allowed to dry in open air, and then placed in tubes to await adult emergence (inspected at least daily except in winter).
Mating trials were conducted by introducing a female to a fed male already present in a clean tube (not the other way round); generally, fresh tubes were used for each pair, and trials were done with newly emerged females (if possible before she had fed). The results of mating trials are given only impressionistically, as no way was found to quantify them satisfactorily.
The collections used for our revision contain the majority of recently collected material of Aleiodes from the western Palaearctic region; collections with type material are separately listed under the description of the species.  (Zoologische Staatssammlung, München). In addition we have seen specimens from various smaller and private collections, which are cited in significant cases.
The number of antennal (i.e. flagellar + 2) segments is frequently an important aid to species recognition and of interest also because in some species the female has more segments on average than the male (while other species are more normal in that the male has the greater number). We give counts of antennal segments for the specimens we have examined, but for some species (especially when the segments did not need to be counted for determination) sometimes only for the first hundred or so of the specimens examined of each sex.
GenBank accession numbers are given for DNA sequences from specimens in the NMS collection bearing a "MRS Aleiodes DNA [number]", in this text simplified to MRS[number] followed in parentheses by the country of origin then GenBank accession number and gene fragment. Not all sequenced samples have been accessioned, but for each species for which we have sequences we give several if available. Further, the DNA data of many taxa and putative taxa have been produced over a long number of years under several auspices: many sequences have been used in other studies of various kinds, often not taxonomic, and often under provisional or unpublished names. Thus many of those previously submitted to GenBank and BOLD had inappropriate names applied, but the names associated with MRS vouchers and accession numbers in this paper supplant previous identifications.
For the recognition of braconid subfamilies, see van Achterberg (1990Achterberg ( , 1993Achterberg ( , 1997, for the identification of Aleiodes Wesmael, see van Achterberg (1991) and Chen and He (1997), and for the terminology used in this paper see Figs 1-6 and in van Achterberg (1988van Achterberg ( , 1993note, however, that in the present work the distance between eye and lateral ocellus is measured differently). For additional references see Yu et al. (2012).
Oviposition. The oviposition behaviour of Aleiodes species is based on the following sequence, from which one or more steps may habitually be eliminated by particular species: (a) antennation of the host, often also investigation using fore and sometimes mid tarsi, during which the host often curls and may be drawn in towards the ventral/ mesosomal region of the parasitoid; (b) a rapid sting (usually less than 0.5 second), executed more or less between the parasitoid's front legs and usually accompanied by a brief fluttering of the wings; (c) waiting motionless by, but often not in physical contact with, the host while temporary paralysis caused by the injected venom takes effect (about 20 to exceptionally 90 seconds); oviposition (a single insertion of the ovipositor, usually about 30-80 seconds duration but regularly much shorter or much longer in certain species); (d) a period (usually about 20-100 seconds) of post-oviposition association, when the parasitoid stands over the host and the host is intermittently antennated, during which time the host recovers from paralysis; (e) abrupt and energetic departure, often by flight. Sluggish hosts are generally unattractive, but superparasitism is frequent if (e) is prevented or if the two come into contact again. In most species host feeding was seen only infrequently or not at all in well-fed parasitoids, but it became commoner Figures 1-6. Terminology and measurements used in this paper. 1 wing venation: pa = parastigma, pt = pterostigma, 1 = marginal cell, 2a, b, c = first, second and third submarginal cell, respectively, 3a, b = first and second discal cell, respectively, 4a = first subdiscal cell, 5 = basal cell, 6 = subbasal cell 2 head dorsal: a = length of eye, b = length of temple 3 head lateral: c = width of temple, d = width of eye, e = height of eye, f = width of malar space (measured as actual true distance in its own plane) 4 head anterior: g = width of face, h = width of hypoclypeal depression 5 fore femur lateral: i = length, j = width 6 first metasomal tergite dorsal: k = length of tergite (measured from adductor), l = apical width of tergite. in aged females; it was always non-destructive and concurrent (i.e using the same host individual as for oviposition) but took place from separate ad hoc wounds made using the ovipositor, usually before but occasionally after oviposition itself. In most species, first instar hosts are oviposited into only with difficulty and even then they frequently die from the trauma, second and early third instars are the most suitable, and from late in the third instar onwards hosts are consistently ignored (a rough guide is that if the host exceeds the length of the parasitoid it will usually be of no interest). In the majority of investigated species the egg floats freely in the haemocoel.
