no Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae) [1 ed.]

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ZooKeys 639: 1–164 (2016) doi: 10.3897/zookeys.639.10893

Revision of western Palaearctic Aleiodes Wesmael, I.

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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 Cornelis van Achterberg1, Mark R. Shaw2 1 Research Associate, Department of Terrestrial Zoology, Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, The Netherlands 2 Honorary Research Associate, National Museums of Scotland, Chambers Street, Edinburgh EH1 1JF, Scotland, U.K. Corresponding author: Cornelis van Achterberg ([email protected]) Academic editor: B. Santos  |  Received 23 October 2016  |  Accepted 27 November 2016  |  Published 12 December 2016 http://zoobank.org/BB23AA3F-DD9E-42CE-92F7-37E047AE80C7 Citation: Achterberg C van, Shaw MR (2016) 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. ZooKeys 639: 1–164. https://doi.org/10.3897/zookeys.639.10893

Abstract Seven new species of the genus Aleiodes Wesmael, 1838 (Braconidae: Rogadinae) are described and illustrated: A. abraxanae sp. n., A. angustipterus sp. n., A. artesiariae sp. n., A. carminatus sp. n., A. diarsianae sp. n., A. leptofemur sp. n., and A. ryrholmi sp. n. A neotype is designated for each of Aleiodes circumscriptus (Nees, 1834) and A. pictus (Herrich-Schäffer, 1838), and both species are redescribed and illustrated. Aleiodes ochraceus Hellén, 1927 (not A. ochraceus (Curtis, 1834)) is renamed as A. curticornis nom. n. & stat. rev., and redescribed and illustrated. Aleiodes bistrigatus Roman, 1917, A. nigriceps Wesmael, 1838, and A. 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, A. albitibia (Herrich-Schäffer, 1838), A. apiculatus (Fahringer, 1932), A. arcticus (Thomson, 1892), A. cantherius (Lyle, 1919), A. esenbeckii (Hartig, 1834), A. jakowlewi (Kokujev, 1898), A. modestus (Reinhard, 1863), A. nigricornis Wesmael, 1838, A. pallidator (Thunberg, 1822), A. praetor (Reinhard, 1863), A. seriatus (Herrich- Schäffer, 1838) sensu lato, A. testaceus (Telenga, 1941), A. ungularis (Thomson, 1892), and A. varius (Herrich-Schäffer, 1838)) which are dealt with in full here (with the exception of A. 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. Copyright C. van Achterberg, M.R. Shaw. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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Keywords Aleiodes, host range, biology, distribution, Europe, phenology

Table of contents Introduction ............................................................................................................. 3 Specimens, methods and presentation of records ...................................................... 4 Taxonomy ................................................................................................................ 8 Aleiodes Wesmael, 1838 ...................................................................................... 8 Key to West Palaearctic species groups, and outlying distinctive species, of the genus Aleiodes Wesmael ........................................................................ 11 Biology and descriptions .................................................................................. 34 Aleiodes abraxanae sp. n. ............................................................................. 34 Aleiodes albitibia (Herrich-Schäffer, 1838) .................................................. 39 Aleiodes angustipterus sp. n. ......................................................................... 44 Aleiodes apiculatus (Fahringer, 1932) .......................................................... 48 Aleiodes arcticus (Thomson, 1892) .............................................................. 54 Aleiodes artesiariae sp. n. ............................................................................. 58 Aleiodes bistrigatus Roman, 1917, stat. rev. ................................................. 62 Aleiodes cantherius (Lyle, 1919) .................................................................. 65 Aleiodes carminatus sp. n. ............................................................................ 69 Aleiodes circumscriptus (Nees, 1834) ........................................................... 74 Aleiodes curticornis nom. n. & stat. rev. ....................................................... 78 Aleiodes diarsianae sp. n. ............................................................................. 82 Aleiodes esenbeckii (Hartig, 1838) ............................................................... 88 Aleiodes jakowlewi (Kokujev, 1898) ............................................................ 94 Aleiodes leptofemur sp. n. ............................................................................ 98 Aleiodes modestus (Reinhard, 1863) ........................................................... 105 Aleiodes nigriceps Wesmael, 1838 .............................................................. 110 Aleiodes nigricornis Wesmael, 1838 ........................................................... 115 Aleiodes pallidator (Thunberg, 1822) ........................................................ 120 Aleiodes pictus (Herrich-Schäffer, 1838) .................................................... 125 Aleiodes praetor (Reinhard, 1863) ............................................................. 130 Aleiodes reticulatus (Noskiewicz, 1956), stat. rev. ...................................... 135 Aleiodes ryrholmi sp. n. ............................................................................. 140 “Aleiodes seriatus (Herrich-Schäffer, 1838)” sensu lato .............................. 143 Aleiodes testaceus (Telenga, 1941) .............................................................. 146 Aleiodes ungularis (Thomson, 1892) ......................................................... 152 Aleiodes varius (Herrich-Schäffer, 1838) ................................................... 156 Acknowledgements ............................................................................................... 159 References ............................................................................................................ 160

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Introduction As defined by van Achterberg (1991), the large genus Aleiodes Wesmael, 1838, has a world-wide distribution and in the western Palaearctic region it forms a moderately prominent element of the braconid fauna. Despite the relatively large size and/ or abundance of several species, and the frequency with which they are reared by lepidopterists, there are no reliable identification keys to the western Palaearctic species. Van Achterberg (1991), for the W. Palaearctic and Afrotropical region, recognised and keyed three subgenera of Aleiodes (Aleiodes (including Heterogamus Wesmael, 1838), Chelonorhogas Enderlein, 1912 and Neorhogas Szépligeti, 1906). Subsequently, from a morphological phylogenetic analysis of the genus Aleiodes as defined by van Achterberg (1991), Fortier and S.R. Shaw (1999) proposed 18 monophyletic species groups, one of which was further divided by Fortier (2006). Zaldívar-Riverón et al. (2008), in a molecular phylogenetic study of the subfamily Rogadinae, defined a tribe Aleiodini to include only two genera, the resurrected Heterogamus and Aleiodes, within which were placed Cordylorhogas Enderlein, 1920, Hemigyroneuron Baker, 1917 and Pholichora van Achterberg, 1991. Within this differently defined Aleiodes two main clades were recovered, one corresponding to Chelonorhogas in van Achterberg’s (1991) sense, and no evidence was found to support the monophyly of most of the species groups proposed by Fortier and S.R. Shaw (1999); although a few proposals (e.g. the pilosus-group (Tetrasphaeropyx Ashmead, 1888) later recognised by Fortier (2006)) may indeed concern monophyletic assemblages embedded within Aleiodes. The conclusion that most of the species groups proposed by Fortier and S.R. Shaw (1999) contain unrelated elements has been further corroborated by more recent molecular work (Butcher et al. 2012, Quicke and M.R. Shaw unpublished), and many characters used by Fortier and S.R. Shaw (1999) for defining species groups, such as pectination of the claws, are homoplastic. In the present work, which is restricted to the W. Palaearctic region, we not only reject the classification proposed by Fortier and S.R. Shaw (1999) but also, indeed, choose to leave the definition of formal species groups aside. For the purposes of the key, we are using informal species groups that can be defined by the characters given in the key, but our concept of the groups differ from those used by Fortier and S.R. Shaw (1999); in particular our circumscriptus- and apicalis-groups are used in a much wider context but, along with our bicolor-group, they do appear each to be monophyletic, at least within the W. Palaearctic (Quicke and M.R. Shaw unpublished). It should also be understood that our use of A. seriatus-agg. applies to a small group of cryptic species, not corresponding to the wider concept of a certainly disparate seriatus-group proposed by Fortier and S.R. Shaw (1999). Despite the findings of Zaldívar-Riverón et al. (2008), we do not use the subgeneric name Chelonorhogas for our apicalis-group partly because the characters by which we recognise the W. Palaearctic elements break down when applied to the World fauna, but also because there are some indications (Butcher et al. 2012) that it may be a basal grade group, paraphyletic with respect to the subgenus Aleiodes as recognised by Zaldívar-Riverón et al. (2008). Further analysis of Aleiodes in the context of the wider Rogadinae on a world basis is in hand (Quicke et al. in prep.).

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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 (1994, 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 and Yu 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

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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 and Yu 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

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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 2nd 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. The following collections and acronyms are used: AAC (A.A. Allen Collection, Dawlish), ALC (A. Lozan Col-

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lection, Institute of Entomology, České Budĕjovice), BMNH (Natural History Museum, London), BZL (Oberösterreichisches Landesmuseum, Biologiezentrum, Linz), CC (M. Čapek Collection, Moravian Museum, Brno), CMIM (C. Morley Collection, Ipswich Museum, Ipswich), CNC (Canadian National Collection of Insects, Ottawa), FC (J.V. Falcó Collection, Valencia), FMNH (Finnish Museum of Natural History, Helsinki), FRAH (Forest Research, Alice Holt Lodge, Farnham), KBIN (Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels), JLC (J. Lukáš Collection, Bratislava), MCZ (Museum of Comparative Zoology, Harvard University, Cambridge, U.S.A.), MHHN (Muséum National d’Histoire Naturelle, Paris), MMUM (Manchester Museum, University of Manchester, Manchester), MSC (M. Schwarz Collection, Linz-Ansfelden), MTMA (Hungarian Natural History Museum, Budapest), NMI (National Museum of Ireland, Dublin), NMS (National Museums of Scotland, Edinburgh), NNHM (National Natural History Museum, Oslo), NRS (Swedish Natural History Museum, Stockholm), OUM (Oxford University Museum of Natural History, Oxford), PAN (Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw & Łomna-Las), RMNH (Naturalis Biodiversity Center, Leiden), SDEI (Senkenberg Deutches Entomologisches Institut, Müncheberg), SMNS (Staatliches Museum für Naturkunde, Stuttgart), SYKE (Finnish Environment Institute, Friendship Park Research Centre, Kuhmo), UMZC (University Museum of Zoology, Cambridge), USNM (U.S. National Museum of Natural History, Washington D.C.), UWIM (University of Wyoming Insect Museum, Laramie), WAE (W.A. Ely Collection, Rotherham), ZILU (Zoological Institute, Lund University, Lund), ZISP (Zoological Institute, Academia NAUK, St. Petersburg), ZJUH (Zhejiang University, Hangzhou), ZMB (Zoologisches Museum, Humboldt Universität, Berlin), ZMC (Zoological Museum, Copenhagen), ZMUO (Zoological Museum, University of Oulu, Oulu), ZMUU (Zoological Museum of Uppsala University, Uppsala), and ZSSM (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

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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 (1990, 1993, 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 (1988, 1993; note, however, that in the present work the distance between eye and lateral ocellus is measured differently). For additional references see Yu et al. (2012).

Taxonomy Aleiodes Wesmael, 1838 Figs 1–378 Aleiodes Wesmael, 1838: 194; Shenefelt 1975: 1163–1185; Marsh 1979: 177–178; Papp 1985a: 143–164 & 1985b: 347–349; Shaw and Huddleston 1991: 95–96 (biology); van Achterberg 1991: 24. Type species (designated by Viereck 1914): Aleiodes heterogaster Wesmael, 1838 [examined; = A. albitibia (Herrich-Schäffer, 1838)]. Petalodes Wesmael, 1838: 123; Tobias 1971: 218 (transl. 1975: 86–87); Shenefelt 1975: 1209–1211; Tobias 1976: 90; Marsh 1979: 179; van Achterberg 1991: 24. Type species (by monotypy): Petalodes unicolor Wesmael, 1838 [examined; = Aleiodes compressor (Herrich-Schäffer, 1838)]. Schizoides Wesmael, 1838: 94. Unavailable name. Nebartha Walker, 1860: 310; Shenefelt 1975: 1216; Marsh 1979: 179; van Achterberg 1991: 24. Type species (by monotypy): Nebartha macropodides Walker, 1860 [examined]. Tetrasphaeropyx Ashmead, 1889: 634; Shenefelt 1975: 1260; Marsh 1979: 181; Fortier 2009: 19 (as subgenus; revision). Type species (by monotypy): Rogas pilosus Cresson, 1872 [examined]. Neorhogas Szépligeti, 1906: 605; Shenefelt 1975: 1205; van Achterberg 1991: 24. Type species (by monotypy): Neorhogas luteus Szépligeti, 1906 [examined; = Aleiodes praetor (Reinhard, 1863)]. Chelonorhogas Enderlein, 1912b: 258; Shenefelt 1975: 1187; van Achterberg 1991: 24. Type species (by monotypy): Chelonorhogas rufithorax Enderlein, 1912 [examined; not Aleiodes rufithorax (Cameron, 1911) = A. convexus van Achterberg, 1991]. Leluthinus Enderlein, 1912c: 96; Shenefelt 1975: 1202–1203; van Achterberg 1991: 24. Type species (by monotypy): Leluthinus lividus Enderlein, 1912 [examined]. Aleirhogas Baker, 1917b: 383, 411; Shenefelt 1974: 1185–1186; van Achterberg 1991: 24. Type species (designated by Viereck 1922): Rhogas (Aleirhogas) schultzei Baker, 1917 [examined]. Heterogamoides Fullaway, 1919: 43; Shenefelt 1975: 1188; van Achterberg 1991: 24. Type species (by monotypy): Heterogamoides muirii Fullaway, 1919 [examined].

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Cordylorhogas Enderlein, 1920: 153; Shenefelt 1975: 1195; van Achterberg 1991: 31. Type species (by monotypy): Cordylorhogas trifasciatus Enderlein, 1920 [examined]. Hyperstemma Shestakov, 1940: 10; Shenefelt 1975: 1200; van Achterberg 1991: 24. Type species (by monotypy): Hyperstemma chlorotica Shestakov, 1940 [examined]. R(h)ogas auctt.; Tobias 1971: 215–217 (transl. 1975: 83–86); Shenefelt 1975: 1215– 1256; Tobias 1976: 81–89; Marsh 1979: 179–181; Tobias 1986: 74–84. Diagnosis. Propodeum with a long median carina dorsally (Figs 11, 52); ovipositor sheath slightly expanded towards apex or parallel-sided and comparatively wide as far as visible (Figs 12, 31, 152, 301); second metasomal tergite with a median carina anteriorly (Figs 193, 291, 320, 331), but absent in part of the genus; hind trochantellus of ♀ normal, at most 2.6 × as long as wide (Figs 55, 104, 183, 194); vein r of fore wing 0.2–0.8 × vein 3-SR (Figs 1, 9, 22, 65, 205), if 0.6–0.8 × (Fig. 343) then precoxal area of mesopleuron granulate or coriaceous, without rugae and second metasomal tergite without triangular area medio-basally. Biology. Very large genus of koinobiont, synovigenic endoparasitoids; in the western Palaearctic of Drepanidae (including Thyratirinae), Erebidae (including Hypeninae, Lymantriinae, Arctiinae, Hypenodinae), Geometridae, Hesperiidae, Lasiocampidae, Lycaenidae, Noctuidae, Nolidae, Notodontidae, Nymphalidae (Satyrinae), Pterophoridae, Sphingidae, Ypsolophidae and Zygaenidae. This list includes only taxa of which we have been able to verify hosts, either by our own rearings or by examination of host remains; there are other host groups recorded in the literature, but we regard many of them as almost certainly erroneous and seek confirmation of others. The caterpillars are killed by the endoparasitoid and “mummified” — i.e. turned into a partly shrunken and hardened structure that is more or less tanned (Figs 8, 244, 259, 287, 342, 354), in most cases before their final instar, and the parasitoid pupates and eventually emerges as an adult from this mummy. Almost all Aleiodes species are strictly solitary (in Europe only two species are gregarious, but neither is treated in this part). 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

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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.

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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 6th, 1838, that the 11th volume of the Nouveau Memoires was in press.

Key to West Palaearctic species groups, and outlying distinctive species, of the genus Aleiodes Wesmael 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 0.8–3.0 × vein 3-SR (d); second submarginal cell of fore wing about as long as high or distinctly shorter (e)........................................ .............................................................genus Heterogamus Wesmael, 1838

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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

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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 (d); [ovipositor with small teeth ventrally and with wide dorsal flange (e)]; parasitoid of Sphingidae................. A. praetor (Reinhard, 1863)

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Ovipositor sheath distinctly setose (aa), but sometimes mainly ventrally so; marginal cell of hind wing subparallel-sided (upper bb), evenly widened (lower bb) apically or somewhat narrowed and distinctly widened apically (bbb); lateral carina of scutellum absent (cc) or if present (ccc) then lunula wide (dd) or narrow (ddd); parasitoids of other families...............................3

3

Apical half of marginal cell of hind wing distinctly gradually widened, its maximum width 1.6 × its width near hamuli or wider (a), if largely parallelsided (aaa) then tarsal claws with coarse blackish pecten (b); second metasomal tergite with wide smooth triangular area medio-basally (c); occipital carina usually reduced ventrally, not reaching hypostomal carina (d); mesopleuron partly smooth and shiny (at least between punctures), but largely densely sculptured in A. krulikowskii and some males of A. ruficornis; (usually macropterous, the known brachypterous specimens also belong here)............ .......................................................................................... A. apicalis-group

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Apical half of marginal cell of hind wing parallel-sided or slightly widened and its maximum width less than 1.8 × its width near hamuli (aa), if 1.7–2.7 × (aaa), then mesopleuron largely coriaceous or granulate and tarsal claws at most yellowish pectinate (bb); second tergite without triangular area mediobasally (cc) or this area is narrow or minute (ccc); occipital carina usually complete ventrally, reaching hypostomal carina (dd); mesopleuron usually extensively coriaceous or finely granulate, but medially coarsely sculptured in A. bicolor-group and a few members of A. circumscriptus-group, rarely largely shiny; (only macropterous specimens known)..............................................4

4

Mesopleuron largely (and often strongly) shiny (a), but may be partly superficially granulate in A. testaceus and A. ungularis, and in A. modestus mesopleuron shiny mainly anteriorly of speculum (aaa); maximum width of hypo­ clypeal depression 0.3–0.6 × minimum width of face (b).............................5

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Mesopleuron largely rather matt and distinctly sculptured (aa; at least coriaceous or granulate, though sometimes only weakly so); maximum width of hypoclypeal depression usually 0.3–0.4 × minimum width of face (bb).......8

5

Inner side of hind tibia with comb of whitish bristles apically (a); mesosoma (except propodeum and metapleuron) largely yellowish or yellowish orange (b); precoxal area impressed medially and finely crenulate (c); metasoma dark brown or blackish medially and largely pale yellow laterally (d)....................... .......................................................................A. ungularis (Thomson, 1892)

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Inner side of hind tibia without comb apically, normally only with bristly setae (aa); mesosoma largely black (bb) except in A. testaceus; precoxal area not impressed and smooth (cc) or rugose (ccc); metasomal tergites practically completely yellowish (dd) or brownish (in A. albitibia sometimes with a large yellow or ivory central patch on second tergite), if blackish or dark brown then also laterally so (ddd)...........................................................................6

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6

OOL 0.4–0.5 × diameter of posterior ocellus (a); pterostigma dark brown (b); inner hind tibial spur 0.40–0.50 × hind basitarsus (c); third antennal segment robust (d); vein 1r-m of hind wing about as long as vein 1-M (e); area in front of anterior ocellus without tubercle (f); inner side of basal half of hind tibia whitish and contrasting with darker apical half (g), rarely largely dark brown; third tergite about as long as second tergite and curved medioposteriorly in dorsal view (h)..............A. albitibia (Herrich-Schäffer, 1838)

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OOL about equal to diameter of posterior ocellus (aa); pterostigma largely yellow (bb), yellowish brown or brown; inner hind tibial spur 0.25–0.30 × hind basitarsus (cc); third antennal segment rather slender (dd); vein 1r-m of hind wing distinctly shorter than vein 1-M (ee); area in front of anterior ocellus with a minute smooth tubercle (ff); inner side of basal half of hind tibia yellowish, similar to apical half (gg); third tergite 0.8 × as long as second tergite and truncate medio-posteriorly in dorsal view (hh)...........................7

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Antennal segments of female 30–35 (of male 34–37); vein r of fore wing 0.7– 0.9 × vein 3-SR (a); vein 1-SR of fore wing rather long (b); ventral margin of clypeus thick (c); maximum width of hypoclypeal depression 0.30–0.35 × minimum width of face (d); head and mesosoma largely yellowish brown (e), but mesopleuron dorsally and propodeum usually more or less dark brown; length of malar space of female 0.3–0.4 × height of eye in lateral view (f); third metasomal tergite with more or less developed diverging striae laterally (g).....................................................................A. testaceus (Telenga, 1941)

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Antennal segments of both sexes (37–)40–45(–47); vein r of fore wing 0.5–0.6 × vein 3-SR (aa); vein 1-SR of fore wing shorter (bb); ventral margin of clypeus thin (cc); maximum width of hypoclypeal depression about 0.5 × minimum width of face (dd); head and mesosoma largely blackish (ee); length of malar space of female 0.5–0.6 × height of eye in lateral view (ff); third tergite without distinct striae (gg)......... A. modestus (Reinhard, 1863)

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Antennal segments of ♀ 50–71 and head usually entirely brownish yellow (a), of ♂ 52–68 (but males of A. pallidator practically unknown); scapus brownish yellow or reddish brown (b) and robust in lateral view (c); [fourth metasomal tergite largely superficially granulate; first tergite lamelliform protruding latero-anteriorly].............................................................................9

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Antennal segments of ♀ 28–51, of ♂ 31–48, if antenna with 45–55 segments then head largely blackish medio-dorsally (aa) and/or scapus largely black or dark brown (bb) and less robust in lateral view (cc)...................................11

9

Vein 2-CU1 of fore wing 0.7–1.2 × as long as vein 1-CU1 (a); vein 1-SR weakly angled with vein 1-M (b) and vein 1-M rather curved (c); vein r of fore wing long and subvertical (d); antennal segments of ♀ 56–62; length of fore wing 6–10 mm; [tarsal claws small (e)]..... A. esenbeckii (Hartig, 1834)

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Vein 2-CU1 of fore wing 1.6‑3.0 × longer than vein 1-CU1 (aa); vein 1-SR distinctly angled (bb) or linearly connected to vein 1-M (bbb) and vein 1-M nearly straight (cc); vein r of fore wing medium-sized and oblique (dd); antennal segments of ♀ 51–71, if with 54–55 segments then length of fore wing 5–7 mm.....................................................................................................10

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Antennal segments of ♀ 66–71; length of malar space of ♀ 0.4 × height of eye (a); vein 2-CU1 of fore wing 1.6–1.8 × vein 1-CU1 (b); occipital carina reduced ventrally (c).............................. A. varius (Herrich-Schäffer, 1838)

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Antennal segments of ♀ 50–55; length of malar space of ♀ 0.3 × height of eye (aa); vein 2-CU1 of fore wing 2.2–3.0 × vein 1-CU1 (bb); occipital carina complete ventrally, reaching hypostomal carina (cc)...................................... ................................................................... A. pallidator (Thunberg, 1822)

11

First metasomal tergite lamelliform protruding latero-anteriorly (a) and hind trochantellus of female slender, its ventral length 2.2–2.9 × its width (b); fore wing rather narrow (c); speculum of mesopleuron often rugose and with satin sheen, largely reticulate or granulate (d).....................................................12

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First tergite less lamelliform protruding latero-anteriorly (aa); if lamella present up to spiracle and rather protruding then hind trochantellus of female moderately robust, its ventral length less than 2.4 × its width (bb), if rarely up to 2.6 × then fore wing moderately wide (cc) and speculum of mesopleuron shiny and (partly) smooth or granulate (dd)...............................................15

12

Tarsal claws with distinct fine pecten (a); scapus and pedicellus of ♀ at least partly blackish, contrasting with yellowish middle of antenna (b); length of malar space of ♀ 0.25–0.30 × height of eye in lateral view (c; of ♂ 0.30 times); [fourth metasomal tergite of ♀ black latero-posteriorly (of ♂ brownish yellow); antenna of ♀ in dorsal view bicoloured, first–eighth and 48th– 49th segments more or less dark brown, remainder of antenna yellowish, of ♂ entire antenna yellowish]........................A. apiculatus (Fahringer, 1932)

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Tarsal claws only bristly setose, without distinct pecten (aa); scapus and pedicellus of ♀ similarly coloured as medial fifth of antenna or paler (bb); length of malar space of ♀ 0.30–0.50 × height of eye in lateral view (cc; of ♂ 0.25 times, but males of A. angustipterus unknown)...........................................13

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Antennal segments of ♀ 49–52; hind tibia infuscate subapically, contrasting with yellowish apex of tibia (a); second tergite comparatively long (b); fourth tergite with distinct sharp lateral crease (c) and basally rugulose or rugose (d); pterostigma bicoloured, with its basal third pale yellow (e); [vein m-cu of fore wing straight and angled to vein 2-CU1; antenna of ♀ sometimes with a narrow white or pale yellowish submedial band]........... A. jakowlewi (Kokujev, 1898)

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Antennal segments of ♀ 34–40; hind tibia subapically and apically similarly coloured and brownish yellow (aa); second tergite comparatively short (bb); fourth tergite partly without distinct sharp lateral crease (cc), partly retracted and tergite largely smooth (dd); pterostigma unicoloured yellowish brown or dark brown (ee).........................................................................................14

14

Length of malar space of ♀ 0.4 × height of eye in lateral view (a); vein m-cu of fore wing slightly curved towards vein 2-CU1, meeting it at about 140° (b); pterostigma dark brown (c); fore wing distinctly infuscate (d); inner side of hind tibia without apical comb (e); speculum sculptured (f); occipital carina complete or narrowly interrupted dorsally (g)............A. angustipterus sp. n.

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Length of malar space of ♀ 0.5 × height of eye in lateral view (aa); vein m-cu of fore wing straight and angled to vein 2-CU1 (bb); pterostigma yellowish (cc); fore wing subhyaline (dd); inner side of hind tibia with weakly developed apical comb (ee); speculum partly smooth and shiny (ff); occipital carina widely interrupted dorsally (gg).......................... A. carminatus sp. n.

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15

Fourth metasomal tergite curved posteriorly in dorsal view (a) and following tergites more or less retracted (b); vein r of fore wing 0.6–0.8 × vein 3-SR (c), precoxal sulcus largely granulate or coriaceous (d); trochanters, trochantelli and pterostigma largely black(ish) (e); [vein m-cu of hind wing distinct; formerly Tetrasphaeropyx Ashmead, 1889. Associated with Macariini (Geometridae)].....................................................................................................16

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Fourth tergite subtruncate medio-posteriorly in dorsal view (aa) and following segments at least partly exposed (bb); vein r of fore wing 0.2–0.6 × vein 3-SR (cc), if 0.6–0.8 × (ccc) then precoxal area coarsely rugose medially (dd) and third tergite enlarged and flattened (A. hergeri Papp); trochanters, and trochantelli usually brownish yellow (ee) and colour of pterostigma variable (ee, eee)......................................................................................................18

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Mesoscutum (a), orbita (b) and malar space (c) largely yellowish brown; all femora and tibiae black or dark brown (d); fore and hind femora slender (e); vein 1-SR of fore wing angled with vein 1-M (f)............ A. artesiariae sp. n.

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Mesoscutum (aa), orbita (bb) and malar space (cc) black; all femora and tibiae reddish or yellowish brown (dd); fore and hind femora less slender (ee); vein 1-SR of fore wing sublinear with vein 1-M (ff)...................................17

17

Vein M+CU1 of fore wing apically at about same level as vein 2-CU1 (a); vein r of fore wing 0.6–0.9 × vein 3-SR (b); length of fore wing 3.4–3.7 mm; arctic and alpine sp......................................... A. arcticus (Thomson, 1892)

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Vein M+CU1 of fore wing apically above level of vein 2-CU1 (aa); vein r of fore wing 0.9–1.1 × vein 3-SR (bb); length of fore wing 3.9–4.7 mm; lowland sp.....................................A. reticulatus (Noskiewicz, 1956), stat. rev.

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18

Head subglobose in dorsal view (a) and high in anterior view (b); dorsal face of propodeum long and (slightly) angularly protruding postero-laterally (c); fore femur stout (d); [antennal segments of ♀ 28–35, stout; strongly sexually dimorphic, male with large ocelli and slender antennal segments and antenna with 39–41 segments; body completely yellowish; length of antenna of ♀ 0.9–1.1 × fore wing, longer in ♂; second submarginal cell of fore wing rather narrow]...................................................................... A. curticornis nom. n.

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Head transverse in dorsal view (aa) and lower in anterior view (bb); dorsal face of propodeum shorter and rounded posteriorly (cc), if more elongate then fore femur slenderer (dd)...................................................................19

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Fourth metasomal tergite with a more or less sharp lateral crease for its whole length (a), if weak or absent then length of malar space of female 0.5–0.6 × height of eye in lateral view (b); precoxal area distinctly (and usually coarsely) rugose medially (c); epicnemial area (d) and propodeum (e) coarsely rugose; fourth metasomal tergite at least basally distinctly sculptured (f)......................A. bicolor-group

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Fourth tergite gently folded laterally, without acute lateral crease or this only anteriorly developed (aa), although rarely present as a simple, non-lamelliform crease to apex of tergite; length of malar space of female less than 0.5 × height of eye in lateral view (bb); precoxal area (cc) and epicnemial area less rugose (dd); propodeum (ee) usually with few rugae or completely coriaceous; fourth tergite usually mainly smooth with some superficial sculpture (ff)............................................................................................................20

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Inner apex of hind tibia with distinct comb (a); surroundings of veins 1-M and 1-SR of fore wing more or less infuscate and darker than surroundings (b); metasoma usually richly patterned (c); fourth tergite of ♀ pale (ivory-) yellowish latero-posteriorly (d), in ♂ usually infuscate; base of hind tibia usually narrowly dark brown (e); [antennal segments of ♀ (35–)44–50, of ♂ (42–)48–54; length of malar space 0.2–0.4 × height of eye in lateral view (f); temple narrow and directly narrowed behind eyes (g); length of hind femur of ♀ 5.1–6.5 × its width (of ♂ up to 8 times)]............................................... A. seriatus (Herrich-Schäffer, 1838) s.l.

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Inner apex of hind tibia without distinct comb (aa); surroundings of veins 1-M and 1-SR of fore wing subhyaline and similar to surroundings (bb); metasoma less patterned (cc); fourth tergite of ♀ dark brown or yellowish brown latero-posteriorly (dd); base of hind tibia usually yellowish brown (ee); [malar space (ff) and head shape variable]; A. circumscriptus-group..........21

21

Antennal segments of ♀ 42–49, of ♂ 42–46; temple directly narrowed behind eyes in dorsal view (a), mesosoma black(ish) dorsally (especially mesoscutum and scutellum (b), but sometimes notaulic area or scutellum brownish) and apical half of metasoma largely blackish (c); hind femur usually orangeish brown (d); [second tergite with (pale) yellowish elliptical patch medially (e). If body pale yellowish, antenna with 47–51 segments, head coarsely sculptured dorsally and fourth and following tergites largely under enlarged and flattened third tergite (f), cf. A. hergeri Papp, 1989]...................................22

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Antennal segments of ♀ 27–46, of ♂ 31–48; if antennal segments of ♀ 42– 46 then temple gradually narrowed behind eyes in dorsal view (aa) or mesosoma partly dorsally (bb) and apical half of metasoma yellowish brown (cc); hind femur variable, often yellowish or partly strongly darkened (dd)........23

22

Fore femur of ♀ less slender, 5.4–5.7 × as long as wide (a) and hardly sculptured, but of males slenderer; scapus and pedicellus (yellowish) brown ventrally (b); precoxal area frequently with some rugae or rugulae (c); propodeum distinctly transversally rugose medially and median carina largely absent on posterior half of propodeum or irregular (d); posterior half of pterostigma of ♀ largely dark brown (e); ivory part of malar space usually reaching clypeus, sometimes extending to lower part of inner orbit (f); mesosternum more or less blackish or dark brown (g), rarely completely reddish; [antennal segments of ♀ 42–47, of ♂ 42–46]...................... A. circumscriptus (Nees, 1834) s.s.

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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 (gg); [antennal segments of ♀ 44–49, of ♂ 43–47]............. ......................................................................A. nigricornis Wesmael, 1838

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23

Hind femur often partly, mesonotum dorsally and/or head largely dark brown or blackish (a); OOL and POL of ♀ 1.2–1.7 × diameter of posterior ocellus (b), but less in A. cantherius and ryrholmi (bbb); antennal segments of ♀ 36–45, of ♂ 34–45; metasoma with medial ivory patch (c), but less developed or absent in A. diarsianae and nigriceps (ccc)......................................24

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Hind femur completely, mesonotum dorsally and head largely yellowish (aa); if these parts are strongly darkened then OOL and POL of ♀ about equal to diameter of posterior ocellus (bb) and/or antennal segments of ♀ (31–)33– 37, rarely up to 40, of ♂ 34–39(–41) and metasoma without ivory patch, brownish yellow, brown or dark brown (cc)...............................................30

24

Head behind eyes directly narrowed in dorsal view (a); ocelli large (b); apex of metasoma of ♀ brownish yellow (c); temple short (d)............................25

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Head behind eyes gradually (roundly) narrowed in dorsal view (aa); ocelli medium-sized (bb); apex of metasoma of ♀ black or dark brown (cc); temple medium-sized (dd).....................................................................................26

25

Face yellowish brown (a); ocelli smaller, POL 0.8 × as wide as diameter of posterior ocellus (b); pale area of second tergite wide (c); mesoscutum with pair of yellowish brown stripes (d; but in males sometimes only vaguely indicated); medio-posterior depression of metanotum wide (e); palpi pale yellowish (f); vein cu-a of fore wing subvertical (g); face mainly transversely rugose (h).......................................................................A. cantherius (Lyle, 1919)

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Face black (aa); ocelli larger, POL 0.6 × as wide as diameter of posterior ocellus (bb); pale area of second tergite narrow (cc); mesoscutum entirely black (dd); medio-posterior depression of metanotum rather narrow (ee); palpi mainly dark brown (ff); vein cu-a of fore wing inclivous (gg); face superficially rugulose (hh).................................................................... A. ryrholmi sp. n.

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Length of fore femur 6.4–8.0 × its maximum width (a) and hind femur (centrally and subapically) parallel-sided (b); hind femur slender basally (c); mesosternum usually black(ish) (d); temple normal (e); scapus ventrally and usually basal half of antenna (dark) brown (f), rarely yellowish; hind femur basally largely yellowish and slightly infuscate subapically (g), paler than ventral side of scapus; if hind femur is distinctly infuscate then often also extreme base of hind tibia infuscate (h); [face usually black or dark brown medially and near eyes yellowish brown, but sometimes completely black or rarely completely yellowish; antennal segments of ♂ 35–40(–41), usually 36–38, less than of ♀, which has usually 37–39 segments]........ A. leptofemur sp. n.

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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 (gg); base of hind tibia yellowish (hh); [face usually completely black or yellowish; POL 1.2–1.7 × diameter of posterior ocellus].............................................................................................27

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Width of hypoclypeal depression of ♀ 0.35–0.40 × minimum width of face (a); antennal segments of ♀ (38–)39–43, of ♂ (38–)39–44; mesoscutum anteriorly and pronotum medio-anteriorly usually black or dark brown (b); medially mesopleuron usually without distinct rugulae or with a few (c); mesosternum yellowish, orange-brown or reddish (d), if darkened then not sharply defined; second metasomal tergite of ♀ with well differentiated median carina (e); lateral margins of third and fourth metasomal tergites of ♂ often completely yellowish, but sometimes darkened; [dark part of hind femur often extended to its apex].........................A. nigriceps Wesmael, 1838

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Width of hypoclypeal depression of ♀ 0.30–0.35 × minimum width of face (aa); antennal segments of ♀ (34–)36–40, of ♂ (36–)37–41; mesoscutum anteriorly and pronotum medio-anteriorly often yellowish (bb); medially mesopleuron of females with several rugulae or rugae (cc), but often lacking in males; colour of mesosternum variable, often strongly darkened or black and this usually sharply defined in N. European specimens (dd; more often orange-brown in S. European and Turkish specimens and in some males only indistinctly darkened); second metasomal tergite of ♀ with less differentiated median carina (ee); lateral margins of third and fourth tergites of ♂ almost always dark brown.....................................................................................28

28

Third and fourth antennal segments of ♀ slenderer (a); ocelli smaller (b); rugosity of face of female less developed (c); palpi slenderer (d); subapical antennal segments of ♀ moderately slender (e); first tergite slenderer (f); propodeum mainly coriaceous and with some rugulae or rugae (g); malar space usually partly or completely and temple near eye yellowish brown (h); fore and hind tarsi slenderer (hind tarsus: i); fore wing subhyaline or somewhat infuscate; mummy slender and light brownish............................................... .........................................................A. pictus (Herrich-Schäffer, 1838) s.s.

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Third and fourth antennal segments of ♀ stout (aa); ocelli somewhat larger (bb); larger part of face of female with distinct rugae (cc); palpi less slender (dd; especially third and fourth maxillary palp segments of ♂ widened); subapical antennal segments of ♀ submoniliform (ee); first tergite robust (ff); propodeum coarsely rugose (gg), but anteriorly less so; malar space and temple near eye usually dark reddish brown or blackish (hh); fore and hind tarsi less slender (hind tarsus: ii); fore wing usually slightly infuscate; mummy swollen and blackish; (unknown of A. bistrigatus)......................................29

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Length of eye in dorsal view about 1.6 × temple (a); 4th–7th antennal segments of both sexes slightly slenderer (b); temple behind eye slightly wider (c); number of antennal segments of ♂ usually less than of ♀, about 38 segments....... .......................................................... A. bistrigatus Roman, 1917, stat. rev.

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Length of eye in dorsal view 2.2–2.5 × temple (aa); 4th–7th antennal segments of both sexes stout (bb); temple behind eyes narrower (cc); number of antennal segments of ♂ usually more than of ♀, 40–45 segments.......................... A. diarsianae sp. n.

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Pterostigma of both sexes blackish or dark brown medially (a), border between dark and pale part well limited, contrasting with each other (b); temples linearly narrowed (c), head trapezoid in anterior view (d), and hind femur rather slender (e); OOL about equal to diameter of ocellus or less (f); antennal segments of ♀ 41–45, of ♂ 40–44; [vein 2-SR of fore wing yellowish; propodeum and first tergite usually yellowish in S. England, almost always moderately darkened in N. England and Scotland; vertex may be distinctly rug(ul) ose; mesopleuron shiny and only superficially granulate; rather long face and malar space; stemmaticum of male black and of female usually partly brownish yellow; a predominantly yellowish orange species]...........A. abraxanae sp. n.

–

Pterostigma of ♀ often largely or completely yellowish (aa), if distinctly infuscate (aaa) then border not well delimited, vague, not or less contrasting with its pale base (bb), antennal segments of ♀ 28–34, temples gradually (roundly) narrowed (cc), hind femur widened (ee) or OOL larger than diameter of ocellus (ff) and/or head nearly globular in anterior view (dd); antenna of ♀ often with less than 41 segments; [antennal segments of ♀ 28–47, of ♂ 30–47; mostly largely yellowish or orange species, but some very dark specimens should also run here]................ remainder of A. circumscriptus-group

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Biology and descriptions Aleiodes abraxanae sp. n. http://zoobank.org/1BE207D8-E7B5-493F-B103-7F5EE6896569 Figs 7–19 Aleiodes abraxanae van Achterberg in Lozan et al. 2010: 19. Nomen nudum. Rogas circumscriptus auct. p.p. (not Nees 1834). Aleiodes armatus auct. p.p. (not Wesmael 1838). Type material. Holotype, ♀ (NMS, Edinburgh), “[England], Otmoor N. R., Oxon., H[ost]: Abraxas grossulariata [on] Prunus spinosa, HLC [= host larva collected] 13.v.[19]79, PLE [parasitoid larva evident = mummification in the case of Aleiodes] 11.vi.[19]79, PIE [= parasitoid imago emerged] 4.vii.[19]79, M.R. Shaw”. Paratypes (74 ♀, 34 ♂): 52 ♀, 25 ♂ reared from larvae of the geometrid Abraxas grossulariata (Linnaeus) collected in v/vi, em (v)vi/vii as follows: 14 ♀, 4 ♂ (NMS, RMNH) England, Oxford, Otmoor, 1972, 1973, 1979, M.R. Shaw; 2 ♀, 3 ♂ (NMS, BMNH) England, West Sussex, Littlehampton, 1978, 1979, A.A. Allen; 1 ♂ (NMS) England, West Sussex, Hove, 1982, A.R. Cronin; 3 ♀, 1 ♂ (NMS, AAC) England, Surrey, Salfords, 1976, A.A. Allen; 1 ♀, 1 ♂ (NMS, RMNH) England, Berks, Maidenhead Thicket, 1979, M.R. Britton; 4 ♀, 4 ♂ (NMS, RMNH) England, Bucks, Butlers Hangings, 1979, M.R. Shaw; 2 ♀ (NMS) England, Bucks, Milton Keynes, 1984, J.P. Brock; 1 ♀ (BMNH) England, Cambridge, 1913, L. Doncaster; 2 ♀ (NMS) England, Westmorland, Beetham, 1991, M.R. Shaw; 11 ♀, 1 ♂ (NMS, BMNH) Scotland, Fife, St Andrews, 1935, 1936, 1938, D.J. Jackson; 1 ♀ (NMS) Scotland, Stirling, D.J. Jackson; 8 ♀, 7 ♂ (NMS, RMNH) Scotland, Orkney, Mainland, Waulkmill Bay, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Mainland, Caldale Bottom, 2009, K.P. Bland; 2 ♀ (NMS) Scotland, Orkney, Mainland, Redland, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Hoy, Nowt Bield, 2009, K.P. Bland; 1 ♂ (NMS) Scotland, Orkney, Hoy, Enegars, 2004, S. Gauld; 2 ♀ (SDEI) Germany, Sachsen-Anhalt, Wolfen, 1957, B. Stehlik; additionally 1 ♀ (NMS) from the Otmoor locality, host larva collected 8.x.1978, mummified 4.vi.1979, emerged 3.vii.1979, M.R. Shaw. Nonreared specimens: 1 ♂ (NMS) England, Cambridge, Chippenham Fen, 9.vii.1983, M.R. Shaw; 1 ♀ (NMS) England, Hunts, Monks Wood, 31.viii.2005, G.R. Broad; 1 ♂ (NMS) England, East Gloucester, Eastleach, 8.viii.2006, M.R. Shaw; 1 ♀ (BMNH) East Cornwall, Botusfleming, Marshall collection; 4 ♀ (BMNH) England, Oxford, Stanton St. John, 19.viii. 1968 (1) and 4.ix.1968 (3), J.P. Brock; 1 ♂ (BMNH) England, Herts, Whetstone, 24.vii.1961, P.H. Ward; 1 ♀, 1 ♂ (BMNH) England, Northampton, Spratton, x.1975 and vii.1976 respectively, I. & P. Gauld; 2 ♀ (BMNH) British Isles, Harwood coll.; 1 ♀ (BMNH) presumed British, A. Matthews in Lyle coll.; 3 ♀ (BMNH) presumed British, Stephens coll.; 1 ♀ (CMIM) England, Dorset, Weymouth 24.vi.1899, Peachell; 1 ♀ (CMIM) England, West Suffolk, Old Newton; 1 ♂ (CMIM) England, East Suffolk, Monk’s Soham, 18.vii.1933; 2 ♀, 1 ♂ (AAC) Eng-

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

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land, South Devon, Shaldon, 7.viii.1978 (1 ♀) and 6.viii.1979 (1 ♀, 1 ♂), A.A. Allen; 2 ♀ (NMS, RMNH) Wales, Anglesey, Llangristiolus, 27.viii–25.ix.1982, S.A. & D.C. Wilkinson; 1 ♀ (NMS) Scotland, West Ross, Sheildaig, viii. 1991, I. MacGowan; 2 ♂ (NMS) Scotland, South Uist, Loch Eynort, vi.1988, D. Whiteley; 2 ♀ (ALC, RMNH) Czech Republic, South Bohemia, Šumava Boubinský Pralés, virgin forest, 1000–1300 m, light trap, 22–24.vii.2003, I. Jaroš & K. Spitzer; 1 ♀ (ALC), Czech Republic, South Bohemia, Šumava Mts, 740 m, peat bogs near Mrtvý, light trap, 21–24.viii.2001, I. Jaroš & K. Spitzer; 1 ♂ (NMS) Finland, Satakunta, Pori, 1991, K. Ruohomäki; 1 ♀ (NMS) Sweden, Gotland, Sundre, Barrshage, vii/viii.2004, N. Ryrholm. Molecular data. MRS391 (Sweden JF962827, CO1), MRS636 (Scotland HQ551278/HQ551264, CO1), MRS637 (Scotland HQ551262, CO1 + KU682263, 28S), MRS694 (Scotland HQ551277, CO1). 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. Diagnosis. Pterostigma of both sexes blackish or dark brown medially, border between dark and pale part sharp, contrasting with each other (Figs 7, 9); temples directly strongly narrowed (Fig. 18) and comparatively wide in lateral view (Fig. 17); OOL about equal to diameter of ocellus; propodeum and first tergite yellowish or medially brown (Fig. 11); vein 2-SR of fore wing yellowish as vein 1-R1 (Fig. 9); head moderately transverse (Fig. 18); antennal segments of ♀ 41–45, of ♂ 40–44; subapical antennal segments slender (Fig. 15); vertex may be distinctly rug(ul)ose and meso-

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

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pleuron only coriaceous medially (Fig. 10); body entirely brownish yellow, at most propodeum and first tergite medially brown. Similar to A. hellenicus Papp, 1985, but A. hellenicus has pterostigma of both sexes completely pale yellowish and precoxal area usually with rugae medially. Description. Holotype, ♀, length of fore wing 5.1 mm, of body 6.2 mm. Head. Antennal segments of ♀ 43, length of antenna 1.3 × fore wing, its subapical segments about 2.3 × as long as wide; frons only coriaceous, matt; OOL 0.9 × diameter of posterior ocellus and coriaceous; vertex coriaceous, matt; clypeus convex, coriaceous; ventral margin of clypeus thick and depressed (Fig. 16); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 16) and face coriaceous; length of eye 3.8 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum coriaceous and occipital carina interrupted by somewhat less than width of ocellus (Fig. 18); clypeus partly above lower level of eyes (Fig. 16); length of malar space 0.3 × height of eye in lateral view; eyes distinctly protruding (Figs 16, 18). Mesosoma. Mesoscutal lobes largely coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli narrow and smooth, posteriorly lost in rugose area; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron and metapleuron coriaceous, matt; mesopleuron above precoxal area (except smooth and shiny speculum) coriaceous, but dorsally rugose; mesosternal sulcus narrow and shallow, impressed and without carina posteriorly; mesosternum angulate posteriorly; scutellum slightly convex, coriaceous, and carinate laterally; propodeum evenly convex and rugose but anteriorly weakly so, median carina complete, without tubercles. Wings. Fore wing: r 0.4 × 3-SR (Fig. 9); 1-CU1 horizontal, 0.35 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Figs 7, 9); cu-a vertical, not parallel with CU1b, straight (Fig. 9); 1-M slightly curved posteriorly. Hind wing: apical half of marginal cell parallel-sided; 2-SC+R short and longitudinal; m-cu present but unpigmented. Legs. Tarsal claws setose; hind coxa coriaceous, largely matt; hind trochantellus 2.6 × longer than wide; length of fore and hind femora 6.1 and 4.9 × their width, respectively (Figs 12–13); inner apex of hind tibia without comb; length of inner hind spur 0.25 × hind basitarsus. Metasoma. First tergite nearly as long as wide posteriorly, moderately convex and latero-posteriorly lamelliform; first and second tergites densely and finely longitudinally rugose, robust (Fig. 11), with distinct median carina; medio-basal area of second tergite obsolescent; second suture shallow and crenulate; basal half of third tergite finely rugose, remainder of metasoma largely superficially coriaceous; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath densely setose. Colour. Brownish yellow; apical fifth of antenna and dorsally propodeum dark brown; ovipositor sheath black; palpi, tegulae, apical 0.4 of first tergite and more or less second tergite pale yellowish; veins (but distally from 2-SR yellowish) and pterostigma (except yellow base and apex) dark brown; border between dark and pale part of pterostigma sharp, contrasting with each other (Figs 7, 9); wing membrane subhyaline.

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Variation. Length of fore wing 4.4–5.3 mm; antennal segments of ♀ 41(10), 42(18), 43(30), 44(6), and 45(1), of ♂ 40(3), 41(7), 42(8), 43(4), 44(4); stemmaticum of male black and of female brownish yellow; basal 0.2–0.5 of pterostigma pale yellow, rarely largely yellow and only medially darkened; first tergite yellowish or infuscate medially. Etymology. Named after the generic name of its host: Abraxas Leach. Distribution. *British Isles (England, Scotland, Wales), *Czech Republic, *Finland, *Germany, *Sweden. 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 1975: 1217; van 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 1975: 1176; Papp 1991: 97 (examined). Rhogas heterogaster; Fahringer 1932: 258–259. Rogas heterogaster; Hammond and Smith 1957: 181; Tobias 1986: 82 (transl.: 136). Type material. Redescribed ♀ and holotype of A. heterogaster (KBIN), “[Belgium], Campine, 1833”, “A. heterogaster mihi, det. C. Wesmael”, “Belgique, Charleroi/teste Papp, J., 1983”, “Holotypus Aleiodes heterogaster Wesm., 1838 / Papp, 1983”. The type series of R. albitibia is lost. Additional material. *Austria, British Isles (England: V.C.s 3, 11, 15, 22, 58, 59, 61, 69; Wales: V.C. 49; Scotland: V.C.s 72, 77, 80, 88, 92, 96, 97, 98, 99; Ireland: (V.C.s H1, H19, H20), *Czech Republic, Finland, France, Germany, Hungary, Netherlands (DR: Wijster, LI: Asselt; Castelre, GE: Heerde; Putten; Tongeren, NB: Tilburg (Kaaistoep), NH: Muiderberg, OV: Buurse (Schipbeek)), Poland, *Spain, Sweden, Switzerland. Specimens in NMS, BMNH, BZL, CNC, OUM, RMNH, SDEI, USNM, ZSSM, I. Kakko collection, and WUIM. Molecular data. MRS210 (Scotland EU979574, CO1), MRS383 (Sweden JF962835/ KU682238, CO1), MRS753 (Sweden KU682248, CO1). Biology. Univoltine parasitoid of arboreal notodontids, overwintering in a highly distinctive mummy (Fig. 21). Specimens (in NMS unless indicated) reared from wild collected arboreal Notodontidae identified as Notodonta dromedarius (Linnaeus) (12 [5 are OUM, 2 are USNM]; T.H. Ford, M.J. Morgan, M.R. Shaw, A.W. Stelfox, W.A. Watson), Eligmodonta ziz-zac (Linneaus) (2:1; M.R. Shaw), Pheosia tremula (Clerck) (8 [1 is AAC]; A.A. Allen, B.T. Parsons, M.R. Shaw), Pheosia gnoma (Fabricius) (2 [BMNH]; G. Graham-Smith). Host range experiments had the following outcomes:

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

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E. ziz-zac 2:32\10\\8+2; Pterostoma palpina (Clerck) 2:10\0\\-; Ptilodon capucina (Linnaeus) 1:5\0\\-; Clostera pigra (Hufnagel) 2:9\0\\-; Phalera bucephala (Linnaeus) 2:2\0\\-. The developmental biology of this species is rather unusual in several respects. On approaching the host (E. ziczac in all the following observations, which are based on two female A. albitibia) the female seems to depend on its fore and middle tarsi more than its antennae for host assessment, although antennation does occur (possibly as much to desensitise the host as to investigate it). The host is not paralysed during attack: the female more or less pounces on the host once accepted and aligns herself along the host’s body, which she grasps with her front and middle legs during oviposition, and fans her wings for short bursts repeatedly during the oviposition process (this may have indicated that venom was being injected, but if so it did not appear to have a significantly paralysing action), which in some cases lasted as long as 10–15 minutes – in these cases usually with more than one insertion. In fact, probably because the host was not temporarily parasitised and so was not sluggish, self-superparasitism happened very easily. The metasoma is only weakly curled downwards during oviposition, after which the female left the host without the usual post-oviposition period of association. When young, the females host-fed non-destructively on occasion. Development of the parasitoid larva was extremely rapid at ca 22–25 °C, with mummification ensuing after as few as 7–10 days from oviposition. The unusual and highly distinctive mummy (Fig. 21) is very shiny and dark mahogany brown in colour. It consists of the caudal portion of the host (usually from abdominal segment 3 onwards), strongly delimited from the anterior portion which usually shrivels up and becomes detached, leaving a sharp rim. The parasitoid pupates in a capsule which is formed in the most caudal half of this structure (usually in abdominal segments 5–8), dorsally raised, thinly lined with silk, and sealed from a more ventral and anterior inflation of the host’s cuticle that is of approximately equal volume but fully open at its anterior end. During the mummification process this area fills with liquid and bubbles, but fluids do not spread to the substrate and the mummy does not become stuck down. Once dried and hardened, the whole structure looks as though it contains two pupating parasitoids, and indeed some authors have been misled by this (Hammond and Smith 1957). It is possible that the empty chamber may serve to decoy pseudo-hyperparasitoids, though this seems unlikely to be its main function. The mummy forms on the host’s food plant, usually on a leaf surface from which it is easily dislodged, and overwinters in the leaf litter. Aleiodes albitibia occurs particularly in wet, bushy places: bearing in mind that its hosts (evidently rather restricted within Notodontidae) feed on trees and shrubs (Salix, Populus, Alnus and Betula) that often overhang water, the form of the mummy may also be an adaptation to prevent submersion and perhaps also results in dispersion by water, as it floats easily and is not wetted. The winter is passed in the mummy, and A. albitibia is univoltine, with a flight period in Britain of roughly 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 (Fig. 24); pterostigma dark brown; vein 1r-m of hind wing about as long as vein 1-M (Fig. 23); inner hind tibial

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

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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. Head. Antennal segments of ♀ 46, long setose, length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide; frons weakly depressed, finely rugose medially, remainder superficially micro-granulate; OOL 0.4 × diameter of posterior ocellus, and finely granulate; vertex finely granulate, with some rugulae posteriorly, rather dull; clypeus normal, micro-granulate; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.5 × minimum width of face (Fig. 29); length of eye 2.8 × temple in dorsal view (Fig. 32); occiput behind stemmaticum granulate with some rugulae, narrow; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view; occipital carina widely interrupted medio-dorsally and ventrally (Fig. 30). Mesosoma. Mesoscutal lobes largely granulate and with punctulation, matt; prepectal carina complete, rather weak; precoxal area of mesopleuron largely smooth (but in other specimens usually micro-granulate with some punctulation); mesopleuron above precoxal area strongly shiny and smooth; metapleuron largely coriaceous; scutellum granulate and finely punctate, no distinct carina; propodeum evenly convex, short, anteriorly granulate, medially and posteriorly rugose, median carina complete, without tubercles. Wings. Fore wing: r 0.5 × 3-SR (Fig. 22); 1-CU1 horizontal, 0.35 × 2-CU1; r-m 0.5 × 3-SR; second submarginal cell rather long (Fig. 22); cu-a inclivous, curved posteriorly; 1-M straight posteriorly. Hind wing: marginal cell subparallel-sided, its apical width 1.0 × width at level of hamuli (Fig. 23); 2-SC+R shortly longitudinal (but in other specimens subquadrate); m-cu absent; M+CU:1-M = 5:3; 1r-m about as long as 1-M. Legs. Tarsal claws yellowish setose; hind coxa sparsely punctulate, and granulate; hind trochantellus robust; length of fore femur, hind femur and basitarsus 5.4, 3.7 and 6.6 × their width, respectively (Figs 26, 28); length of inner hind spur 0.5 × hind basitarsus, as long as outer spur. Metasoma. First tergite robust (Fig. 25); first and second tergites rather coarsely longitudinally (reticulate-)rugose, robust, with distinct median carina; medio-basal area of second tergite minute, triangular; second suture shallow; third tergite as long as second tergite and largely granulate, anteriorly with some rugulae; remainder of metasoma smooth, compressed; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath slender and rather shiny. Colour. Black; malar area, narrow stripe along eyes dorsally, fore and middle legs (but telotarsi infuscate), hind coxa, trochanter trochantellus and femur, palpi and teg-

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ulae yellowish; pterostigma and most veins dark brown; basal 0.6 of hind tibia ivory; remainder of hind tibia and tarsus blackish. Variation. Antennal segments of ♀: 43(8), 44(14), 45(14), 46(3), 47(3), 49(1); of ♂: 39(1), 40(7), 41(7), 42(4), 43(3), 44(2). Second metasomal tergite may be largely yellowish or ivory (except laterally), or only with pale basal patch; hind tibia may be largely dark brown; base of pterostigma and first tergite medio-apically completely black (typical A. heterogaster) or yellowish (typical A. albitibia); width of hypoclypeal depression 0.4–0.5 × minimum width of face; mesopleuron usually with faint brownish longitudinal streak ventrally. 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). Aleiodes angustipterus sp. n. http://zoobank.org/7186C4F3-1A0F-4128-870D-25B7E20DE304 Figs 35–47 Type material. Holotype, ♀ (RMNH, Leiden), “Nederland (Dr.), Wijster, opposite Biol. Station, 12–19.viii.1977, C. v. Achterberg”. Paratypes (15 ♀): 1 ♀ (NMS), England, Cumbria, Whitbarrow, Howe, MV light, 24.viii.1995, M.R. Shaw; 1 ♀ (NMS), England, Norfolk, Scarning, TF981120, 6.vii–1.ix.1988, A.P. Foster/NCC; 1 ♀ (NMS), England, Norfolk, Sutton, TQ373235, water trap, 21.viii–4.ix.1986, A.P. Foster/NCC; 2 ♀ (NMS), Wales, Anglesey, Fedw Fawr, SH6081, MV light, 11.viii.2003, M.R. Shaw; 1 ♀ (NMS), Wales, Gwent, Magor Marsh, ST425865, water trap 8–21.vii.1988, P. Holmes/NCC; 1 ♀ (Tullie House Museum, Carlisle), Scotland, Dumfriesshire, Gretna, Springfield, 17.viii.1939, J. Murray; 1 ♀ (BMNH), Jersey, Trinity, Howard Davis Farm, Rothamsted trap 18.iv–3.xii.2004, A. Vautier/P. Gould; 1 ♀ (FMNH), Finland, U. Vantaa. 6690:384. ex larva Hypenodes humidalis 27.v.1974, cocoon [in which the mummy formed?] 19.vi.1974, em. 3.vii.1974, E.O. Peltonen; 1 ♀ (BZL), Greece, Thráki NW, Mt. Menikio, 12.viii.2010, J. Halada; 1 ♀ (NMS), Lithuania, Cerkelia peat bog, 3.ix.2006, A. Lozan; 1 ♀ (M. Riedel Collection), Russia, E. Siberia, 10 km E Irkutsk, 8.viii.2005, Berlov; 1 ♀ (NMS), Portugal, Azores, ca 2008 [per D.L.J. Quicke, no further data]; 1 ♀ (MRS), China, Yangtze River near Fengdu, 15.vii.2002, M.R. Shaw; 1 ♀ (RMNH), Japan, Kusakai, Kawai V., Iwate, 3–4.viii.1981, A. Takasu. Molecular data. MRS172 (China KU682231, CO1), MRS279 (Wales KU682232, CO1), MRS280 (Wales KU682233, CO1), MRS822 (Azores KU682246, CO1). 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

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Figure 35. Aleiodes angustipterus sp. n., ♀, holotype, habitus lateral.

species, overwintering in the partly fed host larva. Hypenodes humidalis occurs in both acidic and alkali marshy areas, and the larva feeds on plant debris certainly including dead or dying Molinia caerulea (G.M. Haggett, personal communication). Indeed, when known the collecting sites of A. angustipterus have mostly been wet grasslands, including fens and bogs, but at least one specimen was collected in woodland on a limestone hill (Whitbarrow) which may suggest a wider host range. Diagnosis. Head subglobular (Fig. 46) and body slender; antenna of ♀ without a pale submedial band; antennal segments of ♀ 36–40; eye rather small (Fig. 45); OOL 1.2 × posteior ocellus; speculum of mesopleuron rugose or reticulate and dull as is remainder of mesopleuron (Fig. 38); propodeum slightly elongate (Fig. 38); fore wing narrow (Fig. 36); pterostigma brown; hind coxa distinctly shorter than first tergite; hind femur 6–7 × as long as its maximum width; hind trochantellus slender (Fig. 40); dorsal carinae of first metasomal tergite lamelliform protruding basally; second tergite with small smooth triangular area medio-basally and tergite rather short (Fig. 39); third tergite weakly sculptured; fourth tergite partly or entirely without sharp lateral crease, fourth and following tergites partly retracted and largely smooth. Morphologically similar to A. jakowlewi from Finland, Sweden and N. Russia, but A. jakowlewi has the hind coxa about as long as first tergite and second tergite comparatively long (hind coxa distinctly shorter than first tergite in A. angustipterus (Fig. 35) and second tergite comparatively short (Figs 35, 39)); fourth tergite with distinct sharp lateral crease and basally rugulose (fourth tergite partly without distinct sharp lateral crease,

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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). Description. Holotype, ♀, length of fore wing 3.2 mm, of body 4.1 mm. Head. Antennal segments of ♀ 36, length of antenna 1.1 × fore wing, its subapical segments about 1.4 × as long as wide; frons granulate, rather shiny; OOL and POL 1.2 and 1.5 × width of posterior ocellus, respectively; vertex superficially granulatecoriaceous, rather shiny; clypeus convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 44); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 44) and face coriaceous with some rugulae; length of eye 2.4 × temple in dorsal view and temple directly narrowed behind eye; head subglobular (Fig. 46); occiput behind stemmaticum coriaceous with satin sheen; occipital carina complete and dorsally arched (Fig. 46); clypeus partly above lower level of eyes (Fig. 44); length of malar space 0.4 × height of eye in lateral view; eyes somewhat protruding (Figs 44, 46). Mesosoma. Mesoscutal lobes coriaceous-rugulose, matt, but medio-posteriorly longitudinally rugose and anteriorly low; notauli narrow and crenulate, but sculpture largely lost; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron, area below it and mesosternum largely reticulate-rugose; remainder of mesopleuron (including speculum) rugose or rugulose and matt (Fig. 38); metapleuron rugose, matt; mesosternal sulcus deep and sparsely crenulate; mesosternum rounded posteriorly; scutellum slightly convex, rugulose, and laterally with irregular carina; propodeum flattened, without tubercles and coarsely rugose, median carina incomplete, posterior 0.3 absent. Wings. Fore wing: r 0.4 × 3-SR (Fig. 36); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Fig. 36); cu-a vertical, not parallel with CU1b, straight; 1-M straight and 1-SR angled with 1-M. Hind wing: apical half of marginal cell slightly widened; 2-SC+R short; m-cu obsolescent. Legs. Tarsal claws with yellow bristles; hind coxa rugulose and with spaced oblique rugae, with satin sheen and 0.7 × as long as first tergite; hind trochantellus 2.4 × longer ventrally than wide; length of fore and hind femora 5.8 and 6.1 × their width, respectively (Figs 40–41); inner apex of hind tibia without distinct comb; length of inner hind spur 0.25 × hind basitarsus. Metasoma. First tergite 1.1 × as long as wide posteriorly, convex anteriorly and dorsal carinae lamelliform protruding basally; first and second tergites longitudinally rugose, robust (Fig. 39), with distinct median carina; medio-basal area of second tergite minute; second suture narrow and crenulate; basal half of third tergite largely superficially coriaceous, with some fine longitudinal elements; third tergite with complete sharp lateral crease, absent on following tergites; ovipositor sheath densely setose and apically acute.

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

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Colour. Dark brown; head (except stemmaticum), mesoscutum and scutellum medially, tegulum, legs (but femora largely infuscate), patch on posterior third of first tergite, large triangular patch on second tergite (Fig. 39) and anterior patch of third tergite brownish yellow; fourth–seventh tergites yellow; mouthparts, humeral plate and metasoma ventrally pale yellow; ovipositor sheath black; veins and pterostigma dark brown; wing membrane infuscate. Variation. Antennal segments of ♀: 36(2), 37(4), 38(2), 39(3), 40(2). The male is unknown. Pale patches of first and third tergites sometimes absent; hind femur 6.1–7.0 × as long as wide and hind trochantellus 2.4–3.0 × longer ventrally than wide. Central antennal segments vary from 1.2–1.5 × as long as wide, but in one specimen about 2.2 times – although its metasoma (at least posteriorly) is female, it seems possible that this individual is an intersex. Etymology. From “angustus” (Latin for “narrow”) and “pteron” (Greek for “wing”), because of the narrow wings. Distribution. *British Isles (England, Scotland, Wales, Jersey), *Finland, *Greece, *Lithuania, *Netherlands, *Portugal (Azores), *Russia (Siberia), *China (Chongqing), *Japan (Honshu). 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. Aleiodes apiculatus (Fahringer, 1932) Figs 48–62 Rogas apicalis Reinhard, 1863: 266 (not Brullé 1832) (examined). Rhogas apiculatus Fahringer, 1932: 284 (replacement name). Aleiodes apiculatus; Shenefelt 1975: 1165; Papp 1991: 101 (as synonym of A. pallidator); Belokobylskij et al. 2003: 398. Rogas (Aleiodes) negativus Tobias, 1961: 123; Belokobylskij 2000: 60 (as synonym of A. apiculatus (Fahringer, 1932); paratype in BMNH examined). Aleiodes negativus; Shenefelt 1975: 1165. Type material. Holotype of A. apiculatus, ♀ (ZMB), “Type”, “Coll. H. Rhd”, “Germania, [surroundings of] Bautzen”, “Holotypus Rogas apiculatus Reinh., 1863, ♀, Papp, 1983”, “Aleiodes pallidator Thunb., ♀, det. Papp, J., 1984/ var. apiculatus (Fahr.)”. Paratype of A. negativus, ♀ (BMNH) from Russia (Siberia: Tuvinskaya ASSR). Additional material. *England (V.C.s 9, 17, 20, 23, 24, 29, 31), *Poland. Specimens in NMS, BMNH, RMNH, AAC. Molecular data. MRS028 (England EF115455, CO1 + EF115440, 28S), MRS064 (England KU682218, CO1), MRS079 (England KU682222, CO1), MRS407 (England KU682239, CO1).

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

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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. 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

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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 (Fig. 53); hind tibia completely brownish yellow (Fig. 53); tarsal claws with distinct fine pecten (Fig. 54); dorsal face of propodeum medium-sized and rounded posteriorly (Figs 51, 52), first metasomal tergite lamelliform protruding latero-anteriorly (Fig. 62); second metasomal tergite rather stout and with minute smooth triangular area medio-basally (Fig. 52); third tergite finely sculptured; apical half of third and fourth tergite without sharp lateral crease and superficially granulate and with satin sheen; fourth metasomal tergite of ♀ black latero-posteriorly (Figs 52, 53; of ♂ brownish yellow). Description. Redescribed ♀ (NMS) from England, length of fore wing 5.9 mm, of body 6.8 mm. Head. Antennal segments of ♀ 47, length of antenna 1.2 × fore wing, its subapical segments 1.6 × as long as wide; frons granulate, with satin sheen and some rugae; OOL and POL 0.5 and 0.6 × width of posterior ocellus, respectively; stemmaticum strongly protruding; vertex rugulose-granulate, with satin sheen; clypeus convex and punctulate-coriaceous; ventral margin of clypeus thick and convex (Fig. 59); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 59) and face mainly granulate with transverse rugulae; length of eye 3.4 × temple in dorsal view and temple moderately narrowed behind eye; occiput behind stemmaticum rugulose-granulate; occipital carina reduced medio-dorsally and complete ventrally, without crenulae and dorsally curved (Fig. 61); clypeus above lower level of eyes (Fig. 59); length of malar space 0.30 × height of eye in lateral view; eyes rather protruding (Figs 59–61).

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

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Mesosoma. Length of mesosoma 1.7 × its height; mesoscutal lobes finely granulate, matt, but medio-posteriorly irregularly rugose and anteriorly high; notauli mediumsized and crenulate; prepectal carina medium-sized, remaining separate far from anterior border; precoxal area of mesopleuron and area above it distinctly rugose; remainder of mesopleuron (including speculum) granulate and with satin sheen (Fig. 51); metapleuron distinctly granulate and with satin sheen; mesosternal sulcus shallow and sparsely crenulate; mesosternum rather angulate latero-posteriorly but rounded medially; scutellum slightly convex, finely granulate, and antero-laterally with carina; propodeum weakly convex, without tubercles, anteriorly granulate, medially coarsely rugose and posteriorly with longitudinal carinae, median carina complete. Wings. Fore wing: r 0.3 × 3-SR (Fig. 50); 1-CU1 slightly oblique, 0.5 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.45 × 3-SR; second submarginal cell rather long (Fig. 50); cu-a slightly inclivous, not parallel with CU1b, straight (Fig. 50); 1-M straight posteriorly and 1-SR angled to 1-M. Hind wing: marginal cell parallel-sided, but slightly narrowed submedially; 2-SC+R medium-sized, slender; m-cu short and only slightly pigmented; M+CU:1-M = 4:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws rather small and with distinct fine pecten (Fig. 54); hind coxa finely granulate, with satin sheen and 0.8 × as long as first tergite; hind trochantellus 2.6 × longer ventrally than wide; length of fore and hind femora 6.0 and 4.6 × their width, respectively; inner apex of hind tibia without distinct comb; length of inner hind spur 0.35 × hind basitarsus. Metasoma. First tergite 1.1 × as long as wide posteriorly, stout, convex anteriorly and latero-anteriorly distinctly lamelliform; first and second tergites densely coarsely longitudinally rugose (Fig. 52), third tergite basally rugulose and with distinct median carina up to middle of third tergite; medio-basal area of second tergite minute; second suture rather wide and crenulate; remainder of third tergite granulate and following tergites shiny and superficially granulate; apical half of third and fourth tergites without sharp lateral crease; ovipositor sheath densely setose and apically truncate. Colour. Brownish yellow; scapus and pedicellus of ♀ at least partly blackish, contrasting with yellowish middle of antenna and antenna of ♀ in dorsal view bicoloured, first–fifth[–eighth] and 2–3 apical segments more or less dark brown, remainder of antenna yellowish; malar space, mandible, palpi, tegulae, pronotum anteriorly, basal half of pterostigma, trochanters and trochantelli, fore and middle coxae, and ventral half of metasoma ivory or pale yellow; face medially, frons and vertex medially, stemmaticum, occiput dorsally, mesoscutum laterally narrowly, scutellar sulcus, axilla, scutellum posteriorly, metanotum, third (except antero-lateral corner)–sixth tergites, fourth–sixth sternites and ovipositor sheath black or dark brown; telotarsi slightly infuscate; veins and apical half of pterostigma dark brown; wing membrane slightly infuscate. Variation. Antennal segments of ♀ 46(1), 47(10), 48(4), 49(1); ♂ 41(1), 42(2), 43(8), 44(3), 45(6), 46(2). Length of fore wing 5–6 mm, of body 5–7 mm. Males are brownish yellow, but stemmaticum black and antenna apically, occiput dorsally, mesoscutum laterally, scutellum posteriorly, metanotum, propodeum medially, first

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tergite except posteriorly and second tergite laterally somewhat infuscate; malar space, palpi, tegulae, pronotum, fore and middle coxae, trochanters and trochantelli, first tergite medio-apically and middle of second tergite pale yellowish. 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. Aleiodes arcticus (Thomson, 1892) Figs 63–75 Rogas arcticus Thomson, 1892: 1679; Tobias 1986: 83 (transl.: 138) (examined). Rhogas arcticus; Fahringer 1932: 285. Aleiodes arcticus; Shenefelt 1975: 1165–1166; Papp 1985a: 155 (lectotype designation), 1991: 96; Belokobylskij et al. 2003: 398. Type material. Lectotype, ♀ (ZIL), “Lpl” [= Lapland, North Sweden]). Additional material. 3 ♀, 2 ♂ (G. Várkonyi personal coll., NMS) Finland, Ks. Salla 752.61, Värriö H, ex Pygmaena fusca, collection dates (of host larva) between 15.vi and 28.vi.1995, G. Várkonyi; 1 ♂ (G. Várkonyi personal coll.) Finland: Ks. Salla 752.61 Värriö H 21.vi.1995, G. Várkonyi; 1 ♀ (BMNH) Finland, Kuusano, Mäntytunturi, on snow, 29.vi.1935, G.J. Kerrich; 1 ♂ (NMS) S. Norway, Jotunheimen, Giendersheim, 1000–1500 m, 7.vii.[1966], J.E. & R.B. Benson; 2 ♂ (BMNH) S. Norway, Buskerud Fylke Geilo, 1000–1100 m, 16–24.vi.1965, J.E. & R.B. Benson; 1 ♂ (RMNH) Norway, Oppdal, Kongsvoll, Vestbekken, 28.vi.1978, J.O. Solem; 1 ♀

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

(BMNH) Sweden, Torne Lappmark, Tornehamn, 4.vii.1954, J.E. & R. B. Benson; 1 ♀, 1 ♂ (BMNH, NMS) Switzerland, Grisons, Müstairtal, Funt da S. Charl, 2400 m, 9.vi 1960 and 3.vii.1960, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Grisons,

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Engadine National Park, 2380 m, 1.vii.1960, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Valais, Arolla, 8000 ft, 9.vii.1935, J.E. & R.B. Benson; 1 ♂ (BMNH) Switzerland, Valais, near Verbier, 8000–8500 ft, 27.vi.1959, J.E. & R.B. Benson. Biology. This boreoalpine species is univoltine, passing the winter as a mummy. The only known host is the ennomine (Macariini) geometrid Pygmaena fusca (Thunberg) (5:1; G. Várkonyi/Finland), which feeds on Empetrum and Vaccinium (G. Várkonyi personal communication) and probably occurs throughout the range of the parasitoid. The small mummy (Fig. 64) is short, broad and dorsally elevated. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 70); OOL 1.9 × diameter of posterior ocellus; mesoscutum, orbita and malar space black; precoxal sulcus largely granulate; trochanters, trochantelli and pterostigma largely black(ish); mesoscutum without a longitudinal carina on mesoscutum medio-posteriorly; apical half of marginal cell of hind wing parallel-sided or slightly widened; vein M+CU1 of fore wing apically at about same level as vein 2-CU1 (Fig. 65); vein r of fore wing 0.6–0.9 × vein 3-SR (Figs 63, 65); vein 1-SR of fore wing linear with vein 1-M (Fig. 65); all femora and tibiae reddish or yellowish brown; fore and hind femora moderately stout (Figs 68–69); fourth metasomal tergite curved posteriorly in dorsal view (Fig. 67), lateral crease distinct and following tergites more or less retracted; length of fore wing 3.4–3.9 mm. Description. Redescribed ♀ (BMNH) from Müstairtal (Switzerland), length of fore wing 3.7 mm, of body 4.6 mm. Head. Antennal segments 40, length of antenna as long as fore wing, its subapical segments about 1.7 × as long as wide; frons mainly superficially granulate and with some rugulae anteriorly, weakly shiny; OOL 1.9 × diameter of posterior ocellus and granulate as vertex, with satin sheen; clypeus moderately convex, narrow and coriaceous; ventral margin of clypeus thick and depressed (Fig. 70); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 70) and face mainly coriaceous with some rugulae dorsally and long setae; length of eye 1.1 × temple in dorsal view and temple subparallel-sided behind eye; occiput behind stemmaticum granulate and occipital carina absent ventrally and narrowly interrupted dorsally; clypeus below lower level of eyes (Fig. 70); length of malar space 0.5 × height of eye in lateral view; eyes somewhat protruding (Figs 70–72). Mesosoma. Mesoscutal lobes largely granulate-coriaceous, matt and medio-posteriorly rugose, middle lobe without a longitudinal carina; notauli narrow, shallow and very finely crenulate; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) granulate, but dorsally finely rugose; medially metapleuron granulate and with some rugae, rather shiny; mesosternal sulcus narrow and rather deep, with longitudinal carina posteriorly; mesosternum rounded posteriorly; scutellum moderately convex, mainly granulate and largely non-carinate laterally; propodeum rather directly lowered posteriorly and granulate-rugose, median carina complete, without tubercles. Wings. Fore wing: r 0.7 × 3-SR (Fig. 65); 1-CU1 subhorizontal, 0.1 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.8 × 3-SR; second submarginal cell rather small (Fig. 65);

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

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vein M+CU1 of fore wing apically at about same level as vein 2-CU1 (Fig. 65); vein 1-SR of fore wing linear with vein 1-M; cu-a subvertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 65); 2-SC+R short and longitudinal; m-cu present, pigmented. Legs. Tarsal claws setose; hind coxa granulate-coriaceous, with satin sheen; hind trochantellus twice longer ventrally than wide; length of fore and hind femora 5.5 and 4.0 × their width, respectively (Figs 68–69); inner apex of hind tibia without comb; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite 0.7 × as long as wide posteriorly, convex and latero-posteriorly non-lamelliform; first–second tergites finely and densely irregularly rugulose and with median carina (Fig. 67); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite with median carina (except posteriorly), third–fourth tergites very finely rugulose-coriaceous; fourth tergite convex medially and apically; fourth tergite with sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose. Colour. Black (including coxae); palpi basally, tegulae, pterostigma, veins, trochanters and trochantelli dark brown; remainder of palpi and legs yellowish brown; wing membrane slightly infuscate. Variation. Antennal segments of ♀ 39(1), 40(2), 41(0), 42(1), of ♂ 38(1), 39(2), 40(2); length of fore wing 3.4–3.7 mm; maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face; vein r of fore wing 0.6–0.8 × vein 3-SR; median carina of middle mesoscutal lobe absent or weakly indicated; legs (except basally) vary from largely yellowish brown to largely dark brown with base of hind femur and tibiae paler than remainder of legs; second submarginal cell of fore wing rather variable in shape, but some are as trapezoidal as in A. reticulatus, with which this species is closely related. 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. Aleiodes artesiariae sp. n. http://zoobank.org/536DFEB6-65AF-46FC-9F9E-DE22816FD463 Figs 76–88 Type material. Holotype, ♀ (NMS, Edinburgh), “France, Olonne, ex Macaria artesiaria, mummy coll[ected] on Salix repens, vi.1984, N. Hall”.

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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. Diagnosis. Maximum width of hypoclypeal depression 0.3 × minimum width of face (Fig. 84); OOL 2.4 × diameter of posterior ocellus; mesoscutum, orbita and malar space yellowish brown; precoxal sulcus largely granulate; trochanters, trochantelli and pterostigma largely dark brown or black(ish); mesoscutum without a longitudinal carina medio-posteriorly; apical half of marginal cell of hind wing parallel-sided or slightly widened; vein M+CU1 of fore wing apically above level of vein 2-CU1 (Fig. 78); vein r of fore wing about 0.7 × vein 3-SR (Figs 76, 78); vein 1-SR of fore wing angled with vein 1-M (Fig. 78); all femora and tibiae dark brown; fore and hind femora moderately stout (Figs 87–88); fourth metasomal tergite curved posteriorly in dorsal view (Fig. 81), lateral crease distinct and following tergites more or less retracted; length of fore wing about 3 mm. Very similar to A. reticulatus (Noskiewicz) but the latter differs by the black mesoscutum, the reddish or yellowish brown femora and tibiae, the less slender fore and hind femora (Figs 313–314 vs 87–88) and by having vein 1-SR of fore wing linear with vein 1-M (Fig. 305 vs 78). Description. Holotype, ♀, length of fore wing 2.9 mm, of body 3.6 mm. Head. Antennal segments 33+ (incomplete), length of antenna at least as long as fore wing, its subapical segments somewhat longer than wide; frons mainly superficially granulate, moderately shiny; OOL 2.4 × diameter of posterior ocellus and granulate as vertex, with satin sheen; clypeus moderately convex, narrow and coriaceous, 0.4 × width of face; ventral margin of clypeus thick and depressed (Fig. 84); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 84) and face mainly coriaceous with some rugulae dorsally and long setae; length of eye 1.8 × temple in dorsal view and temple sub-parallel-sided behind eye and narrowed posteriorly; occiput behind stemmaticum granulate and occipital carina absent ventrally and complete (but irregular) dorsally; clypeus partly up to lower level of eyes (Fig. 84); length of malar space 0.4 × height of eye in lateral view; eyes slightly protruding (Figs 84–86). Mesosoma. Mesoscutal lobes largely granulate-coriaceous, matt and medio-posteriorly rugose, middle lobe without a longitudinal carina; notauli narrow, shallow and very finely crenulate; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) granulate, but dorsally rugose; medially metapleuron granulate, rather shiny; mesosternal sulcus narrow and rather deep, without longitudinal carina posteriorly; mesosternum rather angulate posteriorly; scutellum moderately convex medially and depressed laterally, mainly granulate and largely non-carinate laterally; lunula moderately wide; propodeum rather directly lowered posteriorly and granulate-rugose, median carina complete, without tubercles.