Distribution. Cosmopolitan. Notes. Two papers with descriptions of the same Aleiodes species appeared in 1838. Most likely Herrich-Schäffer's paper was published earlier (the introduction is dated April, 1838) than Wesmael's paper. Baron de Stassart stated in his presidential report (Bulletins de l'Académie royale des sciences, des lettres et des beaux-arts de Belgique 5: 328) dated May 6 th , 1838, that the 11 th volume of the Nouveau Memoires was in press.

1
Hind trochantellus of female moderately elongate, its ventral length 2.4-4.5 × its width (a), hind wing narrow (b), its vein 1r-m strongly reclivous (c) and vein r of fore wing 0.8-3 -Hind trochantellus of female usually moderately robust (aa), if ventrally 2.4-2.8 × as long as wide, then hind wing wider (bb) and its vein 1r-m moderately reclivous (cc) or vein r of fore wing shorter than 0.6 × vein 3-SR (dd); second submarginal cell of fore wing often longer than high (ee); genus Aleiodes Wesmael, 1838 ....... 2 2 Ovipositor sheath largely glabrous (except apically and ventrally) (a); marginal cell of hind wing narrowed near basal 0.6 and slightly widened apically (b); lateral carina of scutellum strong (c) and lunula rather narrow, but widened medially ( -Fore femur slender, (6.0-)6.7-7.4 × as long as wide (aa) and very finely sculptured; scapus and pedicellus more or less infuscate or black ventrally (bb); precoxal area usually without rugae (cc); propodeum largely coriaceous medially and median carina at least anteriorly present on posterior half of propodeum and regular (dd); posterior half of pterostigma of ♀ more or less yellowish (ee), but usually apical third laterally darkened; pale yellowish part of malar space usually not reaching clypeus (ff); mesosternum frequently reddish or brownish ( -Length of fore femur 5.4-6.4(-8.0) × its maximum width (aa) and hind femur more or less weakly swollen (bb); or fore femur more than 6.4 × (aaa) and hind femur comparatively wide basally (cc), antenna with more than 40 segments and mesosternum yellowish or orange-brown (dd) or temple slightly wider (ee), or scapus ventrally and basal half of antenna yellowish brown (ff); scapus usually yellowish ventrally, if dark brown or blackish then scapus similarly coloured as hind femur subapically; apical half of hind femur (partly) conspicuously dark brown ( Biology. 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.
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.
Note. The males of this species have on average about one antennal segment less than females.

Aleiodes albitibia (Herrich-Schäffer, 1838) Figs 20-34
Rogas albitibia Herrich-Schäffer, [April] 1838: 156;Shenefelt 1975Shenefelt : 1217van Achterberg 1991: 24 (as senior synonym of Aleiodes heterogaster). Aleiodes albitibia; van Achterberg 1991: 24;Belokobylskij et al. 2003: 398. Aleiodes heterogaster Wesmael, [May] 1838: 96;Shenefelt 1975Shenefelt : 1176Papp 1991: 97 (examined). Rhogas heterogaster ;Fahringer 1932: 258-259. Rogas heterogaster;Hammond and Smith 1957: 181;Tobias 1986: 82 (transl.: 136).  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 minutesin 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 June-August.
Diagnosis. 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 (  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.
Description. Holotype of A. heterogaster, length of fore wing 5.4 mm, of body 5.6 mm.
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.
Notes. 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 (pace Hammond and Smith 1957). Biology. 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  (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).
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.
Note. 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. (Fahringer, 1932 (Fahringer, 1932); paratype in BMNH examined). Aleiodes negativus ;Shenefelt 1975Shenefelt : 1165.  Biology. 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.

Aleiodes apiculatus
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 longlived 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.
Diagnosis. 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 ( Description. Redescribed ♀ (NMS) from England, length of fore wing 5.9 mm, of body 6.8 mm.
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.
Notes. 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.
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).
Notes. 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. Biology. 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.
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).
Etymology. From the specific epithet of its host. Distribution. *France.
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.
Note. Possibly a Faroe Islands endemic.

Aleiodes cantherius (Lyle, 1919) Figs 99-112
Rogas cantherius Lyle, 1919: 153-154 (examined 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.
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.
Note. The two sexes have about the same number of antennal segments. Biology. 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.
Diagnosis. 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.
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.
Etymology. From "carmino" (Latin for "comb"), because of the comb on the hind tibia.

Note.
Males have on average about 2-3 more antennal segments than females.  3,4,11,12,14,17,20,22,23,25,26,27,28,30,31,32,33,58,61,62,63,64 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. Diagnosis. 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.
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).
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.
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)  Biology. 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.
Diagnosis. 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.
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.
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.
Etymology. Named after the generic name of its host: Diarsia Hübner. Distribution. *British Isles (England, Wales, Scotland), *France, *Netherlands. Note. Males have on average about 3-4 more antennal segments than females.  Biology. 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) popula- tions 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.
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.

Notes.
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.
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 ; 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.
Note. From limited data males appear to have fewer antennal segments than females.  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. Diagnosis. 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.
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) (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 summergeneration 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.
Etymology. This common and widely distributed species is named after its slender femora ("leptos" = Greek for "thin").
Notes. 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.
Additional material. 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 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.
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.
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.
Note. The number of antennal segments does not differ appreciably between the sexes. The specimens reported from Hungary by Papp (1983Papp ( , 1985aPapp ( , 2005 are misidentified A. fortipes (Reinhard).
Diagnosis. 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).
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.
Notes. 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).
Diagnosis. Antennal segments of female 44-49, of male 43-47; antenna dark brown or black (also scapus and pedicellus more or less infuscate or black ventrally), rarely completely yellowish brown; temples directly narrowed behind eyes; OOL about equal to diameter of posterior ocellus (Fig. 252); pale yellowish part of malar space usually not reaching clypeus (Figs 251, 253); precoxal area usually without rugae; mesosternum usually reddish or brownish; propodeum largely coriaceous medially and median carina at least anteriorly present on posterior half of propodeum and regular; mesosoma (especially mesoscutum and scutellum) black (or blackish) dorsally, but notaulic area may be brownish posteriorly; fore femur of ♀ 6.7-7.4 × as long as wide (Fig. 250) and very finely sculptured; posterior half of pterostigma of ♀ more or less yellowish, but usually apical third laterally darkened; hind femur of ♀ rather reddishbrown, but may be largely infuscate in males; vein 1-CU1 of fore wing horizontal and vein cu-a short, far postfurcal (Fig. 245); metasoma largely blackish with (pale) yellowish elliptical patch medially (Fig. 248).
Metasoma. First tergite 1.2 × as long as wide posteriorly, flattened and latero-posteriorly lamelliform; first-second tergites and base of third tergite coriaceous and finely irregularly longitudinally rugose; second tergite rather robust (Fig. 248), with median carina and rather shiny; medio-basal area of second tergite obsolescent; second suture narrow and distinctly crenulate; remainder of metasoma largely superficially coriaceous and shiny; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate.
Notes. The lectotype has 46 antennal segments. On average males have 1-2 fewer antennal segments than females.

Biology.