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

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

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

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half of marginal cell slightly widened apically (Fig. 78); 2-SC+R medium-sized and longitudinal; m-cu present and slightly pigmented. Legs. Tarsal claws setose; hind coxa granulate-coriaceous, with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 5.4 and 5.2 × their width, respectively (Figs 87–88); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus. Metasoma. First tergite 0.7 × as long as wide posteriorly, convex and non-lamelliform latero-posteriorly and basally; first–second tergites finely and densely irregularly rugulose and with median carina (Fig. 81); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite superficially coriaceous and with median carina (except posteriorly), third–fourth tergites very finely rugulosecoriaceous; fourth tergite convex medially and apically, shiny and with sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose. Colour. Black (including coxae, middle and hind trochanters); palpi, tegulae, pterostigma, veins, first and second tergites and remainder of legs dark brown; malar space, orbita, mesoscutum (but middle lobe somewhat infuscate medio-anteriorly), scutellum laterally, pronotum, mesopleuron (except postero-ventrally) yellowish brown; wing membrane slightly infuscate. Etymology. From the specific epithet of its host. Distribution. *France. Aleiodes bistrigatus Roman, 1917, stat. rev. Figs 89–98 Aleiodes circumscriptus var. bistrigatus Roman, 1917: 9; Shenefelt 1975: 1171; Papp 1991: 109 (as synonym of A. borealis) (examined). Type material. Lectotype here designated, ♀ (NRS), “Färöar [= Faroe Isl., Denmark], Klinck”, “Triangisvaag”, “♀ Aleiodes circumscriptus var. bistrigatus Roman, C. van Achterberg, 1984. Lectotype”, “178, 84”, “Riksmuseum Stockholm”, “NHRSHEVA 000003802”. Paralectotypes: 3 ♂ (NRS) with same locality labels as lectotype. Additional material. None. Biology. Unknown. Diagnosis. Apical half of hind femur (partly) dark brown, darker than hind trochanter and trochantellus; face with distinct rugae; antenna of female with 39 segments and third segment stout, 4th–7th segments moderately stout (but less than in A. diarsianae; Fig. 164), of male with 38 segments; OOL 1.7 × diameter of posterior ocellus (Fig. 97); clypeus 0.4 × as wide as face (Fig. 95); mesopleuron black dorso-posteriorly (Fig. 90); malar space and inner orbita dark brown and temple near eye (= external orbita) reddish-brown; first tergite strongly widened apically; eye (of male) elongate in lateral view; middle third of hind femur yellowish brown or dark brown; vertex

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

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moderately setose, rather shiny and blackish posteriorly; mesosternum dark brown. Close to A. borealis (Thomson, 1892), but this species has less antennal segments (♀: 32–34 segments), palpi and legs more or less infuscate and the clypeus wider (about 0.5 × width of the face). The shape of the subbasal antennal segments is similar to that of series from Scandinavia mentioned under A. diarsianae, but this series has the temples directly narrowed behind the eyes and the males have 42–47 antennal segments and females 44–45. Similar to the A. pictus-aggregate (e.g. A. pictus (Herrich-Schäffer, 1838) and A. nigriceps Wesmael, 1838), but these are usually smaller and less robust species, having the face without distinct rugae or only a few rugae dorsally, the mesosternum usually widely orange-brown (and the mesopleuron usually without rugae in A. nigriceps), the malar space partly or completely yellowish brown, the first tergite less widened apically, the fore and hind tarsi comparatively slender, the eye normal in lateral view and the antenna less robust. Description. Lectotype, ♀, length of fore wing 4.0 mm, of body 5.5 mm. Head. Antennal segments of ♀ 39, length of antenna 1.3 × fore wing, its subapical segments about 1.9 × as long as wide and third segment stout (Fig. 98); frons coriaceous and posteriorly rugulose, with satin sheen; OOL 1.7 × diameter of posterior ocellus and coriaceous; vertex rugulose-coriaceous, rather shiny; clypeus slightly convex, indistinctly sculptured; ventral margin of clypeus rounded and depressed (Fig. 95); width of hypoclypeal depression 0.38 × minimum width of face and face distinctly transversely rugose (Fig. 95); length of eye 1.6 × temple in dorsal view and temple gradually narrowed behind eye (Fig. 97); occiput behind stemmaticum coriaceous and with some rugulae, occipital carina interrupted dorsally; clypeus partly above lower level of eyes and 0.4 × as wide as face (Fig. 95); length of malar space 0.6 × length of eye in lateral view; eyes slightly protruding (Figs 95–97). Mesosoma. Mesosoma 1.7 × as long as high; mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli complete and moderately wide, weakly crenulate and posteriorly widened and rugose; prepectal carina medium-sized and lamelliform, reaching anterior border; precoxal area of mesopleuron very coarsely rugose, connected to rugosity of dorso-anterior part of mesopleuron; speculum nearly smooth and shiny (Fig. 90); metapleuron granulate, matt and posteriorly rather tuberculate; mesosternal sulcus narrow and deep; mesosternum rounded posteriorly; scutellum elongate, slightly convex, coriaceous and laterally largely carinate; propodeum rather flat dorsally, laterally and apically rather rugose, anteriorly only weakly so, median carina complete, but posteriorly irregular. Wings. Fore wing: r 0.3 × 3-SR (Fig. 89); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.65 × 2-SR, and 0.40 × 3-SR; second submarginal cell medium-sized (Fig. 89); cu-a vertical, not parallel with CU1b, straight; 1-M slightly curved posteriorly and not continuous with 1-SR. Hind wing: apical half of marginal cell parallel-sided or nearly so; 2-SC+R short and longitudinal; m-cu present; 1r-m distinctly oblique and 0.7 × 1-M. Legs. Tarsal claws setose; hind coxa coriaceous but partly superficially rugulose, largely matt; hind trochantellus twice longer ventrally than wide; length of fore and

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hind femora 5.4 and 5.0 × their width, respectively (Figs 93–94); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus. Metasoma. First tergite 0.9 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope comparatively wide (Fig. 91); first–third tergites densely and distinctly longitudinally rugose, robust (Fig. 91), with distinct median carina; medio-basal area of second tergite absent; second tergite 1.5 × as long as third tergite; second suture moderately impressed and crenulate; remainder of metasoma largely superficially coriaceous; fourth and apical fifth of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) densely setose. Colour. Black or brownish black; antenna yellowish brown, but scapus dorsally and apical seventh of antenna dark brown; palpi, temple near eyes, legs (except infuscate subapical part of hind femur), tegulae, longitudinal stripe on mesopleuron, mesoscutum posteriorly, metasoma baso-ventrally, first tergite medio-apically, middle of second tergite and third tergite medio-basally largely yellowish; veins and pterostigma (except yellowish basal third and centrally) dark brown; border between dark and pale part of pterostigma diffuse (Fig. 89); wing membrane subhyaline. Variation. The male paralectotypes are very similar to the lectotype; one has a complete antenna with 38 segments and most of the hind femur darkened. One paralectotype has the hind coxa completely yellowish and the mesopleuron less coarsely rugose, but other paralectotypes have the mesopleuron coarsely sculptured and the hind coxa largely infuscate. Note. Possibly a Faroe Islands endemic. Aleiodes cantherius (Lyle, 1919) Figs 99–112 Rogas cantherius Lyle, 1919: 153–154 (examined). Aleiodes cantherius; Shenefelt 1975: 1169; Papp 1991: 112 (as possible synonym of A. nigricornis). Type material. Lectotype here designated, ♀ (BMNH), “2504” [on card], “cotype”, “[England,] New Forest, 4.v.1914, ex Semiothisa liturata, G.T. Lyle”, “G.T. Lyle Coll., B.M. 1930-579”, “Rhogas cantherius Lyle”. Paralectotypes: 4 ♀ + 3 ♂ (BMNH, CMIM), topotypic and from same host, but one non-reared paralectotype from Harwood collection. Additional material. *Austria, British Isles (England: V.C.s 11, 17, 19, 22, 24, 25, 28, 56), *Germany, *Netherlands (Breda; Melissant; Wageningen), *Russia, Slovakia, *Sweden. . Specimens in NMS, BMNH, RMNH, ZISP, FRAH, CC, CMIM, SDEI. Molecular data. MRS777 (Sweden KU682249, CO1)), MRS787 (Sweden KU682253, CO1).

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Biology. A parasitoid of conifer-feeding Macaria species (Geometridae), overwintering as a mummy. Specimens (in NMS unless specified) reared from Macaria liturata (Clerck) (22 [6 are BMNH, 3 CMIM, 2 SDEI, 1 FRAH]; P.E. Hatcher, G.T. Lyle, M.R. Shaw/England); Macaria signaria (Hübner) (2:2 [CC]; M. Čapek/Slovakia). Additionally 5 reared specimens, fortunately accompanied by the host mummy, had been labelled as reared from other conifer-feeding geometrids (Bupalus piniaria (Linnaeus) (2, ZISP), Eupithecia indigata (Hübner) (2, FRAH), Hylaea fasciaria (Linnaeus) (1, ZISP)) but in all cases examination of the host remains established that the host was in fact more consistent with a species of Macaria. Plurivoltine, overwintering in a mummy constructed on a conifer needle. The adult occurs in the field from May until well into September, and it is clear from the rearing data that its conifer-feeding Macaria hosts are likely to be sought across all of their foodplants (Abies cephalonica, Larix decidua, Pinus strobus, Pinus sylvestris and Pseudotsuga menzieseii are indicated on the data labels overall). No experimentation has been undertaken. Diagnosis. Antennal segments of ♀ 39–43, of ♂ 40–43; head strongly narrowed behind eyes (Fig. 110), yellowish anteriorly and mainly dark brown dorsally; OOL 0.6 × diameter of posterior ocellus; length of malar space of ♀ 0.2–0.3 × height of eye in lateral view (Fig. 109); scapus in lateral view rather oblique apically; occipital carina interrupted dorsally and complete ventrally (Fig. 109); eye 3.5–6.5 × as long as temple in dorsal view (Fig. 110); mesosternum and precoxal sulcus superficially granulate and with satin sheen; vein 2-CU1 of fore wing about 3 × vein 1-CU1 (Fig. 101); vein 1-SR narrow and linearly connected to vein 1-M and vein 1-M straight (Fig. 101); hind femur stout (Fig. 104); fourth metasomal tergite largely (superficially) coriaceous and shiny; length of fore wing 4–5 mm. Description. Redescribed ♀ (NMS) from Santon Downham (England), length of fore wing 4.6 mm, of body 5.0 mm. Head. Antennal segments 43, length of antenna 1.3 × fore wing, its subapical segments about 2.3 × as long as wide (Fig. 112) and scapus in lateral view rather oblique apically; frons granulate and rather shiny; OOL and POL 0.6 and 0.8 × diameter of posterior ocellus, respectively and granulate; vertex granulate, dull; clypeus rather high, convex, coriaceous; ventral margin of clypeus thick (Fig. 108); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 108) and face mainly transversely rugose and granulate; length of eye 6.5 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum mainly granulate and occipital carina interrupted medio-dorsally and complete ventrally (Fig. 109); clypeus above lower level of eyes (Fig. 108); length of malar space 0.3 × height of eye in lateral view; eyes protruding (Fig. 109). Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen, but medioposteriorly with some rugae; notauli narrow, shallow and largely smooth; prepectal carina rather lamelliform medio-ventrally, nearly reaching anterior border of mesopleuron and latero-ventrally curved; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) superficially granulate, but dorsally rugulose; medially metapleuron superficially granulate, rather shiny; mesosternal sulcus narrow and rather deep, micro-crenulate, without carina

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

posteriorly; mesosternum rather angulate posteriorly; scutellum finely coriaceous and non-carinate laterally; dorsal face of propodeum medium-sized, convex and rugulose, but posteriorly with some carinae and smooth in between and anteriorly mainly granulate, median carina complete, without tubercles. Wings. Fore wing: r 0.5 × 3-SR (Fig. 101); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell elongate (Fig. 101);

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

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1-SR slightly angled to 1-M and slender; cu-a rather inclivous, not parallel with CU1b, straight; 1-M slightly curved. Hind wing: apical half of marginal cell parallel-sided or nearly so (Fig. 101); 2-SC+R longitudinal; m-cu present as fold, unpigmented; M+CU:1-M = 3:2; 1r-m 0.7 × 1-M. Legs. Tarsal claws yellowish setose; hind coxa superficially finely coriaceous, rather shiny; hind trochantellus 2.4 × longer ventrally than wide; length of fore and hind femora 6.2 and 4.0 × their width, respectively (Figs 104, 111); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.2 × as long as wide posteriorly, flattened and latero-anteriorly narrowly lamelliform; first–second tergites and base of third tergite densely finely longitudinally rugose and with median carina; second tergite stout, 0.8 × longer than wide basally and 1.2 × as long as third tergite (Fig. 103); medio-basal area of second tergite minute; second suture deep and distinctly crenulate; remainder of metasoma largely superficially coriaceous and rather shiny; apical half of third and fourth tergite without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate. Colour. Black or dark brown; palpi, pronotum postero-dorsally and tegulae pale yellowish; scapus and pedicellus ventrally (but dorsally more or less darkened), orbita, two stripes on mesoscutum, legs (but hind coxa more or less dark brown), first tergite medio-apically, second tergite (except postero-lateral corners), third and following tergites mainly yellowish brown (Figs 99, 103); pterostigma and veins dark brown, but base of pterostigma and vein 1-R1 of fore wing brownish yellow; wing membrane slightly infuscate. Variation. Length of fore wing 3.6–4.5 mm, of body 3.8–4.6 mm; antennal segments of ♀ 39(1), 40(2; one is lectotype), 41(6), 42(2), 43(4), of ♂ 39(1), 40(6), 41(8), 42(3), 43(3); specimens have a characteristic pair of more or less obscure dorsal orange brown marks on the otherwise dark mesoscutum. Males examined have the metasoma dark brown apically, hind tibia (except ivory base) and tarsus more or less infuscated. Note. The two sexes have about the same number of antennal segments. Aleiodes carminatus sp. n. http://zoobank.org/5C58514A-72B5-46E9-998D-4B491A436068 Figs 113–124 Type material. Holotype, ♀ (NMS, Edinburgh), “[France:] Corsica: Corte, Val de Restonica (Hôtel Colonna), 500 m, [at] light, 29.vii–3.viii.[20]01, M.R. Shaw”, “MRS Aleiodes DNA 102”. Paratypes (11 ♀ + 23 ♂): 1 ♀ + 1 ♂ (NMS, RMNH), same data as holotype; 1 ♀ (BMNH), “[Spain:] Mallorca, Sa Roca, P.N. de s’Albufera, MV light, 2–27.ix.2013, M.R. Honey BMNH(E) 2013-158”; 1 ♀ (NMS), “Spain: Zaragoza Prov., Los Monegros, Retuerta de Pina, 30TYL 27.94, J. Blasco-Zumeta,

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Figure 113. Aleiodes carminatus sp. n., ♀, holotype, habitus lateral.

5104, 8.viii.[19]92, NMSZ1997.026, swept from Suaeda vermiculata”; 2 ♀ + 1 ♂ (NMS, RMNH), id., but 28.vi.1992 and swept from Suaeda vera, ♂ swept from Osyris alba; 3 ♀ (NMS, RMNH), id., but 12.ix.1991 (1) or 10.ix.1993 (2) and collected at

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light; 2 ♂ (NMS, RMNH), id., but 10.vii.1993, at light; 9 ♂ (NMS, RMNH), id., but 10.ix.1993; 3 ♂ (NMS, RMNH), id., but 20.ix.1993; 2 ♂ (NMS, RMNH),, id., but ?1991; 2 ♂ (NMS), id., but 20.vii.1993; 1 ♂ (NMS), id., but 20.viii.1993; 2 ♀ (FC), “Esp.: Valencia, El Saler (Casal d’Esplai), T.M., 20–27.vii.1992 & 17–24. viii.1992, J.V. Falcó y F. Luna”; 2 ♂ (FC), “Esp.: Valencia, Moncada-TM blanca, 6–13.vii.1992 & 13–20.vii.1992, M.J. Verdú”; 1 ♀, (NMS), “[Spain:] Canary Islands, Tenerife, Anco Viejo, La Sabinita, 20.iii.1999, R.R. Askew”. Molecular data. MRS055 (Corsica JF962818, CO1), MRS098 (Corsica KU682224, CO1), MRS102 (Corsica KU682225, CO1). 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. Description. Holotype, ♀, length of fore wing 3.4 mm, of body 3.9 mm. Head. Antennal segments of ♀ 35, length of antenna 1.2 × fore wing, its subapical segments about 1.7 × as long as wide; frons rugulose-granulate, with satin sheen; OOL and POL 1.0 and 0.8 × width of posterior ocellus, respectively; vertex granulate, rather dull and distinctly depressed near ocelli; clypeus convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 121); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 121) and face coriaceous with superficial rugulae; length of eye 2.4 × temple in dorsal view and temple roundly narrowed behind eye; occiput behind stemmaticum coriaceous with satin sheen; occipital carina widely interrupted medio-dorsally and ventrally weak and irregular (Figs 122–123); clypeus partly above

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

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lower level of eyes (Fig. 121); length of malar space 0.5 × height of eye in lateral view; eyes protruding (Fig. 122). Mesosoma. Mesoscutal lobes coriaceous-granulate, with satin sheen, but medioposteriorly longitudinally rugose and anteriorly steep; notauli obsolescent; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron (except posteriorly) and mesopleuron antero-dorsally distinctly rugose; remainder of mesopleuron (but speculum partly smooth and shiny) granulate and dull; metapleuron largely granulate, matt; mesosternal sulcus shallow and largely smooth; mesosternum rounded posteriorly; scutellum flat, granulate, and laterally with distinct carina, lunula narrow and parallel-sided; propodeum convex, without tubercles, rugulose anteriorly and remainder rugose, median carina complete. Wings. Fore wing: r 0.7 × 3-SR (Fig. 114); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.5 × 3-SR; second submarginal cell medium-sized (Fig. 114); cu-a slightly inclivous, nearly parallel with CU1b, straight; 1-M nearly straight and 1-SR distinctly angled with 1-M. Hind wing: apical half of marginal cell slightly widened; 2-SC+R short; m-cu absent. Legs. Tarsal claws with yellow setae; hind coxa rugulose and with spaced oblique rugae, with satin sheen and 0.8 × as long as first tergite; hind trochantellus 2.8 × longer ventrally than wide (Fig. 117); length of fore and hind femora 6.6 and 4.8 × their width, respectively (Figs 117–118); inner apex of hind tibia with distinct comb (Fig. 119); length of inner hind spur 0.2 × hind basitarsus. Metasoma. First tergite as long as wide posteriorly, convex anteriorly and dorsal carinae lamelliform protruding basally; first and second tergites longitudinally striate, robust (Fig. 116), with distinct median carina; medio-basal area of second tergite absent; second suture narrow and crenulate; third tergite largely longitudinally rugulose, but smooth posteriorly; third tergite with complete sharp lateral crease but this absent from following tergites; ovipositor sheath ventrally densely setose and remainder smooth, shiny and apically acute. Colour. Yellowish brown; antenna (except scapus and pedicellus ventrally), ovipositor sheath and most of ventral part of metasoma dark brown; stemmaticum black; tegulae, pronotum partly and legs brownish yellow; veins brown; pterostigma pale yellowish, but slightly darkened laterally; wing membrane subhyaline. Variation. Antennal segments of ♀: 34(5), 35(3), 36(2), 37(2), of ♂: 35(1), 36(1), 37(8), 38(3), 39(1), 40(2). In many specimens fore wing 2-SR is strikingly longer than r-m, but in others this is less distinctive. Hind femur sometimes brown, 4.5–4.9 × as long as wide and hind trochantellus 2.6–2.9 × longer ventrally than wide; occipital carina ventrally sinuate and reduced or complete; colour of body varies from nearly completely yellowish brown to largely brown. The female from Canary Islands is the darkest specimen examined with metasoma (except medial pale patch) and hind leg largely brown. Etymology. From “carmino” (Latin for “comb”), because of the comb on the hind tibia. Distribution. *France (Corsica), *Spain (mainland, Balearic and Canary Islands). Note. Males have on average about 2–3 more antennal segments than females.

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Aleiodes circumscriptus (Nees, 1834) Figs 125–137 Rogas circumscriptus Nees, 1834: 216 (syntypes lost). Aleiodes circumscriptus; Shenefelt 1975: 1170–1171 (p.p.); Papp 1991: 113 (p.p.); Belokobylskij et al. 2003: 398. Type material. Neotype here designated, ♀ (NMS, Edinburgh), “[Scotland], Rowardennan, Stirlings., Hypena proboscidalis [on] Urtica, 2.ix.[19]89, mum. 17.iv. [19]90, em. 10.vi.[19]90, M.R. Shaw (♀ 2 in 1990 Expts)”. Additional material. Widespread in western Europe: *Austria, Belgium, British Isles (England: V.C.s 1, 3, 4, 11, 12, 14, 17, 20, 22, 23, 25, 26, 27, 28, 30, 31, 32, 33, 58, 61, 62, 63, 64; Wales: V.C. 52; Scotland: V.C.s 72, 77, 84, 86, 87, 89, 99, 111; Ireland: Co. Cork), Bulgaria, Czech Republic, *Finland, Germany, Hungary, Italy, *Lichtenstein, Netherlands (FL: Lelystad (Oostvaardersplassen), FR: Ried, GE: Heerde; Tongeren; Brummen (Voorstonden), LI: Kerkrade; St. Pietersberg; Tegelen; Wrakelberg, NB: Bergen op Zoom, ZH: Asperen; Waarder; Lexmond), Norway, Spain, Slovakia, *Sweden. Specimens in NMS, BMNH, OUM, BZL, RMNH, MTMA, ZSSM, ZISP, World Museum Liverpool, CNC, USNM, UWIM, M. Riedel collection, H. Schnee collection, MSC, JLC WAE, and I. Kakko collection. Molecular data. MRS062 (England EU979579, CO1 + KU682264, 28S), MRS073 (England KU682256, CO1), MRS074 (England KU682220, CO1). Biology. Plurivoltine parasitoid of larvae of Hypena proboscidalis (Linnaeus, 1758) (Erebidae: Hypeninae), overwintering in the host larva. Mummy (Fig. 15) brown and moderately slender. Specimens (in NMS unless indicated) reared from Hypena proboscidalis (Linnaeus) (15 [1 BMNH, 1 OUM, 1 AAC, 3 H. Schnee collection]; A.A. Allen, G.M. Haggett, A. Hawkins, R.J. Heckford, S. Ratering, M.R. Shaw). It may be an absolute specialist on Hypena proboscidalis; related species that similarly feed on Urtica (H. obsitalis (Hübner) and H. obesalis (Treitschke)) overwinter as adults and would not (by themselves) be capable of supporting the parasitoid’s annual life cycle: indeed, it has been absent from several large collections of H. obesalis made in various localities in the Alps (M.R. Shaw). The oviposition sequence (observations from two females, and H. proboscidalis) is abnormal in that there is no separate pre-oviposition sting inducing temporary paralysis, nor is there a post-oviposition period of association. The host is scarcely antennated, but quickly recognised and pounced upon or snatched with the front two pairs of legs, and held aligned with the parasitoids’s body while the ovipositor is inserted for a prolonged period – sometimes there are several insertions, with self-superparasitism then often occurring. The egg is strongly attached to internal organs (gut and malpighian tubules both observed) at its narrow end. Because of subsequent disease in the stock, it is not possible to give quantitative results, but it appeared that enthusiastic attack on fourth instar hosts resulted in oviposition but no development, and that only earlier instar hosts were suitable. Mummification takes place on the host’s food plant, usually in a semi-concealed position.

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

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.

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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. Description. Neotype, ♀, length of fore wing 4.9 mm, of body 5.3 mm. Head. Antennal segments 46, length of antenna 1.3 × fore wing, its subapical segments about 1.6 × as long as wide; frons coriaceous and posteriorly rugulose, weakly shiny; OOL equal to diameter of posterior ocellus and coriaceous; vertex coriaceous, with satin sheen; clypeus moderately convex, coriaceous; ventral margin of clypeus thick and depressed (Fig. 134); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 134) and face mainly coriaceous with some rugae dorsally; length of eye 3.2 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum coriaceous and occipital carina nearly complete, interrupted dorsally by somewhat less than width of ocellus (Fig. 136); clypeus partly above lower level of eyes (Fig. 134); length of malar space 0.4 × height of eye in lateral view; eyes moderately protruding (Fig. 136). Mesosoma. Mesoscutal lobes largely coriaceous, matt, but medio-posteriorly with a few longitudinal rugae; notauli narrow, shallow and crenulate, but posterior half absent; prepectal carina lamelliform medio-ventrally, reaching anterior border; precoxal area of mesopleuron coriaceous and with some rugae medially; mesopleuron above precoxal area (except large smooth and shiny speculum) coriaceous, but dorsally rugose; medially metapleuron coriaceous, matt; mesosternal sulcus narrow and rather deep, with carina posteriorly; mesosternum rather angulate posteriorly; scutellum nearly flat, coriaceous and largely non-carinate laterally; propodeum rather flat and coriaceous but posteriorly with some rugae, median carina present but absent on posterior half, without tubercles. Wings. Fore wing: r 0.2 × 3-SR (Fig. 127); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 127); cu-a vertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell parallel-sided or nearly so (Fig. 127); 2-SC+R short and longitudinal; short stub of m-cu present, unpigmented. Legs. Tarsal claws setose; hind coxa superficially coriaceous, with satin sheen; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 5.7 and 4.3 × their width, respectively (Figs 130–131); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus.

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

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Metasoma. First tergite as long as wide posteriorly, flattened and latero-posteriorly lamelliform; first tergite coriaceous and finely irregularly longitudinally rugose; second tergite robust (Fig. 129), without distinct median carina, with satin sheen and superficially rugulose; medio-basal area of second tergite absent; second suture shallow and largely crenulate; basal half of third tergite indistinctly rugulose, remainder of metasoma largely superficially coriaceous and rather shiny; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath largely densely setose. Colour. Black or brownish black; antenna brown, but scapus dorsally and laterally dark brown; palpi, malar space up to eyes, mandible, tegulae, fore and middle coxae, trochanters and trochantelli, bases of fore and middle femora, medio-apical fifth of first tergite, medially second tergite and medio-basal patch of third tergite pale yellowish (Fig. 129); orbita (except latero-ventrally) brownish yellow (Figs 134–136) and remainder of head dark brown; mesopleuron ventrally yellowish brown with darker mesosternum; hind femur (except basally) fuzzy brown (Fig. 130), remainder of legs brownish yellow; veins and pterostigma (except yellow basal 0.4 and apex) dark brown; border between dark and pale part of pterostigma fairly sharp, contrasting with each other (Fig. 127); wing membrane subhyaline. Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀: 42(1), 43(1), 44(17), 45(46), 46(28), 47(4); of ♂: 42(13), 43(30), 44(38), 45(25), 46(1); notauli absent posteriorly or shallowly impressed; mesoscutum sometimes with weak diffuse reddish colouration posteriorly, along notaulic courses; orbita sometimes completely yellowish; mesosternum varying from (frequently) almost black, and then strongly contrasting with the reddish lower third of the mesopleuron, to reddish brown; median carina of propodeum sometimes traceable to posterior margin. Note. Males have on average about one fewer antennal segment than females. Aleiodes curticornis nom. n. & stat. rev. Figs 138–151 Aleiodes ochraceus Hellén, 1927: 24, 32 (not Rogas ochraceus Curtis, 1834); Shenefelt 1975: 1179; Papp 1985a: 154 (as possible synonym of A. gastritor (Thunberg, 1822); Koponen and Tobias 1989: 24 (lectotype deposition) (examined). Rhogas ochraceous; Fahringer 1932: 305. Type material. Lectotype of A. ochraceus Hellén here designated (FMNH), ♀, “[Finland,] Jomala”, “Hellén”, “829”, “Coll. Hellén: Aleiodes ochraceus Hellén”, “http:// id.luomus.fi./GL3421”; one ♀ paralectotype (topotypic, GL3420) and one ♂ paralectotype (Nystad, GL3419). Additional material. *Austria, Finland, France (*mainland and *Corsica), *Hungary, Italy (*mainland and *Sicily), *Romania, *Spain, *Slovakia, *Slovenia, *Turkey. Specimens in NMS, BMNH, RMNH, MTMA, FMNH, ZSSM, FC, JLC.

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Figure 138. Aleiodes curticornis nom. n., ♀, Italy, Tyrol, near Laudes, habitus lateral.

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

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Biology. Unknown. Most specimens have been collected at night in July and August in open situations. It may be univoltine and have hosts in low vegetation. Diagnosis. Head subglobose (Fig. 150); antennal segments of ♀ 28–35, and stout (Fig. 147), of ♂ 39–42; antenna of ♀ 0.8–1.1 × as long as fore wing, longer in ♂; OOL of ♀ 1.2 × diameter of posterior ocellus; occiput coriaceous-rugose or -rugulose (Fig. 150); second submarginal cell of fore wing rather narrow (Fig. 139); dorsal face of propodeum long and (slightly) angularly protruding postero-laterally (Fig. 142); fore femur stout (Fig. 145); hind femur rather wide (Fig. 143); sexes strongly dimorphic, the ♂ having larger ocelli, and slender and more numerous antennal segments; body completely yellowish brown, except black stemmaticum. Description. Redescribed ♀ (RMNH) from Agri (Turkey), length of fore wing 3.2 mm, of body 3.7 mm. Head. Antennal segments 28, length of antenna 0.8 × as long as fore wing, its subapical segments slightly longer than wide (Fig. 147); head subglobose in dorsal view (Fig. 150); frons granulate-rugulose and slightly shiny; OOL 1.2 × diameter of posterior ocellus and very finely rugulose as is vertex, with satin sheen; clypeus moderately convex and coriaceous; ventral margin of clypeus thick and depressed (Fig. 148); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 148) and face mainly transversely rugulose and with medium-sized setae; length of eye 2.2 × temple in dorsal view and temple subparallel-sided behind eye; occiput behind stemmaticum rugulose and occipital carina present ventrally and widely interrupted dorsally; clypeus near lower level of eyes (Fig. 148); length of malar space 0.5 × height of eye in lateral view (Fig. 149); eyes somewhat protruding (Figs 148–150). Mesosoma. Pronotum medio-anteriorly distinctly convex; mesoscutal lobes largely coriaceous, matt and medio-posteriorly rugulose, notauli narrow and sparsely finely crenulate and posteriorly absent; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron finely rugose; mesopleuron above precoxal area (except partly smooth and shiny speculum) superficially granulate, but dorsally rugose; medially metapleuron granulate and matt; mesosternal sulcus narrow and rather shallow, without carina posteriorly; mesosternum rounded posteriorly; scutellum slender, moderately convex, mainly granulate and non-carinate laterally; dorsal face of propodeum largely rugose, long and (slightly) angularly crest-like or tuberculate protruding postero-laterally, median carina present but irregular and similar to surrounding sculpture. Wings. Fore wing: r 0.5 × 3-SR (Fig. 139); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell rather slender (Figs 138–139); vein M+CU1 of fore wing apically at same level as vein 2-CU1 (Fig. 139); vein 1-SR of fore wing short and linear with vein 1-M (Fig. 139); cu-a subvertical, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 140); 2-SC+R narrow and longitudinal; m-cu present, but unpigmented. Legs. Tarsal claws setose; hind coxa coriaceous, with some oblique striae and satin sheen and about reaching apex of first tergite; hind trochantellus 2.4 × longer ventrally

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

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than wide; length of fore and hind femora 4.5 and 4.2 × their width, respectively (Figs 143, 145); inner apex of hind tibia without comb; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite 0.9 × as long as wide posteriorly, rather flattened medially and latero-posteriorly non-lamelliform; first–second tergites longitudinally rugose and with median carina (Fig. 142); medio-basal area of second tergite absent; second suture narrow, deep and finely crenulate; third tergite without median carina; third–fourth tergites finely coriaceous; fourth tergite flat medially and apically truncate; fourth tergite without sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose. Colour. Yellowish brown; palpi, tegulae, pterostigma, veins (but parastigma and part of basal veins dark brown) and legs yellow; stemmaticum and ovipositor sheath black; wing membrane subhyaline. Variation. Sexual dimorphism is unusually pronounced in this species, in respect of the large ocelli and the slenderer and much higher number of antennal segments of the male. Antennal segments of ♀ 28(2), 30(6), 31(2), 32(1), 33(1), 34(3), 35(1) and of ♂ 39(2), 40(1), 41(5), 42(1); antenna of ♀ 0.8–1.1 × as long as fore wing; stemmaticum black or brown; hind femur of ♀ moderately robust (Fig. 143) to rather swollen. Aleiodes diarsianae sp. n. http://zoobank.org/04A7F1AC-F831-4ED3-BABC-287C8C42BBF9 Figs 152–164 Type material. Holotype, ♀ (NMS), “[U.K.], Wales: Anglesey, Fedw Fawr, ex indet. Noctuid swept at night ex Calluna etc., 23.v.[19]97, mum. 29.v.[19]97, em. 16.vi.[19]97, died 1.ii.[19]98, M.R. Shaw, ♀ A. diarsianae in 1997 expts.”, “Host remains compatible with Diarsia sp., possibly brunnea or mendica det M.R. Shaw, 2013”. Paratypes (3 ♀ + 55 ♂): 42 ♂ (NMS, RMNH, BMNH) progeny of the holotype, 29 cultured in the noctuid Diarsia mendica (Fabricius), oviposition in range 26.vii–3.viii.1997, mummification 18–27.v.1998, emergence 25.vi–4.vii.1998 and 13 in Diarsia rubi (Vieweg), oviposition 23–30.viii.1997, mummification 4.iv–9.v.1998, emergence 7–25.vi.1998; 1 ♂ (NMS) England: Westmorland, Arnside Knott, ex Diarsia ?brunnea (Denis & Schiffermüller) on Calluna, coll. 5.v.1984, mum. 16.v.1984, em. 19.vi.1984, M.R. Shaw; 1 ♀ + 1♂ (NMS) Scotland: Orkney, ex Diarsia brunnea on Calluna, coll. v.1977, em. vi.1977, R.I. Lorimer; 1 ♀ (NMS) Scotland, East Perth, Drumderg, NO2055, ix.2012, A. Huff; 1 ♂ (NMS) Scotland, South Aberdeen, Glen Tanar 16.vii–4.viii.1986, I. MacGowan; 1 ♂ (NMS) Scotland, South Aberdeen, Braemar, Morrone Birkwood, 12.vii–6.viii.1984, B.D. Batty; 1 ♂ (NMS) Scotland, Easterness, Loch Garten, vi.1984, J.A. Owen; 3 ♂ (NMS) Scotland, Easterness, river Nethy shingle bank, NJ0214, 19.vi–5.vii.1999 (1 ♂) and 5–19.vii.1999 (2 ♂), M. Edwards); 2 ♂ (NMS) Scotland, Elgin, Bognacruie, NJ0415 19.vii–3.viii.1999 (1 ♂) and 3–23.viii.1999 (1 ♂), M. Edwards; 1 ♂ (NMS) Scotland, Elgin, Elchies, NJ2146,

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

27.vii–9.viii.1999, B. Hicks; 2 ♂ (NMS) Scotland, Shetland, HU335730, 15–19. vii.2004, C. Sullivan; 1 ♀ (ZSSM), Netherlands: Nijmegen, ex Diarsia rubi, Bauer; 1 ♀ (RMNH), France: Besse en Chande SSE, Puy de Dôme, 13.vi.1976, H. Teunissen.

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Molecular data. MRS030 (Wales JF962600, CO1), MRS135 (Scotland KU682257, CO1 + EU854345, 28S). 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, 4th–7th 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, 4th–7th 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

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

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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. Description. Holotype, ♀, length of fore wing 4.0 mm, of body 5.5 mm. Head. Antennal segments of ♀ 40, length of antenna 1.3 × fore wing, its subapical segments about 1.5 × as long as wide and third segment stout (Figs 159–160, 164); frons coriaceous and posteriorly rugulose, with satin sheen; OOL 1.4 × diameter of posterior ocellus and coriaceous; vertex coriaceous, matt; clypeus slightly convex, indistinctly sculptured; ventral margin of clypeus rounded and depressed; width of hypoclypeal depression 0.36 × minimum width of face and face distinctly transversely rugose (Fig. 161); length of eye 2.5 × temple in dorsal view and temple rather directly narrowed behind eye (Fig. 163); occiput behind stemmaticum coriaceous and with some rugulae, occipital carina interrupted dorsally by somewhat less than width of ocellus (Fig. 163); clypeus partly above lower level of eyes and 0.4 × as wide as face (Fig. 161); length of malar space 0.5 × length of eye in lateral view; eyes moderately protruding (Figs 161–163). Mesosoma. Mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose; notauli complete and moderately wide, weakly crenulate and posteriorly widened and rugose; prepectal carina medium-sized and lamelliform, reaching anterior border; precoxal area of mesopleuron largely widely rugose, mesopleuron above precoxal area (except nearly smooth and shiny speculum) largely rugose (Fig. 155); metapleuron coriaceous, matt and posteriorly rather tuberculate; mesosternal sulcus narrow and deep, absent and replaced by carina medio-posteriorly; mesosternum rounded posteriorly; scutellum elongate, slightly convex, coriaceous and laterally largely carinate; propodeum rather flat dorsally (depressed laterally and posteriorly, rather tuberculate latero-posteriorly) and strongly rugose but anteriorly weakly so, median carina complete, but posteriorly irregular. Wings. Fore wing: r 0.3 × 3-SR (Fig. 154); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.65 × 2-SR, and 0.55 × 3-SR; second submarginal cell comparatively large (Figs 152, 154); cu-a weakly oblique, not parallel with CU1b, straight; 1-M slightly curved posteriorly. Hind wing: apical half of marginal cell parallel-sided or nearly so; 2-SC+R short and longitudinal; m-cu present and weakly pigmented (Fig. 154). Legs. Tarsal claws setose; hind coxa coriaceous but partly superficially rugulose, largely matt; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 5.1 and 4.3 × their width, respectively (Figs 157–158); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus. Metasoma. First tergite 0.8 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope comparatively wide (Fig. 156); first–third tergites densely and distinctly longitudinally rugose, robust (Fig. 156), with distinct median carina; medio-basal area of second tergite absent; second suture moderately impressed and crenulate; remainder of metasoma largely superficially coriaceous; fourth and apical fifth of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) densely setose. Colour. Black or brownish black; antenna pale brown, but scapus dorsally and apical seventh of antenna dark brown; palpi, and tegulae pale yellowish (Fig. 152);

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orbita posteriorly and dorsally brownish yellow (Figs 162–163); mesosoma orange brown, but propleuron, mesoscutal lobes medially, metanotum laterally, anterior half of propodeum and metapleuron, mesopleuron dorso-posteriorly and mesosternum black; metasoma largely dark orange brown (Figs 152, 156); hind femur (except basally) fuzzy dark brown (Fig. 157) and remainder of legs yellowish brown; veins and pterostigma (except yellow basal 0.2 and apex) dark brown; border between dark and pale part of pterostigma sharp, contrasting with each other (Fig. 154); wing membrane subhyaline. This specimen had lived in a humid tube for 7 months, and its colour had deepened considerably over this time. Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀ 36(1), 38(3), 39(1), 40(1), of ♂ 40(2), 41(2), 42(4), 43(13), 44(16), 45(14), 46(3); mesosoma largely black to largely orange-brown; OOL of male slightly longer than diameter of posterior ocellus and apical half of antenna dark brown; fifth maxillary palp segment slender to moderately widened and rather long; first tergite (except medio-posteriorly) black (♂) or entirely dark reddish-brown (♀) and second tergite black or reddish laterally; in British females only posterior segments somewhat darkened; in British males first tergite more or less blackish in anterior half as well, but second and third tergites usually (almost) fully orange, sometimes with infuscation sublaterally on second tergite (especially anteriorly).; mesopleuron medially and propodeum rugose or superficially rugulose; few females seen, but in one very extensively orange specimen the legs are almost completely orange, with only slight infuscation in the apical half of hind femur. May be confused with A. borealis (Thomson, 1892), but this species has less antennal segments (♀: 32–34 segments), palpi and legs more or less infuscate and the clypeus wider (about 0.5 × width of the face). We have seen 3 ♀ + 11 ♂ (NMS) from Sweden (Bohuslän and Västerbotten) and Finland (Kuusamo and Saarijärvi) that come close to A. diarsianae, but differ in being less robust (T1 less expanded apically; antennal segments longer in relation to width, especially basally), less strongly sculptured (fewer rugae on face; mesopleuron with only weak rugae), and having somewhat larger eyes. They also have slightly more antennal segments, at least in the female sex (the two females with intact antennae have 44 and 45 segments, the males have 42(1), 43(1), 44(1), 45(6), 47(1)), and the females have T1 more or less extensively blackish in anterior half, unlike the British A. diarsianae females seen, in which it is uniformly orange. CO1 sequences have been obtained for two localities (Västerbotten and Kuusamo; respectively MRS304 GenBank KU682234, and MRS692 GenBank KU682247): they form a well-isolated clade with A. diarsianae but differ from it by 8 fixed base-pairs. One of the Finnish males was reared from a noctuid mummy collected on a twig in a bog (N.R. Fritzén) later kindly identified from its CO1 sequence as Coenophila subrosea (Stephens) by Dr Katja Kramp (SDEI). Another male (in NMS) from Norway (Turtagrö, Sogn og Fjordane) has 42 antennal segments and probably belongs to the same species; it was reared from an unidentified dark noctuid mummy on Betula nana (K.P. Bland), which, like the Finnish one, is somewhat swollen but not as extensively so as in the considerable number of British A. diarsianae mummies we have seen. Both of these specimens, in

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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. Aleiodes esenbeckii (Hartig, 1838) Figs 165–189 Rogas esenbeckii Hartig, 1838: 255; Tobias 1986: 81 (transl.: 135) (examined). Rhogas esenbeckii; Kokujev (in Serebryanikova), 1901: 100. Aleiodes esenbecki; Papp 1991: 93 (as synonym of A. procerus). Aleiodes esenbeckii; Belokobylskij et al. 2003: 398. Rhogas corsicus Szépligeti, 1906: 616 (examined). Aleiodes corsicus; Papp 1991: 93 (as synonym of A. procerus), 2004: 215 (holotype). Rogas gastropachae Kokujev (in Serebryanikova), 1901: 100–101. Aleiodes gastropachae; Papp 1991: 93 (as synonym of A. procerus). Phanomeris dendrolimi Matsumura, 1926: 41; Chen and He 1997: 50 (as synonym of A. esenbeckii). Aleiodes dendrolimi; Shenefelt 1975: 1172–1173. Phanomeris dendrolimusi Matsumura, 1926: 32 (invalid emendation). Phanomeris spectabilis Matsumura, 1926: 33; Chen and He 1997: 50 (as synonym of A. esenbeckii). Rhogas metanastriae Rohwer, 1934: 47; Chen and He 1997: 50 (as synonym of A. esenbeckii). Type material. Holotype of R. esenbeckii, ♂ (ZSSM), “715, [Germany, Charlottenburg]”, “Esenbeckii n.”, together with mummy of Dendrolimus pini (L.); holotype of R. corsicus, ♀ (MTMA), “[France, Corsica,] Ajaccio”, “praetor Reinh.? (Corsica)”. Additional material. f. esenbeckii: Austria, *Croatia, Czech Republic, France (*mainland and Corsica) *Netherlands (Muiderberg), Germany, Spain (*Mallorca); f. dendrolimi: China, *Finland, Russia, Switzerland. Specimens in NMS, ZSSM, RMNH, BZL, MTMA, BMNH, SDEI, ZISP Molecular data. MRS180 (Finland EU979581,CO1 + EU854329, short 28S), MRS500 (Mallorca JF962845/KU682240, CO1). 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-

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

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

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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. Diagnosis. Antennal segments of ♀ 56–62; head entirely brownish yellow apart from stemmaticum (Figs 173–175): f. esenbeckii) or blackish (Figs 185–187: part of f. dendrolimi); scapus in lateral view ratherly oblique apically; OOL 0.3 × diameter of posterior ocellus; occipital carina complete ventrally (Fig. 174); length of malar space 0.25 × height of eye in lateral view; vein 2-CU1 of fore wing 0.7–1.2 × vein 1-CU1 (Fig. 167); vein 1-SR angled to vein 1-M and vein 1-M distinctly curved (Fig. 167); vein r of fore wing gradually merging into vein 3-SR; tarsal claws small (Fig. 170); fourth metasomal tergite superficially coriaceous; length of fore wing 6–10 mm. Often confused with A. varius (as A. procerus), but differs by the relative lengths of veins 1and 2-CU1 and by the number of antennal segments. Description. Redescribed ♀ (NMS) from Mallorca (Spain), length of fore wing 7.6 mm, of body 9.5 mm. Head. Antennal segments 59, length of antenna 1.3 × fore wing, its subapical segments about 1.9 × as long as wide and scapus in lateral view rather oblique apically; frons superficially granulate, rather shiny; OOL 0.3 × diameter of posterior ocellus and granulate; vertex superficially coriaceous, with satin sheen; clypeus rather high, convex dorsally and flattened ventrally, coriaceous and with long setae; ventral margin of clypeus thick and gradually depressed (Figs 173, 185); width of hypoclypeal depression 0.4 × minimum width of face (Figs 173, 185) and face mainly rugose with interspaces coriaceous; length of eye 4.7 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum finely rugose and occipital carina curved and interrupted medio-dorsally and complete ventrally (Figs 174, 186); clypeus entirely above lower level of eyes (Figs 173, 185); length of malar space 0.25 × height of eye in lateral view; eyes strongly protruding (Figs 173–175, 185–187). Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen; notauli narrow, shallow and mainly coriaceous; prepectal carina rather lamelliform medio-ventrally, almost reaching anterior border of mesopleuron and latero-ventrally curved; precoxal area of mesopleuron coriaceous, without fine rugae medially; mesopleuron above precoxal area (including shiny and granulate speculum) coriaceous, but dorsally finely rugose (cf. Fig. 181); medially metapleuron superficially granulate and shiny; mesosternal sulcus narrow and deep, without carina posteriorly; mesosternum angulate posteriorly; scutellum superficially coriaceous and carinate antero-laterally; dorsal face of propodeum rather long and coriaceous, posterior face short, hardly differentiated, with some short carinae and smooth in between, median carina complete and with weak tubercles postero-laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 167); 1-CU1 horizontal, 1.1 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.45 × 3-SR; second submarginal cell stout (Fig. 167);

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

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

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

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1-SR rather angled to 1-M; cu-a somewhat reclivous and curved; 1-M slightly curved. Hind wing: marginal cell parallel-sided submedially and slightly widened apically (Fig. 167); 2-SC+R short and subquadrate; m-cu present as slightly pigmented vein; M+CU:1-M = 6:5; 1r-m 0.7 × 1-M. Legs. Tarsal claws with fine brownish pecten basally; hind coxa finely coriaceous, with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 5.7 and 4.5 × their width, respectively; inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus; hind basitarsus wider than following segments. Metasoma. First tergite 1.3 × as long as wide posteriorly, flattened and latero-anteriorly widely lamelliform; first–second tergites densely finely irregularly rugulose and with fine median carina; second tergite as long as wide basally and 1.4 × as long as third tergite (Fig. 169); minute medio-basal area of second tergite present; second suture rather deep, widened medially and distinctly crenulate; third and following tergites superficially coriaceous and shiny; fourth tergite largely without sharp lateral crease; ovipositor sheath largely densely setose and apically truncate. Colour. Yellowish brown; antenna (except yellow scapus and pedicellus) dark brown; stemmaticum black; hypopygium, middle and hind tarsi more or less infuscate; pterostigma and veins of middle third of wings dark brown (Fig. 167); other veins brownish yellow; wing membrane subhyaline. Variation. Length of fore wing 6–10 mm, of body 7.5–11.5 mm; antennal segments of ♀ 58(1), 59(2), 60(3), (and of f. dendrolimi: 60(2), 61(2), 62(1), 63(1)), of ♂ 55(1), 56(2), 57(1), 58(2), 59(2), 61(1) (of f. dendrolimi: 54(1), 61(1)); latero-anterior lamella of first tergite rather wide or narrow; marginal cell of hind wing parallel-sided or slightly narrowed submedially; f. dendrolimi has head partly, palpi, mesosoma ventrally and posteriorly, metasoma and legs more or less dark brown or blackish; rarely nearly entire head black. 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. Aleiodes jakowlewi (Kokujev, 1898) Figs 190–202 Rhogas (Aleiodes) jakowlewi Kokujev, 1898: 307. Aleiodes jakowlewi; Shenefelt 1975: 1176; Papp 1991: 112. Rogas jakowlewi; Tobias 1986: 82 (transl.: 136).

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Figure 190. Aleiodes jakowlewi (Kokujev), ♀, Finland, Kangaslampi, habitus lateral.

Material. *Finland: 2 ♀ (NMS), Pohjois-Savo, Kangaslampi, Malaise trap, 26.vi– 17.vii.2004 and 19.vii–1.viii.2005, N.M. Laurenne; 1 ♀ (I. Kakko collection) Finland, Loppi Topeno, 67410xx: 33468xx, 24.viii.2006, I. Kakko; *Sweden: 2 ♀, 1 ♂ (BMNH), Skåne, Röstanga 6.vii.1938, D.M.S & J.F. Perkins; *Slovakia: 1 ♀ (MTMA), Smolnieká Huta, 2.vii.1956, M. Čapek. Molecular data. MRS355 (Finland JF962849, CO1). Biology. Nothing is known of the biology of this predominantly boreal species. Diagnosis. Head transverse in dorsal view and directly narrowed ventrally; eye rather large; antenna of ♀ sometimes with a narrow white or pale yellowish submedial band, scapus and pedicellus of ♀ similarly coloured as medial fifth of antenna; antennal segments of ♀ 49–52; OOL equal to width of posterior ocellus; length of malar space of ♀ 0.30–0.40 × (of ♂ 0.25 times) height of eye in lateral view (Fig. 200); speculum of mesopleuron rugose or reticulate and dull as remainder of mesopleuron; propodeum distinctly elongate (Fig. 190); fore wing narrow (Fig. 191); vein m-cu of fore wing straight and angled to vein 2-CU1 (Fig. 191); pterostigma dark brown with its basal third pale yellow; hind femur 4.9–5.5 × as long as its maximum width; hind trochantellus 2.2–2.7 × as long ventrally as wide (Fig. 194); hind tibia infuscate subapically, contrasting with yellowish apex of tibia (Fig. 194); tarsal claws only bristly

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setose, without distinct pecten; first metasomal tergite with lamella latero-anteriorly (Fig. 202), second metasomal tergite rather long; second tergite with small smooth triangular area medio-basally (Fig. 193); third tergite coarsely sculptured; fourth tergite with distinct sharp lateral crease and basally rugulose. Very similar to the East Palaearctic A. parentalis Belokobylskij, 2000; this species differs mainly by the wider hind femur, the subbasally yellowish first tergite (except two dark brown patches) and the more robust apical antennal segments. Description. Redescribed ♀ (NMS) from Kangaslampi (Finland), length of fore wing 4.5 mm, of body 5.5 mm. Head. Antennal segments of ♀ 49, length of antenna 1.2 × fore wing, its subapical segments 1.7–1.8 × as long as wide; frons granulate, with satin sheen; OOL and POL 1.0 and 0.8 × width of posterior ocellus, respectively; vertex distinctly rugulose-granulate, with satin sheen; clypeus convex and coriaceous; ventral margin of clypeus thick and convex (Fig. 199); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 199) and face mainly coarsely granulate with some rugulae; length of eye 3 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum rugulose-coriaceous; occipital carina complete, with short crenulae and dorsally curved (Figs 200–201); clypeus partly above lower level of eyes (Fig. 199); length of malar space 0.35 × height of eye in lateral view; eyes rather protruding (Figs 199–201). Mesosoma. Length of mesosoma 1.8 × its height; mesoscutal lobes coriaceous, matt, but medio-posteriorly longitudinally rugose and anteriorly low; notauli narrow and crenulate; prepectal carina medium-sized, remaining separate from anterior border; precoxal area of mesopleuron and area above it largely rugose (Fig. 192); remainder of mesopleuron (including speculum) granulate and with satin sheen; metapleuron distinctly granulate and with satin sheen; mesosternal sulcus deep and sparsely crenulate; mesosternum rather angulate posteriorly; scutellum slightly convex, coriaceous, and laterally with carina; propodeum flattened, without tubercles and largely coarsely rugose, median carina complete. Wings. Fore wing: r 0.5 × 3-SR (Fig. 191); 1-CU1 horizontal, 0.3 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 191); cu-a inclivous, parallel with CU1b, straight (Fig. 191); 1-M straight and 1-SR angled to 1-M. Hind wing: marginal cell parallel-sided; 2-SC+R short; m-cu short and only weakly pigmented; M+CU:1-M = 15:10; 1r-m 0.7 × 1-M. Legs. Tarsal claws with yellow bristles and small; hind coxa rugose-granulate, with satin sheen and 0.9 × as long as first tergite; hind trochantellus 2.7 × longer ventrally than wide; length of fore and hind femora 7.0 and 4.9 × their width, respectively (Figs 194–195); inner apex of hind tibia without distinct comb; length of inner hind spur 0.25 × hind basitarsus. Metasoma. First tergite 1.2 × as long as wide posteriorly, stout, convex anteriorly and latero-anteriorly distinctly lamelliform; first–fourth tergites densely finely rugose (Fig. 193), with distinct median carina up to middle of third tergite; medio-basal area of second tergite minute; second suture medium-sized and crenulate; third and fourth tergites with complete sharp lateral crease; fifth and following tergites retracted; ovipositor sheath mainly densely setose and apically acute.

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

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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 23rd–27th 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 leptofemur sp. n. http://zoobank.org/B3F2C302-894D-4DF9-89A0-3C9737CFC373 Figs 203–214 Aleiodes ?nigriceps; Shaw 1983: 319, 321. Aleiodes borealis; Shaw 1994: 134, 136, 137. Aleiodes borealis; Papp and Rezbanyai-Reser 1996: 73, 96. Aleiodes borealis; Belokobylskij et al. 2003: 398. Aleiodes nigriceps auctt. p.p. (not Wesmael 1838). Type material. Holotype, ♀ (NMS, Edinburgh), “[England:] Norfolk, Santon, ex Stilbia anomala on Deschampsia flexuosa, 18.xii.[20]01, mum. ?ii.[20]02, em. 12.v.[20]02, G. M. Haggett”, “MRS Aleiodes DNA 154”. Paratypes (475 ♀, 412 ♂) from England (West Cornwall, East Cornwall, South Devon, North Devon, Isle of Wight, North Somerset, North Wilts, South Wilts, Dorset, North Hants, South Hants, West Sussex, East Sussex, East Kent, West Kent, Surrey, South Essex, Herts, Middlesex, Berks, Oxford, Bucks, West Suffolk, East Norfolk, West Norfolk, Cambridge, Bedford, Hunts, Northampton, East Gloucester, West Gloucester, Warwick, Stafford, Leicester, Derby, Chester, South Lancaster, Mid-west York, North-west York, Westmorland), Scotland (Dumfries, Ayr, Lanark, Berwick, Haddington, Edinburgh, Linlithgow, Fife, Stirling, West Perth, East Perth, Elgin, Easterness, Westerness, Dunbarton, Kintyre, West Ross), Wales (Glamorgan, Merioneth, Caernarvon, Monmouth, Pembroke, Anglesey), Isle of Man, Guernsey, Jersey, Andorra (St. Julia), Austria (Niederösterreich: Raglitz; Poysdorf; Hainburg; Oberösterreich: Linz, Kirchschlag; Sensengebirge Bärenriedlau), Belgium (Mt. St. Pierre), Bulgaria (Rodopi: Galabovo), Cyprus (Yermasoyia Riv.; Yeroskipon; Cherkes; Limasol; Paramytha), Czech Republic (Sumava, 1000–1300m; Moravia: Mikulov), France (Orsay; Hautes-Alpes: Briançon), Finland (Kainuu: Kuhmo), Germany (Bonn: Rheinhöhenweg im Kottenforst; Schleswig-Holstein: Lübeck; Niedersachsen: Berkhof; Gottingen: Hann-Munden; Bonn; Rheinland, Köln-Flittard; Bavaria: Wiesen/Spessart; Lower Saxony: Harzburg; Mullingen; Alfeld; Oberhaverbeck; Hürth-Fischenich; Ahlem; Saxony: Zöbigker; Baden-Württemberg:

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

Heidel/Jagdl; Goslar, Astfeld), Gibraltar (Botanical Garden), Greece (Lakonia, Parnon Oros, 1700 m; id., Taygetos, 1000–1200 m; Thessalia: Mt Olympos; Pisadia; Kozani). Hungary (Budapest: Budaliget; Budakeszi, Hársbokorhegy; Pécs, Tettye), Ireland (WI:

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Ballinclea; SK: Caragh Lake; DU: Phoenix Park; Carlow, Antrim), Italy (Tuscany: Gorgona Isl.; Sicily: Catania; Mt. Etna, Milo Fornazzo, 800 m; Bolzano, Sarntal; 1250 m; Südtirol: Ahrntal; Trentino: Riva s. Garda; Trentino: Tremalzo), Netherlands (DR: Wijster; Borger, FL: Lelystad (Oostvaardersplassen & Jagersbos), GE: Gortel; Ede; Epe; Heerde; Nunspeet; Tongeren, NB: Baarle-Nassau; Eindhoven; Geertruidenberg; Hoogerheide; Nijmegen, Berg en Dal; Vierlingsbeek; Etten-Leur; Hilvarenbeek; Rijen; Bergen op Zoom; Oss; Raamdonksveer; Tilburg (Kaaistoep), LI: Arcen; Geulle; Lemelerberg; Neercanne; Wrakelberg; Grubbenvorst; St. Pietersberg; Vilt; Wolder, NH: Weesp; Texel, Oudeschild, UT: Linschoten, ZH: Meijendel; Oostvoorne; Ouddorp; Lexmond; Melissant; Nieuwkoopse Plassen; Noordwijk; Rotterdam; Voorschoten; Waarder, FR: Ried; Terschelling, ZE: Haamstede; Westenschouwen), Norway (Oppland, Lom-Lia), Slovakia (B. Karpaty-Jaktar, Drietoma), Spain (Teruel, Tramacastilla; Navarra, Alsasua, 600 m; Mallorca, Porto Cristo; Zaragoza: Juslibol), Sweden (Skåne: Järahusen; Ystad; Böste; Spraggehusen; Ö. Väringe; Spukke; Härjedalen: Tänndalen; Duvberget; Halland: Åsa Närsbokrok), and Switzerland (GR: Sent Surains, Val Gronda, 1500 m). Paratypes in NMS, AAC, BMNH, BZL, M. Riedel collection, RMNH, H. Schnee collection, SYKE, MSC, ZJUH, MCZ and ZSSM. Molecular data. MRS154 (England KU682229, CO1), MRS156 (England JF962813, CO1), MRS157 (England KU682230, CO1), MRS515 (Netherlands KU682260, CO1). Biology. A parasitoid of a wide range of low feeding noctuid larvae, as listed below. Overwinters as a small larva in the host, which is killed before it is in its final instar. Mummy (Fig. 204) largely dark brown (summer generations paler) and slender, usually formed in a prominent position at least in spring. Specimens (in NMS unless indicated) reared from wild-collected Noctuidae identified as Abrostola triplasia (Linnaeus) (1; J.L. Gregory), Ammoconia caecimacula (Denis & Schiffermüller) (4:1; J. Connell/Austria), Autographa gamma (Linneaus) (2; G.E. King/Spain, E. Haeselbarth/ Germany; 1 (RMNH), G. Peters/Germany), ?Cerastis rubricosa (Denis & Schiffermüller) (1; J.L. Gregory), Cucullia chamomillae (Denis & Schiffermüller) (2; A.A. Allen [1 is AAC]), ?Diarsia rubi (Vieweg) (1; T.H. Ford), Dicestra trifolii (Hufnagel) (1; G.M. Haggett), Euplexia lucipara (Linnaeus) (6:1; P. Baker), Lacanobia oleracea (Linnaeus) (2:1; P. Baker), Melanchra pisi (Linnaeus) (5; P. Baker, A. Lord, M.R. Shaw), Mythimna ferrago (Fabricius) (1; J. L. Gregory), ?Mythimna impura (Hübner) (3:1; M. R. Hall), ?Mythimna littoralis (Curtis) (1; F.D. Bennett), Noctua comes Hübner (14; J. Connell[?], D. Hackett[?], G.M. Haggett, E. Haeselbarth, R. Hinz, R.A. Softly), Noctua fimbriata (Schreber) (2; G.M. Haggett, R.A. Softly), Noctua interjecta Hübner (2; M.R. Hall, G.M. Haggett), Noctua janthina (Denis & Schiffermüller) (1; G.M. Haggett), Noctua orbona (Hufnagel) (3; G.M. Haggett), Noctua pronuba (Linnaeus) (5 [2 are BMNH]; R.A. Softly), Orthosia gracilis (Denis & Schiffermüller) (1; M.R. Shaw), Paradiarsia glareosa (Esper) (1; G.M. Haggett), Phlogophora meticulosa (Linnaeus) (10; M.R. Shaw, R.A. Softly, J. Voogd), Shargacucullia verbasci (Linnaeus) (24 [5 are OUM, 1 is AAC]; A.A. Allen, F.C. Woodforde, M.R. Shaw), Standfussiana lucernea (Linnaeus) (1; R.F. Logan), Stilbia anomala (Haworth) (4; G.M. Haggett), Xestia agathina (Duponchel) (12 [6 are ZSSM, 2 are AAC]; A.A. Allen, E. Bauer, A. Dobson,

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

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M.R. Shaw), Xestia baja (Denis & Schiffermüller) (1; J.L. Gregory), Xestia castanea (Esper) (8 [5 are ZSSM]; E. Bauer, K.P. Bland[?], M.R. Shaw), Xestia xanthographa (Denis & Schiffermüller) (31 [1 is ZSSM, 1 is RMNH]; E. Bauer, M.R. Britten, J. Connell, G.M. Haggett, M.R. Hall, N. Hall, R. Hinz, M.R. Shaw, R.A. Softly), and unidentified noctuids (81 [14 are BMNH], mostly collected as mummies). Specimens in NMS reared in culture experiments are included in the type material (but not in the above host list), and these experimental results (using females from the overwinter generation to parasitize summer hosts) are as follows: Diarsia rubi 1:16\16\\7+4 [several others were retarded but died]; Dicestra trifolii 5:28\28\\24+4; Lacanobia oleracea 6:28\22\\0+16 [several others were retarded but died]; Melanchra persicariae (Linnaeus) 4:22\13[several others pricked and paralysed without oviposition]\\0+13; Orthosia cerasi (Fabricius) 7:55\6\\5+1; Orthosia gothica (Linneaus) 6:40\1\\0+1; Orthosia gracilis 8:27\16\\14+2; Orthosia incerta (Hufnagel) 6:23\4\\2+1; Phlogophora meticulosa 9:49\44\\29+14: Shargacucullia verbasci 1:1\1\\1+0. In Britain adult flight times peak around May, July–August and October. However, in common with most Aleiodes, the adults are very long-lived and can be found in most months (but we have no British records from January to March). Both sexes come feely to light, but it is also active by day. The males tend to court non-conspecific females (at least of some species) with as much – though unsuccessful – vigour as with conspecifics. All host records are from Noctuidae feeding on low plants, but within that group this species has an unusually broad host range. Many (but not all) of the overwintering hosts, such as Noctua and Xestia species (parasitized from September to November), feed on Poaceae and grow slowly during mild periods in the winter, with mummification by the parasitoid in the host’s 3rd or 4th 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

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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. Description. Holotype, ♀, length of fore wing 4.5 mm, of body 4.4 mm. Head. Antennal segments 39, length of antenna 1.1 × fore wing, its subapical segments about 1.8 × as long as wide (Fig. 211) and basal segments comparatively wide (Fig. 210); frons granulate-coriaceous, with satin sheen; OOL 1.5 × diameter of posterior ocellus and coriaceous; vertex granulate-coriaceous, with satin sheen; clypeus distinctly convex (Fig. 213), coriaceous; ventral margin of clypeus rounded and depressed (Fig. 212); width of hypoclypeal depression 0.4 × minimum width of face and face coriaceous, dorsally somewhat rugulose (Fig. 212); length of eye 2.5 × temple in dorsal view and temple gradually roundly narrowed behind eye (Fig. 214); occiput

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behind stemmaticum granulate-coriaceous, occipital carina interrupted by somewhat more than width of ocellus (Fig. 214); clypeus partly above lower level of eyes (Fig. 212); length of malar space 0.4 × length of eye in lateral view; eyes moderately protruding (Figs 212–214). Mesosoma. Mesoscutal lobes finely coriaceous, with satin sheen, medio-posteriorly with a few rugulae; notauli complete and narrow, largely smooth and posteriorly reduced; prepectal carina narrow lamelliform, reaching anterior border; precoxal area of mesopleuron granulate-coriaceous, mesopleuron with superficially granulate and shiny speculum and rugose dorso-anteriorly (Fig. 206); metapleuron granulate-coriaceous, matt and posteriorly not tuberculate; mesosternal sulcus narrow and rather deep, shallow medio-posteriorly and no carina; mesosternum angulate posteriorly; scutellum elongate, slightly convex, granulate-coriaceous and laterally largely without carina; propodeum rather flat dorsally, not tuberculate latero-posteriorly, and coriaceous with median carina complete. Wings. Fore wing: r 0.3 × 3-SR (Fig. 205); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.6 × 2-SR, and 0.35 × 3-SR; second submarginal cell comparatively large (Fig. 205); cu-a weakly oblique, not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apically marginal cell nearly twice as wide as its minimum width; 2-SC+R short and longitudinal; m-cu short and not pigmented (Fig. 205). Legs. Tarsal claws setose; hind coxa finely coriaceous, largely matt; hind trochantellus 2.4 × longer than wide; length of fore and hind femora 6.9 and 6.9 × their width, respectively (Figs 208–209); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.1 × as long as wide posteriorly and latero-posteriorly narrowly lamelliform, moderately convex and flattened posteriorly, dorsope mediumsized (Fig. 207); first and second tergites densely and longitudinally rugulose (Fig. 207), with distinct median carina; medio-basal area of second tergite absent; second suture narrow and finely crenulate; third tergite coriaceous and remainder of metasoma largely smooth and shiny; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath (except dorsally) moderately setose. Colour. Black; antenna dark brown; palpi largely brown; tegulae, malar space ventrally and triangular patch on second tergite pale yellowish (Fig. 207); inner orbita as dark as face centrally; outer orbita posteriorly and dorsally, malar space dorsally and mesoscutum medio-posteriorly brownish yellow (Fig. 213); mesopleuron with ventral brownish yellow stripe (Fig. 206); hind coxa largely, apical half of hind femur, base of hind tibia and tarsi rather fuzzy dark brown (Figs 203, 208) and remainder of legs yellowish brown; veins and pterostigma (except yellow basal 0.3 and slightly apex) dark brown; border between dark and pale part of pterostigma rather sharp, contrasting with each other (Fig. 205); wing membrane subhyaline. Variation. Length of fore wing 4.5–5.0 mm; antennal segments of ♀ 35(3), 36(16), 37(79), 38(98), 39(70), 40(20), 41(1), 42(1), of ♂ 35(22), 36(64), 37(89), 38(60), 39(17), 40(4), 41(1)); mesosoma largely black to largely orange brown; medial length of second tergite 0.8–0.9 × its basal width; OOL of male slightly longer than diameter

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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”). Distribution. *Andorra, *Austria, *Belgium, *British Isles (England, Wales, Isle of Man, Scotland, Ireland, Guernsey, Jersey), *Bulgaria, *Cyprus, *Czech Republic, *France, *Finland, *Germany, *Gibraltar (British territory), *Greece, *Hungary, *Italy, *Netherlands, *Norway, *Slovakia, *Spain, *Sweden, *Switzerland. The southern European countries are included provisionally (see above under variation). 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. Aleiodes modestus (Reinhard, 1863) Figs 215–229 Rogas modestus Reinhard, 1863: 271; Shenefelt 1975: 1177; Papp 1985a: 160 (lectotype designation); Tobias 1986: 83 (transl.: 138). Aleiodes modestus; Papp 1991: 99; Belokobylskij et al. 2003: 398. Rhogas (Aleiodes) modestus var. piceus Fahringer, 1932: 302–303; Shenefelt 1975: 1177. Type material. Lectotype ♀ (ZMB) from Germany examined. 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; Wales:

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V.C.s 48, 52: Scotland: V.C.s 88, 96, 97), Bulgaria, *Czech Republic, *Denmark, Finland, *France, Germany, Italy, Netherlands (DR: Borger; Wijster, GE: Heerde; Brummen (Voorstonden), FR: Terschelling (Midsland-Noord, dunes), LI: Epen, ZH: Asperen; Meijendel (dunes)), Poland, *Romania, Russia, Slovakia, Sweden, Switzerland. Specimens in NMS, MNHN, BMNH, RMNH, NRSM, BZL, MTMA, USNM, CNC, CMIM, M. Riedel coll., JLC, Delémont, ZSM, MCZ, SDEI, FRAH, WAE, UWIM, ZMC, ZMUO. Molecular data. MRS282 (Wales JF962850, CO1 + KU682267, 28S). Biology. A univoltine parasitoid of a wide range of Eupithecia species (Geometridae: Larentiinae), mummifying the host in its pupation chamber and overwintering in the mummy. Specimens reared from wild collected hosts determined as follows: Eupithecia absinthiata (Clerck) (2 NRS), Eupithecia exiguata (Hübner) (1 ZMUH), Eupithecia gelidata hyperboreata Staudinger (2 SDEI), Eupithecia goossensiata Mabille (2 NMS, 1 OUM; T.H. Ford), Eupithecia innotata (Hufnagel) (2 ZMUH, 1 ZMUO); I. Itämes), Eupithecia lariciata (Freyer) (1 NMS, 2 Copenhagen, 2 Delémont, 10 FRAH), Eupithecia nanata (Hübner) (22 NMS, 5 BMNH, 5 OUM]; T.H. Ford, G.T. Lyle), Eupithecia satyrata (Hübner) (1 NRS), Eupithecia subfuscata (Haworth) (3 OUM; T.H. Ford), Eupithecia succenturiata (Linnaeus) (1 NMS, 1 OUM; T.H. Ford), Eupithecia vulgata (Haworth) (6 NMS, 2 OUM; T.H. Ford, M.R. Shaw) and 13 (NMS) from undetermined Eupithecia spp on various low plants (including Artemisia, Lotus and Pimpinella). Experimental culture result in E. vulgata 2:16\16\\7+1. While it has been reared particularly from hosts feeding on field layer plants (as is the habit of the majority of European Eupithecia species), the rearing records also include a substantial number from E. lariciata on Larix; however, it seems probable that many of the specimens seen were reared in a nursery context, in which young Larix would have presented as part of the field layer, and it is perhaps significant that A. modestus was not found to be a prominent parasitoid of E. lariciata on mature Larix sampled in the Alps (although it was indeed reared from it in small numbers at most sampling sites: Kenis et al. 2005). The host larva is usually fully grown and cocooned in the soil as a prepupa by the time mummification occurs, and the wide and sharp-rimmed clypeus of the adult probably reflects the need to chew its way through the host’s cocoon and soil. The winter is passed in the mummy, which is rather tough and dark, slightly dorso-ventrally flattened but not keeled (Fig. 216), and sometimes weakly upcurled. Typically the thoracic and first two abdominal segments are contracted, with the parasitoid occupying approximately abdominal segments 3–8 in a thin silken lining. The mummy is not stuck down, though probably the usual ventral opening for the expulsion of fluid occurs none the less (however, this is unconfirmed). The adult flight time in Britain is approximately from late June through August. Diagnosis. Antennal segments of both sexes 37–(40–45)–47 and third segment rather slender (Fig. 223); ventral margin of clypeus thin and protruding (Fig. 226); maximum width of hypoclypeal depression about 0.5 × minimum width of face (Fig. 225); length of malar space of female 0.2–0.3 × height of eye in lateral view (Fig. 226); OOL about equal to diameter of posterior ocellus (Fig. 227); area in front of

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

anterior ocellus with a minute smooth tubercle (Fig. 227); vein r of fore wing 0.5–0.6 × vein 3-SR; vein 1-SR of fore wing short (Fig. 217); vein 1r-m of hind wing distinctly shorter than vein 1-M (Fig. 217); mesopleuron above precoxal area strongly shiny and superficially sculptured; third tergite without distinct striae; pterostigma largely yellow

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or yellowish brown; inner side of basal half of hind tibia yellowish; head and mesosoma largely blackish. Description. Figured ♀ (RMNH) from Wijster (Netherlands), length of fore wing 5.4 mm, of body 5.6 mm. Head. Antennal segments of ♀ 43, moderately setose, length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide (Fig. 224); frons rather flat and granulate; OOL equal to diameter of posterior ocellus and finely granulate; vertex finely granulate, with satin sheen; clypeus rather large, micro-granulate; ventral margin of clypeus thin and rather protruding forwards (Fig. 226); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 225); length of eye 1.9 × temple in dorsal view (Fig. 227); occiput behind stemmaticum mainly granulate with some rugulae; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view; occipital carina complete; with a minute smooth tubercle in front of anterior ocellus. Mesosoma. Mesoscutal lobes densely and finely granulate and with punctulation, matt; prepectal carina complete, distinct; precoxal area of mesopleuron with some rugulae medially; mesopleuron above precoxal area strongly shiny, sparsely punctate and with some superficial micro-granulation (Fig. 218); metapleuron largely granulate; scutellum flat, micro-granulate and without lateral carina; propodeum convex, finely rugose and interspaces micro-granulate (Fig. 219), without tubercles, median carina complete, weak and rather irregular. Wings. Fore wing: r 0.6 × 3-SR (Fig. 217); 1-CU1 horizontal, slender, 0.35 × 2-CU1; r-m 0.5 × 3-SR; second submarginal cell rather long (Fig. 217); cu-a vertical, straight posteriorly; 1-M slightly curved posteriorly. Hind wing: marginal cell subparallel-sided but slightly constricted medially, its apical width equal to width at level of hamuli (Fig. 217); 2-SC+R subquadrate; m-cu faintly indicated; M+CU:1-M = 5:4; 1r-m 0.6 × as long as 1-M. Legs. Tarsal claws yellowish setose; hind coxa granulate; hind trochantellus rather robust; length of fore femur, hind femur and basitarsus 5.7, 5.2 and 8.6 × their width, respectively (Figs 220–221); length of inner hind spur 0.25 × hind basitarsus. Metasoma. First tergite rather robust (Fig. 219); first and second tergites densely and rather finely rugose, with rather weak median carina, reduced posteriorly; medio-basal area of second tergite absent; length of second tergite 0.9 × its basal width; second suture rather deep and distinctly crenulate; third tergite 0.8 × as long as second tergite, anterior two-thirds rugose and remainder of metasoma superficially granulate and punctate, somewhat compressed; fourth and apex of third tergite without sharp lateral crease; ovipositor sheath widened and setose. Colour. Black; palpi, tegulae and pterostigma yellow; veins (except basally) dark brown; malar area ventrally, orbita dorsally and posteriorly, pronotum anteriorly and ventrally, mesopleuron antero-dorsally and postero-ventrally, hind coxa (but basally black), medio-posterior part of mesoscutum, first tergite latero-posteriorly, second and third tergites dark reddish brown; apical half of hind femur largely dark brown, except

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

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apically; antenna, telotarsi and clypeus dark brown; remainder of legs yellowish brown or blackish; wing membrane subhyaline. Variation. Antennal segments of ♀ 38(1), 39(0), 40(0), 41(1), 42(18), 43(27), 44(12), 45(4) and of ♂ 40(1), 41(1), 42(2), 43(37), 44(18), 45(2); length of antenna of ♂ 1.3 × fore wing; length of eye of ♀ 1.2–1.9 × temple in dorsal view, of ♂ 1.6–2.5 times; dark specimens have mesosoma, metasoma and hind coxa nearly completely black; pale specimens have scapus, pedicellus, clypeus, entire orbita, notaulic area of mesoscutum, metapleuron posteriorly, first tergite apically and most of second–fourth tergites dark reddish brown; pterostigma sometimes slightly infuscate laterally, but remaining largely yellow; the infuscation of the hind femur is sometimes diffuse; vein cu-a of fore wing vertical or oblique; rugae of precoxal area may be entirely absent. Note. The number of antennal segments does not differ appreciably between the sexes. The specimens reported from Hungary by Papp (1983, 1985a, 2005) are misidentified A. fortipes (Reinhard). Aleiodes nigriceps Wesmael, 1838 Figs 230–242 Aleiodes nigriceps Wesmael, 1838: 109; Papp 1985a: 156 (lectotype designation). Aleiodes circumscriptus var. nigriceps; Shenefelt 1975: 1171. Type material. Lectotype, ♀ (KBIN) from Belgium examined. Additional material. *Austria, Belgium, British Isles (*England: V.C.s 3, 8, 12, 19, 20, 22, 23, 24, 25, 27, 28, 29, 30, 31, 33, 40, 58, 62, 65; *Wales: V.C.s 41, 45, 52; *Isle of Man: V.C. 71; *Scotland: V.C.s 75, 77, 83, 84, 89, 97, 99, 101, 105; *Ireland: V.C. H5), *Jersey, *Bulgaria, *Czech Republic, *Finland, France, Germany, Hungary, *Italy, *Lithuania, *Netherlands (FR: Terschelling (Formerum), GE: Putten, NB: Geertruidenberg; Oploo; Udenhout (de Brand), NH: Overveen; Muiderberg, LI: Tegelen; St. Pietersberg; Venlo; Maastricht, UT: Harmelen, ZE: Cadzand; Oostkapelle; ZH: Delft; Den Haag; Wassenaar; Leidschendam), *Norway, *Poland, *Portugal, Russia, *Slovakia, Spain, *Sweden and Switzerland. Specimens in NMS, BMNH, RMNH, BZL, SDEI, ZSSM, FMNH, MTMA, CNC, P.-N. Libert collection, JLC, WAE, H. Haraldseide collection, and MSC. Molecular data. MRS075 (England KU682221, CO1), MRS144 (Scotland KU682228, CO1), MRS613 (Wales KU682243, CO1), MRS783 (Austria KU682250, CO1). Biology. At least partly a plurivoltine parasitoid of larvae of low-feeding noctuine Noctuidae inhabiting grassy sites; most frequent in humid situations. Mummy (Fig. 231) dark brown or brown and slender, only slightly or not swollen, forming low down in concealment. Specimens (in NMS unless stated otherwise) reared from wild-collected Noctuidae identified as Diarsia rubi (Vieweg) (10 [2 are ZSSM]; G.M. Haggett, M.R. Shaw), Lycophotia porphyyrea (Dennis and Schiffermüller) (1 [ZSSM/

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

Netherlands]), Noctua fimbriata (Schreber) (1; J. Connell/Austria), Xestia c-nigrum (Linnaeus) (1; J. Connell/Austria), Xestia sexstrigata (Haworth) (3; G.M. Haggett), and indet. green noctuid (1). A further specimen was reared from a noctuid mummy

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collected in a Carex tussock (D.G. Notton). Specimens in NMS reared in culture experiments as follows: Xestia sexstrigata 6:20\20\\15+2, Xestia xanthographa (Denis & Schiffermüller) 4:47\47\\3+30 [several others were retarded but died], Noctua fimbriata 6:45\40\\24+0 [several others were retarded but died], Diarsia rubi 6:89\83\\74+9, Phlogophora meticulosa (Linnaeus) 7:43\19\\0+12 [several others were retarded but died]. No ovipositions resulted from the following culture experiments: Diarsia mendica (Fabricius) 1:5\0\\-, Xanthorhoe fluctuata (Linnaeus) 3:19\0\\-, Xanthorhoe montanata (Denis & Schiffermüller) 3:15\0\\-, Camptogramma bilineata (Linnaeus) 6:13\0\\-. While many of its hosts are univoltine with overwintering larvae, in which the parasitoid overwinters as an early instar larva, A. nigriceps has the capacity to be plurivoltine (e.g. in D. rubi), and its long summer flight period, including abundant males in late summer, indicates that it regularly is so. Experimental rearings suggest that even within the genera it uses as hosts it specialises narrowly, and these experiments also clearly distinguish between it and both A. pictus and A. leptofemur. The mummy forms low down, more or less in concealment, and is very seldom collected (unlike that of A. leptofemur). The adult emerges soon after. In captivity adult females of this species are long-lived, but have always died off during the winter, and the flight period is ca May–September. Males are sluggish in their courtship, even towards conspecific females which, however, typically offer themselves for mating immediately. 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

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

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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). Description. Redescribed ♀ (NMS), “Culture: [Wales], Anglesey, Llangristiolus, from ♀ 3:2 [ex] Diarsia rubi in Xestia sexstrigata, ovip. 28.v.[20]08, mum. 4.iv.[20]09, em. 22.v.2009, died viii.2009, ♀ 2/09, M.R. Shaw”. Length of fore wing 4.4 mm, of body 5.5 mm. Head. Antennal segments 43, length of antenna 1.3 × fore wing, its subapical segments about 1.6 × as long as wide; frons granulate-coriaceous, with satin sheen; OOL 1.6 × diameter of posterior ocellus (POL 1.4 times); vertex granulate-coriaceous, with satin sheen; clypeus distinctly convex and largely smooth; ventral margin of clypeus thick and depressed (Fig. 239); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 239) and face mainly coriaceous with some rugulae medio-dorsally; length of eye 2.3 × temple in dorsal view and temple roundly narrowed behind eye (Fig. 241); occiput behind stemmaticum coriaceous and occipital carina absent medio-dorsally, its interruption slightly less than width of stemmaticum (Fig. 241); clypeus partly above lower level of eyes (Fig. 239); length of malar space 0.4 × height of eye in lateral view; eyes moderately protruding (Figs 239–241). Mesosoma. Mesoscutal lobes finely coriaceous, with satin sheen, but medio-posteriorly longitudinally rugose; notauli narrow, moderately impressed and finely crenulate, but posteriorly lost in rugose area; prepectal carina narrow lamelliform and reaching anterior border; precoxal area of mesopleuron coriaceous and without rugae medially; mesopleuron above precoxal area (except small smooth and shiny speculum) coriaceous, but antero-dorsally rugose; medially metapleuron coriaceous, matt; mesosternal sulcus narrow and moderately deep, but posteriorly shallow and with a carina; mesosternum rather angulate posteriorly; scutellum nearly flat, coriaceous and largely non-carinate laterally; propodeum rather flat and coriaceous but posteriorly with some rugae, median carina complete, without tubercles. Wings. Fore wing: r 0.2 × 3-SR (Fig. 232); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell medium-sized (Fig. 232); cu-a oblique, but not parallel with CU1b, straight; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell somewhat widened apically (Fig. 232); 2-SC+R short and longitudinal; short stub of m-cu present, pigmented. Legs. Tarsal claws setose; hind coxa superficially coriaceous, with satin sheen; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 6.3 and 5.6 × their width, respectively (Figs 235–236); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.2 × as long as wide posteriorly, convex, but posteriorly flattened; first and second tergites and base of third tergite densely longitudinally rugose; second tergite robust (Fig. 234), with distinct median carina, matt; medio-basal area of second tergite obsolescent; second suture shallow and crenulate; third tergite (except basally) largely coriaceous, remainder of metasoma largely smooth and rather

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shiny; fourth and apical third of third tergite without sharp lateral crease; ovipositor sheath largely densely setose. Colour. Black or brownish black; antenna brown, but scapus and pedicellus dorsally and laterally and apical third of antenna dark brown; palpi and tegulae pale yellowish (Fig. 230); outer orbita posteriorly and dorsally brownish yellow (Figs 240–241) and remainder of head black; mesopleuron (except dorsally), mesosternum and metapleuron medially orange brown; hind femur (except basally) dark brown (Fig. 235); scutellum, first tergite medio-apically, second and third tergites, part of fourth tergite and remainder of legs brownish yellow; veins (except dark brown veins 1-SR, 1-M, r and CU1) brown; pterostigma yellowish, but anteriorly and posteriorly darkened (Fig. 232); wing membrane subhyaline. Variation. Length of fore wing 4.3–4.4 mm; antennal segments of ♀ 38(1), 39(9), 40(44), 41(44), 42(27), 43(13), 44(2), 45(1); of ♂ 37(1), 38(4), 39(31), 40(82), 41(113), 42(33), 43(12), 44(7), 45(3); vein r of fore wing 0.2–0.5 × vein 3-SR; length of first tergite of ♀ 1.1–1.2 × its apical width; pterostigma medially yellow or dark brown; medio-posteriorly mesoscutum black or brownish yellow; second and third tergites yellowish, infuscate or dark brown laterally, mesosternum orange brown or infuscate. 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). Aleiodes nigricornis Wesmael, 1838 Figs 243–256 Aleiodes nigricornis Wesmael, 1838: 105; Shenefelt 1975: 1178; Papp 1991: 112, 1985: 160 (lectotype designation); Belokobylskij et al. 2003: 398. Type material. Lectotype, ♀ (KBIN) from Belgium examined. Additional material. *Austria, Belgium, British Isles (England: V.C.s 2, 3, 7, 11, 16, 17, 20, 22, 23, 24, 25, 29, 30, 31, 34, 38, 57, 58, 59, 60, 63, 64, 65, 66, 67, 69; Wales: V.C.s 35, 44, 48, 52; Scotland: V.C.s 72, 75, 78, 79, 83, 85, 86, 87, 88, 89, 90, 95, 96, 97, 98, 99, 101, 102, 105, 107, 108, 111; Ireland: V.C.s H21, H28, H29, “Westport”), Czech Republic, *Denmark, France, Finland, Germany, Italy, Netherlands (GE: Ede; Nunspeet; Heerde; Otterlo, Velp, LI: Grubbenvorst, NB: Geertruidenberg; Helvoirt; Bergen op Zoom; Etten-Leur, NH: Overveen; Muiderberg, ZH: Den Haag; Meijendel; Oegstgeest; Voorschoten; Asperen; Waarder), *Norway, Poland, Russia, Slovakia, Sweden, Switzerland. Specimens in NMS, BMNH, RMNH, BZL, MTMA, NRS, SDEI, CC, FMNH, ZSSM, OUM, NMI, CMIM,

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Figures 243–244. Aleiodes nigricornis Wesmael, ♀, England, Gait Barrows. 243 habitus lateral 244 mummy of Orthosia gothica (Linnaeus).

Sheffield Museum, SMNS, ZISP, SYKE, ZMUO, USNM, CNC, UWIM, M. Riedel collection, JLC, MSC, AAC, WAE, I. Kakko collection, H. Haraldseide collection. Molecular data. MRS216 (Scotland EU979585, CO1 + AJ784934, 28S), MRS373 (Sweden KU682258, CO1), MRS790 (Scotland KU682254, CO1), MRS794 (England KU682255, CO1). Biology. A plurivoltine parasitoid of Noctuidae, using Apamea species in which to overwinter, and (possibly exclusively) Orthosia gothica (Linnaeus) in early summer. The mummy is moderately dark brown, rather elongate (Fig. 244) and often found in exposed positions. Specimens of the overwintering generation reared from noctuids

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identified as Apamea crenata (Hufnagel) (2 ZSSM, 1 NMS, 1 NRS; E. Haeselbarth, R.I. Lorimer), Apamea epomidion (Haworth) (2 ZSSM, 1 OUM; E. Haeselbarth), Apamea ?monoglypha (Hufnagel) (1 NMS; M.R. Britton), Apamea ?remissa (Hübner) (1 H. Haraldseide/Norway) and ?Apamea sp. (11, hosts unidentified or misidentified but with preserved mummies that are consistent with Apamea species (det. M.R. Shaw), having at least moderately large shining warts and a well-developed prothoracic plate: 7 (2 as Xestia xanthographa (Denis & Schiffermüller)) NMS, 4 (as Polia spp.) OUM, 1 (as Noctua fimbriata (Schreber)) RMNH, 1 AAC). Specimens of the summer generation reared from Orthosia gothica (3 NMS; R.I. Lorimer, M.R. Shaw), and unidentified noctuid mummies compatible with O. gothica (8 NMS, 1 ZMUO, 1 ZSSM, 2 H. Haraldseide collection). Other specimens (both generations) with a host recorded (Mythimna ferrago (Fabricius) in ZSSM, Epirrita autumnata (Borkhausen) in CC) have not been accompanied by mummies and we regard the records as dubious. Experimental results from female ex Apamea ?monoglypha; with 2nd instar hosts: Orthosia gothica 1:9\9\\6\3, Orthosia cerasi (Fabricius) 1:10\ 0\\-, Orthosia incerta (Hufnagel) 1:4\0\\-; with 3rd instar hosts O. gothica 1:8\8\\0\8. The single female trialed with hosts undertook considerable non-destructive concurrent host feeding on O. gothica. Adult flight time approximately May to August, with females persisting until October. 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). Description. Redescribed ♀ (RMNH) from Helvoirt, length of fore wing 5.6 mm, of body 6.2 mm. Head. Antennal segments 46, length of antenna 1.2 × fore wing, its subapical segments about twice as long as wide; frons mainly granulate, posteriorly with some rugulae, rather shiny and mainly flat; OOL 0.8 × diameter of posterior ocellus and granulate-coriaceous; vertex superficially granulate-coriaceous, with satin sheen; clypeus convex, coriaceous; ventral margin of clypeus thick and depressed (Fig. 253); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 251) and face coriaceous and transversely rugose, except ventrally; length of eye 2.2 × 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

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Figures 245–256. Aleiodes nigricornis Wesmael, ♀, England, Gait Barrows. 245 wings 246 mesosoma lateral 247 mesosoma dorsal 248 propodeum and metasoma dorsal 249 hind leg lateral 250 fore femur lateral 251 head anterior 252 head dorsal 253 head lateral 254 apical segments of antenna 255 basal segments of antenna 256 antenna.

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ocellus (Fig. 252); clypeus above lower level of eyes (Fig. 251); length of malar space 0.25 × height of eye in lateral view; eyes moderately protruding (Figs 251–253). Mesosoma. Mesoscutal lobes finely granulate and with satin sheen; notauli narrow, moderately deep and smooth; prepectal carina lamelliform medio-ventrally, reaching anterior border; precoxal area of mesopleuron finely granulate and with some rugulae medially (Fig. 246); mesopleuron above precoxal area (except small and nearly smooth speculum) granulate, but dorsally rugose; medially metapleuron granulate and with satin sheen; mesosternal sulcus narrow and rather deep, without carina posteriorly; mesosternum rather angulate posteriorly (Fig. 246); scutellum nearly flat, granulate and largely non-carinate laterally; propodeum rather flat and granulate but posteriorly with some rugae, median carina complete and regular, evenly convex laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 245); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.4 × 3-SR; second submarginal cell rather long (Fig. 245); cu-a slightly oblique, not parallel with CU1b, straight; 1-M straight posteriorly. Hind wing: apical half of marginal cell slightly widened (Fig. 243); 2-SC+R short and longitudinal; m-cu absent, except for a faint trace; M+CU:1-M = 6:5; 1r-m 0.65 × 1-M. Legs. Tarsal claws setose; hind coxa superficially coriaceous, with satin sheen; hind trochantellus 2.3 × longer than wide; length of fore and hind femora 6.2 and 5.0 × their width, respectively (Figs 249–250); hind femur granulate-coriaceous; inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.2 × as long as wide posteriorly, flattened and latero-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. Colour. Black or brownish-black; antenna dark brown; palpi, tegulae, tibiae (except apically), medio-apical fifth of first tergite and medially second tergite pale yellowish (Fig. 248); malar space nearly up to eyes, orbita dorsally and posteriorly, mesopleuron (except dorsally), mesosternum, metapleuron largely, mandible and remainder of legs, more or less reddish brown; veins (but of middle third of wing mainly dark brown) and pterostigma (but posterior border somewhat darkened) yellowish brown; wing membrane subhyaline. Variation. Antennal segments of ♀ 42(2), 43(4), 44(10), 45(44), 46(73), 47(59), 48(12), 49(5); of ♂ 42(3), 43(24), 44(20), 45(30), 46(15), 47(9), 48(1); mesosternum reddish brown or partly fuzzy dark brown; precoxal area of mesopleuron medially entirely granulate or rarely with some weak rugulae; scapus entirely dark brown or partly brown. Notes. The lectotype has 46 antennal segments. On average males have 1–2 fewer antennal segments than females.

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Aleiodes pallidator (Thunberg, 1822) Figs 257–271 Ichneumon pallidator Thunberg, 1822: 259. Aleiodes pallidator; Shenefelt 1975: 1179; Papp 1991: 101; Belokobylskij et al. 2003: 398. Rogas pallidator; Tobias 1986: 81 (transl.: 135). Rogas ochraceus Curtis, 1834: 512.4; Shenefelt 1975: 1182 (as synonym of A. testaceus) (examined). Aleiodes ochraceus; Papp 1991: 101 (as synonym of A. pallidator); Belokobylskij et al. 2003: 399 (as synonym of A. similis). Aleiodes unicolor Wesmael, 1838: 111; Roman 1912: 271 (synonym of A. pallidator); Shenefelt 1975: 1179–1180; Papp 1985a: 160 (lectotype designation) (examined). Type material. Lectotype of I. pallidator here designated, ♀ (ZMUU) from Sweden (“α”, “Rhogas (Aleiod.) pallidator Thbg”). Holotype of R. ochraceus, ♀ (Melbourne) from England (Regent’s Park), “Type”, “Rhogas ochraceus Curtis, type, J.F. Perkins, 1948”. Lectotype of A. unicolor, ♀ (KBIN) from Belgium and 2 paralectotypes examined. Additional material. British Isles (England: V.C.s 11, 15, 24, 59, 60), Bulgaria, Netherlands (FL: Bant; Lelystad (Jagersbos), FR: Ried, GE: Ede (Maanderbroek); Zaltbommel (Kerkwijk), GR: Scheemda, LI: Reuven; Vlodrop, NB: Nederweert; Geffen; Valkenswaard; Heusden, ZH: Rotterdam; Lexmond; Melissant), Germany, Hungary, *Romania, Russia, Serbia, Slovakia, Sweden, Turkey. Specimens in NMS, BMNH, RMNH, OUM, ZSSM, MCZ, CC, CNC, UWIM, R. van der Hout collection. Molecular data. MRS001 (Turkey EU979586, CO1 + EU854333, 28S). 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

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Figure 257. Aleiodes pallidator (Thunberg), England, penultimate instar mummies in precocious pseudococoons of Leucoma salicis (Linnaeus).

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

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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. Diagnosis. Antennal segments of ♀ 50–55 and head (except stemmaticum) entirely brownish yellow; scapus in lateral view distinctly oblique apically (Fig. 258); OOL 0.6 × diameter of posterior ocellus; occipital carina complete ventrally or nearly so (Fig. 268); vein 2-CU1 of fore wing 2.2–3.0 × vein 1-CU1 (Fig. 260); vein 1-SR linearly connected to vein 1-M and vein 1-M straight or slightly curved (Fig. 260); fourth metasomal tergite largely (superficially) granulate; length of fore wing 5–7 mm. Easily confused with A. varius (Herrich-Schäffer), but this species has more antennal segments (♀: 66–71), malar space 0.6 × as long as height of eye, vein 2-CU1 of fore wing 1.6–1.8 × vein 1-CU1 and the occipital carina is reduced ventrally. Often also confused with A. gastritor (Thunberg) s. lat., but specimens of this species-group have many fewer antennal segments, the segments are more elongate and the pterostigma is often more or less dark brown or infuscate. Description. Redescribed ♀ (RMNH) from Nederweert, length of fore wing 5.9 mm, of body 7.1 mm. Head. Antennal segments 54, length of antenna 1.2 × fore wing, its subapical segments about 1.8 × as long as wide and scapus in lateral view distinctly oblique apically (Fig. 258); frons superficially granulate and rugose, shiny; OOL 0.6 × diameter of posterior ocellus and rugulose; vertex granulate-rugulose, with satin sheen; clypeus rather high, convex, coriaceous; ventral margin of clypeus thick and gradually depressed (Fig. 268); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 267) and face mainly coriaceous dorsally; length of eye 2.8 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum finely rugulose and occipital carina nearly complete medio-dorsally and ventrally (Figs 268–269); clypeus partly above lower level of eyes (Fig. 267); length of malar space 0.3 × height of eye in lateral view; eyes distinctly protruding (Figs 267–269). Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen, but medioposteriorly rugose; notauli narrow, shallow and crenulate; prepectal carina rather lamelliform medio-ventrally, nearly reaching anterior border of mesopleuron and latero-ventrally angulate; precoxal area of mesopleuron with some short rugae medially (Fig. 262); mesopleuron above precoxal area (except large smooth and shiny speculum) superficially granulate, but dorsally extensively rugose; medially metapleuron coriaceous-rugulose, rather shiny; mesosternal sulcus narrow and rather deep, without carina posteriorly; mesosternum rather angulate posteriorly; scutellum finely coriaceous and non-carinate laterally; dorsal face of propodeum rather long and densely finely rugose, posterior face with some carinae and smooth in between, median carina complete, without tubercles, but somewhat angulate postero-laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 260); 1-CU1 horizontal, 0.35 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell elongate (Fig. 260); 1-SR angled with 1-M; cu-a rather oblique, not parallel with CU1b, slightly curved;

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Figures 258–259. Aleiodes pallidator (Thunberg), ♀, Netherlands, Nederweert. 258 habitus lateral 259 mummy of Leucoma salicis (Linnaeus).

1-M slightly curved posteriorly. Hind wing: apical half of marginal cell parallel-sided or nearly so (Fig. 261); 2-SC+R short and longitudinal; m-cu present as fold, unpigmented; M+CU:1-M = 15:12; 1r-m 0.7 × 1-M.

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Figures 260–271. Aleiodes pallidator (Thunberg), ♀, Netherlands, Nederweert. 260 fore wing 261 hind wing 262 mesosoma lateral 263 propodeum and metasoma dorsal 264 hind leg lateral 265 antenna 266 fore femur lateral 267 head anterior 268 head lateral 269 head dorsal 270 basal segments of antenna 271 apical segments of antenna.

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Legs. Tarsal claws yellowish setose; hind coxa superficially finely coriaceous, with satin sheen; hind trochantellus 2.7 × longer ventrally than wide; length of fore and hind femora 6.2 and 4.9 × their width, respectively (Figs 264, 266); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. 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. Colour. Yellowish brown; apical third of antenna and stemmaticum (except medially) dark brown; palpi, malar space up to eyes, mandible, tegulae, fore and middle legs, hind trochanter and trochantellus and pterostigma pale yellowish (Fig. 260); medial veins dark brown and other veins brownish yellow; wing membrane subhyaline. Variation. Length of fore wing 5.6–6.5 mm, of body 6.3–7.5 mm; antennal segments of ♀ 50(3), 51(13), 52(43), 53(79), 54(44), 55(7); colour and shape are very uniform in this species, probably because of absence of sexual propagation. 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 Papp and 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. Aleiodes pictus (Herrich-Schäffer, 1838) Figs 272–284 Rogas pictus Herrich-Schäffer, 1838: 156 (type series lost); Shenefelt 1975: 1171 (as synonym of A. circumscriptus). Aleiodes pictus; Papp 1991: 113 (as synonym of A. circumscriptus). Type material. Neotype of A. pictus here designated, ♀ (NMS), “Lower Austria, Raglitz, J. Connell, [ex] Camptogramma bilineata, [coll.] 29.iii.2011, mum. 4.iv.[20]11, em. 29.iv.[20]11, [J. Connell reference number] XI 2.05.05 ♀4, died ca. 10.vii.2011”. “Voucher: BFW Sparkling Science Schwarzes C”. Additional material. *Austria, British Isles (*England V.C.s 3, 15, 16, 20, 22, 23, 28, 29, 30, 31, 40, 59, 65, 69; *Scotland: V.C.s 77, 83, 84, 88, 89, 96, 101, 105,

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111; Ireland: “Westport”), *Bulgaria, Czech Republic, *Finland, *France, Germany, Gibraltar (British territory), *Greece, Hungary, *Iceland, *Italy, Netherlands (DR: Drijber, LI: St. Pietersberg, ZH: Den Haag; Meijendel; Goeree; Ouddorp), *Norway, *Poland, *Portugal, *Romania, *Russia, *Serbia, *Slovakia, Spain, Sweden, Turkey. Specimens in NMS, BMNH, OUM, RMNH, ZSSM, SDEI, BZL, ZISP, CMIM, MSC, CNC, UWIM, JLC, H. Schnee collection, H. Haraldseide collection, World Museum Liverpool. Molecular data. MRS518 (Austria KU682261, CO1), MRS549 (Austria KU682241, CO1), MRS556 (Austria KU682242, CO1), MRS719 (Austria KU682245, CO1), MRS784 (Austria KU682251, CO1), MRS785 (Austria KU682252, CO1). Biology. This is a plurivoltine parasitoid, abundant in grassland habitats, of larvae of both geometrids and noctuids feeding in low vegetation, overwintering as a small larva in that of the host. Mummy made low down, more or less in concealment, brown and not swollen (Fig. 273). Specimens (in NMS unless stated otherwise) reared from wild-collected larvae of the larentiine geometrids Camptogramma bilineata (Linnaeus) (78:2; J. Connell [77 specimens from Austria resulting from a long term survey], A.R. Cronin), Epirrhoe alternata (Müller) (1; G.M. Haggett), Lithostege griseata (Denis & Schiffermüller) (3:1; G.M. Haggett), Xanthorhoe fluctuata (Linneaus) (3:2; G.M. Haggett, G.E. King), and from the diverse range of noctuids Hoplodrina ambigua (Denis & Schiffermüller) (2:1; J. Connell/Austria), Hoplodrina sp. (blanda (Denis & Schiffermüller), octogenaria (Goeze) and superstes (Ochsenheimer) co-occurring but indistinguishable when small) (17:1; J. Connell/Austria), Agrotis exclamationis (Linnaeus) (1; M.R. Shaw/France); Noctua fimbriata (Schreber) (1; J. Connell/Austria); Xestia xanthographa (Denis & Schiffermüller) (1; J. Connell/Austria), Phlogophora meticulosa (Linnaeus) (1; M.R. Shaw); Syngrapha interrogationis (Linnaeus) (1 ZMUO; J. Itämes/ Finland). The Austrian material reared by J. Connell resulted from a prolonged survey of grassland caterpillars at a single site. Experimental cultures (specimens in NMS) as follows: Epirrhoe alternata 6:27\1\\1+0, Xanthorhoe fluctuata 2:45\40\\25+14 [other females even more willing to oviposit (6: 86\84) but cultures lost to disease], Xanthorhoe montanata (Denis & Schiffermüller) 3:15\15\\8+2, Xestia xanthographa 2:2\0\\-, Xestia sexstrigata (Haworth) 2:3\1\\0+1, Noctua fimbriata 1:2\0\\- [This female was then boxed with 10 host larvae for 5 days: 6 hosts pupated; the other 4 died and on dissection were found to contain living parasitoid larvae], Diarsi rubi (Vieweg) 3:37\18\\0+12, Phlogophora meticulosa 2:17\0\\-. The males are aggressive and rather indiscriminate in courtship, seeming as interested in females of A. nigriceps as their own species, but the A. nigriceps females always repelled them successfully. Males of A. pictus are, however, somewhat less interested in females of A. leptofemur. Mating trials between specimens from Hoplodrina spp. and Camptogramma bilineata (all from the same site in Austria) repeatedly indicated that females from both series were less willing to mate with males from the other than with males reared from the same host as themselves; however, several cooperative matings (that then seemed normal, though of often shorter duration) were obtained and (from very limited trials) no difference was

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Figures 272–273. Aleiodes pictus (Herrich-Schäffer), ♀, neotype. 272 habitus lateral 273 mummy of Camptogramma bilineata (Linnaeus).

seen in behaviour towards hosts by females of different host origin (but Hoplodrina larvae were not available for experiments). However, the possibility that specimens from Hoplodrina (which differ slightly in their morphology, particularly colour) are at least some way towards representing a genetic isolate cannot be ruled out, as most of the experimental results involved females from C. bilineata (but also the female from Noctua fimbriata whose behaviour was essentially the same). A host range encompassing both Geometridae and Noctuidae is unusual in the genus Aleiodes, but the species is evidently rather specialised to a narrow range of taxa in each family. The adult flight period is May to September in Britain. Diagnosis. Antennal segments of ♀ (35–)36–40, of ♂ (36–)37–41; pale area of orbita of ♀ rather wide (Figs 281–282); mesopleuron with some rugulae; mesoster-

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num almost always strongly darkened or black and usually sharply defined; width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 280); OOL 1.3  × diameter of posterior ocellus; mesoscutum antero-laterally and pronotum medio-anteriorly very often brownish yellow; length of first tergite of ♀ 1.0–1.1 × its apical width; second tergite often less wide than in A. nigriceps (Fig. 276); fore and hind tarsi comparatively slender (Figs 277–278); pterostigma yellow anteriorly; middle to distal third of hind femur partly or entirely dark brown; mummy not swollen, usually light brown (Fig. 273). Description. Neotype, ♀, length of fore wing 4.6 mm, of body 4.9 mm. Head. Antennal segments 37 (right) and 38 (left), length of antenna 1.2 × fore wing, its subapical segments about 1.6 × as long as wide; frons granulate and some rugulae, with satin sheen; OOL 1.3 × diameter of posterior ocellus (POL equal to ocellus); vertex superficially granulate, with satin sheen; clypeus distinctly convex and largely nearly smooth; ventral margin of clypeus thick and depressed (Fig. 281); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 280) and face mainly granulate-coriaceous with some rugulae medially; length of eye 2.8 × temple in dorsal view and temple gradually narrowed behind eye (Fig. 282); occiput behind stemmaticum granulate-coriaceous and occipital carina absent medio-dorsally and interruption slightly less than width of stemmaticum (Fig. 282); clypeus partly above lower level of eyes (Fig. 280); length of malar space 0.4 × length of eye in lateral view; eyes moderately protruding (Figs 280–282). Mesosoma. Mesoscutal lobes finely granulate-coriaceous, with satin sheen, but medio-posteriorly rugose; notauli narrow, moderately impressed and finely crenulate, but posteriorly merging in rugose area; prepectal carina narrow lamelliform and reaching anterior border; precoxal area of mesopleuron coriaceous and with some rugae medially (Fig. 275); mesopleuron above precoxal area (except small nearly smooth and shiny speculum) antero-dorsally distinctly rugose and granulate-coriaceous; medially metapleuron granulate, matt; mesosternal sulcus narrow and rather deep, but posteriorly shallow and with a fine carina; mesosternum rather angulate posteriorly; scutellum nearly flat, superficially coriaceous and only antero-laterally carinate; propodeum rather flat and coriaceous with rugae, median carina nearly complete, without tubercles. Wings. Fore wing: r 0.3 × 3-SR (Fig. 274); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.7 × 2-SR, and 0.4 × 3-SR; second submarginal cell slender (Fig. 274); cu-a slightly oblique, not parallel with CU1b, straight; 1-M slightly curved posteriorly. Hind wing: apical half of marginal cell somewhat widened apically (Fig. 274); 2-SC+R subquadrate; m-cu medium-sized, unpigmented; M+CU:1-M = 50:39; 1r-m 0.6 × as long as 1-M. Legs. Tarsal claws setose; hind coxa granulate-coriaceous, matt; hind trochantellus 2.5 × longer than wide; length of fore and hind femora 6.1 and 4.9 × their width, respectively (Figs 277–278); hind femur pimply; inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.1 × as long as wide posteriorly, convex, but posteriorly flattened; first and second tergites densely longitudinally rugose; second tergite slen-

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Figures 274–284. Aleiodes pictus (Herrich-Schäffer), ♀, neotype. 274 wings 275 mesosoma lateral 276 propodeum and metasoma dorsal 277 hind leg lateral 278 fore femur lateral 279 antenna 280 head anterior 281 head lateral 282 head dorsal 283 basal segments of antenna 284 apical segments of antenna.

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

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rather shiny; fourth and apical half of third tergite without sharp lateral crease; ovipositor sheath moderately setose. Colour. Black or brownish black; antenna brown, but scapus and pedicellus dorsally and laterally and apical half of antenna dark brown; malar space (except near eye), palpi, tegulae, fore and middle coxae, trochanters and trochantelli, first tergite medioposteriorly, second tergite medially and third tergite narrowly medially pale yellowish (Fig. 276); orbita posteriorly and dorsally widely brownish yellow (Figs 281–282) and remainder of head black; dorso-posteriorly pronotum brown; mesoscutum with notaulic and medio-posterior area brownish yellow; mesosternum narrowly anteriorly behind prepectal carina, ventral half of mesopleuron and posteriorly orange brown; hind femur (except basally and apex) dark brown (Fig. 272); scutellum dark reddish brown medially; veins (except dark brown veins 1-SR, 1-M, r and CU1) brown; pterostigma pale yellowish, but posteriorly somewhat darkened (Figs 272, 274); wing membrane subhyaline. Variation. Length of fore wing 3.8–4.6 mm; antennal segments of ♀ 34(1), 35(5), 36(27), 37(52), 38(64), 39(38), 40(9) and of ♂ 36(9), 37(22), 38(39), 39(34), 40(18), 41(4); vein r of fore wing 0.4 × vein 3-SR; clypeus 0.3–0.4 × as wide as face; length of first tergite of ♀ 1.0–1.1 × its apical width; pterostigma medially and anteriorly yellow; malar space largely dark brown to largely pale yellow; pronotum medio-anteriorly and scutellum brownish yellow or dark brown; medially metapleuron black or orange brown. 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 1975: 1244; Tobias 1986: 78 (transl.: 128). Aleiodes praetor; Papp 1991: 73; Belokobylskij et al. 2003: 398. Neorhogas luteus Szépligeti, 1906: 606; Shenefelt 1975: 1205; Papp 1977: 115 (as synonym of A. praetor); 1991: 73; 2004: 215 (lectotype designation); Chen and He 1997: 37. Type material. Holotype of A. praetor ♀ (MNHN) from France (“Mout. [= Moutiers, Savoie]”, “Moutiers”, “Muséum Paris, 1867, Coll. O. Sichel”, “Rogas praetor Rhd.”) and lectotype of N. luteus ♂ (MTMA) from Serbia examined. Additional material. Austria, Belgium, British Isles (England: V.C.s 5, 11, 16, 17, 19, 20, 21, 22, 24, 30, 31, 34, 38, 62, 64), *Bulgaria, Croatia, Finland, France, Germany, Hungary, Netherlands (GE: Heerde; LI: Stein, Epen, Tegelen; NH: Naardermeer; UT: Bilthoven, Leersum; ZH: Melissant, Oostkapelle), Serbia, Spain, Sweden, Switzerland. Specimens in NMS, MNHN, BMNH, CMIM, OUM,

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Figure 285. Aleiodes praetor (Reinhard), ♀, hanging from a leaf edge.

RMNH, FMNH, NRSM, MTMA, S. Dodd collection, P. McMullen collection, WAE, UWIM. Molecular data. MRS067 (England KM067256/KU682219, CO1 + EU854334, 28S), MRS654 (Bulgaria HQ551265/KU682244, CO1). 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

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Figures 286–287. Aleiodes praetor (Reinhard), ♀, Netherlands, Epen. 286 habitus lateral 287 mummy of Mimas tiliae (Linnaeus) from England.

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

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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. Description. Redescribed ♀ (RMNH) from Naardermeer, length of fore wing 8.7 mm, of body 8.8 mm. Head. Antennal segments of ♀ 72, with many tyloids and apex of subbasal segments oblique (Fig. 296), length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide and apical segment with long spine (Fig. 297); frons narrow, rather flat and largely micro-granulate; OOL 0.3 × diameter of posterior ocellus and micro-sculptured; vertex flat, micro-sculptured and shiny; clypeus convex, microgranulate and near lower level of eyes; ventral margin of clypeus not differentiated (Fig. 299); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 298); face micro-sculptured and partly transversely rugulose; length of eye 3.2 × temple in dorsal view (Fig. 300); occiput behind stemmaticum superficially sculptured; length of malar space 0.2 × length of eye in lateral view; occipital carina strong, but medio-dorsally absent; eyes distinctly protruding (Figs 298–300). Mesosoma. Mesoscutal lobes densely punctate, micro-sculptured and shiny; prosternum rather large and distinctly concave; prepectal carina complete, distinct; precoxal area of mesopleuron with some striae medially; mesopleuron above precoxal area strongly shiny, punctate medio-posteriorly and remainder smooth (Fig. 290); metapleuron punctate dorsally and rugose ventrally; scutellar sulcus wide, deep and with 7 carinae; scutellum flat, densely punctate, with striae medio-posteriorly and lateral carina largely present and lunula narrow; propodeum convex, dorsal face about as long as posterior face, densely rugose, tuberculate protruding latero-dorsally (Fig. 290), propodeal spiracle large and median carina of propodeum complete and regular. Wings. Fore wing: r 0.6 × 3-SR (Fig. 288); 1-CU1 nearly horizontal, slender, 0.2 × 2-CU1; r-m 0.4 × 3-SR and not pigmented; second submarginal cell rather long (Fig. 288); cu-a inclivous, curved posteriorly; 1-M straight posteriorly. Hind wing: marginal cell subparallel-sided basally, constricted near basal 0.7 and its apical width nearly equal to width at level of hamuli (Fig. 289); 2-SC+R short; m-cu absent; M+CU:1-M = 33:16; 1r-m 1.1 × as long as 1-M.

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Figures 288–301. Aleiodes praetor (Reinhard), ♀, Netherlands, Epen. 288 fore wing 289 hind wing 290 mesosoma lateral 291 propodeum and metasoma dorsal 292 hind leg lateral 293 fore femur lateral 294 antenna 295 ovipositor lateral 296 basal segments of antenna 297 apical segments of antenna 298 head anterior 299 head lateral 300 head dorsal 301 hypopygium and ovipositor sheath lateral.

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Legs. Tarsal claws yellowish setose; hind coxa punctate and micro-sculptured dorso-basally and remainder largely smooth and punctulate; hind trochantellus ventrally twice as long as wide; length of fore femur, hind femur and basitarsus 6.3, 4.2 and 7.8 × their width, respectively (Figs 292–293); length of inner hind spur 0.35 × hind basitarsus; inner apex of hind tibia with indistinct weak comb. 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). Colour. Yellowish brown; antenna (but scapus brownish basally), stemmaticum, apical third of hind tibia (except spurs) and hind tarsus largely, black; base of hind tibia pale yellowish; pterostigma and veins brownish yellow; wing membrane largely subhyaline, but basally slightly pigmented and near veins 1-SR and 1-M slightly infuscate. Variation. Antennal segments of European ♀ 67(2), 68(7), 69(7), 70(7), 71(3), 72(4), 73(4); of ♂ 62(3), 63(5), 64(5), 65(5), 66(1), 67(1); males have fifth–seventh tergites moderately setose; vein m-cu of fore wing sometimes slightly curved and gradually merging into 3-CU1; precoxal sulcus entirely smooth or with some striae; scapus and pedicellus partly yellowish brown or entirely black. 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. Aleiodes reticulatus (Noskiewicz, 1956), stat. rev. Figs 302–315 Rhogas reticulatus Noskiewicz, 1956: 176 (examined). Aleiodes reticulatus; Shenefelt 1975: 1181; Papp 1991: 96 (as synonym of A. arcticus). Type material. Holotype, ♀ (PAN), “[Poland], 15/1 [19]48 Itame fulvaria Vill. 1.II. [19]49”, “Rhogas - ♂ reticularis [sic] Nosk.”, “Holotyp. (lgz. pnedui)”, “Holotypus ♀ % Rhogas reticulatus Nosk. 1956. Papp 1983”. Paratypes: 1 ♀ (glued on same card as holotype, with one emerged mummy of geometrid (compatible with Macaria sp.) with same labels and “Aleiodes ♀ arcticus Th. det. Papp J., 1983”, “Syntypus Rhogas reticulatus Noskiewicz 1954”; 2 ♂ “15/1/ [19]48 Karczewski Jędrzejów 15.xii.48”, “Rhogas - ♂ reticularis [sic] Nk”, “Allotypus Rhogas ♂ reticulatus Nosk. 1956. Papp 1983”, “Aleiodes ♂ arcticus Th. det. Papp J. 1983”, “Syntypus Rhogas reticulatus Noskiewicz, 1954”.

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Figures 302–304. Aleiodes reticulatus (Noskiewicz), ♀, Russia, Rostchino, but 304 from Poland. 302 habitus lateral 303 mummy of cf. Macaria wauaria (Linnaeus) 304 mummy of Macaria brunneata (Thunberg) forming, at end will resemble 303.

Additional material. 2 ♀, 2 ♂ (3 ZISP, 1 NMS), *Belarus, Zubky, 120 km W of Minsk, ex Macaria wauaria on Ribes nigrum, em. 1–8.v.1984 (Silvanovich); 1 ♀ (SDEI), *Germany, Mecklenburg, Fürstenberg, 21.v.1888 (Konow); 1 ♂ (SDEI)

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Germany, Mecklenberg, Kalkhof, 30.iv.1890 (F.W. Konow); 1 ♀ (NMS) Germany, Saxony, Tharandt, Fichtenwald, 18.v.1980 (Walter); 2 ♂ (NMS), Poland, Pomeranian, Czarne, ex Macaria brunneata on Vaccinium myrtillus, coll. 24.v.2014, mummies 27.v.2014, em. 27.iii.2015 (M.R. Shaw); 1 ♂ (ZISP) *Russia, Voronezezhskij zapovednik ex geometrid [host mummy compatible with Macaria wauaria] on Ribes nigrum, 18.iv.1950 (Donvar); 1 ♀ (ZISP) Russia, Rostchino, NW of St Petersburg, 15.vi.1966 (V.I. Tobias). In addition several mummies were obtained from M. brunneata larvae collected from Vaccinium myrtillus in Polish conifer forests in v. 2014, both at the type locality (Kielce: Jędrzejów, Lasków forest), and also at Ruciane-Nida (Warmian-Masurian) and Biebrza (Podalaskie), but the adults failed to develop (mummies in NMS). 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). Diagnosis. Maximum width of hypoclypeal depression 0.4 × minimum width of face (Fig. 311); OOL twice diameter of posterior ocellus; mesoscutum, orbita and malar space black; precoxal sulcus granulate; trochanters, trochantelli and pterostigma largely black(ish); mesoscutum with a fine longitudinal carina on mesoscutum medioposteriorly and more or less anteriorly, but sometimes absent; apical half of marginal cell of hind wing parallel-sided or slightly widened; vein M+CU1 of fore wing apically above level of vein 2-CU1 (Fig. 305); vein r of fore wing 0.9–1.1 × as long as vein 3-SR (Fig. 305); vein 1-SR of fore wing slightly angled with vein 1-M; all femora and tibiae dark reddish brown; fore and hind femora moderately stout (Figs 313–314); fourth metasomal tergite curved posteriorly in dorsal view (Fig. 308), its lateral crease distinct and following tergites more or less retracted (Fig. 302); length of fore wing 3.9–4.7 mm.

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Figures 305–315. Aleiodes reticulatus (Noskiewicz), ♀, Russia, Rostchino. 305 wings 306 mesosoma dorsal 307 mesosoma lateral 308 metasoma dorsal 309 antenna (tip absent) 310 basal segments of antenna 311 head anterior 312 head lateral 313 fore femur lateral 314 hind leg lateral 315 head dorsal.

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Description. Redescribed ♀ (NMS) from Zubky (Belarus), length of fore wing 4.7 mm, of body 5.1 mm. 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). Mesosoma. Mesoscutal lobes largely granulate-coriaceous, matt and medio-posteriorly rugose, middle lobe with a complete longitudinal carina, but weakly developed anteriorly (Fig. 306); notauli narrow, rather shallow and finely crenulate; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron granulate; mesopleuron above precoxal area (except large smooth and shiny speculum) granulate, but dorsally finely rugose; medially metapleuron granulate and with some rugae, rather shiny; mesosternal sulcus narrow and rather shallow; mesosternum rounded posteriorly; scutellum moderately convex, shallowly impressed medio-anteriorly, mainly granulate and non-carinate laterally; propodeum rather directly lowered posteriorly and granulate-rugose, median carina complete, without tubercles. Wings. Fore wing: r nearly as long as 3-SR (Fig. 305); 1-CU1 oblique, 0.3 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.9 × 3-SR; second submarginal cell rather short (Fig. 305); vein M+CU1 of fore wing apically above level of vein 2-CU1 (Fig. 305); vein 1-SR of fore wing slightly angled with vein 1-M; cu-a slightly oblique, not parallel with CU1b; 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically (Fig. 305); 2-SC+R short and longitudinal; m-cu present, pigmented; M+CU:1-M = 25:14; 1r-m slightly oblique and 0.6 × as long as 1-M. Legs. Tarsal claws setose; hind coxa granulate-coriaceous, with satin sheen and nearly reaching apex of first tergite; hind trochantellus 1.8 × longer ventrally than wide; length of fore and hind femora 5.3 and 4.5 × their width, respectively (Figs 313–314); inner apex of hind tibia without comb; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite 0.7 × as long as wide posteriorly, convex and latero-posteriorly non-lamelliform; first–second tergites finely and densely irregularly rugulose and with median carina (Fig. 308); medio-basal area of second tergite absent; second suture medium-sized, deep and distinctly crenulate; third tergite with median carina (but obsolescent posteriorly), third–fourth tergites very finely rugulose-coriaceous; fourth tergite convex medially and apically; fourth tergite with sharp lateral crease; remainder of metasoma largely retracted; ovipositor sheath truncate apically and moderately setose.

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Colour. Black (including coxae); palpi basally, tegulae, pterostigma, veins, trochanters, middle and hind femora dorso-apically and more or less trochantelli dark brown; remainder of palpi and legs brown; wing membrane slightly infuscate, especially near basal veins. Variation. Antennal segments of ♀ 40(1), 41(1), 43(1), 44(1); of ♂ 40(1), 42(1), 43(3), 44(3); male has shape of head just like the examined specimens of true A. arcticus but females have the temple slightly longer and more narrowed; mesoscutum black but one ♀ vaguely brownish near origin of notauli; pale parts of legs brown or orange brown; mesopleuron black or more or less brownish. 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. Aleiodes ryrholmi sp. n. http://zoobank.org/2ED2733A-AA4C-4ECD-9A56-983CA62F5B10 Figs 316–327 Type material. Holotype, ♀ (NMS, Edinburgh), “Sweden: Hr., Sveg, Duvberg, 16.vii.–12.viii.2004, N. Ryrholm, NMSZ 2004.167”, “MRS Aleiodes DNA 395”. Molecular data. MRS395 (Sweden JF962792, CO1). Biology. Unknown. Diagnosis. Antennal segments of ♀ about 40, of ♂ unknown; head strongly directly narrowed behind eyes (Fig. 324); OOL 0.6 × diameter of posterior ocellus; length of malar space of ♀ 0.35 × height of eye in lateral view (Fig. 325); palpi mainly dark brown; face superficially rugulose (Fig. 323); occipital carina interrupted dorsally and complete ventrally (Figs 324–325); eye about 4.5 × as long as temple in dorsal view (Fig. 324); face black; POL 0.6 × as wide as diameter of posterior ocellus (Fig. 324); mesoscutum entirely black (Fig. 319); medio-posterior depression of metanotum rather narrow (Fig. 319); mesosternum and precoxal sulcus superficially granulate and with satin sheen; vein 2-CU1 of fore wing about twice vein 1-CU1 (Fig. 317); vein cu-a of fore wing inclivous; vein 1-SR narrow and linearly connected to vein 1-M and vein 1-M straight (Fig. 317); hind femur stout (Fig. 321); pale area of second metasomal tergite narrow (Fig. 320); fourth tergite largely (superficially) coriaceous and shiny; length of fore wing about 4.5 mm. Very similar to A. cantherius (Lyle), but the new species has a much darker body, larger ocelli and superficially sculptured face. Description. Holotype, ♀, length of fore wing 4.4 mm, of body 4.9 mm. Head. Antennal segments 40, length of antenna 1.2 × fore wing, basal segments rather robust (Fig. 326), its subapical segments about 2.3 × as long as wide (Fig. 327) and scapus in lateral view oblique apically; frons granulate and rather shiny; OOL and POL 0.6 and 0.6 × diameter of posterior ocellus, respectively, and granulate; vertex coriaceous, with satin sheen; clypeus rather high, convex, coriaceous, 0.4 × as wide as face; ventral margin of clypeus thick (Fig. 325); width of hypoclypeal depression

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Figure 316. Aleiodes ryrholmi sp. n., ♀, holotype, habitus lateral.

0.35 × minimum width of face (Fig. 323) and face mainly granulate, dorsally superficially rugulose; length of eye 4.5 × temple in dorsal view and temple directly narrowed behind eye; occiput behind stemmaticum mainly granulate-coriaceous and occipital carina widely interrupted medio-dorsally and nearly complete ventrally (Fig. 325); clypeus above lower level of eyes (Fig. 323); length of malar space 0.35 × height of eye in lateral view; eyes protuberant (Figs 323–325). Mesosoma. Mesoscutal lobes very finely granulate-coriaceous, with satin sheen, but medio-posteriorly rugulose; notauli narrow, shallow and anterior half largely smooth; prepectal carina narrow lamelliform medio-ventrally, nearly reaching anterior border of mesopleuron and latero-ventrally curved; precoxal area of mesopleuron granulate and shiny; mesopleuron above precoxal area (except large smooth and shiny speculum) superficially granulate, but dorsally rugose; medially metapleuron nearly smooth, superficially granulate, shiny; mesosternal sulcus narrow and rather deep, smooth, without carina posteriorly; mesosternum rather angulate posteriorly; scutellum finely coriaceous and non-carinate laterally; lunula (= smooth lateral part of scutellum) wide medially; medio-posterior depression of metanotum rather narrow (Fig. 319); dorsal

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Figures 317–327. Aleiodes ryrholmi sp. n., ♀, holotype. 317 wings 318 mesosoma lateral 319 mesosoma dorsal 320 propodeum and metasoma dorsal 321 hind leg lateral 322 fore femur lateral 323 head anterior 324 head dorsal 325 head lateral 326 basal segments of antenna 327 apical segments of antenna.

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face of propodeum medium-sized, convex and coriaceous, but posteriorly with some carinae and smooth in between, without tubercles, median carina complete. Wings. Fore wing: r 0.55 × 3-SR (Fig. 317); 1-CU1 horizontal and slender, 0.55 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.5 × 3-SR; second submarginal cell elongate (Fig. 317); 1-SR slightly angled to 1-M and slender; cu-a inclivous, subparallel with CU1b, straight; 1-M slightly curved. Hind wing: apical half of marginal cell parallelsided or nearly so (Fig. 317); 2-SC+R longitudinal; m-cu present as weakly pigmented fold; M+CU:1-M = 10:7; 1r-m 0.7 × 1-M. Legs. Tarsal claws yellowish setose; hind coxa superficially finely coriaceous, rather shiny; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 5.2 and 3.9 × their width, respectively (Figs 321–322); inner apex of hind tibia without distinct comb; length of inner hind spur 0.4 × hind basitarsus. 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. Colour. Black; antenna, palpi, basal two-thirds of third tergite, apices of femora, bases of tibiae and hind coxa dark brown; pronotum postero-dorsally, trochanters, trochantelli and tegulae pale yellowish; fore and middle coxae, remainder of tibiae and tarsi brown; orbita (except ventrally and sides of face), femora (except apically), first tergite medio-apically, second tergite narrowly medially, apical third of third and following tergites mainly orange brown (Fig. 320); pterostigma and veins dark brown; wing membrane slightly infuscate. 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. “Aleiodes seriatus (Herrich-Schäffer, 1838)” sensu lato Figs 328–340 Rogas seriatus Herrich-Schäffer, 1838: 156–12, Fig. [type series lost]. Aleiodes seriatus; Papp 1991: 107; Belokobylskij et al. 2003: 399. Aleiodes vittiger Wesmael, 1838: 112; Shenefelt 1975: 1185; Papp 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).

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Figure 328. Aleiodes seriatus (Herrich-Schäffer) s.l., ♀, Netherlands, Wijster, habitus lateral.

Diagnosis. Antennal segments of ♀ (35–)44–50, of ♂ (42–)48–53; length of malar space of ♀ 0.3–0.4 × (of ♂ 0.25 times) height of eye in lateral view (Fig. 338); OOL 0.8 × diameter of posterior ocellus; temple narrow (Fig. 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).

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Figures 329–340. Aleiodes seriatus (Herrich-Schäffer) s.l., ♀, Netherlands, Wijster. 329 wings 330 mesosoma lateral 331 propodeum and metasoma dorsal 332 hind leg lateral 333 fore femur lateral 334 antenna 335 basal segments of antenna 336 apical segments of antenna 337 head anterior 338 head lateral 339 head dorsal 340 inner apex of hind tibia and basitarsus lateral.

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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). Aleiodes testaceus (Telenga, 1941) Figs 341–352 Heterogamus testaceus Telenga, 1941: 134; Shenefelt 1975: 1202; Tobias 1986: 85 (transl.: 142; lectotype designation); Papp 1991: 115 (examined). Aleiodes testaceus; Bergamesco et al. 1995: 5; Papp and Rezbanyai-Reser 1996: 83, 87, 1997: 116; Fortier and Shaw 1999: 221; van Achterberg and Aguiar 2009: 793–794. 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; *Wales: V.C.s 48, 52; *Ireland: H5, H30 and Co. Cork), Bulgaria, *Cyprus, France (*mainland and *Corsica), Greece (mainland), Italy, Morocco, *Netherlands (NB: Tilburg (Kaaistoep)), Portugal (Madeira), Spain (mainland, *Mallorca and *Canary Islands (Gomera; Gran Canaria; Tenerife)). Specimens in NMS, BMNH, RMNH, CNC, BZL, OUM, ZISP, ZSSM, CMIM, UWIM, AAC. Molecular data. MRS057 (Corsica JF973344, CO1) MRS084 (England JF962864, CO1), MRS086 (England KU682223, CO1), MRS103 (Corsica KU682226, CO1), MRS106 (Corsica KU682227, CO1), MRS261 (France JF962863, CO1). 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 (10; T.H. Ford/France, M.R. Shaw/France), Eupithecia sp. (6), Chloroclystis v-ata (Haworth) (9; G.M. Haggett, M.R. Shaw), Gymnoscelis rufifasciata (Haworth) (2; G.M. Haggett, M.R. Shaw). 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

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Figures 341–342. Aleiodes testaceus (Telenga), ♀, England, Bodney. 341 habitus lateral 342 mummy of Gymnoscelis rufofasciata (Haworth).

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

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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. Diagnosis. Antennal segments of female 30–35 (of male 34–37); third antennal segment rather slender (Fig. 348); OOL about equal to diameter of posterior ocellus (Fig. 351); area in front of anterior ocellus with a minute smooth tubercle; ventral margin of clypeus thick (Fig. 350); maximum width of hypoclypeal depression 0.30– 0.35 × minimum width of face; length of malar space of female 0.3–0.4 × height of eye in lateral view (Fig. 350); occipital carina very widely effaced dorsally (Fig. 351); vein r of fore wing 0.7–0.9 × vein 3-SR; vein 1-SR of fore wing rather long; vein 1r-m of hind wing distinctly shorter than vein 1-M (Fig. 343); marginal cell of hind wing somewhat widened apically; inner hind tibial spur 0.25–0.30 × as long as hind basitarsus; tarsal claws setose; pterostigma, inner side of basal half of hind tibia, head and mesosoma largely yellowish, but mesopleuron dorsally and propodeum usually more or less dark brown; third metasomal tergite with more or less developed diverging striae laterally Description. Redescribed ♀ (RMNH) from Garde Freinet (France: Var), length of fore wing 3.3 mm, of body 3.7 mm. 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). Mesosoma. Mesoscutal lobes finely granulate, matt and medio-posteriorly rugose; anterior half of notauli narrow, shallow and smooth and posterior half obsolescent; prepectal carina narrow lamelliform medio-ventrally, not reaching anterior border of mesopleuron; precoxal area of mesopleuron finely rugulose medially (Fig. 344); remainder of mesopleuron above precoxal area nearly smooth or superficially shiny granulate,

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Figures 343–352. Aleiodes testaceus (Telenga), ♀, France, Var, Garde Freinet. 343 wings 344 mesosoma lateral 345 propodeum and metasoma dorsal 346 hind leg lateral 347 fore femur lateral 348 basal segments of antenna 349 head anterior 350 head lateral 351 head dorsal 352 apical segments of antenna.

but dorsally finely rugose; medially metapleuron distinctly granulate and with some rugae, rather shiny; mesosternal sulcus narrow and shallow; mesosternum rounded posteriorly; scutellum nearly flat, densely granulate and laterally non-carinate; propodeum convex, and rugulose-granulate, median carina complete, without tubercles.

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Wings. Fore wing: r nearly as long as 3-SR (Fig. 343); 1-CU1 horizontal, 0.5 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.7 × 3-SR; second submarginal cell short (Fig. 343); vein M+CU1 of fore wing apically at level of vein 2-CU1; vein 1-SR of fore wing rather long and nearly linear with vein 1-M (Fig. 343); cu-a short and vertical, first subdiscal cell narrow (Fig. 343); 1-M nearly straight posteriorly. Hind wing: apical half of marginal cell slightly widened apically; 2-SC+R short and longitudinal; m-cu present as unpigmented fold; M+CU:1-M = 7:5; 1r-m moderately oblique and 0.7 × as long as 1-M. Legs. Tarsal claws yellowish setose; hind coxa granulate and with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 6.4 and 4.7 × their width, respectively (Figs 346–347); inner apex of hind tibia without comb; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite 1.1 × as long as wide posteriorly, convex and latero-posteriorly narrowly lamelliform; first–second tergites and basal 0.6 of third tergite finely longitudinally rugose, interspaces granulate and with median carina (Fig. 345); remainder of metasoma superficially micro-sculptured; medio-basal area of second tergite absent; second suture medium-sized, rather shallow and crenulate; third tergite 0.8 × as long as second tergite; fourth tergite without sharp lateral crease; remainder of metasoma partly retracted; ovipositor sheath truncate apically and moderately setose. Colour. Brownish yellow; stemmaticum, patch on outer side of scapus and pedicellus, apical third of antenna, pronotum dorsally, mesopleuron dorsally, metanotum, metapleuron dorsally, propodeum largely and veins around 1-M of fore wing dark brown; remainder of veins brown or yellowish; palpi, tegulae and pterostigma pale yellowish; wing membrane subhyaline; first tergite somewhat infuscate basally; ovipositor sheath black. Variation. Antennal segments of ♀ 30(1), 31(9), 32(24), 33(78), 34(30), 35(6); of ♂ 34(7), 35(16), 36(18), 37(9); side of pronotum and metasoma of both sexes sometimes largely dark brown or brown except ivory patch of second and third tergites and yellowish brown apex of metasoma. 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. The name “testaceus” (or the invalid emendation “testaceator” by Thunberg (1822)) has been used for 3 different taxa or groups of taxa in the genera Rogas/Rhogas, Aleiodes or Heterogamus (Braconidae: Rogadinae). Most taxa included in the past in Rogas/ Rhogas are now included in the genus Aleiodes Wesmael, 1838 s. l. (Yu et al. 2012) as well as some taxa formerly included in Heterogamus Wesmael, 1838 (including Heterogamus testaceus Telenga, 1941).

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The oldest name is Ichneumon testaceus Fabricius, 1798, not Gmelin, 1790. As a junior homonym Ichneumon testaceus Fabricius is unavailable, and the oldest available name for this taxon is Rogas luteus Nees, 1834 (see van Achterberg 1982). In 1822 Thunberg emended the Fabrician name to “testaceator”, but this emendation is invalid. The senior homonym Ichneumon testaceus Gmelin is a synonym of Monoblastus brachyacanthus (Gmelin, 1790) and belongs to the Ichneumonidae: Tryphoninae (Yu & Horstmann 1997). 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).

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Aleiodes ungularis (Thomson, 1892) Figs 353–365 Rogas (Aleiodes) ungularis Thomson, 1892: 1677 (examined). Aleiodes ungularis; Shenefelt 1975: 1184; Papp 1991: 98, 1985: 163 (lectotype designation). Type material. Lectotype ♀ (ZIL) from Sweden (“Pål”, “ungularis”, “Funnen vid Pålsjo I Skåne, Sverige, teste Papp, 1983”). Additional material. British Isles (*England: V.C. 18; *Wales: V.C. 49; *Ireland: V.C. H19) *Finland, France, Germany, Greece, Hungary, *Romania, Slovakia, Sweden, Switzerland. Specimens in NMS, BMNH, RMNH, FMNH, Doncaster Museum, MTMA, ZSSM, JLC. Molecular data. MRS604 (France JF962867, CO1 + KU682268, 28S). Biology. A probably monophagous and at least partly plurivoltine parasitoid of Pseudopis prasinana (Linnaeus) (Nolidae: Chloephorinae), overwintering as a mummy. The mummy is cylindrical and dark brown (Fig. 354). This is evidently a rare species because its common host is quite often collected in the larval stage, yet we have seen little material. Reared specimens seen from Pseudoips prasinana (5:2; M.R. Shaw/France, T. Weber/Germany). From four host larvae collected in late July (France: Haute-Marne, Fayl Billot) mummification took place over the next few days and all adults emerged in August. Only incomplete observations of its performance with its host in culture could be made, but there was no period of post-oviposition association in the otherwise normal oviposition sequences observed. Two mummies resulted in September from ovipositions obtained in August, the adults emerging the following May and June. Diagnosis. Maximum width of hypoclypeal depression 0.4 × minimum width of face (Fig. 361); OOL 0.6 × diameter of posterior ocellus; mesopleuron largely (and often strongly) shiny; mesosoma (except propodeum and metapleuron) largely yellowish (or orange); precoxal area impressed medially and crenulate (Fig. 356); inner side of hind tibia with whitish comb apically; metasoma dark brown or blackish medially and largely pale yellow laterally (Fig. 357). A distinctive species because of the combination of the shiny mesopleuron with the presence of the hind tibial comb. Description. Redescribed ♀ (NMS) from Ardscull Moat (Ireland), length of fore wing 5.5 mm, of body 5.5 mm. Head. Antennal segments of ♀ 46, with medium-sized setae, length of antenna 1.2 × fore wing, its subapical segments distinctly longer than wide (Fig. 365); frons weakly depressed and with coarse oblique rugae; OOL 0.6 × diameter of posterior ocellus and superficially granulate; vertex granulate, rather dull; clypeus convex, finely granulate and punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 362); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 361); length of eye 3.2 × temple in dorsal view and temple directly narrowed

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Figures 353–354. Aleiodes ungularis (Thomson), ♀, France, Fayl Billot. 353 habitus lateral 354 mummy of Pseudopis prasinana (Linnaeus).

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Figures 355–365. Aleiodes ungularis (Thomson), ♀, France, Fayl Billot. 355 wings 356 mesosoma lateral 357 propodeum and metasoma dorsal 358 hind leg lateral 359 fore femur lateral 360 basal segments of antenna 361 head anterior 362 head lateral 363 head dorsal 364 antenna 365 apical segments of antenna.

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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. Wings. Fore wing: r 0.5 × 3-SR (Fig. 355); 1-CU1 horizontal, 0.45 × 2-CU1; r-m 0.7 × 3-SR; second submarginal cell rather large (Fig. 355); cu-a slightly inclivous, straight; 1-M nearly straight posteriorly. Hind wing: marginal cell subparallel-sided, its apical width 1.6 × width at level of hamuli (Fig. 355); 2-SC+R shortly longitudinal; m-cu obsolescent; M+CU:1-M = 10:7; 1r-m long (Fig. 355) and nearly as long as 1-M. Legs. Tarsal claws yellowish setose; telotarsi 1.5 × wider than fourth tarsal segment in dorsal view; hind coxa superficially granulate; hind trochantellus 1.8 × longer ventrally than wide; length of fore femur, hind femur and basitarsus 5.5, 5.1 and 6.8 × their width, respectively (Figs 358–359); length of inner hind spur 0.4 × hind basitarsus, as long as outer spur; inner apex of hind tibia with whitish comb. 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. Colour. Yellowish brown; head (except orbita dorsally and part of malar space), metapleuron, propodeum and metasoma dorsally (except lateral margins) blackish; antenna, pterostigma, most veins, tibiae (except narrow pale basal ring), hind coxa largely, apical half of middle and hind femora largely, narrowly base of tibiae and telotarsi dark brown; palpi and tegulae yellowish brown; wing membrane subhyaline. Variation. Antennal segments of ♀ 44(1), 45(1), 46(4), 47(2); of ♂ 41(1), 42(2), 43(5), 44(2); head mainly blackish or dark brown; metapleuron blackish or largely yellowish brown; hind coxa and tarsi nearly entirely dark brown or largely (except telotarsus) yellowish brown. In males the metasoma is sometimes entirely dark (without a paler lateral margin) and the hind coxae are dark brown. Note. Males have about three fewer antennal segments than females. In MTMA is a ♀ from Korea with 43 antennal segments and the apical half of hind coxa, hind and middle femora and tibiae dark brown, similar to males from Switzerland.

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Aleiodes varius (Herrich-Schäffer, 1838) Figs 366–378 Rogas varius Herrich-Schäffer, 1838: 156-7, fig.; Shenefelt 1975: 1244–1245 (as synonym of A. procerus). Aleiodes varius; Belokobylskij et al. 2003: 399. Aleiodes procerus Wesmael, 1838: 104; Papp 1985a: 161 (lectotype designation), 1991: 93 (examined). Rogas procerus; Shenefelt 1975: 1244–1245; Tobias 1986: 81 (transl.: 135). Type material. Type series of Rogas varius is lost. Lectotype of Aleiodes procerus ♀ (KBIN), “Belgique, Charleroi/teste Papp J. 1983”, “A. procerus”, “dét. C. Wesmael”, “Lectotypus”, “Aleiodes procerus Wesm. 1838, ♀, Papp, 1983”. One ♀ paralectotype with same label data. Additional material. *Austria, *Finland, *Netherlands (Oisterwijk), Russia. There is a specimen in BMNH labelled “British Isles: Devignes Coll. B.M. 1868–52” but we have seen no other evidence of its occurrence in Britain, and it is probably extinct if indeed it ever occurred. Specimens in NMS, BMNH, FMNH, MSC, I. Kakko collection. Molecular data. MRS446 (Russia HQ551275, CO1). Biology. The only reared specimen seen is from Euthrix potatoria (Linnaeus) (Lasiocampidae) (E.O. Peltonen/Finland). The single date (6.vii.1987) on the label does not suggest voltinism or how the winter is passed, but the specimen is accompanied by a mummy (Fig. 367) whose elongate shape is very different from the relatively short stout mummies of the common Aleiodes alternator (Nees) (in the A. bicolor-group, not treated here) which parasitises the same host (among others). Diagnosis. Antennal segments of ♀ 66–71; head (except stemmaticum) entirely brownish yellow; scapus in lateral view distinctly oblique apically; occipital carina reduced ventrally (Fig. 375); length of malar space 0.4 × height of eye in lateral view; vein 2-CU1 of fore wing 1.6–1.8 × vein 1-CU1 (Fig. 368); vein 1-SR slightly angled to vein 1-M and vein 1-M slightly curved (Fig. 368); vein r of fore wing gradually merging into vein 3-SR; fourth metasomal tergite superficially coriaceous; length of fore wing 8.0–8.5 mm. Description. Redescribed ♀ (NMS) from Lappeenranta (Finland), length of fore wing 8.2 mm, of body 10.1 mm. Head. Antennal segments 69, length of antenna 1.4 × fore wing, its subapical segments about 1.9 × as long as wide and scapus in lateral view distinctly oblique apically (Figs 377–378); frons superficially granulate and with oblique rugae, rather shiny; OOL 0.7 × diameter of posterior ocellus and granulate; vertex granulate-rugulose, with satin sheen; clypeus rather high, convex dorsally but flattened ventrally, coriaceous and with long setae; ventral margin of clypeus thick and gradually depressed (Fig. 375); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 374) and face mainly coriaceous dorsally; length of eye 2.4 × temple in dorsal view and temple directly narrowed behind eye (Fig. 376); occiput behind stemmaticum finely rugose and occipital carina arched and complete medio-dorsally and reduced ventrally

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Figures 366–367. Aleiodes varius (Herrich-Schäffer), ♀, Finland, Lappeenranta. 366 habitus lateral 367 mummy of Euthrix potatoria (Linnaeus).

(Figs 375–376); clypeus partly above lower level of eyes (Fig. 374); length of malar space 0.6 × height of eye in lateral view; eyes distinctly protruding (Figs 374–376). Mesosoma. Mesoscutal lobes very finely coriaceous, with satin sheen, but medioposteriorly with some rugae; notauli narrow, shallow and mainly coriaceous; prepectal carina rather lamelliform medio-ventrally, reaching anterior border of mesopleuron and latero-ventrally angulate; precoxal area of mesopleuron with some fine rugae medially (Fig. 369); mesopleuron above precoxal area (including hardly shiny speculum) superficially coriaceous, but dorsally coarsely rugose; medially metapleuron superficially granulate, rather shiny; mesosternal sulcus narrow and rather deep, without carina posteriorly; mesosternum angulate posteriorly; scutellum coriaceous-rugulose and carinate antero-laterally; dorsal face of propodeum rather long and densely moderately rugose, posterior face with some carinae and smooth in between, median carina complete, without tubercles, but somewhat angulate postero-laterally.

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Figures 368–378. Aleiodes varius (Herrich-Schäffer), ♀, Finland, Lappeenranta. 368 wings 369 mesosoma lateral 370 propodeum and metasoma dorsal 371 hind leg lateral 372 fore femur lateral 373 antenna 374 head anterior 375 head lateral 376 head dorsal 377 basal segments of antenna 378 apical segments of antenna.

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Wings. Fore wing: r 0.3 × 3-SR (Fig. 368); 1-CU1 horizontal, 0.55 × as long as 2-CU1; r-m 0.8 × 2-SR, and 0.5 × 3-SR; second submarginal cell stout (Fig. 368); 1-SR slightly angled to 1-M; cu-a oblique, not parallel with CU1b, straight; 1-M slightly curved posteriorly. Hind wing: marginal cell slightly narrowed submedially and slightly widened apically (Fig. 368); 2-SC+R short and longitudinal; m-cu present as slightly pigmented vein; M+CU:1-M = 17:15; 1r-m 0.6 × 1-M. Legs. Tarsal claws with fine brownish pecten; hind coxa superficially finely coriaceous, with satin sheen; hind trochantellus 2.2 × longer ventrally than wide; length of fore and hind femora 7.5 and 5.3 × their width, respectively (Figs 371–372); inner apex of hind tibia without comb; length of inner hind spur 0.35 × hind basitarsus. 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. Colour. Yellowish brown; apical 0.6 of antenna dark brown; stemmaticum, mesosoma (except largely brownish pronotum, mesoscutum medio-posteriorly, scutellum, metanotum posteriorly) and base of first tergite black or nearly so; pterostigma pale yellowish but apical third infuscate (Fig. 368); medial veins dark brown and other veins brownish yellow; wing membrane subhyaline. Variation. Length of fore wing 8.0–8.5 mm, of body 10.1–10.3 mm; antennal segments of ♀ 67(1), 68(1), 69(1), 70(1), 71(2), of ♂ 65(2), 66(1), 68(2), 69(1); marginal cell of hind wing parallel-sided or slightly narrowed submedially. Note. Females have on average 1–2 more antennal segments than males.

Acknowledgements We thank Pekka Malinen and Marti Koponen (FMNH) for the loan of types and gift of specimens, the hospitality to the first author during his visit to Helsinki, and supplying pictures; for the latter also Stefan Schmidt (ZSSM). Sergey Belokobylskij (ZISP) for his help with type specimens and supplying copies of Russian taxonomic literature. Many entomologists, most of whose names appear in the rearing data, have often gone considerably out of their way to contribute reared (often living) and other material to the NMS collection, and/or have helped MRS by providing host eggs for experimental cultures, all of which has been invaluable for this study. Donald Quicke has arranged for DNA sequencing and analysis of a large number of samples provided by MRS, and given much help in relation to depositing sequences in GenBank. We also thank Jose Fernandez-Triana for additional CO1 sequencing.

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Marsh PM (1979) Braconidae. Aphidiidae. Hybrizontidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America north of Mexico. Washington, 144–313. Matsumura S (1926) On the three species of Dendrolimus (Lepidoptera) which attack spruceand fir-trees in Japan, with their parasites and predaceous insects. Ezhegodnik Zoologicheskago Muzeya [Annales du Musée Zoologique, Leningrad] 26(1925): 27–50. Morley C (1916) Notes on Braconidae XI. The tribe Rhogadides, with additions to the British list. Entomologist 49: 109–112, 125–130. Nees von Esenbeck CG (1834) Hymenopterorum Ichneumonibus affinium monographiae, genera Europaea et species illustrantes 1: 1–320. Stuttgartiae et Tubingae. Noskiewicz J (1956) [Deux espèces nouvelles du genre Rhogas Nees (Hymenoptera, Braconidae) et quelques remarques sur les espèces voissines.] Polskie Pismo Entomologiczne 24: 173–186. [In Polish with French summary] Papp J (1977) Contributions to the Braconid fauna of Yugoslavia. III. Folia Entomologica Hungarica 30: 105–117. Papp J (1983) A survey of the Braconid fauna of the Hortobágy National Park (Hymenoptera, Braconidae), II. In: Mahunka S (Ed.) The Fauna of the Hortobágy National Park. II. Budapest, 315–337. Papp J (1985a) Contributions to the braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae). Folia Entomologica Hungarica 46: 143–164. Papp J (1985b) Braconidae (Hymenoptera) from Korea. VII. Acta Zoologica Hungarica 31: 341–365. Papp J (1991) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae. Magyarorszag Allatvilaga [Fauna Hungariae] 167: 1–122. Papp J, Rezbanyai-Reser L (1996) Contributions to the braconid fauna of Monte Generoso, Canton Ticino, southern Switzerland (Hymenoptera: Braconidae). Entomologische Berichte Luzern 35: 59–134. Papp J, Rezbanyai–Reser L (1997) Zur Brackwespenfauna der Insel Brissago, Kanton Tessin (Hymenoptera: Braconidae). Entomologische Berichte Luzern 38: 113–120. Papp J (2004) Type specimens of the braconid species by Gy. Szépligeti deposited in the Hungarian Natural History Museum (Hymenoptera: Braconidae). Annales Historico-Naturales Musei Nationalis Hungarici 96: 153–223. Papp J (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica. 66: 137–194. Quicke DLJ (2015) The Braconid and Ichneumonid Parasitoid Wasps – Biology, Systematics, Wvolution and Ecology. Wiley Blackwell, 681 pp. Reinhard H (1863) Beiträge zur Kenntniss einiger Braconiden-Gattungen. Berliner Entomologische Zeitschrift 7: 248–274. https://doi.org/10.1002/mmnd.47918630304 Rohwer SA (1934) Descriptions of five parasitic Hymenoptera. Proceedings of the Entomological Society of Washington 36(2): 43–48. Roman A (1912) Die Ichneumonidentypen C.P. Thunbergs. Zoologiska Bidrag fran Uppsala 1: 229–293. Roman A (1917) Braconiden aus den Färöern. Arkiv för Zoologi 11(7): 1–10.

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Tobias VI (1986) Rogadinae: 72–85. In: Medvedev GS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4. Opredeliteli po Faune SSSR 145: 1–501. [In Russian; translation (1995): 512–588. Lebanon, U.S.A.] Viereck HL (1914) Type species of the genera of Ichneumon flies. United States National Museum Bulletin 83: 1–186. https://doi.org/10.5479/si.03629236.83.1 Viereck HL (1922) First supplement to “Type species of the genera of Ichneumon-flies.” Proceedings of the United States National Museum 59: 129–150. https://doi.org/10.5479/ si.00963801.2364.129 Walker F (1860) Characters of some apparently undescribed Ceylon insects. Annals and Magazine of Natural History (3)5: 304–311. Wesmael C (1838) Monographie des Braconides de Belgique 4. Nouveaux Mémoires de l’Academie Royale des Sciences et Belles-lettres de Bruxelles 11: 1–166. Yu DSK, Achterberg C van, Horstmann K (2012) Taxapad 2012, Ichneumonoidea 2011. Database on flash-drive. www.taxapad.com, Ottawa, Ontario, Canada. Zaldívar-Riverón A, Shaw MR, Sáez AG, Mori M, Belokobylskij SA, Shaw SR, Quicke DLJ (2008) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Biology 8: 329.

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Revision of western Palaearctic Aleiodes Wesmael, II.

doi: 10.3897/zookeys.919.39642

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Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis group Cornelis van Achterberg1, Mark R. Shaw2, Donald L.J. Quicke3 1 State Key Laboratory of Rice Biology, Ministry of Agriculture Key Lab of Agricultural Biology of Crop Pathogens and Insects, and Institute of Insect Sciences, Zhejiang University, Hangzhou 310058, China 2 National Museums of Scotland, Chambers Street, Edinburgh EH1 1JF, Scotland, UK 3 Department of Biology, Faculty of Life Sciences, Chulalongkorn University, Bangkok, Thailand Corresponding author: Cornelis van Achterberg ([email protected]) Academic editor: B. Santos  |  Received 2 September 2019  |  Accepted 28 November 2019  |  Published 16 March 2020 http://zoobank.org/0CC5169A-2325-41AD-938F-179FCB056381 Citation: van Achterberg C, Shaw MR, Quicke DLJ (2020) Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis group. ZooKeys 919: 1–259. https://doi.org/10.3897/zookeys.919.39642

Abstract The West Palaearctic species of the Aleiodes apicalis group (Braconidae: Rogadinae) as defined by van Achterberg & Shaw (2016) are revised. Six new species of the genus Aleiodes Wesmael, 1838, are described and illustrated: A. carbonaroides van Achterberg & Shaw, sp. nov., A. coriaceus van Achterberg & Shaw, sp. nov., A. improvisus van Achterberg & Shaw, sp. nov., A. nigrifemur van Achterberg & Shaw, sp. nov., A. turcicus van Achterberg & Shaw, sp. nov., and A. zwakhalsi van Achterberg & Shaw, sp. nov. An illustrated key to 42 species is included. Hyperstemma Shestakov, 1940, is retained as subgenus to accommodate A. chloroticus (Shestakov, 1940) and similar species. Fourteen new synonyms are proposed: Rogas bicolor Lucas, 1849 (not Spinola, 1808), Rogas rufo-ater Wollaston, 1858, Rhogas bicolorinus Fahringer, 1932, Rhogas reticulator var. atripes Costa, 1884, and Rhogas similis Szépligeti, 1903, of Aleiodes apicalis (Brullé, 1832); Rogas (Rogas) vicinus Papp, 1977, of Aleiodes aterrimus (Ratzeburg, 1852); Rogas affinis Herrich-Schäffer, 1838, of Aleiodes cruentus (Nees, 1834); Bracon dimidiatus Spinola, 1808, and Rhogas (Rhogas) dimidiatus var. turkestanicus Telenga, 1941, of Aleiodes gasterator (Jurine, 1807); Rogas alpinus Thomson, 1892, of Aleiodes grassator (Thunberg, 1822); Rhogas jaroslawensis Kokujev, 1898, of Aleiodes periscelis (Reinhard, 1863); Rhogas carbonarius var. giraudi Telenga, 1941, of Aleiodes ruficornis (Herrich-Schäffer, 1838); Ichneumon ductor Thunberg, 1822, of Aleiodes unipunctator (Thunberg, 1822); Rogas heterostigma Stelfox, 1953, of Aleiodes pallidistigmus (Telenga, 1941). Neotypes are designated for Rogas affinis Herrich-Schäffer, 1838; Rogas nobilis Haliday (in Curtis), 1834; Rogas pallidicornis Herrich-Schäffer, 1838; Rogas ruficornis Herrich-Schäffer, 1838. Lectotypes are designated for Rhogas (Rhogas) dimidiatus var. turkestanicus Telenga, 1941, and Rhogas hemipterus Marshall, 1897. Copyright Cornelis van Achterberg et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original authors and source are credited.

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Keywords Aleiodes apicalis group, key, new species, host range, biology, distribution, West Palaearctic, Europe, phenology

Table of content Introduction.............................................................................................................. 3 Specimens, methods, and presentation of records...................................................... 4 Molecular methods............................................................................................... 6 Phylogeny............................................................................................................. 8 Taxonomy............................................................................................................... 12 Aleiodes Wesmael, 1838..................................................................................... 12 Aleiodes apicalis group........................................................................................ 14 Key to West Palaearctic species of the Aleiodes apicalis group............................... 16 Biology and descriptions..................................................................................... 44 Aleiodes aestuosus (Reinhard, 1863).................................................................... 44 Aleiodes agilis (Telenga, 1941)............................................................................ 48 Aleiodes apicalis (Brullé, 1832)........................................................................... 52 Aleiodes arnoldii (Tobias, 1976)......................................................................... 59 Aleiodes aterrimus (Ratzeburg, 1852)................................................................. 63 Aleiodes carbonarius Giraud, 1857..................................................................... 71 Aleiodes carbonaroides van Achterberg & Shaw, sp. nov...................................... 76 Aleiodes caucasicus (Tobias, 1976)...................................................................... 80 Aleiodes coriaceus van Achterberg & Shaw, sp. nov............................................. 84 Aleiodes cruentus (Nees, 1834)........................................................................... 88 Aleiodes desertus (Telenga, 1941)........................................................................ 94 Aleiodes dissector (Nees, 1834)............................................................................ 98 Aleiodes diversus (Szépligeti, 1903)................................................................... 104 Aleiodes eurinus (Telenga, 1941)...................................................................... 109 Aleiodes fahringeri (Telenga, 1941)................................................................... 113 Aleiodes fortipes (Reinhard, 1863).................................................................... 117 Aleiodes gasterator (Jurine, 1807)...................................................................... 123 Aleiodes grassator (Thunberg, 1822)................................................................. 129 Aleiodes hemipterus (Marshall, 1897)............................................................... 134 Aleiodes hirtus (Thomson, 1892)...................................................................... 139 Aleiodes improvisus van Achterberg & Shaw, sp. nov......................................... 144 Aleiodes krulikowskii (Kokujev, 1898).............................................................. 149 Aleiodes miniatus (Herrich-Schäffer, 1838)...................................................... 153 Aleiodes morio (Reinhard, 1863)...................................................................... 157 Aleiodes nigrifemur van Achterberg & Shaw, sp. nov......................................... 162 Aleiodes nobilis (Haliday [in Curtis], 1834)...................................................... 165 Aleiodes pallidicornis (Herrich-Schäffer, 1838)................................................. 171

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Aleiodes pallidistigmus (Telenga, 1941)............................................................. 174 Aleiodes periscelis (Reinhard, 1863).................................................................. 178 Aleiodes pulchripes Wesmael, 1838................................................................... 184 Aleiodes quadrum (Tobias, 1976)..................................................................... 190 Aleiodes ruficeps (Telenga, 1941)...................................................................... 195 Aleiodes ruficornis (Herrich-Schäffer, 1838)...................................................... 199 Aleiodes rufipes (Thomson, 1892)..................................................................... 206 Aleiodes rugulosus (Nees, 1811)........................................................................ 210 Aleiodes schewyrewi (Kokujev, 1898)................................................................ 218 Aleiodes schirjajewi (Kokujev, 1898)................................................................. 221 Aleiodes sibiricus (Kokujev, 1903).................................................................... 225 Aleiodes turcicus van Achterberg & Shaw, sp. nov............................................. 230 Aleiodes unipunctator (Thunberg, 1822)........................................................... 233 Aleiodes venustulus (Kokujev, 1905)................................................................. 239 Aleiodes zwakhalsi van Achterberg & Shaw, sp. nov.......................................... 242 Erratum for Part 1................................................................................................. 246 Acknowledgements................................................................................................ 246 References............................................................................................................. 247 Appendix 1............................................................................................................ 257 Introduction In this 2nd part of a revision of western Palaearctic species of Aleiodes Wesmael we treat the group identified in Part 1 (van Achterberg and Shaw 2016) as the A. apicalis group. It should be noted that our A. apicalis group is constituted in a different (wider) way than of Shaw et al. (1998), Fortier and Shaw (1999), and Garro et al. (2017). The Aleiodes apicalis group as interpreted here is easily recognised from the key given by van Achterberg and Shaw (2016) and includes a majority of species that are rather large for the genus as a whole. From data presented in this paper, some of the commonest species, as well as being rather morphologically isolated, can now be said to be well-understood biologically and appear to be taxon-specialists (see Shaw, 1994, 2017), which suggests to us that they have not been involved in recent speciation events (see Shaw, 2003). However, for others, including a substantial number of rare, or at any rate rarely collected species, there is practically no biological information, with the result that this generalisation cannot be extended: indeed, there are some groups of apparently closely related species that are much more difficult to separate and in these parts of the A. apicalis group speciation has probably been more recent. We deal here with 42 species, a few of which are included only because they have been recorded from the region by others (i.e., the relevant specimens not examined by us) and/or are considered likely to occur in the eastern part of the area. Several of the most seldom-collected species occur as adults early in the year and may not in reality be as rare as they seem.

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Specimens, methods, and presentation of records The biological data from rearings of wild-collected hosts is in some cases supplemented by experimentation, and the protocols and means of scoring results are as outlined in van Achterberg and Shaw (2016). The rather full introductory sections of that paper apply here and are not repeated except when not to do so would leave this paper difficult to use by itself. The term plurivoltine is used to indicate more than one generation in the year (very often this would be only two, but it could be more under favourable circumstances). Overall, many of the species treated here have been widely misinterpreted in the literature and, as in Part 1 of our revision (van Achterberg and Shaw 2016), we have ignored published records when compiling host and distributional data, depending only on specimens we have actually seen ourselves. As previously, we have simply updated the nomenclature of hosts rather than transcribing obsolete names or obvious misspellings from data labels, and unless stated otherwise the reared material cited is in National Museum of Scotland, Edinburgh (NMS). From the host data we are able to give, indicating compact host ranges, in comparison with that expressed in Yu et al. (2016) it should be clear that this was a wise action. By similarly ignoring distribution data (e.g., from Yu et al. (2016)) we do not suggest that published distribution records are necessarily wrong: simply that we are unable to confirm them from the many thousands of specimens we have examined. The sheer number of these prevented us from listing specimen data in full, except for the few species of which we have seen only a very few specimens. The countries we list from the area under consideration (the western Palaearctic) are followed by a list of extralimital countries, in square brackets, from which we have also examined the species in question. All available collections containing recently collected material of Aleiodes from the western Palaearctic region were used for our revision; collections with type material are separately listed under the description of the species. The following collections and acronyms are used: AAC ALC BMNH BZL CC CMIM CNC FC FMNH FRAH HHC HSC IKC KBIN JLC

A.A. Allen Collection, Dawlish, A. Lozan Collection, Institute of Entomology, České Budĕjovice, Natural History Museum, London, Oberösterreichisches Landesmuseum, Biologiezentrum, Linz, M. Čapek Collection, Moravian Museum, Brno, C. Morley Collection, Ipswich Museum, Ipswich, Canadian National Collection of Insects, Ottawa, J.V. Falcó Collection, Valencia, Finnish Museum of Natural History, Helsinki, Forest Research, Alice Holt Lodge, Farnham, H. Haraldseide Collection, H. Schnee Collection, I. Kakko Collection, Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels, J. Lukáš Collection, Bratislava,

Revision of western Palaearctic Aleiodes Wesmael, II.

MCZ MNHN MMUM MRC MSC MSNV MTMA NMI NMS NNHM NRS OUM PAN RMNH SDEI SMNS SYKE UMZC UNS USNM UWIM WAE ZIL ZISP ZJUH ZMB ZMC ZMUO ZMUU ZSSM

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Museum of Comparative Zoology, Harvard University, Cambridge, U.S.A., Muséum national d’Histoire naturelle, Paris, Manchester Museum, University of Manchester, Manchester, M. Riedel Collection, M. Schwarz Collection, Linz, Museo de Storia Naturale, Venice, Hungarian Natural History Museum, Budapest, National Museum of Ireland, Dublin, National Museums of Scotland, Edinburgh, National Natural History Museum, Oslo, Swedish Natural History Museum, Stockholm, Oxford University Museum of Natural History, Oxford, Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw & Łomna-Las, Naturalis Biodiversity Center, Leiden, Senkenberg Deutches Entomologisches Institut, Müncheberg, Staatliches Museum für Naturkunde, Stuttgart, Finnish Environment Institute, Friendship Park Research Centre, Kuhmo, University Museum of Zoology, Cambridge, Department of Biology and Ecology, University of Niš, Serbia, U.S. National Museum of Natural History, Washington D.C., University of Wyoming Insect Museum, Laramie, W.A. Ely Collection, Rotherham, Zoological Institute, Lund University, Lund, Zoological Institute, Academia NAUK, St. Petersburg, Zhejiang University, Hangzhou, Zoologisches Museum, Humboldt Universität, Berlin, Zoological Museum, Copenhagen, Zoological Museum, University of Oulu, Oulu, Zoological Museum of Uppsala University, Uppsala, Zoologische Staatssammlung, München; including E. Haeselbarth Collection.

In addition, we have examined specimens from various smaller and private collections, which are cited in significant cases. Unless otherwise specified, reared material is in NMS. 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 (males have a greater number in other species, which is the normal condition seen in Braconidae). 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. Attention has been paid to the apical tergites of males. The medial dorsal pores of A. fortipes (Reinhard), which are unique to this species within the A. apicalis group

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as treated here, are described and discussed in the entry for that species (note that the unknown male of A. caucasicus (Tobias) is likely to be similar). In the remainder of the species group there is either no evident modification, or a different development is evident to a greater or lesser extent. In some species specialised setae are present on tergites 4–6(7), presumably connected with pheromone dispersal from tergal glands. Broadly, two kinds of specialised setae can occur on these tergites. First, a fringe of short backwards-projecting setae (hereafter “fringe”), possibly associated with pores, originate from close to an apparent sulcus near the extreme base of the tergite (which is normally concealed). The presence and nature of the fringe varies between species, and even when present, it may not be visible in a given specimen owing to telescoping of the tergites. Second, there may be backwards-directed and more or less dense patches of longer setae (hereafter “setal patches”) on each side of the mid-line, the setae to some extent being adpressed in their anterior part but tending to be raised posteriorly (in extreme cases giving the tergites a concave appearance) and appearing different from the arrangement of setae on the more anterior tergites. A median glabrous area is left between the paired setal patches on each tergite, which collectively present as a glabrous and often shiny dorsal stripe along the length of these tergites (hereafter “glabrous stripe”). There is considerable variation in the extent to which these features are developed in the species keyed here, and indeed in some species they are scarcely present or wholly absent. In the species accounts given below we attempt to give a score from 1 to 4 for the development of the setal patches and glabrous stripe in males, with minimal elaboration (but including also mention of the setal fringe in cases for which we have been able to observe it). Type 1 = not at all developed, setae as on anterior tergites and evenly distributed. Type 2 = setal patches hardly developed, but glabrous stripe evident to some extent. Type 3 = setal patches clearly developed but relatively weak or sparse, glabrous stripe strong. Type 4 = setal patches strongly developed, making the tergites appear concave, glabrous stripe also strong. It should be borne in mind that there is some intraspecific variation, much of which may be artefactual (i.e., the condition of the specimen may make it hard to assess and score accurately). For the recognition of braconid subfamilies, see van Achterberg (1990, 1993, 1997), for the identification of Aleiodes Wesmael, see van Achterberg (1991) and Chen and He (1997). For additional references see Yu et al. (2016). For the terminology used in this paper see Figs 1–6 or van Achterberg (1988, 1993; note, however, that in the present work the distance between eye and lateral ocellus is measured differently). An asterisk indicates a new country record according to Yu et al. (2016).

Molecular methods A molecular dataset of the barcode region of cytochrome oxidase c subunit 1 (CO1) was compiled for a total of 141 Aleiodes specimens and three of Heterogamus (Fig. 1) which are the sister group of Aleiodes (Zaldivar-Riveron et al. 2008, Quicke et al. unpublished) and were used for rooting the trees. Most of the recent DNA extractions were carried out using normal procedures for 96-well plates (Ivanova et al. 2006), and PCR

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

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and sequencing reactions were carried out using standard protocols (Hajibabaei et al. 2005). Most sequences were obtained using the LCO-HCO primer pair combination (Folmer et al. 1994: LCO 5’- GGT CAA CAA ATC ATA AAG ATA TTG G-3’, HCO 5’ - TAA ACT TCA GGG TGA CCA AAA AAT CA-3’) or, less often, LepF1-LepR1 (Smith et al. 2005: LepF1 5’-ATT CAA CCA ATC ATA AAG ATA TTG G-3’, LepR1 5’-TAA ACT TCT GGA TGT CCA AAA AAT CA-3’). Sequence alignment was carried out manually and was largely trivial as there was no length variation apart from a three base pair deletion uniting most species of the A. risaae Quicke and Butcher species group as previously noted (Butcher et al. 2012) and its precise location determined by reference to amino acid identities and the known codon positions. Sequences were analysed using maximum likelihood with the programme RAxML (v.8) (Stamatakis 2014), using a GTR + G rate model with three data partitions corresponding to the three codon positions. Each analysis comprised 100 replicates with two threads. Trees were visualised using Figtree (1.4.3) (Rambaut 2016). GenBank accessions numbers are given in Appendix 1. Specimens with an identifier code comprising MRS followed by a number are deposited in NMS, with the exception of the paratype of A. coriaceus (MRS311) which is in RMNH. The specimens indicated CollHH with a number are retained in the personal collection of Håkon Haraldseide (Norway); voucher locations of samples prefixed by BCLDQ are as follows: for Thai specimens depositories are given in Butcher et al. (2012), USA specimens are in University of Wyoming collection; others and that of A. mexicanus (BMNHE897778) are in the collection of the Natural History Museum, London; the voucher of A. cameroniiJanz01 (DHJPAR0021064) is in the collection of Prof. Dan Janzen (Philadelphia); the voucher of A. trianguliscleroma (CCDB27844-E03) is in the collection of Tel Aviv University, Tel Aviv, Israel; the voucher of A. aff. wyomingensis (BIOUG01036-F12) is in the collection of the Center for Biodiversity Genomics, University of Guelph, Canada.