A univoltine, thelytokous specialist parasitoid of the erebid lymantriine Leucoma salicis (Linnaeus), overwintering in the host. More than 200 reared specimens seen from L. salicis (most in NMS, others in BMNH, RMNH, ZSSM, OUM, CC, MCZ). Its biology has been studied by Dowden (1938) in the course of its attempted introduction to North America for the control of the introduced L. salicis: the notes given here supplement rather than repeat his findings. Based on English data the adult flight time is from the last few days of June through to early September. In experimental rearings the females were slow to accept L. salicis larvae, but always did so eventually after repeated contact (being especially attracted to traces of silk), and second instars were only marginally more acceptable than firsts. This may suggest some adaptation to the essentially gregarious nature of early stage L. salicis larvae. Oviposition was brief, usually taking about 5 but sometimes up to 20 seconds: although there was a pre-oviposition sting the female usually did not wait for paralysis to take effect before ovipositing, but would then avoid superparasitism of hosts thus rendered sluggish for a short period, though not subsequently. Partly because of winter mortality exact quantitative data are not available, but from about 100 observed single ovipositions the success rate in L. salicis (partly judged from living established larvae within overwintering hosts) was at least 90%. In one experiment, already briefly reported by Askew & Shaw (1986), a cohort of L. salicis larvae (N = 57) parasitised by A. pallidator and kept under outdoor conditions came out of their hibernacula a mean of 9.2 days later than controls (N = 66) from the same host egg batch (t = 9.18, P < 0.001). This delay was interpreted as extending the range of host plants suitable for the development of parasitised hosts, as spring bud burst varies greatly between the Salicaceae present at the English site of origin (Ainsdale, Lancashire); in addition, however, this behaviour by a monophagous parasitoid might be an adaptation to ensure that there is reasonably good synchrony with the next host generation. In experiments to test host range extension, inexperienced females would not oviposit into other species of Lymantriinae, but females that had already oviposited into L. salicis often would do so quite readily into both Euproctis similis (Fuessly) and Dicallomera fascelina (Linnaeus), although attempts were often at least for a time thwarted by the longer setae of the trial hosts. In all cases in which oviposition occurred, hosts were later dissected and found to contain encapsulated parasitoids (usually eggs; possibly in some cases first instar larvae). The results of trials were E. similis 2:15\8\\0, D. fascelina 3:12\6\\0, Orgyia antiqua (Linnaeus) 4:7\0\\-, Lymantria dispar (Linnaeus) 1:5\0\\-. The penultimate (or earlier) instar L. salicis larva is induced to prepare a frail cocoon, as though for pupation, within a leaf package just before being mummified, and the mummy forms within that structure with only its setae in contact (Fig. 257). Although the parasitoid larva does make a ventral opening in the host's thoracic region, through which fluid escapes and dries, the mummy is not thereby stuck down.
Metasoma. First tergite 0.9 × as long as wide posteriorly, flattened and lateroposteriorly lamelliform; first-second tergites and base of third tergite densely finely irregularly rugose and with median carina; second tergite robust and 1.5 × as long as third tergite (Fig. 263); medio-basal area of second tergite absent; second suture deep and distinctly crenulate; apical half of third tergite granulate-coriaceous, remainder of metasoma largely superficially coriaceous and rather shiny; fourth tergite largely with sharp lateral crease; ovipositor sheath largely densely setose and apically rounded.
Notes. Apart from a single specimen (reared from L. salicis probably in Russia, with its mummy present) in poor condition in NMS, no males of A. pallidator have been seen; examined males identified by J. Papp or V.I. Tobias as A. pallidator belong to A. gastritor (Thunberg) s. lat. or to a species near A. abraxanae with darkened pterostigma, black stemmaticum and widened hind femur (e.g. females reported by Rezbanyai-Reser (1996, 1997)). The lectotype of I. pallidator has 53 antennal segments and the ocelli somewhat larger than OOL. The holotype of R. ochraceus has the ocelli nearly twice OOL. Papp (1985) synonymised A. apiculatus (Fahringer, 1932) with A. pallidator, but we consider it to be a valid species.
Metasoma. First tergite 1.1 × as long as wide posteriorly, convex, but posteriorly flattened; first and second tergites densely longitudinally rugose; second tergite slen- derer than is usual in A. nigriceps (Fig. 276), with distinct median carina, with satin sheen; medio-basal area of second tergite absent; second suture distinctly impressed and crenulate; third tergite largely coriaceous (but anteriorly rugose) and medially 0.7 × as long as second tergite (Fig. 276); remainder of metasoma largely nearly smooth and rather shiny; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath moderately setose.
Note. Males have on average about one more antennal segments than females. In some populations pale specimens (including hind femur and much of face) occur that superficially resemble A. leptofemur, but can be distinguished by their more robust femora. For further notes see A. nigriceps.