Phylogeny Three datasets were investigated with different levels of taxonomic and sequence inclusion. Firstly, we conducted an overview analysis including representatives of a wide range of extra-limital species groups of Aleiodes, single representatives of the species treated in this paper for which molecular data were available (22 of the 42 species), and representatives of other West Palaearctic species groups, with three members of the genus Heterogamus used as outgroups (Fig. 7). In close agreement with the molecular tree presented for Thai Aleiodes (Butcher et al. 2012: fig. 5 loc. cit.), our results show that most of the A. apicalis species group sensu van Achterberg and Shaw (2016) (= Chelonorhogas auctt.) form a grade together with various generally large bodied extralimital species, notable among which are the Oriental and East Palaearctic A. coronarius group which are characterised by having a deep pronope. Immediately basal to this grade are two large bodied species (A. melanopterus (Erichson) and A. mexicanus Cresson)

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Figure 7. Maximum likelihood tree based on DNA barcode sequence data for representatives of taxa included in this paper (‘Chelonorhogas’ group) together with data from additional West Palaearctic and extra-limital species showing broad picture of relationships. Terminal text show specimen voucher code and provenance (when known).

that had previously been included in a separate genus, Eucystomastax Brues, but which Shaw (1993) showed to be a distinctive monophyletic species group of Aleiodes within which he treated them as a subgenus. The sister to all other Aleiodes as recovered in this analysis are a group of species most members of which have males with metasomal

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Figure 8. Maximum likelihood tree based on DNA barcode sequence data for taxa included in this paper (“Chelonorhogas” group – the dark red grade) together with a larger subset of West Palaearctic species from other species groups.

tergal glands that open at a single medial subposterior pore on one or more of tergites four to six, although they are lacking, for example, in A. miniatus. This clade includes the West Palaearctic A. fortipes (Reinhard), the Nearctic A. cameronii (Dalla Torre) and some other species within the A. pulchripes group sensu Shaw et al. (1997), and the Palaeotropical subgenus Hemigyroneuron Baker (Shaw et al. 1997, Delfin and Wharton 2000, Butcher and Quicke 2011). From within this large basal grade emerges, on a relatively long branch, a monophyletic group which includes the vast majority of Aleiodes species. The West Palaearctic and Nearctic members of this clade have been placed in various species groups including the A. bicolor, A. circumscriptus and A. gastritor complexes. However, many species even within the Palaearctic fauna fall outside of these as isolated groups, often more closely related to extralimital taxa. Secondly, we analysed a matrix comprising the most complete available sequence for each West Palaearctic species and using A. fortipes as the outgroup based on the results of the first analysis (Fig. 8). The A. circumscriptus group in the sense of van Achterberg and Shaw (2016) (including the A. similis and A. gastritor subgroups and the rather isolated A. circumscriptus (Nees) itself, as well as some other species) and the A. bicolor group sensu van Achterberg and Shaw (2016) were recovered together as a monophyletic clade but without strong indication of its comprising two separate groups. Indeed, the previous concept of the A. circumscriptus group was challenged by its paraphyly with respect to the A. bicolor group. The clade comprising the A. bicolor

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Figure 9. Maximum likelihood tree based on all DNA barcode sequence data for taxa included in this paper (“Chelonorhogas” group).

group and the A. similis subgroup (and a few extraneous species including A. circumscriptus) was not treated as a unit by van Achterberg and Shaw (2016) because of rather clear apparent differences: the clade indicated both in that work and here as the A. bicolor group comprises rather stocky and heavily sculptured species that have various morphological features (such as a long malar space and margined T4) in common and they, and several additional similar species, will be dealt with in Part 3 of this work. The species indicated as the A. similis subgroup and the A. gastritor subgroup will be treated together in Part 4. In practice, these two subgroups are less easy to separate on morphological grounds, and again there are many additional species. Throughout most of this large clade radiation seems to have been relatively recent and/or rapid with many

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morphologically and biologically clearly distinguishable species often having CO1 sequences that differ from one another by less than 1 %. The remainder of the species outside the A. apicalis grade are rather well characterised and indeed isolated, with little evidence of recent radiation, and the species for which host relations are known are taxon-specialists (cf. Shaw 2003). All of these species (with the exception of spM3) have already been treated in Part 1 of this work (van Achterberg and Shaw 2016). Thirdly, we constructed a tree for the available barcodes for the species treated in this paper (Fig. 9). Each species represented by multiple sequences is recovered as a monophyletic cluster, mostly with relatively little intraspecific variation. Some of the differences observed are likely due to particular sequences being quite short compared to the full-length barcode, others no doubt due to reading errors particularly for those samples that were sequenced more than ten years ago with different methodologies. Re-examination of existing electropherograms has usually confirmed the generality of this and such reading errors are typically at the 5’ or 3’ ends of sequence reads. The most notable exception is provided by the three specimens of A. hirtus (Thompson), with the sequence from the British specimen differing at 15 positions (2.5 %) along the 606-base region of overlap with the two Romanian individuals (which were identical). These specimens are briefly discussed in the species entry for A. hirtus.

Taxonomy Aleiodes Wesmael, 1838 Figs 10–840 Aleiodes Wesmael, 1838: 194; Shenefelt 1975: 1163–1185; Marsh 1979: 177–178; Papp 1985a: 143–164 & 1985b: 347–349; Shaw and Huddleston 1991: 95–96 (biology); van Achterberg 1991: 24; Zaldivar-Riverón et al. 2004: 225 & 2008: 329 (phylogeny); van Achterberg and Shaw 2016: 8–11 (groups). Type species (designated by Viereck, 1914): Aleiodes heterogaster Wesmael, 1838 [examined; = A. albitibia (Herrich-Schäffer, 1838)]. Petalodes Wesmael, 1838: 123; Tobias 1971: 218 (transl. 1975: 86–87); Shenefelt 1975: 1209–1211; Tobias 1976: 90; Marsh 1979: 179; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838); van Achterberg and Shaw 2016: 8. Type species (by monotypy): Petalodes unicolor Wesmael, 1838 [examined; = Aleiodes compressor (Herrich-Schäffer, 1838)]. Schizoides Wesmael, 1838: 94. Unavailable name. Nebartha Walker, 1860: 310; Shenefelt 1975: 1216; Marsh 1979: 179; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Nebartha macropodides Walker, 1860 [examined]. Tetrasphaeropyx Ashmead, 1889: 634; Shenefelt 1975: 1260; Marsh 1979: 179; Fortier and Sherman 2008: 445 (as subgenus of Aleiodes Wesmael, 1838); Zaldivar-

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Riverón et al. 2008: 329 (as synonym of subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Rogas pilosus Cresson, 1872 [examined]. Neorhogas Szépligeti, 1906: 605; Shenefelt 1975: 1205; van Achterberg 1991: 24 (as subgenus of Aleiodes Wesmael, 1838); Zaldivar-Riverón et al. 2008: 329 (included in subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Neorhogas luteus Szépligeti, 1906 [examined; = Aleiodes praetor (Reinhard, 1863)]. Chelonorhogas Enderlein, [Sept. 1st] 1912a: 258; Shenefelt 1975: 1187; van Achterberg 1991: 24 (as subgenus of Aleiodes Wesmael, 1838); Zaldivar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838). Type species (by monotypy): Chelonorhogas rufithorax Enderlein, 1912 [examined; not Aleiodes rufithorax (Cameron, 1911) = A. convexus van Achterberg, 1991]. Eucystomastax Brues, [(end of?) Sept.] 1912: 223; Shaw 1993: 5 (as subgenus of Aleiodes Wesmael, 1838); Zaldivar-Riverón et al. 2004: 225 (included in Aleiodes Wesmael, 1838); Shimbori & Penteado-Dias 2011: 17 (as subgenus of Aleiodes Wesmael, 1838). Type species (by monotypy): Eucystomastax bicolor Brues, 1912 (= Rogas melanopterus Erichson, 1848). Leluthinus Enderlein, 1912b: 96; Shenefelt 1975: 1202–1203; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Leluthinus lividus Enderlein, 1912 [examined]. Aleirhogas Baker, 1917b: 383, 411; Shenefelt 1975: 1185–1186; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (designated by Viereck, 1921): Rhogas (Aleirhogas) schultzei Baker, 1917 [examined]. Hemigyroneuron Baker, 1917a: 284, 322–327; Zaldivar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838); Butcher and Quicke 2011: 1405 (as subgenus of Aleiodes Wesmael, 1838, and Hemigyroneuron sensu Zaldivar-Riverón et al. (2008) is not Hemigyroneuron)); Butcher and Quicke 2015: 275–279. Type species (original designation): Hemigyroneuron speciosus Baker, 1917 [examined]. Heterogamoides Fullaway, 1919: 43; Shenefelt 1975: 1188; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Heterogamoides muirii Fullaway, 1919 [examined]. Cordylorhogas Enderlein, 1920: 153; Shenefelt 1975: 1195; van Achterberg 1991: 31; Zaldivar-Riverón et al. 2004: 232 & 2008: 329 (as synonym of subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Cordylorhogas trifasciatus Enderlein, 1920 [examined]. Hyperstemma Shestakov, 1940: 10; Shenefelt 1975: 1200; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Hyperstemma chlorotica Shestakov, 1940 [examined]. Dimorphomastax Shenefelt, 1979: 131–133; Shaw et al. 1998: 66 (as synonym of Aleiodes Wesmael, 1838). Type species (by original designation): Dimorphomastax peculiaris Shenefelt, 1979 [examined; = Aleiodes atriceps Cresson, 1869]. Pholichora van Achterberg, 1991: 48–53; Quicke and Shaw 2005: 532; Zaldivar-Riverón et al. 2008: 329 (as synonym of Aleiodes Wesmael, 1838); Butcher and Quicke 2011: 1405 (as synonym of subgenus Hemigyroneuron Baker, 1917); Butcher et al.

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2012: 9 (id.). Type species (original designation): Hemigyroneuron madagascariensis Granger, 1949 [examined]. Arcaleiodes Chen & He, 1997: 60–62; Zaldivar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838); Butcher et al. 2012: 18–19 (id.). Type species (original designation): Aleiodes unifasciatus Chen & He, 1991 [examined]. Vietorogas Long & van Achterberg, 2008: 313–314; Butcher et al., 2012: 15–17 (as synonym of Aleiodes Wesmael, 1838). Type species (original designation): Vietorogas bachma Long, 2008 [examined]. R(h)ogas auct; Tobias, 1971: 215–217 (transl. 1975: 83–86); Shenefelt, 1975: 1215– 1256; Tobias, 1976: 81–89; Marsh, 1979: 179–181; Tobias, 1986: 74–84. Notes. Hyperstemma Shestakov, 1940, is traditionally included in the genus Heterogamus Wesmael, 1838 (Shenefelt 1975) or in the subgenus Heterogamus of the genus Aleiodes Wesmael (e.g., Belokobylskij 2000), but differs by the shape of the head (Figs 17–19) and of the tarsal claws (Fig. 20), the position of the clypeus (Fig. 17), the elongate 2nd submarginal cell of the fore wing (but folded in Fig. 10), the widened 1st subdiscal cell of the fore wing and distinctly widened marginal cell of the hind wing (Fig. 10). Therefore, we retain the subgenus Hyperstemma Shestakov of Aleiodes Wesmael for at least the following species: A. chloroticus (Shestakov, 1940) from China (Palaearctic and Oriental), *Japan (RMNH), Korea, and Russia (Far East), A. albigenus Chen & He, 1997, from China (Oriental) and Vietnam, A. crassinervis Chen & He, 1997, from China (Oriental) and Vietnam, A. naevius Chen & He, 1997, from China (Oriental), and A. pallidinervis (Cameron, 1910) from China (Palaearctic and Oriental), Japan, Korea, and Russia (Far East). Aleiodes apicalis group Diagnosis. Apical half of marginal cell of hind wing distinctly widened, its maximum width 1.6 × its width near hamuli or wider (Fig. 27) and vein r of fore wing shorter than vein 3-SR (Figs 180, 608), if marginal cell largely parallel-sided (Figs 506, 609, 704) then tarsal claws comparatively robust and with often blackish pecten (Figs 517, 621, 716) or brachypterous (Fig. 390); occipital carina usually reduced ventrally, not reaching hypostomal carina (Figs 600, 663, 713, 788); mesopleuron partly smooth (at least between punctures), but largely densely sculptured in both sexes of A. hemipterus and A. krulikowskii, as well in some males of A. ruficornis and allies; lateral carina of scutellum absent or if present then weakly developed and lunula wide (Fig. 508); 2nd metasomal tergite with distinct and smooth triangular area medio-basally (Fig. 509); ovipositor sheath distinctly setose apically (Fig. 483); males are often darker than females, most extremely so in A. arnoldii, A. carbonarius and A. carbonaroides; brachypterous specimens of Aleiodes are included in this group. Biology. All species of the A. apicalis group for which host data exist are parasitoids of Noctuidae. However, the putatively more basal A. fortipes belonging to the Hemi-

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Figures 10–21. Aleiodes chloroticus (Shestakov), ♂, Japan 10 wings 11 mesosoma lateral 12 mesosoma dorsal 13 1st–3rd metasomal tergites dorsal 14 fore femur lateral 15 hind femur lateral 16 base of antenna 17 head anterior 18 head dorsal 19 head lateral 20 outer hind tarsal claw 21 apex of antenna.

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gyroneuron clade (see below) is a parasitoid of Geometridae. Also, only A. fortipes and A. sibiricus are known to parasitise hosts only in spring although these hosts would have been available in autumn of the previous year. Possibly others in the A. apicalis group will be found to do this too, and we consider the habit putatively as ancestral, in contrast with the more derived A. circumscriptus and A. bicolor groups in which species using hosts that overwinter as larvae invariably (as far as known) parasitise the host in the autumn and overwinter as a young larva inside it. While we have no host data for a disappointingly large number of species of the A. apicalis group, the form of the clypeus may give important clues as to the site at which host mummification occurs, as those species in which mummification is known to take place in open situations (e.g., on a twig or in a leaf curl) invariably have a relatively small hypoclypeal depression and the clypeal margin blunt (A. apicalis, A. aterrimus, A. fortipes, A. nobilis, A. pulchripes, A. rugulosus) while species known to cause their hosts to mummify in concealed situations tend to have the hypoclypeal opening wider and the margin sharper (e.g., A. cruentus, A. dissector, A. ruficornis, A. sibiricus, A. unipunctator). Notes. According to the 28S + COI analysis by Zaldivar-Riverón et al. (2008) the following former subgenera or genera belong to this group: Chelonorhogas Enderlein, [1st Sept.] 1912 (worldwide), Eucystomastax Brues, [(end of?) Sept.] 1912 (Neotropical group with 2nd and 3rd maxillary palp segments enlarged), Hemigyroneuron Baker, 1917 (Old World group with distal half of subbasal cell of fore wing modified and glabrous), and Dimorphomastax Shenefelt, 1979 (males of this monotypic Neotropical group have a large curved tooth near the base of the mandible (an outgrowth of the condylar carina) and the hind tibial spurs are blunt apically; females have the tooth smaller and triangular, and the hind tibial spurs are acute). Butcher et al. (2012) indicate in their cladogram based on the analysis of COI sequences that A. fortipes (Reinhard) forms together with Hemigyroneuron Baker and Arcaleiodes Chen & He the most basal clade of Aleiodes Wesmael, and it is noteworthy that all known hosts of this clade are Geometridae (see species entry for A. fortipes). According to the same analysis the A. apicalis group consists of three clades: (i) the Hemigyroneuron clade (see above; likely also includes A. caucasicus (Tobias)), (ii) the A. rugulosus clade (including the Asian group with modified pronotum), and (iii) the A. gasterator clade. Since we do not have the COI sequences of all species, we unite these three clades in the A. apicalis group to allow identification based on their morphology.

Key to West Palaearctic species of the Aleiodes apicalis group 1

Basal half of fore wing (except anteriorly) largely glabrous (a), or rather inconspicuously setose as remainder of wing; width of hypoclypeal depression 0.8–1.0 × minimum width of face (b) and anterior part of clypeus very narrow (c); vein r of fore wing 0.5–0.7 × vein 3-SR (d); mandibles massive triangular and coarsely punctate (e); [mandible with thick ventral lamella; antennal segments 47–63 and 4th segment of ♀ 1.1–1.3 × longer than wide; head

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(except clypeus and mandible) and mesosoma (except partly prothorax and mesoscutum) black; tarsal claws slender and only setose].............................2

–

Basal half of fore wing normally setose (except sometimes near veins) as remainder of wing (aa) or brachypterous (♀ A. hemipterus), if rarely with reduced setosity (A. venustulus) then width of hypoclypeal depression less than 0.7 × width of face (bb) and/or anterior part of clypeus moderately wide (cc), or vein r of fore wing 0.2–0.4 × vein 3-SR (dd); shape of mandible variable, often less massive and largely smooth (ee)....................................................3

2

Lateral lobes of mesoscutum whitish setose and with satin sheen (a); flagellum of antenna somewhat darker than scapus and pedicellus (b); middle lobe of mesoscutum distinctly punctate (c; more or less obscured by setosity); height of eye approx. 6 × length of malar space (d); basal half of metasoma dark brown, but laterally more or less yellowish (e)......... A. agilis (Telenga, 1941)

–

Lateral lobes of mesoscutum largely glabrous or sparsely setose and strongly shiny (aa); flagellum of antenna brownish yellow, similar to colour of scapus and pedicellus (bb); middle lobe of mesoscutum largely smooth (cc); height of eye

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nearly 7 × length of malar space (dd); basal half of metasoma brownish yellow, at most 1st tergite darker brown medio-basally (ee)......A. desertus (Telenga, 1941)

3

Mesoscutum densely rugose or rugulose (a), with medio-longitudinal ridge or carina (b); mesopleuron mainly rugose (c); 3rd metasomal tergite densely sculptured (d) and convex posteriorly (e); propodeum angulate posteriorly (f ); ♀ brachypterous and ♂ macropterous; N Africa..................................... ...................................................................A. hemipterus (Marshall, 1897)

–

Mesoscutum largely smooth and punctate or punctulate, mainly granulate or coriaceous (aa), usually without medio-longitudinal ridge or carina (bb); mesopleuron at most medially and antero-dorsally rugose (cc); 3rd metasomal tergite truncate posteriorly or nearly so (dd) and/or largely smooth posteriorly (ee); if mesopleuron largely sculptured (ccc) combined with 3rd tergite convex (ddd) and densely sculptured (eee) posteriorly, then propodeum rounded posteriorly (ff); both sexes macropterous......................................................4

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4

Anterior part of clypeus short and subparallel-sided, near lower level of eyes (a) and hind femur slender (b); antenna with 65–72 segments and 5th–10th segments approx. as wide as long (c); tarsal claws slender (d) and without distinct pecten (e); marginal cell of hind wing slightly constricted subbasally (f) or subparallelsided (fff below); [temple behind eye densely setose, convex and curved in dorsal view; 2nd–5th metasomal tergites more or less yellowish to reddish brown and head except mouthparts black]..............................A. sibiricus (Kokujev, 1903)

–

Anterior part of clypeus medially distinctly wider than laterally (aa); if intermediate then partly above lower level of eyes or hind femur inflated (bb); antennal segments usually 62 or less, 4th–10th segments variable, often longer than wide (cc); tarsal claws often rather robust (dd), if slender (ddd) then either with distinct pecten (ee) and/or marginal cell of hind wing directly widened subbasally (ff).................................................................................5

5

Temples extremely short (a), approx. 0.2 × as long as eye in dorsal view; basal half of marginal cell of hind wing parallel-sided (b) and pterostigma pale-yellowish or light brown (c); malar space slightly shorter than basal width of mandible (d); hind tibial spurs of ♂ obtuse apically (e); [OOL distinctly less than diameter of posterior ocellus; tarsal claws with coarse pecten; mesopleuron, mesosternum and scutellum brownish yellow]........................... A. pulchripes Wesmael, 1838

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–

Temples medium-sized to long (aa), at least 0.3 × as long as eye in dorsal view; basal half of marginal cell of hind wing gradually widened (bb), if parallelsided (bbb) then pterostigma dark brown (cc); malar space usually as long as basal width of mandible (dd) or longer; hind tibial spurs of ♂ usually acute apically (ee)..................................................................................................6

6

Vein 2-SR+M of fore wing 0.8–1.0 × vein m-cu (a); pronotum and meso­ scutum similarly coloured; clypeus width 0.3 × minimum width of face (b); length of fore wing 3.7–5.0 mm; length of hind femur 3.5–3.9 × its maximum width (c) and occipital carina reduced or anteriorly angled medio-dorsally (d); [4th–6th metasomal tergites of ♂ with setose round pits (but ♂ of A. caucasicus unknown); vein m-cu of fore wing more or less subvertical and relatively short; 3rd–10th antennal segments of ♀ pale yellowish, contrasting with entirely dark brown scapus].................................................................7

–

Vein 2-SR+M of fore wing 0.2–0.6 × vein m-cu (aa), if more (some A. nobilis) then pronotum orange in contrast with blackish mesopleuron; clypeus width 0.4–0.8 × minimum width of face (bb); fore wing almost always longer than 4.9 mm; length of hind femur either more than 3.9 × its maximum width (cc) or occipital carina complete medio-dorsally (dd)..........................................8

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7

Posterior half of mesosoma largely black or dark brown (a); precoxal area largely smooth, at most with some aciculae or punctures medially (b); tegulae brown (c); antero-dorsally mesopleuron coarsely rugose (d); [body of ♂ completely black and antenna completely blackish, dark brown or with some segments yellowish subbasally]; N & C Europe................. A. fortipes (Reinhard, 1863)

–

Posterior half of mesosoma largely yellowish brown (aa); precoxal area more or less vertically striate (bb); tegulae usually yellow (cc), but sometimes dark brown; mesopleuron antero-dorsally moderately rugose (dd); SE Europe [♂ unknown]........................................................ A. caucasicus (Tobias, 1976)

8

Tarsal claws gradually narrowed submedially, slender and hardly curved (a) and 4th hind tarsal segment brownish yellow and 1.8–2.0 × as long as wide (c); clypeus yellowish brown, distinctly protruding anteriorly and ventrally thick (b); tarsal segments ventrally with long apical spiny bristles (d); [4th antennal segment of ♀ distinctly longer than wide; basal half of antenna and mesosoma anteriorly of ♀ largely yellowish brown, in ♂ more or less dark brown or infuscated; clypeus of ♂ yellowish and contrasting with black face]............................................................. A. schewyrewi (Kokujev, 1898)

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–

Tarsal claws more directly narrowed submedially, moderately robust and apically curved (aa); if slender (ccc) and hardly curved (aaa) then clypeus black, with thin ventral margin (bb) or 4th hind tarsal segment dark brown or infuscate and at most 1.5 × longer than its maximum width (cc) and tarsal segments ventrally with shorter apical bristles (dd)...........................................9

9

Vein 1-CU1 of fore wing 0.7–1.5 × as long as vein m-cu (a); if 0.7–0.9 × (A. aestuosus, A. zwakhalsi) then base of hind tibia yellowish dorsally (b), hind trochanter orange or yellowish (c) and pecten up to apical tooth of tarsal claw (d)...................................................................................................10

–

Vein 1-CU1 of fore wing 0.3–0.8 × as long as vein m-cu (aa); if 0.7–0.8 × then base of hind tibia with dark brown patch dorsally (bb) or hind trochan­ ter dark brown (cc) and in both cases pecten remaining removed from apical tooth of tarsal claw (dd).............................................................................19

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23

10

Pronotum orange (except antero-medially), distinctly contrasting with black posterior half of mesosoma in lateral view (a, rarely black) and 3rd–6th antennal segments of ♀ pale yellowish, contrasting with dorsally entirely dark brown scapus (b) and angle of vein m-cu of fore wing with vein 3-CU1 distinctly larger than 90° (c) and apex of hind femur black or dark brown (e); palpi yellow (d); [head black; hind basitarsus brownish yellow, contrasting with dark brown telotarsus].......................................................................11

–

Pronotum black or reddish brown and less contrasting with posterior half of mesosoma in lateral view (aa); if pronotum orange brown and contrasting with dark posterior parts (A. venustulus) then 3rd–6th antennal segments of ♀ and scapus similarly dark brown (bb), angle of vein m-cu of fore wing with vein 3-CU1 closer to 90° (cc) and palpi dark brown (dd) or apex of hind femur yellowish or reddish brown (ee).......................................................12

11

Mesoscutum and scutellum black (a); temple rather mat and mainly granulate between punctulation (b); frons mat and strongly granulate (c); base of hind tibia pale yellowish (d); [precoxal area usually with some very superficial short rugulae or crenulae medially]......................A. nobilis (Haliday, 1834)

–

Mesoscutum and scutellum orange brown (aa); temple shiny and smooth between punctures, striae or rugae (bb); frons shiny and with distinct striae

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or rugae (cc); base of hind tibia more or less infuscate (dd)............................ .................................................................... A. schirjajewi (Kokujev, 1898)

12

Tarsal claws without pecten near apical tooth (a); vein m-cu of hind wing absent (b); wing membrane subhyaline to slightly infuscate (c); [5th–10th antennal segments of ♀ distinctly longer than wide]......................................13

–

Tarsal claws with pecten near apical tooth (aa); vein m-cu of hind wing at least weakly present (bb); wing membrane moderately infuscate or brownish (cc)..................................................................................... 14

13

Head of ♀ entirely yellowish brown or orange (a); ventral margin of clypeus thick and not protruding (b); vertex and OOL with smooth interspaces between punctures (c); mesopleuron remotely punctate and precoxal area coarsely punctate (d); 3rd tergite nearly flat in lateral view (e) and medioposteriorly nearly truncate in dorsal view (f ); [fore femur elongate and hind femur 4.3–4.7 × longer than wide; vertex at least partly densely punctate]..... ..................................................................... A. venustulus (Kokujev, 1905)

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–

Head of ♀ largely black (aa); ventral margin of clypeus thin and protruding anteriorly (bb); vertex and OOL without distinct smooth interspaces, rugose (cc); mesopleuron very densely and coarsely punctate and precoxal area rugose-punctate (dd); 3rd tergite convex in lateral view (ee) and medioposteriorly convex in dorsal view (f ); [3rd tergite coarsely punctate; fore and middle femora with dark patch].................A. krulikowskii (Kokujev, 1898)

14

Head of ♀ entirely yellowish brown or orange (a); ventral margin of clypeus thin and protruding anteriorly (b); eye 0.8–1.2 × temple in dorsal view (c); apical third of metasoma of ♀ completely yellowish (d; but ♂ often with 1st tergite partly and 4th–6th tergites blackish); hind femur of ♀ distinctly inflated (e), but sometimes less so; [antenna of ♀ with 49–56 segments; hind tibia of ♀ ivory except dark brown apex].................. A. aestuosus (Reinhard, 1863)

–

Head of ♀ black (aa); ventral margin of clypeus thick and hardly protruding anteriorly (bb); eye 1.0–1.9 × temple in dorsal view (cc); apical third of metasoma of ♀ black (dd); hind femur of ♀ slender to moderately wide (ee)....15

15

Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (a); 5th–10th antennal segments of ♀ distinctly longer than wide (b); posterior half of mesoscutum black (c); 1st metasomal tergite robust (d); [surroundings of veins M+CU1 and 1-+2-CU1 largely setose; vein M+CU of

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hind wing distinctly longer than vein 1-M; apical fifth of hind femur always blackish; 4th–6th metasomal tergites of ♂ appearing concave and with conspicuous setosity]; C Europe, Mediterranean area, Central Asia. Examined specimens from S England (BMNH), C Netherlands (RMNH) and S Sweden (NMS) are almost certainly passive migrants and do not represent breeding populations..................................................... A. apicalis (Brullé, 1832)

–

Maximum width of hypoclypeal depression 0.5–0.7 × minimum width of face (aa); 5th–10th antennal segments of ♀ approx. as long as wide (bb); posterior half of mesoscutum at least partly red (cc); 1st metasomal tergite rather slender (dd); [surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein r of fore wing 0.3–0.4 × vein 3-SR]....................................................16

16

Female: 2nd metasomal tergite of ♀ as long as wide basally (a; of ♂ 0.9 ×); 1st tergite only slightly widened posteriorly and 1.3–1.4 × as long as wide posteriorly (b; of ♂ 1.2 ×); 3rd tergite largely smooth basally, only sparsely punctulate (c; of ♂ rugose); OOL distinctly rugose anteriorly (d); [hind femur ca 4.5 × longer than wide; ♂ may be easily confused with A. cruentus]....A. quadrum (Tobias, 1976)

–

Both sexes: 2nd tergite of ♀ 0.7–0.9 × as long as wide basally (aa); 1st tergite distinctly widened posteriorly and 1.0–1.1 × as long as wide posteriorly (bb); 3rd tergite distinctly punctate or punctate-rugulose medio-basally (cc); OOL

Revision of western Palaearctic Aleiodes Wesmael, II.

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usually densely and coarsely punctate anteriorly (dd), rarely striate or rugose (ddd), but less sculptured in males and in A. zwakhalsi..............................17

17

Ocelli medium-sized to large (a), OOL of ♀ 0.5–0.8 × diameter of posterior ocellus, rarely up to 1.0 times; length of eye 1.5–1.9 × temple in dorsal view (b); 1st and 2nd metasomal tergites at least partly reddish or orange brown (c); [hypoclypeal depression usually 0.6–0.7 × width of face; hind femur 3.1–4.0 × longer than wide. If hind femur is 5 × longer than wide and hypoclypeal depression 0.5 × width of face, cf. A. parvicauda (Tobias, 1985) from Afghanistan].............................................................. A. cruentus (Nees, 1834)

–

Ocelli smaller (aa), OOL of ♀ 0.9–1.2 × diameter of posterior ocellus; length of eye 1.0–1.3 × temple in dorsal view (bb); 1st and 2nd metasomal tergites entirely black or dark brown (cc)...............................................................18

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18

Vein 1-CU1 of fore wing distinctly shorter than vein m-cu (a); hind femur 4.0–4.2 × longer than wide (b); vein cu-a inclivous and parallel with vein 3-CU1 (c); 5th–10th antennal segments of ♀ as long as wide (d); vertex and OOL remotely punctate (e); width of hypoclypeal depression 0.7 × minimum width of face (f ); [1st metasomal tergite slender and rounded latero-basally; 3rd tergite densely punctulate basally; metasoma of ♀ strongly compressed posteriorly; if body completely black, precoxal sulcus extensively rugose, pterostigma medially pale brown and OOL densely rugulose, cf. A. morio (Reinhard)].......................................................................... A. zwakhalsi sp. nov.

–

Vein 1-CU1 of fore wing approx. as long as vein m-cu (aa); hind femur 3.0– 3.3 × longer than wide (bb); vein cu-a vertical and vein 3-CU1 diverging posteriorly (cc); 5th–10th antennal segments of ♀ shorter than wide (dd); vertex and OOL moderately to densely punctate (ee); width of hypoclypeal depression 0.5–0.7 × minimum width of face (ff); [metasoma of ♀ less compressed posteriorly; ovipositor sheath rather robust]....A. diversus (Szépligeti, 1903)

19

Third metasomal tergite largely coarsely punctate and yellowish brown (a); 2nd submarginal cell of fore wing short and square (b); medio-longitudinal carina at least in middle part of propodeum absent, obsolescent or incomplete (c); eye much narrower than temple in lateral view (d) and 4th–10th antennal segments of ♀ distinctly longer than wide (e); [clypeus distinctly protruding in lateral view and ventrally thin (Fig. 478); vein 1r-m of hind wing much longer than vein 1-M; OOL twice as long as diameter of posterior ocellus; antennal segments of ♀ 64–70; 2nd tergite coarsely reticulate-punctate; 4th–6th metasomal tergites of ♂ flat and with longer (than of basal tergites) backwards directed setae with a narrow glabrous stripe centrally].............A. miniatus (Herrich-Schäffer, 1838)

Revision of western Palaearctic Aleiodes Wesmael, II.

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–

Third tergite rugose, striate, rugulose or smooth, if punctate then black (aa); 2nd submarginal cell longer than high (bb); medio-longitudinal carina of posterior half of propodeum complete or nearly so (cc); eye usually approx. as wide as temple in lateral view (dd), if distinctly narrower (ddd) then 4th–10th antennal segments of ♀ approx.as long as wide (ee)...................................20

20

Pecten of hind tarsal claws of ♀ robust (a), close to apical tooth (b) and often dark brown or blackish (c); [pecten of ♂ sometimes less developed than in ♀ (e.g., of A. periscelis) but then with some robust teeth medially (aaa)]........21

–

Pecten of hind tarsal claws absent or inconspicuous (aa), if present then remaining removed from apical tooth (bb) and often yellowish or brownish (cc), but dark brown in A. hirtus................................................................26

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21

Ventral margin of [anterior part of ] clypeus comparatively sharp (a), clypeus more or less protruding anteriorly (b); palpi yellowish (c); vein 1-M of fore wing dark brown (d); basal half of metasoma weakly sculptured (e); hind femur largely or completely reddish or brownish (f ); width of hypoclypeal depression 0.6–0.7 × minimum width of face (g); [precoxal area completely smooth or nearly so; length of malar space 0.2 × length of eye in lateral view; outer side of posterior ocellus with deep groove; vertex flattened. If palpi black or dark brown, temple with long setae, width of hypoclypeal depression 0.5 × minimum width of face, OOL more than diameter of posterior ocellus, 1st tergite coarsely sculptured, and length of malar space 0.40–0.45 × length of eye in lateral view, cf. A. hirtus (Thomson)]........ A. dissector (Nees, 1834)

–

Ventral margin of clypeus (rather) obtuse apically (aa) and clypeus hardly protruding anteriorly (bb); palpi dark brown at least basally (cc) or vein 1-M of fore wing yellowish brown (dd; A. rugulosus); basal half of metasoma distinctly sculptured (ee); hind femur dark brown or black dorso-apically (ff; but yellowish in A. rugulosus); width of hypoclypeal depression usually 0.3–0.4 × minimum width of face (gg)......................................................................22

22

Only apical two fifths of marginal cell of hind wing distinctly widened and remainder parallel-sided or nearly so (a), rarely hardly widened apically; vertex flattened behind ocelli (b) and apex of hind femur yellowish or reddish (c); first metasomal tergite with coarse sublongitudinal rugae (d); ovipositor sheath distinctly narrowed apically (e); [vein 1-M of fore wing yellowish brown; basal half of hind tibia pale yellowish/ivory or orange and its apical half black; mesopleuron nearly or completely smooth medio-ventrally; propodeum with pair of crest-like protuberances laterally]............................ A. rugulosus (Nees, 1811)

Revision of western Palaearctic Aleiodes Wesmael, II.

31

–

At least apical half of marginal cell of hind wing gradually widened (aa); if less distinctly so, then vertex declivous behind ocelli (bb) and hind femur black or dark brown apically (cc); 1st tergite moderately striate, rugulose or vermiculaterugose (dd); ovipositor sheath truncate apically or nearly so (ee)...................23

23

Second metasomal tergite of both sexes black (a); scutellum densely and finely coriaceous (b); hind tibia largely black (c), dorsally paler at extreme base; 3rd metasomal tergite of ♀ mainly punctate (d), but basal half more or less rugose in ♂; mesoscutum with satin sheen (e); [vein 2-SC+R of hind wing subquadrate or vertical; 4th–6th tergites of ♂ with medium-sized dense setosity and with narrow glabrous central stripe; mesosternal sulcus shallow, obsolescent or absent. If hind tibia completely dark brown basally and temple roundly narrowed in dorsal view, cf. A. sapporensis (Watanabe) from East Palaearctic region]........................................................A. aterrimus (Ratzeburg, 1852)

–

Second tergite of ♀ yellowish or dark reddish brown (aa; up to almost black in ♂ of A. periscelis); scutellum partly smooth and punctate (bb); basal half of hind tibia (largely) pale yellowish or ivory (cc), rarely brownish; 3rd tergite largely rugulose-striate basally (dd); mesoscutum rather shiny (ee).............24

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24

Antennal segments of ♀ 39–45 (of ♂ 50–56) and subbasal segments of ♀ yellowish (a; of ♂ darkened but basal half of hind tibia ivory); fore femur of ♀ more robust (b); antenna of ♀ robust (c), 0.8–1.0 × longer than fore wing (= 0.7–0.8 × body length); fore coxa dark brown (d); vertex of ♀ coarsely rugose laterally (e); [mandible blackish basally; fore femur 4.8 × as long as wide]............................................................. A. periscelis (Reinhard, 1863)

–

Antennal segments of both sexes 52–62 and subbasal segments dark brown or blackish (aa); basal half of hind tibia of ♂ reddish to dark brown; fore femur of ♀ slenderer (bb); antenna of ♀ elongate (cc), 1.0–1.1 × longer than fore wing; fore coxa (brownish) orange (dd); vertex punctate-rugulose to coriaceous laterally (ee)......................................................................................25

25

Mesoscutum largely matt (a); base of fore femur, fore trochanter and trochantellus at least partly dark brown or infuscate (b); 2nd and 3rd metasomal tergites comparatively slender (maximum width of 2nd tergite ca 1.5 × its median length; c); OOL of ♀ 0.9–1.1 × longer posterior ocellus (d); posterior half of hind femur blackish dorsally (e).................................... A. coriaceus sp. nov.

Revision of western Palaearctic Aleiodes Wesmael, II.

33

–

Mesoscutum rather shiny (aa); base of fore femur, fore trochanter and trochantellus yellowish brown (bb); 2nd and 3rd metasomal tergites robust (maximum width of 2nd tergite ca 1.6 × its median length; cc); OOL of ♀ 1.1–1.5 × longer posterior ocellus (dd); posterior half of hind femur partly yellowish brown dorsally (ee)............................................ A. rufipes (Thomson, 1892)

26

Length of malar space of ♀ 0.45–0.70 × height of eye (a) and clypeus below lower level of eye in lateral view (b), if intermediate (in A. ruficornis) then basal antennal segments of ♀ very short (4th segment approx. as long as wide; c); lateral lobes of mesoscutum mainly smooth, (rather) densely punctate, punctulate or rugose-punctate, interspaces (as far as present) between punctures usually largely smooth and shiny (d), but sometimes distinctly granulate; marginal cell of fore wing of ♀ usually robust and ending further removed from wing apex (e); [wing membrane more or less infuscate; precoxal area coarsely vermiculate-rugose medially; hind femur at least apico-dorsally dark brown or black; maximum width of hypoclypeal depression usually 0.3–0.4 × minimum width of face, if 0.5 × then ventral margin of clypeus thick; vein 1-R1 of fore wing 1.0–1.2 × length of pterostigma]...................................27

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–

Length of malar space of ♀ 0.20–0.45 × height of eye (aa) and clypeus near lower level of eye in lateral view (bb); basal antennal segments of ♀ usually moderately slender (with 4th segment distinctly longer than wide; cc); lateral lobes of mesoscutum finely granulate, punctulate or moderately punctate, and often with a satin sheen (dd), but sometimes shiny (A. hirtus); marginal cell of fore wing of ♀ slender and ending closer to wing apex (ee), except in A. morio (eee); [wing membrane usually subhyaline; basal half of hind tibia largely pale yellowish or reddish, but less so in A. morio; vein 1-R1 of fore wing usually at least 1.4 × length of pterostigma, but approx. equal in A. morio (eee)].................................................................35

27

Area between ocellus and eye, vertex and temple sparsely punctate (a); head of ♀ entirely brownish yellow (b; of ♂ variable, at least stemmaticum black); 1st metasomal tergite 1.5–1.7 × wider posteriorly than subbasally (c); length of eye 1.0–1.2 ×temple in dorsal view (d)..................................................28

–

Area between ocellus and eye, vertex and temple at least moderately densely punctate or rugose (aa); head of ♀ black (bb) or more or less dark red (bbb; of ♂ black); 1st tergite 1.3–1.4 × wider posteriorly than subbasally (cc); length of eye 1.2–1.4 × temple in dorsal view (dd)...............................................29

Revision of western Palaearctic Aleiodes Wesmael, II.

35

28

Eye small (a) and in lateral view maximum width of temple 1.5–1.6 × maximum width of eye (b); ventral margin of clypeus thin and protruding anteriorly (c); antennal segments of ♀ 45–47 (of ♂ 56–58); mesoscutum of ♀ comparatively convex (d); [apex of hind tibia and basal part of palpi of ♀ dark brown]............................................A. ruficeps (Telenga, 1941)

–

Eye medium-sized (aa) and in lateral view temple hardly wider than eye (bb); ventral margin of clypeus thick and hardly protruding anteriorly (cc); antennal segments of ♀ 35–37; mesoscutum of ♀ less convex (dd); [apex of hind tibia and palpi of ♀ yellowish brown]..................A. arnoldii (Tobias, 1976)

29

Area between posterior ocellus and eye moderately punctate (a); posterior half of notauli shallow (b); head in anterior view rather trapezoid (c); [antenna of ♀ 1.1–1.2 × fore wing; 4th antennal segment of ♀ moderately robust; ptero­ stigma blackish; antennal segments of ♀ approx. 47].......A. turcicus sp. nov.

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–

Area between posterior ocellus and eye densely (and finely) rugose (aa), sometimes superficially so and rugulose or with some punctures; posterior half of notauli deep (bb); head in anterior view less trapezoid (cc); [antenna of ♀ with 30–47 segments; if antenna of ♀ with 54–64 segments, cf. A. ferrugiteli (Shenefelt, 1975) from C. Asia].................................................................30

30

Fore femur of ♀ subparallel-sided and 3.9–4.0 × longer than wide (a; this character is less reliable for ♂); antenna of ♀ 0.8–0.9 × fore wing (b); hypo­ clypeal depression usually slightly wider, 0.45–0.50 × minimum width of face (c); head of ♀ largely black (d), rarely face partly reddish; antennal segments of ♂ 36–46(–51) (usually 39–44); [tegulae usually (partly) dark brown; antennal segments of ♀ 29–41].....................................................................31

–

Fore femur of ♀ inflated and 3.0–3.6 × longer than wide (aa); antenna of ♀ 0.9–1.2 × fore wing (bb); hypoclypeal depression usually narrower, ca 0.40 × minimum width of face (cc); head of ♀ at least partly reddish brown (dd); antennal segments of ♂ 47–63 (usually 48–54); [pale males have whole frons and stemmaticum yellowish; palpi dark brown or blackish, rarely brown; OOL of ♂ 1.5–2.0 × diameter of ocellus]..................................................32

Revision of western Palaearctic Aleiodes Wesmael, II.

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31

Antennal segments of ♀ ca 41; subbasal antennal segments of ♀ dark brown and robust, with 4th segment as long as wide (a); basal half of 3rd tergite entirely coarsely striate (b; of ♂ sometimes with curved striae posteriorly); hind trochanter and trochantellus largely dark brown (c); inner and dorsal side of hind tibia apically dark brown (d); parastigma mostly brown (e); vein 1-CU1 of fore wing slightly longer than vein cu-a (f); [palpi dark brown, if largely ivory, cf. A. periscelis; 3rd tergite only anteriorly reddish or yellowish; marginal cell of ♂ wide (Fig. 445); if slender, cf. A. ruficornis]; C. Europe (Alpine).................... A. improvisus sp. nov.

–

Antennal segments of ♀ 29–39; subbasal antennal segments of ♀ yellow and comparatively slender, with 4th segment ca 1.2 × as long as wide (aa); 3rd tergite weakly sculptured, with (faint) curved or antero-medially transverse rugulae or striae (bb) or largely smooth (but sometimes with basal longitudinal striae laterally and often with distinct punctures laterally); hind trochanter and trochantellus yellowish or reddish brown (cc); inner and/or dorsal side of hind tibia (largely) yellowish or red apically (dd); parastigma mostly yellowish (ee); vein 1-CU1 distinctly longer than vein cu-a (ff); [palpi usually brownish or yellowish, but sometimes dark brown; pale males nearly always have frons medially and stemmaticum black; hind tibial spurs of male are usually blunt apically]; Mediterranean, C. Asia...................... A. gasterator (Jurine, 1807)

32

OOL of ♀ approx. 2.6 × diameter of posterior ocellus (a); vein 3-SR of fore wing of ♀ 1.7–2.0 × vein 2-SR (b; of ♂ 1.2–1.5 ×); penultimate antennal segment of ♂ ca 1.2 × longer than wide (c); stemmaticum of ♀ usually black or dark brown (d), rarely reddish; telotarsi of ♀ dark brown (e); scapus of ♀ often black dorsally (f ); [antenna of ♂ 0.9 × as long as body; inner side of hind tibia of ♀ dark brown apically]............... A. carbonarius Giraud, 1857

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–

OOL of ♀ 1.4–2.3 × diameter of posterior ocellus (aa); vein 3-SR of fore wing of ♀ 1.5–1.6 × vein 2-SR (bb; of ♂ 1.0–1.4 ×); penultimate antennal segment of ♂ approx. as long as wide (cc); stemmaticum of ♀ yellowish brown or reddish (dd); telotarsi of ♀ yellowish brown or reddish (ee); scapus of ♀ variable, brownish yellow dorsally (ff) to blackish..............................33

33

Length of eye 1.5–2.0 × temple in dorsal view (a; if measured with posterior ocelli up to posterior level of eyes); OOL of ♀ 1.2–1.8 × diameter of posterior ocellus (b); subbasal antennal segments of ♀ slightly less moniliform (c); [inner side of hind tibia of ♀ usually dark brown or blackish apically; colour of legs of ♂ usually similar to legs of ♀ and usually partly yellowish; antenna of ♂ approx. as long as body and 1.2–1.4 × fore wing; antennal segments of ♀ 34–39(–47), of ♂ (44–)47–60]........A. ruficornis (Herrich-Schäffer, 1838)

–

Length of eye 1.1–1.5 × temple in dorsal view (aa; if measured with posterior ocelli up to posterior level of eyes); OOL of ♀ usually 1.9–2.3 × diameter of posterior ocellus (bb); subbasal antennal segments of ♀ distinctly submoniliform (cc); [inner side of hind tibia of ♀ yellowish; hind femur and basitarsus of both sexes more robust (but in ♂ sometimes rather slender); legs of males of W. European specimens strongly infuscate, darker than legs of females, but legs of N. European specimens paler; antenna of ♂ 0.8–0.9 × body and 1.0–1.1 × fore wing; antennal segments of ♀ 35–45(–46), of ♂ (44–)47–61]..............34

Revision of western Palaearctic Aleiodes Wesmael, II.

39

34

Apical tooth of hind tarsal claws of ♀ robust (a); 2nd metasomal tergite of ♂ orange brown (b); hind femur (c), tibia and basitarsus (d) of ♂ more robust and femur basally yellowish brown (e), if rarely almost black then hind tibia basally yellowish brown (f ); clypeus less protruding in front of face (g); boreal and highland species.....................................A. grassator (Thunberg, 1822)

–

Apical tooth of hind tarsal claws of ♀ slender (aa); 2nd metasomal tergite of ♂ black (bb); hind femur (cc), tibia and basitarsus (dd) of ♂ comparatively slender and femur basally black (ee); hind tibia basally black (ff); clypeus more protruding in lateral view (gg); lowland species..... A. carbonaroides sp. nov.

35

Head brownish yellow (a); ventral margin of clypeus thin and distinctly protruding anteriorly (b); vertex (c) and mesoscutum (d) shiny; maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (e); pterostigma brownish yellow (f ); [tarsal claws medium-sized and yellowish pectinate; body laterally and dorsally (except more or less dark brown propodeum and 1st tergite) yellowish brown; occipital carina weakly indicated medio-dorsally]........................................... A. fahringeri (Telenga, 1941)

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–

Head black (aa); ventral margin of clypeus thick and usually hardly protruding (bb); vertex (cc) and mesoscutum (dd) usually rather dull and with satin sheen (cc); if shiny (ccc) then maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (ee) and pterostigma dark brown (ff); [pterostigma yellowish (fff ) in A. pallidistigmus].........................................36

36

Vertex (a) and mesoscutum (b) distinctly shiny because of smooth interspaces between punctures or rugae; head conspicuously setose because of long setae (c; less distinctive in ♀), as are propodeum and first tergite laterally; trochan­ ters (and often also trochantelli) nearly always at least somewhat infuscate, darker than orange part of femora (d); subbasal antennal segments of ♀ dark brown; [palpi blackish or dark brown; tarsal claws with small pecten; 3rd–6th antennal segments robust, hardly longer than wide; fore femur slender]........ .......................................................................... A. hirtus (Thomson, 1892)

–

Vertex (aa) and mesoscutum (bb) rather dull and with satin sheen, interspaces finely coriaceous-granulate between punctures or rugulae; head usually less conspicuously setose (cc); if sculpture and setosity are intermediate then trochanters and trochantelli have same colour as basal part of femora (dd) and subbasal antennal segments of ♀ brownish yellow.....................................37

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37

Vein cu-a of fore wing at least as long as vein 1-CU1 (a); 1st tergite of ♀ strongly widened apically (b); marginal cell of fore wing shorter (c); subbasal antennal segments of ♀ subquadrate (d); hind coxa (as femur) completely black (e); clypeus distinctly protruding anteriorly (f ); fore femur largely dark brown (g); [OOL distinctly longer than diameter of posterior ocellus; 2nd tergite finely and densely sculptured; if fore and hind femora yellowish brown, clypeus hardly protruding and marginal cell of fore wing elongate, cf. A. sibiricus (Kokujev)]......................................................................................38

–

Vein cu-a of fore wing distinctly shorter than vein 1-CU1 (aa); 1st tergite moderately widened apically (bb); marginal cell of fore wing long (cc); subbasal antennal segments of ♀ longer than wide (dd); hind coxa orange or yellowish brown (ee); clypeus hardly or not protruding anteriorly (ff); fore femur brownish yellow (gg).......................................................................39

38

Wings infuscate apically (a); 2nd submarginal cell of fore wing less widened posteriorly (b); pterostigma medially dark brown (c); basal 0.4 of hind ti­ bia yellowish (d); 2nd–5th metasomal tergites orange brown (e); middle lobe of mesoscutum densely punctate, without distinct granulation in between punctures (f ); [vein 2-1A of hind wing comparatively long; pecten of tarsal claws present and claws rather robust].........................A. nigrifemur sp. nov.

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–

Wings subhyaline apically (aa); 2nd submarginal cell of fore wing widened posteriorly (bb); pterostigma medially pale or yellowish brown (cc); hind tibia mainly dark brown, only basally narrowly pale yellowish (dd); 2nd–5th tergites black (ee); middle lobe of mesoscutum coriaceous (ff); [basal half of 3rd tergite and OOL rugulose].............................A. morio (Reinhard, 1863)

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Apex of hind tibia reddish or yellowish (a); pterostigma brownish yellow medially (b), rarely darkened; 3rd tergite dull (c); [antenna of ♀ with 54–64 segments]................................................ A. pallidistigmus (Telenga, 1941)

–

Apex of hind tibia dark brown or infuscated (aa); pterostigma more or less dark brown medially (bb); 3rd tergite usually shiny (cc), but sometimes rather dull (ccc)....................................................................................................40

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Tarsal claws with medium-sized pecten (a); precoxal area of mesopleuron smooth medially, but sometimes with sparse weak punctures or some rugae below it (b); temple shiny (c); basal half of antenna of ♀ largely yellowish brown (d); [tegula and humeral plate equally yellowish orange; hind tarsus partly yellowish or brownish; hind tibia dorsally dark brown at extreme base, then pale subbasally and infuscate apically].................................................... ....................................................A. pallidicornis (Herrich-Schäffer, 1838)

–

Tarsal claws without pecten (aa) or with fine pale pecten; precoxal area of mesopleuron moderately rugose medially (bb); temple rather dull (cc); basal half of antenna blackish (dd)......................................................................41

41

Area between posterior ocellus and eye mainly granulate or coriaceous, at most with some punctures or rugulae (a); eyes larger in lateral view (b) and less protuberant in dorsal view (c); 3rd metasomal tergite usually largely smooth, especially in ♀ (but basal half in ♂ sometimes extensively striate-rugulose) and as strongly glossy as following tergite (d); precoxal area comparatively narrow and posteriorly largely or completely smooth (e); antennal segments of ♀ 47–57(–58); hind femur largely reddish apically, with only slight infuscation (f ); N & C Europe...................... A. unipunctator (Thunberg, 1822)

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Area between posterior ocellus and eye densely rugose, coarsely punctate or rugulose (aa); eyes smaller in lateral view (bb) and more protuberant in dorsal view (cc); basal half of 3rd tergite distinctly striate or densely rugulose and less shiny (dd), but intermediates occur; precoxal area comparatively wide and usually posteriorly rugose or distinctly punctate (ee); antennal segments of ♀ 54–62; hind femur apically more or less smudged dark brown or black (ff), but sometimes very indistinct; [if mesoscutum rather steep anteriorly and width of hypoclypeal depression 0.4–0.5 × minimum width of face, and long malar space, cf. A. gasterator (Jurine) and related spp.]; S & E Palaearctic......A. eurinus (Telenga, 1941)

Biology and descriptions Aleiodes aestuosus (Reinhard, 1863) Figs 22–37 Rogas aestuosus Reinhard, 1863: 265; Shenefelt, 1975: 1216–1217; Zaykov, 1980a: 112; Tobias, 1976: 84, 1986: 78 (transl.: 129); Kotenko, 1992: 96 [examined]. Rhogas aestuosus; Fahringer, 1931: 232–234. Aleiodes (Neorhogas) aestuosus; Papp, 1985a: 152, 1989: 52, 1990: 90, 1991a: 67–68. Aleiodes (Chelonorhogas) aestuosus; Chen & He, 1997: 38; He et al., 2000: 665; Belokobylskij, 2000: 26; Chen et al., 2003: 211; Papp, 2012: 187; Farahani et al., 2015: 238–240. Aleiodes aestuosus; Fortier & Shaw, 1999: 230; Zaldivar-Riverón et al., 2004: 234. Rhogas (Rhogas) aestuosus var. desertus Telenga, 1941: 152–153, 404 (not Rhogas (R.) desertus Telenga, 1941). Type material. Holotype, ♀ (MNHN), “Cipro [= Cyprus]”, “Muséum Paris, 1867, coll. O. Sichel”, “Rogas aestuosus Rhd.”. Additional material. Albania, Bulgaria, Cyprus, Greece, Russia, Turkey, Tunisia, [Azerbaijan, Georgia, Iran, Iraq, Israel, Jordan, Syria, Turkmenistan, Uzbekistan]. Specimens in BMNH, BZL, CNC, HSC, MRC, MTMA, NMS, RMNH, ZSSM, ZISP. Distributed principally in Asia Minor, extending to Cyprus where it has been collected plentifully. Only single specimens examined from Albania (MTMA) and mainland Greece (BZL), but in North Africa it apparently extends westwards to Tunisia (one specimen in BZL). Molecular data. MRS004 (Turkey).

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Figures 22–25. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River, but 25 from Uzbekistan, Qamashi 22 ovipositor sheath lateral 23 habitus lateral 24 apex of antenna 25 mummy of Heliothis peltigera (Denis & Schiffermüller).

Biology. Collected March–July, often at light, but it is not clear how many generations are represented nor how the winter is passed. Reared from Noctuidae: Heliothis peltigera (Denis & Schiffermüller) (4 [1 CNC/Iraq, 1 ZISP (with mummy)/Uzbekistan, 2 MTMA/Iraq]), Sesamia sp. (2 [BMNH/Iran]). This indicates a host range of both endophagous and exophagous larvae, but the individuals purporting to be from Sesamia are labelled [no doubt incorrectly] “ex pupa” and lack mummies, suggesting that they may have resulted from substrate rearings (presumably from stems of crop species of Poaceae, inside which Sesamia larvae feed and pupate) rather than from isolated hosts, with a consequent reduction in the reliability of the host determination and suspicion that mummies of other hosts could have been overlooked on the

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stems (see also remark under A. apicalis). On the other hand, the large hypoclypeal depression and somewhat protruding clypeus does indicate that A. aestuosus adults are equipped to chew their way out of mummies made in concealed sites. The hosts given above are regular crop pests, but the paucity of reared material examined may suggest that A. aestuosus is not especially associated with cultivated habitats. The single mummy seen (Fig. 25) is rather elongate, scarcely arched, and the cocoon occupies most of the host abdomen. It has the appearance of not being securely stuck to the substrate. Diagnosis. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 34); clypeus rather protruding anteriorly and rather thick ventrally (Fig. 36); head brownish yellow; vertex finely punctate; lateral lobes of mesoscutum sparsely and finely punctate, with wide smooth interspaces; precoxal sulcus absent, area only sparsely finely punctate or smooth; 1-CU1 of fore wing subequal to vein 2-CU1 (Fig. 26); hind tarsal claws with brownish pecten (Fig. 37); only apex of hind tibia dark brown; metasoma of ♀ completely yellowish and distinctly depressed subapically, 1st tergite partly and 4th–6th tergites of ♂ often blackish. Sometimes entire body (including propodeum and 1st metasomal tergite) yellowish (“var. desertus”). Description. Redescribed ♀ (RMNH) from Turkey (Icil). Length of fore wing 6.8 mm, of body 8.3 mm. Head. Antennal segments of ♀ 52, length of antenna 1.1 × fore wing, its subapical segments approx. as long as wide; frons with irregular curved rugae, shiny, and rugose behind antennal sockets; OOL 2.4 × diameter of posterior ocellus, and finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate, shiny; clypeus short, coarsely and densely punctate; ventral margin of clypeus thick and rather protruding forwards (Fig. 36); width of hypoclypeal depression 0.65 × minimum width of face (Fig. 34); length of eye 0.8 × temple in dorsal view (Fig. 35); vertex behind stemmaticum sparsely punctate; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely smooth, shiny, sparsely and finely punctate; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron and metapleuron remotely punctate, interspaces much wider than diameter of punctures, shiny; mesopleuron above precoxal area (except speculum) sparsely punctate; scutellum slightly convex, remotely punctate and evenly rounded laterally, no carina; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete, but irregular posteriorly, without tubercles. Wings. Fore wing: r 0.4 × 3-SR (Fig. 26); 1-CU1 horizontal, nearly as long as (0.9 x) 2-CU1; r-m 0.9 × 2-SR, and 0.7 × 3-SR; second submarginal cell medium-sized (Fig. 26); cu-a vertical, not parallel with CU1b, straight; 1-M rather curved posteriorly. Hind wing: marginal cell gradually and evenly widened, its apical width 2.3 × width at level of hamuli (Fig. 27); 2-SC+R shortly longitudinal; m-cu distinct; M+CU:1-M = 23:19; 1r-m 0.7 × 1-M. Legs. Tarsal claws subpectinate, with four brown medium-sized pectinal bristles (Fig. 37); hind coxa remotely punctate, shiny; hind trochantellus robust; length of hind femur and basitarsus 3.0 and 3.5 × their width, respectively; length of inner hind spur 0.55 × hind basitarsus; hind tibia slender (Fig. 23).

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Figures 26–37. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River 26 fore wing 27 hind wing 28 mesosoma lateral 29 mesosoma dorsal 30 metasoma dorsal 31 fore femur lateral 32 hind femur lateral 33 base of antenna 34 head anterior 35 head dorsal 36 head lateral 37 inner hind tarsal claw.

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Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites coarsely and densely rugose, robust, with distinct median carina; medio-basal area of 2nd tergite wide and short; 2nd suture deep medially and shallow laterally; basal half of 3rd tergite finely rugose, remainder of metasoma largely smooth, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with mediumsized setae and apically rounded (Fig. 22). Colour. Brownish yellow; antenna, mesosternum (except anteriorly) and mesopleuron (except anteriorly and dorsally), metapleuron, propodeum, ovipositor sheath and stemmaticum black; hind tibia (except apically) pale yellowish; apices of femora (dorsally) and tibiae, palpi, tarsi (except basally), veins and pterostigma dark brown; wing membrane rather infuscate. Variation. Size of eyes and ocelli rather variable. Mesopleuron, mesosternum, metapleuron and propodeum brownish yellow or black; 1st tergite entirely brownish yellow or with dark brown patch basally; in desert areas body can be wholly orange. Antennal segments: ♀ 49 (1), 50 (3), 51 (9), 52 (13), 53 (10), 54 (3), 55 (5), 56 (2); ♂ 51 (10), 52 (11), 53 (5), 54 (4), 55 (3), 56 (1). The two sexes have comparable numbers of antennal segments. Apical tergites of ♂ type 3 and fringe moderately strong; inner hind tibial spur 0.50 × as long as hind basitarsus. Distribution. Albania, Azerbaijan, Bulgaria, Cyprus, Georgia, *Greece, Iran, *Iraq, Israel, *Jordan, Russia, Syria, Turkey, Tunisia, *Turkmenistan, Uzbekistan. Aleiodes agilis (Telenga, 1941) Figs 38–49 Rhogas (Rhogas) agilis Telenga, 1941: 165–166, 417. Rogas agilis; Shenefelt, 1975: 1217. Rogas (Rogas) agilis; Tobias, 1976: 83, 1986: 76 (transl. 122, 124) (lectotype designation). Aleiodes agilis; Fortier & Shaw, 1999: 230. Aleiodes (Chelonorhogas) agilis; Ma et al., 2002: 98; Farahani et al., 2015: 240. Rhogas desertus var. armenica Telenga, 1959: 85; Tobias, 1976: 83 (as synonym of A. agilis (Telenga, 1941)), 1986: 76 (transl. 122, 124; id.). Type material. Lectotype, ♀ (ZISP; examined via photos), “Persiya [= Iran], Tavriz, 21.iii.[19]14, Andrievskij”, “Rhogas agilis sp. n., N. Telenga det.”, “Syntypus agilis Tel.”, “Lectotypus Rogas agilis Tel., design. Tobias, 1980”; paralectotype, ♀ (ZISP; id.), “Armenia, pr. Eriwan [= Yerevan], A. Schelkovnikow / Ragakag, 19.iii.[19]25”, “Paralectotypus Rhogas agilis Telenga, design. Tobias, 1986”. In the original description the latter date is incorrectly cited as 24.vii.1925. Molecular data. None. Biology. Unknown. It appears to fly very early in the year (March). Diagnosis. Maximum width of hypoclypeal depression approx. 0.8 × minimum width of face; anterior part of clypeus very narrow (Fig. 46); OOL 1.0–1.3 × dia­

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meter of posterior ocellus and coarsely remotely punctate with some weak rugulosity; head and mesosoma (except pronotal side partly and mesoscutum medio-posteriorly and laterally) blackish; mandible massive triangular, coarsely punctate and with thick ventral lamella (Fig. 46); face largely transversely rugose and conspicuously whitish setose; frons rugose and shiny; vertex and temple coarsely remotely punctate and shiny; area of precoxal sulcus (but posteriorly superficially) distinctly rugose; lateral lobes of mesoscutum largely smooth (anteriorly becoming densely punctate and somewhat rugose), whitish setose and with satin sheen, middle lobe distinctly punctate; basal half of wings (except anteriorly) largely glabrous and remainder of wing inconspicuously setose; vein r of fore wing approx. 0.6 × vein 3-SR (Fig. 40); vein 1-CU1 0.2–0.3 × as long as 2-CU1, narrow and subhorizontal; tarsal claws long, slender, hardly bent and simple (Fig. 44); 1st and base of 2nd tergite weakly longitudinally rugulose with some superficial punctures; metasoma dark brown but with yellow patches (Fig. 38), clypeus and antenna (except yellow scapus and pedicellus) yellowish brown; legs and palpi pale yellowish, but hind coxa and most of middle coxa dark brown. Description. Paralectotype, ♀, length of fore wing 6.6 mm, of body 7.0 mm. Head. Antennal segments of ♀ 47, antenna as long as body and its subapical segments moderately slender; frons rugose, shiny; OOL 1.3 × diameter of posterior ocellus; OOL and vertex remotely punctate, with satin sheen, OOL also with some rugulae; anterior part of clypeus 9 × wider than high, coarsely punctate and rather convex; clypeus above lower level of eyes; ventral margin of clypeus thick and not protruding forwards (Fig. 48); width of hypoclypeal depression 0.8 × minimum width of face (Fig. 46); length of eye 1.8 × temple in dorsal view; vertex behind stemmaticum convex and sparsely punctate; length of malar space 0.19 × length of eye in lateral view; occipital carina nearly complete, fine and ventrally strongly curved; mandible massive triangular, coarsely punctate and with thick ventral lamella (Fig. 46). Mesosoma. Lateral lobes of mesoscutum largely smooth, with satin sheen and whitish setose, middle lobe distinctly punctate and setose; prepectal carina complete and lamelliform; precoxal area of mesopleuron widely rugose, but posterior 0.2 narrowly striate; mesopleuron largely weakly and sparsely punctate, shiny, but anteriorly becoming densely punctate and somewhat rugulose; scutellum largely smooth, with some punctures; propodeum evenly convex, finely rugose and with medio-longitudinal carina, without tubercles. Wings. Fore wing: basal half largely glabrous; r 0.6 × 3-SR (Fig. 40); 1-CU1 subhorizontal, 0.25 × as long as 2-CU1; r-m 0.7 × as long as 3-SR; 2ndsubmarginal cell robust (Fig. 40); cu-a distinctly inclivous; 1-M weakly curved posteriorly. Hind wing: basal 0.4 of marginal cell slightly widened and distally strongly widened, its apical width approx. twice width at level of hamuli; 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 24:19; 1r-m 0.7 × 1-M. Legs. Tarsal claws slender, slightly curved and only setose (Fig. 44); hind coxa partly obliquely striate dorsally; tarsi slender, segments (except telotarsus) with long apical spiny bristles; length of hind femur and basitarsus 5.0 and 6.0 × their width, respectively; length of inner hind spur 0.3 × hind basitarsus.

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Figures 38, 39. Aleiodes agilis (Telenga), ♀, paralectotype 38 habitus lateral 39 ovipositor sheath lateral. Photographs by K. Samartsev.

Metasoma. First tergite robust, as long as wide apically, distinctly narrowed anteriorly and rather flat posteriorly; 1st and 2nd tergites finely longitudinally striate-rugulose; medio-longitudinal carina of 1st and 2nd tergites indistinct; 2nd tergite 0.7 × longer than its basal width; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture shallow and narrow; 3rd tergite mainly smooth and with satin sheen; 4th and apical half

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Figures 40–49. Aleiodes agilis (Telenga), ♀, paralectotype, but 47 of lectotype 40 wings 41 mesosoma lateral 42 fore femur lateral 43 hind femur lateral 44 outer hind tarsal claw 45 fore leg 46 head anterior 47 head dorso-lateral 48 head lateral 49 base of antenna. Photographs by K. Samartsev.

of 3rd tergite without sharp lateral crease; ovipositor sheath rather slender, with short setae and apically truncate (Fig. 39). Colour. Black; pronotal side largely yellowish brown; mesoscutum medio-posteriorly and postero-laterally partly chestnut brown; tegulae, clypeus and antenna (except

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yellow scapus and pedicellus) yellowish brown; mandible, legs and palpi pale yellowish, but hind coxa and most of middle coxa dark brown; metasoma dark brown but with yellow patches (Fig. 38); pterostigma brown medially and dark brown laterally; ovipositor sheath dark brown; veins of fore wing (but pale yellow in basal 0.2 of fore wing) brown; wing membrane hyaline. Distribution. Armenia, Iran. Included in this revision, because it may occur in Turkey. Aleiodes apicalis (Brullé, 1832) Figs 50–71 Bracon apicalis Brullé, 1832: 381 [examined]. Rhogas apicalis; Fahringer, 1932: 317–318. Rogas apicalis; Shenefelt, 1975: 1218. Aleiodes apicalis; Shaw et al., 1998: 63; Fortier & Shaw, 1999: 227; Zaldivar-Riverón et al., 2004: 234, 2008: figs 2–6. Aleiodes (Chelonorhogas) apicalis; Falco et al., 1997: 60; Farahani et al., 2015: 240–242; Abdolalizadeh et al., 2017: 36. Rogas reticulator Nees, 1834: 211; Shenefelt, 1975 (as synonym of A. ductor); Papp, 2005: 176 (id.). Syn. nov. Aleiodes reticulator; Papp, 1991a: 70 (as synonym of A. ductor). Rogas bicolor Lucas, 1849: 336–337 (not Spinola, 1808); Shenefelt, 1975: 1219; Papp, 1985a: 157 (lectotype designation), 2005: 176 (as synonym of A. ductor). Syn. nov. Rogas rufo-ater Wollaston, 1858: 24; Shenefelt, 1975: 1247; Papp, 1990: 90 (as synonym of A. ductor) [examined]. Syn. nov. Rhogas rufoater; Fahringer, 1934: 321. Rhogas bicolorinus Fahringer, 1932: 318 (replacement name for Rogas bicolor Lucas). Syn. nov. Rhogas reticulator var. atripes Costa, 1884: 13; Papp, 1990: 90 (as synonym of R. rufoater). Syn. nov. Rhogas ductor var. atripes; Fahringer, 1932: 244. Aleiodes (Neorhogas) ductor var. atripes; Papp, 1985a: 157. Rhogas similis Szépligeti, 1903: 114 (not Curtis, 1834); Papp, 1985a: 157–158 (lectotype designation and as synonym of A. ductor), 2005: 176 (id.). Syn. nov. Rhogas ductor var. similis; Fahringer, 1932: 245. Rogas ductor auct. p.p.; Shenefelt, 1975: 1226–1227; Zaykov, 1980a: 112; Tobias, 1976: 85, 1986: 80 (transl.: 133); Samartsev & Belokobylskij, 2013: 765. Aleiodes ductor auct. p.p.; Bergamasco et al., 1995: 5. Type material. Holotype of B. apicalis, ♂ (MNHN) “[Greece], Morée, Muséum Paris, Brullé 4187-33”, “Type”, “Bracon apicalis Brullé, Type”. Lectotype of R. similis, ♂ (MTMA) “[Hungary], Kecskemét, Szépligeti”, “Hym. Typ. No. 7021, Mus. Budapest”, “Lectotypus”, “Rhogas similis Szépl. 1903 ♂, Papp, 1984”, “Rhogas reticulator

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Figures 50–53. Aleiodes apicalis (Brullé), ♀, Greece, Thimiana Chios, but 52 mummies of Autographa gamma (Linnaeus) from Malta and 53 of undetermined plusiine host from Cyprus 50 habitus lateral 51 ovipositor sheath lateral 52 mummy dorsal 53 mummy covered by silk of host.

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var. similis Sz., det. Szépligeti, 1906”, “Aleiodes ductor Thunbg., det. Papp, J., 1983”. Holotype of R. rufo-ater, ♂ (BMNH) “[Portugal], Madeira, Wollaston, 55.7”, “Rogas rufo-ater, W.”, Type, H.T.”, “B.M. Type Hym., 3.c.241”. Additional material. Albania, Austria, Bosnia & Herzegovina, Bulgaria, Croatia, Cyprus, Czech Republic, France (including Corsica), Germany, Greece (including Chios, Corfu, Crete, Lesbos, Rhodes), Hungary, Italy (including Sardinia, Sicily), Malta, Moldova, Montenegro, Morocco, North Macedonia, Portugal (including Madeira), Romania, Russia (including Dagestan), Serbia, Slovakia, Spain (including Mallorca and Canary Islands: Tenerife, Fuerteventura), Switzerland, Tunisia, Turkey, Ukraine, [Georgia, Kazakhstan, Oman, Iran, Iraq, Israel, Syria, Turkmenistan]. Specimens in BMNH, BZL, CMIM, CNC, HSC, MRC, MSC, MTMA, NMS, RMNH, SDEI, UNS, ZISP, ZSSM. This is a mainly Mediterranean species, extending into Central Europe and West Asia, and one of the commonest species of the group in the Mediterranean area. One surprising female from Sweden (Skåne, Käseberga, MV light 17-vii-14.ix.2013, N. Ryrholm & C. Källander, in NMS) is presumed, like two British specimens (England, V.C. 3, S. Devon, Slapton Ley 7–14.vi.1932, H.St.J. Donisthorpe, in BMNH; V.C. 22, Berkshire, Beale Park, 25–27.vii.2018, Rothamsted trap, in coll. A.C. Galsworthy destined for BMNH) and one specimen from Netherlands (Lexmond, ZH, 10.viii.2004, C. Gielis in RMNH) to have been deposited there by winds from southern Europe or N. Africa rather than representing an established breeding population. Whether or not A. apicalis can eventually establish permanent populations, i.e., with winter survival, in these relatively northerly parts of Europe may depend on whether its host can do likewise. Molecular data. MRS008 (Turkey), MRS111 (Turkey), MRS112 (Turkey), MRS181 (Russia), MRS869 (Sweden). Biology. Time of flight varies according to harshness of summer. In its more temperate sites plurivoltine April-September(October), overwintering in the mummy, but in Cyprus (and presumably other places with extremely hot dry summers) it appears to be most active from autumn to spring (October–May), with a prolonged summer diapause (June–October or later) in the mummy (reared series ex “Plusia” in BMNH and NMS, W.R. Ingram, six with mummification dates recorded in May or June and adult emergence in the following October–December, further specimens in the series have only one date, which is ambiguous). Reared from Noctuidae: Autographa gamma (Linnaeus) (6 [4 ZISP/Moldova, 1 HSC/Germany, 1 NMS/Malta]; J.L. Gregory, H. Schnee), indet. Plusiinae (14). There is no reason to suppose that the hosts recorded as indet. Plusiinae are anything except A. gamma. A further specimen labelled as ex Peribroma [sic] saucia is accompanied by a clearly Plusiinae mummy (Sicily, NMS). Also, one labelled as from Anarsia lineatella Zeller (Gelechiidae) (Ukraine, ZISP), but without a mummy and clearly in error on grounds of size alone. Another specimen labelled as “ex Sesamia pupa” (Iran, BMNH) lacks its mummy but accompanies two individuals of A. aestuosus (q. v.) from the same source, and the remarks made under that species apply also to this record – but with the added objection that the small hypoclypeal opening and flat clypeus of A. apicalis strongly suggest that its hosts do not

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Figures 54–65. Aleiodes apicalis (Brullé), ♀, Bulgaria, Rodopi 54 wings 55 mesosoma lateral 56 mesosoma dorsal 57 1st –3rd metasomal tergites dorsal 58 fore femur lateral 59 hind femur lateral 60 head anterior 61 head dorsal 62 head lateral 63 outer hind tarsal claw 64 base of antenna 65 apex of antenna.

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mummify in deep concealment. The mummy (Fig. 52) is of a pale chalky buff colour, and the cocoon occupies approx. abdominal segments 4–7 of the host larva. Several of the mummies examined, all of which seem to be penultimate instar, have been formed in a more or less curled leaf beneath a web (Fig. 53) that the host had been induced to spin before being mummified, and were weakly stuck to the substrate. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 60); antennal segments of ♀ 44–51 and flagellar segments moderately robust (Figs 64, 65); ventral margin of clypeus thick and obtuse apically and clypeus not protruding outwards (Fig. 62); vertex, mesoscutum, metapleuron and scutellum normally shiny and without dense granulation, at most with some superficial microsculpture; frons (and more or less vertex) with striae (Fig. 61) or rugae; scutellum largely smooth and shiny; mesopleuron largely smooth; vein 2-CU1 of fore wing approx. as long as vein 1-CU1 (Fig. 54); vein M+CU of hind wing distinctly longer than vein 1-M (Fig. 54); hind tarsal claws of ♀ with rather slender and brownish pecten (Fig. 63); basal half of hind tibia (largely) pale yellowish, or if black (var. rufoater) then also fore femur black; 3rd tergite (except basally) largely smooth; medially 4th–6th tergites of ♂ slightly concave and with dense band of medium-sized setae (Figs 68, 69); head, mesoscutum and scutellum black; 2nd tergite yellowish or reddish. Description. Redescribed ♀ (RMNH) from Hungary (Budapest), length of fore wing 5.1 mm, of body 6.7 mm. Head. Antennal segments of ♀ more than 40, but apical segments missing (length of antenna of ♀ from Lesbos 1.4 × fore wing and its subapical segments robust); frons with coarse curved rugae, shiny; OOL 1.5 × diameter of posterior ocellus, and distinctly striate; vertex transversely striate, rather weak; clypeus normal, punctulate and convex; ventral margin of clypeus thick and not protruding forwards; width of hypo­ clypeal depression 0.3 × minimum width of face (Fig. 60); length of eye 1.6 × temple in dorsal view (Fig. 61); vertex behind stemmaticum transversely striate; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view; occipital carina complete, fine. Mesosoma. Mesoscutal lobes largely smooth, punctulate, shiny; prepectal carina complete, rather strong; precoxal area of mesopleuron largely smooth; mesopleuron above precoxal area weakly and sparsely punctate, especially posteriorly; scutellum largely smooth, with striae laterally; propodeum evenly convex, coarsely vermiculaterugose, only anteriorly with median carina, without tubercles. Wings. Fore wing: r 0.6 × 3-SR (Fig. 54); 1-CU1 horizontal, equal to or slightly longer than 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell comparatively short (Fig. 54); cu-a vertical, slightly curved posteriorly; 1-M straight posteriorly. Hind wing: marginal cell basally slightly and distally strongly widened, its apical width 2.6 × width at level of hamuli (Fig. 54); 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 5:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather slender and medium-sized brownish pecten (Fig. 63); hind coxa largely densely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 5.1 and 6.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus.

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Figures 66–71. Aleiodes apicalis (Brullé), ♂, Turkey, Sivas 66 habitus lateral 67 head dorsal 68 3rd –7th metasomal tergites dorsal 69 3rd –7th metasomal tergites lateral 70 head anterior 71 outer hind tarsal claw.

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Metasoma. First tergite robust, evenly convex; 1st and 2nd tergites rather coarsely obliquely rugose; 1st tergite and basal half of 2nd tergite with median carina; 2nd tergite robust and with striae diverging posteriorly; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture rather deep medially; 3rd tergite largely smooth, except anteriorly with some striae; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with rather long setae and apically rounded (Fig. 51). Colour. Black; scapus, pedicellus, tegulae (but humeral plate brownish yellow), base of hind tibia narrowly, apical half of hind tibia, telotarsi, hind tarsus largely, ventral apical half of metasoma, pterostigma and veins (except C+SC+R of fore wing) dark brown; remainder of basal half of antenna and palpi yellowish brown; remainder of legs (but apical two-fifths of hind femur black), 1st and 2nd tergites, 3rd tergite basally and laterally orange brown; remainder of hind tibia pale yellowish; apex of middle femur and wing membrane somewhat infuscate. Variation. A. apicalis is very variable in colour and the colour patterns are not restricted to certain areas, but in general southern Palaearctic specimens are darker than northern ones (or specimens from high altitudes). The tegula is dark brown or black, and the humeral plate usually paler than the tegula or equally black, but both usually yellowish in southern specimens; the hind tarsus is dark brown or black, but sometimes 3rd and 4th segments yellowish; the hind tibia variably reddish to black, but palest at extreme base; the pronotum is very occasionally reddish. The extent of black colouration of the legs is especially variable, and sometimes all legs are entirely black (var. rufoater (Wollaston, 1858)). Antenna, especially in females, can be more or less light reddish brown, especially basally, or dark brown/black throughout. Antennal segments: ♀ 44(1), 46(3), 47(11), 48(20), 49(31), 50(41), 51(19), 52(10), 54(3), 55(1), 57(1); ♂ 46(3), 47(7), 48(17), 49(29), 50(30), 51(32), 52(11), 53(5), 54(1). Males have on average approx. one antennal segment more than females. Apical tergites of ♂ type 4, setosity dense (making the tergites appear concave; Figs 68, 69) and fringe weak. Distribution. *Albania, Austria, *Bosnia & Herzegovina, *Bulgaria, *Croatia, Cyprus, *Czech Republic, *France (including Corsica), *Georgia, *Germany, Greece (including Chios, Corfu, Crete, Lesbos, Rhodes), *Hungary, Iran, *Iraq, *Israel, *Italy (including Sardinia, Sicily), *Kazakhstan, *Malta, *Moldova, *Montenegro, *Morocco, *North Macedonia, *Oman, *Portugal (including Madeira), *Romania, *Russia (including Dagestan), *Serbia, *Slovakia, Spain (including Mallorca and Canary Islands: Tenerife, Fuerteventura), *Syria, Switzerland, *Tunisia, Turkey, *Turkmenistan. New synonymy. The synonymy of Rogas rufo-ater Wollaston, 1858, and Rhogas similis Szépligeti, 1903, are based on examination of the types listed above. The lectotype of Rogas bicolor Lucas, 1849 (not Spinola, 1808) and of Rhogas bicolorinus Fahringer, 1932, has been examined by Dr Jenö Papp and we agree with his opinion that it is a synonym of A. ductor auct. (= A. apicalis). The types of Rogas reticulator Nees, 1834, and Rhogas reticulator var. atripes Costa, 1884, are lost or unavailable and their synonymy is based on the original description and the interpretation by later authors.

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Aleiodes arnoldii (Tobias, 1976) Figs 72–92 Rogas (Rogas) arnoldii Tobias, 1976: 84, 222, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) arnoldi [sic!]; Papp, 1985a: 152. Aleiodes (Neorhogas) arnoldii; Papp, 1991a: 87. Type material. Holotype, ♀ (ZISP) “[Azerbaijan], Kosmoljan, Zuvan, 19.v.[1]936, Arnoldi”, “Holotypus Rogas arnoldii Tobias”. Additional material. 1 ♂ (RMNH), “Turkey, Hakkâri, Tanin Tanin Pass, 25.vi.1985, 2200 m, C.J. Zwakhals”. Male is provisionally associated with this species; it may belong to a related species. Molecular data. None. Biology. Unknown. The holotype was collected in May. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 78); clypeus obtuse apically and not protruding in lateral view (Fig. 77); length of malar space of ♀ 0.5–0.6 × height of eye in lateral view; antennal segments of ♀ 35–37 and length of antenna of ♀ 0.8–0.9 × fore wing; OOL sparsely punctate; lateral lobes of mesoscutum largely smooth; posterior half of notauli shallow; precoxal

Figures 72, 73. Aleiodes arnoldii (Tobias), ♀, holotype 72 habitus lateral 73 ovipositor sheath lateral. Photographs: K. Samartsev.

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Figures 74–79. Aleiodes arnoldii (Tobias), ♀, holotype 74 wings 75 mesosoma lateral 76 antenna 77 head lateral 78 head anterior 79 head dorsal. Photographs: K. Samartsev.

area coarsely vermiculate-rugose medially; head, palpi and part of mesosoma of ♀ yellowish brown; pterostigma dark brown; apex of hind tibia of ♀ yellowish; hind tarsal claws yellowish or brownish setose (Fig. 72); 4th–6th tergites of ♂ flat and normally setose, but setae slightly longer than on basal tergites (Fig. 92). Description. Holotype, ♀, length of fore wing 4.4 mm, of body 5.7 mm. Head. Antennal segments of ♀ 37, length of antenna 0.85 × fore wing, its subapical segments quadrate; frons with rather coarse curved rugae, shiny, and rugose behind antennal sockets; OOL 2.0 × diameter of posterior ocellus, and finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate, shiny; face transversely rugose; clypeus finely rugulose and with long setae; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression

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Figure 80. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass, habitus lateral.

0.45 × minimum width of face; length of eye 1.1 × temple in dorsal view (Fig. 79); vertex behind stemmaticum rugulose; clypeus near lower level of eyes; length of malar space 0.55 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely smooth, shiny, sparsely and finely punctate; precoxal area of mesopleuron coarsely rugose, but absent posteriorly; metapleuron remotely punctate, interspaces much wider than diameter of punctures, shiny; mesopleuron above precoxal area (except speculum) punctate and dorsally rugose; scutellum sparsely punctate or punctulate, medio-posteriorly rugulose and with some striae laterally, no carina; propodeum evenly convex and coarsely vermiculate-rugose, mediolongitudinal carina strong in basal 0.6, and without tubercles. Wings. Fore wing: just reaching apex of metasoma; r 0.35 × 3-SR (Fig. 74); 1-CU1 horizontal, 0.45 × 2-CU1; r-m unsclerotized; 2nd submarginal cell mediumsized (Fig. 74); cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR wide. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 72); 2-SC+R subquadrate; m-cu distinct, but unsclerotized and as long as cu-a; M+CU:1-M = 15:9; 1r-m 0.7 × 1-M. Legs. Tarsal claws subpectinate, with six yellowish medium-sized pectinal bristles; hind coxa obliquely striated dorsally, punctulate laterally; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 4.6 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus.

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Figures 81–92. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass 81 wings 82 mesosoma lateral 83 meso­ soma dorsal 84 metasoma dorsal 85 fore femur lateral 86 hind femur lateral 87 inner hind tarsal claw 88 head anterior 89 head dorsal 90 head lateral 91 base of antenna 92 apex of metasoma lateral.