Aleiodes praetor (Reinhard, 1863) Figs 285-301
Rogas praetor Reinhard, 1863: 264;Shenefelt 1975Shenefelt : 1244Tobias 1986: 78 (transl.: 128). Aleiodes praetor ;Papp 1991: 73;Belokobylskij et al. 2003: 398. Neorhogas luteus Szépligeti, 1906Shenefelt 1975Shenefelt : 1205Papp 1977: 115 (as synonym of A. praetor);1991: 73;2004: 215 11,16,17,19,20,21,22,24,30,31,34,38,62,64) Biology. Aleiodes praetor is a univoltine parasitoid of at least some arboreal Sphingidae, and overwinters in the host mummy. Reared specimens seen were from Lathoe populi (Linnaeus) (1 CMIM; C. Morley), Mimas tiliae (Linnaeus) (2 OUM, 1 NMS, 1 BMNH, 1 RMNH; J.C. Fraser, J. Koorneef, R.A. Softly). In Britain the flight period is from late June through August. A series of males was reared in culture in both L. populi and M. tiliae parented by a virgin female from M. tiliae. The female was often seen hanging from a leaf edge by only a few of her legs (Fig. 285). Most ovipositions, into late first instar and more particularly second instar hosts, occurred in a rearing cage overnight and were unobserved; however two of these remarkable occasions were witnessed (into late first instar L. populi). The host, which rests and feeds from the under surface of the leaf, was approached and repeatedly touched with the outstretched extreme tips of the antennae, causing the host to twitch more or less violently from side to side. It is noteworthy that the apical segment of the antenna in this species has a well-pronounced nipple-like tip (Fig. 297). When this reaction wore out, the parasitoid suddenly jumped on the host and rapidly inserted her ovipositor, with the metasoma scarcely curled; she then immediately straightened her body and released her footing completely so that she hung from the host with only her ovipositor touching it, and all legs completely free of any support. After 30 seconds she jerked free, and took flight as she fell from the host, which was apparently not paralysed to any extent although it was quiescent during oviposition. The extraordinary oviposition behaviour is clearly facilitated by the unusual flange and teeth at the apex of the ovipositor (Fig. 295), and may be completely constraining: otherwise suitable hosts on the floor of containers (i.e. lacking a drop) were consistently just walked over or otherwise ignored. The lack of paralysis ensures that the host maintains its footing, without risk that it would fall and be unable to rediscover its food source. The mummy appears to be highly adapted for a lengthy persistence in crevices in tree bark. It is very hard, matt, and predominantly light greyish brown in colour but with darker transverse variegation and sometimes small dark grey dorsal patches (Fig. 287). The parasitoid occupies abdominal segments (4-)5-8 which become thinly lined with silk and weakly arched. This structure is strongly stuck down ventrally at about the fourth abdominal segment, with the anterior part of the host becoming physically detached at an oblique angle by the action of the parasitoid larva. In captivity the stricken hosts sought refuge in paper tissues at the base of the rearing boxes, where mummies were made glued firmly in surface folds, and, despite Morley's (1916) finding a mummy on a Populus leaf, it is clear that the penultimate instar host larva is normally induced to descend and find a crevice before perishing; indeed, a partly grown M. tiliae larva which was collected on the bark of a Tilia tree was mummified within a few hours (R.A. Softly, personal communication). Before the widespread use of UV lights by lepidopterists A. praetor was rarely collected in Britain; now, however, specimens turn up quite regularly in light traps.
Diagnosis. Large yellowish brown species with antennal segments of female 67-77 and of male 62-75; OOL 0.3 × diameter of posterior ocellus; lateral carina of scutellum strong and lunula rather narrow; marginal cell of hind wing narrowed near basal 0.6 and slightly widened apically (Fig. 289); inner apex of hind tibia with weak and indistinct comb; tarsal claws yellowish setose; ovipositor sheath largely glabrous (except apically and ventrally; Fig. 301); ovipositor with small teeth ventrally and with wide dorsal flange (Fig. 295); length of fore wing 8-10 mm; parasitoid of Sphingidae.
Metasoma. First tergite as long as wide apically (Fig. 291); first and second tergites densely and coarsely longitudinally rugose, with distinct median carina, reduced near apex of second tergite; medio-basal area of second tergite absent; length of second tergite 0.7 × its basal width; second suture deep and distinctly crenulate; third tergite 0.9 × as long as second tergite, anterior half largely densely and finely punctate and remainder of metasoma largely smooth and depressed; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath largely glabrous (except apically and ventrally; Fig. 301); ovipositor with small teeth ventrally and with wide dorsal flange (Fig. 295).
Notes. European males have approximately four fewer antennal segments than females. Antenna of possibly conspecific Chinese and Japanese females consists of 70-77 segments and of males 62-75 segments and they have the pterostigma darker compared to the veins below it. (Noskiewicz, 1956), stat. rev.