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Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites coarsely longitudinally and densely rugose, robust and posterior corners of 1st protruding outside base of 2nd tergite, with distinct median carina; medio-basal area of 2nd tergite wide and short; 2nd suture moderately deep and crenulate; basal half of 3rd tergite longitudinally striate, remainder of metasoma largely smooth, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, setose and apically truncate (Fig. 73). Colour. Yellowish brown; mesosoma (except mesoscutum, scutellum medially, pronotum anteriorly and dorsally), ovipositor sheath, 3rd tergite (except antero-lateral corners) and following segments black; apical half of antenna, pedicellus, palpi, hind femur apico-dorsally, telotarsi, veins, parastigma basally and pterostigma dark brown; wing membrane rather brownish infuscate. Variation. Antennal segments of ♀ 37(1). Male is largely black, except for 2nd tergite and anterior half of 3rd tergite (Fig. 80). Distribution. Azerbaijan, *Turkey. Notes. Easily confused with A. ruficornis (Herrich-Schäffer); the relative size of the clypeus (wider and somewhat shorter in A. arnoldii than in A. ruficornis) seems to be the main difference in both sexes. In addition, the female of A. arnoldii has the temple ventrally and the malar space yellowish brown (dark brown in A. ruficornis). The male has darker legs and 1st metasomal tergite than the female (the sexes more similar in A. ruficornis). Also reported from Uzbekistan (Yuldashev, 2006); the record from Poland (Huflejt, 1997) most likely concerns A. ruficornis (Herrich-Schäffer). Aleiodes arnoldii sensu Farahani et al. (2015) concerns a species closely related to A. gasterator (Jurine) but has basal half of 3rd tergite coarsely longitudinally rugose, antenna of ♀ with 30–35 segments (of ♂ 36), head linearly narrowed ventrally and subbasal antennal segments of ♀ slightly slenderer. Aleiodes aterrimus (Ratzeburg, 1852) Figs 93–115 Bracon aterrimus Ratzeburg, 1852: 35; Shenefelt, 1978: 1467. Aleiodes aterrimus; Belokobylskij et al., 2003: 398; Zaldivar-Riverón et al., 2004: 234. Aleiodes grandis Giraud, 1857: 178; Papp, 1991a: 77; Bergamasco et al., 1995: 5; Belokobylskij et al., 2003: 398; Papp, 2005: 176 (as valid species) [examined]. Aleiodes (Neorhogas) grandis; Papp, 1985a: 159 (lectotype designation and as synonym of A. aterrimus); Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) aterrimus; Falco et al., 1997: 60. Rogas grandis; Shenefelt, 1975: 1232. Rogas (Rogas) grandis; Tobias, 1976: 87, 1986: 81 (transl.: 134). Rhogas malaisei Shestakov, 1940: 7. Rogas malaisei; Shenefelt, 1975: 1237. Aleiodes malaisei; Shaw et al., 1998: 63 (as synonym of A. grandis Giraud); Belokobylskij et al., 2003: 398 (as synonym of A. aterrimus (Ratzeburg)); Papp, 2005: 176 (as synonym of A. grandis Giraud).

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Rogas (Rogas) vicinus Papp, 1977a: 114, 115 [examined]. Syn. nov. Aleiodes (Neorhogas) vicinis; Papp, 1991a: 78. Type material. Lectotype of A. grandis, ♂ (MNHN), “[Austria:] environs de Vienne”. Holotype of R. vicinus (MTMA), ♀, “Yugoslavia, [Serbia:] Vojvonida, Fruška Gora Mts., Sremska Kamenica, 1–2.v.1972, Papp & Horvatovich”, “Holotypus ♀ Rogas vicinus sp. n., Papp, 1977”, “Hym. Typ. No. 2375, Mus. Budapest”; paratype of R. vicinus, ♀ (MTMA), “[Romania:] Transylvania, Szászkezd%, Silbernagel”, “Paratypus ♀ Rogas vicinus sp. n., Papp, 1977”, “Hym. Typ. No. 2376, Mus. Budapest”; 1 ♂ (MTMA), id., but No. 2375. Additional material. Austria, Belgium, British Isles (England V.C.s 8, 9, 10, 11, 12, 14, 15, 20, 22, 28, 29, 39), Czech Republic, Finland, Germany, Hungary, Netherlands (GE: Brummen (Leuvenheim); LI: Epen; ZH: Schoonrewoerd), Poland, Romania, Russia, Slovakia, Spain, Switzerland. Specimens in BMNH, BZL, CNC, FMNH, HSC, MRC, MSC, MTMA, NMS, OUM, RMNH, SDEI, ZSSM. Molecular data. MRS024 (UK), MRS147 (UK). Biology. Univoltine, spending ca ten months of the year in the exposed mummy on an aerial twig. Collected from April–June, among broadleaved trees (but see paragraph below). Reared from arboreal Amphipyra spp.: A. pyramidea (Linnaeus) (29; M.G. Bloxham, C. Bystrowski, J. Connell, A.P. Fowles, G.M. Haggett, B.T. Parsons, D.L.J. Quicke, M.R. Shaw); A. berbera (Rungs) (5:1 [5 OUM]; G.C. Varley); Amphipyra sp. (8). Some of the forgoing specimens were reared and labelled in the period before it was known that there are two closely related and sympatric arboreal species of Amphipyra in Britain, and it is possible that British records from A. pyramidea (especially when collected on Quercus; cf. Shaw, 1981) have been overstated at the expense of A. berbera; however, both certainly serve as host. An account of frequency at one site is given by Shaw (1981). Before becoming mummified the host moves to a narrow twig, to which the mummy will be very strongly glued. In the early stage of the mummification process (Fig. 99), in which the anterior end of the host is particularly contracted, the parasitoid larva strongly protrudes anteriorly to spread the necessary glue (Fig. 101). The resultant almost semi-circularly domed and hard mummy (Fig. 100), in which the parasitoid occupies approximately abdominal segments 4–7 of the host, forms in ca May–June and persists through the remainder of the summer and the following winter until the adult emerges in ca April–May. (The univoltine hosts overwinter in the egg stage.) The swollen part of the mummy, which is moderately densely lined with silk, is externally usually matt chalky buff in colour, but dark brown diamond-shaped patches centred dorsally on intersegmental areas tend to remain (Fig. 97), and sometimes (perhaps especially when the mummy is unable to dry as it forms) these are coalesced to leave a single shiny dark brown patch covering most of the dorsal surface. Some of the mummies examined might be of somewhat stunted final instar hosts, but others are more clearly penultimate instar. The outcomes of an experiment involving six females and cultured A. pyramidea larvae were unfortunately marred by unavoidably high temperatures and then disease overcoming the cultures so that no mummies resulted, but the following observations

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Figures 93–97. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 93 habitus lateral 94 detail of fore wing with arrow indicating lost vein r-m 95 ovipositor sheath lateral 96, 97 mummies of Amphipyra sp. showing variation in markings.

were made: (i) 2nd to 5th instar hosts were potentially attractive; (ii) 2nd instar hosts were, however, often ignored or else tended to be abandoned after being paralysed with a single jab (i.e., without oviposition subsequently taking place); (iii) 3rd instar hosts were

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Figures 98–100. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 98 parasitised caterpillar of Amphipyra sp. 99 early stage of mummy 100 later stage of mummy.

often ignored, but when attacked seemed the most smoothly parasitised, sometimes with a single paralysing jab being followed, after a short pause, by a single insertion of the ovipositor for presumed oviposition, although the pattern observed for 4th and 5th instars also occurred with 3rd instars; (iv) 4th and (2 only) 5th instar hosts were embraced the most enthusiastically, but it required several (3–5) injections to subdue them, and then there were usually several (3–4) separate sequential and lengthy (often as long as 80 seconds) insertions of the ovipositor (which may or may not all have been actual ovipositions), the parasitoid turning between insertions and always grasping the host

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Figure 101. Larva of Aleiodes aterrimus (Ratzeburg) mummifying Amphipyra pyramidea (Linnaeus), with its anterior (indicated by the arrow) projecting from the ventral opening in the host to spread adhesive over a wide area.

with all six legs during the insertion; (v) antennation of the host was minimal, and there was no post-oviposition association; (vi) all temporarily paralysed hosts hung from the substrate by one or usually more prolegs until they recovered, presumably preventing their falling from their pabulum; (vii) there is no long-term physiological venom effect. The behaviour of adults observed toward the different instars is intriguing, and the experiment would be well worth repeating under better circumstances. Although the above is a consistent pattern for this species, it does not account for a small number of specimens (14 ♀, 4♂ in BZL, MRC, MTMA, NMS, SDEI, ZSSM) examined from various localities in central Europe (Czech Republic, Germany, Hungary) and S. Russia. These specimens share small but rather consistent morphological differences from the usual form, in particular tendencies towards: more intense sculpture on the metasomal tergites (T3 being more or less strongly punctate or even rugose-punctate); the hind wing marginal cell parallel-sided in basal three fifths; shorter 3-SR in fore wing; basal cell of fore wing with more, and more evenly distributed, setae; fewer antennal segments; wing membrane slightly brownish. These differences are not absolutely consistent and would be easy to let pass without comment were it not for the fact that they are correlated with an apparently different phenology, as (of the ten specimens with dates recorded) five ♀ were collected in July and one in August, with only three ♀ in May and one in June (none in April). This is in marked contrast

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with the earlier flight time of the usual form, and the usual hosts (arboreal Amphipyra spp.) are not available after early June. A further ♀ specimen (MTMA) examined and returned in 1997 by MRS but apparently no longer in the main MTMA collection was labelled “Hungaria, Fót, Somlyó-hegy, 30.vii.1958, Ehik”; “Ex Panchrysia deaurata Esp [J. Papp’s handscript]”; “ex Pytometra deautate [sic]”. Unfortunately, no mummy had been preserved, but this plusiine noctuid feeds on Thalictrum (a low plant, not a tree) and it is unlikely for an arboreal Amphipyra, even if fallen from a tree above, to have been mistaken for it. The date, whether referring to collection of the host larva or emergence of the adult parasitoid, is also out of step with arboreal Amphipyra species. We considered but rejected the possibility that these specimens belong to a separate species, and instead conclude that under certain circumstances A. aterrimus can have a partial 2nd brood (in the southern part of its range) which uses different hosts, and that the morphological variation is merely seasonal. The material (which does not conform to A. sapporensis (Watanabe), see below) is being returned to holding institutions determined as A. aterrimus but with “var: T3 sculpture etc.” appended to facilitate recall if necessary. It should be added that this form has (on account of its heavy metasomal sculpture and extensively parallel-sided marginal cell in the hind wing) sometimes been misidentified as A. rugulosus, but the two species are always easily separated by the sculpture of the mesoscutum and scutellum, as well as by leg colour. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 109); ventral margin of clypeus obtuse apically and not protruding outwards (Fig. 111); OOL of ♀ distinctly longer than diameter of posterior ocellus; meso­ scutal lobes densely and finely punctate-coriaceous, rather matt; scutellum densely and finely coriaceous; mesosternal sulcus shallow, obsolescent or absent; vein 1-CU1 of fore wing 0.2 × vein 2-CU1 (Fig. 102); vein 2-SC+R of hind wing subquadrate or vertical (Fig. 102); hind tarsal claws with conspicuous and robust blackish pecten (Fig. 113); head black; hind tibia largely to completely black; metasoma of both sexes black; 4th– 6th tergites of ♂ flat and densely short setose, except a narrow glabrous strip centrally. Dr K. Samartsev (in litt.) kindly brought to the first author’s attention that the East Palaearctic A. sapporensis (Watanabe, 1937) occurs in southern European Russia (Middle and Lower Volga territories). Aleiodes aterrimus and A. sapporensis differ only slightly, mainly by the colour of the extreme base of the hind tibia (completely dark brown in A. sapporensis and usually narrowly pale yellowish in A. aterrimus) and by the shape of temple in dorsal view (roundly narrowed in A. sapporensis and rather linearly narrowed in A. aterrimus). There is also a slight difference in the proportions of the face (A. sapporensis has facial width 1.50–1.60 × medial height including clypeus and A. aterrimus 1.65–1.75 ×). A. sapporensis seems to have the lateral carinae of propodeum more protruding and has 58–66 antennal segments. Description. Redescribed ♀ (RMNH) from England (Pamber Forest). Length of fore wing 7.3 mm, of body 8.6 mm. Head. Antennal segments of ♀ 59, length of antenna 1.1 × fore wing, its subapical segments rather robust; frons largely superficially granulate; OOL 1.8 × diameter of posterior ocellus, and superficially rugulose-granulate and shiny; vertex superficially

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Figures 102–115. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest, but 102 from Austria, Wien 102 fore wing 103 hind wing 104 mesosoma lateral 105 mesosoma dorsal 106 metasoma dorsal 107 fore femur lateral 108 hind femur lateral 109 head anterior 110 head dorsal 111 head lateral 112 hind tibia and tarsus lateral 113 outer hind tarsal claw 114 base of antenna 115 apex of antenna.

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rugulose-granulate, rather shiny; clypeus with some punctures; ventral margin of cly­ peus thick and not protruding forwards (Fig. 111); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 109); length of eye 1.3 × temple in dorsal view (Fig. 110); vertex behind stemmaticum superficially granulate-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate-coriaceous, rather matt; precoxal area of mesopleuron largely smooth medially, densely punctate anteriorly and posteriorly; metapleuron densely punctate; metanotum with nearly complete median carina; scutellum punctate-coriaceous; propodeum rather convex and coarsely reticulate-rugose, medio-longitudinal carina nearly complete, and with slightly protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 102); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 102); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 103); 2-SC+R short and vertical; m-cu absent; M+CU:1M = 12:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws with conspicuous and robust blackish pecten (Fig. 113); hind coxa largely densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 106); 2nd suture deep and narrow; basal half of 3rd tergite finely punctate-rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 95). Colour. Black; antenna (except scapus and pedicellus), palpi, tegulae, fore and middle telotarsi, veins and pterostigma dark brown; coxae, trochanters and trochantelli, apical third of hind femur (ventrally extended to its apical two-thirds), hind tibia (except pale yellowish basal ring) and hind tarsus black, remainder of legs yellowish brown; wing membrane subhyaline. Variation. Hind femur usually only apically dark brown, but sometimes entirely dark brown; coxae black or sometimes largely yellowish brown. Two females (both NMS, from different localities) have vein r-m of fore wing absent. Males are very similar, apical tergites type 3, with fringe very weak to negligible; hind femur often only apically blackish, but sometimes up to apical 0.6 darkened. Antennal segments: ♀ 57(3), 58(1), 59(5), 60(7), 61(3), 62(6), 63(5), 64(1); ♂ 51(1), 52(1), 53(3), 54(1), 55(5), 56(7), 57(6), 58(5), 59(3), 60(1), 62(2); females have on average ca four more antennal segments than males. The antennal segments for the specimens of the abnormal series (see above) are scored separately here: ♀ 54(2), 55(1), 56(2), 57(3), 58(3), 60(1); ♂ 54(1).

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Distribution. Austria, *Belgium, British Isles (England), Czech Republic, *Finland, Germany, Hungary, *Netherlands, Poland, *Romania, Russia, Serbia, Slovakia, Spain, *Switzerland. New synonymy. The synonymy of Rogas vicinus Papp, 1977, with Aleiodes aterrimus (Ratzeburg, 1852) is based on the examination of the types listed above. The differences between R. vicinus and R. grandis (= A. aterrimus) listed in the original description (head less constricted posteriorly, apical antennal segments more robust, 1st metasomal tergite less robust and 2nd tergite somewhat longer) fall within the normal variation of A. aterrimus. Aleiodes carbonarius Giraud, 1857 Figs 116–137 Aleiodes carbonarius Giraud, 1857: 177–178 [examined]. Aleiodes (Neorhogas) carbonarius; Papp, 1985a: 156 (lectotype designation), 1991a: 88. Aleiodes carbonarius; Papp, 2005: 176. Rogas carbonarius; Shenefelt, 1975: 1220–1221. Rhogas (Rhogas) carbonarius ab. giraudi Fahringer, 1931: 236; Shenefelt, 1975: 1221 (invalid name). Type material. Lectotype of A. carbonarius, ♂ (MNHN), “Hunga[ry]”, “2”, “Hungaria”, “Neusiedlersee/teste Papp J., 1979”, “Lectotypus”, “Aleiodes carbonarius Gir., 1857, ♂, Papp, 1979”. Paralectotype ♂ (MNHN) from Austria (near Vienna). Additional material. 3 ♀ (NMS), “Hungary: Veszprém, nr Tótvázsony, larva coll. 21.v.2001, Tholera decimalis, mum. c. 12.vi.[20]01, em. 19.v., 24.v. and 25.v.[20]02, M.R. Shaw”; 1 ♂ (MSC), “A[ustria], Oberösterreich, Wels, Flughaven, 48°10'N, 14°2'E, 30.iv.2012, M. & J. Schwarz”; 1 ♂ (MTMA), “Hungaria, Csákvár”, “Vértes Hgs., Hajduvágás”, “12.v.1961, Sólymosné”, “Rogas carbonarius Gir. ♂, det. Papp, 1979 / compared with lectotype ♂”; 1 ♂ (NMS), “[Hungaria,] P. Szt. Lelek, Ujhelyi”, “Rogas morio Reinh. ♂, det. Szépligeti”, “Rogas carbonarius Gir. ♂, det. Papp, 1979”; 1 ♂ (MTMA), id., but Budapest, Svabhegy; 2 ♂ (MRC) “Russia, E. Siberia Lake Baikal, Biakalo-Lenskiy res. 20.vi. and 19.vii. [19]05, leg. Berlov”; 1 ♂ (BZL), “CSR [Czech Rep.], envir. Prague, 1968, Dr. Pádr”. This species appears to be sporadic in central and eastern Europe. The specimens from which Morley (1937) recorded this species as new to Britain have been examined and belong to A. carbonaroides sp. nov. Molecular data. MRS162 (Hungary), MRS163 (Hungary), MRS 164 (Hungary). Biology. Adults of this lowland species have been collected from the very end of April to July (see also Papp, 1999), and it is found in grassland habitats. Reared from the noctuid Tholera decimalis (Poda) (3:1; M.R. Shaw/Hungary). The decidedly large mummy is very similar to that of A. grassator and forms underground (Fig. 118). Univoltine, overwintering in the mummy.

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Figures 116–118. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 116 habitus lateral 117 ovipositor sheath lateral 118 mummy of Tholera decimalis (Poda).

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Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 126); OOL of ♀ ca 2.6 × as long as diameter of posterior ocellus (Fig. 127) and distinctly rugose; length of 4th antennal segment of ♀ ca 0.9 × its width (Fig. 129; in ♂ 0.9–1.0 times; Fig. 135); clypeus thick apically and not protruding anteriorly (Fig. 128); lobes of mesoscutum densely punctate, interspaces superficially granulate and with satin sheen; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending near level of apex of vein 3-M (Fig. 119); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 119); vein 3-SR of ♀ 1.7–2.0 × as long as vein 2-SR; vein 3-SR ca 0.7 × vein SR1 (Fig. 119; of ♂ ca 0.5×); hind tarsal claws yellowish or brownish bristly setose (Fig. 131); inner side of hind tibia of ♀ yellowish; tegulae yellowish brown; 4th and 5th tergites black. Probably a lowland species in C. Europe. Description. Redescribed ♀ (NMS) from Hungary (Veszprém). Length of fore wing 4.1 mm, of body ca 6.0 mm. Head. Antennal segments of ♀ 46, 4th segment 0.9 × longer than wide (Fig. 129); length of antenna 1.15 × fore wing, its subapical segments robust (Fig. 130); frons with coarse curved rugae and rather shiny; OOL 2.6 × diameter of posterior ocellus and rugulose; vertex rugose and shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 128); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 126); length of eye 1.4 × temple in dorsal view (Fig. 127); vertex behind stemmaticum rugose; clypeus distinctly below lower level of eyes; length of malar space 0.7 × length of eye in lateral view (Fig. 128). Mesosoma. Mesoscutal lobes densely punctate, interspaces superficially granulate and with satin sheen; precoxal area of mesopleuron coarsely rugose medially and punctate posteriorly; remainder of mesopleuron mainly coarsely punctate; scutellum flat, sparsely finely punctate and with lateral carina; propodeum coarsely rugose, mediolongitudinal carina indistinct, rounded posteriorly and dorsal part rather short. Wings. Fore wing: r 0.4 × 3-SR; marginal cell ends near level of apex of 3-M (Fig. 119); 1-CU1 horizontal and slightly widened, 0.45 × 2-CU1; r-m 0.3 × 3-SR; 2nd submarginal cell elongate (Fig. 119), 3-SR twice as long as 2-SR; cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR slender and medium-sized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 120); 2-SC+R slightly longer than wide; m-cu short, postfurcal; M+CU:1-M = 61:36; 1r-m 0.75 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 131); hind coxa largely rather densely punctate; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 4.5 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 0.9 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and short (Fig. 123); 2nd suture deep and crenulate; basal third of 3rd tergite finely longitudinally striate, remainder of metasoma superficially microsculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 117).

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Figures 119–131. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 119 fore wing 120 hind wing 121 mesosoma lateral 122 mesosoma dorsal 123 metasoma dorsal 124 fore femur lateral 125 hind femur lateral 126 head anterior 127 head dorsal 128 head lateral 129 base of antenna 130 apex of antenna 131 inner hind tarsal claw.

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Figures 132–137. Aleiodes carbonarius Giraud, ♂, Hungary, Csákvár 132 habitus lateral 133 head dorsal 134 wings 135 base of antenna 136 apex of antenna 137 head anterior.

Colour. Dark orange brown; apical two-thirds of antenna, patch on hind femur dorso-apically, and telotarsi, dark brown; temple ventrally, malar space, mesosternum, mesopleuron, metapleuron, propodeum, pair of patches on 2nd tergite and most of apical 0.4 of tergite, and 3rd–7th tergites black; palpi (especially labial palp), veins and ptero­ stigma dark brown, basal third of antenna (but scapus dorsally blackish) rather pale yellowish brown; tegulae and remainder of legs; yellowish brown; wings strongly infuscate. Variation. Antennal segments: ♀ 46(2), 49(1); ♂ 47(1), 50(1), 52(1), 54(1), 56(1), 57(1); length of fore wing of ♀ ca two-thirds of body length (0.8 × in ♂); males

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always darker than females; mainly black with legs mainly dark brown or blackish, but male from Austria has basal half of metasoma orange brown and legs partly yellowish brown. Males have 2nd submarginal cell distinctly shorter than in females (as in A. grassator), antenna 0.9 × length of body and slightly less robust subapically, temple and face long setose and malar space 0.5–0.7 × length of eye in lateral view; metasoma black or 1–2 basal tergites reddish and apical tergites type 1, fringe not observed (Fig. 132). Distribution. Austria, Czech Republic, Hungary, *Russia (Lake Baikal). Notes. Very similar to A. grassator (Thunberg), and especially A. carbonaroides; males of A. carbonarius and carbonaroides are normally black but males with partly orange brown metasoma occur. The three species exhibit sexual dimorphism of the 2nd submarginal cell (less robust (and also longer in A. carbonarius) in female than in male). Giraud (1857) gave an incomplete description of the only two males he possessed, but clearly indicated that the antenna is slightly shorter than the body. The female of this species is reported for the first time. Aleiodes carbonaroides van Achterberg & Shaw, sp. nov. http://zoobank.org/0BE2C69B-E310-4DFB-BE5C-07218AC6F018 Figs 138–160 Type material. Holotype, ♀ (NMS), “[Netherlands: Friesland], Holland [sic!], Schier­ monnikoog, em. 20.v.[19]82”, “ex Cerapteryx graminis larva”. Paratypes: 2 ♀ (NMS, RMNH), 3 ♂ (NMS, RMNH), topotypic and from same host, em. 19 or 20.v.1982; 1 ♂ (ZSSM) “[Germany], Münehey, 26.iv.[18]85 R7”, “1-653”; 2 ♂ (CMIM) “[England] 25.v.[19]22, Bdn. [= Brandon, Suffolk] HF”, “Named by Claude Morley 2 Rhogas carbonarius Giraud. NEW TO BRIT. CM V.22”; 1 ♂ (BMNH) “[England], Totternhoe, [Bedfordshire], 30.v.[19]64 [V.H. Chambers]”. Sporadic in western Europe. Molecular data. None. Biology. Adults of this lowland species have been collected in April and May. The two paratypes from Suffolk were swept from Breck grassland (Morley, 1937, misidentified as A. carbonarius). Reared from the grass-feeding noctuid Cerapteryx graminis (Linnaeus) (6 [2 are RMNH]; K.P. Carl/Netherlands). If it is a specialist, it is presumably univoltine and overwinters in the mummy (the univoltine known host overwinters in the egg stage). Mummy similar to that of the closely related A. carbonarius and A. grassator, but slightly smaller. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 149); OOL of ♀ 1.8–2.0 × as long as diameter of posterior ocellus (Fig. 150) and distinctly rugose or rugulose; length of 4th antennal segment of ♀ 0.7–0.9 × its width (Fig. 152; in ♂ up to 1.0 times); clypeus thick apically and not protruding anteriorly (Fig. 151); lobes of mesoscutum punctate, interspaces largely coriaceous and superficially coriaceous; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 142); vein 1-CU1 of

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Figures 138–141. Aleiodes carbonaroides sp. nov., ♀, holotype 138 habitus lateral 139 ovipositor sheath lateral 140 outer hind tarsal claw lateral 141 mummy of Cerapteryx graminis (Linnaeus).

fore wing 0.5–0.6 × as long as vein 2-CU1 (Fig. 142); 2nd submarginal cell of fore wing medium-sized (Fig. 142); hind tarsal claws slender and yellowish or brownish bristly setose; hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; head and mesoscutum of ♀ reddish; palpi and tegulae of ♀ brownish yellow; males entirely black, with palpi, tegulae and antenna dark brown or blackish. Description. Holotype, ♀, length of fore wing 4.2 mm, of body 7.1 mm. Head. Antennal segments of ♀ 45, 4th segment 0.9 × longer than wide (Fig. 152); length of antenna 1.1 × fore wing, its subapical segments robust (Fig. 153) and scapus oblique apically; frons with coarse curved rugae and shiny; OOL 1.8 × diameter of posterior ocellus and rugulose; vertex rugose and shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 151); width of hypo­ clypeal depression 0.4 × minimum width of face (Fig. 149); length of eye 1.2 × temple in dorsal view (Fig. 150); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view.

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Figures 142–153. Aleiodes carbonaroides sp. nov., ♀, holotype 142 fore wing 143 hind wing 144 mesosoma lateral 145 mesosoma dorsal 146 metasoma dorsal 147 fore femur lateral 148 hind femur lateral 149 head anterior 150 head dorsal 151 head lateral 152 base of antenna 153 apex of antenna.

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Mesosoma. Mesoscutal lobes moderately punctate, interspaces superficially granulate-coriaceous and with satin sheen; precoxal area of mesopleuron coarsely rugose medially, but largely smooth posteriorly; remainder of mesopleuron mainly punctate; scutellum flat, sparsely finely punctate and with irregular lateral carina; propodeum coarsely rugose, medio-longitudinal carina complete, rounded posteriorly and dorsal part approx. as long as posterior part. Wings. Fore wing: r 0.4 × 3-SR (Fig. 142); marginal cell ends basad of level of apex of 3-M; 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell robust (Fig. 142), 3-SR 1.4 × as long as 2-SR; cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR similar to 1-M and medium-sized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 1.7 × width at level of hamuli (Fig. 143); 2-SC+R subquadrate; m-cu short; M+CU:1-M = 27:15; 1r-m 0.7 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 140); hind coxa largely rugulose dorsally; hind trochantellus robust; length of hind femur and basitarsus 3.2 and 4.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather flattened, 0.7 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite without medio-longitudinal carina; medio-basal area of 2nd tergite triangular and short; 2nd suture deep and crenulate; basal half of 3rd tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured; 4th and apical third of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 139). Colour. Dark orange brown; apical half of antenna, patch on hind femur dorsoapically, and telotarsi apically, dark brown; mesosternum, mesopleuron (except dorsally and postero-ventrally), metapleuron (except medio-dorsally), propodeum (except pair of posterior patches), 3rd–7th tergites (except antero-lateral corners of 3rd tergite) black; palpi, basal half of antenna, tegulae and remainder of legs rather pale yellowish brown; veins and pterostigma dark brown; wings strongly infuscate but hind wing less than fore wing. Variation. Basal third or half of antenna of ♀ pale yellowish brown; vein 3-SR 1.4–1.6 × as long as vein 2-SR; hind femur of ♀ 3.2–3.5 × longer than wide; 1st metasomal tergite 0.7–0.8 × its apical width; temple and occiput ventrally, and malar space ventrally orange brown or black. Antennal segments: ♀ 43(1), 45(1); ♂ 48(1), 49(2), 51(1), 50(1), 53(2); males clearly have many more antennal segments than females. Males are much darker than females; body black with palpi and legs mainly dark brown or blackish (Fig. 154). Males have 2nd submarginal cell slightly smaller than females (Fig. 158), temple and face long setose, malar space 0.5–0.7 × length of eye in lateral view, and apical tergites type 1 and fringe not observed (Fig. 154); sometimes superficial granulosity of 3rd tergite and of mesoscutum are absent. Distribution. Germany, Netherlands, U.K. Etymology. The suffix “-oides” indicates similar to; in this case the high similarity to A. carbonarius Giraud.

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Figures 154–160. Aleiodes carbonaroides sp. nov., ♂, paratype 154 habitus lateral 155 apex of antenna 156 antenna 157 base of antenna 158 wings lateral 159 head anterior 160 head dorsal.

Aleiodes caucasicus (Tobias, 1976) Figs 161–177 Rogas (Rogas) caucasicus Tobias, 1976: 86, 222, 1986: 81 (transl.: 133) [examined]. Aleiodes (Neorhogas) caucasicus; Papp, 1985a: 152. Aleiodes caucasicus; Papp, 1991a: 75 (as synonym of A. fortipes), 2005: 176 (id.); Fortier & Shaw, 1999: 227; Žikić et al., 2002: 108; Aydogdu & Beyarslan, 2005: 191. Type material. Holotype, ♀ (ZISP), “[Russia], Sotchi, Lazarevskoe [terras], 26.iv.[1]973, V. Tobias”, “Holotypus Rogas caucasicus Tobias”; 2 ♀, paratype (MTMA), id., but 29.iv.1973.

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Additional material. Figured ♀ (NMS), “[Russia], Sotchi, Lazarevskoe terras. Sklony, les [= forest], 25.iv.1988, V. Tobias”, “Rogas caucasicus Tob.”, “Aleiodes caucasicus (Tobias), det. Belokobylskij, 2005. ♀ Ant. 40”; 2 ♀ (ALC, RMNH), id., but 7.v.1975; 1 ♀ (MTMA), “Bulgaria”, “Rhodopi, St[ara] Zagora, 17.iv.1977, J. Kolarov”, “Rogas sp. n.?, det. Zaykov, 1983”, “Aleiodes fortipes Rh. ♀, det. Papp J., 1985”. Molecular data. None. Biology. Unknown. Specimens collected in April-May and flight time probably April–May. We have not seen reared material. Probably, like A. fortipes, it will be found to be univoltine, overwintering in the mummy, but direct evidence is lacking. Diagnosis. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 171); antenna of ♀ with 38–41 segments and 2nd – 10th antennal segments yellowish, contrasting with remaining segments; OOL coarsely transversely striate; clypeus obtuse apically and not protruding in lateral view (Fig. 173); precoxal area finely striate (Fig. 166); tegulae yellow; lobes of mesoscutum finely coriaceousgranulate and rather dull, with satin sheen; vein 1-CU1 of fore wing much shorter than vein 2-CU1 (Fig. 164); posteriorly vein m-cu of fore wing diverging from anterior half of vein 1-M; length of hind femur 3.6–3.8 × its maximum width (Fig. 170); hind tarsal claws brownish setose (Fig. 177); length of fore wing 3.7–5.0 mm. Very similar to A. fortipes (Reinhard) and differs mainly by its body colour and sculpture of mesopleuron. Description. Holotype, ♀, length of fore wing 3.7 mm, of body 4.6 mm. Head. Antennal segments of ♀ 41, length of antenna 1.3 × fore wing, its subapical segments rather robust; frons largely finely rugulose medially; OOL 2.2 × diameter of posterior ocellus, and coarsely transversely striate; vertex transversely striate and rather shiny; clypeus rugulose, but ventrally depressed and smooth; ventral margin of clypeus thick and not protruding forwards (Fig. 173); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 171); length of eye twice temple in dorsal view (Fig. 172); vertex behind stemmaticum rugulose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view; occipital carina largely absent dorsally and weakly developed ventrally. Mesosoma. Mesoscutal lobes largely rugulose-granulate, rather matt; precoxal area of mesopleuron transversely striate medially, distinctly rugose antero-dorsally and remainder largely punctulate; pleural sulcus moderately crenulate (Fig. 166); ventral half of metapleuron rugose; metanotum with nearly complete median carina; scutellum coriaceous; propodeum densely and finely granulate-rugose and medio-longitudinal carina medium-sized. Wings. Fore wing: r 0.6 × 3-SR; 1-CU1 horizontal, 0.5 × 2-CU1; r-m unsclerotized and 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 164); cu-a vertical, straight and rather short; 1-M slightly curved posteriorly; posteriorly vein m-cu diverging from anterior half of vein 1-M. Hind wing: marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 165); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 5:3; 1r-m 0.7 × 1-M.

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Figures 161–163. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 161 habitus lateral 162 ovipositor sheath lateral 163 apex of antenna (of paratype).

Legs. Tarsal claws robust and with brownish bristles (Fig. 177); hind coxa densely rugulose and rather dull; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 5.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, 0.9 × longer than wide apically; 1st and 2nd tergites with indistinct medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 168); 2nd suture rather shallow and crenulate; medio-basally 3rd tergite striate, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 162). Colour. Orange brown; head, 3rd tergite (except antero-laterally) and subsequent tergites black; scapus, pedicellus basally, 11th and following antennal segments, palpi, veins, parastigma, pterostigma and femora apico-dorsally, tibia and tarsal segments apically, ventral half of metasoma and ovipositor sheath dark brown; tegulae, 3rd–10th antennal segments brownish yellow; wing membrane subhyaline. Variation. Head black or mainly dark brown, specimen from Bulgaria also anterior half of mesosoma; antenna of ♀ with 38 or 41 segments according to the original description; 11th and 12th antennal segments of ♀ dark brown or brownish yellow;

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Figures 164–177. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 164 fore wing 165 hind wing 166  mesosoma lateral 167 mesosoma dorsal 168 metasoma dorsal 169 fore femur lateral 170 hind femur lateral 171 head anterior 172 head dorsal 173 head lateral 174 base of antenna 175 antenna 176 hind tibia and tarsus lateral 177 outer hind tarsal claw.

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hind femur 3.6–3.8 × as long as wide. The male is unknown, or possibly has not been distinguished from that of A. fortipes. Distribution. *Bulgaria, Russia (SW). Notes. It remains unclear whether this predominantly rather yellowish orange species is distinct from A. fortipes, which in its more western localities is a much darker insect. Females intermediate in colour (and included in A. fortipes) seem to predominate in eastern Europe. More material (preferably with biological data) is needed to clarify the status of A. caucasicus. Aleiodes coriaceus van Achterberg & Shaw, sp. nov. http://zoobank.org/EA99A74A-AA7C-460F-85F5-AC7405FE67B9 Figs 178–195 Type material. Holotype, ♀ (NMS), “Sweden: Hr, Sveg, Duybergshammaren, 17.vii.2004, N. Ryholm, NMSZ 2004.167”, “MRS Aleiodes DNA 377”, “COI worked”. Paratypes: 1 ♂ (NMS), same label data as holotype; 1 ♀ (RMNH), “Sweden: Ås. Lilla, Vammasj. Window trap on Betula F2, 8, vii.2003, J. Hilszczanski”, “MRS Aleiodes DNA 311”, “COI worked”. Molecular data. MRS311 (Sweden), MRS377 (Sweden). Biology. Unknown. The available specimens were collected in July, and it is almost certainly univoltine, but we have not seen reared material. Diagnosis. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Fig. 186); OOL of ♀ 0.9–1.1 × as long as diameter of posterior ocellus (Fig. 187), and rugulose-coriaceous or only coriaceous; ventral margin of clypeus rather thin or blunt and not protruding forwards (Fig. 188); vertex mainly coriaceous and rather dull; mesoscutal lobes coriaceous and largely matt; scutellum remotely punctate; area of precoxal sulcus largely smooth, with some punctulation; length of vein 1-CU1 of fore wing 0.3–0.4 × vein 2-CU1 and 0.4–0.5 × vein m-cu; marginal and 2nd submarginal cells of fore wing elongate (Fig. 180); tarsal claws with robust apical tooth and with medium-sized dark brown pecten (Fig. 190); hind femur and basitarsus slender (Figs 178, 185); 1st metasomal tergite comparatively steep anteriorly (Fig. 178); basal half of 3rd tergite with posteriorly diverging rugulae; head black; dorsal half of hind femur largely black dorsally; basal half of hind tibia largely dark brown; fore and middle trochanters and trochantelli infuscate or dark brown; 2nd tergite yellowish or reddish and rather slender (Fig. 183); 5th–7th tergites of ♂ medially glabrous and convex, and laterally with long setae (Figs 194, 195). Closely related to A. rufipes (Thomson) and differs mainly by the sculpture of the mesoscutum (matt instead of rather shiny), darker colour of legs, different COI and less robust 2nd and 3rd metasomal tergites. Description. Holotype, ♀, length of fore wing 6.1 mm, of body 6.7 mm. Head. Antennal segments of ♀ 54, antenna 1.1 × as long as fore wing, its basal segments robust, subapical segments medium-sized and apical segment with spine; frons largely smooth, except for some micro-sculpture; OOL 0.9 × diameter of poste-

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Figures 178, 179. Aleiodes coriaceus sp. nov., ♀, holotype 178 habitus lateral 179 ovipositor sheath lateral.

rior ocellus, rugulose-coriaceous and rather dull, groove beside posterior ocellus deep and smooth; vertex coriaceous with some rugulae, rather dull; face transversely rugose; clypeus densely rugulose; ventral margin of clypeus thin and not protruding forwards (Fig. 188); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 186); length of eye 2.1 × temple in dorsal view (Fig. 187); vertex behind stemmaticum coriaceous; clypeus partly above lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely coriaceous and matt; precoxal area of mesopleuron partly remotely punctulate and superficially micro-sculptured; medio-longitudinal carina of metanotum distinct posteriorly; scutellum punctate and with lateral carina; propodeum convex and rugose, medio-longitudinal carina absent posteriorly, and without protruding carinae laterally. Wings. Fore wing: r 0.35 × 3-SR (Fig. 180); 1-CU1 slightly oblique, 0.35 × 2-CU1; r-m 0.4 × 3-SR; 2nd submarginal cell long (Fig. 180); cu-a slightly inclivous, straight but posteriorly slightly curved; 1-M nearly straight posteriorly; 1-SR widened; surroundings of M+CU1, 1-M and 1-CU1 densely setose. Hind wing: marginal cell linearly widened, its apical width 2.3 × width at level of hamuli (Fig. 180); 2-SC+R slightly longer than wide; m-cu absent; M+CU:1-M = 50:46; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized dark brown pecten (Fig. 189); hind coxa (except depression) coriaceous and with some rugulae dorsally;

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Figures 180–191. Aleiodes coriaceus sp. nov., ♀, holotype 180 wings 181 mesosoma lateral 182 mesosoma dorsal 183 propodeum and 1st–3rd metasomal tergites dorsal 184 fore femur lateral 185 hind femur lateral 186 head anterior 187 head dorsal 188 head lateral 189 outer hind tarsal claw 190 base of antenna 191 apex of antenna.

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Figures 192–195. Aleiodes coriaceus sp. nov., ♂, paratype 192 habitus lateral 193 inner hind claw lateral 194 3rd–7th metasomal tergites dorsal 195 3rd –7th metasomal tergites lateral.

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hind trochantellus robust and with long setae; length of hind femur and basitarsus 4.5 and 5.8 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex and basally rather steep, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and longitudinally rugose; maximum width of 2nd tergite 1.5 × its median length; medio-basal area of 2nd tergite mediumsized triangular and rather short (Fig. 183); 2nd suture distinct and moderately crenulate; basal half of 3rd tergite finely rugulose and rugulae diverging posteriorly, remainder of metasoma nearly smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 179). Colour. Black; mesoscutum posteriorly, legs (but fore and middle telotarsi, fore and middle femora basally and apically, fore and middle trochanters and trochantelli, hind tarsus dark brown or infuscate, posterior half of hind femur dorsally and hind tibia largely blackish), propodeum and 1st –3rd metasomal tergites (but posterior half of 3rd tergite blackish posteriorly) reddish brown; tegulae brownish yellow, but humeral plate largely dark brown; palpi, pterostigma and veins dark brown; wing membrane slightly infuscate. Variation. Antennal segments: ♀ 52(1), 54(1); ♂ 53(1). Length of fore wing 5.3– 6.1 mm. Male is very similar to female (Figs 195–195). Apical tergites of male type 1–2, and fringe scarcely visible in the single male seen. Distribution. Sweden. Etymology. Coriaceus is Latin for leathery, because of the coriaceous sculpture of vertex and mesoscutum. Aleiodes cruentus (Nees, 1834) Figs 196–216 Rogas cruentus Nees, 1834: 212; Shenefelt, 1975: 1222; Zaykov, 1980a: 112; Kotenko, 1992: 96. Rogas (Rogas) cruentus; Tobias, 1976: 85, 1986: 80 (transl.: 130). Aleiodes (Neorhogas) cruentus; Papp, 1985a: 156–157 (neotype designation), 1987b: 35, 1991a: 83; Belokobylskij, 1996: 6; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) cruentus; Chen & He, 1997: 39; Belokobylskij, 2000: 32. Aleiodes cruentus; Bergamasco et al., 1995: 5; Belokobylskij et al., 2003: 398; Papp, 2005: 176. Rhogas cruentus ab. nigricans Fahringer, 1932: 238; Papp, 1991a: 83 (invalid name). Rhogas cruentus ab. basalis Hellén, 1927: 22 (invalid name). Rhogas cruentus ab. nigromaculata Hellén, 1927: 22 (invalid name). Rhogas cruentus ab. rufofasciata Hellén, 1927: 22 (invalid name). Rogas dorsalis Herrich-Schäffer, 1838: 154; Shenefelt, 1975: 1222 (as synonym of A. cruentus); Papp, 2005: 176 (id.). Rogas affinis Herrich-Schäffer, 1838: 124 (key only); Shenefelt, 1975: 1174–1175 [neotype designated below]. Syn. nov. Aleiodes affinis; Belokobylskij et al., 2003: 398.

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Type material. Neotype of A. affinis here designated, ♀ (RMNH), “Museum Leiden, Nederland, Melissant (ZH), [at light], 10.viii.1980, K.J. Huisman”. It is important for nomenclatorial stability to fix our interpretation of A. affinis because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; the first author could not find any specimen in ZMB), the original description is rudimentary and there are very similar species in Europe. The specimen from Netherlands is selected because it fits best the original description, Netherlands is relatively close to the probable German (but unknown) type location and it is in good condition. Another complication is that the neotype of A. cruentus by Papp (1985) is an old male from uncertain origin in the Gravenhorst Collection (Wroclaw). Additional material. Austria, Bulgaria, Croatia, Czech Republic, Finland, France, Germany, Greece, Italy (including Sicily), Moldova, Netherlands (FR: Ried, GE: Beusichem; Heerde; Voorst (Twello), LI: Thorn, NB: Eindhoven; Tilburg (Kaaistoep), OV: Buurse; Hasselt, ZH: Lexmond; Melissant; Middelharnis; Oostvoorne, ZL: Oostkapelle), Norway, Romania, Slovakia, Slovenia, Spain, Sweden, Ukraine, [Mongolia]. Specimens in BMNH, BZL, FMNH, HSC, IKC, MSC, MTMA, NMS, NRS, RMNH, SDEI, ZSSM. Widespread in the region but rather sporadic. The specimen (CMIM) from which Morley (1915) recorded this species as new to Britain has been examined and proves to be A. alternator (Nees). A further specimen in CMIM recorded by Lyle (1919) as A. cruentus has been examined and belongs to A. diversus (Szépligeti), q. v., as do another three British specimens in BMNH and one in NMS, and there is no evidence that A. cruentus has ever occurred in Britain. Biology. Probably univoltine, certainly overwintering as a mummy. Collected June-August, often at light and including around Dianthus barbatus harbouring larvae of the noctuid Hadena confusa (Hufnagel) (H. Schnee/Germany). In Austria it has been collected up to 2000 m. Only one reared specimen seen, from H. confusa [FMNH], the adult emerging in June in the year following host mummification. Extensive rearings of this host in various parts of Britain in recent years by one of us (MRS) has not produced A. cruentus, strengthening the view that it does not occur in Britain. The predominantly dark mummy seen (Fig. 198) is stout, rather short and weakly swollen dorsally, and has a paler and moderately strong lateral keel. The cocoon is substantially silk-lined and occupies most of the host’s abdomen (approx. 2nd–7th abdominal segments). The mummy probably forms underground, albeit from penultimate instar hosts, and the somewhat reflexed and sideways twisted head suggests that it is not or scarcely stuck down; the caudal segments are also somewhat recurved ventrally. Although oviposition has not been witnessed, the somewhat laterally compressed apex of the female’s metasoma appears to be an adaptation for attacking the host at rest or feeding within the seed capsules of its food plants (Dianthus, Silene, etc.). Molecular data. MRS558 (France), MRS624 (Germany), MRS625 (Germany). Diagnosis. Maximum width of hypoclypeal depression (0.5–)0.6–0.7 × minimum width of face (Fig. 206); OOL of ♀ coarsely punctate and 0.5–0.8(–1.0) × diameter of posterior ocellus; ventral margin of clypeus (rather) obtuse apically and not protruding (Fig. 208), but sometimes intermediate; length of eye 1.5–1.9 × temple in dorsal view;

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Figures 196–198. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg, but 198 from Finland, Mäntyharju 196 habitus lateral 197 ovipositor sheath lateral 198 mummy of Hadena confusa (Hufnagel).

lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures; precoxal area with some rugae medially; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein r of fore wing 0.3– 0.4 × vein 3-SR (Fig. 199); vein 1-CU1 of fore wing 0.8–1.1 × vein 2-CU1 (Fig. 199), rarely shorter; hind tarsal claws with conspicuous dark brown pecten (Fig. 205); 1st tergite widened apically; 2nd tergite 0.7–0.9 × as long as wide (Fig. 202), its colour variable,

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Figures 199–211. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg 199 wings 200 mesosoma lateral 201 mesosoma dorsal 202 metasoma dorsal 203 fore femur lateral 204 hind femur lateral 205 outer hind tarsal claw 206 head anterior 207 head dorsal 208 head lateral 209 base of antenna 210 apex of antenna 211 hind tarsus lateral.

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often reddish; head black; vein 1-M of fore wing brownish; wing membrane subhyaline; 4th–6th tergites of ♂ with long setae, but flattened and narrowly glabrous medially. Description. Neotype of A. affinis, ♀, length of fore wing 7.3 mm, of body 10.2 mm. Head. Antennal segments of ♀ 61, length of antenna 1.2 × fore wing, its subapical segments rather robust; frons largely smooth and shiny, but rugulose near stemmaticum; OOL 0.6 × diameter of posterior ocellus, and coarsely punctate, interspaces approx. equal to diameter of punctures; vertex mainly densely punctate, shiny; clypeus coarsely punctate-rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 208); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 206); length of eye 1.9 × temple in dorsal view and temple rather long and densely setose (Fig. 207); vertex behind stemmaticum punctate-rugose; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view (Fig. 208). Mesosoma. Mesoscutal lobes densely and finely punctate, with satin sheen; precoxal area of mesopleuron with some rugae medially, rather densely punctate anteriorly and posteriorly; metapleuron mainly sparsely punctate, shiny; scutellum rather weakly punctate and slightly convex; propodeum evenly convex and coarsely rugose, mediolongitudinal carina complete and straight. Wings. Fore wing: r 0.4 × 3-SR (Fig. 199); 1-CU1 horizontal, as long as 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell rather short (Fig. 199); cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR wide; anterior half of subbasal and of subdiscal cells largely glabrous. Hind wing: basal half of marginal cell slightly widened, but apical half wide, apical width of cell 2.5 × width at level of hamuli (Fig. 199); 2-SC+R subquadrate; m-cu short and obsolescent; surroundings of M+CU and 1-M glabrous; M+CU:1-M = 75:47; 1r-m 0.8 × 1-M. Legs. Tarsal claws with conspicuous and robust dark brown pecten (Fig. 205); hind coxa largely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.3 and 5.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and largely coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 202); 2nd suture deep medially, shallow laterally and crenulate; 2nd tergite 0.7 × as long as wide (Fig. 202); anterior 0.7 of 3rd tergite densely and finely punctate, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically rather rounded (Fig. 197). Colour. Black; posterior half of mesoscutum, scutellum largely, apical rim of 1st tergite and basal rim of 2nd tergite reddish brown; fore coxa, bases of middle and hind coxae blackish; apex of hind tibia, telotarsi, hind tarsus, palpi, veins and pterostigma dark brown; tegulae and remainder of hind tibia pale yellowish; remainder of legs reddish brown; wing membrane subhyaline. Variation. Vein 1-CU1 of fore wing 0.8–1.1 × as long as 2-CU1; mesoscutum, scutellum, metanotum, 1st and 2nd metasomal tergites are most often entirely reddish or orange brown but variably partly blackish, in particular 1st tergite sometimes with

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Figures 212–216. Aleiodes cruentus (Nees), ♂, Hungary, Hársbokorhegy, but 213–215 from Germany, Markkleeberg 212 habitus lateral 213 1st–3rd metasomal tergites dorsal 214 4th–7th metasomal tergites lateral 215 id. dorsal 216 basal antennal segments.

dark medial patch; pronotum and mesopleuron black or reddish dorsally; parastigma narrowly dark brown or yellowish brown; coxae entirely reddish to entirely dark brown. Antennal segments: ♀ 53(1), 55(1), 56(3), 57(5), 58(9), 59(9), 60(10), 61(9), 62(3), 63(1), 65(2), 67(1). ♂ 60(6), 61(7), 62(2), 63(5), 64(3), 65(1), 66(1), 67(5), 69(1). The males have on average approx. three more antennal segments than females. Males are very similar but often darker than females, 2nd tergite 0.9–1.0 × as long as basal width of tergite and apical tergites type 1 and (usually) type 2, with fringe present in the latter (Fig. 215); hind femur at most apically blackish, and hind tibial spurs sometimes blunt. Distribution. *Austria, Bulgaria, Croatia, Czech Republic, Finland, France, Germany, *Greece, Italy, *Moldova, Mongolia, *Netherlands, Norway, *Romania, Slovakia, *Slovenia, Spain, Sweden, Ukraine. Notes. An examined female (NMS) from Albania (Mt Mali me Gropa, above Shengiergi, 1400 m, 13.viii.2019, MV light, C.W. Plant) has a CO1 sequence

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(MRS940) 3 % different from A. cruentus (19 differences in 626 bp of overlap) and although superficially similar in colour is clearly distinct in having OOL shorter (0.5 × lateral ocellus), a smaller hypoclypeal depression (0.5 × width of face), slenderer hind femur (5 × as long as wide), and several other differences. It may be A. parvicauda (Tobias, 1985) described from Afghanistan, but it has more (64; 58–60 in type series) and somewhat more elongate antennal segments than described for A. parvicauda, as well as other small deviations. Additional material as well as comparison with the type series of A. parvicauda are needed to settle the status of the Albanian species. Aleiodes desertus (Telenga, 1941) Figs 217–233 Rhogas (Rhogas) desertus Telenga, 1941: 184–185, 423 (not R. aestuosus var. desertus Telenga, 1941, from China) [examined]. Rogas desertus; Shenefelt, 1975: 1223. Rogas (Rogas) desertus; Tobias, 1986: 76 (transl. 124) (lectotype designation). Aleiodes desertus; Fortier & Shaw, 1999: 230. Type material. Lectotype, ♀ (ZISP), “[Uzbekistan:] Khiva, 30.iv.[1]927, V. Gussakovskij/ S.Kh.Op.Ot., at light”, “Lectotypus Rogas desertus Tel., design. [V.I.] Tobias, 1980”. Paralectotypes: 1 ♀ (ZISP), “[Turkmenistan:] Ashkhabad [= Ashgabat], 25.iii. [1]905, S. Ahnger”, “Paralectotypus Rogas desertus Tel., design. [V.I.] Tobias, 1980”; 1 ♀ (BMNH, figured), “Khiva, Rabat, 3.v.[1]927, V. Gussakovskij/collected at light”, “Paratypus Rogas desertus Telenga”, “Rec[eived] in exchange Academy of Science, Leningrad, B.M.1963-211”. Molecular data. None. Biology. Unknown. It seems to fly in spring (March–May) and may be univoltine. Diagnosis. Maximum width of hypoclypeal depression 0.9–1.0 × minimum width of face; anterior part of clypeus very narrow, most of clypeus depressed (Fig. 229); OOL approx. 0.9 × diameter of posterior ocellus and remotely punctate; mandible massive triangular, coarsely punctate and with thick ventral lamella (Figs 229, 231); face largely transversely rugose; malar space 0.15 × as long as height of eye and 0.27 × basal width of mandible; area of precoxal sulcus (but posteriorly superficially) and anteriorly area above it distinctly rugose; lateral lobes of mesoscutum largely smooth, strongly shiny and glabrous, middle lobe remotely punctulate and with satin sheen; basal half of wings (except anteriorly) largely glabrous and remainder of wing inconspicuously setose; vein r of fore wing 0.7–0.8 × vein 3-SR (Fig. 221) vein 1-CU1 0.1 × as long as 2-CU1, narrow and oblique; tarsal claws long, slender, hardly bent and simple (Fig. 232); tarsal segments (except telotarsus) with four apical spines; 1st and base of 2nd tergite aciculate-rugulose, 3rd tergite micro-sculptured and matt, remainder of metasoma shiny and rather smooth; head and mesosoma (except prothorax anteriorly and mesoscutum posteriorly) black; pterostigma dark brown; legs and palpi pale yellowish. According to original description antenna of ♀ with 50–52 segments, but BMNH paralectotype has 63 segments.

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Figures 217–220. Aleiodes desertus (Telenga), ♀, paralectotype 217 habitus lateral 218 antenna lateral 219 detail of fore wing 220 ovipositor sheath lateral.

Description. Lectotype, ♀, length of fore wing 7.5 mm, of body 8.2 mm. Head. Antennal segments of ♀ more than 45, but apical segments missing, length of antenna of paralectotype 1.1 × body and its subapical segments moderately slender; frons rugose, shiny; OOL 0.9 × diameter of posterior ocellus; OOL and vertex remotely punctate, shiny; anterior part of clypeus 9 × wider than high, coarsely punctate and rather convex; clypeus above lower level of eyes; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.9 × minimum width of face (Fig. 229); length of eye 1.7 × temple in dorsal view (Fig. 230); vertex behind stemmaticum convex and sparsely punctate; length of malar space 0.15 × length of eye in lateral view; mandible massive triangular, coarsely punctate and with thick ventral lamella (Figs 229, 231); occipital carina nearly complete, fine and ventrally strongly curved.

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Figures 221–233. Aleiodes desertus (Telenga), ♀, paralectotype, but 224 and 230 of lectotype 221 fore wing 222 hind wing 223 mesosoma lateral 224 mesosoma dorsal 225 metasoma dorsal 226 fore femur lateral 227 hind femur lateral 228 apex of antenna 229 head anterior 230 head dorsal 231 head lateral 232 outer hind tarsal claw 233 base of antenna.

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Mesosoma. Lateral lobes of mesoscutum largely smooth, strongly shiny and glabrous, middle lobe remotely punctulate and with satin sheen; prepectal carina complete and lamelliform; precoxal area of mesopleuron widely rugose, but posterior 0.2 narrowly striate; mesopleuron above precoxal area anteriorly rugose and remainder weakly and sparsely punctate, shiny; axilla crenulate but posteriorly densely and coarsely rugose; scutellum largely smooth, with some punctures; propodeum evenly convex, finely rugose and with strong medio-longitudinal carina, without tubercles. Wings. Fore wing: basal half largely glabrous; r 0.7 × 3-SR (Fig. 219); 1-CU1 oblique, 0.1 × as long as 2-CU1; r-m nearly as long as 3-SR; 2nd submarginal cell comparatively short (Fig. 221); cu-a inclivous; 1-M nearly straight posteriorly. Hind wing: basal 0.4 of marginal cell slightly widened and distally strongly widened, its apical width 2.7 × width at level of hamuli (Fig. 222); 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 3:2; 1r-m 0.8 × 1-M. Legs. Tarsal claws slender, slightly curved and only setose (Fig. 232); hind coxa partly obliquely striate dorsally; tarsi slender, segments (except telotarsus) with long apical spines; length of hind femur and basitarsus 5.0 and 6.8 × their width, respectively; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite robust, 0.9 × longer than wide apically, strongly narrowed anteriorly (Fig. 225) and rather flat posteriorly; 1st and 2nd tergites finely longitudinally striate-rugulose; medio-longitudinal carina of 1st and 2nd tergites indistinct; 2nd tergite 0.6 × longer than its basal width; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture shallow and narrow; 3rd tergite matt and micro-sculptured, anteriorly finely striate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with rather short setae and apically truncate (Fig. 220). Colour. Black; mesoscutum posteriorly partly chestnut brown; antenna, clypeus, malar space ventrally, mandible, pronotum and propleuron anteriorly and metasoma, brownish yellow; tegulae, legs and palpi pale yellowish; pterostigma and ovipositor sheath dark brown; veins of fore wing (but pale in basal 0.3 of fore wing) brown; wing membrane hyaline. Variation. Length of body 7.0–8.2 mm, of fore wing 7.5–7.9 mm; temple punctate to smooth; precoxal sulcus area finely to rather coarsely rugose; pronotal side largely black (except ventrally) black or brownish yellow; lateral lobes of mesoscutum entirely dark chestnut brown or only posteriorly so, or mesoscutum largely yellowish brown posteriorly and prolonged to base of notauli; first tergite usually entirely brownish yellow, but sometimes dark brown and only posteriorly and laterally yellowish; pterostigma dark brown or brown. Antennal segments: ♀ 63(1). Distribution. Turkmenistan, Uzbekistan. Notes. We have included this extralimital species from Central Asia because we suspect it may occur in Turkey. It should not be confused with Rogas aestuosus var. desertus Telenga, 1941, described from China in the same paper. The latter is an unavailable name (a primary homonym) and most likely a colour variety of R. aestuosus.

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Aleiodes dissector (Nees, 1834) Figs 234–263 Rogas dissector Nees, 1834: 208; Shenefelt, 1975: 1225–1226; Papp, 1977a: 110. Rogas (Rogas) dissector; Tobias, 1976: 81, 1986: 75 (transl.: 121). Aleiodes (Neorhogas) dissector; Papp, 1985a: 145, 1987b: 35, 1991a: 74, 1991d: 5, 1999: 550; Belokobylskij, 1996: 9; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) dissector; Belokobylskij, 2000: 34; Ku et al., 2001: 234, 235. Aleiodes dissector; Fortier & Shaw, 1999: 230; Belokobylskij et al., 2003: 398; ZaldivarRiverón et al., 2004: 234, 2008: 392; Papp, 2005: 176; Belokobylskij et al., 2008: 136–137. Phylax aestivalis Snellen van Vollenhoven, 1858: 282; Shenefelt, 1975: 1226 (as synonym of A. dissector); van Achterberg, 1992: 363 (id.); Papp, 2005: 176 (id.) [examined]. Type material. Holotype of A. aestivalis, ♀ (RMNH), “[Netherlands], Haag [= near The Hague], 6 [= June], v.Voll.”. According to the original description the ♂ holotype of R. dissector from Germany should be in the Gravenhorst collection (Museum of Natural History, University of Wrocław, Wrocław), but so far it has not been found. Additional material. Austria, British Isles (England: V.C.s 15, 17, 20, 22, 23, 24, 30, 31, 34, 37, 58; Scotland: V.C.s 73, 88, 89, 95, 96, 97, 107), Croatia, Czech Republic, Finland, France, Germany, Greece, Hungary, Montenegro, Italy, Netherlands (FL: Lelystad, GE: Barneveld, OV: Raalte (Heino), ZH: Wassenaar), Norway, Russia, Serbia, Slovakia, Switzerland, Ukraine, [Armenia]. Specimens in BMNH, BZL, CNC, IKC, MRC, MSC, MSNV, MTMA, NMS, OUM, RMNH, SDEI, UNS, UWIM, ZSSM. Molecular data. MRS007 (Turkey), MRS025 (Turkey), MRS145 (UK), MRS146 (UK). Biology. Univoltine, collected in May and June in deciduous scrub and woodland. In Britain it is widespread but particularly common in birch-dominated woodland in upland Scotland. Reared from the noctuids Orthosia incerta (Hufnagel) (17, M.R. Shaw), O. gothica (Linnaeus) (1, J.L. Yela) and Orthosia sp. (3), overwintering in the concealed mummy. An additional specimen, lacking a mummy but labelled as reared doubtfully from the sesiid Paranthrene tabaniformis (Rottemburg) (RMNH), which normally feeds under Populus bark at ground level or below, can be discounted as a probable substrate rearing in which the mummy of the true host was overlooked. Parasitised host larvae in their penultimate instar leave their feeding sites and enter the soil or other site of moderate concealment (including below loose bark), where they prepare a chamber as though to pupate. At this time the parasitoid larva within the strongly retarded host (Fig. 234) is around half its final length, and the host lies quiescent for approx. a week until the parasitoid has completed its feeding (Fig. 235). During mummification (Figs 236, 237) the caudal end of the host recurves ventrally as the host’s body becomes weakly retracted. A ventral opening at the head end is made, but the head (as with the caudal segments) is usually tucked downwards rather than becoming raised, and so the resulting expelled fluid (Fig. 237) usually dries without the mummy becoming stuck down. The eventual

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Figures 234–239. Aleiodes dissector (Nees), U.K., Scotland (in culture) parasitising Orthosia incerta (Hufnagel) 234 pre-mummy, removed from its hideaway, with unparasitised control from the same egg batch (below) 235 pre-mummy 236 early mummification 237 mummy with ventral ooze 238 three fully hard mummies 239 emerged mummy, cut open to expose silken lining.

outcome is a rather distinctive (Fig. 238) elongate and curved dark brown structure with a paler and weakly raised lateral keel. The parasitoid’s pupation chamber occupies ca 2nd–8th abdominal segments of the host, which are moderately strongly lined with silk (Fig. 239). The moderately large hypoclypeal opening and protruding sharp-rimmed clypeus of A. dissector is seen in some other species (e.g., A. modestus (Reinhard), treated in part 1 of this work) whose hosts also pupate in shallow soil. In culture experiments A. dissector was found to prefer hosts in the early to middle part of the 3rd instar, although late 2nd instar host were often also acceptable. Oviposition into suitable hosts was rapid (1–2 seconds) and accomplished with a single insertion of the ovipositor, following only brief

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Figures 240–242. Aleiodes dissector (Nees), ♀, Switzerland, Tessin, but 242 from Scotland (culture) 240 habitus lateral 241 ovipositor sheath lateral 242 mummy of Orthosia incerta (Hufnagel).

antennation and no use of the legs. There was no clear temporary paralysis. Experimental rearings from O. incerta (6:107\85\\75+10) and O. gothica (6:61\49\\34+15) were comparable (given that some insertions of less than a full second might have been scored as ovipositions incorrectly; and furthermore that some failures to oviposit into these hosts might be ascribed to temporary egg depletion, as the protocol of normally ceasing

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Figures 243–255. Aleiodes dissector (Nees), ♀, Switzerland, Tessin 243 fore wing 244 hind wing 245 mesosoma lateral 246 mesosoma dorsal 247 propodeum and 1st –4th metasomal tergites dorsal 248 fore femur lateral 249 hind femur lateral 250 outer hind tarsal claw 251 head anterior 252 head dorsal 253 head lateral 254 base of antenna 255 apex of antenna.

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to offer hosts to a particular female after four apparent ovipositions on the day had not been developed until after the experiments were undertaken), and clearly demonstrated the suitability of both species as hosts. In contrast, no parasitoids developed (and indeed possibly no ovipositions occurred) in the other species of Orthosia tested, which were all found to be clearly outside the host range: O. cerasi (Fabricius) (3:32\?3\\0+3); O. cruda (Denis & Schiffermüller) (2:12\0\\-); O. munda (Denis & Schiffermüller) (3:10\0\\-); O. gracilis (Denis & Schiffermüller) (2:11\?1\\0+1). Of these four, only O. gracilis is not fully arboreal. There is no adverse venom effect on host development. Diagnosis. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 251); OOL of ♀ 0.6–0.7 × diameter of posterior ocellus (Fig. 252) and sparsely punctate; ventral margin of anterior part of clypeus comparatively sharp and more or less protruding outwards (Fig. 253); length of malar space 0.2 × length of eye in lateral view (Fig. 253); head transverse in dorsal view and eye 1.5–2.0 × as long as temple in dorsal view (Fig. 252); lobes of mesoscutum punctulate, with interspaces smooth to superficially micro-sculptured; precoxal area completely smooth or nearly so; vein 1-CU1 of fore wing 0.2–0.3 × vein 2-CU1 and horizontal (Fig. 243); hind tarsal claws with conspicuous dark brown pecten close to apical tooth (Fig. 250); 1st tergite rounded antero-laterally and 1.0–1.1 × as long as wide apically; basal half of metasoma black and weakly sculptured; 3rd tergite smooth; head black; palpi yellowish; basal half of hind tibia pale yellowish, but in some males almost uniformly dark; 4th–6th tergites of males depressed medially and conspicuously setose (Fig. 258). Description. Redescribed ♀ (RMNH) from Austria (Burgenland, Winden am See). Length of fore wing 8.5 mm, of body 9.0 mm. Head. Antennal segments of ♀ 60, antenna as long as fore wing, its subapical segments rather slender, slightly longer than wide; frons largely smooth; OOL 0.7× diameter of posterior ocellus, sparsely punctate, shiny and with deep groove near posterior ocellus (Fig. 252); vertex sparsely punctate, rather shiny; clypeus coarsely punctate; ventral margin of clypeus rather thin and forward protruding (Fig. 253); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 251); length of eye 1.5 × temple in dorsal view (Fig. 252); vertex behind stemmaticum superficially rugose-punctate; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Mesoscutal lobes punctulate with interspaces superficially micro-sculptured and shiny; precoxal area of mesopleuron smooth except some punctulation, mesopleuron punctulate anteriorly and posteriorly; metapleuron densely punctate; metanotum with nearly complete median carina; scutellum flat (but with rugulose depression medio-posteriorly), remainder punctulate and with weak lateral carinae; propodeum evenly convex and coarsely rugose, and medio-longitudinal carina absent posteriorly. Wings. Fore wing: r 0.4 × 3-SR; 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.3 × 3-SR; 2nd submarginal cell medium-sized (Fig. 243); cu-a inclivous, straight; 1-M straight posteriorly; 1-SR medium-sized; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell rather narrow basally, apical half gradually widened, its apical width 3.1 × width at level of hamuli (Fig. 244); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 35:33; 1r-m 0.7 × 1-M.

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Figures 256–263. Aleiodes dissector (Nees), ♂, Austria, Kärnten 256 habitus lateral 257 head dorsal 258 3rd–7th tergites lateral 259 mesosoma dorsal 260 1st–6th metasomal tergites dorsal 261 fore femur lateral 262 head anterior 263 hind femur lateral.

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Legs. Tarsal claws with conspicuous and robust dark brown pecten (Fig. 250); hind coxa distinctly punctate and with some oblique striae postero-dorsally; hind trochantellus robust; length of hind femur and basitarsus 3.8 and 5.3 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite flattened, basally narrowed, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and largely finely punctate-rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite wide and triangular, distinct (Fig. 247); 2nd suture rather deep and micro-sculptured; 3rd and subsequent tergites largely smooth; apical half of 3rd and 4th tergites without sharp lateral crease; ovipositor sheath wide, with long and mediumsized setae and apically truncate (Fig. 241). Colour. Black; apical half of hind tibia and hind tarsus blackish; basal half of hind tibia pale yellowish; remainder of legs, palpi and tegulae yellowish brown; most veins and pterostigma dark brown; wing membrane slightly yellowish basally and remainder slightly infuscate. Variation. Interspaces between punctulation of mesoscutum smooth to superficially micro-sculptured; medio-longitudinal carina of propodeum complete or absent posteriorly; 3rd metasomal tergite largely finely sculptured (except posteriorly) to largely smooth; mesopleuron black or with brownish longitudinal stripe; hind tibia usually ivory or pale yellowish basally. Antennal segments: ♀ 51(2), 55(2), 56(7), 57(4), 58(7), 59(12), 60(14), 61(18), 62(6), 63(4); ♂ 51(1), 53(2), 54(2), 55(4), 56(8), 57(29), 58(29), 59(27), 60(15), 61(4), 62(4), 63(2), 64(2). Females have on average ca one to two more antennal segments than males. Males are very similar but hind femur more or less blackish and, in some males, hind tibia almost uniformly dark, OOL approx. as long as diameter of posterior ocellus (Fig. 257) and apical tergites type 3–4 with fringe long and strong (Figs 258, 260). Distribution. *Armenia, *Austria, British Isles (England, Scotland), Croatia, Czech Republic, Finland, France, Germany, *Greece, Hungary, *Montenegro, *Italy, Netherlands, Norway, Russia, *Serbia, Switzerland, Ukraine. Aleiodes diversus (Szépligeti, 1903) Figs 264–288 Rhogas diversus Szépligeti, 1903: 114; Papp, 2004: 216 (as synonym of A. dissector) [examined]. Rogas dissector var. diversus; Shenefelt, 1975: 1226 (lectotype designation). Aleiodes (Neorhogas) diversus; Papp, 1977a: 110–112 (re-instated), 1985: 145, 1991a: 81. Aleiodes diversus; Belokobylskij et al., 2003: 398 (as synonym of A. dissector); Papp, 2005: 176; Merz & Pasche, 2012: 244; van Achterberg, 2014: 209. Type material. Lectotype, ♀ (MTMA), “Croatia, Buccari [= Bakar], 1893, Pavel”, “Lectotypus, ♀”, Rogas (s. str.) diversus Szépligeti, 1906 [sic!], Papp, 1968”, “Hym. Typ. No. 1011, Mus. Budapest”.

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Figures 264, 265. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 264 habitus lateral 265 ovipositor sheath lateral.

Additional material. Austria, British Isles (England: V.C.s 8, 25, 70), Bulgaria, Croatia, Hungary, Italy (Sicily), Norway, Switzerland. Specimens in BMNH, BZL, CMIM, MHNG, MRC, MTMA, NMS, RMNH, SDEI, ZSSM. The most recent of the five English specimens seen is dated 1931, and it seems likely that this rather large and showy insect is extinct in Britain. Molecular data. None. Biology. Unknown. Female specimens have been collected in (May–)June, and also September, suggesting that it may be plurivoltine. This is reinforced by the date of capture of the two available males (which would not have hibernated as an adult) in Sicily on 30.iv.1965 (BMNH) and 1.v.1994 (RMNH). There is no indication of habitat on data labels and we have not seen reared material. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 273); OOL of ♀ coarsely punctate and 1.0–1.2 × diameter of posterior ocellus; ventral margin of clypeus (rather) obtuse apically and clypeus not protruding outwards (Fig. 275), but sometimes intermediate; length of eye 1.0–1.2 × temple in dorsal view; lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures, shiny and smooth; precoxal area with some rugae medially; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein 1-CU1 of fore wing 0.7–1.1 × vein 2-CU1 and approx. as long as vein m-cu (Fig. 266), rarely shorter; hind femur 3.0–3.3 × longer than wide; hind tarsal claws with medium-sized dark brownish pecten up to apical tooth (Fig. 272); 1st tergite widened apically and moderately wide basally (Fig. 269); 2nd tergite 0.7–0.8 × as long as wide (Fig. 269) and black; 4th–7th tergites of males flat and with long yellowish setae (Figs 279, 282); head black; vein 1-M of fore wing brownish; wing membrane subhyaline. Description. Lectotype, ♀, length of fore wing 7.0 mm, of body 10.0 mm. Head. Antennal segments of ♀ 56, antenna as long as fore wing, its subapical segments robust; frons largely smooth behind antennal sockets; OOL 1.2 × diameter of

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Figures 266–277. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 266 wings 267 mesosoma lateral 268 mesosoma dorsal 269 propodeum and metasoma dorsal 270 fore femur lateral 271 hind femur lateral 272 outer hind tarsal claw 273 head anterior 274 head dorsal 275 head lateral 276 base of antenna 277 apex of antenna.

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posterior ocellus, and coarsely punctate, interspaces less than diameter of puncture; vertex coarsely punctate; clypeus rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 275); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 273); length of eye 1.2 × temple in dorsal view (Fig. 274); vertex behind stemmaticum superficially punctate-rugose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, interspaces largely smooth, shiny; precoxal area of mesopleuron coarsely punctate and without rugae medially, mesopleuron coarsely punctate anteriorly and posteriorly; metapleuron moderately punctate; scutellum remotely punctate; propodeum rather convex and coarsely rugose. Wings. Fore wing: r 0.5 × 3-SR (Fig. 266); 1-CU1 horizontal, 0.7 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell rather long (Fig. 266); cu-a vertical, straight; 1-M rather curved posteriorly; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened, its apical width 2.3 × width at level of hamuli (Fig. 266); 2-SC+R transverse; m-cu largely absent, only as short antefurcal remnant (Fig. 266); M+CU:1-M = 35:23; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather conspicuous, medium-sized dark brown pecten up to apical tooth (Fig. 272); hind coxa largely punctate; hind trochantellus robust; length of hind femur and basitarsus 3.1 and 4.4 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely vermiculate-rugose; medio-basal area of 2nd tergite triangular and distinct (Fig. 269); 2nd suture deep; 2nd tergite 0.7–0.8 × as long as wide (Fig. 269); 3rd tergite densely punctate and interspaces largely smooth, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; apical third of metasoma rather compressed; ovipositor sheath wide, with rather short setae and apically truncate (Fig. 265). Colour. Black; mesoscutum (except anterior third), scutellum, clypeus ventrally, mandible, tegulae and legs largely brownish red; palpi, fore coxa largely, telotarsi, hind tarsus and apex of hind tibia (excluding spurs) dark brown; pterostigma blackish brown; veins dark brown, but near wing base yellowish; wing membrane slightly infuscate. Variation. OOL 1.0–1.2 × diameter of posterior ocellus; mesoscutum of ♀ entirely brownish red or yellowish brown, or anteriorly black; 1st tergite 1.0–1.1 × longer than wide apically; metasoma rarely partly obscurely reddish dark brown; mesopleuron may be just punctate or may have some rugae in lower half. Antennal segments ♀: 55(3), 56(3), 57(3), 58(1), 59(1); ♂ 58(1). Males have mesosoma black (Fig. 278), 2nd tergite 0.8–0.9 × as long as basal width of tergite (Fig. 282) and apical tergites type 2, setae rather long, fringe long and strong (Fig. 279). Distribution. *Austria, *British Isles (England; probably extinct), *Bulgaria, Croatia, Hungary, *Italy (Sicily), *Norway, *Switzerland. Notes. Close to A. cruentus which, however, almost always has much or all of 1st and 2nd metasomal tergites orange-red (usually wholly black or dark brown in

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Figures 278–288. Aleiodes diversus (Szépligeti), ♂, Italy, Sicily 278 habitus lateral 279 3rd–7th tergites lateral 280 wings 281 mesosoma lateral 282 metasoma dorsal 283 outer hind claw 284 hind tibial spurs lateral 285 head anterior 286 head dorsal 287 base of antenna 288 apex of antenna.

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A.  diversus). In addition to characters given in the key A. diversus is a more robust insect, and females have broader antennal segments (distinctly transverse near middle of flagellum) and on average they are fewer in number (although with overlap). Aleiodes eurinus (Telenga, 1941) Figs 289–306 Rhogas (Rhogas) eurinus Telenga, 1941: 422. Rogas eurinus; Shenefelt, 1975: 1228; Papp, 1971: 359. Rogas (Rogas) eurinus; Tobias, 1976: 85, 1986: 80 (transl.: 130; lectotype designation). Aleiodes (Neorhogas) eurinus; Papp, 1985a: 145; 1991a: 94. Aleiodes (Chelonorhogas) eurinus; Chen & He, 1997: 39; He et al., 2000: 667; Belokobylskij, 2000: 49; Ku et al., 2001: 235; Farahani et al., 2015: 242–243; Beyarslan et al., 2017: 330. Aleiodes eurinus; Fortier & Shaw, 1999: 223, 230; Belokobylskij et al., 2003: 398; Papp, 2005: 176. Rogas eurinus ab. nigratus Papp, 1967: 223 (invalid name). Rogas eurinus ab. nigrimaculatus Papp, 1967: 223 (invalid name). Rogas eurinus ab. nigripes Papp, 1967: 223 (invalid name). Type material. None seen. Additional material. Italy, Russia (Siberia and Far East), Spain, Turkey, [China, Mongolia]. Specimens in BMNH, BZL, MRC, MSNV, MTMA, NMS, RMNH, SDEI, ZISP. Molecular data. None. Biology. Specimens have been collected from April to August, and the presence of males in both April and July clearly demonstrates that it is plurivoltine and overwinters in the mummy. We have not seen reared material, but specimen labelling indicates that it occurs among Ammophila and Schoenus in the Venice Lido and Triticum (presumably cultivated wheat) in Turkey, suggesting that its hosts will occur in open grassland habitats. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 298); OOL of ♀ approx. as long as diameter of posterior ocellus (Fig. 299) and densely rugose; clypeus rather thin apically and rather protruding anteriorly (Fig. 300); eyes prominent (Fig. 299); lobes of mesoscutum distinctly punctate-granulate and rather matt; precoxal area more or less rugose and comparatively wide medially, and posteriorly punctate; vein 1-CU1 of fore wing 0.3 × vein 2-CU1 (Fig. 292); hind tarsal claws slender, brownish setose and without pecten (Fig. 302); basal half of 3rd tergite striate; 3rd antennal segment of ♀ dark brown; basal half of hind tibia pale yellowish or ivory, at least inner side contrasting with reddish or dark brown colour of basal half of hind femur (usually less pronounced in ♂).

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Figures 289–291. Aleiodes eurinus (Telenga), ♀, Mongolia, but 291 Russia, Chelyabinskoi Obl. 289 habitus lateral 290 ovipositor sheath lateral 291 apex of antenna.

Description. Redescribed ♀ (RMNH) from Turkey (Ankara). Length of fore wing 7.0 mm, of body 8.0 mm. Head. Antennal segments of ♀ 51 remaining, but apical segments missing, length of antenna 1.2 × fore wing; frons with coarse curved rugae and dorsally coarsely rugose; OOL equal to diameter of posterior ocellus, and densely rugose; vertex spaced rugose, rather dull; clypeus medium-sized and coarsely rugose (as face); ventral margin of clypeus rather thin and rather protruding forwards (Fig. 300); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 298); length of malar space in anterior view 0.7–1.0 × maximum width of hypoclypeal depression (Fig. 298); head in anterior view trapezoid; length of eye 1.3 × temple in dorsal view and temples directly narrowed behind eyes (Fig. 299); vertex behind stemmaticum densely rugose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes distinctly punctate-granulate, and with satin sheen; precoxal area of mesopleuron coarsely rugose, rather wide medially and posteriorly coarsely punctate and some short rugae, densely punctate; remainder of mesopleuron mainly sparsely and finely punctate; metapleuron densely punctate; metanotum with nearly complete median carina; scutellum punctulate and weakly granulate; propodeum coarsely vermiculate-rugose, medio-longitudinal carina irregular. Wings. Fore wing: r 0.4 × 3-SR (Fig. 292); m-cu far antefurcal; 1-CU1 horizontal, slightly widened, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 292); cu-a inclivous, somewhat curved posteriorly; 1-M rather curved posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing:

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Figures 292–302. Aleiodes eurinus (Telenga), ♀, Mongolia 292 wings 293 mesosoma lateral 294 mesosoma dorsal 295 1st–3rd metasomal tergite dorsal 296 fore femur lateral 297 hind femur lateral 298 head anterior 299 head dorsal 300 head lateral 301 base of antenna 302 outer hind tarsal claw.

marginal cell gradually widened, its apical width 2.6 × width at level of hamuli (Fig. 292); 2-SC+R subquadrate; m-cu medium-sized and only pigmented; M+CU:1-M = 50:43; 1r-m 0.7 × 1-M.

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Figures 303–306. Aleiodes eurinus (Telenga), ♂, Mongolia 303 habitus lateral 304 head anterior 305 apical half of metasoma lateral 306 head dorsal.

Legs. Tarsal claws slender and brownish setose (Fig. 302); hind coxa rather finely and densely punctate; hind trochantellus robust; length of hind femur and basitarsus 5.0 and 6.5 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus.

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Metasoma. First tergite rather flattened, 1.1 × longer than wide apically; 1st and 2 tergites with medio-longitudinal carina and rather regularly longitudinally rugose; medio-basal area of 2nd tergite narrow triangular (Fig. 295); 2nd suture deep and crenulate; basal half of 3rd tergite longitudinally striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 290). Colour. Black; palpi and basal half of antenna (except scapus and pedicellus) brown; scapus, pedicellus, clypeus largely, apex of hind femur (but ventrally reddish), apex of hind tibia, hind tarsus, all telotarsi, pterostigma (but basally narrowly pale) and veins (except yellowish veins of basal quarter of wings) dark brown; remainder of legs and 1st–3rd tergites orange brown; tegulae and hind tibia (except apically) pale yellowish; wing membrane subhyaline. Variation. Coxae and hind femur (except its basal third) largely dark brown, black or orange brown; apical half of hind tibia dark brown or only apically so; 1st tergite largely dark brown (except posteriorly), with pair of dark brown spots or entirely orange or reddish brown; apical half of 3rd tergite orange brown or largely black. Antennal segments: ♀ 54(1), 55(2), 57(2), 58(3), 59(1), 60(2); ♂ 52(1), 60(1). Male is very similar and has apical tergites type 1–2, setae moderately dense, glabrous stripe only rarely evident and fringe very short, negligible (Figs 303, 305). Distribution. China, *Italy, Mongolia, Russia (Siberia and Far East), Spain, *Turkey. nd

Aleiodes fahringeri (Telenga, 1941) Figs 307–321 Rhogas (Rhogas) fahringeri Telenga, 1941: 173. Rogas fahringeri; Shenefelt, 1975: 1228. Rogas (Rogas) fahringeri; Papp, 1977b: 113. Aleiodes (Chelonorhogas) fahringeri; Chen & He, 1997: 40; He et al., 2000: 666; Belokobylskij, 2000: 39 (lectotype designation); Papp, 2009: 149. Aleiodes fahringeri; Chen & He, 1992: 125; Fortier & Shaw, 1999: 230. Rhogas (Rhogas) flavipennis Telenga, 1941: 174, 419. Rogas flavipennis; Shenefelt, 1975: 1229. Aleiodes (Chelonorhogas) flavipennis; Belokobylskij, 2000: 39 (lectotype designation and synonymised with A. fahringeri (Telenga, 1941)). Type material. None examined. Additional material. 3 ♀ (MTMA, NMS, RMNH), “Mongolia: Südgobi aimak, Somon Bulgan, Talyn bulag, 1350 m, Exp. Dr. Z. Kaszab, 1967”, “Nr. 889, 5.vii.1967”, “Rogas fahringeri Tel., ♀, det. Papp, 1976”; 1 ♀ (RMNH, ZJUH), “China: Ningxia, Yinchuan, 6.vii.1983, no. 840994, Xu Wenzhong, RMNH’99”. Molecular data. None.

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Figures 307–309. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 307 habitus lateral 308 ovipositor sheath lateral 309 detail of fore wing.

Biology. Unknown. Specimens have been collected in June–August. Presumed to be univoltine, but we have not seen reared material and the means of overwintering is unclear. Diagnosis. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face; OOL 0.9 × diameter of posterior ocellus, largely smooth with spaced punctures; ventral margin of clypeus thin, anterior part shiny and distinctly protruding anteriorly (Fig. 320); mesoscutum shiny and moderately punctulate; precoxal area with only some rugulae medially; vein r of fore wing 0.5–0.6 × as long as vein 3-SR; tarsal claws rather slender and with yellowish or brown pecten, pecten remains removed from apical tooth (Fig. 321); hind tarsus fairly elongate and segment with medium-sized apical spines (Figs 307, 321); pterostigma brownish yellow; wings subhyaline; head and mesosoma laterally and dorsally (except more or less dark brown propodeum) yellowish brown; fore wing longer than 5 mm. Description. Redescribed ♀ (RMNH) from Mongolia (Somon Bulgan). Length of fore wing 6.9 mm, of body 7.7 mm. Head. Antennal segments of ♀ 58, length of antenna 1.1 × fore wing, its basal and subapical segments slender (Figs 316, 317); frons largely smooth anteriorly and rugulose posteriorly; OOL 0.9 × diameter of posterior ocellus, largely smooth with spaced punctures, (but superficially coriaceous near eye) and with satin sheen; vertex largely smooth, but superficially rugulose behind ocelli; anterior part of clypeus nearly 5 × wider than long, medially distinctly wider than laterally, largely smooth, punctulate,

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Figures 310–321. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 310 wings 311 mesosoma lateral 312 mesosoma dorsal 313 propodeum and 1st–3rd metasomal tergites dorsal 314 fore femur lateral 315 hind femur lateral 316 base of antenna 317 apex of antenna 318 head anterior 319 head dorsal 320 head lateral 321 inner hind tarsal claw

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its ventral margin thin and protruding forwards (Fig. 320); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 318); length of eye 1.6 × temple in dorsal view (Fig. 319); clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Pronotum medio-dorsally flat, shiny and largely smooth; mesoscutal lobes largely smooth except for punctulation, shiny and densely setose; precoxal area of mesopleuron largely smooth medially, with only some superficial rugulae; remainder of mesopleuron finely punctate and antero-dorsally rugose; metapleuron remotely punctate and largely smooth medially; scutellum remotely punctulate; metanotum with fine complete median carina; propodeum weakly convex and densely rugose, its medio-longitudinal carina complete and fine. Wings. Fore wing: r 0.5 × 3-SR (Fig. 310); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 310); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR somewhat widened; surroundings of M+CU1, 1-M and 1-CU1 evenly setose but setae pale and easily overlooked. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 310); 2-SC+R short and longitudinal; m-cu vaguely indicated; M+CU:1-M = 10:7; 1r-m 0.8 × 1-M. Legs. Tarsal claws with rather inconspicuous and pale brownish pecten remaining far removed from apical tooth (Fig. 321); hind coxa largely superficially finely punctate, but dorso-anteriorly densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.4 and 6.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flat, 1.1 × as long as wide apically; 1st and 2nd tergites with fine medio-longitudinal carina and finely longitudinally (1st) or irregularly (2nd) densely rugose; medio-basal area of 2nd tergite triangular and medium-sized (Fig. 313); 2nd suture rather deep, finely crenulate and narrow; basal half of 3rd tergite finely rugulose, remainder of metasoma superficially micro-sculptured or nearly smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 308). Colour. Yellowish brown; antenna (except dark brown scapus and pedicellus), stemmaticum and ovipositor sheath black; tarsi, medio-posterior patch of propodeum, basal patch of 1st tergite and apex of hind tibia dark brown; veins rather dark brown at medial third of fore wing, remainder of veins pale brown or yellowish; pterostigma brownish yellow; wing membrane subhyaline. Variation. Scapus entirely dark brown or largely yellowish brown; dark patches of propodeum and 1st tergite sometimes absent (♀ RMNH from Ningxia). Antennal segments: ♀ 56(2), 58(2), 59(1); ♂ 57(1), 59(1). Male is very similar with apical tergites type ?1–2, setae short, sparse and hard to see, with fringe very short and negligible. Distribution. China (Ningxia), Mongolia. Notes. This Asian species is included here because it was reported from Poland (Huflejt, 1997). The record needs confirmation to rule out confusion with a similar European species.

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Aleiodes fortipes (Reinhard, 1863) Figs 322–342 Rogas fortipes Reinhard, 1863: 272; Shenefelt, 1975: 1229 [examined]. Aleiodes (Neorhogas) fortipes; Papp, 1985a: 158, 1987a: 333, 1987b: 35, 1991a: 75. Aleiodes fortipes; Papp, 2005: 176; Lozan et al. 2010: 17; Butcher et al., 2012: 14. Rhogas freyi Hellén, 1927: 25–26; Papp, 1985a: 158 (unnecessary lectotype designation and as synonym of A. fortipes), 2005: 176 [examined]. Rogas freyi; Shenefelt, 1975: 1229–1230; Tobias, 1986: 75 (transl.: 121). Type material. Holotype of A. fortipes, ♂ (ZMB), “Gallia [France]”, “Type”, “Coll. H. Rhd.”, “26723”, “fortipes Rhd.”, “Holotypus”, “Rogas fortipes Reinh., 1863, ♂, Papp, 1983.”. Holotype of A. freyi, ♂ (ZMH), “[Finland], Nagu”, “R. Frey”, “Freyi n. sp., Hellén det.”, “Mus. Zool. H:fors, sp. typ. No. 5363, Rhogas Freyi Hellén”, “Lectotypus Rogas freyi Hellén, design. Tobias”, “Aleiodes % ♂ fortipes Rh., det. Papp J., 1983/ compared with ♂ holotype of A. fortipes”. The lectotype designation is superfluous because it is evident from the description that the author had only one male. Additional material. Austria, British Isles (England: V.C.s 16, 26, 28), Bulgaria, Czech Republic, Finland, France, Germany, Hungary, Netherlands (GE: ‘t Harde, Nunspeet), Poland, Spain, Sweden, Turkey. Specimens in BMNH, BZL, CMIM, FC, MTMA, NMS, RMNH, SDEI. It has been collected in open or understory habitats, including (but not exclusively) growths dominated by Vaccinium and/or Calluna below sparse conifers. Generally, found on sandy well-drained soils in England (Breck heaths of East Anglia) and the Netherlands (Veluwe). Molecular data. MRS650 (France), MRS807 (Poland). Biology. The flight time of this univoltine species is (April)May–June, and ca 10 months of the year is spent as an exposed mummy. The only mummy seen (Fig. 324) formed in captivity firmly attached beneath a thin stem and would have been positioned low down in the vegetation, but probably aerially. It is light brown, moderately slender, and the parasitoid occupied approximately abdominal segments 3–8. The host was Idaea sp. (Geometridae), either I. aversata (Linnaeus) or I. straminata (Borkhausen), and the rearing arose when a few larvae of the foregoing were collected (MRS) along with an adult female of A. fortipes at the same site in Poland (22.v.2016) and offered to the parasitoid, which had been fed honey water, on 24.v.2016. Although two of the caterpillars were well-grown, in their final instars and at least twice as long as the parasitoid, one was accepted avidly. This host was first pricked several times, at intervals. Paralysis was rather slow to take effect and not complete until after the host was revisited for oviposition: a single insertion of ca 30 seconds duration, with no postoviposition association (the parasitoid simply walked away after oviposition). The host mummified on 9.vi.2016 and an adult female emerged on 22.v.2017. The other host was rejected after being pricked just once, and later died. A penultimate instar caterpillar of the same host aggregate was also parasitised but died after an ecdysis. Subsequent

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

barcoding (through the kindness of Axel Hausmann, ZSSM) of the dead caterpillars revealed one specimen each of I. aversata and I. straminata, leaving the precise determination of the successful host unclear. It is possible that the parasitised host had already been attacked before it was collected, but the rather long time before mummification occurred suggests not. In any case, at least one Idaea species in the aversata/ straminata group clearly serves as host. Some individuals of the long and slender, morphologically

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very different, larvae of Idaea muricata (Hufnagel) were also offered. Although possibly of less interest to the parasitoid, one penultimate instar larva (1.7 times the length of the female parasitoid) was immediately parasitised (a single prick for eventual paralysis, followed after an interval by a single insertion for oviposition lasting just more than a minute), but this larva later produced a moth. Final instars of this very elongate species of caterpillar were generally ignored, but one did elicit a downwards curl of the metasoma without, however, being stung. There are two particularly significant aspects to the successful rearing. The first is that these Idaea species overwinter as quite well-grown larvae, so during the flight period of the parasitoid they are in late instars, and attacking hosts at this stage is an unusual strategy for Aleiodes (but see A. aterrimus and A. sibiricus). The second is that we know of no other Aleiodes species apart from A. sibiricus (q. v.) among those whose host overwinters as a larva that fails to take advantage of that to overwinter as an early instar larva within it. The apparently riskier strategy taken by A. fortipes, in both respects, may be plesiomorphic. Aleiodes fortipes is the only known West Palaearctic species in which males have small, subapical setose pore (probably associated with pheromone release) situated middorsally on each of the 4th–6th metasomal tergites (Fig. 340). We also expect these pores to be present in A. caucasicus, which is only doubtfully distinct from A. fortipes, but we have not seen the male of A. caucasicus. Similar, probably homologous, pores are also a feature of males of Aleiodes (Hemigyroneuron) species which are found in the near East, Oriental and Afrotropical regions (Butcher & Quicke, 2015). Outside of Hemigyroneuron, metasomal pores are also found the New World Aleiodes cameronii (Dalla Torre) species complex and in a number of undescribed Madagascan Aleiodes. In Hemigyroneuron the pores have been shown to connect with large sub-tergal glands (Butcher & Quicke, 2011). Collectively these taxa form a basal clade in our molecular phylogeny (Fig. 1). Aleiodes fortipes is the only species among those treated in this part of our revision with known hosts outside the Noctuidae and, although no host is known for rather a lot of these species, the apparently basal position of A. fortipes in the group is noteworthy and using geometrid hosts may also be plesiomorphic. The rather slender ovipositor sheath (Fig. 322) is another indication for its basal position. It is interesting that the known hosts of both A. fortipes and of the subgenus Hemigyroneuron are all Geometridae (two species of Hemigyroneuron with examined mummies, India and S. Africa, cited by Butcher & Quicke, 2011 [a label record indicating a pierid host of a 3rd species is also cited in that paper but is discounted here because no mummy was present]). An Australian species described under Hemigyroneuron with examined mummy reported to be that of a geometrid by Butcher & Quicke (2016) is probably (a) actually not a member of A. (Hemigyroneuron) and (b) may be from a lasiocampid (W. Moore in litt.). The hosts of members of the Aleiodes cameronii complex, based on multiple rearings in both North America and Costa Rica include both Geometridae and Erebidae (Eiseman & Charney, 2010; http://v4.boldsystems.org). Diagnosis. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 332); 2nd–10th antennal segments yellowish, contrasting with re-

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

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maining darker segments; clypeus obtuse apically and not protruding in lateral view (Fig. 334); precoxal area largely smooth, at most with some aciculae or punctures medially (Fig. 327); tegulae brown; lobes of mesoscutum finely coriaceous-granulate and rather dull, with satin sheen; vein 1-CU1 of fore wing 0.4–0.6 × vein 2-CU1 (Fig. 332); length of hind femur 3.5–3.8 × its maximum width (Fig. 331); hind tarsal claws small and only yellowish or brownish setose (Fig. 338); body of ♂ completely black, antenna completely blackish, dark brown or with some segments yellowish subbasally and 4th–6th tergites with a setose medio-dorsal depression; length of fore wing 3.7–5.0 mm. Description. Holotype of A. freyi, ♂, length of fore wing 4.5 mm, of body 5.3 mm. Head. Antenna incomplete, (length of antenna of ♀ from Santon Downham 1.2 × fore wing, its subapical segments rather robust: Fig. 336); frons smooth anteriorly and with coarse curved striae posteriorly; OOL 2.7 × diameter of posterior ocellus, and rather regularly and rather coarsely striate; vertex transversely striate, rather shiny; clypeus narrow, strongly curved dorsal margin, rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 334); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 332); length of eye 1.4 × temple in dorsal view (Fig. 333); vertex behind stemmaticum transversely rugose; clypeus near lower level of eyes; face coarsely transversely rugose; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes very densely coriaceous-granulate, with vague micro-reticulate sculpture, matt; precoxal area of mesopleuron largely smooth (except some micro-sculpture) medially, rather depressed; remainder of mesopleuron largely smooth, except some punctures and antero-dorsally coarsely rugose; scutellum superficially granulate and with some punctures; propodeum coarsely rugose-reticulate and medio-longitudinal carina nearly complete. Wings. Fore wing: r 0.6 × 3-SR (Fig. 325); 1-CU1 horizontal and somewhat widened, 0.45 × 2-CU1; r-m 0.65 × 3-SR; 2nd submarginal cell rather short (Fig. 325); cu-a vertical, straight; 1-M rather curved posteriorly; 1-SR short and narrow; surroundings of M+CU1, 1-M and 1-CU1 evenly setose; m-cu subvertical, slightly diverging from 1-M posteriorly. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 326); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 80:57; 1r-m 0.6 × 1-M; 1-M straight. Legs. Tarsal claws small but robust and only yellowish setose (Fig. 338); hind coxa largely rugulose-granulate; hind trochantellus medium-sized; length of hind femur and basitarsus 3.8 and 7.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus; fore femur 4.8 × as long as wide. Metasoma. First tergite evenly convex, 1.3 × as long as wide apically; 1st and 2nd tergites with weak medio-longitudinal carina and together with basal half of 3rd tergite densely and finely longitudinally rugose; medio-basal area of 2nd tergite narrow but rather distinct (Fig. 329); 2nd suture rather shallow; remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; 4th–6th tergites with a setose medio-dorsal depression. Colour. Dark brown or blackish; palpi dark brown; mesopleuron with reddish brown streak; legs yellowish brown but tarsi, apex of hind femur (and indistinctly api-

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

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ces of fore and middle femora, and of tibiae) and base of hind coxa infuscate; tegulae and pterostigma brown; wing membrane slightly infuscate. Variation. Maximum width of marginal cell of hind wing 2.0–2.6 × its width near hamuli (Fig. 326); vein 1-CU1 of fore wing 0.4–0.6 × vein 2-CU1; length of hind femur 3.5–3.8 × its maximum width; length of 1st tergite 1.0 (♀)–1.3 (♂) × its apical width. Propodeum and metapleuron posteriorly, 1st and 2nd tergites and base of 3rd tergite of ♀ more or less brown, basal third of antenna (except scapus and base of pedicellus) brownish yellow or yellowish brown and ovipositor sheath rather slender, with long setae and apically narrowed (Fig. 323). Antennal segments: ♀ 38(1), 39(1), 41(1), 43(1), 44(1), 45(1); ♂ 36(1), 39(3), 40(6), 41(3), 42(7), 43(2), 44(2), 45(1). The number of antennal segments appear to be comparable between the sexes. Males have 1st–3rd metasomal tergites completely black, and basal third of antenna completely blackish, dark brown or with some subbasal segments yellowish. The male apical tergites (besides pores, see above) are type 1, setae rather dense, no fringe observed and probably absent (Fig. 341). Distribution. *Austria, *British Isles (England), Bulgaria, Czech Republic, Finland, France, *Germany, Hungary, *Netherlands, *Poland, *Spain, *Sweden, *Turkey. Aleiodes gasterator (Jurine, 1807) Figs 343–364 Bracon gasterator Jurine, 1807: 118, pl. 8. [examined]. Rogas gasterator; Shenefelt, 1975: 1230–1231; Zaykov, 1980a: 112. Rogas (Rogas) gasterator; Tobias, 1976: 86, 1986: 81 (transl.: 133) p.p. Aleiodes (Neorhogas) gasterator; Papp, 1991a: 91 p.p. Aleiodes (Chelonorhogas) gasterator; Falco et al., 1997: 60; Ghahari et al., 2011: 267; Rastegar et al., 2012: 3; Farahani et al., 2015: 243. Aleiodes gasterator; Bergamasco et al., 1995: 5; Zaldivar-Riverón et al., 2004: 234; Papp, 2005: 176. Bracon dimidiatus Spinola, 1808: 123–124. Syn. nov. Aleiodes dimidiatus; Bergamasco et al., 1995: 5. Rogas (Rogas) dimidiatus: Tobias, 1976: 86; 1986: 81 (transl.: 134) p.p. Rogas dimidiatus; Zaykov, 1980a: 112. Aleiodes (Neorhogas) dimidiatus; Papp, 1991a: 90 p.p. Aleiodes (Chelonorhogas) dimidiatus; Samartsev & Belokobylskij, 2013: 765; Farahani et al., 2015: 242. Rhogas (Rhogas) dimidiatus var. turkestanicus Telenga, 1941: 184, 409; Shenefelt, 1975: 1225 [examined]. Syn. nov. Type material. Holotype of A. gasterator, ♀ (Museum Genève), “[? Switzerland], gasterator J.”, “Typus”, “Bracon gasterator Jur., Type”, “Type du g. Rogas [= incorrect]”, “vu par [R.D.] Shenefelt, U.S.A., 1967” (metasoma on separate card and pin). Lectotype of A. turkestanicus here designated, ♀ (ZISP), “[Turkmenistan], Transcaspia,

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Bajram-aly, 17.viii.1930, T. Boguj/311, i.s.”, “Rhogas dimidiatus Spin. var. turkestanica [sic!] nov., N. Telenga det.”. Additional material. Albania, Cyprus, France (including Corsica), Greece (including Crete), Italy (including Sardinia, Sicily), North Macedonia, Portugal (including Madeira), Spain (including Mallorca, Menorca, Tenerife), Tunisia, Turkey, [Iraq, Jordan, Syria]. Specimens in BMNH, BZL, CNC, MSC, MTMA, NMS, RMNH, ZSSM. Widespread in the Mediterranean region, where it tends to replace A. ruficornis. Molecular data. MRS046 (France), MRS048 (France), MRS892 (Spain). Biology. Collected chiefly in May–July and September–November, but specimens have occurred in every month of the year. Plurivoltine; there is no indication of a unique overwintering mode in the material seen. Reared from low-feeding Noctuidae: Agrotis segetum (Denis & Schiffermüller) (6 [6 BMNH], Spain), Agrotis sp. (1 [BMNH], Cyprus; W.R. Ingram), mixed Agrotis and Spodoptera littoralis (Boisduval) (1 [BMNH], Cyprus; W.R. Ingram). The two mummies seen are rather different (Figs 345, 346), though it may be that neither overwintered; a small, pale and relatively slender one from Agrotis sp. produced a small male, while a more normal sized individual emerged from the larger, dark and stout mummy whose host was (even) less certain. Even in the latter case, the mummy is less keeled, less lined with silk and much more in relation to the size of the emerging adult than is the situation with the overwintering mummies of the A. grassator/ carbonarius/ carbonaroides/ ruficornis complex. The appearance of both mummies suggest that they would normally form below ground. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 355); OOL of ♀ 1.2–1.6 × as long as diameter of posterior ocellus (Fig. 356) and distinctly rugose or rugulose; antennal segments of ♀ 29–39, of ♂ 36–46(–51) (usually 39–43); antenna of ♀ 0.8–0.9 × fore wing; length of malar space of ♀ 0.5–0.6 × height of eye in lateral view (Fig. 357); clypeus thick apically and not protruding anteriorly (Fig. 357); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; posterior half of notauli deep; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 347); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 347); hind tarsal claws yellowish or brownish bristly setose; third tergite with (faint) curved or antero-medially transverse rugulae or striae (Fig. 364) or largely smooth (sometimes with only longitudinal striae baso-laterally), often with distinct punctures laterally; hind femur at least apico-dorsally dark brown or black; inner and/ or dorsal side of hind tibia (largely) yellowish or red; tegulae usually (partly) dark brown; pale males have nearly always frons medially and stemmaticum black; palpi usually brownish or yellowish, sometimes dark brown; 3rd metasomal tergite frequently partly or completely reddish or yellowish; 4th and 5th tergites black. Description. Redescribed ♀ (RMNH) from France (Isle sur le Sorque). Length of fore wing 4.9 mm, of body 6.1 mm. Head. Antennal segments of ♀ 35, length of antenna 0.9 × fore wing, its subapical segments robust (Fig. 359), 4th segment 1.2 × longer than wide; frons with coarse curved rugae; OOL 1.6 × diameter of posterior ocellus, and densely rugose or rugulose;

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Figures 343–346. Aleiodes gasterator (Jurine), ♀, France, Les Constants, but 345 and 346 from Cyprus 343 habitus lateral 344 ovipositor sheath lateral 345 mummy of Agrotis sp. 346 mummy of Agrotis sp. or Spodoptera littoralis (Boisduval).

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Figures 347–359. Aleiodes gasterator (Jurine), ♀, France, Les Constants 347 fore wing 348 hind wing 349 mesosoma lateral 350 mesosoma dorsal 351 metasoma dorsal 352 fore femur lateral 353 hind femur lateral 354 base of antenna 355 head anterior 356 head dorsal 357 head lateral 358 outer hind tarsal claw 359 apex of antenna.

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vertex densely rugose, shiny; clypeus densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 357); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 355); length of eye 1.3 × temple in dorsal view (Fig. 356); vertex behind stemmaticum punctate-rugose; clypeus just below lower level of eyes; length of malar space 0.5 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, interspaces largely smooth with superficial granulation, shiny; precoxal area of mesopleuron evenly vermiculate-rugose medially, but only sparsely punctate posteriorly; metanotum without median carina; scutellum rather flat, sparsely punctate, but rugose laterally; propodeum coarsely vermiculaterugose, medio-longitudinal carina nearly complete, and angulate latero-posteriorly. Wings. Fore wing: r 0.35 × 3-SR (Fig. 347); marginal cell distinctly ending basad of level of apex of vein 3-M; 1-CU1 horizontal, slender, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 347); cu-a inclivous, posteriorly curved; 1-M rather curved posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened, but slightly basally, its apical width 2.3 × width at level of hamuli (Fig. 348); 2-SC+R subquadrate; m-cu largely absent; M+CU:1-M = 28:19; 1r-m 0.6 × 1-M. Legs. Tarsal claws robust and with only bristly brownish setae (Fig. 358); hind coxa largely densely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.2 and 5.8 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex medially and 0.9 × as long as wide apically; 1st and nd 2 tergites with medio-longitudinal carina and coarsely longitudinally rugose; mediobasal area of 2nd tergite short and rather distinct (Fig. 351); 2nd suture deep; subbasally 3rd tergite with faint curved striae and medially transverse (Fig. 364); remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically oblique, dorsally longer than ventrally (Fig. 344). Colour. Black; face (except medio-dorsally), malar space, dorsal half of temple, frons largely laterally, notauli, mesoscutum laterally, scutellum, pronotum postero-dorsally, mesopleuron dorsally and posteriorly, metapleuron largely, 1st and 2nd metasomal tergites and base of 3rd tergite orange brown; palpi and humeral plate and veins of hind wing yellowish brown; tegula rather dark brownish; ventral half of temple largely, dorso-apical patch of hind femur, pterostigma and veins of fore wing dark brown; fore wing membrane slightly infuscate, of hind wing subhyaline. Variation. A very colour-variable species; head and mesoscutum of female may be largely black (nominate form) or reddish (= “A. dimidiatus/var. turkestanicus”, but especially the mesoscutum may be intermediate). Especially males may have the hind coxa black and most of hind tibia dark brown, sometimes the entire leg is nearly completely black or dark brown. Antennal segments: ♀ 29(1), 31(2), 32(3), 33(13), 34(13), 35(15), 36(9), 37(9), 38(3), 39(2); ♂ 36(2), 37(3), 38(4), 39(8), 40(13), 41(9), 42(5), 43(11), 44(7), 45(2), 46(2). Additionally, an exceptionally large male with 50 segmented antennae from Cyprus (BMNH) appears to belong to this species, as does a female from Spain (RMNH) with 41 antennal segments, basal half of 3rd tergite largely obliquely rugose

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Figures 360–364. Aleiodes gasterator (Jurine), ♂, Italy, Livorno, but 364 of ♀, France 360 habitus lateral 361 antenna 362 base of antenna 363 apex of antenna 364 3rd metasomal tergite dorsal.

and blackish scapus. On average males have ca 7 more antennal segments than females. Males are very similar with apical tergites type 1–2, setae rather sparse and with evident

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but short fringe (Fig. 360), hind tibial spurs often blunt apically and 3rd tergite remotely punctate basally. Distribution. *Albania, *Cyprus, France (including Corsica), Greece (including Crete), *Iraq, Italy (including Sardinia, Sicily), *Jordan, *North Macedonia, *Portugal (including Madeira), Spain (including Mallorca, Menorca and Tenerife), *Syria, *Tunisia, *Turkmenistan, Turkey. New synonymy. The new synonymy of Rhogas dimidiatus var. turkestanicus Telenga, 1941, is based on direct comparison of the types of both taxa. The identity of Bracon dimidiatus Spinola, 1808, is problematic because the holotype from Italy (Genoa) is lost and the original description is far too incomplete for an easy identification. Its colour pattern (head completely yellowish, hind tibia and 3rd tergite reddish) does not fit with A. ruficornis (Herrich-Schäffer); if the head is largely reddish brown then the temple ventrally and malar space remain blackish. This pattern agrees better with that of pale specimens of A. gasterator (named as A. dimidiatus var. turkestanicus Telenga, 1941). Aleiodes ruficornis occurs also in Italy, but its females have the head partly black ventrally, the apex of the hind tibia dark brown and most of the 3rd metasomal tergite black. Therefore, we synonymise Bracon dimidiatus with A. gasterator (syn. nov.). The holotype of Bracon gasterator Jurine, 1807, has the 3rd metasomal tergite finely curved (nearly circular) aciculate or striate basally, palpi (as far as present) pale brownish, maximum with of hypoclypeal depression 0.45 × minimum width of face, vein 1-CU1 of fore wing half as long as vein 2-CU1 and 4th antennal segment 1.3 × as long as wide. Aleiodes arnoldii sensu Farahani et al. (2015) concerns a species closely related to A. gasterator (Jurine) having basal half of 3rd tergite coarsely longitudinally rugose, antenna of ♀ with 30–35 segments (of ♂ 36), head linearly narrowed ventrally in anterior view and subbasal antennal segments of ♀ slightly slenderer. Aleiodes grassator (Thunberg, 1822) Figs 365–384 Ichneumon grassator Thunberg, 1822: 256 [examined]. Rogas grassator; Shenefelt, 1975: 1232. Aleiodes (Neorhogas) grassator; Papp, 1991a: 86. Aleiodes grassator; Zaldivar-Riverón et al., 2004: 234; Quicke et al., 2014: 240; Butcher et al., 2014: 458. Rhogas grassator ab. thoracicus Hellén, 1927: 24; Shenefelt, 1975: 1232 (unavailable name). Rogas (Rogas) flavipalpis Thomson, 1892: 1672 [examined]. Aleiodes flavipalpis; Papp, 1991a: 86 (as synonym of A. grassator). Rogas alpinus Thomson, 1892: 1671; Shenefelt, 1975: 1217 [examined]. Syn. nov. Aleiodes alpinus; Papp, 1991a: 90 (as synonym of A. dimidiatus). Type material. Holotype of A. grassator, ♀ (ZMUU), unlabelled. Lectotype of A. flavipalpis, ♀ (ZIL), “åre”, “Sverige, Ǻreskutan I Jemtland/teste Papp, 1983”, “Lectotypus Rogas flavipalpis Thomson, 1899 [sic!], Papp, 1983”, “Aleiodes grassator Thb., det. Papp J., 1983”. Lectotype of A. alpinus, ♀ (ZIL), “[Norway:] Dovre”, “alpinus m.”,

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Figures 365–367. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 365 habitus lateral 366 antenna 367 ovipositor sheath lateral.

“Lectotypus Rogas alpinus Thoms., 1891, ♀. Papp, 1983”, “Aleiodes dimidiatus var. alpinus Th., ♀, det. Papp J., 1983”. Additional material. Austria, British Isles (England: V.C. 70; Scotland: V.C.s 83, 85, 88, 89, 97, 103; Finland, France (both Alps and Pyrenees), Italy, Germany, Norway, Romania, Sweden, Switzerland. Specimens in BMNH, BZL, MRC, MTMA, NMS, RMNH, SDEI, ZIL, ZMUU, ZSSM. This is essentially a montane grassland species, though occurring at low altitudes in northern Europe. Molecular data. MRS215 (UK), MRS721 (UK), MRS725 (UK). Biology. Collected in (April)May–July. Univoltine, overwintering in the mummy. Reared from the noctuid Cerapteryx graminis (Linnaeus) (9: K.P. Bland, M.J.W.

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Cock, M.R. Shaw) and from mummies compatible with that (3), and it may be strictly monophagous. The known host overwinters in the egg stage, and feeds on Poaceae near ground level. The tough dark brown mummy is formed on or below the soil surface and seems spectacularly too large for the adult that will emerge from it (Fig. 384). It is more or less cylindrical, though with a pronounced lateral keel, and well-lined with silk. The cocoon chamber occupies most of the abdominal segments. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 375); OOL of ♀ ca twice as long as diameter of posterior ocellus (Fig. 376) and distinctly rugose or rugulose; length of 4th antennal segment of ♀ 0.7–0.9 × its width (Fig. 378; in ♂ 0.9–1.0 ×); clypeus thick apically and not protruding anteriorly (Fig. 377); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 368); vein 1-CU1 of fore wing 0.5–0.6 × as long as vein 2-CU1 (Fig. 368); hind tarsal claws robust (Fig. 380) and yellowish or brownish bristly setose; hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; pale males have whole frons and stemmaticum yellowish; palpi dark brown or blackish, rarely brown; 3rd metasomal tergite only antero-laterally reddish or yellowish; 4th and 5th tergites black. Description. Redescribed ♀ (RMNH) from Finland (Sb: Leppävirta). Length of fore wing 4.6 mm, of body 5.7 mm. Head. Antennal segments of ♀ 39, 4th segment 0.8 × longer than wide (Fig. 378); antenna as long as fore wing, its subapical segments robust (Fig. 379); frons with coarse curved rugae and shiny; OOL 1.9 × diameter of posterior ocellus and rugose; vertex rugose and shiny; face rugose-punctate; clypeus rugose; ventral margin of clypeus rather thick and not protruding forwards (Fig. 377); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 375); length of eye 1.1 × temple in dorsal view (Fig. 376); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view. Mesosoma. Mesoscutal lobes moderately punctate, laterally interspaces mainly smooth, medially superficially granulate and rather shiny; precoxal area of mesopleuron coarsely rugose medially and largely smooth posteriorly; remainder of mesopleuron mainly punctate, but dorsally coarsely rugose; scutellum flat, sparsely finely punctate and only anteriorly with lateral carina; propodeum coarsely rugose, medio-longitudinal carina present on anterior half, rounded posteriorly and dorsal part approx. as long as posterior part. Wings. Fore wing: r 0.3 × 3-SR (Fig. 368); marginal cell ends basad of level of apex of 3-M; 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell robust (Fig. 368), 3-SR 1.3 × as long as 2-SR; cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR similar to 1-M and medium-sized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width twice width at level of hamuli (Fig. 369); 2-SC+R subquadrate; m-cu short; M+CU:1-M = 27:18; 1r-m 0.7 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 380); hind coxa largely densely punctate, but dorsally with some rugae; hind trochantellus robust;

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Figures 368–380. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 368 fore wing 369 hind wing 370 mesosoma lateral 371 mesosoma dorsal 372 metasoma dorsal 373 fore femur lateral 374 hind femur lateral 375 head anterior 376 head dorsal 377 head lateral 378 base of antenna 379 apex of antenna 380 outer hind tarsal claw.

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Figures 381–384. Aleiodes grassator (Thunberg), ♂, Scotland, Isle of Coll 381 habitus lateral 382 base of antenna 383 apex of antenna 384 mummy of ?Cerapteryx graminis (Linnaeus).

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length of hind femur and basitarsus 3.1 and 3.9 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite rather flattened, 0.8 × as long as wide apically; 1st and nd 2 tergites with medio-longitudinal carina and coarsely longitudinally rugose; medio-basal area of 2nd tergite wide triangular and short (Fig. 372); 2nd suture deep and crenulate; basal half of 3rd tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured; 4th tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 367). Colour. Orange brown; apical two thirds of antenna, labial palp, patch on hind femur dorso-apically, posterior patch of 2nd tergite and telotarsi, dark brown; head, mesosoma (except side of pronotum postero-dorsally and pair of latero-posterior patches of propodeum), 3rd–7th tergites (except antero-lateral corners of 3rd tergite) black; maxillary palp, basal third of antenna, tegulae and remainder of legs rather pale yellowish brown; veins and pterostigma dark brown; wings distinctly infuscate but hind wing less than fore wing. Variation. Basal third or half of antenna of ♀ pale yellowish brown; head partly and mesosoma anteriorly of ♀ dark orange brown or both entirely black; 3rd tergite longitudinally striate or rugulose basally (sometimes narrowly so), without curved sculptural elements (Fig. 372), except sometimes some weak transverse striae occasionally present at extreme apex. Males are always darker than females; mainly black with legs mainly dark brown or blackish (Fig. 381). Antennal segments: ♀ 36(2), 37(4), 38(6), 39(10), 40(6), 41(1); ♂ 47(1), 48(2), 49(2), 50(3), 51(2), 52(4), 53(1), 60(1). On average males have ca 12 more antennal segments than females. Males have 2nd submarginal cell slightly shorter than of females, temple and face long setose, malar space 0.5–0.7 × length of eye in lateral view, apical tergite type 1, rarely type 2, setae rather dense, fringe not observed and probably absent (Fig. 381). The superficial granulosity of 3rd tergite and mesoscutum may be absent. Distribution. *Austria, British Isles (England, Scotland), Finland, *France, *Ireland, *Italy, *Germany, Norway, *Romania, Sweden, *Switzerland. New synonymy. The synonymy of Rogas alpinus Thomson, 1892, with Aleiodes grassator (Thunberg, 1822) is based on direct comparison of the types listed above. Notes. Although males of A. carbonaroides are generally easily distinguished from A. grassator through being black, it is possible that lighter forms occur which would be difficult to recognise. Also, females of A. carbonaroides are similar in colour to those of A. grassator. Therefore, specimens collected at low altitude away from northern areas that appear, on other characters, to be A. grassator might well really be A. carbonariodes. See also remarks under A. carbonarius and A. ruficornis. Aleiodes hemipterus (Marshall, 1897) Figs 385–408 Rhogas hemipterus Marshall, 1897: 137. Rogas hemipterus; Shenefelt, 1975: 1233.

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Aleiodes hemipterus; Papp, 1990: 90, 2003: 138 (lectotype listed). Aleiodes (Chelonorhogas) hemipterus; Belokobylskij and Kula 2012: 35–38. Type material. Lectotype here designated, ♀ (BMNH), “Tunisie, Sicard”, “Type, H.T.”, “B.M. Type Hym. 3.c.243”, “B.M. Type Hym., Rhogas hemipterus Marshall, 1896”, “hemipterus Marsh.”, “Marshall coll. 1904-120”. Paralectotype: 1 ♀ (MTMA), “Tunisie”, “hemipterus M. coll. Marshall”, “Paratypus Rhogas hemipterus Marshall 1897 sp. n. % des Papp J. 1986”, “Hym. Type No 10582 Museum Budapest”, “Aleiodes ♀ hemipterus Mshl. Det. Papp J. 1991”. Additional material. 1 ♀ + 1 ♂ (MNHN), “[N. Tunisia:] Teboursouk”, “Rhogas hemipterus Mrsh.”, “Muséum Paris, Coll. J. de Gaulle, 1919” [figured specimens]; 1 ♂ [but metasoma missing] (BMNH) “Rabat, Maroc [= Morocco], coll. Thery”. Molecular data. None. Biology. Unknown. The specimens seen do not have phenological data, and we have not seen reared material. As the female is brachypterous it is likely that the host will be found near the ground. Diagnosis. Maximum width of hypoclypeal depression approx. 0.6 × minimum width of face (Fig. 396); OOL of ♀ 1.2–1.3 × as long as diameter of posterior ocellus, of ♂ 0.9 × (Figs 397, 408) and finely reticulate-rugose; stemmaticum protruding dorsally; antennal segments of ♀ 46–50 (of ♂ unknown), penultimate segments approx. as long as wide and antenna 0.8 × as long as body; length of malar space 0.3–0.4 × height of eye; mesoscutal lobes densely rugose or rugulose and rather matt, middle lobe with medio-longitudinal ridge or carina, of ♀ surrounded by shallow grooves (Fig. 392); propodeum angulate posteriorly (Fig. 392); ♀ brachypterous and ♂ macropterous; marginal cell of hind wing of ♀ hardly widened (Fig. 390) and of ♂ distinctly widened apically (Fig. 403); hind tarsal claws with rather conspicuous pale brown pecten (Fig. 399); 1st–3rd metasomal tergites very densely and finely longitudinally rugose; 1st tergite of ♀ 1.0–1.1 × its apical width, of ♂ approx. 1.4 ×; 2nd metasomal suture of ♀ hardly impressed but densely costate, of ♂ medium-sized. Description. Lectotype, ♀, length of hind wing 1.7 mm (fore wing missing, but in other specimens ca one-third longer than hind wing and 2.2 mm, brachypterous), of body 7.8 mm. Head. Antenna incomplete, segments robust; frons largely striate-rugose (but transversely costate in figured ♀); OOL 1.2 × diameter of posterior ocellus, (as vertex) rather finely and densely reticulate-rugose and rather dull; clypeus rugose; ventral margin of clypeus rather thick ventrally and rather forward protruding (Fig. 398); width of hypoclypeal depression 0.6 × minimum width of face and long (Fig. 396); length of eye 1.2 × temple in dorsal view (Fig. 397); vertex behind stemmaticum rather coarsely reticulate-rugose; clypeus near lower level of eyes; length of malar space 0.35 × length of eye in lateral view. Mesosoma. Antescutal depression distinct; mesoscutal lobes coarsely rugose-punctate (but superficial in figured ♀) and rather matt, middle lobe of pair of submedian grooves (Fig. 392); nearly entire mesopleuron (except minute smooth speculum)

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Figures 385–389. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 385 habitus lateral 386 antenna 387 apex of antenna 388 base of antenna 389 ovipositor sheath lateral.

densely and coarsely reticulate-rugose; scutellum coarsely rugose and without lateral carinae; propodeum coarsely vermiculate-rugose, dorsal face long and rectangularly angulate postero-laterally (Fig. 392). Wings. Fore wing brachypterous, hardly surpassing propodeum (Marshall, 1897): (of ♀ from Tunisia r 0.2 × 3-SR; 1-CU1 distinctly widened and oblique, 0.4 × 2-CU1; r-m 0.8 × 3-SR; 2nd submarginal cell medium-sized (Fig. 390); cu-a short, vertical, straight; 1-M straight posteriorly; 1-SR widened; 1st subdiscal cell open apically and posteriorly; surroundings of M+CU1, 1-M and 1-CU1 setose; 2m-cu present as curved and only pigmented vein). Hind wing brachypterous: marginal cell reduced, sinuate and apically narrowed (Fig. 390); 2-SC+R quadrate and widened; m-cu absent; M+CU:1-M = 27:13; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous pale brown pecten, remaining far from apical tooth and much shorter (Fig. 399); hind coxa finely and densely reticulate-

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Figures 390–399. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 390 wings 391 mesosoma lateral 392 mesosoma dorsal 393 metasoma dorsal 394 fore femur lateral 395 hind femur lateral 396 head anterior 397 head dorsal 398 head lateral 399 outer hind tarsal claw.

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Figures 400–408. Aleiodes hemipterus (Marshall), ♂, Tunisia, Teboursouk 400 habitus lateral 401 fore femur lateral 402 fore wing 403 hind wing 404 mesosoma dorsal 405 metasoma dorsal 406 hind femur lateral 407 head anterior 408 head dorsal.

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rugose; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 6.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st–3rd tergites regularly finely and very densely longitudinally rugose, rather matt and medio-longitudinal carina rather weak; medio-basal area of 2nd tergite triangular and short (Fig. 393); 2nd suture shallow; 4th and subsequent tergites superficially punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with mediumsized setae and apically truncate (Fig. 389). Colour. Brown; stemmaticum and ovipositor sheath black; frons, vertex medially, occiput, femora, propodeum, 1st and 2nd tergites somewhat infuscate; wing membrane subhyaline. Variation. Length of body 7.8–8.8 mm. Antennal segments: ♀ 46(1), 50(1); ♂ unknown. Male is normally winged (vein 3-SR of fore wing 1.5 × vein 2-SR, vein r 0.3 × 3-SR, vein cu-a oblique, vein 1-CU1 narrow and 0.3 × vein 2-CU1) and has marginal cell of hind wing 2.2 × wider than width at level of hamuli (with vein m-cu present anteriorly, 2-SC+R quadrate and M+CU:1-M:1r-m = 40:30:26). Apical metasomal segments of ♂ type 1 and sparsely setose. Distribution. Morocco, Tunisia. Notes. Marshall (1897) based his description on three females from Tunisia. Papp (2003) listed a female in BMNH as lectotype, but this was not accepted as a designation by Belokobylskij & Kula (2012). Therefore, the redescribed female above is here designated formally as lectotype and is the same specimen intended to become lectotype by Papp (2003). Aleiodes hirtus (Thomson, 1892) Figs 409–427 Rogas hirtus Thomson, 1892: 1672; Shenefelt, 1975: 1233. Aleiodes (Neorhogas) hirtus; Papp, 1985a: 153, 155, 161 (lectotype designation and as synonym of A. pallidicornis), 1991a: 75 (id.). Aleiodes hirtus; Papp, 2005: 177 (as synonym of A. pallidicornis). Rhogas hirtus ab. coloratus Hellén, 1927: 23; Shenefelt, 1975: 1233; Papp, 2005: 177 (as synonym of A. pallidicornis) (unavailable name; not Rogas coloratus Motschulsky, 1863). Type material. Lectotype of A. hirtus, ♂ (ZIL), “[Norway], Norl.”, “hirtus m.”, “Funne I Norrland, teste Papp J., 1983”, “Lectotypus”, “Rogas hirtus Thms. 1891, ♂, Papp J., 1983”, “Aleiodes pallidicornis HS ♂, det. Papp J., 1983”. Additional material. Austria, Belgium, British Isles (England: V.C.s 26, 29, 32, 62; Scotland: V.C. 78; Ireland: V.C. H12), Bulgaria, Czech Republic, Finland, France, Germany, Hungary, Netherlands (DR: Borger), Norway, Romania, Russia, Serbia, Slovakia, Switzerland, Ukraine, [? Mongolia]. Specimens in BMNH, BZL, CMIM, CNC, FMNH, MRC, MSC, MTMA, NMS, RMNH, SDEI, UNS, USNM, UWIM, ZIL, ZSSM.

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Figures 409, 410. Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 409 habitus lateral 410 ovipositor sheath lateral.

Molecular data. MRS619 (UK), MRS882 (Romania), MRS883 (Romania). Biology. Unknown. Collected in June–August, presumably univoltine but the mode of overwintering is unclear. Most British sites are more or less damp and calcareous grasslands, approaching fens. We have not seen reared material, but the clypeus suggests that the mummy will form in the soil. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 419); OOL of ♀ approx. 1.3 × as long as diameter of posterior ocellus (Fig. 420; in ♂ approx.1.6 ×) and punctate-rugose; ventral margin of clypeus rather thick but rather strongly protruding forwards (Fig. 421; stronger in in ♂: Fig. 426); mesoscutal lobes largely smooth, only indistinctly punctulate and shiny; precoxal area finely punctate and often with some rugulae (Fig. 413); vein 1-CU1 0.3–0.6 × vein 2-CU1 of fore wing (Fig. 411); hind tarsal claws with rather conspicuous brownish pecten (Fig. 424); length of inner spur of hind tibia 0.5–0.7 × hind basitarsus; palpi dark brown; basal half of metasoma at least partly reddish or orange and 1st tergite rather

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Figures 411–424. Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 411 fore wing 412 hind wing 413 mesosoma lateral 414 mesosoma dorsal 415 metasoma dorsal 416 fore femur lateral 417 hind femur lateral 418 antenna 419 head anterior 420 head dorsal 421 head lateral 422 base of antenna 423 apex of antenna 424 inner hind tarsal claw.

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coarsely sculptured; setae of body of ♂ (but of ♀ mainly its head) conspicuous and dense (Fig. 426); hind coxa black; hind femur largely or completely reddish or brownish; basal half of hind tibia usually (pale) yellowish or yellowish brown, but sometimes uniformly reddish and of ♂ ivory. In the past this species has been frequently misidentified as “Rogas dimidiator” or “Rogas gasterator”. Description. Redescribed ♂ (RMNH) from Germany (Graswang). Length of fore wing 6.2 mm, of body 8.0 mm. Entire body with long whitish setae. Head. Antennal segments of ♂ 60, length of antenna 1.3 × fore wing, its subapical segments somewhat longer than wide; frons medially largely smooth, laterally with some fine curved rugae; OOL 1.6 × diameter of posterior ocellus, and punctate-rugose, POL approx. half as long as diameter of ocellus; vertex spaced rugose, shiny; clypeus punctate; ventral margin of clypeus rather thick but distinctly protruding forwards (Fig. 421); width of hypoclypeal depression 0.55 × minimum width of face (Fig. 419); length of eye 1.1 × temple in dorsal view (Fig. 420), temples conspicuously setose (Figs 419, 426); vertex behind stemmaticum rugose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Pronotum rugose and anteriorly oblique, without antescutal depression; mesoscutal lobes large smooth and shiny, only indistinctly punctulate and densely setose; precoxal area of mesopleuron punctulate and medially with some superficial rugulae; remainder of mesopleuron sparsely punctate; scutellum sparsely punctate and largely smooth, posteriorly with lateral rugae; propodeum rather convex and coarsely rugose, its medio-longitudinal carina only in anterior half of propodeum. Wings. Fore wing: r 0.3 × 3-SR (Fig. 411); 1-CU1 horizontal, 0.45 × 2-CU1; r-m 0.7 × 3-SR and as long as 2-SR; 2nd submarginal cell medium-sized (Fig. 411); cu-a vertical, straight; 1-M straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened (but less so basally: Fig. 412), its apical width 2.5 × width at level of hamuli; 2-SC+R short longitudinal; m-cu narrowly indicated; M+CU:1-M = 9:7; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized brownish pecten (Fig. 424); hind coxa pimply punctate and shiny; hind trochantellus robust; length of hind femur and basitarsus 3.8 and 6.0 × their width, respectively; length of inner hind spur 0.65 × hind basitarsus (0.6 × in ♀). Metasoma. First tergite evenly convex, approx. as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and densely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite minute (Fig. 415); 2nd suture deep and moderately crenulate; basal half of 3rd tergite finely rugose, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; 4th – 6th tergites with long setae and flat. Colour. Black; legs (except black coxae, trochanters and trochantelli, 1st and 2nd metasomal tergites (but base of 1st tergite partly infuscate) and base of 3rd tergite orange brown; vaguely near base of femora, telotarsi, apex of hind femur, apical half of hind tibia, hind tarsus largely black or blackish; basal half of hind tibia pale yellow; palpi, tegulae, veins and pterostigma dark brown; wing membrane slightly infuscate.

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Figures 425–427. Aleiodes hirtus (Thomson), ♂, England, Chippenham Fen 425 habitus lateral 426 head and anterior part of mesosoma lateral 427 propodeum and 1st –3rd metasomal segments lateral.

Variation. Hind femur varies from apically black to entirely orange. Propodeum can be partly orangish in posterior part. Usually both sexes have hind trochanter (often also trochantellus) more or less infuscate and darker than the orange part of the hind femur, but this is scarcely evident in a series from S. Russia (MRC, NMS). Hind coxa varies from orange to black. Female is similar to the more distinctive male but is less conspicuously setose (Figs 419–421) and its ovipositor sheath is wide, with long setae and apically truncate (Fig. 410). Precoxal sulcus smooth to superficially rugulose medially. A female and a male from Romania (NMS) are slightly different from British ones; ocelli approx. 1/5 larger and frons coarsely rugose posteriorly. This appears to be reflected by a small divergence in CO1 (2.75 %), but for the moment we treat them

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as belonging to A. hirtus. Antennal segments of ♀ 54(1), 55(2), 56(6), 57(6), 59(2), 60(2), 61(1), of ♂ 56(1), 58(1), 59(3), 60(3), 61(1). Apical tergites of male type (1–)2, setae rather sparse but long and glabrous stripe consequently not always evident and fringe present but poorly differentiated. A female from Mongolia (BZL) with completely black hind femur and base of hind tibia pale yellowish and with dark basal ring may be another very similar species. Distribution. *Austria, *Belgium, *British Isles (England, Scotland, Ireland), *Bulgaria, *Czech Republic, Finland, *France, *Germany, *Hungary, *Netherlands, Norway, *Romania, *Russia, *Serbia, *Slovakia, *Switzerland, *Ukraine. Aleiodes improvisus van Achterberg & Shaw, sp. nov. http://zoobank.org/BA464FC8-D95C-4CF8-A27F-788BB7D2D3C4 Figs 428–452 Type material. Holotype, ♀ (BMNH), “Austria: Tirol, Obergurgl, 2000 m, vii. [19]81, Day & Fitton”. Paratypes: 1 ♂ (NMS), “Switzerland: Valais, Aletschwald, 6000–7000 ft, 7–17.vi.1959, J.E. & R.B. Benson”; 1 ♂ (BMNH), “Switzerland: Valais, J.E. & R.B. Benson, B.M. 1935-581”, “Arolla, 6500 ft, 12.vi.1935”; 3 ♂ (BMNH, NMS), “Austria: Tirol Vent., 1860 m, vii.1981, Fitton & Day”; 1 ♂ (BMNH), “Austria: Tirol Vent (Winterstallen), 1750 m. vii.1981, Fitton & Day”. Molecular data. None. Biology. Unknown. Collected above the tree line in the Alps in June–July, and presumably univoltine. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 439); OOL of ♀ 1.8 × as long as diameter of posterior ocellus and densely rugose (Fig. 440); antenna of ♀ as long as fore wing; clypeus thick apically and not protruding anteriorly in lateral view (Fig. 441); lobes of mesoscutum densely punctate, interspaces largely finely coriaceous and with satin sheen; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 432); vein 1-CU1 of fore wing 0.3–0.6 × as long as vein 2-CU1; fore femur subparallel-sided (Fig. 437); hind tarsal claws yellowish or brownish bristly setose and with few dark brown or brown pectinal teeth submedially (Fig. 442); 3rd tergite longitudinally rugulose basally, without curved sculptural elements (Fig. 436); head of ♀ black; inner side of hind tibia of ♀ dark brown ventrally; palpi dark brown or blackish; hind trochanter and trochantellus largely dark brown; 2nd metasomal tergite of both sexes orange or dark reddish brown; 4th and 5th tergites black. Description. Holotype, ♀, length of fore wing 5.5 mm, of body 7.7 mm. Head. Antennal segments of ♀ 41, antenna as long as fore wing, its subbasal and subapical segments rather robust (Fig. 444); frons with curved rugae; OOL 1.8 × diameter of posterior ocellus, densely rugose and with satin sheen; vertex densely rugose (also behind stemmaticum), with satin sheen; clypeus transversely rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 441); width of hypoclypeal

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Figures 428–431. Aleiodes improvisus sp. nov., ♀, holotype 428 habitus lateral 429 ovipositor sheath lateral 430 right fore femur lateral 431 right hind femur lateral.

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depression 0.4 × minimum width of face (Fig. 439); eye as long as temple in dorsal view (Fig. 440); clypeus below lower level of eyes; length of malar space 0.55 × length of eye in lateral view; temple striate near eye, and remainder rugose; head with long setae. Mesosoma. Mesoscutal lobes densely punctate, interspaces largely finely coriaceous and with satin sheen; precoxal area of mesopleuron coarsely vermiculate-rugose medially, but posteriorly rugose; mesopleuron remotely punctate and shiny medially; metapleuron densely rugose and rather dull; scutellum largely smooth (except for spaced punctures), shiny and nearly flat, with lateral carina; propodeum coarsely rugose but antero-laterally rugulose, laterally dorsal face longer than posterior one, somewhat angulate laterally but without tubercles, and with complete medio-longitudinal carina. Wings. Fore wing: r 0.4 × 3-SR; marginal cell fairly short (Fig. 432); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.8 × 3-SR; 2nd submarginal cell medium-sized (Fig. 432); cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell linearly widened, its apical width 2.1 × width at level of hamuli (Fig. 433); 2-SC+R quadrate; m-cu narrowly pigmented; M+CU:1-M = 30:21; 1r-m 0.7 × 1-M. Legs. Tarsal claws mainly setose but submedially with four rather short and dark brown pectinal teeth (Fig. 442); fore femur largely parallel-sided and rather slender (Fig. 437); hind coxa punctate and shiny, but dorsally mainly rugose; hind trochantellus rather robust; length of hind femur and basitarsus 3.3 and 5.7 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite distinctly convex medially, its length 0.8 × apical width, robust and irregularly longitudinally rugose as 2nd tergite; both tergites with mediolongitudinal carina; medio-basal area of 2nd tergite triangular and small (Fig. 436); 2nd suture moderately deep and crenulate; basal half of 3rd tergite largely longitudinally striate, remainder of metasoma superficially micro-sculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide medially, with long setae and apically truncate (Fig. 429). Colour. Black; antenna (but only scapus partly yellowish), right fore coxa, trochanter, trochantellus, and femur (but left all yellowish brown except dark base of coxa and infuscated apex of femur), middle femur dorso-apically, middle coxa basally, hind trochanter, trochantellus and femur (but dorso-basally yellowish and left femur also ventrally), apical third of hind tibia (but left tibia yellowish ventrally), tegulae, pterostigma, veins largely, and metasoma ventrally largely dark brown; dorsal part of scutellum, 1st tergite laterally and narrowly medially and posteriorly, 2nd tergite and antero-laterally 3rd tergite orange brown; right fore tibia (except basally and left one yellowish brown) and tarsi more or less infuscate (but left fore tarsus only telotarsus dark brown); fore wing membrane somewhat infuscate, but hind wing nearly subhyaline. Variation. Eye of ♀ as long as temple in dorsal view (of ♂ 1.0–1.4 ×); length of malar space 0.5–0.6 × length of eye in lateral view; palpi black or largely dark brown; 1-CU1 0.3–0.6 × 2-CU1; length of fore wing 4.0–6.5 mm. Antennal segments: ♀ 41(1); ♂ 44(2), 49(2), 51(2). Male often has much darker legs (largely dark brown with coxae black as right legs of holotype, but legs are more extensively orange, including basal half

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Figures 432–444. Aleiodes improvisus sp. nov., ♀, holotype 432 fore wing 433 hind wing 434 mesosoma lateral 435 mesosoma dorsal 436 propodeum and 1st –3rd metasomal tergites dorsal 437 left fore femur lateral 438 left hind femur lateral 439 head anterior 440 head dorsal 441 head lateral 442 inner hind tarsal claw 443 base of antenna 444 apex of antenna.

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Figures 445–452. Aleiodes improvisus sp. nov., ♂, paratype, Switzerland (Arolla) 445 habitus lateral 446 fore femur lateral 447 hind femur lateral 448 base of antenna 449 head dorsal 450 4th–7th metasomal tergites dorsal 451 apex of antenna 452 4th–7th metasomal tergites lateral.

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of hind femur, in two paratypes) than female and scutellum black; metasoma similarly sculptured and coloured or also basal half of 3rd tergite orange brown or 1st tergite only posteriorly orange or only 2nd and 3rd tergites (except posteriorly) dark reddish brown; in the largest male paratype (Winterstallen) traces of inwardly curved sculpture are discernible posteriorly on the almost completely longitudinally rugose 3rd tergite; marginal cell of fore wing similar to ♀, with apical tergites type 1 and fringe not observed (Figs 450, 452). Distribution. Austria, Switzerland. Etymology. Improvisus is Latin for unexpected, unforeseen, because at first sight the specimens were expected to belong to A. gasterator or A. ruficornis. Notes. As suggested by its name this species can be easily confused with A. gasterator or A. ruficornis. It differs from A. gasterator mainly by being darker (subbasal antennal segments of ♀, hind trochanter and trochantellus, inner and dorsal side of hind tibia, parastigma) and somewhat higher number of antennal segments of ♀ (41 vs 29–39). Aleiodes ruficornis has an inflated fore femur (hardly or not inflated in A. improvisus), antenna of ♀ medium-sized (1.0–1.2 × fore wing vs 0.8–0.9 ×) and head of ♀ at least partly reddish brown. Aleiodes krulikowskii (Kokujev, 1898) Figs 453–466 Rhogas (Rhogas) krulikowskii Kokujev, 1898: 302; Telenga, 1941: 157–158. Rogas krulikowskii; Papp, 1971: 360; Shenefelt, 1975: 1235. Rogas (Rogas) krulikovskii; Tobias, 1976: 83. Rogas (Rogas) krulikowskii; Tobias, 1986: 77 (transl. 125) (lectotype designation). Aleiodes (Neorhogas) krulikowskii; Papp, 1985a: 153, 1991a: 84; Belokobylskij, 1996: 9. Aleiodes (Chelonorhogas) krulikowskii; Belokobylskij, 1996: 2000: 31; Chen & He, 1997: 40; He et al., 2000: 665. Aleiodes krulikowskii; Fortier & Shaw, 1999: 230; Papp, 2004: 153. Rhogas csikii Szépligeti, 1901 150. Rogas csikii; Shenefelt, 1975: 1223. Aleiodes csikii; Papp, 1991: 84 (as synonym of A. jaroslawensis); 2004: 216 (as synonym of A. krulikowskii). Type material. Lectotype of A. krulikowskii, ♀ (ZISP), “[Russia:] Kirovsk ts., Malmyzh, L.K. Krulikovsk, N. 1906”, “Rh. Krulikowskii Kokw., No. 1906”, “Lectotypus Rogas krulikowskii Kok., design. [V.I.] Tobias, 1980”. Holotype of A. csikii, ♀ (MTMA), “[Russia:] Siberia, Minusinsk, 30.vii.[18]98, Exp. Zichy, leg. Csiki”, “Holotypus ♀ Rhogas csikii sp. n. Szépl., 1901/ des. Papp J, 1967”, “Hym. Typ. No. 403, Museum Budsapest”, “Aleiodes ♀ krulikowskii Kok., det. Papp J., 1983”. Additional material. 1 ♀ (MTMA), “[Romania: N Siebenburgen,] Radnai havas, Páváy V.F/ 26.vii.1906, 1400”; 1 ♂ (MTMA), “[Hungary:] Jaruer, 20.vi.”, “Rhogas carbonarius Gir. var. det. Szépligeti”; 1 ♀ (MTMA), “[Russia:] Ussuri, Kasakewitsch, 1907,

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Figures 453, 454. Aleiodes krulikowskii (Kokujev), ♀, lectotype 453 habitus lateral 454 ovipositor sheath lateral.

Korb”; 1 ♀ (MTMA), “Mongolia: Suchebaator aimak, 44 km SSW von Baruun urt, 1050 m, Exp. Dr. Z. Kaszab, 1965, nr. 349, 2–3.viii.1965”; 1 ♂ (MTMA), “Mongolia: Cojbalsan aimak, Somon Chalchingol, 600 m, Exp. Dr. Z. Kaszab, 1965, nr. 409, 13.viii.1965”. Molecular data. None. Biology. Unknown. The collection dates (June–August) suggest that it is univoltine, but there is nothing to suggest how it overwinters. Diagnosis. Maximum width of hypoclypeal depression 0.7–0.8 × minimum width of face (Fig. 463); OOL of ♀ approx. 1.3 × longer than diameter of posterior ocellus and coarsely rugose (Fig. 464); ventral margin of anterior part of clypeus thin, clypeus approx. 5 × wider than long medially (Fig. 463) and more or less protruding in lateral view (Fig. 465); head robust in anterior view (Fig. 463); lateral mesoscutal lobes

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densely and coarsely punctate, with interspaces narrower than punctures but interspaces becoming wider posteriorly, middle lobe coriaceous, but punctate near narrow and distinctly impressed notauli; mesopleuron densely and coarsely punctate, interspaces approx. equal to diameter of punctures or narrower; vein 1-CU1 of fore wing 0.8 × vein 2-CU1, widened and 1.1 × longer than vein m-cu; hind tarsal claws robust and with inconspicuous fine subbasal brownish pecten (Fig. 466); 1st and 2nd metasomal tergites comparatively slender and 1st tergite moderately widened (Fig. 459); 2nd tergite basally and 3rd tergite apically distinctly convex in lateral view (Fig. 453); 3rd tergite coarsely punctate, with complete lamelliform lateral margin (Fig. 453); hind coxa and femur completely dark brown or blackish; hind tibia usually ivory or pale yellowish basally; first and 2nd metasomal tergites reddish or orange. Description. Lectotype, ♀, length of fore wing 6.9 mm, of body 9.6 mm. Head. Antenna incomplete, 32 segments remaining; frons mainly with curved or oblique rugae; OOL 1.3 × diameter of posterior ocellus, coarsely rugose and rather matt; vertex densely and rather finely rugose, hardly shiny; anterior part of clypeus densely punctate and flat; ventral margin of clypeus thin and rather forward protruding (Fig. 465); clypeus approx. 5 × wider than long medially; width of hypoclypeal depression 0.8 × minimum width of face (Fig. 463); length of eye 1.4 × temple in dorsal view (Fig. 464); vertex behind stemmaticum coarsely rugose; clypeus largely above lower level of eyes; length of malar space 0.2 × length of eye in lateral view; mandible punctate-rugose and with long setae (Fig. 465). Mesosoma. Lateral mesoscutal lobes densely and coarsely punctate, with interspaces narrower than punctures but interspaces becoming wider posteriorly, middle lobe coriaceous, but punctate near narrow and distinctly impressed notauli; precoxal area of mesopleuron and metapleuron coarsely and densely rugose punctate; remainder of mesopleuron densely punctate; metanotum with incomplete median carina; scutellum coarsely punctate; axilla largely densely rugose; propodeum rather flat and coarsely reticulate-rugose, medio-longitudinal carina on only anterior half. Wings. Fore wing: r 0.6 × 3-SR (Fig. 455); 1-CU1 horizontal, 0.8 × 2-CU1 and widened; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 455); cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR widened; surroundings of M+CU1, 1-M and 1-CU1 evenly setose; M+CU1 curved distally. Hind wing: marginal cell linearly widened, its apical width 2.8 × width at level of hamuli (Fig. 456); 2-SC+R quadrate; m-cu absent; M+CU:1-M = 40:21; 1r-m 1.1 × 1-M. Legs. Hind tarsal claws robust and with inconspicuous fine subbasal brownish pecten (Fig. 466); hind coxa largely densely finely punctate, dorso-posteriorly with oblique rugae; hind trochantellus slender; length of hind femur and basitarsus 4.3 and 5.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather convex and moderately widened (Fig. 459), 1.1 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose-punctate; medio-basal area of 2nd tergite triangular and minute (Fig. 459); 2nd suture deep and rather wide; 1st tergite; 2nd tergite basally and 3rd tergite apically distinctly convex in lateral view (Fig. 453); 3rd tergite coarsely punctate,

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Figures 455–466. Aleiodes krulikowskii (Kokujev), ♀, lectotype 455 fore wing 456 hind wing 457 mesosoma lateral 458 mesosoma dorsal 459 metasoma dorsal 460 fore femur lateral 461 hind femur lateral 462 base of antenna 463 head anterior 464 head dorsal 465 head lateral 466 outer middle tarsal claw.

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interspaces approx. equal to diameter of punctures, with complete lamelliform lateral margin; remainder of metasoma smooth and shiny; ovipositor sheath rather slender, with long setae and apically truncate (Fig. 454). Colour. Black; orbit near ocelli reddish brown; fore and middle legs (except blackish or dark brown coxae, trochanters and trochantelli), apex of hind trochantellus and basal third of hind tibia brownish yellow; tarsi darkened and remainder of legs dark brown; palpi (except basally) pale yellowish; mandible yellowish but basally largely dark brown; propleuron and tegula anteriorly dark brown and tegula posteriorly brown; 1st and 2nd metasomal tergites and metasoma ventrally (except apically) orange brown; pterostigma dark brown; veins brown; wing membrane subhyaline. Variation. Orbit near hind ocellus sometimes only very slightly lighter in colour. Antennal segments: ♂ 68(1); according to the original descriptions of A. krulikowskii and A. csikii, the female types have 60 and 62 antennal segments, respectively. Apical tergites of ♂ type 2 and no fringe observed. Distribution. *Hungary, Mongolia, *Romania, Russia (Central and Far East). Aleiodes miniatus (Herrich-Schäffer, 1838) Figs 467–481 Rogas miniatus Herrich-Schäffer, 1838: 156; Shenefelt, 1975: 1238–1239 (type series lost). Rogas (Rogas) miniatus; Tobias, 1976: 81, 1986: 75–76 (transl.: 122). Aleiodes (Neorhogas) miniatus; Papp, 1987b: 36, 1991a: 88. Aleiodes (Chelonorhogas) miniatus; Belokobylskij et al., 2003: 398. Aleiodes miniatus; Bergamasco et al., 1995: 5; Papp, 2005: 177. Rogas bicoloratus Boheman, 1853: 180; Shenefelt, 1975: 1239 (as synonym of A. miniatus); Papp, 2005: 177 (id.). Aleiodes formosus Giraud, 1857: 177; Shenefelt, 1975: 1239; (as synonym of A. miniatus); Papp, 1985a: 159 (lectotype designation and as synonym of A. miniatus), 2005: 177 [examined]. Type material. Lectotype of A. formosus, ♀ (MNHN), “[Austria, Wien,] Prata 16 juin”, “Austria, Vienne, Prater, 16 juin/Papp 1979”, “Lectotypus Aleiodes formosus Gir., 1857, ♀, Papp, 1979”, “Rogas miniatus HS ♀, det. Papp J., 1979”. Additional material. Austria, Czech Republic, France, Finland, Germany, Hungary, Romania, Russia, Sweden, Ukraine, [Kazakhstan, Kyrgyzstan]. Specimens in BMNH, BZL, SDEI, MNHN, MTMA, NMS, OUM, RMNH, ZSSM. The OUM specimen is labelled “Litchfield L.A. Carr 23” but there are very evidently numerous non-British specimens in the (now somewhat dispersed) Carr collection labelled Litch­ field, and good reasons for discounting them as British are given by Perkins (1953). Such labelling may have been a means of identifying ownership of specimens at a time of considerable exchange and identification by others, and there is no evidence that

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Figures 467, 468. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 467 habitus lateral 468 ovipositor sheath lateral.

this species has ever been collected in the British Isles. Material examined from central Europe (often labelled “Germany” or “Bohemia”) is mostly much more than 100 years old, when it seems to have been quite readily collected. Three recent specimens (NMS) from different sites in Sweden (Öland: Halltorp, 2015, 2017 and Skåne: Ravlunda, 2018, all N. Johansson) were swept from herb-rich sandy or gravelly grasslands overlying calcareous bedrock, with outstanding biodiversity partly maintained by grazing (Niklas Johansson, pers. comm.). The evident decline of A. miniatus in central Europe, as evidenced by specimen data showing a declining number of specimens collected in that region through time, probably reflects the loss of similar steppe habitat and, although a fairly recent (1994) specimen from Romania is in MTMA, it may now be extinct in large parts of central Europe. Molecular data. MRS950 (Sweden), MRS951 (Sweden). Biology. Unknown, but it seems to inhabit herb-rich calcareous steppe grasslands. Collected in (May)June–August; presumably univoltine, but we have not examined reared material of this large and distinctive species and there is no indication of how it may overwinter. A series in BZL (one now in NMS) is labelled “Wien D. Au” which can be interpreted as [? wet] woodland near the Danube (M. Schwarz, pers. comm.), which would probably be well under 200 m a.s.l. In contrast, a recent specimen (also in BZL) from Kyrgyzstan was collected higher at 2550 m. Diagnosis. Maximum width of hypoclypeal depression approx. 0.5 × minimum width of face (Fig. 469); OOL of ♀ approx. twice as long as diameter of posterior ocellus and punctate (Fig. 477); ventral margin of clypeus thin and distinctly protruding

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in lateral view; length of malar space approx. equal to height of eye in lateral view (Fig. 478); mesoscutal lobes densely punctate; area of precoxal sulcus wide and coarsely rugose; length of vein 1-CU1 of fore wing 0.4 × vein 2-CU1; 2nd submarginal cell of fore wing short and square (Fig. 469); vein 1r-m of hind wing longer than vein 1-M; vein 2-SC+R of hind wing subquadrate; 3rd tergite densely punctate (Fig. 473); head and mesoscutum orange or brownish yellow; basal half of hind tibia (largely) pale yellowish; metasoma (except part of 1st tergite) orange or brownish yellow. Description. Redescribed ♀ (RMNH) from Russia (Yaaseni). Length of fore wing 6.5 mm, of body 7.9 mm. Head. Antennal segments of ♀ 65, length of antenna 1.1 × fore wing, its subapical segments somewhat longer than wide; frons with coarse curved rugae; OOL 2.3 × diameter of posterior ocellus, and punctate; vertex densely punctate and shiny; clypeus densely punctate; ventral margin of clypeus thin and distinctly protruding forwards (Fig. 478); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 476); length of eye as long as temple in dorsal view (Fig.477); vertex behind stemmaticum densely punctate; clypeus just below lower level of eyes; malar space 0.5 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, with minute interspaces and rather shiny; precoxal area of mesopleuron wide and coarsely rugose medially, mesopleuron above it coarsely and densely punctate, even speculum with some punctures; scutellum convex and punctate; propodeum evenly convex and coarsely reticulate-rugose, mediolongitudinal carina incomplete. Wings. Fore wing: r 0.7 × 3-SR (Fig. 469); 1-CU1 horizontal, 0.4 × 2-CU1; r-m 1.2 × 3-SR; 2nd submarginal cell short (Fig. 469); cu-a inclivous, straight and rather short; 1-M rather curved posteriorly; 1-SR slender and short; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell evenly widened, its apical width 2.7 × width at level of hamuli (Fig. 470); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 35:16; 1r-m 1.5 × 1-M. Legs. Tarsal claws with only three conspicuous brownish and widened bristles basally (Fig. 481); hind coxa densely and rather finely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 4.0 and 4.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite evenly convex, 0.9 × as long as wide apically; 1st tergite coarsely reticulate-rugose, 2nd tergite coarsely and densely rugose-punctate, without median carina; medio-basal area of 2nd tergite short triangular (Fig. 473); 2nd suture deep and finely crenulate; basal half of 3rd tergite densely punctate, remainder of metasoma superficially micro-sculptured; apical half of 3rd tergite with sharp lateral crease; ovipositor sheath moderately wide, with long setae and apically rounded (Fig. 468). Colour. Brownish yellow; antenna, mesosternum, mesopleuron (except anterodorsally), metapleuron, propodeum, 1st tergite, and ovipositor sheath black; propleuron, small patch on middle mesoscutal lobe anteriorly, apices of femora, fore and middle tibiae, tarsi, apical half of hind tibia, veins, and pterostigma dark brown; wing membrane subhyaline; basal half of hind tibia pale yellowish.

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Figures 469–481. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 469 fore wing 470 hind wing 471 mesosoma lateral 472 mesosoma dorsal 473 metasoma dorsal 474 fore femur lateral 475 hind femur lateral 476 head anterior 477 head dorsal 478 head lateral 479 base of antenna 480 apex of antenna 481 inner hind tarsal claw.

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Variation. Second submarginal cell square or somewhat narrower; propleuron dark brown or yellowish; mesopleuron black or yellowish anteriorly and dorsally; medio-longitudinal carina of posterior half of propodeum absent, obsolescent or incomplete. Antennal segments: ♀ 64(5), 65(3), 66(2), 67(3), 68(2), 70(1); ♂ 61(1), 64(2), 66(1), 67(2), 68(1), 69(1), 70(1). On this limited evidence there seems to be little, if any, difference in the number of antennal segments between the sexes. Males are very similar but have the metasoma infuscated apically and the apical tergites are type 3, setae short and dense, glabrous stripe rather narrow and fringe not observed. Distribution. Austria, Czech Republic, Finland, *France, Germany, Hungary, Kazakhstan, *Kyrgyzstan, *Romania, Russia, Sweden, *Ukraine. Aleiodes morio (Reinhard, 1863) Figs 482–503 Rogas morio Reinhard, 1863: 255; Shenefelt, 1975: 1239. Rogas (Rogas) morio; Tobias, 1976: 83, 1986: 76 (transl.: 124). Aleiodes (Neorhogas) morio; Papp, 1985a: 160 (♀ type lost, designation as lectotype), 1991a: 92. Aleiodes (Chelonorhogas) morio; Belokobylskij et al., 2003: 398. Aleiodes morio; Papp, 2005: 177. Type material. Lectotype ♀ from southern Germany most probably lost (Papp 1985a); the only paralectotype (♂, ZMB) is a melanistic male of A. coxalis (Spinola, 1808) which does not fit the original description. The paralectotype was listed as A. tristis Wesmael by Papp (1985a). Additional material. 1 ♀ (MTMA), “[Hungary:] Budapest, Rákospalita, 4.iv.%”, “Rhogas morio Reinh. ♀, det. Szépligeti”, “Aleiodes morio Reinh., ♀, det. Papp J., 1984”; 1 ♂ (MTMA), “[Hungary:] Nadap, 19–21.iv.1951, Móczár Bajári”; 1 ♂ (ZSSM), “[? Germany:] Rogas n. sp.?, /: Ruthe, Berlin[?]/”; 1 ♀ (FMNH), “Suomi [= Finland:] U: Nurmijärvi, 6712:373, 13.v.1984, M. Koponen”; 2 ♀ + 1 ♂ (FMNH, RMNH), id., but 15.v.1984. Molecular data. None. Biology. Unknown. Specimens of both sexes have been collected in April and May, from which from which it is safe to surmise that the winter is passed in the mummy as the male would not hibernate. But we have seen no reared material nor any indication of habitat for this central European species. Its early flight time might be one reason why it is seldom collected and apparently rare. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face; OOL of ♀ 0.8 × (of ♂ 0.9 ×) diameter of posterior ocellus and rugose; ventral margin of anterior part of clypeus comparatively sharp and more or less protruding in lateral view (Fig. 493); head rather transverse (Fig. 492); mesoscutal lobes coriaceous;

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Figures 482, 483. Aleiodes morio (Reinhard), ♀, Hungary, Budapest 482 habitus lateral 483 ovipositor sheath lateral.

precoxal area of mesopleuron rugose medially; vein 1-CU1 0.3–0.4 × vein 2-CU1 and 0.3 × vein m-cu; hind tarsal claws with conspicuous and robust brownish pecten (Figs 494, 503); posterior orbit black; pterostigma of ♀ pale brown medially, of ♂ dark brown; coxae and femora completely black or dark brown; hind tibia usually ivory or pale yellowish basally; 1st and 2nd metasomal tergites of both sexes black. According to Papp (1985) most closely related to A. sibiricus (Kokujev), but that species does not have all black females and has the shape of the clypeus different. According to the original description the pterostigma is yellowish and laterally darkened, 1st subdiscal cell of the fore wing rather short, because vein cu-a distinctly more postfurcal than its own length (ca twice its own length) and meaning vein 1-CU1 of intermediate [approx. 0.6 ×] length of 2-CU1 and 0.8 × vein m-cu] [= “discoidali posterior brevior” as indicated for A. pallidicornis], precoxal sulcus area rugose medially, 1st tergite twice wider posteriorly than basally, hind leg black, except pale yellowish dorso-basal area of hind tibia and palpi dark brown. Here we accept the interpretation of the first reviser (Szépligeti 1906) despite the difference in the shape of the 1st subdiscal cell, because it may be part of intraspecific variation.

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Figures 484–495. Aleiodes morio (Reinhard), ♀, Hungary, Budapest, but 490 of ♂ from Nadap 484 fore wing 485 hind wing 486 mesosoma lateral 487 mesosoma dorsal 488 1st–3rd metasomal tergites dorsal 489 fore femur lateral 490 hind femur lateral 491 head anterior 492 head dorsal 493 head lateral 494 outer middle tarsal claw 495 base of antenna.

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Figure 496. Aleiodes morio (Reinhard), ♂, Hungary, Nadap, habitus lateral.

Description. Redescribed ♀ (MTMA) from Hungary (Budapest). Length of fore wing 7.5 mm, of body 9.4 mm. Head. Antenna incomplete, 47 segments remaining (54 in lectotype), length of complete antenna approx. 0.9 × fore wing, its subbasal and subapical segments short; frons largely rugose; OOL 0.8 × diameter of posterior ocellus, and mainly rugulose and with satin sheen; depression near posterior ocellus smooth; vertex densely rugulose and with satin sheen; clypeus with some punctures; ventral margin of clypeus rather thin and protruding forwards (Fig. 493); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 491); length of eye 1.7 × temple in dorsal view (Fig. 492); vertex behind stemmaticum punctate-rugulose; clypeus largely above lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, with satin sheen; precoxal area of mesopleuron widely and densely rugose, but densely punctate posteriorly; middle of mesopleuron densely rugulose and dorsally coarsely rugose; metapleuron largely rugose; scutellum punctate-coriaceous; propodeum rather flat and densely rugose or rugulose, medio-longitudinal carina complete, and without protruding carinae laterally. Wings. Fore wing: r 0.5 × 3-SR (Fig. 484); 1-CU1 slightly oblique, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell robust and posteriorly somewhat diverging

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Figures 497–503. Aleiodes morio (Reinhard), ♂, Hungary, Nadap 497 wings 498 apex of metasoma lateral 499 apex of metasoma dorsal 500 base of antenna 501 head anterior 502 head dorsal 503 inner hind tarsal claw.

(Fig. 484); cu-a inclivous, straight; 1-M slightly curved posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 densely but inconspicuously setose. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 485); 2-SC+R short and subquadrate; m-cu present anteriorly; M+CU:1-M = 4:3; 1r-m 0.6 × 1-M. Legs. Tarsal claws bristly setose, medium-sized, and with robust pecten basally (cf. Fig. 494); hind leg missing in redescribed specimen. Metasoma. First tergite evenly convex, 0.9 × as long as wide apically; 1st tergite with medio-longitudinal carina; 1st and 2nd tergites and basal half of 3rd tergite finely and densely longitudinally rugulose; medio-basal area of 2nd tergite short triangular (Fig. 488); 2nd suture distinct and finely crenulate; remainder of metasoma superficially micro-sculptured; 4th tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Fig. 483).

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Colour. Black; palpi brownish yellow, but basally dark brown; tegulae pale yellowish; legs (except pale base of tibiae), metasoma ventrally and veins dark brown; pterostigma brown and medially yellowish brown; wing membrane subhyaline. Variation. Clypeus distinctly to moderately protruding and ventrally rather thin to thick. Antennal segments of ♀ 51(1), 52(1), 54(1), of ♂ 55(1). Males are very similar to the redescribed female (including the wing venation: Fig. 497), apical tergites type 1 with fringe not observed and probably absent (Fig. 498), mesopleuron rugulose or punctate medially and pterostigma entirely dark brown. Distribution. *Finland, Germany, Hungary. Notes. The lost lectotype from Germany had hyaline wings (which separates it from the A. carbonarius/ grassator/ carbonaroides complex), the pterostigma paler medially than laterally (entirely dark brown), base of the hind tibia pale yellow (black in ♂) and the body of ♀ entirely black (more or less yellowish or reddish). Aleiodes nigrifemur van Achterberg & Shaw, sp. nov. http://zoobank.org/2535423C-36E7-4ECC-9FC3-362039FC4928 Figs 504–518 Type material. Holotype, ♀, (RMNH), “Greece, Peloponn[esus], Chelmos, 1700 m, 29.v.1987, H. Teunissen”. Molecular data. None. Biology. Unknown; the only known specimen was collected at the end of May which gives no clue of voltinism or how the winter is passed. Diagnosis. Maximum width of hypoclypeal depression 0.6 × minimum width of face (Fig. 513); OOL of ♀ 1.6 × longer than diameter of posterior ocellus and rugulose (Fig. 514); width of clypeus intermediate apically, but strongly protruding in lateral view (Fig. 515); lobes of mesoscutum densely punctate, coriaceous between punctures; precoxal area widely rugose, and posteriorly punctate; vein 1-CU1 of fore wing 0.2 × vein 2-CU1 (Fig. 506); hind tarsal claws rather robust and with few yellowish pectinal teeth (Fig. 517); 3rd antennal segment of ♀ and basal third of hind femur black; basal third of hind tibia pale yellowish, contrasting with black basal half of hind femur. Similar to A. morio (Reinhard), but has pterostigma black (pale brown in A. morio), fore wing darkened apically (subhyaline), vein 1-M of hind wing linear with M+CU (angled); metasoma largely yellowish brown (entirely blackish) and eye in lateral view comparatively small (eye larger). Description. Holotype, ♀, length of fore wing 7.2 mm, of body 8.2 mm. Head. Antennal segments of ♀ 62, length of antenna 1.1 × fore wing, its subapical segments medium-sized (Fig. 516); frons largely rugose; OOL 1.6 × diameter of posterior ocellus, rugulose and shiny; depression near posterior ocellus rugose; vertex largely rugose, rather shiny; clypeus rugulose; ventral margin of clypeus intermediate and distinctly protruding forwards (Fig. 515; as face dorsally); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 513); length of eye 1.3 × temple in dor-

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Figures 504, 505. Aleiodes nigrifemur sp. nov. ♀, holotype 504 habitus lateral 505 ovipositor sheath lateral.

sal view (Fig. 514); vertex behind stemmaticum rugulose; clypeus largely above lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, rather shiny and interspaces coriaceous; precoxal area of mesopleuron widely rugose but posteriorly punctate, and area above it densely punctate or rugulose; metapleuron densely punctate dorsally and rugose ventrally; metanotum with short median carina anteriorly; scutellum remotely punctate, with some lateral rugae; propodeum rather short and flat, coarsely reticulaterugose, medio-longitudinal carina complete, and without protruding carinae laterally. Wings. Fore wing: r 0.5 × 3-SR (Fig. 506); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 506); cu-a inclivous, straight; 1-M curved posteriorly; 1-SR wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 1.9 × width at level of hamuli (Fig. 506); 2-SC+R short and longitudinal; m-cu present anteriorly; vein 2-1A comparatively long (Fig. 506); M+CU:1-M = 24:37; 1r-m 0.65 × 1-M. Legs. Tarsal claws rather robust, bristly setose and few small yellowish teeth (Fig. 517); hind coxa largely punctate and with some oblique striae dorsally; hind trochantellus rather robust; length of hind femur and basitarsus 4.5 and 5.1 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite rather flat medially, 0.8 × as long as wide apically; 1st tergite and anterior half of 2nd tergite with medio-longitudinal carina; 1st–2nd tergites densely longitudinally rugose; 3rd tergite (except posterior third) mainly rugulose; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 509); 2nd suture rather deep and crenulate; remainder of metasoma superficially micro-sculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 505). Colour. Black; maxillary palp apically, basal 0.4 of hind tibia and tegulae pale yellowish; mandible (but with dark brown patch), side of pronotum dorso-posteriorly, fore and middle tibiae, hind basitarsus basally, 1st tergite apically, 2nd–5th tergites or-

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Figures 506–518. Aleiodes nigrifemur sp. nov. ♀, holotype 506 wings 507 mesosoma lateral 508 mesosoma dorsal 509 1st–3rd metasomal tergites dorsal 510 fore femur lateral 511 hind femur lateral 512 base of antenna 513 head anterior 514 head dorsal 515 head lateral 516 apex of antenna 517 outer hind tarsal claw 518 antennae.