Molecular data. MRS808 (Poland KU682262, CO1). Biology. The type series was reared in Poland from the ennomine gemetrid Macaria brunneata (Thunberg) (3 ♀, 3 ♂) feeding on Vaccinius myrtillus -and supposedly also Arctia caja (Linnaeus) (1 ♂) collected from the same plant, but we discount that as a presumed error (the specimen can no longer be found in PAN, its supposed depository). It is not surprising to add Macaria wauaria (Linnaeus) (4:1 [Belarus]; Silvanovich) and another possibly from this host (Russia) to the known host range, especially as these two Macaria species both overwinter in the egg stage (unlike many others). From material recently obtained from M. brunneata in the type locality and five other sites in Poland (M.R. Shaw), it is clear that A. reticulatus is a regular univoltine parasitoid of M. brunneata feeding on Vaccinium myrtillus growing as understory in conifer forests especially on infertile sandy soils (on one of the German specimens "Fichtenwald" translates as spruce forest), flying in early spring (April and May) which is no doubt why it has remained poorly understood until now. The small mummy (Fig.  303) forms at about the end of May and is firmly glued to a twig of the foodplant well below the crown, or frequently on twigs and conifer needles in the litter. It is brownish grey, with a dark brown posterodorsal patch corresponding to the site of eventual adult emergence, and rather short, broad and dorsally elevated. After summer diapause A. reticulatus overwinters as a prepupa (without defaecation but with well-formed eyes: ascertained by opening a mummy with living contents in December).
Head. Apical antennal segments missing, remaining 37 segments, length of antenna about as long as fore wing, its subbasal segments about 1.2 × as long as wide; frons mainly superficially granulate and with some rugae anteriorly, weakly shiny; OOL twice diameter of posterior ocellus and granulate as is vertex, with satin sheen; clypeus weakly convex, narrow and coriaceous; ventral margin of clypeus thick and depressed (Fig.  312); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 311) and face mainly coriaceous with some rugulae dorsally and long setae; length of eye 1.1 × temple in dorsal view and temple sub-parallel-sided behind eye (Fig. 315); occiput behind stemmaticum granulate and occipital carina nearly complete ventrally and narrowly interrupted dorsally; clypeus near lower level of eyes (Fig. 311); length of malar space 0.6 × height of eye in lateral view; eyes somewhat protruding (Figs 311-312, 315).
Note. This species is very close to A. arcticus but, in addition to small morphological differences, the fact that A. reticulatus is a lowland species while A. arcticus is boreoalpine is also regarded as significant.
Metasoma. First tergite as long as wide posteriorly, flattened and latero-anteriorly lamelliform near dorsope; first-second tergites and base of third tergite mainly coriaceous with superficial rugulosity, and with median carina weakly developed on apical half of first tergite and middle of second tergite; second tergite 0.9 × longer than wide basally and 1.3 × as long as third tergite (Fig. 320); medio-basal area of second tergite minute; second suture rather deep and distinctly crenulate; remainder of metasoma largely smooth and shiny; fourth tergite without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate.
Etymology. This species from Sweden is named after the collector of the type specimen, Swedish lepidopterist Nils Ryrholm, whose generous donations of Swedish parasitic wasps to the NMS collection have enriched it greatly.
Distribution. *Sweden.  Wesmael, 1838: 112;Shenefelt 1975Shenefelt : 1185Papp 1991: 107;Belokobylskij et al. 2003: 399 (as synonym of A. seriatus) [examined]. Rogas kuslitzkyi Tobias, 1976: 88, 223-224;1986: 83 (transl.: 137). Aleiodes kuslitzkyi; Belokobylskij et al. 2003: 399 (as synonym of A. seriatus).  . 339); surroundings of veins 1-M and 1-SR of fore wing more or less infuscate (Fig. 329); inner apex of hind tibia with comb (Fig. 340); metasoma of ♀ maculate ( Fig. 331; less so in males); fourth tergite of ♀ pale (ivory) yellowish latero-posteriorly, in ♂ usually infuscate or this tergite uniformly pale brown; base of hind tibia usually narrowly dark brown; length of hind femur of ♀ 5.1-6.5 × its width (of ♂ up to 8 x); fourth tergite gently folded laterally, without acute lateral crease or this only anteriorly developed, although rarely present as a simple, non-lamelliform crease to apex of tergite; precoxal area, epicnemial area and propodeum laterally, rugose; fourth tergite superficially transversely rugulose or aciculate; setose part of ovipositor sheath 0.6 × as long as hind basitarsus. The patterning of the metasoma is characteristic but very variable in extent (Figs 328, 331). Notes. We include this taxon only in outline, because DNA evidence (Quicke & Shaw, unpublished) suggest that an aggregate of two species in Europe and another in the Russian Far East currently going under this name remains unresolved. This will be addressed in a future paper (van Achterberg, Shaw & Quicke, in prep.), but here we include A. seriatus sensu lato in the key because the aggregate is morphologically isolated (within the region covered), easily recognised and does not fall logically into any of the species groups represented. The aggregate is very widespread in Europe, and at least one segregate is associated with Lithosiini (Erebidae: Arctiinae).
Type material. Lectotype ♀ (ZISP) from southern Russia (Astrakhanskaya Oblast) examined. Additional material. *Austria, British Isles (*England: V.C.s 2, 3,5,17,21,22,23,28,29,31,32,38,39,57,58,63 Biology. This is a plurivoltine parasitoid of Eupithecia (Geometridae: Larentiinae) and close relatives feeding on flowers, overwintering as an adult. Specimens in NMS reared from wild collected larvae identified as Eupithecia dodoneata Guenée ( Hosts were always associated with flowers, especially of trees (Quercus) and shrubby plants (Ligustrum, Clematis), but including field layer plants (Solidago, Torilis). Adults always emerge in the year of formation of the mummy, and females of this species overwinter as adults. In addition to a torpid specimen having been beaten from conifers in late December and another taken in early March (both M.W.R. de Vere Graham, southern England), this has been confirmed experimentally. However, in captivity the females tend to become active and require food above about 4 °C, indicating that the species is best adapted to areas where the winter is relatively short, as in southern Europe. Dissection of 5 (out of 51) female specimens collected by fogging Quercus canopy in mid September (southern England, N.E. Stork) revealed neither mature nor developing eggs, only moderately extensive fat bodies. As the species had been virtually absent from other fortnightly samples fogged from the same trees, it seems that they were feeding, perhaps on honeydew, prior to overwintering elsewhere. Because males are found late in the summer as well as early, it appears to be a plurivoltine species; however, it is unclear whether the autumnal specimens lacking mature eggs had resorbed them or refrained from developing them. Light trapped specimens tend to be female, but this is probably just a consequence of the much longer life span of the female sex, and in reared series the sex ratio is more equal. The light brown, banana-shaped, apically attenuated and short-lived mummy is interesting in not being stuck down at the thoracic area; instead (at least when forming from relatively elongate and fully exposed Eupithecia larvae such as those of E. dodoneata) it projects outwards at an angle similar to that normally adopted by the host at rest. None the less, a ventral opening is made by the parasitoid larva in the host's thoracic region at the time of mummification, through which a small quantity of fluid is expelled and quickly dries.
Head. Antennal segments 32, antenna as long as fore wing, its subbasal and subapical segments about 2.4 and 1.7 × as long as wide, respectively; frons mainly granulate and flat; OOL equal to diameter of posterior ocellus and very finely granulate as vertex, with satin sheen; clypeus weakly convex, medium-sized and granulate; ventral margin of clypeus thick and depressed (Fig. 350); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 349) and face granulate and with medium-sized setae; length of eye 3.2 × temple in dorsal view and temple gradually narrowed behind eye; occiput behind stemmaticum very finely granulate and occipital carina complete ventrally and widely interrupted dorsally; clypeus near lower level of eyes (Fig. 349); length of malar space 0.35 × height of eye in lateral view; eyes rather protruding (Figs 349-351).
Notes. This species is distinctive and (with the wide use of UV light traps by lepidopterists) proving to be rather common in southern England, but it seems to have been very rarely collected and generally overlooked as a British species until quite recently. However, it has probably been present for a long time; a British specimen in the Dale collection (OUM) is dated 1892. In The Netherlands known since 2006 but only from one locality in the southern province of Noord-Brabant. On average the males have 2-3 antennal segments more than females.