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ange brown; remainder of legs dark brown; remainder of palp, veins and pterostigma dark brown; lateral lobes of mesoscutum (except anteriorly and medially) dark reddish brown; wing membrane subhyaline, but apically infuscated (Fig. 506). Distribution. Greece (main). Etymology. The species is named after its black femur; niger is Latin for black, dark, dusky. Aleiodes nobilis (Haliday [in Curtis], 1834) Figs 519–542 Rogas nobilis Haliday [in Curtis], 1834: 512; Papp, 2005: 176 (as syn. of A. ductor). Rogas ductor var. nobilis; Shenefelt, 1975: 1227. Aleiodes (Neorhogas) nobilis; Papp, 1991a: 70 (as synonym of A. ductor). Aleiodes (Chelonorhogas) nobilis; van Achterberg, 1997: 62 (both syntypes lost); Belokobylskij et al., 2003: 398. Aleiodes nobilis; Bergamasco et al., 1995: 5; O’Connor et al., 1999: 91–92; Papp, 2005: 177. Rogas medianus Thomson, 1892: 1668; Shenefelt, 1975: 1237; van Achterberg, 1997: 62 (as synonym of A. nobilis); Belokobylskij et al., 2003: 398 (id.); Papp, 2005: 177 (id.) [examined]. Rogas (Rogas) medianus; Tobias, 1976: 85, 1986: 80 (transl.: 133). Aleiodes (Neorhogas) medianus; Papp, 1991a: 69; Belokobylskij, 1996: 13. Aleiodes medianus; Papp & Vas, 2016: 152. Aleiodes ductor; auct. p.p. Type material. Neotype of A. nobilis here designated: ♀ (NMS), “[Scotland:] W. Ross, Coppachy, Letterewe Estate, ix.2007, Mal. trap, P. Tinsley-Marshall”, “BCLDQR _00123”. Lectotype of A. medianus, ♀ (ZIL), “[Sweden:] Scan”, “medianus m.”, “Funnen vid Esperöd I Skåne, teste Papp J., 1983”, “Lectotypus Rogas medianus Thoms., 1891, ♀, Papp, 1983”, “Aleiodes medianus Th., ♀, det. Papp J., 1983”. The lectotype designation for A. nobilis is necessary for nomenclatural stability, because the type series is lost (van Achterberg, 1997) and the species has been confused with similar species in the past. The specimen from Scotland is selected neotype because it fits well the original description, Scotland is relatively close to both type localities (near Holywood in Ireland and Monk’s Wood in England) and it is in good condition. Additional material. Austria, British Isles (Scotland: V.C.s 72, 88, 105; Ireland: V.C. H29), Bulgaria, Croatia, Czech Republic, Finland, Germany, Hungary, Italy, Moldova, Netherlands (LI: Gulpen; St. Pietersberg; Geulle (Bunderbos); NB: Udenhout (“de Brand”), OV: Voorst (Twello), ZH: Lexmond), Poland, Romania, Russia, Serbia, Slovakia, Sweden. Specimens in ALC, BMNH, BZL, HSC, MTMA, NMS, NRS, RMNH, SDEI, Tullie House Museum Carlisle, USNM, ZSSM.

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Figures 519–521. Aleiodes nobilis (Haliday), ♀, neotype 519 habitus lateral 520 ovipositor sheath lateral 521 mummy of Autographa gamma Linnaeus (Germany, Lindenhayn).

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Figures 522–535. Aleiodes nobilis (Haliday), ♀, neotype 522 wings 523 mesosoma lateral 524 mesosoma dorsal 525 1st–4th metasomal tergites dorsal 526 fore femur lateral 527 hind femur lateral 528 hind tibia lateral 529 head anterior 530 head dorsal 531 head lateral 532 base of antenna 533 apex of antenna 534 outer hind tarsal claw 535 outer fore tarsal claw.

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Molecular data. MRS401 (Finland), MRS880 (Russia), MRS881 (UK). Biology. Collected predominantly in grassy places, June–October. Reared from the noctuid Autographa gamma (Linnaeus) (4 [1 NRS, 2 HSC], Germany, Sweden; H. Schnee) but, in view of its moderately northern areas of occurrence, it seems very likely that other plusiine noctuids would play an important part in its host range. The rearing data indicate that it is plurivoltine, and adult emergence in November from mummies forming in October suggests that it overwinters in the host larva. Diagnosis. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 529); OOL of ♀ approx. as long as diameter of posterior ocellus and granulate (Fig. 530); ventral margin of clypeus thick and not protruding in lateral view (Fig. 531); mesoscutal lobes (as vertex) very finely and densely granulate, with satin sheen; precoxal area smooth; vein 1-CU1 0.7–1.3 × vein 2-CU1 and vein 1-CU1 wider than 2-CU1 (Fig. 522); tarsal claws with distinct dark brown pecten (Figs 534, 535); hind femur and basitarsus slender (Figs 519, 527); 1st metasomal tergite comparatively slender (Fig. 525); at least basal half of 4th–6th tergites of ♂ usually with long dense setosity (Figs 537, 538); head black; pronotum usually (partly) orange brown; both tegula and humeral plate equally yellowish; base of hind tibia pale yellowish; hind basitarsus brownish yellow, strongly contrasting with dark brown telotarsus; 2nd tergite yellowish or reddish. Description. Redescribed ♀ (RMNH) from Slovakia (Kubrica). Length of fore wing 5.1 mm, of body 5.9 mm. Head. Antennal segments of ♀ 48, length of antenna 1.25 × fore wing, its subapical segments slender (Fig. 533); frons matt and granulate; OOL equal to diameter of posterior ocellus, and coriaceous-granulate; vertex coriaceous-granulate and rather dull; clypeus punctate-coriaceous; ventral margin of clypeus thick and not protruding forwards (Fig. 531); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 529); length of eye 2.5 × temple in dorsal view (Fig. 530); vertex behind stemmaticum granulate; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely granulate, rather shiny near tegulae; precoxal area of mesopleuron smooth, surroundings sparsely punctulate; metapleuron mostly granulate; metanotum without median carina; scutellum granulate and with lateral carina; propodeum slightly convex, granulate with spaced rugosity, mediolongitudinal carina only anteriorly present, and no protruding carinae laterally. Wings. Fore wing: r 0.6 × 3-SR (Fig. 522); 1-CU1 straight, 1.2 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell short (Fig. 522); cu-a vertical, nearly straight; 1-M slightly curved posteriorly; 1-SR rather narrow; surroundings of M+CU1, 1-M and 1-CU1 evenly setose. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 522); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 14:13; 1r-m 0.6 × 1-M. Legs. Tarsal claws with conspicuous and robust dark brown pecten (Figs 534, 535); hind coxa sparsely finely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.7 and 8.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus.

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Figure 536. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen, habitus lateral.

Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites rather regularly sublongitudinally striate, without medio-longitudinal carina on 2nd tergite; medio-basal area of 2nd tergite wide triangular and rather distinct (Fig. 525); 2nd suture rather deep and narrow; basal quarter of 3rd tergite finely striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath rather long and slender, with long setae and apically rounded (Fig. 520). Colour. Black; pterostigma (except yellowish extreme base and apex), veins (except brown vein C+SC+R), clypeus, apical third of hind tibia and telotarsus dark brown; pal-

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Figures 537–542. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen 537 apex of metasoma dorsal 538 apex of metasoma lateral 539 apex of antenna 540 head anterior 541 head dorsal 542 base of antenna.

pi, tegulae, remainder of tibiae and tarsi, pale yellowish; apex of middle femur and apical half hind femur, black; remainder of legs, antenna (but apical segments and to some degree scapus infuscate) yellowish brown; 1st–3rd metasomal tergites (except black medial patch of 1st tergite), propleuron and pronotum orange; wing membrane subhyaline. Variation. 1-CU1 0.7–1.2 × 2-CU1; striae of 2nd tergite regularly sublongitudinal or somewhat diverging posteriorly (Fig. 525), but in male sometimes only granulate; basal third or half of 3rd tergite finely striate, rarely completely smooth; fore and middle femora black or dark brown apically or brownish yellow; pronotal side orange to dark brown dorsally; dark patch of 1st tergite absent (e.g., lectotype of A. medianus), small, large or occupying most of tergite; posterior half of 3rd tergite orange or black. Antennal segments: ♀ 46(3), 47(8), 48(3), 49(5), 50(5); ♂ 45(2), 46(2), 47(6), 48(5), 49(6); with little difference in the number of antennal segments between the sexes. Males are very similar, but apical tergites type 4, dense setae (making the tergites appear concave) and fringe strong (Figs 537, 538). Distribution. *Austria, British Isles (Scotland, Ireland), Bulgaria, *Croatia, Czech Republic, Finland, Germany, Hungary, Italy, Moldova, Netherlands, *Poland, *Romania, Russia, *Serbia, *Slovakia, Sweden.

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Aleiodes pallidicornis (Herrich-Schäffer, 1838) Figs 543–557 Rogas pallidicornis Herrich-Schäffer, 1838: 156; Shenefelt, 1975: 1241; Zaykov, 1980b: 87. Rhogas pallidicornis; Fahringer, 1932: 266. Rogas (Rogas) pallidicornis; Tobias, 1976: 84, 1986: 80 (transl.: 130). Aleiodes (Neorhogas) pallidicornis; Papp, 1987b: 36, 1991a: 70 (as senior synonym of A. hirtus). Aleiodes (Chelonorhogas) pallidicornis; Belokobylskij, 2000: 42; Ku et al., 2001: 236; Belokobylskij et al., 2003: 398. Aleiodes pallidicornis; Papp, 2005: 177. Rhogas pallidipennis Dalla Torre, 1898: 221 [invalid emendation]. Rogas ductor auctt. p.p. [North & Central Europe, e.g., Lozan et al. 2010: 17]. Type material. Neotype of A. pallidicornis here designated, ♀ (RMNH), “[Netherlands], [Zuid-]Holland, Asperen, 6.viii.1972, C.J. Zwakhals”. The neotype designation is necessary for nomenclatorial stability, because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; no specimens could be found by the first author in ZMB), and the species has been confused with similar species in the past. The specimen is selected because it fits well the original description, Netherlands is relatively close to the probable type location in Germany and it is in excellent condition. Additional material. Austria, Belarus, British Isles (Scotland: V.C. ?92), Bulgaria, Germany, Hungary, Italy, Montenegro, Netherlands (ZH: Asperen; Schoonrewoerd; Waarder), Romania, Russia, Slovakia, Switzerland, Turkey [Iran, North Korea]. Specimens in BMNH, BZL, MRC, MSC, MTMA, NMS, RMNH, UNS, ZSSM. Molecular data. MRS885 (Russia). Biology. Very little is known. Specimens collected in (May) June–August (September), the great majority in June-July strongly suggesting that it is at least largely univoltine. The Dutch specimens were collected in fairly humid coppice woods. The single British specimen (BMNH; G.T. Lyle) was reared (emergence 20.vi.1926) from a “noctua” caterpillar collected by E.A. Cockayne in Aberdeenshire. The mummy is lost. At that time, the term “noctua” was used generally for Noctuidae rather than in the restricted sense of the genus of that name, and it would appear (as Cockayne was by then a distinguished amateur lepidopterist) that the host larva did not belong to an obviously identifiable species. Otherwise we have not seen reared material, and there is no indication of how the winter is passed. Diagnosis. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 552); OOL of ♀ approx. as long as diameter of posterior ocellus and remotely punctate with interspaces superficially granulate (Fig. 553); ventral margin of cly­ peus thick and not protruding in lateral view (Fig. 554); mesoscutal lobes and vertex very finely and densely granulate, with satin sheen; precoxal area smooth medially, but sometimes some rugae below it; vein 1-CU1 0.4–0.6 × vein 2-CU1 and equally slender (Fig. 545); tarsal claws with distinct dark brown pecten (Fig. 557); hind femur and basitarsus

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Figures 543, 544. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 543 habitus lateral 544 ovipositor sheath lateral.

slender (Fig. 543); basal quarter of 3rd tergite largely finely striate; at least basal half of 4th– 6th tergites of ♂ usually with long dense setosity; head and pronotum black; both tegula and humeral plate equally yellowish; base of hind tibia with narrow dark brown band; hind femur and tibia at least partly black or dark brown; 2nd tergite yellowish or reddish. Description. Neotype, ♀, length of fore wing 5.9 mm, of body 6.6 mm. Head. Antennal segments of ♀ 54, length of antenna 1.3 × fore wing, its subapical segments rather robust (Fig. 556); frons largely superficially granulate, anteriorly with some weak striae; OOL 1.4 × diameter of posterior ocellus, and punctate, interspaces granulate; vertex spaced punctate, shiny; clypeus densely and coarsely punctate, with granulate interspaces; ventral margin of clypeus thick and not protruding forwards (Fig. 554); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 552); length of eye 2.1 × temple in dorsal view (Fig. 553); vertex behind stemmaticum granulate with some transverse rugae; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate with largely granulate interspaces, with satin sheen; precoxal area of mesopleuron distinctly remotely punctate, interspaces larger than punctures; metapleuron densely punctate-granulate; metanotum with median carina; scutellum punctate-granulate; propodeum evenly convex and coarsely rugose, its medio-longitudinal carina complete.

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Figures 545–557. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 545 fore wing 546 hind wing 547 mesosoma lateral 548 mesosoma dorsal 549 1st–3rd metasomal tergites dorsal 550 fore femur lateral 551 hind femur lateral 552 head anterior 553 head dorsal 554 head lateral 555 base of antenna 556 apex of antenna 557 inner hind tarsal claw.

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Wings. Fore wing: r 0.4 × 3-SR (Fig. 545); 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 545); cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 evenly setose. Hind wing: marginal cell gradually widened, its apical width 2.6 × width at level of hamuli (Fig. 546); 2-SC+R short and longitudinal; m-cu present basally; M+CU:1-M = 15:14; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather small dark brownish pecten, absent near apical tooth (Fig. 557); hind coxa largely densely and finely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 5.0 and 7.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely irregularly rugose, but posteriorly 2nd tergite largely smooth and no median carina; medio-basal area of 2nd tergite triangular and rather large (Fig. 549); 2nd suture rather deep and finely crenulate; basal half of 3rd tergite smooth (except for punctuation) and shiny as remainder of metasoma; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath moderately wide, with long setae and apically truncate (Fig. 544). Colour. Black; hind tarsus largely infuscate, but 3rd and 4th segments paler than other segments; apices of fore and middle tibiae slightly infuscate, base of middle and hind tibiae and telotarsi dark brown; apical two-fifths of hind femur and hind tibia (except a pale yellowish band subbasally) black; remainder of legs, 1st and 2nd tergites, and 3rd tergite antero-laterally orange brown; palpi and tegulae brownish yellow; most of veins and pterostigma dark brown; wing membrane subhyaline. Variation. Antennal segments: ♀ 49(2), 50(2), 51(2), 52(6), 53(6), 54(1), 56(1); ♂ 50(1), 51(3), 52(1), 53(2), 54(2), 55(1), 56(3). On average males have ca two more antennal segments than females. Males are similar but have a large dark brown patch on 1st tergite, hind tarsus largely dark brown and apical tergites type 3, positioned rather posteriorly, setae long and fringe not observed. Distribution. Austria, *Belarus, *British Isles (Scotland), Bulgaria, Hungary, *Iran, *Italy, *Montenegro, *Netherlands, North Korea, *Romania, Russia, *Slovakia, Switzerland, *Turkey. Notes. The type of Rogas pallidicornis Herrich-Schäffer, 1838, has been lost. Traditionally, it has been considered to belong to Aleiodes ductor (Thunberg, 1822), but the latter species is a synonym (see under A. unipunctator). The inadequate original description indicates that the 2nd tergite has diverging rugae, which excludes part of A. ductor auctt. Female specimens with yellowish or brownish palpi, basal half of the antenna yellowish and blackish hind tibia (except its pale yellowish base) fit well the original description of A. pallidicornis. Aleiodes pallidistigmus (Telenga, 1941) Figs 558–572 Rhogas (Rhogas) pallidistigmus Telenga, 1941: xii, 143, 177 (but also as palidistigma (p. 409) and pallidistigma (p. 420)) [examined].

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Rogas pallidistigmus; Shenefelt, 1975: 1241. Aleiodes (Neorhogas) pallidistigmus; Papp, 1985b: 348. Aleiodes (Chelonorhogas) pallidistigmus; Chen & He, 1997: 41; Fortier & Shaw, 1999: 228; Belokobylskij, 2000: 47; Ku et al., 2001: 237. Aleiodes pallidistigma; He et al., 2000: 667. Rogas heterostigma Stelfox, 1953: 149; Shenefelt, 1975: 1233 [examined]. Syn. nov. Aleiodes (Neorhogas) heterostigma; Papp, 1985a: 143, 146–147, 153, 1991a: 95. Aleiodes heterostigma; O’Connor et al., 1999: 91; Papp, 2005: 177. Type material. Paratypes of A. heterostigma, 4 ♀ + 1 ♂ (BMNH, USNM), “[Ireland], Rye Water, Co. KD, 5/9.vii.[19]42, AWS[telfox]”; 1 ♀ (RMNH), id., but 5.vii.1942; 1 ♀ (NMI), id., but 8.vii.1948; 1 ♂ (NMI), “[Ireland], Woodbrook, OC, 26.vi.[19]38”. Holotype of A. pallidistigmus (♀, ZISP) from Far East Russia (Primorsky Krai, Ussuri area, Vinogradovka, 10.viii.1929, Kiritshenko). Additional material. 1 ♀ (RMNH), “Belgium: Liège, Mt. Rigi, 650 m, 2.viii.1986, at light, C. v. Achterberg, RMNH”; 1 ♀ (MTMA), “Dania [= Denmark]: S-Jutland, Kragelund Mose, near Baekke, 11.viii.1973, [T.] Munk”; 1 ♀ (NMS), “Wales: Cereigion, Rhôs Rhydd, SN572738, Molinia bog, 30.vii.1987, NCC Welsh Peatland Survey, P. Holmes, NMSZ 1996.023”; 1 ♂ (NMS) “Wales; Ceredigion, Comin Esgair Maen, SN652649, Equisetum bog, 23.vii.1987, NCC Welsh Peatland Survey, P. Holmes, NMSZ 1996.023”; 1 ♂ (FMNH), “Fennia [= Finland]: Helsinki, 27.vii.1978, O. Ranin”; 1 ♀ (NMS) Far East Russia, Anismovka v. Shkotova/S Primorje reg., 11–13.viii.2003, leg. Osipov”; 2 ♀ (MRC) “Far East Russia, S. Primorje reg., Lazo distr. Valentin 17–18.vii.2003 leg. Osipov”: 1 ♀ (NMS) [Russian, Far East]; 2 ♀ (RMNH), “China: Jilin, Gomngzhuling, 43°5'N, 124°8'E, viii–ix.1983, Wang Chenghun”. Molecular data. None. Biology. Unknown for West Palaearctic populations. Specimens have been collected in (June)July-August in open boggy areas, certainly at least sometimes over limestone. Presumably univoltine, but we have not seen reared material from West Palaearctic and the overwintering mode is unclear. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 568); OOL of ♀ approx. 1.3 × as long as diameter of posterior ocellus and densely granulate (Fig. 569); penultimate segments rather slender and antenna 1.5 × as long as fore wing; ventral margin of clypeus thick, not protruding in lateral view; mesoscutal lobes finely granulate-punctulate and matt; precoxal area coarsely rugose; marginal cell of fore wing of ♀ ending rather close to wing apex (Fig. 560); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 560); hind tarsal claws rather robust and only brownish setose (Fig. 571); 3rd tergite and basal half of 4th tergite coriaceous and dull; labial palp yellowish brown or brown; basal half of hind tibia reddish or yellowish, slightly paler than basal half of hind femur, and its apex reddish or yellowish; 4th and 5th tergites black. Description. Redescribed ♀ paratype of A. heterostigma (RMNH) from Ireland (Rye Water). Length of fore wing 4.9 mm, of body 6.3 mm.

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Figures 558, 559. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 558 habitus lateral 559 ovipositor sheath lateral.

Head. Antennal segments of ♀ 58, length of antenna 1.1 × fore wing, its subapical segments rather robust (Fig. 567); frons largely superficially granulate; OOL 1.8 × diameter of posterior ocellus, and superficially rugulose-granulate and shiny; vertex superficially rugulose-granulate, rather shiny; clypeus with some punctures; ventral margin of clypeus thick and not protruding forwards (Fig. 570); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 568); length of eye 1.3 × temple in dorsal view (Fig. 569); vertex behind stemmaticum superficially granulate-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate-granulate, matt; precoxal area of mesopleuron coarsely rugose medially; remainder of mesopleuron coarsely punctate, with some rugae near speculum and interspaces superficially granulate; scutellum rather flat, punctulate-granulate and with weak lateral carinae; propodeum rather convex, shiny and coarsely rugose, medio-longitudinal carina distinct only on its anterior half. Wings. Fore wing: r 0.3 × 3-SR (Fig. 560); 1-CU1 horizontal, 0.45 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell short (Fig. 560); cu-a inclivous, straight; 1-M slightly

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Figures 560–572. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 560 fore wing 561  hind wing 562 mesosoma lateral 563 mesosoma dorsal 564 1st–3rd metasomal tergites dorsal 565 fore femur lateral 566 hind femur lateral 567 apex of antenna 568 head anterior 569 head dorsal 570 head lateral 571 outer hind tarsal claw 572 base of antenna.

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curved posteriorly; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell gradually widened with basal half rather narrow (Fig. 561), its apical width 2.6 × width at level of hamuli; 2-SC+R short longitudinal; m-cu weakly developed; M+CU:1-M = 35:26; 1r-m 0.5 × 1-M. Legs. Tarsal claws rather robust and only brownish setose (Fig. 572); hind coxa coarsely punctate, dorsally with oblique striae; hind trochantellus robust; length of hind femur and basitarsus 4.1 and 4.6 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites and base of 3rd tergite finely and irregularly longitudinally rugose, with mediolongitudinal carina weak; medio-basal area of 2nd tergite triangular and short (Fig. 564); 2nd suture rather deep and crenulate; apical half of 3rd tergite punctate-granulate, remainder of metasoma smooth except for some superficial micro-sculpture; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 559). Colour. Black; telotarsi largely and basal quarter of antenna dark brown; palpi, tegulae and pterostigma pale yellow; remainder of legs, 1st and 2nd tergites, basal half of 3rd tergite largely and pronotum orange brown; veins brown; wing membrane subhyaline. Variation. Pronotum anteriorly and basal half of antenna orange brown to dark brown, pterostigma is yellowish to largely (except base) rather dark brown; length of malar space 1.0–1.4 × basal width of mandible; OOL 0.7–1.8 × diameter of ocellus and metapleuron medially more or less punctate, rugulose-coriaceous or rugose. Antennal segments: ♀ 54(1), 58(1), 59(4), 60(2), 62(1), 64(1); ♂ 59(1), 60(1), 62(1). Apical tergites of ♂ type 1, fringe absent. Distribution. *Belgium, British Isles (Ireland, Wales), China, *Denmark, *Finland, Russia (Far East). New synonymy. We tried to separate the East Palaearctic A. pallidistigmus from the West Palaearctic A. heterostigma, but efforts were in vain. The differences such as the colour of the basal half of the antenna (dark brown in A. heterostigma and usually yellowish or brown in A. pallidistigmus), the eyes and ocelli often smaller, OOL 1.1– 1.8 × diameter of ocellus (0.7–1.4 ×), malar space 1.2–1.3 × basal width of mandible (1.0–1.4 ×) and metapleuron with a shiny and more or less punctate area (less shiny and rugulose-coriaceous or rugose) are too variable to justify separation of A. heterostigma. Therefore, we synonymise A. heterostigma with A. pallidistigmus (syn. nov.). Aleiodes periscelis (Reinhard, 1863) Figs 573–603 Rogas periscelis Reinhard, 1863: 254; Shenefelt, 1975: 1242; Kotenko, 1992: 96 [examined]. Rhogas (Rhogas) periscelis var. charkowensis Kokujev, 1898: 297. Rhogas (Rhogas) periscelis var. charkoviensis [sic!]; Telenga, 1941: 163, 164.

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Rogas (Rogas) periscelis; Tobias, 1976: 86, 1986: 81 (transl. 133); Papp, 1983: 330. Aleiodes (Neorhogas) periscelis; Papp, 1985a: 161 (lectotype designation), 1985b: 348; 1991a: 79, 1991c: 641, 1994: 307. Aleiodes (Chelonorhogas) periscelis; Belokobylskij, 2000: 36. Aleiodes periscelis; Fortier & Shaw, 1999: 230; Belokobylskij & Taeger, 2001: 115; Belokobylskij et al., 2003: 400 (excluded from German checklist); Papp, 2005: 177. Rhogas jaroslawensis Kokujev, 1898: 302. Syn. nov. Rhogas (Rhogas) jaroslavensis; Telenga, 1941: 176, 408 (invalid emendation). Rogas jaroslawensis; Shenefelt, 1975: 1235. Rogas (Rogas) jaroslavensis; Tobias, 1976: 85. Rogas (Rogas) jaroslawensis; Tobias, 1986: 81 (transl.: 133; as synonym of A. rufipes (Thomson)). Aleiodes (Neorhogas) jaroslawensis; Papp, 1985a: 153, 1991a: 80. Aleiodes jaroslawensis; Papp, 2005: 177 (as valid species). Type material. Lectotype of R. periscelis, ♂ (ZMB), “[Austria:] Neusiedler See”, “Type”, “Coll. H. Rhd.”, “26696”, “periscelis Gir. [= from Giraud]”, “Lectotypus Rogas periscelis Rhd., 1863, ♂, Papp, 1982”, “Aleiodes periscelis Rhd., ♂, det. Papp J., 1983”; 2 ♂ paralectotypes (MNHN), one with “[Austria:] Prater, Mai”, “ex coll. Giraud”. Holotype of R. jaroslawensis, ♀ (ZISP; examined photos made by K. Samartsev), “[S. Russia:] Berditsino [Yaroslavskiy rayon, 57.454N, 40.108E], 22.vi.1892, A.M. Yakovlev, 1909”, “Rh. jaroslawensis Kokw., No. 1909” and with a round golden label. Additional material. Czech Republic, Germany, Hungary, Russia. Specimens in ALC, BZL, MTMA, NMS, SDEI, ZISP, ZMB, ZSSM. Molecular data. None. Biology. Unknown but presumably univoltine. Specimens of both sexes collected in April and May suggest that the winter is passed in the mummy. We have not seen reared material, but several Hungarian specimens appear to have been collected in Quercus-dominated woodland, but without indication of any association with Quercus as such. Diagnosis. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Figs 573, 584); OOL of ♀ 1.5 × as long as diameter of posterior ocellus (Fig. 585; of ♂ 1.4 ×; Fig. 599), rugulose or rugose and with satin sheen; ventral margin of clypeus thick and not protruding anteriorly (Fig. 586); mesoscutum remotely punctulate and with satin sheen, interspaces of lateral lobes largely smooth, of middle lobe superficially coriaceous; area of precoxal sulcus smooth and shiny; length of vein 1-CU1 of fore wing 0.3–0.4 × vein 2-CU1 and 0.5 × vein m-cu; vein 2-SC+R of hind wing subquadrate; hind basitarsus robust; head (including basal half of mandible) black; antenna of ♀ (except scapus and pedicellus, and apically darkened) brownish yellow; apex of hind femur usually largely black dorsally; basal half of hind tibia (largely) pale yellowish; fore coxa dark brown; 2nd tergite of ♀ orange or dark reddish brown, of ♂ largely black; 4th–6th tergites of males flat, and with long dense setosity (Fig. 590). Description. Redescribed ♀ (BZL), Czech Republic (Pisek); length of fore wing 5.8 mm, of body 8.1 mm.

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Figures 573–576. Aleiodes periscelis (Reinhard), ♀, 573, 574 Russia (holotype R. jaroslawensis Kokujev) and 575, 576 Czech Republic, Pisek 573 head anterior 574, 575 habitus lateral 576 ovipositor sheath lateral. Photographs 573, 574 by K. Samartsev.

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Figures 577–589. Aleiodes periscelis (Reinhard), ♀, Czech Republic, Pisek 577 fore wing 578 hind wing 579 mesosoma lateral 580 mesosoma dorsal 581 metasoma dorsal 582 fore femur lateral 583 hind femur lateral 584 head anterior 585 head dorsal 586 head lateral 587 base of antenna 588 apex of antenna 589 outer middle tarsal claw.

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Figure 590. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov, habitus lateral.

Head. Antennal segments 45 (holotype ♀ of A. jaroslawensis: 42), length of antenna approx. as long as fore wing, its subbasal and subapical segments robust (Figs 587, 588); frons largely smooth anteriorly and with curved rugae posteriorly; OOL 1.5 × diameter of posterior ocellus, rugulose and shiny; vertex finely rugose and with satin sheen; clypeus slightly convex and mainly transversely aciculate; ventral margin of cly­ peus thick and not protruding anteriorly (Fig. 586); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 584); length of eye 1.5 × temple in dorsal view (Fig. 585); vertex behind stemmaticum rugulose; clypeus below lower level of eyes; length of malar space 0.45 × length of eye in lateral view and 1.3 × basal width of mandible. Mesosoma. Mesoscutum remotely punctulate and with satin sheen, interspaces of lateral lobes largely smooth, and of middle lobe superficially coriaceous; scutellum superficially punctate, laterally rugose; precoxal area of mesopleuron smooth and shiny; metapleuron largely densely punctate, but ventrally coarsely rugose; metanotum with distinct median carina anteriorly; propodeum rather flat and coarsely vermiculate rugose, medio-longitudinal carina complete, and slightly tuberculate laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 577); 1-CU1 horizontal, 0.3 × as long as 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 577); cu-a vertical, straight; 1-M nearly straight posteriorly and subparallel; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 densely setose. Hind wing: marginal cell gradually widened

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Figures 591–603. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov 591 wings 592 mesosoma lateral 593 mesosoma dorsal 594 1st –5th metasomal tergites dorsal 595 fore femur lateral 596 hind femur lateral 597 hind tarsus lateral 598 head anterior 599 head dorsal 600 head lateral 601 base of antenna 602 apex of antenna 603 inner hind tarsal claw.

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(but less so in its basal third) and apical width 2.2 × width at level of hamuli (Fig. 578); 2-SC+R quadrate; m-cu short; M+CU:1-M = 40:33; 1r-m 0.5 × 1-M. Legs. Tarsal claws with conspicuous and medium-sized brownish pecten, remaining removed from tarsal tooth (Fig. 589); hind coxa largely coriaceous-punctate, but dorsal besides smooth depression rugose; hind trochantellus rather robust; length of hind femur 3.7 × its width. Metasoma. First tergite rather flat posteriorly, wide subbasally and 0.9 × longer than wide apically; 1st and 2nd tergites with coarse medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite rather finely rugose; medio-basal area of 2nd tergite triangular and wide (Fig. 581); 2nd suture moderately deep, finely crenulate and narrow; basal two-thirds of 3rd tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured and with satin sheen; 4th and apical half of 3rd tergites without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Figs 574, 576). Colour. Black; palpi dark brown basally and remainder pale brown; antenna (except dark brown scapus and pedicellus), tegulae (but anteriorly dark brown), middle and hind trochanters and trochantelli brownish yellow; fore coxa, trochanter and femur dark brown; basal 0.4 of hind tibia ivory and remainder black; remainder of legs (but hind femur with a blackish patch dorso-apically), 1st and 2nd tergites and basal two thirds of 3rd tergite, largely dark reddish brown; pterostigma dark brown; veins mainly yellowish brown, but medially brown (Figs 577, 578); wing membrane subhyaline. Variation. Holotype of A. jaroslawensis has apex of hind femur yellowish brown (Fig. 574). Antennal segments: ♀ 42(1), 45(1); ♂ 50(1), 53(1), 54(2), 56(1). Males appear to have ca ten more antennal segments than females. Male has apical tergites type 1, setae moderately long and fringe not observed, probably absent (Fig. 590); antenna rather dark brown, but scapus largely blackish brown and antennal segments slightly slenderer than of female and 1.1–1.2 × as long as fore wing; metasoma black, but extreme apex of 1st tergite yellowish brown; OOL approx. 1.4 × width of posterior ocellus. Distribution. Austria, Czech Republic, Germany, Hungary, Russia. New synonymy. The ♀ holotype of Rhogas jaroslawensis lacks the antennae, but according to the original description the antenna was 42-segmented, distinctly shorter than the body, reddish brown, except for the darkened apex and the black scapus. This and the other characters still visible agree well with our interpretation of A. periscelis (except that the hind femur is yellowish brown apically); therefore, we synonymise R. jaroslawensis with A. periscelis (syn. nov.). Aleiodes pulchripes Wesmael, 1838 Figs 604–626 Aleiodes pulchripes Wesmael. 1838: 102; Čapek & Lukás, 1989: 31; Papp, 1991a: 73, 2005: 177; Fortier & Shaw, 1999: 224; O’Connor et al., 1999: 92; Belokobylskij et al., 2003: 398 [examined].

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Rogas pulchripes; Shenefelt, 1975: 1245. Rogas (Rogas) pulchripes; Tobias, 1976: 83, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) pulchripes; Papp, 1985a: 149, 153, 161, 1991a: 73. Aleiodes pulchricornis Kolubajiv, 1962: 27; Shenefelt, 1975: 1245 (invalid emendation; not A. pulchricornis (Szépligeti, 1902)); Papp, 2005: 177 (as synonym of A. pulchripes). Type material. Holotype of A. pulchripes, ♂ (KBIN), “A. pulchripes ♂ mihi”, “A. pulchripes mihi, dét. C. Wesmael”, “Coll. Wesmael”, “Belgique, Charleroi/ teste Papp J., 1983”, “Holotypus”, “Aleiodes pulchripes Wesm., 1838, ♂, Papp, 1983”. Additional material. Austria, British Isles (England: V.C. 59; Isle of Man: V.C. 71: Ireland: V.C. H21), Czech Republic, Finland, Germany, Hungary, Netherlands (GE: Vierhouten; ZH: Leiden; NH: Amsterdam; Sloten), Russia, Sweden. Specimens in BMNH, CNC, IKC, KBIN, MSC, MTMA, NMS, NRS, RMNH, USNM, UWIM, ZISP, ZSSM. Molecular data. MRS847 (Sweden), MRS873 (Sweden). Biology. Collected in (June)July and August. Univoltine, overwintering in an exposed mummy. Reared from the following arboreal acronictine Noctuidae: Acronicta aceris (Linnaeus) (2 [CNC, MSC], Austria, Germany; J. Schwarz), Acronicta psi (Linnaeus) (22 [1 NRS, 2 ZISP]; M.R. Shaw), Acronicta leporina (Linnaeus) (1 [IKC], Finland; M.J. Pellinen), Acronicta tridens (Dennis & Schiffermüller) (4:2; M.R. Shaw), Acronicta psi or tridens (2), indet. Acronictinae (1). A quantitative account of rearing this species at its only known English site, comprising old hedges rich in Sorbus aucuparia bordering a largely reclaimed peat bog, is given by Shaw (1979). Experimental rearings were unfortunately limited to unobserved exposures of multiple hosts in closed boxes; extremely hot weather marred the results, but from one box containing 15 of each of Subacronicta megacephala (Dennis & Schiffermüller) and A. tridens, the surviving 13 S. megacephala were dissected after three days of exposure and contained no hosts, while at least eight of the A. tridens contained parasitoids (two found by dissection + six mummies formed; of the other seven, one contained no parasitoid on dissection + six resulted in moths). This suggests that S. megacephala is outside the host range. Similar but less well quantified experiments also excluded the low-feeding Acronicta rumicis (Linnaeus) and the arboreal lymantriine Erebidae Euproctis similis (Fuessly). It is worth adding that the rather frequent citation of lymantriine hosts in the literature can undoubtedly be explained by misidentification of the setose and rather colourful larvae of most species of arboreal acronictine noctuids. The mummy is dark grey in colour, leaving only little evidence of the patches of bright colour that had been a feature of the host larva. It forms in the caudal part of the host, the anterior segments of which strongly contract towards the extensive point of attachment, and the cocoon occupies approx. 4th–7th abdominal segments. As mummification approaches, the host aligns itself on a narrow aerial twig to which the mummy becomes ventrally adpressed, thus leaving a weakly arched dorsal profile bearing a strong resemblance to an overwintering lateral bud (e.g., of Sorbus aucuparia: Fig. 605).

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Figures 604–607. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 604 habitus lateral 605 mummy of Acronicta psi (Linnaeus) in winter 606 ovipositor sheath lateral, 607 mummy of Acronicta psi (Linnaeus) after emergence of parasitoid.

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Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 616); OOL distinctly less than diameter of posterior ocellus, largely smooth but micro-sculptured near eyes; ventral margin of clypeus thick and not protruding in lateral view (Fig. 618); mesoscutal lobes coriaceous; mesopleuron (including precoxal sulcus area) nearly or completely smooth; propodeum with pair of crest-like protuberances laterally; vein 1-CU1 of fore wing much shorter than vein 2-CU1; basal half of marginal cell of hind wing parallel-sided (Fig. 609); tarsal claws with large dark brown pecten up to apical tooth of claw (Fig. 621); hind spurs dark brown; hind tibial spurs of ♂ obtuse apically (Fig. 624); head black; pterostigma pale yellowish or light brown; mesopleuron, mesosternum and scutellum brownish yellow; apex of hind femur yellowish or reddish; basal half of hind tibia pale yellowish. Description. Redescribed ♀ (RMNH) from England (Chat Moss). Length of fore wing 5.3 mm, of body 6.5 mm. Head. Antennal segments of ♀ 56, length of antenna 1.3 × fore wing, its subapical segments slender (Fig. 620); frons largely smooth; OOL 0.3 × diameter of posterior ocellus, and smooth near ocelli, but micro-sculptured near eye, shiny; vertex largely smooth, with few punctures, shiny; face crest-like protruding medio-dorsally; clypeus densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 618); width of hypoclypeal depression 0.35 × minimum width of face (Fig. 616); length of eye 5.6 × temple in dorsal view (Fig. 617); vertex behind stemmaticum mainly smooth but partly rugulose; clypeus above lower level of eyes; length of malar space 0.15 × length of eye in lateral view. Mesosoma. Mesoscutal lobes coriaceous, rather shiny; precoxal area of mesopleuron smooth as most of mesopleuron; metanotum with medio-longitudinal carina anteriorly; scutellum finely punctate, interspaces smooth, but posteriorly coriaceous; propodeum rather flat medially and rather remote rugose, medio-longitudinal carina nearly complete, and with slightly protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 608); 1-CU1 slightly oblique, 0.4 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell rather robust (Fig. 608); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 sparsely setose. Hind wing: basal half of marginal cell parallel-sided, apical half linearly widened, its apical width twice width at level of hamuli (Fig. 609); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 31:26; 1r-m 0.7 × 1-M and 1-M oblique. Legs. Tarsal claws with conspicuous and robust dark brown pecten up to apical tooth of claw (Fig. 621); hind coxa dorsally largely smooth and remainder remotely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.3 and 5.0 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite evenly convex medially, 0.9 × longer than wide apically, wider than base of 2nd tergite; 1st and 2nd tergites with medio-longitudinal carina and coarsely irregularly sublongitudinally rugose; medio-basal area of 2nd tergite triangular and rather large (Fig. 612); 2nd suture deep and coarsely crenulate; basal half of 3rd tergite rugulose, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 606).

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Figures 608–621. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 608 fore wing 609 hind wing 610 mesosoma lateral 611 mesosoma dorsal 612 1st–3rd metasomal tergites dorsal 613 fore femur lateral 614 hind femur lateral 615 hind tarsus lateral 616 head anterior 617 head dorsal 618 head lateral 619 base of antenna 620 apex of antenna 621 outer hind tarsal claw.

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Figures 622–626. Aleiodes pulchripes Wesmael, ♂, England (ex Acronicta culture) 622 habitus lateral 623 outer hind claw lateral 624 hind tibial spurs and basitarsus lateral 625 metasoma dorsal 626 4th–7th metasomal tergites lateral.

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Colour. Blackish or dark brown; telotarsi, apical 0.4 of hind tibia, hind tibial spurs and hind tarsus dark brown; remainder of hind tibia and palpi yellowish; remainder of legs, pterostigma and tegulae pale brownish yellow; veins brown; mesoscutum mediodorsally, scutellum, metanotum, mesopleuron (except partly antero-dorsally), meso­ sternum and metapleuron orange yellow; wing membrane subhyaline. Variation. Scutellum largely finely punctate, coriaceous medio-posteriorly, but may be striate. Specimens from Sweden are appreciably darker than those from Britain. Vein m-cu of hind wing absent or faintly indicated. Antennal segments: ♀ 51(1), 52(2), 53(2), 54(3), 55(4), 56(6), 57(7), 58(7), 59(2), 60(1), 62(1); ♂ 49(1), 51(1), 53(2), 54(9), 55(6), 56(6), 57(2). Females have on average ca three more antennal segments than males. Males have obtuse hind tibial spurs and the tarsal pecten less developed than in females, propleuron and pronotum yellowish or blackish posteriorly; posterior half of mesoscutum largely yellowish or blackish; apical tergites type 2, somewhat sparse setose, glabrous stripe broad but with some setae directed into it and fringe rather weak (Figs 625, 626). Distribution. *Austria, British Isles (England, Isle of Man, Ireland), Czech Republic, Finland, Germany, Hungary, *Netherlands, Russia, Sweden. Aleiodes quadrum (Tobias, 1976) Figs 627–651 Rogas (Rogas) quadrum Tobias, 1976: 83, 221, 1986: 76 (transl.: 125). Aleiodes (Neorhogas) quadrum; Papp, 1985a: 162, 1991a: 83. Aleiodes quadrum; Papp, 2005: 177. Rogas (Rogas) illustris Papp, 1977a: 112, 1985a: 162 (as synonym of A. quadrum), 1991a: 83 (id.), 2005: 176 (id.) [examined]. Type material. Holotype of A. illustris, ♀ (MTMA), “Yugoslavia, [Croatia:] Kostrena, Rijeka, 12.viii.1966, Uremović”, “Holotypus ♀ Rogas illustris sp. n., Papp, J., 1977”, “Hym. Typ. No. 2378, Mus. Budapest”; paratype, ♀ (MTMA), “[Hungary], Hársbokorhegy, Nagykovacsi”, “1.viii.1952, Bajári”, “Paratypus ♀ Rogas illustris sp. n., Papp, J., 1977”, “Hym. Typ. No. 2380, Mus. Budapest”. Additional material. 1 ♀ (NMS), “France: Ardèche, Accons, UV light, 24.vi.2013, M.R. Shaw”, “MRS Aleiodes DNA 796”; 1 ♀ (NMS), “France: Savoie, Queige, Le Villaret, 700m., 19.vi. 2019, C.W. Plant”; 1 ♀ (BZL), “Turkey, 15 km W Refahye, W of Erzincan, 1600 m, 7.vii.2000, M. Halada”; 1 ♀ (BZL), “GRC [= Greece], Westmakadonien, Florina, Aussichtsplatz SE Karies, 40°45'2"N, 21°10'39"E, 1080 m msl, 27.vi.2016, 2016/31, LF, H. u. R. Rausch”; 1 ♀ (MTMA), “[North] Macedonia, Skopje Prov., Mt. Vodno, 16.vii.1997, Gy. Rozner”; 1 ♀ (NMS), “N. Macedonia, Vardar river valley, above Demir Kapiya, N41°22'58", E22°11'45", 244m, 13.vii.2019 S. Beshkov & A. Nahirnic”; 1 ♂ (NMS), “Bulgaria: Haskovo, E. Rhodopes, SW Mezek, 450 m, MV light, 17.vii.2015, C.W. Plant”.

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Figures 627, 628. Aleiodes quadrum (Tobias), ♀, France, Accons 627 habitus lateral 628 ovipositor sheath lateral.

Molecular data. MRS796 (France), additionally MRS824 (Bulgaria) likely to be a male of this species. Biology. Unknown. Collected in June–July, likely to be univoltine, but there is nothing to suggest how the winter is passed. We have not seen reared material, but the elongate and strongly apically compressed metasoma suggests that the host would be concealed, perhaps between spun leaves, in a leaf sheath, in a seed capsule or in some similar situation. Diagnosis. Maximum width of hypoclypeal depression approx. 0.6 × minimum width of face (Fig. 636); OOL of ♀ 0.6–0.7 × diameter of posterior ocellus and rugose; ventral margin of clypeus obtuse apically and clypeus hardly protruding anteriorly (Fig. 638); lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures, smooth and shiny; precoxal area distinctly rugose, but posteriorly only punctate; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein 1-CU1 of fore wing approx. 0.8 × vein 2-CU1 and as long as m-cu (Fig. 629); surroundings of veins M+CU and 1-M of hind wing largely glabrous; hind tarsal claws with conspicuous dark brown pecten close to apical tooth (Fig. 635); 1st tergite parallel-sided and longer than wide apically (Fig. 632); 2nd tergite of ♀1.0–1.2 × as long as wide basally and black; head black; vein 1-M of fore wing dark brown; wing membrane slightly infuscate. Redescription. ♀ (NMS) from France (Accons). Length of fore wing 6.9 mm, of body 9.6 mm.

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Figures 629–641. Aleiodes quadrum (Tobias), ♀, France, Accons 629 wings 630 mesosoma lateral 631 mesosoma dorsal 632 metasoma dorsal 633 fore femur lateral 634 hind femur lateral 635 outer hind tarsal claw 636 head anterior 637 head dorsal 638 head lateral 639 base of antenna 640 apex of antenna 641 antenna.

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Figure 642. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo, habitus lateral.

Head. Antennal segments of ♀ 53, antenna as long as fore wing, its subapical segments rather robust (Fig. 640); frons with curved striae but medially largely smooth; OOL 0.7 × diameter of posterior ocellus, finely rugose and shiny; vertex coarsely punctate but behind ocelli rugose, rather shiny; clypeus nearly flat and coarsely rugosepunctate; ventral margin of clypeus thick and hardly protruding anteriorly (Fig. 638); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 636); length of eye 1.6 × temple in dorsal view (Fig. 637); clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, with interspaces approx. equal to diameter of punctures, smooth and shiny; scutellum finely and densely punctate; precoxal sulcus area of mesopleuron distinctly rugose but posteriorly only punctate, remainder of mesopleuron distinctly but remotely punctate; metapleuron remotely punctate, but ventrally rugose; propodeum evenly convex and coarsely vermiculate-rugose and medio-longitudinal carina nearly complete. Wings. Fore wing: r 0.4 × 3-SR (Fig. 629); 1-CU1 horizontal, 0.8 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 629); cu-a nearly vertical

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Figures 643–651. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo 643 wings 644 1st–3rd metasomal tergites dorsal 645 3rd–7th metasomal tergites lateral 646 id. dorsal 647 basal antennal segments 648 head anterior 649 head dorsal 650 head lateral 651 inner hind tarsal claw.

and straight; 1-M curved posteriorly; 1-SR rather slender; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 629); 2-SC+R short longitudinal; m-cu narrowly present; M+CU:1-M = 50:33; 1r-m 0.8 × 1-M. Legs. Tarsal claws with conspicuous and robust blackish pecten, close to level of apical tooth (Fig. 635); hind coxa largely densely punctate; hind trochantellus rather

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robust; length of hind femur and basitarsus 4.3 and 5.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, 1.3 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and finely longitudinally rugose, but posterior quarter of 2nd tergite smooth and no median carina; medio-basal area of 2nd tergite triangular and wide (Fig. 632); 2nd tergite as long as wide basally and with shallow transverse impression; 2nd suture shallow and narrowly crenulate; 3rd and subsequent tergites finely punctulate and strongly shiny; apical half of 3rd and 4th tergites without sharp lateral crease; ovipositor sheath widened apically, with medium-sized setae and apically truncate (Fig. 628). Colour. Black; hind tibia dark brown apically and remainder pale yellowish; middle and hind tarsi, and fore telotarsus blackish or dark brown; mandible, remainder of legs, mesoscutum, pronotum postero-dorsally, mesopleuron dorsally, scutellum, metanotum, ovipositor sheath and 1st tergite (except pair of dark patches apically) orange; tegulae yellowish; palpi and pterostigma blackish; vein dark brown, but vein at base of wings yellowish; wing membrane largely slightly infuscate. Variation. OOL of ♀ 0.6–0.7 × diameter of posterior ocellus. The female from Turkey is very similar but has 1st tergite 1.4 × as long as wide apically and 2nd tergite 1.2 × longer than wide basally. Antennal segments: ♀ 53(1), 56(1), 57(1), 58(1). Apical tergites of male type 2 with fringe rather strong (Figs 645, 646). The figured male from Bulgaria has OOL rugose anteriorly, 1st tergite 1.2 × as long as wide posteriorly, 2nd tergite completely parallel-sided and 0.9 × as long as wide, 3rd tergite rugose-striate in anterior half and parallel-sided 0.7 × as long as wide, fore femur 4.5 × longer than wide and hind femur 4.1 × longer than wide. Distribution. Azerbaijan, *Bulgaria, Croatia, *France, *Greece, Hungary, *North Macedonia, *Turkey. Notes. The holotype of A. quadrum is a male and it is less reliable to identify this species from it than from the holotype female of A. illustris; nevertheless, we accept the synonymy proposed by Papp (1985a). The figured male from Bulgaria (NMS; Figs 642–651) is considered to be this species (initially through its CO1 sequence); it is morphologically very similar to A. cruentus and there is a possibility that some similar males have been returned to depositories determined as A. cruentus with no recognition that they might belong to A. quadrum. However, the matter remains unresolved until more males of A. quadrum become available. Aleiodes ruficeps (Telenga, 1941) Figs 652–666 Rhogas (Rhogas) ruficeps Telenga, 1941: 179, 421, Fig. [examined]. Rogas ruficeps; Shenefelt, 1975: 1146; Zaykov, 1980b: 87. Rogas (Rogas) ruficeps; Tobias, 1976: 81, 1986: 76 (transl.: 122; lectotype designation). Aleiodes ruficeps; Papp, 1991a: 88; Fortier & Shaw, 1999: 230. Rogas gasterator auctt. p.p.

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Type material. Lectotype of A. ruficeps, ♀ (ZISP), “[Russia: Crimea,] Eupatoria [= Eupatoriya], Tavrits, gub, 7.v.1907, V.E. Jakovlev”, “Rhogas ruficeps sp. n., Telenga det.”, “Lectotype Rogas ruficeps Tl., design. Tobias, 1980”. Additional material. Bulgaria, Russia, Turkey, [Armenia, Iran]. Specimens in BMNH, BZL, NMS, RMNH, ZISP, ZSSM. Molecular data. None. Biology. Unknown. Specimens collected in April-May; presumably univoltine, but there is nothing to suggest how it overwinters. We have not seen reared material. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 661); OOL ca twice diameter of posterior ocellus and moderately punctate (Fig. 662); 4th–10th antennal segments approx.as long as wide (Figs 652, 664); ventral margin of clypeus thick to rather sharp and distinctly protruding in lateral view (Fig. 663); mesoscutal lobes punctate and interspaces largely smooth and shiny, lobes rather convex; scutellum sparsely punctate; precoxal sulcus coarsely vermiculate-rugose; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 654); length of vein r of fore wing 0.3 × vein 3-SR (Fig. 654); vein 1-CU1 of fore wing 0.3–0.5 × vein 2-CU1 (Fig. 654); hind tarsal claws rather slender, hardly curved and only brownish setose (Fig. 666); head completely or largely orange or yellowish; palp dark brown basally; hind femur apico-dorsally dark brown or black; hind tibia pale yellowish but apically darkened; 4th and 5th tergites black; wing membrane nearly entirely infuscate. Resembles A. grassator because of the robust antennal segments and dark wings, but A. ruficeps has frons, OOL, vertex, malar space, and third tergite less sculptured, apex of hind tibia and palpi dark brown, basal antennal segments of ♀ somewhat less robust, hypoclypeal depression wider, marginal cell of fore wing slenderer and vein r of fore wing shorter. Differs from the similar A. ruficornis by having hypoclypeal depression wider, clypeus wider and lower, apical antennal segments of ♀ slenderer, OOL less sculptured and more antennal segments (♀: 45–47 vs 35–39(–41) of A. ruficornis). Description. Lectotype, ♀, length of fore wing 7.3 mm, of body 8.6 mm. Head. Antennal segments of ♀ 45, length of antenna 1.1 × fore wing, its subapical segments moderately robust (Fig. 665); frons with few rugae, remainder smooth; OOL 2.0 × diameter of posterior ocellus, and moderately densely punctate; vertex spaced punctate and shiny; clypeus punctate; ventral margin of clypeus thick and distinctly protruding forwards (Fig. 663); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 661); length of eye 1.1 × temple in dorsal view (Fig. 662), temples subparallel-sized behind eyes; vertex behind stemmaticum punctate; clypeus distinctly below lower level of eyes; occipital carina widely reduced ventrally (Fig. 663); length of malar space 0.7 × length of eye in lateral view. Mesosoma. Mesoscutal lobes punctate and interspaces smooth, shiny; precoxal area of mesopleuron coarsely vermiculate-rugose, near precoxal area mesopleuron distinctly punctate; scutellum rather flattened, sparsely punctate; propodeum coarsely vermiculate-rugose, medio-longitudinal carina incomplete, absent posteriorly and propodeum rounded laterally.

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Figures 652, 653. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 652 habitus lateral 653 ovipositor sheath lateral.

Wings. Fore wing: r 0.3 × 3-SR (Fig. 654); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell rather long (Fig. 654); cu-a slightly inclivous, straight; 1-M slightly curved posteriorly; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell linearly widened, its apical width 2.5 × width at level of hamuli (Fig. 655); 2-SC+R longitudinal; m-cu pigmented only basally; M+CU:1-M = 7:5; 1r-m 0.7 × 1-M. Legs. Tarsal claws rather slender, hardly curved and with six brownish bristles (Fig. 666); hind coxa rather weakly punctate; hind trochantellus robust; hind femur distinctly punctate; fore femur 3.3 × longer than wide; length of hind femur and basitarsus 3.2 and 5.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 0.9 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and regularly longitudinally rugose; mediobasal area of 2nd tergite triangular and short (Fig. 658); 2nd suture rather deep and finely crenulate; basal half of 3rd tergite finely striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 653).

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Figures 654–666. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 654 fore wing 655 hind wing 656 mesosoma lateral 657 mesosoma dorsal 658 1st –3rd metasomal tergites dorsal 659 fore femur lateral 660 hind femur lateral 661 head anterior 662 head dorsal 663 head lateral 664 base of antenna 665 apex of antenna 666 inner hind tarsal claw.

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Colour. Orange brown; antenna, palpi, apices of femora, telotarsi, bases of fore and middle coxae, pterostigma, and veins (but 1-SR much paler than 1-M) dark brown; mesosoma (except for mesoscutum and scutellum), ovipositor sheath, 3rd tergite (except basally) and subsequent tergites black or blackish; apex of hind tibia only narrowly dark brown; wing membrane nearly entirely infuscate. Variation. Vein 1-CU1 of fore wing 0.3–0.5 × vein 2-CU1. Females may have the flagellum extensively pale basally, or entirely dark. Antennal segments: ♀ 45(1), 46(1), 47(1); ♂ 56(1), 58(1). Males appear to have ca ten more antennal segments than females. Male has apical tergites type 1, setae rather long and sparse, and fringe not observed. Distribution. *Armenia, Bulgaria, *Iran, Russia, *Turkey. Aleiodes ruficornis (Herrich-Schäffer, 1838) Figs 667–685 Rogas ruficornis Herrich-Schäffer, 1838: 156, fig.; Shenefelt, 1975: 1224 (as synonym of A. dimidiatus) [neotype designated below]. Aleiodes (Neorhogas) ruficornis; Papp, 1985a: 152 (as synonym of A. dimidiatus), 1991a: 90 (id.). Aleiodes (Chelonorhogas) ruficornis; Belokobylskij et al., 2003: 398; van Achterberg, 2014: 209; Abdolalizadeh et al., 2017: 37. Aleiodes ruficornis; Bergamasco et al., 1995: 5; Zaldivar-Riverón et al., 2004: 234; Papp, 2005: 176 (as synonym of A. dimidiatus). Aleiodes brevicornis Wesmael, 1838: 98; Shenefelt, 1975: 1224; Papp, 1985a: 152, 157 (as synonym of A. dimidiatus), 2005: 176 (id.); Belokobylskij et al. 2003 (as synonym of A. ruficornis) [examined]. Aleiodes (Neorhogas) brevicornis; Papp, 1991a: 90 (as synonym of A. dimidiatus). Aleiodes nigripalpis Wesmael, 1838: 97; Shenefelt, 1975: 1224; Papp, 1985a: 152, 157 (lectotype designation; as synonym of A. dimidiatus), 1991a: 90, 2005: 176 (id.); Belokobylskij et al. 2003 (as synonym of A. ruficornis) [examined]. Rhogas dimidiatus ab. nigrobasalis Hellén, 1927: 24 (invalid name). Rhogas dimidiatus ab. ruficollis Hellén, 1927: 24 (invalid name). Rhogas carbonarius ab. giraudi Fahringer, 1931: 236 [unavailable name for melanistic males]. Rhogas carbonarius var. giraudi Telenga, 1941: 168. Syn. nov. Rogas dimidiatus ab. infuscatus Hellén, 1957: 49 (invalid name). Rogas dimidiatus ab. nigripes Hellén, 1957: 49 (invalid name). Aleiodes (Aleiodes) arnoldii; Farahani et al., 2015: 232–233 (but see note under A. arnoldii). Rogas gasterator auctt. p.p. Rogas dimidiatus auctt. p.p. Type material. Neotype of A. ruficornis here designated, ♀ (NMS), “[Germany,] Einbeck, L. 31.v.[19]85, [R. Hinz]”, “ex: Hoplodrina blanda Schiff. (Lep.)”. Holotype of A. brevicornis, ♀ (KBIN), “A. brevicornis ♀ mihi. 3.”, “Coll. Wesmael”, “A. brevicornis

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Figures 667–669. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 667 habitus lateral 668 ovipositor sheath lateral 669 mummy of Hoplodrina blanda (Denis & Schiffermüller).

mihi, dét. C. Wesmael”, “Belgique, Liège, leg. Robert/ teste Papp J., 1983”, “Holotypus Aleiodes brevicornis Wesm., 1838 ♀, det. Papp, 1983”, “Aleiodes dimidiatus Spin. ♀, det. Papp J., 1983”. Holotype of A. nigripalpis, ♂ (KBIN), “A. nigripalpis ♂ mihi. 2.”, “Coll. Wesmael”, “A. nigripalpis mihi, dét. C. Wesmael”, “Belgique, Liège/ teste Papp J., 1983”, “Lectotypus Aleiodes nigripalpis Wesm., 1838 ♂, det. Papp, 1983”, “Aleiodes dimidiatus Spin. ♂, det. Papp J., 1983”. The neotype designation for A. ruficornis is necessary for nomenclatural stability, because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; the first author could not find any specimen in ZMB) and the species has been confused with similar species in the past. The specimen is selected because it fits well the original description, the probable type location was in Germany, it has been reared and it is in good condition. Additional material. Andorra, Austria, British Isles (England (V.C. 5, 11, 13, 15, 17, 19, 20, 22, 23, 24, 28, 29, 31, 33, 38, 39, 60, 63); Wales (V.C. 52) [no specimens seen from Scotland]), Bulgaria, Croatia, Czech Republic, Finland, France, Germany, Greece, Hungary, Italy (including Sicily), Moldova, Montenegro, Netherlands (DR: Borger; LI: Venlo, ZH: Oostkapelle), North Macedonia, Norway, Romania,

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Russia (including Far East), Serbia, Slovakia, Sweden, Switzerland, Turkey, Ukraine, [Afghanistan, Dagestan, Iran, Kazakhstan, Kyrgyzstan]. Specimens in ALC, BMNH, BZL, FMNH, MMUM, MRC, MSC, MSNV, MTMA, NMS, NRS, OUM, RMNH, SDEI, UNS, UWIM, ZISP, ZSSM. This is a widespread species, generally common, but partly replaced by A. gasterator in the Mediterranean region. Molecular data. MRS140 (UK), MRS877 (Sweden), MRS888 (UK), MRS890 (UK), MRS891 (UK). Biology. Specimens collected from April–September; probably plurivoltine in the southern part of its range but in Britain univoltine, flying from June–August with a varied means of overwintering (see below). Reared from the noctuids Agrotis clavis (Hufnagel) (4 [4 ZISP]/ Russia), Agrotis segetum (Dennis & Schiffermüller) (1 [FMNH]/Finland), Agrotis sp. (3), Euxoa nigricans (Linnaeus) (1 [FMNH]/Finland), Euxoa sp. (3:1 [3 FMNH/Finland], Hoplodrina blanda (Dennis & Schiffermüller) (4:1, Germany; R. Hinz), Hoplodrina octogenaria (Goeze) (1; W.A. Watson), Mythimna impura (Hübner) (1 [ZSSM]/Germany; E. Haeselbarth). A further mummy from H. blanda failed to emerge (M.R. Shaw), but was no doubt of this species. The above hosts belong to three different subfamilies of Noctuidae, all feeding and resting close to ground level. In addition, we have seen a specimen labelled as ex the nymphalid Brenthis ino (Rottemburg) but accompanied by a mummy of a noctuid, probably Hoplodrina sp. (det. M.R. Shaw). In experiments a range of hosts recorded for this species in the literature (several arctiine and lymantriine Erebidae and the lasiocampid Lasiocampa quercus (Linnaeus)) that are actually hosts of superficially similar species such as A. alternator (Nees) were offered to the female reared ex H. octogenaria but, as expected, they were firmly rejected. However, this female readily accepted late 1st instar larvae of Agrotis exclamationis (Linnaeus), from which adult progeny resulted very smoothly (1:16\13\\12\12+0. The few failures to oviposit were almost certainly due to egg depletion). Searching in the vicinity of hosts included antennal drumming (the tips curled downwards) and indeed the antennae seemed to be the only proximal means of locating and assessing the host. Once the host was found it was immediately accepted, rapidly jabbed and stood over or often withdrawn from (1.0–1.5 cm) while the venom took affect (20–40 secs), then relocated via antennal searching (when it had been withdrawn from this might take up to a minute, but it was always eventually successful) scooped in with the fore legs (the antennae only slightly involved), positioned and held between the mid legs for the duration of oviposition (20–30 secs). Frequently the host larva was kicked free of the ovipositor by the parasitoid’s hind leg(s) and the parasitoid rapidly left without any period of post-oviposition association. Recovery from the venom was rather protracted (up to 20 mins), during which time hosts were rejected if rediscovered. Towards the end of successful oviposition runs it was evident that venom depletion ran ahead of egg depletion, resulting in erratic (but nevertheless successful) oviposition sequences. No host feeding took place. In this experimental series oviposition took place in mid-July with mummification at the end of August and adult emergence in late May of the following year. However, although the winter was passed in the mummy in this entire series (and probably also the case for the other,

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Figures 670–681. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 670 wings 671 mesosoma lateral 672 mesosoma dorsal 673 metasoma dorsal 674 fore femur lateral 675 hind femur lateral 676 head anterior 677 head dorsal 678 head lateral 679 base of antenna 680 apex of antenna 681 outer hind tarsal claw.

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natural, Agrotis hosts), it is clear that the rearings from Hoplodrina and probably also Mythimna involved overwintering in the host larva with adult emergence in the year of mummification. This host-related difference in overwintering is not inconsistent with univoltinism in Britain (where the experiments and other observations were done) but it is certainly an interesting quirk of its host range and might be of significance in suggesting one way in which a temporal isolating mechanism could potentially arise as a forerunner to speciation (cf. Shaw, 2003). The mummy is formed in the soil and is not strongly (if at all) glued to the substrate. It is predominantly dark brown in colour, very large in relation to the size of the insect that will emerge and, although basically cylindrical, somewhat flattened in appearance owing to a pronounced but blunt lateral keel (Fig. 669). It is more or less strongly contracted at the anterior end, markedly less so caudally, and the copiously silken lining typically occupies 3rd–8th abdominal segments. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 676); OOL of ♀ 1.4–1.6 × as long as diameter of posterior ocellus and distinctly rugose or rugulose (Fig. 677); length of 4th antennal segment of ♀ 1.0–1.4 (of ♂ 1.1–1.4) × its width (Fig. 679); ventral margin of clypeus thick and not protruding anteriorly (Fig. 678); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 670); vein 1-CU1 of fore wing 0.4–0.6 × as long as vein 2-CU1; hind tarsal claws yellowish or brownish bristly setose and with few yellowish pectinal teeth (Fig. 681); hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; pale males have whole frons and stemmaticum yellowish; palpi dark brown or blackish, rarely brown; 3rd metasomal tergite only antero-laterally reddish or yellowish; 4th and 5th tergites black. Specimens from high altitude have the head conspicuously long setose and the tarsal claws brownish pectinate basally. In this respect males are similar to A. hirtus, but A. hirtus has precoxal area and mesoscutum largely smooth and clypeus distinctly protruding in lateral view. Description. Neotype, ♀, length of fore wing 3.9 mm, of body 6.5 mm. Head. Antennal segments of ♀ 35, antenna as long as fore wing, its subbasal and subapical segments robust (Figs 679, 680); frons with curved rugae; OOL 1.2 × diameter of posterior ocellus, rugose and moderately shiny; vertex rugose, rather shiny; cly­ peus rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 678); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 676); length of eye twice temple in dorsal view (Fig. 677); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.5 × length of eye in lateral view; temple punctate and shiny, but rugulose near occipital carina. Mesosoma. Mesoscutal lobes distinctly punctate, interspaces of lateral lobes smooth and shiny; precoxal area of mesopleuron coarsely vermiculate-rugose medially, but posteriorly punctate; mesopleuron punctate medially; metapleuron distinctly rugose ventrally and dorsally punctate; scutellum largely smooth (except for punctulation), rather shiny and nearly flat, with lateral carina; propodeum coarsely reticulaterugose, laterally dorsal face longer than posterior one, somewhat angulate laterally but without tubercles, and with complete medio-longitudinal carina.

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Figures 682–685. Aleiodes ruficornis (Herrich-Schäffer), ♂, U.K. (culture) 682 habitus lateral 683 apex of antenna 684 base of antenna 685 mummy of Hoplodrina octogenaria (Goeze).

Wings. Fore wing: r 0.3 × 3-SR; marginal cell short (Fig. 670); 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 670); cu-a inclivous, straight; 1-M rather curved posteriorly; 1-SR slightly wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell lin-

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early widened, its apical width 2.2 × width at level of hamuli (Fig. 670); 2-SC+R subquadrate; m-cu narrowly pigmented; M+CU:1-M = 30:19; 1r-m 0.6 × 1-M. Legs. Tarsal claws mainly setose and medially with 4 yellowish rather short pectinal teeth (Fig. 681); hind coxa punctate and shiny; hind trochantellus robust; length of hind femur and basitarsus 3.3 and 4.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather convex medially, 0.9 × longer than wide apically, robust and coarsely irregularly longitudinally rugose as 2nd tergite; 1st tergite and basal half of 2nd tergite with medio-longitudinal carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 673); 2nd suture deep and crenulate; basal half of 3rd tergite largely longitudinally striate, remainder of metasoma superficially micro-sculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 668). Colour. Reddish or orange-brown; stemmaticum medially, malar space largely, temple and occiput ventrally, mesosternum, mesopleuron ventrally, 3rd tergite (except anterolaterally), 4th–7th tergites black; palpi (only labial palp darkened basally) brown; basal half of antenna, tegulae, parastigma, and base of pterostigma pale yellowish; apical half of antenna, pedicellus dorsally, propodeum dorsally and medially, middle and hind femora apico-dorsally, and telotarsi dark brown; remainder of pterostigma and veins dark brown or brown (Fig. 670); fore wing membrane rather infuscate, but hind wing nearly subhyaline. Variation. Female: mesosoma occasionally wholly black. Male face and mesosoma usually black but can be variably marked with red; scape and pedicel usually (partly) reddish in central and southern populations but most often entirely black in more north-western ones (e.g., British Isles, Sweden); hind coxa varies from black to red. Length of malar space 0.5–0.6 × length of eye in lateral view; head black or largely reddish brown (except temple ventrally and malar space); interspaces of mesoscutal lobes smooth to micro-sculptured; 1-CU1 0.4–0.6 × 2-CU1; 3rd tergite longitudinally striate or rugulose basally (sometimes narrowly so), without curved sculptural elements (Fig. 673), except sometimes some weak transverse striae occasionally present at extreme apex; males from montane habitats are generally darker than lowland males. Antennal segments: ♀ 34(6), 35(12), 36(15), 37(22), 38(17), 39(13), 40(2), 41(3), 42(2), 43(1); ♂ 43(1), 44(1), 45(1), 46(1), 47(5), 48(13), 49(32), 50(19), 51(36), 52(36), 53(28), 54(11), 55(4), 56(4), 57(2). On average males have ca 14 more antennal segments than females. Male has marginal cell of fore wing less robust than in ♀, with apical tergites type 1–2, density of setae rather variable and fringe evident but sparse (Fig. 682). Distribution. *Afghanistan, *Andorra, Austria, British Isles (England, Wales), *Bulgaria, *Croatia, *Czech Republic, *Finland, *France, Germany, *Iran, *Kazakhstan, *Kyrgyzstan, *Montenegro, *Netherlands, *North Macedonia, *Norway, *Romania, *Russia (including Dagestan and Far East), *Serbia, *Slovakia, *Sweden, Switzerland, *Turkey, *Ukraine. Notes. An examined female (NMS) from Hungary, Borzsony Mts., 140 m altitude, 20–30.vii.2005 (unfortunately, too damaged for description) represents a very similar but new species. The 4th–10th antennal segments are not moniliform, slenderer

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than in typical A. ruficornis, the fore femur is more robust than in A. ruficornis, and the COI sequence (MRS886) is different (2.1 %). Aleiodes ruficornis is the commonest and most widespread of a small group of related species parasitising grassland and “cutworm” hosts, exhibiting strong sexual dimorphism with unremarkable males but the more extensively orange females having a stronger build and much shorter antennae. The least extreme in these respects is A. gasterator, which largely (but not completely) replaces A. ruficornis in the Mediterranean region. Aleiodes grassator is similar to A. ruficornis, but it appears to be restricted to montane and northern habitats where it might be thought to replace A. ruficornis. However, some males that morphologically agree best with A. ruficornis have been collected at high altitude in the Alps (up to 2550 m), where A. improvisus also occurs, but whether these highaltitude A. ruficornis males are parts of breeding populations or have simply been carried up in thermals is impossible to say. The females in this group (excluding A. gasterator) are scarcer in collections than males, as they fly very little and rarely enter Malaise traps. Aleiodes rufipes (Thomson, 1892) Figs 686–700 Rogas rufipes Thomson, 1892: 1669; Shenefelt, 1975: 1224; Kotenko, 1992: 96 [examined]. Rogas (Rogas) rufipes; Tobias, 1986: 81 (transl.: 133). Aleiodes (Neorhogas) rufipes; Papp, 1985a: 162, 1987b: 36, 1991a: 88; Belokobylskij, 1996: 14; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) rufipes; Chen et al., 1992: 496; Belokobylskij, 2000: 40; Chen & He, 1997: 42; He et al., 2000: 665; Ghahari et al., 2011: 4; Farahani et al., 2015: 229, 244. Aleiodes rufipes; Fortier & Shaw, 1999: 228; Papp, 2002: 562, 2005: 177; Aydogdu & Beyarslan, 2005: 191, 192. Type material. Holotype, ♀ (ZIL), “Lap”, “rufipes m”, “Sverige [= Sweden], Lappland, teste Papp J., 1983”, “Holotypus”, “Rogas rufipes Thoms., 1891, ♀, Papp, 1983”. Additional material. Finland, Norway, Sweden. Specimens in FMNH, NMS, MTMA, RMNH, ZIL. Molecular data. MRS294 (Sweden), MRS312 (Sweden), MRS314 (Sweden), MRS673 (Finland), MRS674 (Finland), MRS676 (Finland), MRS680 (Finland). Biology. Unknown. Collected from July–August; presumably univoltine. We have not seen reared material and there is no indication of how the winter may be passed. Diagnosis. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Fig. 695); OOL of ♀ 1.1–1.5 × as long as diameter of posterior ocellus and punctate-rugulose to coriaceous-rugose (Fig. 696); length of antenna of ♀ 1.0–1.1 × length of fore wing; ventral margin of clypeus thin and not protruding in lateral view (Fig. 697); mesoscutal lobes remotely punctulate and with satin sheen; area

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Figures 686, 687. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 686 habitus lateral 687 ovipositor sheath lateral.

of precoxal sulcus smooth; length of vein 1-CU1 of fore wing 0.2–0.3 × vein 2-CU1 and 0.4 × vein m-cu; vein 2-SC+R of hind wing subquadrate; tarsal claws with robust apical tooth and with medium-sized yellowish brown pecten (Fig. 698); hind femur and basitarsus slender (Fig. 686); 1st metasomal tergite comparatively steep anteriorly (Fig. 686); head (largely) black; apex of hind femur usually largely black dorsally; basal half of hind tibia (largely) pale yellowish; 2nd tergite yellowish or reddish; males usually with dense and long setosity on at least basal half of 4th–6th tergites.

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Figures 688–700. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 688 fore wing 689 hind wing 690 mesosoma lateral 691 mesosoma dorsal 692 1st–3rd metasomal tergites dorsal 693 fore femur lateral 694 hind femur lateral 695 head anterior 696 head dorsal 697 head lateral 698 outer hind tarsal claw 699 base of antenna 700 apex of antenna.

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Description. Redescribed ♀ (RMNH) from Finland (Enntekiö). Length of fore wing 5.3 mm, of body 5.8 mm. Head. Antennal segments of ♀ 51, antenna as long as fore wing, its subapical segments rather robust and apical segment with short spine (Fig. 700); frons largely smooth, except for some micro-sculpture; OOL 1.2 × diameter of posterior ocellus, coriaceous-rugose and slightly shiny, groove beside posterior ocellus rather shallow and crenulate; vertex coriaceous with some rugulae, rather dull; clypeus rugose; ventral margin of clypeus thin and not protruding forwards (Fig. 697); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 695); length of eye 1.7 × temple in dorsal view (Fig. 696); vertex behind stemmaticum coriaceous-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely punctate-coriaceous, with satin sheen; precoxal area of mesopleuron partly remotely punctate as its surroundings; medio-longitudinal carina of metanotum distinct posteriorly; scutellum punctate; propodeum convex and coarsely rugose, medio-longitudinal carina absent posteriorly, and without protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 688); 1-CU1 slightly oblique, 0.3 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell long (Fig. 688); cu-a slightly inclivous, straight but posteriorly slightly curved; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 densely setose. Hind wing: marginal cell linearly widened, its apical width 1.7 × width at level of hamuli (Fig. 689); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 26:25; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized brownish pecten (Fig. 698); hind coxa densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.8 and 6.4 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex and basally rather steep, 0.9 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and longitudinally rugose; maximum width of 2nd tergite 1.6 × its median length; medio-basal area of 2nd tergite medium-sized triangular and rather short (Fig. 692); 2nd suture deep and finely crenulate; basal half of 3rd tergite rugulose, remainder of metasoma superficially microsculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 687). Colour. Black; mesoscutum largely, legs, and 1st–3rd metasomal tergites (but 3rd tergite narrowly infuscate posteriorly) reddish brown; tegulae brownish yellow; ovipositor sheath, palpi, pterostigma and veins dark brown; telotarsi partly infuscate; wing membrane subhyaline. Variation. Legs usually largely reddish, but telotarsi, apices of hind femur and tibia frequently dark brown and sometimes most of hind tibia and apical half of hind femur black; clypeus blunt to rather acute ventrally; depression near posterior ocelli smooth or finely crenulate; mesoscutum of ♀ usually partly reddish brown, but sometimes largely or entirely black; ventral third of mesopleuron regularly and finely punctate.

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Antennal segments: ♀ 57(2), 59(1); ♂ 57(1), 59(1), 60(5), 61(1), 62(1). Male is very similar with mesoscutum black (rarely partly reddish) and at least 2nd tergite orange brown (sometimes with pair of dark brown patches), apical tergites type 1–2 with fringe rather strong when visible, and also often evident on tergites following the third. Distribution. Finland, Norway, Sweden. Aleiodes rugulosus (Nees, 1811) Figs 701–727 Bracon rugulosus Nees, 1811: 32; Papp, 1985a: 162 (neotype designation). Rogas rugulosus; Shenefelt, 1975: 1247–1248. Rogas (Rogas) rugulosus; Tobias, 1976: 84, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) rugulosus; Papp, 1987b: 36, 1991a: 79; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) rugulosus; Belokobylskij et al., 2003: 398. Aleiodes rugulosus; Bergamasco et al., 1995: 5; Zaldivar-Riverón et al., 2004: 234; Papp, 2005: 177. Rhogas rugulosus var. pictus Kokujev, 1898: 296; Shenefelt, 1975: 1247–1248 (not Herrich-Schäffer, 1838). Type material. Neotype, ♀ (KBIN), “A. rugulosus”, “dét. C. Wesmael”, “Coll. Wesmael”, “Belgique, Bruxelles”/ teste Papp J., 1983”, “Neotypus, Bracon rugulosus Nees, 1812 [sic!], ♀, Papp 1983”, “Aleiodes rugulosus Ns. ♀, det. Papp J., 1983”. Additional material. Albania, Austria, Belgium, Bulgaria, Czech Republic, British Isles (England: V.C.s 1, 3, 4, 11, 17, 25, 27, 29, 69; Wales: V.C.s 41, 48; Scotland: V.C.s 78, 81, 88, 98; Ireland: V.C. H30), Finland, France, Germany, Hungary, Moldova, Netherlands (DR: Borger; GE: ‘t Harde; Heerde; Otterlo, NB: Kampina; NH: Crailo; OV: Buurserzand), North Macedonia, Norway, Poland, Romania, Russia, Slovakia, Spain, Sweden. Specimens in ALC, BMNH, BZL, HHC, IKC, MMUM, MRC, MSC, MSNV, MTMA, NMS, NRS, OUM, RMNH, SDEI, UWIM, ZSSM. Molecular data. MRS191 (Hungary), MRS217 (UK), MRS398 (France), MRS884 (Poland). Biology. Collected in (May)July–August(September), usually in open habitats such as mosses, heaths, herb-rich grasslands and fens. Partly plurivoltine, at least in the southern part of its range, but largely univoltine in the north (in a UK culture only one female out of 20 reared, and two males out of 69, emerged in the same year as mummy formation). Reared from Noctuidae, Acronictinae: Acronicta auricoma (Denis & Schiffermüller) (7 [1 BZL, 1 MSC, 1 ZMUO]; J. Voogd/Netherlands, M & J. Schwarz/Austria, Finland), Acronicta euphorbiae (Dennis & Schiffermüller)/cinerea (Hufnagel) (5 [2 IKC, 1 BZL, 1 FMNH]; M.J. Pellinen, D. & J. Steedan), Acronicta menyanthidis (11; R.P. Knill-Jones, W.A. Watson), Acronicta rumicis (Linnaeus) (5 [1 BMNH, 1 IKC, 1 MTMA]; R.J. Heckford, M.J. Pellinen, M.R. Shaw), Oxicestra geographica (Fabricius) (17 [12 BZL, 5 MTMA]; Hungary), Acronicta sp. on low

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Figures 701–703. Aleiodes rugulosus (Nees), ♀, U.K., culture 701 habitus lateral 702 ovipositor sheath lateral 703 mummy of Acronicta rumicis (Linnaeus).

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plants (3), Simyra albovenosa (Goeze) (14 [5 BMNH, 4 FMNH, 2 UMZC, 1 NRS]; M.R. Shaw); A. Lozan, Romania), Acronicta euphorbiae (Dennis & Schiffermüller) on Euphorbia sanguinea Hochst (5 [2 RMNH, 3 ALC]). These species all feed on low plants. The mummy is moderately arched, very strongly glued down (usually to a narrow twig or stem low in the vegetation), and persists through the winter. The pupation chamber, occupying approximately abdominal segments 3–7 of the host, is rather densely lined with silk which is laid down after the mummy has hardened suggesting that the larva within can turn easily. Rearing experiments, undertaken using stock originally reared from A. menyanthidis, suggested that this host and A. rumicis were equally suitable, but most experiments were not conducted in a way to provide clear data in this respect. The behaviour of the adult females towards these hosts indicated some adaptation to use of highly aggregated species (i.e., that lay large batches of eggs) as, firstly there was a habituation process whereby repeated contacts with hosts generally preceded oviposition, and secondly there was only weak displacement following oviposition (resulting in rather frequent super-parasitism). The antennae were used to locate hosts with wide sweeping motions, and usually the host curled up and was manipulated backwards against the hind tarsi before the ovipositor was inserted and the egg was laid. Generally, there was no pre-oviposition sting and post-oviposition association with the only slightly subdued host was minimal, but the oviposition process was variable and occasionally there was a brief jab, but no subsequent waiting period, before oviposition. Less enthusiasm for sub-active hosts, such as those oviposited into a few seconds or minutes earlier, provided a short-lived impediment to super-parasitism, although sometimes two (on one occasion four, confirmed by dissection) eggs were laid into a single host in separate consecutive bouts without the parasitoid really relinquishing the host. First instar hosts were less easy than 2nd or 3rd instars for the parasitoid to deal with, and although oviposition into 2nd instar hosts was somewhat more successful than into 3rd instars, occasionally successful oviposition into early 4th instar hosts occurred. Mean development times from oviposition to mummification in different instar hosts (A. rumicis) under the same ambient conditions (Reading, S. England, July) varied as might be expected given that mummification was always at essentially the same (penultimate instar) stage of the host’s larval life: for 1st (N = 23), 2nd (N = 40) and 3rd (N = 7) instars, 27.0, 25.5 and 20.7 days, respectively. There is no venom effect to influence successful host development. Opportunities to offer other hosts were limited but it was clear that, although oviposition into larvae of the closely related arboreal species Subacronicta megacephala (Dennis & Schiffermüller) was fairly readily obtained (N = 15), though slightly inhibited by the host’s adherence to its silken pad rather than curling up, the parasitoid was always encapsulated (as a 1st instar larva in observed cases) and no progeny resulted. No rearings of A. rugulosus from arboreal Acronictinae have been seen, although these conspicuous larvae are often collected and reared. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 711); OOL approx. equal to diameter of posterior ocellus and coarsely punctate (Fig. 712); vertex flattened behind ocelli; ventral margin of clypeus thick and

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Figures 704–716. Aleiodes rugulosus (Nees), ♀, U.K., culture 704 wings 705 mesosoma lateral 706  mesosoma dorsal 707 propodeum and 1st–3rd metasomal tergites dorsal 708 fore femur lateral 709 hind femur lateral 710 antenna 711 head anterior 712 head dorsal 713 head lateral 714 base of antenna 715 apex of antenna 716 outer hind tarsal claw.