Second is "Aleiodes testaceus" of Spinola (1808) of which the original combination is Bracon testaceus. It has so often been considered to be a valid taxon that it is included as such in Taxapad (Yu et al. 2012). However, in the supposed description by Spinola (1808), the author refers to Ichneumon testaceus Fabricius, 1798, and it is, therefore, not a valid description of a new taxon. Nomenclaturally it is the same as Rogas luteus Nees, 1834 (see above), but subsequent authors have incorrectly interpreted it as another (or other) species. Rogas luteus Nees is a valid name and refers to the type species of the genus Rogas Nees, 1819, and its (unjustified) emendation Rhogas Agassiz, 1849. "Aleiodes testaceus (Spinola)" of authors usually refers to one or another of several brownish yellow ("testaceous") species of Aleiodes, most often the taxon which we consider to be correctly named as Aleiodes similis (Curtis, 1834), or taxa included in Taxapad under the species-aggregate of Aleiodes gastritor (Thunberg, 1822) s. l. (Yu et al. 2012). According to Art. 11.5 of the ICZN Code (1999), to be available a name must be used as valid for a taxon when proposed, unless it was first published as a junior synonym and subsequently made available under the provisions of Art. 11.6.1. The status of a previously unavailable name is not changed by its mere citation as a valid taxon (Art. 11.5.2) and Bracon testaceus was not described by Spinola as a new taxon. A name could become available according to Art. 11.10, if it was deliberately employed for a misidentified type species, but this does not apply to the Aleiodes testaceus of Spinola. In principle, to maintain prevailing usage of a misinterpreted name, the case could be brought to the Commission for a ruling, but in our opinion there are no good reasons to do this because the intention of Spinola (1808) is unambiguous, the name has been used for several taxa, and it is not an iconic name that should be preserved because of many unambiguous published citations outside of the field of taxonomy.
Finally, Heterogamus testaceus Telenga, 1941, was (correctly in our view) included in the genus Aleiodes by several authors, including Shenefelt (1975) and Bergamesco et al. (1995). Thus Aleiodes testaceus (Telenga) could be confused, nomenclaturally, with "A. testaceus (Spinola)" of authors (belonging mainly to the A. gastritor-aggregate or other taxa of what we refer to in this paper as the Aleiodes circumscriptus-group). The two are only distantly related according to Fortier and Shaw (1999), a conclusion with which we concur (including from unpublished molecular data; Quicke et al. in prep.). Included among the purposes of this paper is to clarify the differences between these taxa and to suppress the incorrect usage of the name "A. testaceus (Spinola)" of authors not (Telenga 1941), as for example by Quicke (2015).  behind eye (Fig. 363); occiput behind stemmaticum superficially granulate, narrow; clypeus largely above lower level of eyes; length of malar space 0.25 × length of eye in lateral view (Fig. 362); occipital carina nearly complete medio-dorsally, complete and strong ventrally.
Mesosoma. Mesoscutal lobes finely and densely granulate and with fine punctation, matt; prepectal carina strongly developed, lamelliform, not reaching anterior border of mesopleuron; precoxal area of mesopleuron medially impressed and crenulate, remainder of mesopleuron (except dorsal rugose area) superficially granulate, shiny and with some superficial punctures (but in other specimens largely smooth); metapleuron granulate-punctate; scutellum finely granulate and rather flat, only antero-laterally with distinct carina; propodeum convex, rather short, distinctly rugose and its median carina complete, without tubercles.
Metasoma. First tergite robust; first and second tergites and basal half of third tergite finely and densely rugose, with distinct median carina; medio-basal area of second tergite minute; second suture narrow and rather shallow; third tergite nearly as long as second tergite; remainder of metasoma micro-sculptured, depressed; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath shiny, setose and apically truncate.
Metasoma. First tergite 1.3 × as long as wide posteriorly, flattened and latero-anteriorly distinctly lamelliform; first-second tergites and base of third tergite densely finely regularly rugose and with median carina; second tergite slender, 1.1 × longer than wide basally and 1.4 × as long as third tergite (Fig. 370); medio-basal area of second tergite absent; second suture rather deep and distinctly crenulate; apical half of third tergite mainly rugulose-coriaceous, remainder of metasoma superficially coriaceous and rather shiny; fourth tergite largely without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate.
Note. Females have on average 1-2 more antennal segments than males.