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Figures 717, 718. Aleiodes rugulosus (Nees), dark form, ♀, U.K., Meathop Moss 717 habitus lateral 718 mummy of Acronicta menyanthidis (Esper).

not protruding in lateral view (Fig. 713); mesoscutal lobes coriaceous; mesopleuron (including precoxal sulcus area) nearly or completely smooth; propodeum with pair of crest-like protuberances laterally; vein 1-CU1 of fore wing much shorter than vein

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2-CU1; basal half of marginal cell of hind wing parallel-sided and subapically widened (Fig. 704); tarsal claws with large dark brown pecten up to apical tooth of claw (Fig. 716); hind spurs (dark) reddish brown; hind tibial spurs of ♂ acute apically (Fig. 722); head black; dorsal 0.4 of mesopleuron, mesosternum and scutellum black; metasoma entirely black (typical) or 1st and 2nd tergites orange or yellowish brown; apex of hind femur yellowish or reddish; basal half of hind tibia pale yellowish. Description. Redescribed ♀ (RMNH) from Netherlands (Buurserzand). Length of fore wing 6.3 mm, of body 7.7 mm. Head. Antennal segments of ♀ 65, length of antenna 1.3 × fore wing, its subapical segments rather slender (Fig. 716); frons largely smooth except few striae; OOL equal to diameter of posterior ocellus, coarsely punctate and shiny; vertex coarsely punctate with some rugae, shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 713); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 711); length of eye twice temple in dorsal view (Fig. 712); vertex behind stemmaticum flattened and punctate-rugose; clypeus between eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate with very finely granulate interspaces, rather matt; precoxal area of mesopleuron smooth, mesopleuron densely punctate posteriorly; scutellum coarsely punctate and rather flat; propodeum rather flattened medially, very coarsely reticulate-rugose, medio-longitudinal carina nearly complete, and with small crest-like protuberances laterally. Wings. Fore wing: r 0.6 × 3-SR (Fig. 704); 1-CU1 horizontal, 0.4 × 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell short (Fig. 704); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: basal 0.6 of marginal cell subparallel-sided and remainder linearly widened, its apical width 1.8 × width at level of hamuli (Fig. 704); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 37:28; 1r-m 0.8 × 1-M. Legs. Tarsal claws with conspicuous and robust blackish pecten (Fig. 716); hind coxa largely densely and coarsely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 3.9 and 5.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite moderately convex, 0.9 × longer than wide apically; 1st and 2nd tergites with coarse medio-longitudinal carina and very coarsely and irregularly longitudinally rugose; medio-basal area of 2nd tergite large and distinct (Fig. 707); 2nd suture deep, rather wide and coarsely crenulate; basal half of 3rd tergite punctate-rugose, remainder of metasoma finely punctate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically narrowed and rounded (Fig. 702). Colour. Black (including fore and middle telotarsi, apical half of hind tibia and hind tarsus); basal half of hind tibia pale yellowish; palpi (but basally somewhat infuscate) and remainder of legs reddish brown; tegulae yellowish brown; hind tibial spurs and pterostigma dark brown; veins brown; mesopleuron with broad dark reddish longitudinal band; wing membrane slightly infuscate. Variation. Micro-sculpture of lateral lobes of mesoscutum very finely granulate or absent and resulting in a largely smooth surface; maximum width of marginal cell

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Figure 719. Aleiodes rugulosus (Nees), Netherlands, Ede, mummy of Acronicta auricoma (Denis & Schiffermüller). Photograph: J. Voogd.

of hind wing 1.5–2.3 × its width near hamuli; body entirely black (both sexes) or 1st and 2nd metasomal tergites and mesosoma partly dark reddish (both sexes, but 2nd and 3rd tergites of males more or less darkened); hind tibial spurs dark brown or reddish brown. Antennal segments: ♀ 60(4), 61(7), 62(10), 63(18), 64(14), 65(2), 66(8), 67(4), 68(3), 69(1); ♂ 53(1), 56(2), 57(1), 58(7), 59(6), 60(15), 61(15), 62(19), 63(28), 64(12), 65(10), 66(4), 67(2), 68(1). Females have on average ca one more antennal segment than males. Male is similar to the dark female form, with acute hind tibial spurs (Fig. 722), 3rd tergite convex in lateral view (Fig. 720), with dense and long setosity and apical tergites type 1 and no fringe observed (Figs 721, 724). Aleiodes rugulosus is a very colour-variable species; the 1st and 2nd metasomal tergites are quite frequently mostly or entirely dark red or orange brown (Fig. 701) in populations in which entirely black females (Fig. 717) also often occur. The variation is not geographical, since most populations definitely have both forms. In rearing experiments, a strong genetic basis for this feature became evident: from a virgin female with completely black metasoma, all five males reared were black, and a cross between one of them and a (wild-reared) black female produced eleven female progeny, all black. In contrast, a lineage from a pairing between a weakly red male and a red female comprised three red males and eight red females, separate individuals of which produced a single red and a single black male as progeny. When sufficient material from single sites is available it is usual to see a clear predominance of one form or the other. Sometimes the lower part of mesopleuron is pale (often looking unpigmented there, but narrow-

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Figures 720–727. Aleiodes rugulosus (Nees), ♂, Hungary (Halastó) 720 habitus lateral 721 metasoma lateral 722 hind tibial spurs and basitarsus lateral 723 base of antenna 724 metasoma dorsal 725 fore femur lateral 726 head dorsal 727 hind femur lateral.

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ly), and in extreme reddish specimens the scutellum, much of the mesopleuron (but usually the mesosternum remains darkish), the metanotum dorsally, the metapleuron in part and much of the propodeum are also reddish. Distribution. Albania, Austria, Belgium, Bulgaria, Czech Republic, British Isles (England, Wales, Scotland, Ireland), Finland, France, Germany, Hungary, *Netherlands, *North Macedonia, *Moldova. Norway, Poland, *Romania, Russia, *Slovakia, *Spain, Sweden. Aleiodes schewyrewi (Kokujev, 1898) Figs 728–742 Rhogas (Rhogas) schewyrewi Kokujev, 1898: 304. Rhogas (Rhogas) schevyrevi [sic!]; Telenga, 1941: 186. Rogas schewyrewi; Shenefelt, 1975: 1248 Rogas (Rogas) schewyrewi; Papp, 1977b: 116. Aleiodes (Chelonorhogas) schewyrewi; Belokobylskij, 2000: 46. Rhogas (Rhogas) schewyrewi var. zaydamensis Kokujev, 1898: 305; Telenga, 1941: 186 [examined]. Rogas schewyrewi var. zaydamensis; Shenefelt, 1975: 1248–1249. Rogas (Aleiodes) schewyrewi var. zaydamensis; Papp, 1977b: 116, 117. Type material. Holotype of A. schewyrewi zaydamensis, ♀ (ZISP), “[Mongolia], Kerijsk Kr., Ruio Zaydam, Przewalski”, “1910a”, [illegible handwritten label], “Rh. Schewyrewi Kokw. var. zaydamensis Kokw., No.1910a”, “♀ Rhogas schewyrewi var. zaydamensis Kok., C. van Achterberg, 1992, holotype”. Holotype of A. s. schewyrewi not found, according to the original description with same label data and with a larger part of the body blackish. Additional material. 1 ♀ (BZL), “S. Russia, [Volgograd obl.], Elton Lake env., 20.v.2001, J. Miatleuski”; 1 ♂ (MTMA), “Mongolia, Gobi Altay aimak, Mongol els, 10 km SO von Somon Chechmort, 1600 m, Exp. Dr. Z. Kaszab, 1966”, “Nr. 684, 13.vii.1966”, “Rogas schewyrewi Kok., det. Papp J., 1977/ compared with ♀ det. Kokujev, Papp, 1983, 57”. Molecular data. None. Biology. Unknown. A female collected in May and a male in July may suggest that it is plurivoltine (or, less probably, that the female overwinters as an adult). Diagnosis. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 737); OOL of ♀ approx. as long as diameter of posterior ocellus and densely rugose (Fig. 738); head in anterior view rather robust (Fig. 737); clypeus distinctly protruding anteriorly in lateral view, thick apically and with long setae on medium-sized anterior part (Fig. 739); lobes of mesoscutum largely superficially punctate, interspaces finely granulate or smooth and with satin sheen; precoxal area densely rugose, but posterior third only finely punctate; vein 1-CU1 of fore wing 0.3 × vein

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Figures 728, 729. Aleiodes schewyrewi (Kokujev), ♀, holotype var. zaydamensis (Kokujev) 728 habitus lateral 729 ovipositor sheath lateral.

2-CU1 and 0.5 × vein m-cu (Fig. 730); hind tarsal claws long and slender, nearly straight and only brownish bristly setose (Fig. 740); tarsal segments (except telotarsus) with long apical spiny bristles (Fig. 728); basal half of hind tibia pale yellowish, contrasting with dark brown colour of basal half of hind femur. Description. Holotype, ♀, length of fore wing 5.8 mm, of body 7.5 mm. Head. Antennal segments of ♀ 48, length of antenna 1.1 × fore wing, its subapical segments moderately slender (Fig. 742); frons largely with fine curved rugae; OOL equal to diameter of posterior ocellus, and densely rugose; vertex superficially rugosepunctate, rather shiny; clypeus convex and densely punctate; ventral margin of clypeus thick and protruding forwards (Fig. 740); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 737); length of eye 1.3 × temple in dorsal view (Fig. 738);

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Figures 730–742. Aleiodes schewyrewi (Kokujev), ♀, holotype var. zaydamensis (Kokujev) 730 fore wing 731 hind wing 732 mesosoma lateral 733 mesosoma dorsal 734 metasoma dorsal 735 fore femur lateral 736 hind femur lateral 737 head anterior 738 head dorsal 739 head lateral 740 outer hind tarsal claw 741 base of antenna 742 apex of antenna.

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vertex behind stemmaticum finely rugose-punctate and with long setae; clypeus largely above lower level of eyes; length of malar space 0.3 × height of eye in lateral view. Mesosoma. Mesoscutal lobes largely superficially punctate, interspaces finely granulate and with satin sheen; precoxal area of mesopleuron largely smooth medially, densely punctate anteriorly and posteriorly densely rugose, but posterior 0.3 only finely punctate; metapleuron spaced coarsely punctate; metanotum with fine and nearly complete median carina; scutellum sparsely punctate, shiny; propodeum rather convex and rather coarsely rugose, medio-longitudinal carina on anterior 0.4 of propodeum. Wings. Fore wing: r curved and 0.4 × 3-SR (Fig. 730); 1-CU1 horizontal and slightly widened, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 730); cu-a slightly inclivous, straight; 1-M slightly curved; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: basal third of marginal cell subparallel-sided and remainder linearly widened; 2-SC+R short and longitudinal; m-cu slightly indicated; M+CU:1-M = 10:7; 1r-m 0.6 × 1-M. Legs. Tarsal claws long and slender, nearly straight and only brownish bristly setose (Fig. 740); tarsal segments (except telotarsus) with long apical spiny bristles (Figs 728, 740); hind coxa largely punctate, but dorsally punctate-rugose; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex and strongly widened posteriorly, 0.9 × longer than wide apically; 1st and 2nd tergites with weak medio-longitudinal carina (absent posteriorly) and finely longitudinally rugose, but 2nd tergite smooth medioposteriorly; medio-basal area of 2nd tergite wide triangular and rather distinct (Fig. 734); 2nd suture deep and narrow; basal half of 3rd tergite aciculate, remainder of metasoma smooth and shiny; 2nd and 3rd tergites with sharp lateral crease; ovipositor sheath moderately widened, with medium-sized setae and apically truncate (Fig. 729). Colour. Yellowish brown (including basal half of antenna); apical half of antenna, frons largely, stemmaticum, occiput dorso-laterally, pronotal side medially, axilla, mesopleuron (except antero-dorsally), mesosternum, metapleuron, propodeum, hind femur and apical third of hind tibia, 5th–7th tergites, last two posterior sternites, ovipositor sheath and pterostigma dark brown or blackish brown; veins brown; wing membrane subhyaline. Variation. Length of 4th hind tarsal segment 1.8–2.0 × longer than wide; malar space and temple ventrally largely dark brown or yellowish brown. Antennal segments: ♀ 45(1); ♂ 54(1); according to original description ♀ type has 58 segments. Male has clypeus yellowish and contrasting with black face, apical tergites type 1, and no fringe observed. Distribution. *Iran, Mongolia, *Russia (European part). Aleiodes schirjajewi (Kokujev, 1898) Figs 743–757 Rhogas reticulator var. schirjajewi Kokujev, 1898: 299 [examined].

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Rogas schirjajewi; Shenefelt, 1975: 1249. Rogas (Rogas) schirjaevi [sic!]; Tobias, 1976: 85. Rogas (Rogas) schirjaewi [sic!]; Tobias, 1986: 80 (transl.: 132). Aleiodes (Neorhogas) schirjajewi; Papp, 1991a: 71, 2002: 562. Aleiodes (Chelonorhogas) schirjajewi; Samartsev & Belokobylskij, 2013: 766. Aleiodes schirjajewi; Shaw et al., 1998: 63; Papp, 2005: 177. Type material. Holotype, ♂ (ZISP), “[Kazakhstan], Kemropavl., Akmolin, 908a”, “K. Kokujeva”, “908a, Rh. reticulator Nees v. schirjajewi Kokw.”, “Holotypus”. Additional material. Bulgaria, Hungary, Italy, Moldova, Russia, Serbia, Ukraine [Dagestan, Kazakhstan]. Specimens in BZL, BMNH, MRC, MSC, MTMA, NMS, RMNH, SDEI, ZISP. Molecular data. None. Biology. Unknown. Specimens collected throughout April–September, presumably plurivoltine. We have not seen reared material and it is unclear how the winter is passed. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 751); length of antenna of ♀ 1.1–1.4 × fore wing; ventral margin of clypeus thick and obtuse apically and clypeus not protruding in lateral view (Fig. 753); vertex and frons with strong striae or rugae; mesoscutum, metapleuron and scutellum normally shiny and without dense granulation, at most with some superficial microsculpture; precoxal area of mesopleuron smooth; vein 2-CU1 of fore wing approx. as long as vein 1-CU1 or shorter (Fig. 744); vein M+CU of hind wing distinctly longer than vein 1-M (Fig. 745); hind tarsal claws with medium-sized dark brown pecten (Fig. 756); head black; mesoscutum and scutellum orange brown; fore and middle femora distinctly black or dark brown apically; basal half of hind tibia dark brown; anterior half of mesosoma, 1st and 2nd metasomal tergites yellowish or orange brown; at least basal half of 4th–6th tergites of ♂ with long and dense setosity. Description. Holotype, ♂, length of fore wing 5.0 mm, of body 5.8 mm. Head. Antennal segments of ♂ 50, length of antenna 1.3 × fore wing, its subapical segments rather robust; 4th segment of maxillary palp slender and cylindrical; frons with rather coarse curved rugae and interspaces smooth; OOL 1.1 × diameter of posterior ocellus, coarsely rugose and shiny; coarsely transversely rugose and shiny; clypeus punctate-rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 753); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 751); length of eye 3.2 × temple in dorsal view (Fig. 752); vertex behind stemmaticum coarsely rugose; clypeus below lower level of eyes; occipital carina complete; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate, shiny, interspaces micro-sculptured; precoxal area of mesopleuron smooth medially except for some crenulations and punctures, its surroundings smooth; scutellum moderately punctate; propodeum rather convex and coarsely reticulate, coarse medio-longitudinal carina present anteriorly, carinae not protruding laterally.

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Figure 743. Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, habitus lateral.

Wings. Fore wing: r 0.7 × 3-SR (Fig. 744); 1-CU1 horizontal, as long as 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell short (Fig. 744); cu-a vertical, largely straight; 1-M nearly straight posteriorly; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 745); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 15:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws with medium-sized dark brown pecten (Fig. 756); hind coxa largely densely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 4.0 and 7.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite moderately flattened, 0.9 × longer than wide apically; 1st and nd 2 tergites with medio-longitudinal carina and densely vermiculate-rugose; medio-basal area of 2nd tergite wide triangular and distinct (Fig. 748); 2nd suture deep and moderately crenulate; 3rd tergite finely striate basally, remainder of metasoma largely smooth; 4th without sharp lateral crease; basal half of 4th–6th tergites of ♂ with long and dense setosity. Colour. Orange brownish; basal half of antenna, palpi largely and parastigma (except base) yellowish brown; head, mesosternum (except anteriorly), mesopleuron (except anteriorly and antero-dorsally), apical 0.4 of hind femur, 3rd tergite (except antero-laterally) and subsequent tergites black; scapus, pedicellus basally, apical half of antenna, apex of fore and middle femora, apex of middle and hind tibiae, hind basi-

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Figures 744–757. Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, but 756 ♀, Ukraine 744 fore wing 745 hind wing 746 mesosoma lateral 747 mesosoma dorsal 748 metasoma dorsal 749 fore femur lateral 750 hind femur lateral 751 head anterior 752 head dorsal 753 head lateral 754 base of antenna 755 apex of antenna 756 outer hind tarsal claw 757 antenna.

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tarsus, 2nd hind tarsal segment apically, telotarsi, pterostigma, parastigma basally and veins dark brown; wing membrane subhyaline. Variation. Vein 1-CU1 of fore wing 1.0–2.3 × vein 2-CU1; maximum width of marginal cell of hind wing 1.6–2.6 × its width near hamuli (Fig. 745). Antennal segments: ♀ 46(1), 48(3), 49(1), 50(2), 51(1); ♂ 47(1), 48(2), 49(2), 50(1), 51(1). The sexes have comparable numbers of antennal segments. Apical tergites of ♂ type 4, dense, making the tergites look concave and fringe not observed. Female is very similar to the redescribed male; ovipositor sheath wide, with long setae and apically truncate. Distribution. *Bulgaria, Hungary, *Italy, Kazakhstan, Moldova, Russia (including Dagestan and Far East), Serbia, Ukraine. Aleiodes sibiricus (Kokujev, 1903) Figs 758–776 Rhogas sibiricus Kokujev, 1903: 286 [examined]. Rogas sibiricus; Shenefelt, 1975: 1250. Rogas (Rogas) sibiricus; Tobias, 1976: 83, 84, 1986: 76, 78 (transl.: 124, 128; lectotype designation). Aleiodes (Neorhogas) sibiricus; Papp, 1985a: 150, 153, 162, 1991a: 92; Belokobylskij, 1996: 15. Aleiodes sibiricus; Papp, 2005: 177. Rhogas hungaricus Szépligeti, 1906: 616; Papp, 1985a: 150, 153, 162 (as synonym of A. sibiricus; lectotype designation), 2005: 177 (id.); 2004: 216 (id.) [examined]. Rogas hungaricus; Shenefelt, 1975: 133. Rhogas reinhardi Fahringer, 1931: 221 (description in key only), 1932: 275 (full description; as R. rheinhardi); Papp, 1985a: 153, 162 (as synonym of A. sibiricus; holotype examined), 2005: 177 (id.). Type material. Paralectotype of A. sibiricus, ♀, (BMNH), “[Russia], Irkutsk, v., I. Jakovlev”, “K. Kokujeva”, “Paratypus Rogas sibiricus Kokujev”, “Rec. in exchange [from] Academy of Science, Leningrad, BM.1963.211”, “Ant. 69”. Lectotype of A. hungaricus, ♀, (MTMA), “[Hungary], Budapest, Szépligeti”, “Lectotypus Rhogas hungaricus Szépl. 1906, ♀, Papp, 1966”, “Hym. Typ. No. 401, Mus. Budapest”, “Aleiodes sibiricus Kok., ♀, det. Papp J., 1983/compared with ♀ paralectotype”. Additional material. Albania, Austria, Bulgaria, France, Germany, Greece, Hungary, Italy, North Macedonia, Sweden, Turkey. Specimens in BMNH, BZL, MTMA, NMS, RMNH, ZSSM. Molecular data. MRS310 (Sweden), MRS313 (Sweden), MRS805 (France). Biology. Collected in April and May, and presumably univoltine, but 2 ♀ from Sweden: Ångermanland, Lillavammasjon were collected apparently in July in window traps set on the trunks of Betula and Picea. We have examined four males collected in April which suggests spring emergence from the mummy rather than overwintering as

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Figures 758, 759. Aleiodes sibiricus (Kokujev), ♀, lectotype 758 habitus lateral 759 head anterior. Photographs: K. Samartsev.

an adult. This is corroborated by the data with the single reared specimen examined (MTMA), from the noctuid Noctua comes Hübner collected 7.iv.1961 and emerging on 3.iv.1962 (Germany; [R.] Hinz). The rearer was widely experienced with caterpillars, and the host determination is unlikely to be wrong (the other caterpillar species with which it might conceivably be confused all have similar biology and phenology in any case). This host initiates its overwintering as a small larva, feeding in mild weather through the winter and normally being well-grown by April, by then in its penultimate or final instar. The rearing is of great interest because it shows that A. sibiricus, like A. fortipes (q. v.), not only parasitises a host that has overwintered as a larva, but also must habitually attack late instar hosts. The reared specimen is accompanied by a stout mummy, large but not unduly so for the size of the adult that emerged, lacking its anterior portion to leave a partitioned chamber comprising abdominal segments 4 onwards, which is well lined with silk and would presumably normally form in the soil (Fig. 762). Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Figs 759, 770); antenna of ♀ with 65–72 segments and 5th–10th segments wider than long; anterior part of clypeus short and transverse, its height 0.2–0.3 × height of hypoclypeal depression (Fig. 770); ventral margin of clypeus rather thin and slightly protruding in lateral view (Fig. 772); mesoscutal lobes densely punctate and interspaces smooth; precoxal area (rather) coarsely vermiculate-rugose medially; length of vein r of fore wing 0.3–0.5 × vein 3-SR (Fig. 763); vein 1-CU1 horizontal and 0.2–0.3 × vein 2-CU1; hind tarsus and claws slender and claws with inconspicuous brownish

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Figures 760–762. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves, but 762 Germany, Freiburg 760 habitus lateral 761 ovipositor sheath lateral 762 mummy of Noctua comes Hübner.

teeth (Fig. 775); 4th and 5th metasomal tergites more or less yellowish to reddish brown; head, mesoscutum, scutellum, mesopleuron and apex of metasoma black. Description. Paralectotype of A. sibiricus, ♀, length of fore wing 9.1 mm, of body 10.0 mm. Head. Antennal segments of ♀ 69, antenna as long as fore wing, its subapical segments medium-sized; frons smooth; OOL 1.1 × diameter of posterior ocellus, and finely coriaceous-rugulose; vertex rugulose and rather dull; clypeus coriaceous and

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strongly transverse (4–6 × wider than high; Figs 759, 770); ventral margin of clypeus rather thin and slightly protruding forwards (Fig. 772); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 770); length of eye 1.1 × temple in dorsal view (Fig. 771); vertex behind stemmaticum rugulose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate and interspaces smooth, rather matt; precoxal area of mesopleuron rugose medially and anteriorly, its surroundings moderately punctate; scutellum sparsely punctate and no lateral carina; propodeum rather convex and densely and finely rugose, medio-longitudinal carina complete and no protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 763); 1-CU1 horizontal, 0.2 × 2-CU1; r-m unsclerotized, 0.7 × 3-SR; 2nd submarginal cell rather short (Fig. 763); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: basal half of marginal cell slightly widened, but apical half distinctly linearly widened, its apical width 2.5 × width at level of hamuli (Fig. 764); 2-SC+R subquadrate; m-cu slightly indicated; M+CU:1-M = 51:38; 1r-m 0.7 × 1-M. Legs. Tarsal claws with four inconspicuous brownish pecten-teeth (Fig. 775); hind coxa punctulate; hind trochantellus robust; length of hind femur and basitarsus 4.1 and 6.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather flattened, 0.8 × longer than wide apically; 1st and nd 2 tergites with weak medio-longitudinal carina and densely finely rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite wide and short (Fig. 767); 2nd suture deep, rather wide medially and finely crenulate; basal half of 3rd tergite finely rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 761). Colour. Black; palpi and tegulae pale yellowish; legs (except black coxae; apex of hind femur dorsally, inner side of hind tibia apically (not outer side!) and telotarsi infuscated), apex of first tergite, 2nd–5th tergites and metasoma ventrally, yellowish brown; ovipositor sheath largely, pterostigma and most veins dark brown; vein 1-R1 of fore wing yellowish brown; wing membrane subhyaline. Variation. Face, clypeus, mesoscutum, propleuron, upper part of mesopleuron, and first tergite partly, or rarely entirely, reddish brown. Usually in males and rarely in females mesoscutum wholly black; vein r of fore wing 0.3–0.5 × vein 3-SR; clypeus flattened and subparallel-sided or convex and ventrally concave; pterostigma medially dark brown or yellowish brown. Antennal segments ♀: 65(4), 66(8), 67(6), 68(4), 69(2), 70(5), 71(2), 72(2); ♂ 66(1), 71(1). Male apical tergites of type 1 and fringe not observed. Distribution. *Albania, *Austria, *Bulgaria, *France, Germany, *Greece, Hungary, Italy (main), *North Macedonia, Russia (Siberia), *Sweden, *Turkey. Notes. The holotype of A. reinhardi (Fahringer, 1931) from Bolzano (N Italy) was examined by Papp (1985a) and directly compared with the lectotype of A. hungaricus. Unfortunately, the holotype could not be located in NHMW, but there is no obvi-

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Figures 763–776. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves 763 fore wing 764 hind wing 765 mesosoma lateral 766 mesosoma dorsal 767 metasoma dorsal 768 fore femur lateral 769 hind femur lateral 770 head anterior 771 head dorsal 772 head lateral 773 base of antenna 774 apex of antenna 775 outer hind tarsal claw 776 antenna.

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ous reason not to follow the synonymy with A. sibiricus (Kokujev) proposed by Papp (1985a). Aleiodes agilis (Telenga, 1941) from China, Iran, and Caucasus is very similar to A. sibiricus, but A. agilis has antenna of ♀ with ca 48 segments (69–72 segments in A. sibiricus), 2nd tergite narrowly smooth posteriorly (finely sculptured), pronotum largely yellow (black), clypeus not protruding in lateral view (somewhat protruding) and is often smaller (body length 7–8 mm vs 7–11 mm). Aleiodes turcicus van Achterberg & Shaw, sp. nov. http://zoobank.org/885C0189-5A7B-4D35-B7D3-82F1094633AA Figs 777–791 Type material. Holotype, ♀ (NMS), “Turkey: Sivas, v.2001, D.L.J. Quicke”, “MRS Aleiodes DNA 126 [one middle leg]”. Paratype: 1 ♂ (RMNH), “Turkey, Hakkâri, [20 km S Siirt, 500 m, 23.vi.1985], C.J. Zwakhals”. Molecular data. MRS126 (Turkey). Biology. Unknown. The material examined was collected in the period May–June. It is not clear how many generations occur, or how the winter is passed. Diagnosis. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 786); OOL approx. 2.3 × diameter of posterior ocellus and sparsely punctate (Fig. 787); ventral margin of clypeus obtuse and not protruding in lateral view (Fig. 788); length of antenna of ♀ 1.1–1.2 × fore wing; lateral lobes of mesoscutum largely smooth; precoxal area coarsely vermiculate-rugose medially; vein 1-CU1 of fore wing approx. 0.4 × as long as vein 2-CU1; hind tarsal claws yellowish or brownish setose (Fig. 791); head and part of mesosoma black; palpi, pterostigma and apical 0.2–0.3 of hind tibia of ♀ blackish; wing membrane distinctly infuscate. Description. Holotype, ♀, length of fore wing 5.3 mm, of body 7.9 mm. Head. Antennal segments of ♀ 47, length of antenna 1.15 × fore wing, length of 4th segment 1.1 × its width, and its subapical segments 1.2 × as long as wide (Figs 789, 790); frons with regular curved rugae, shiny, and rugose behind antennal sockets; OOL 2.3 × diameter of posterior ocellus, and area mostly finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate laterally, densely punctate and with transverse rugae medially, shiny; clypeus medium-sized, coarsely and densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 788); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 786); length of eye 1.3 × temple in dorsal view (Fig. 787); vertex behind stemmaticum sparsely punctate; clypeus near lower level of eyes; length of malar space 0.5 × length of eye in lateral view; eyes medium-sized, elliptical (Fig. 788). Mesosoma. Mesoscutal lobes smooth between rather remote punctures, strongly shiny, more densely punctate on middle lobe; notauli distinct but shallow, especially posterior half; mesoscutum short setose, widely and strongly rugose medio-posteriorly; scutellum slightly convex, remotely punctate and evenly rounded laterally, no carina; prepectal carina strong, reaching anterior border; precoxal area coarsely vermiculate

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Figures 777, 778. Aleiodes turcicus sp. nov., ♀, holotype 777 habitus lateral 778 ovipositor sheath lateral.

rugose anteriorly and medially, posteriorly absent; mesopleuron above precoxal area (except speculum) remotely punctate, shiny, and antero-dorsally coarsely vermiculaterugose; metapleuron densely rugose, but dorsally punctate, interspaces approx. equal to diameter of punctures; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete, but irregular, no tubercles.

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Figures 779–791. Aleiodes turcicus sp. nov., ♀, holotype 779 fore wing 780 hind wing 781 mesosoma lateral 782 mesosoma dorsal 783 metasoma dorsal 784 fore femur lateral 785 hind femur lateral 786 head anterior 787 head dorsal 788 head lateral 789 base of antenna 790 apex of antenna 791 outer hind tarsal claw.

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Wings. Fore wing: r 0.35 × 3-SR (Fig. 779); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 779); cu-a slightly oblique, approx. parallel with CU1b, straight; 1-M rather curved posteriorly. Hind wing: marginal cell gradually and evenly widened, its apical width 1.9 × width at level of hamuli (Fig. 780); 2-SC+R subquadrate; m-cu distinct, shorter than cu-a. Legs. Tarsal claws subpectinate, with four brown medium-sized pectinal bristles and some finer ones basally (Fig. 791); hind coxa moderately coarsely punctate, with several long oblique rugae, shiny; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 4.9 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus; hind tibia slender (Fig. 777). Metasoma. First tergite rather flattened; 1st and 2nd tergites coarsely and densely rugose, robust, with distinct median carina; medio-basal area of 2nd tergite wide and short (Fig. 783); 2nd suture deep medially and shallow laterally; basal 0.4 of 3rd tergite finely striate, remainder of metasoma largely smooth, strongly shiny, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath rather wide, with long setae and apically rounded (Fig. 778). Colour. Black; palpi, base of middle coxa, apical 0.2 (dorsally)–0.3 (inner side) of hind tibia, apex of hind femur and telotarsi blackish; remainder of tarsi more or less darkened and base of hind tibia dark brown; basal seven segments of antenna (remainder more or less dark brown), 1st and 2nd tergites and antero-lateral corners of 3rd tergite, and remainder of legs orange brown; humeral plate pale yellowish but partly darkened; pterostigma blackish; veins dark brown; wing membrane blackish infuscate. Variation. Apical metasomal tergites of ♂ type 2; inner hind tibial spur 0.50 × hind basitarsus; mesopleuron, metapleuron and propodeum may be largely yellowish. Distribution. Turkey. Aleiodes unipunctator (Thunberg, 1822) Figs 792–812 Ichneumon unipunctator Thunberg, 1822: 267 [examined]. Rogas unipunctator; Shenefelt, 1975: 1254–1255; Zaykov, 1980c: 229; Jakimavicius, 1974: 96. Rogas (Rogas) unipunctator; Tobias, 1976: 84, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) unipunctator; Papp, 1985a: 151, 163, 1991a: 86, 1996: 456; Belokobylskij, 1996: 18; Papp & Rezbanyai-Reser, 1996: 71, 73, 95, 96; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) unipunctator; Belokobylskij, 2000: 44, 2003: 399; Chen & He, 1997: 43; He et al., 2000: 667; Rastegar et al., 2012: 3; Farahani et al., 2015: 229, 244. Aleiodes unipunctator; Čapek & Lukás, 1989: 31; Bergamasco et al., 1995: 6; O’Connor et al.: 1999: 92; Fortier & Shaw, 1999: 230; Belokobylskij & Taeger, 2001: 115;

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Marsh & Shaw, 2001: 303; Belokobylskij et al., 2003: 399; Zaldivar-Riverón et al., 2004: 234; Papp, 2005: 177; Lozan et al. 2010: 17. Ichneumon ductor Thunberg, 1822: 269; Papp, 1985a: 157 (not auctt.) [examined]. Syn. nov. Aleiodes irregularis Wesmael, 1838: 101; Shenefelt, 1975: 1255 (as synonym of A. unipunctator); Papp, 1985a: 163 (id.) [examined]. Rhogas unipunctator ab. nigrescens Hellén, 1927: 23; Shenefelt, 1975: 1255 (excluded name). Type material. Holotype of A. unipunctator, ♂ (ZMUU) with holotype label by CvA. Holotype of A. ductor, ♂ (ZMUU), “α”, “Rhogas ductor Thbg”, “Uppsala Univ. Zool Mus., Thunbergsaml. Nr. 25332, Ichneumon ductor Sv. Type”. Holotype of A. irregularis, ♂ (KBIN), “[Belgium], 11 Juin, Brig.”, “A. irregularis ♂ mihi 5”, “dét. C. Wesmael”, “Belgique, Bruxelles/teste Papp J., 1983”, “Holotype”, “Aleiodes irregularis Wesm., 1838, ♂, Papp, 1983”, “Aleiodes unipunctator Thb. ♂, det. Papp J., 1984”. Additional material. Austria, Belgium, British Isles (England: V.C.s 1, 3, 5, 6, 8, 15, 16, 17, 18, 20, 21, 23, 25, 26, 27, 28, 29, 32, 33, 40, 53, 55, 58, 59, 63, 64, 65, 66, 69; Wales: V.C.s 35, 52 ; Scotland: V.C.s 72, 75, 76, 77, 79, 80, 82, 83, 85, 86, 87, 88, 89, 90, 91, 95, 96, 97, 98, 99, 100, 101, 103, 105, 108, 109, 110, 111, 112; Ireland: V.C.s H5, H19, H20, H21, H22, H28, H30), Bulgaria, Czech Republic, Denmark, Finland, Germany, Greece (mainland, Corfu), Hungary, Italy, Montenegro, Netherlands (DR: Wijster; Borger; GE: Nunspeet; Kemperberg, NB: Bergen op Zoom, ZH: Arkel; Melissant; Oostvoorne; ZE: Oostkapelle), Norway, Romania, Russia, Sweden, [Kazakhstan, Tadzhikistan, W. Caucasus]. Specimens in AAC, ALC, BMNH, BZL, CNC, FMNH, HHC, HSC, IKC, OUM, MMUM, MRC, MSC, MSNV, MTMA, NMI, NMS, RMNH, SDEI, UMZC, UNS, UWIM, ZMUU, ZSSM. Molecular data. MRS211 (UK), MRS221 (Germany), MRS354 (UK), MRS893 (UK). Biology. Univoltine, flying from May to August. Reared from the noctuids Apa­ mea unanimis (Hübner) (23 [1 BMNH, 1 FMNH, 1 NRS, 1 MTMA]; M.R. Shaw), Apamea crenata (Hufnagel) (2:1, BMNH; G.T. Lyle), Apamea ?sordens (Hufnagel) (3:1; K.P. Bland), and from mummies compatible with Apamea spp. (8). These closely related hosts all live in the shoots or leaf sheaths of Poaceae. The large number reared from A. unanimis may be at least partly the result of a sampling bias, as the larvae of that species are so readily detected and collected when feeding on Phalaris. The winter is passed in the relatively slender brown mummy, which is nearly cylindrical and only weakly keeled laterally (Fig. 794). It probably usually forms at or below soil level and is rather weakly contracted at the head end (which is bent sideways in a high proportion of cases), with the thinly silken cocoon occupying abdominal segments (2–)3–8. This species is widespread and often abundant, especially in rank or damp grassland habitats, in the northern part of its range. Diagnosis. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 802); OOL of ♀ approx. 0.8 × as long as diameter of posterior

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Figures 792–794. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 792 habitus lateral 793 ovipositor sheath lateral 794 mummy of Apamea unanimis Hübner.

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ocellus and mainly granulate mixed with some punctures (Fig. 803); ventral margin of clypeus thick, not protruding in lateral view (Fig. 804); mesoscutal lobes finely granulate-punctulate and matt; precoxal area comparatively narrow and moderately rugose medially; marginal cell of fore wing of ♀ ending rather close to wing apex (Fig. 795); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 795); hind tarsal claws rather robust and only brownish setose (Fig. 807); 1st tergite rather slender basally (Fig. 798); whole 4th and part of 3rd metasomal tergite smooth and very glossy (Fig. 798); labial palp yellowish brown or brown; basal half of hind tibia pale yellowish or ivory, at least inner side contrasting with reddish or dark brown colour of basal half of hind femur (usually less pronounced in ♂), and its apex dark brown or black; 4th and 5th tergites black; wings rather slender and their membrane subhyaline. Description. Redescribed ♀ (RMNH) from Sweden (Storbacken). Length of fore wing 5.8 mm, of body 5.7 mm. Head. Antennal segments of ♀ 50, length of antenna 1.2 × fore wing, its subapical segments rather robust; frons largely smooth anteriorly (except some fine sculpture) and densely rugulose posteriorly; OOL 0.8 × diameter of posterior ocellus, granulate with some punctures and matt; vertex granulate with some rugulosity and rather dull; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 804); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 802); length of eye 1.7 × temple in dorsal view (Fig. 803); vertex behind stemmaticum rugulose-granulate; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view. Mesosoma. Mesoscutal lobes moderately punctate and interspaces distinctly granulate, with satin sheen; precoxal area of mesopleuron moderately rugose medially, sparsely punctulate posteriorly as surroundings of precoxal area; scutellum rather sparsely punctate, but medio-posteriorly rugulose, shiny; propodeum rather convex and moderately rugose, medio-longitudinal carina complete, and no protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 795); 1-CU1 narrow and horizontal, 0.4 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell medium-sized and 1st subdiscal cell slender (Fig. 795); cu-a nearly vertical, straight; 1-M slightly curved posteriorly; 1-SR narrow posteriorly and widened anteriorly; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: basal half of marginal cell slightly wider and its apical half distinctly gradually widened, its apical width 2.2 × width at level of hamuli (Fig. 795); 2-SC+R short longitudinal; m-cu indistinct; M+CU:1-M = 4:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws yellowish bristly setose, without distinct pecten (Fig. 807); hind coxa largely distinctly punctate; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 6.3 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 1.2 × as long as wide apically; 1st and 2nd tergites with distinct medio-longitudinal carina and longitudinally striate; medio-basal area of 2nd tergite wide triangular and short (Fig. 798); 2nd suture rather deep and finely crenulate; 3rd tergite nearly entirely smooth and strongly shiny, as remainder of metasoma; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Fig. 793).

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Figures 795–807. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 795 wings 796 mesosoma lateral 797 mesosoma dorsal 798 metasoma dorsal 799 fore femur lateral 800 hind femur lateral 801 antenna 802 head anterior 803 head dorsal 804 head lateral 805 base of antenna 806 apex of antenna 807 inner hind tarsal claw lateral.

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Figures 808–812. Aleiodes unipunctator (Thunberg), ♂, Scotland, Edinburgh, but 808 Sweden, SäröHamra 808 habitus lateral 809 apex of metasoma dorsal 810 apex of metasoma lateral 811 head dorsal 812 base of antenna.

Colour. Black; pronotum dorso-posteriorly, telotarsi largely, hind tarsus, apical half of hind tibia, pterostigma (except paler extreme base), most veins and apical fifth of 2nd tergite dark brown; palpi brown; tegulae and basal half of hind tibia pale yellowish; apical third of 1st tergite and 2nd tergite (except apically) and remainder of legs, orange brown; wing membrane subhyaline. Variation. Propodeum and pronotum sometimes weakly marked with orange. One male seen with vein r-m of fore wing absent. Antennal segments: ♀ 47(1), 48(7),

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49(12), 50(21), 51(26), 52(40), 53(40), 54(15), 55(5), 56(3), 57(1); ♂ 47(1), 48(1), 49(1), 50(8), 51(7), 52(22), 53(27), 54(34), 55(35), 56(21), 57(6), 58(8), 59(1). On average males have ca one to two more antennal segments than females. Male is very similar with apical tergites type 2, setae rather sparse and fringe short (Figs 809, 810). Melanistic females and males occur, metasoma entirely blackish or with only apex of 1st tergite narrowly and 2nd tergite medially and antero-laterally dark orange; clypeus reddish brown or blackish; apex of hind femur often somewhat infuscate. Distribution. Austria, British Isles (England, Wales, Scotland, Ireland), Bulgaria, Czech Republic, *Denmark, Finland, Germany, *Greece (mainland and Corfu), Hungary, Italy, *Kazakhstan, *Montenegro, Netherlands, Norway, *Romania, Russia (including W. Caucasus), Sweden, *Tadzhikistan. New synonymy. Both male holotypes of I. unipunctator and I. ductor are preserved in the Thunberg collection, but are severely damaged. From the holotype of A. ductor only the head, fore coxa, mesoscutum and metasoma remain (Roman, 1912). The holotype of A. unipunctator has the head and the metasoma separately glued on a card and the mesosoma is still attached to the pin. Judging from these remnants (especially the mainly smooth and very shiny 3rd tergite, the wide hypoclypeal depression (0.6 × minimum width of face) and the coriaceous vertex), it is obvious that both belong to the same species. Consequently, A. ductor (Thunberg) is synonymised with A. unipunctator (Thunberg) (syn. nov.). Aleiodes ductor auctt. is divided among A. pallidicornis (Herrich-Schäffer, 1838) (N and C European populations) and A. apicalis (Brullé, 1832) (Mediterranean and southern C European populations). Aleiodes venustulus (Kokujev, 1905) Figs 813–826 Rhogas venustulus Kokujev, 1905: 15. Rogas venustulus; Shenefelt, 1975: 1255–1256; Tobias, 1986: 78 (transl.: 129). Aleiodes venustulus; Fortier & Shaw, 1999: 230; Aydogdu & Beyarslan, 2005: 192, 2006: 87. Rhogas (Rhogas) robustus Telenga, 1941: 151, 415. Rogas robustus; Tobias, 1986: 78 (transl.: 129; as synonym of A. venustulus). Type material. Holotype of A. venustulus, ♀ (ZISP), “[Kyrgyzstan], Ushch. Kyzylsu[u], [= village south of Lake Issyk Kul], 7–9.000’[ft], 5.vii.[19]03, E. Pojarkov”, No. 2273, Rh. venustulus Kok.”, “K. Kokujeva”. Additional material. 1 ♀ + 1 ♂ (MTMA), “[Kazakhstan:] Turkestan, Almásy, Naryn-kol, 1906, Tekkes”. Molecular data. None. Biology. Unknown. Diagnosis. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 822); OOL twice as long as diameter of posterior ocellus and coarsely rugose; clypeus rather short, rather flat, weakly protruding anteriorly and its ventral

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Figures 813, 814. Aleiodes venustulus (Kokujev), ♀, holotype 813 habitus lateral 814 ovipositor sheath lateral.

margin thick (Fig. 824); mesoscutum coarsely and remotely punctate, with interspaces smooth and wider than punctures; precoxal sulcus area coarsely and densely punctate; vein 1-CU1 of fore wing approx. 0.8 × as long as vein 2-CU1 and as long as vein m-cu (Fig. 815); membrane near veins M+CU1 and 1-CU1 of fore wing sparsely setose; tarsal claws rather robust and with some fine dark brown spiny bristles subbasally (Fig. 825); head brownish yellow; vein 1-M of fore wing brown; apical half of hind tibia dark brown; metasoma of ♀ yellowish, but anterior 0.6 of first tergite black; wing membrane subhyaline. Description. Holotype of A. venustulus, ♀, length of fore wing 6.7 mm, of body 8.6 mm. Head. Antenna incomplete, with eight segments remaining; frons rugose and shiny; OOL twice diameter of posterior ocellus, mainly rugose and shiny; stemmaticum densely punctate; vertex remotely punctate and shiny; clypeus punctate and slightly convex; ventral margin of clypeus thick and anterior part weakly protruding (Fig. 824); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 822); length of eye 1.3 × temple in dorsal view (Fig. 823); vertex behind stemmaticum convex and remotely punctate; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view and temple as wide as eye.

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Figures 815–826. Aleiodes venustulus (Kokujev), ♀, holotype 815 fore wing 816 hind wing 817 mesosoma lateral 818 mesosoma dorsal 819 metasoma dorsal 820 fore femur lateral 821 hind femur lateral 822 head anterior 823 head dorsal 824 head lateral 825 inner hind tarsal claw lateral 826 base of antenna.

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Mesosoma. Mesoscutum coarsely and remotely punctate, with interspaces smooth and wider than punctures; precoxal area coarsely and densely punctate, remainder of mesopleuron remotely punctate and antero-dorsally rugose; metapleuron densely and coarsely punctate; metanotum with coarse medio-longitudinal carina anteriorly; scutellum punctate; propodeum convex and coarsely rugose, its medio-longitudinal carina present only on anterior third of propodeum. Wings. Fore wing: r 0.4 × 3-SR (Fig. 815); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 815); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width twice width at level of hamuli (Fig. 816); 2-SC+R short and vertical; m-cu absent; M+CU:1M = 12:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws rather robust and with some fine dark brown spiny bristles subbasally (Fig. 825); hind coxa largely densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely rugose-reticulate; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 819); 2nd suture deep and narrow; basal half of 3rd tergite punctate-rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 814). Colour. Black; antenna (except scapus and pedicellus), palpi, tegulae, fore and middle telotarsi, veins, and pterostigma dark brown; coxae, trochanters and trochantelli, apical third of hind femur (ventrally extended to its apical two-thirds), hind tibia (except pale yellowish basal ring), fore and middle femora apically, and hind tarsus black; remainder of legs yellowish brown; wing membrane subhyaline. Variation. No specimens with intact antennae examined; 2nd tergite coarsely rugose-reticulate or coarsely longitudinally rugose. Male is very similar and with apical tergites type 3, setae quite dense, glabrous stripe narrow, and fringe very weak. Distribution. Kazakhstan, Kyrgyzstan. Notes. This Central Asian species bears a superficial resemblance to A. miniatus and A. aestuosus. It is included in this revision, because it has been reported twice from Turkey (Aydogdu and Beyarslan 2005, 2006). Aleiodes zwakhalsi van Achterberg & Shaw, sp. nov. http://zoobank.org/3C42EDAB-B9DB-45DB-884C-B23A630B249F Figs 827–840 Type material. Holotype, ♀ (RMNH), “Turkey, Ankara, Kizilcahaman, 1100 m, 17.vi.1985, C.[J.] Zwakhals”. Paratypes: 1 ♂ (RMNH), “Turkey, Agri, Mt. Ararat, 1800 m, 3.vii.1985, C.J. Zwakhals”; 1 ♀ (RMNH), “Turkiye, Gümüshane, 40–46 km

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Figures 827–829. Aleiodes zwakhalsi sp. nov., ♀, holotype 827 habitus lateral 828 detail of fore wing 829 ovipositor sheath lateral.

E [of ] Bagburt, 1500 m, 19.vii.1989, J.A.W. Lucas”; 1 ♀ (NMS), “Turkey: Zigana Dagi, 5,000 ft., SW of Tabzon, 10.viii.1959, K.M. Guichard”. Molecular data. None. Biology. Unknown. Probably univoltine as all known adults were collected in the period June to August, but there is no indication of its means of overwintering.

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Figures 830–840. Aleiodes zwakhalsi sp. nov., ♀, holotype 830 wings 831 mesosoma lateral 832 mesosoma dorsal 833 metasoma dorsal 834 fore femur lateral 835 hind femur lateral 836 base of antenna 837 head anterior 838 head dorsal 839 head lateral 840 outer hind tarsal claw lateral.

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Diagnosis. Maximum width of hypoclypeal depression approx. 0.7 × minimum width of face (Fig. 837); OOL of ♀ finely remotely punctate and 0.8–1.0 × diameter of posterior ocellus; ventral margin of clypeus thick apically and not protruding in lateral view (Fig. 839); lobes of mesoscutum densely finely punctate, with interspaces shiny; precoxal area densely punctate and with some rugae medially; vein cu-a of fore wing oblique, parallel with vein 3-CU1; surroundings of veins M+CU1 and 1-+2-CU1 largely setose; vein r of fore wing 0.4–0.5 × vein 3-SR (Fig. 830); vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1 (Fig. 830); hind tarsal claws with rather conspicuous pale brown pecten (Fig. 840); 1st tergite gradually narrowed basally (Fig. 833); 2nd tergite 0.7 × as long as wide basally and black; 3rd tergite densely punctulate basally, and sparsely so apically; head black; vein 1-M of fore wing brown; wing membrane subhyaline. This new species is similar to A. cruentus (Nees), but it differs by having the surroundings of veins M+CU1 and 1-+2-CU1 partly setose (Fig. 828; largely glabrous in A. cruentus), vein M+CU1 of fore wing with bend near its distal fifth (Fig. 828; straight or slightly curved), vein cu-a of fore wing oblique, parallel with vein 3-CU1 (vertical or nearly so, rarely oblique), vein r of fore wing 0.4–0.5 × vein 3-SR (0.3–0.4 ×), vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1 (0.8–1.1 ×, rarely less), OOL of ♀ 0.8–1.0 × diameter of posterior ocellus (0.5–0.8 ×, rarely longer), length of eye 1.1–1.6 × temple in dorsal view (1.5–1.9 ×), ovipositor sheath comparatively slender and area between ocelli and eyes moderately punctate (coarsely punctate). Also very similar to A. diversus (Szépligeti), it differs from the latter by having vein 1-CU1 of fore wing distinctly shorter than vein m-cu, hind femur approx. 4 × as long as wide, vein cu-a inclivous (parallel with vein 3-CU1; vertical and vein 3-CU1 diverging posteriorly in A. diversus), 5th–10th antennal segments of ♀ as long as wide (shorter than wide), vertex and OOL remotely punctate (densely punctate) and ovipositor sheath slender (robust). Description. Holotype, ♀, length of fore wing 7.0 mm, of body 9.0 mm. Head. Antennal segments of ♀ 59, length of antenna 1.1 × fore wing, its subapical segments moderately slender; frons largely smooth; OOL equal to diameter of posterior ocellus, finely remotely punctate and shiny; vertex distinctly punctate and shiny; clypeus punctate-rugose, wide and short; ventral margin of clypeus thick and not protruding forwards (Fig. 839); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 837); length of eye 1.1 × temple in dorsal view (Fig. 838); vertex behind stemmaticum densely punctate; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely densely and finely punctate, shiny; precoxal area of mesopleuron densely punctate, medially with few rugae; surroundings of pre­ coxal area densely punctate; scutellum sparsely and finely punctate, rather flat, shiny and laterally rugose-punctate; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete but irregular posteriorly, and no protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 830); 1-CU1 horizontal, 0.5 × 2-CU1 and 0.7 × m-cu; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 830); cu-a inclivous, straight; 1-M slightly curved posteriorly; vein M+CU1 of fore wing with distinct bend near its distal fifth (Fig. 828); 1-SR widened; surroundings of M+CU1,

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1-M and 1-CU1 largely setose. Hind wing: marginal cell gradually widened, its apical width 2.1 × width at level of hamuli (Fig. 830); 2-SC+R subquadrate; short m-cu weakly developed; M+CU:1-M = 7:4; 1r-m 0.9 × 1-M. Legs. Tarsal claws with rather conspicuous pale brownish pecten (Fig. 840); hind coxa largely densely punctulate; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 5.0 × their width, respectively; length of inner hind spur 0.55 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically and distinctly narrowed basally (Fig. 833); 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose; medio-basal area of 2nd tergite wide triangular and short (Fig. 833); 2nd tergite 0.7 × as long as its basal width; 2nd suture deep and finely reticulate; basally 3rd tergite densely punctulate and apically (as remainder of metasoma) sparsely punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath slender, with medium-sized setae and apically rounded (Fig. 829). Colour. Black (including fore coxa anteriorly and basally); apex of hind tibia, telotarsi, hind tarsus, palpi, pterostigma and veins, dark brown; hind tibia (except apex) brownish yellow; clypeus narrowly ventrally, remainder of legs, pronotum (except ventrally), mesopleuron dorsally, mesoscutum, scutellum, and metanotum, orangebrown; tegulae pale brownish yellow; wing membrane subhyaline. Variation. Antennal segments of ♀ 58(1), 59(1), 60(1), of ♂ 62(1); vein r of fore wing 0.4–0.5 × vein 3-SR; vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1; OOL of ♀ 0.8–1.0 × diameter of posterior ocellus; length of eye 1.1–1.6 × temple in dorsal view; clypeus ventrally orange brown or black; mesopleuron dorsally or largely orange brown; basal half of third tergite rugose, punctate-rugose or punctulate; entire 1st tergite orange brown, posterior half of 1st tergite brownish and rest of tergite blackish (as base of middle coxa) or entirely black. Male is very similar, apical tergites type 1 and no fringe observed. Distribution. Turkey (Asian part: 1100–1800 m).

Erratum for Part 1 In the key given by van Achterberg and Shaw (2016) the newly described species A. carminatus van Achterberg & Shaw was inserted at a late stage, which led to an error in couplet 14, as there is also a weak apical comb at the apex of the inner side of the hind tibia present in A. angustipterus van Achterberg & Shaw. The other characters provided should easily separate the two species.

Acknowledgements We thank Sergey Belokobylskij and Konstantin Samartsev (ZISP) for their exceptional help with type specimens and additional material; without this, the present extensive revision would have been impossible. Pekka Malinen and Marti Koponen (FMNH)

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for the loan of types and specimens, gifts of specimens, and the hospitality to the first author during his visit to Helsinki. Gavin Broad (BMNH), Claire Villemant and Agnièle Touret-Alby (MNHN), Martin Schwarz (BZL), Andrew Liston (SDEI), Stefan Schmidt (ZSSM), Gergely Várkonyi (SYKE), Kathryn Riddington (CNIM), and Zoltán Vas (MTMA) for the loan of valuable specimens, and also the many individuals who have loaned specimens from their personal collections (see list of depositories). Jeroen Voogd (Ede) for the permission to use his excellent photograph of the mummified Acronicta auricoma. Many entomologists, most of whose names appear in the rearing data, have often gone considerably out of their way to contribute reared (often living) and other material to the NMS collection, and/or have helped MRS by providing host eggs for experimental cultures, all of which has been invaluable for this study. Funding for sequencing came from a variety of sources including NERC grant NDC519583 to D.L.J. Quicke (and A. Purvis) and Government of Canada through Genome Canada and the Ontario Genomics Institute (2008-0GI-ICI-03). We are grateful for considerable assistance provided by Paul Hebert and M. Alex Smith at the Biodiversity Institute of Ontario, University of Guelph, Guelph, Canada. We are grateful for a Senior Postdoctoral Fellowship from the Rachadapisek Sompote Fund, Graduate School, Chulalongkorn University to DLJQ. We thank Dan Janzen and Winnie Hallwachs for allowing us to use a sequence from their projects; Håkon Haraldseide for the sequences he gave us that were funded through Norwegian Barcode of Life Network (NorBOL), funded by the Research Council of Norway and the Norwegian Biodiversity Information Centre; Julia Stigenberg for managing to obtain the sequence for Heterogamus excavatus on our behalf; Stephan Schmidt for providing a sequence for A. pulchripes which was funded by the German Federal Ministry of Education and Research (Bundesministerium für Bildung und Forschung, Berlin, Germany, project German Barcode of Life); and Sophie Cardinal and José Fernandez-Triana (Canadian National Collection of Insects, Ottawa Research and Development Centre, Agriculture and Agri-Food Canada) for sequencing numerous specimens through funding from Project J-001283 “Arthropod Systematics” from Agriculture and Agri-Food Canada.

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Appendix 1 List of barcoded specimens. Genus Aleiodes

Species or species code abraxanae

Voucher code/ Sample ID MRS636

adorabelleae aestuosus aff. wyomingensis albitibia alternator angustipterus antescutum apicalis

BCLDQ0252 MRS004 BIOUG01036-F12 MRS383 MRS161 MRS276 BCLDQ00210 MRS008 MRS111 MRS112 MRS181 MRS869 MRS079 MRS843 MRS024 MRS147 MRS197 BCLDQ0730 BCLDQ00454 BCLDQ00927 BCLDQ01479 DHJPAR0021064 MRS777 MRS162 MRS163 MRS164 MRS055 BCLDQ0245 MRS693 MRS073 MRS170 BCLDQ01515 MRS311 MRS377 BCLDQ00764 BCLDQ01555 MRS606 MRS558 MRS624 MRS625 MRS343

apiculatus assimilis aterrimus bicolor bobwhartoni bucculentus buoculus buzuritriplus cameronii Janz01 cantherius carbonarius

carminatus castaneus caudalis circumscriptus compressor concoronarius coriaceus coronopus corrusciput coxalis cruentus

curticornis damus diarsianae dissector

BCLDQ00126 MRS030 MRS007 MRS025 MRS145

Country

BOLD Process ID (if GenBank accession applicable) number (s) UK ASQSP978-10 HQ551278/ HQ551264 Thailand ASQSP222-08 JF963436 Turkey GBAH4104-09 EU979573 USA JSHYN023-11 KR791670 Sweden ASQSP075-08 JF962835 UK ASQBR258-09 MK585882 UK – KU682232 Thailand ASQSP462-08 JF962536 Turkey ASQBR099-09 EU979575 Turkey ASQBR099-09 JF962839 Turkey ASQBR100-09 MK585857 Russia ASQBR101-09 MK585870 Sweden BCNCA217-18 MK585872 UK – KU682222 France BCNCA194-18 MK585863 UK ASQBR103-09 JN000875 UK GBAH4101-09 EU979577 UK ASQBR086-09 MK585862 Thailand ASQSQ011-09 JF271188 USA ASQSP644-08 JF962486 USA ASQSQ223-09 MH272394 Thailand ASQSR027-11 JN278254 Costa Rica – JF792897 Sweden GBMIN74555-17 KU682249 Hungary ASQBR120-09 MK585853 Hungary ASQAS220-11 JF962848 Hungary ASQBR121-09 MK585851 France ASQBR152-09 JF962818 Thailand ASQSP215-08 JQ388461 France ASQSP1001-10 HQ551216 UK – KU682256 UK GBAH4098-09 EU979580 Thailand ASQSR079-11 JN278271 Sweden ASQBR140-09 JF962853 Sweden ASQBR141-09 MK585885 Thailand ASQSQ045-09 JQ388389 Thailand ASQSR119-11 JN278306 UK ASQSP553-08 MK585874 France ASQSP119-08 MK585876 Germany ASQSP989-10 HQ551274 Germany ASQSP977-10 HQ551263 Italy ASQBR105-09 JF962826/ KU682237 Thailand ASQSP378-08 JQ388354 UK ASQSP006-08 JF962600 Turkey ASQBR115-09 JF957045 Turkey ASQBR116-09 MK585881 UK ASQBR117-09 MK585849

258 Genus Aleiodes

Cornelis van Achterberg et al. / ZooKeys 919: 1–259 (2020) Species or species code dissector esenbeckii

Voucher code/ Sample ID MRS146 MRS500

Country

esenbeckii f. dendrolimi flavostriatus fortipes

MRS180

Finland

BCLDQ0628 MRS650 MRS807 MRS046 MRS048 MRS892/ BF002924 MRS225 MRS017 MRS351 MRS153 MRS208/AL422 MRS215 MRS721 MRS725 MRS619 MRS882/ BCLDQ0505 MRS883 MRS355 MRS156 BCLDQ0799 AL0037 BMNHE897778 MRS950 MRS951 MRS282 AL0323 MRS613 MRS216 MRS401 MRS880/ BCLDQ0277 MRS881/ BCLDQ00123 MRS723 MRS001 MRS885 CCDB-27844-E04 CK0002 MRS556 MRS654

Thailand France Poland France France Spain UK UK UK UK UK UK UK UK UK Romania

ASQSP849-08 ASQSP966-10 ASQBR957-18 ASQAS218-11 ASQBR118-09 ASQBR119-09 ASQBR273-09 ASQBR285-09 ASQSP101-08 ASQBR296-09 ASQBR959-19 ASQBR122-09 ASQBR960-19 ASQBR961-19 ASQSP1000-10 ASQSP695-08

JF962636 HQ551254 MH272207 EU979582 MK585879 JF962819 MK585878 MK585867 MK585847 MK585887 MK585859 JF957046 MK585855 MK585886 HQ551215 MK585888

Romania Finland UK Brazil Thailand Belize Sweden Sweden UK Uganda UK UK Finland Russia

ASQSP694-08 ASQBR123-09 ASQBR196-09 ASQSQ080-09 ASQBR476-09 ASQSQ642-10 ALEIO030-19 ALEIO031-19 ASQBR124-09 GBAH2900-07 ASQSP769-08 GBAH4093-09 ASQBR093-09 ASQSP247-08

MK585850 JF962849 JF962813 MH272348 JF962707 HQ551341 MN968689 MN968690 JF962850 EF115447 KU682243 EU979585 MK585856 MK585871

UK

ASQSP375-08

JF962562

Austria Turkey Russia Kenya Thailand Austria Bulgaria

ASQBR930-18 GBAH4092-09 ASQSP742-08 BBTH766-17 ASQSP942-10 GBMIN74565-17 ASQSP979-10

BCLDQ1202 MRS847 MRS873/ BC-ZSMHYM-27497-A05 MRS212 MRS824 MRS796

Thailand Sweden Sweden

ASQSQ410-09 BCNCA197-18 BCHYM15694-17

MH272254 EU979586 JF957043 MH272335 HQ551236 KU682242 HQ551265/ KU682244 HM435162 MK585848 MK585880

UK Bulgaria France

GBAH4091-09 AAHYM658-16 ASQBR939-18

EU979587 MK585865 MH272232

gasterator

gastritor agg spG1 gastritor agg spG2 gastritor agg spG3 gastritor agg spG4 gastritor agg spG5 grassator

hirtus

hirtus jakowlewi leptofemur melanopterus mellificus mexicanus miniatus modestus mubfsi nigriceps nigricornis nobilis

nunbergi pallidator pallidicornis pappi paulmarshi pictus praetor probuzurae pulchripes

punctipes quadrum

UK Spain

BOLD Process ID (if GenBank accession applicable) number (s) GBAH2876-07 EF115471 ASQBR092-09 JF962845/ KU682240 GBAH4097-09 EU979581

Revision of western Palaearctic Aleiodes Wesmael, II. Genus Aleiodes

Species or species code reticulatus risaae rivulus ruficornis

nr ruficornis rufipes

rugulosus

ryrholmi seriatus agg sibiricus

signatus similis sophieae sp M3 spurivena terminalis testaceus trevelyanae trianguliscleroma trisphaeropyx turcicus ungularis unipunctator

Heterogamus

valinus varius dispar excavatus fasciatipennis

Voucher code/ Sample ID MRS808 BCLDQ1268 BCLDQ01646 MRS140 MRS877 MRS887/ AL0149 MRS888 MRS889 MRS890 MRS891 MRS886 MRS294 MRS312 MRS314 MRS673/ SwedFin2 MRS674/ SwedFin5 MRS676/ SwedFin17 MRS680/ SwedFin28 MRS191 MRS217 MRS398 MRS884/ BCLDQ0509 CollHH1599 MRS395 MRS616 MRS310 MRS313 MRS805 MRS844 MRS696 BCLDQ01065 MRS703/ MRSA 703 BCLDQ00003 BCLDQ0692 MRS072 AL0226 CCDB27844-E03 BCLDQ01643 MRS126 MRS604 MRS211 MRS221 MRS354 MRS893 CollHH1603 CollHH1604 CollHH1605 BCLDQ0267 MRS446 MRS066 MRS717 MRS669

Country Poland Thailand Thailand UK Sweden UK UK ? UK UK Hungary Sweden Sweden Sweden Finland Finland Finland Finland Hungary UK France Poland Norway Sweden UK Sweden Sweden France Austria Austria Thailand Hungary Vietnam USA UK Uganda Malawi Thailand Turkey France UK Germany UK UK Norway Norway Norway Thailand Russia UK Sweden Sweden

259

BOLD Process ID (if GenBank accession applicable) number (s) GBAHB1497-18 KU682262 ASQSQ476-09 HM435190 ASQSR210-11 JN278372 ASQBR132-09 MK585892 BCNCA221-18 MK585860 ASQBR131-09 MK585846 GBAH2871-07 EF115476 ASQBR127-09 EF115475 ASQBR962-19 MK585883 GBAH5262-09 EF115477 ASQSP806-08 JF962480 ASQBR134-09 MK585884 ASQSP069-08 MH272380 ASQBR135-09 MK585873 ASQBR137-09 MK585864 ASQBR138-09 MK585854 ASQBR136-09 JF962855 ASQBR139-09 JF962854 ASQBR142-09 JF962857 ASQBR143-09 JF962856 ASQSP011-08 MH272286 ASQSP699-08 MK585875 COLHH1790-18 ASQBR322-09 ASQSP754-08 ASQBR157-09 ASQBR158-09 BCNCA185-18 BCNCA195-18 ASQSQ727-10 ASQSQ362-09 ASQSR240-11 – ASQSP913-08 ASQBR963-19 – BBTH765-17 ASQSR207-11 ASQAS219-11 ASQSP757-08 ASQAS221-11 ASQBR171-09 ASQSP035-08 BCNCA223-18 COLHH1794-18 COLHH1795-18 COLHH1796-18 ASQSP237-08 – ASQBR042-09 ASQBR935-18 ASQBR044-09

MK585877 JF962792 MH272311 JF962862 MH272159 MK585861 MK585889 HQ551413 JQ388368 MK585852 KY621612 JF962663 MK585866 EF115433 MH272236 JQ388329 JF962613 JF962867 JF962868 MK585858 MK585890 MK585891 MK585893 MK585868 MK585869 JF963430 HQ551275 JF963405 MH272379 MH272347

zookeys.919.39642-treatment1-44.xml 1. XML treatment for Aleiodes. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396428EE21655-5B9C-5413-B4C9D48B32191822 Aleiodes Wesmael, 1838Figs 10–21, 22–25, 26–37, 38, 39, 40–49, 50–53, 54–65, 66–71, 72, 73, 74–79, 80, 81–92, 93–97, 98–100, 101, 102–115, 116–118, 119–131, 132–137, 138– 141, 142–153, 154–160, 161–163, 164–177, 178, 179, 180–191, 192–195, 196–198, 199–211, 212– 216, 217–220, 221–233, 234–239, 240–242, 243–255, 256–263, 264, 265, 266–277, 278–288, 289– 291, 292–302, 303–306, 307–309, 310–321, 322–324, 325–338, 339–342, 343–346, 347–359, 360– 364, 365–367, 368–380, 381–384, 385–389, 390–399, 400–408, 409, 410, 411–424, 425–427, 428– 431, 432–444, 445–452, 453, 454, 455–466, 467, 468, 469–481, 482, 483, 484–495, 496, 497–503, 504, 505, 506–518, 519–521, 522–535, 536, 537–542, 543, 544, 545–557, 558, 559, 560–572, 573– 576, 577–589, 590, 591–603, 604–607, 608–621, 622–626, 627, 628, 629–641, 642, 643–651, 652, 653, 654–666, 667–669, 670–681, 682–685, 686, 687, 688–700, 701–703, 704–716, 717, 718, 719, 720–727, 728, 729, 730–742, 743, 744–757, 758, 759, 760–762, 763–776, 777, 778, 779–791, 792– 794, 795–807, 808–812, 813, 814, 815–826, 827–829, 830–840 Aleiodes Wesmael, 1838: 194; Shenefelt 1975: 1163–1185; Marsh 1979: 177–178; Papp 1985a: 143–164, 1985b: 347–349; Shaw and Huddleston 1991: 95–96 (biology); van Achterberg 1991: 24; Zaldívar-Riverón et al. 2004: 225, 2008: 329 (phylogeny); van Achterberg and Shaw 2016: 8–11 (groups). Type species (designated by Viereck 1914): Aleiodes heterogaster Wesmael, 1838 [examined; = A. albitibia (Herrich-Schäffer, 1838)]. Petalodes Wesmael, 1838: 123; Tobias 1971: 218 (transl. 1975: 86–87); Shenefelt 1975: 1209–1211; Tobias 1976: 90; Marsh 1979: 179; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838); van Achterberg and Shaw 2016: 8. Type species (by monotypy): Petalodes unicolor Wesmael, 1838 [examined; = Aleiodes compressor (Herrich-Schäffer, 1838)]. Schizoides Wesmael, 1838: 94. Unavailable name. Nebartha Walker, 1860: 310; Shenefelt 1975: 1216; Marsh 1979: 179; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Nebartha macropodides Walker, 1860 [examined]. Tetrasphaeropyx Ashmead, 1889: 634; Shenefelt 1975: 1260; Marsh 1979: 179; Fortier and Sherman 2008: 445 (as subgenus of Aleiodes Wesmael, 1838); Zaldívar-Riverón et al. 2008: 329 (as synonym of subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Rogas pilosus Cresson, 1872 [examined]. Neorhogas Szépligeti, 1906: 605; Shenefelt 1975: 1205; van Achterberg 1991: 24 (as subgenus of Aleiodes Wesmael, 1838); Zaldívar-Riverón et al. 2008: 329 (included in subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Neorhogas luteus Szépligeti, 1906 [examined; = Aleiodes praetor (Reinhard, 1863)]. Chelonorhogas Enderlein, [Sept. 1st] 1912a: 258; Shenefelt 1975: 1187; van Achterberg 1991: 24 (as subgenus of Aleiodes Wesmael, 1838); Zaldívar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838). Type species (by monotypy): Chelonorhogas rufithorax Enderlein, 1912 [examined; not Aleiodes rufithorax (Cameron, 1911) = A. convexus van Achterberg, 1991]. Eucystomastax Brues, [(end of?) Sept.] 1912: 223; Shaw 1993: 5 (as subgenus of Aleiodes Wesmael, 1838); Zaldívar-Riverón et al. 2004: 225 (included in Aleiodes Wesmael, 1838); Shimbori & Penteado-Dias 2011: 17 (as subgenus of Aleiodes Wesmael, 1838). Type species (by monotypy): Eucystomastax bicolor Brues, 1912 (= Rogas melanopterus Erichson, 1848). Leluthinus Enderlein, 1912b: 96; Shenefelt 1975: 1202–1203; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Leluthinus lividus Enderlein, 1912 [examined]. Aleirhogas Baker, 1917b: 383, 411; Shenefelt 1975: 1185–1186; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (designated by Viereck 1921): Rhogas (Aleirhogas) schultzei Baker, 1917 [examined]. Hemigyroneuron Baker, 1917a: 284, 322–327; Zaldívar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838); Butcher and Quicke 2011: 1405 (as subgenus of Aleiodes Wesmael, 1838, and HemigyroneuronsensuZaldívar-Riverón et al. (2008) is not Hemigyroneuron)); Butcher and Quicke 2015: 275–279. Type species (original designation): Hemigyroneuron speciosus Baker, 1917 [examined]. Heterogamoides Fullaway, 1919: 43; Shenefelt 1975: 1188; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Heterogamoides muirii Fullaway, 1919 [examined]. Cordylorhogas Enderlein, 1920: 153; Shenefelt 1975: 1195; van Achterberg 1991: 31; Zaldívar-Riverón et al. 2004: 232, 2008: 329 (as synonym of subgenus Aleiodes Wesmael, 1838). Type species (by monotypy): Cordylorhogas trifasciatus Enderlein, 1920 [examined]. Hyperstemma Shestakov, 1940: 10; Shenefelt 1975: 1200; van Achterberg 1991: 24 (as synonym of Aleiodes Wesmael, 1838). Type species (by monotypy): Hyperstemma chlorotica Shestakov, 1940 [examined]. Dimorphomastax Shenefelt, 1979: 131–133; Shaw et al. 1998: 66 (as synonym of Aleiodes Wesmael, 1838). Type species (by original designation): Dimorphomastax peculiaris Shenefelt, 1979 [examined; = Aleiodes atriceps Cresson, 1869]. Pholichora van Achterberg, 1991: 48–53; Quicke and Shaw 2005: 532; Zaldívar-Riverón et al. 2008: 329 (as synonym of Aleiodes Wesmael, 1838); Butcher and Quicke 2011: 1405 (as synonym of subgenus Hemigyroneuron Baker, 1917); Butcher et al. 2012: 9 (id.). Type species (original designation): Hemigyroneuron madagascariensis Granger, 1949 [examined]. Arcaleiodes Chen & He, 1997: 60–62; Zaldívar-Riverón et al. 2008: 329 (as subgenus of Aleiodes Wesmael, 1838); Butcher et al. 2012: 18–19 (id.). Type species (original designation): Aleiodes unifasciatus Chen & He, 1991 [examined]. Vietorogas Long & van Achterberg, 2008: 313–314; Butcher et al. 2012: 15–17 (as synonym of Aleiodes Wesmael, 1838). Type species (original designation): Vietorogas bachma Long, 2008 [examined]. R (h)ogas auct; Tobias, 1971: 215–217 (transl. 1975: 83–86); Shenefelt, 1975: 1215–1256; Tobias, 1976: 81–89; Marsh, 1979: 179–181; Tobias, 1986: 74–84. Hyperstemma Shestakov, 1940, is traditionally included in the genus Heterogamus Wesmael, 1838 (Shenefelt 1975) or in the subgenus Heterogamus of the genus Aleiodes Wesmael (e.g., Belokobylskij 2000), but differs by the shape of the head (Figs 17–19) and of the tarsal claws (Fig. 20), the position of the clypeus (Fig. 17), the elongate 2nd submarginal cell of the fore wing (but folded in Fig. 10), the widened 1st subdiscal cell of the fore wing and distinctly widened marginal cell of the hind wing (Fig. 10). Therefore, we retain the subgenus Hyperstemma Shestakov of Aleiodes Wesmael for at least the following species: A. chloroticus (Shestakov, 1940) from China (Palaearctic and Oriental), *Japan (RMNH), Korea, and Russia (Far East), A. albigenus Chen & He, 1997, from China (Oriental) and Vietnam, A. crassinervis Chen & He, 1997, from China (Oriental) and Vietnam, A. naevius Chen & He, 1997, from China (Oriental), and A. pallidinervis (Cameron, 1910) from China (Palaearctic and Oriental), Japan, Korea, and Russia (Far East). 10.3897/zookeys.919.39642.figures10-2114D64090-0E55-56E6-8539-3322ECC56CB9Figures 10– 21. Aleiodes chloroticus (Shestakov), ♂, Japan 10 wings 11 mesosoma lateral 12 mesosoma dorsal 13 1st–3rd metasomal tergites dorsal 14 fore femur lateral 15 hind femur lateral 16 base of antenna 17 head anterior 18 head dorsal 19 head lateral 20 outer hind tarsal claw 21 apex of antenna. https://binary.pensoft.net/fig/39004910.3897/zookeys.919.39642.figures22-2591C13739-EA905B88-BA5E-C363154C28E1Figures 22–25. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River, but 25 from Uzbekistan, Qamashi 22 ovipositor sheath lateral 23 habitus lateral 24 apex of antenna 25 mummy of Heliothis peltigera (Denis & Schiffermüller). https://binary.pensoft.net/fig/39005010.3897/zookeys.919.39642.figures26-3779D0915E-1C6A5272-A38E-E5E71FA2DAC4Figures 26–37. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River 26 fore wing 27 hind wing 28 mesosoma lateral 29 mesosoma dorsal 30 metasoma dorsal 31 fore femur lateral 32 hind femur lateral 33 base of antenna 34 head anterior 35 head dorsal 36 head lateral 37 inner hind tarsal claw. https://binary.pensoft.net/fig/39005110.3897/zookeys.919.39642.figures38-3959BD7BC5-62D75AC8-B75B-1B987E9AD7A4Figures 38, 39. Aleiodes agilis (Telenga), ♀, paralectotype 38 habitus lateral 39 ovipositor sheath lateral. Photographs by K. Samartsev. https://binary.pensoft.net/fig/39005210.3897/zookeys.919.39642.figures40-49BD25508C-E43B52D7-84C7-A56A662B24A1Figures 40–49. Aleiodes agilis (Telenga), ♀, paralectotype, but 47 of lectotype 40 wings 41 mesosoma lateral 42 fore femur lateral 43 hind femur lateral 44 outer hind tarsal claw 45 fore leg 46 head anterior 47 head dorso-lateral 48 head lateral 49 base of antenna. Photographs by K. Samartsev. https://binary.pensoft.net/fig/39005310.3897/zookeys.919.39642.figures50-5325D7F282-B3A75EF0-A169-0A257A48DF74Figures 50–53. Aleiodes apicalis (Brullé), ♀, Greece, Thimiana Chios, but 52 mummies of Autographa gamma (Linnaeus) from Malta and 53 of undetermined plusiine host from Cyprus 50 habitus lateral 51 ovipositor sheath lateral 52 mummy dorsal 53 mummy covered by silk of host. https://binary.pensoft.net/fig/39005410.3897/zookeys.919.39642.figures54-65FB7FA51D-63DD5751-B715-879670F8A4E0Figures 54–65. Aleiodes apicalis (Brullé), ♀, Bulgaria, Rodopi 54 wings 55 mesosoma lateral 56 mesosoma dorsal 57 1st –3rd metasomal tergites dorsal 58 fore femur lateral 59 hind femur lateral 60 head anterior 61

head dorsal 62 head lateral 63 outer hind tarsal claw 64 base of antenna 65 apex of antenna. https://binary.pensoft.net/fig/39005510.3897/zookeys.919.39642.figures66-71143A68FD-09935A13-ACDF-DC57416186CBFigures 66–71. Aleiodes apicalis (Brullé), ♂, Turkey, Sivas 66 habitus lateral 67 head dorsal 68 3rd –7th metasomal tergites dorsal 69 3rd –7th metasomal tergites lateral 70 head anterior 71 outer hind tarsal claw. https://binary.pensoft.net/fig/39005610.3897/zookeys.919.39642.figures72-73D8339728-0BB75DE2-AF96-7F6763CD34ECFigures 72, 73. Aleiodes arnoldii (Tobias), ♀, holotype 72 habitus lateral 73 ovipositor sheath lateral. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39005710.3897/zookeys.919.39642.figures74-791EB71D01-9D245985-8EC6-CB0EAF5EA6DAFigures 74–79. Aleiodes arnoldii (Tobias), ♀, holotype 74 wings 75 mesosoma lateral 76 antenna 77 head lateral 78 head anterior 79 head dorsal. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39005810.3897/zookeys.919.39642.figure809FE171F2-0F5F-5F6F90BE-4550C5CAD1C1Figure 80. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass, habitus lateral. https://binary.pensoft.net/fig/39005910.3897/zookeys.919.39642.figures81-92D7B63730-B1C95937-9156-0402627E8E5FFigures 81–92. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass 81 wings 82 mesosoma lateral 83 mesosoma dorsal 84 metasoma dorsal 85 fore femur lateral 86 hind femur lateral 87 inner hind tarsal claw 88 head anterior 89 head dorsal 90 head lateral 91 base of antenna 92 apex of metasoma lateral. https://binary.pensoft.net/fig/39006010.3897/zookeys.919.39642.figures93-977E59AE8D-3A52583D-BBEE-A55931BCD1A8Figures 93–97. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 93 habitus lateral 94 detail of fore wing with arrow indicating lost vein r-m 95 ovipositor sheath lateral 96, 97 mummies of Amphipyra sp. showing variation in markings. https://binary.pensoft.net/fig/39006110.3897/zookeys.919.39642.figures98-100051EF52B-1F9E5B88-BA41-4A1940BB491FFigures 98–100. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 98 parasitised caterpillar of Amphipyra sp. 99 early stage of mummy 100 later stage of mummy. https://binary.pensoft.net/fig/39006210.3897/zookeys.919.39642.figure1019D618773-4828-506AA8FF-0BAEB8C42B2AFigure 101. Larva of Aleiodes aterrimus (Ratzeburg) mummifying Amphipyra pyramidea (Linnaeus), with its anterior (indicated by the arrow) projecting from the ventral opening in the host to spread adhesive over a wide area. https://binary.pensoft.net/fig/39006310.3897/zookeys.919.39642.figures102-115A19714FD-6AB9522E-9BF7-1EADD8FAF96EFigures 102–115. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest, but 102 from Austria, Wien 102 fore wing 103 hind wing 104 mesosoma lateral 105 mesosoma dorsal 106 metasoma dorsal 107 fore femur lateral 108 hind femur lateral 109 head anterior 110 head dorsal 111 head lateral 112 hind tibia and tarsus lateral 113 outer hind tarsal claw 114 base of antenna 115 apex of antenna. https://binary.pensoft.net/fig/39006410.3897/zookeys.919.39642.figures116-1187A65E536-5B4558A6-A395-B61CA19A0A9DFigures 116–118. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 116 habitus lateral 117 ovipositor sheath lateral 118 mummy of Tholera decimalis (Poda). https://binary.pensoft.net/fig/39006510.3897/zookeys.919.39642.figures119-131C42B5EB2-8FBC5583-A967-300046F11C24Figures 119–131. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 119 fore wing 120 hind wing 121 mesosoma lateral 122 mesosoma dorsal 123 metasoma dorsal 124 fore femur lateral 125 hind femur lateral 126 head anterior 127 head dorsal 128 head lateral 129 base of antenna 130 apex of antenna 131 inner hind tarsal claw. https://binary.pensoft.net/fig/39006610.3897/zookeys.919.39642.figures132-13714702ABD-1CCA5F18-87A4-7B99B412D4F1Figures 132–137. Aleiodes carbonarius Giraud, ♂, Hungary, Csákvár 132 habitus lateral 133 head dorsal 134 wings 135 base of antenna 136 apex of antenna 137 head anterior. https://binary.pensoft.net/fig/39006710.3897/zookeys.919.39642.figures138-141A0138053-F6345848-AFE7-BB186A4EAA4FFigures 138–141. Aleiodes carbonaroides sp. nov., ♀, holotype 138 habitus lateral 139 ovipositor sheath lateral 140 outer hind tarsal claw lateral 141 mummy of Cerapteryx graminis (Linnaeus). https://binary.pensoft.net/fig/39006810.3897/zookeys.919.39642.figures142-153E65C2ACC-84B65071-AB21-E6A3AE3F6212Figures 142–153. Aleiodes carbonaroides sp. nov., ♀, holotype 142 fore wing 143 hind wing 144 mesosoma lateral 145 mesosoma dorsal 146 metasoma dorsal 147 fore femur lateral 148 hind femur lateral 149 head anterior 150 head dorsal 151 head lateral 152 base of antenna 153 apex of antenna. https://binary.pensoft.net/fig/39006910.3897/zookeys.919.39642.figures154-16086D778D8-38175FEF-9743-028C28D1C1ECFigures 154–160. Aleiodes carbonaroides sp. nov., ♂, paratype 154 habitus lateral 155 apex of antenna 156 antenna 157 base of antenna 158 wings lateral 159 head anterior 160 head dorsal. https://binary.pensoft.net/fig/39007010.3897/zookeys.919.39642.figures161-1633948DC74-1B565A6F-A324-8F80387803A4Figures 161–163. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 161 habitus lateral 162 ovipositor sheath lateral 163 apex of antenna (of paratype). https://binary.pensoft.net/fig/39007110.3897/zookeys.919.39642.figures164-17726C7F91D-55A352EE-A04C-89C66E6AAC9FFigures 164–177. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 164 fore wing 165 hind wing 166 mesosoma lateral 167 mesosoma dorsal 168 metasoma dorsal 169 fore femur lateral 170 hind femur lateral 171 head anterior 172 head dorsal 173 head lateral 174 base of antenna 175 antenna 176 hind tibia and tarsus lateral 177 outer hind tarsal claw. https://binary.pensoft.net/fig/39007210.3897/zookeys.919.39642.figures178-179A24DA8CB-17665059-A4FE-A315916CCE31Figures 178, 179. Aleiodes coriaceus sp. nov., ♀, holotype 178 habitus lateral 179 ovipositor sheath lateral. https://binary.pensoft.net/fig/39007310.3897/zookeys.919.39642.figures180-1915548F088-03BA5D78-BB9B-82B3B3E62130Figures 180–191. Aleiodes coriaceus sp. nov., ♀, holotype 180 wings 181 mesosoma lateral 182 mesosoma dorsal 183 propodeum and 1st–3rd metasomal tergites dorsal 184 fore femur lateral 185 hind femur lateral 186 head anterior 187 head dorsal 188 head lateral 189 outer hind tarsal claw 190 base of antenna 191 apex of antenna. https://binary.pensoft.net/fig/39007410.3897/zookeys.919.39642.figures192-195212724B8-BD435195-A895-702B10123BABFigures 192–195. Aleiodes coriaceus sp. nov., ♂, paratype 192 habitus lateral 193 inner hind claw lateral 194 3rd–7th metasomal tergites dorsal 195 3rd –7th metasomal tergites lateral. https://binary.pensoft.net/fig/39007510.3897/zookeys.919.39642.figures196-198FD1C577A-699A5351-9934-D041B273800AFigures 196–198. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg, but 198 from Finland, Mäntyharju 196 habitus lateral 197 ovipositor sheath lateral 198 mummy of Hadena confusa (Hufnagel). https://binary.pensoft.net/fig/39007610.3897/zookeys.919.39642.figures199-21167A8AF3F-595956F5-96CE-E198218BF417Figures 199–211. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg 199 wings 200 mesosoma lateral 201 mesosoma dorsal 202 metasoma dorsal 203 fore femur lateral 204 hind femur lateral 205 outer hind tarsal claw 206 head anterior 207 head dorsal 208 head lateral 209 base of antenna 210 apex of antenna 211 hind tarsus lateral. https://binary.pensoft.net/fig/39007710.3897/zookeys.919.39642.figures212-2160F129638-7BE65A59-9DF4-74DA62458E82Figures 212–216. Aleiodes cruentus (Nees), ♂, Hungary, Hársbokorhegy, but 213–215 from Germany, Markkleeberg 212 habitus lateral 213 1st–3rd metasomal tergites dorsal 214 4th–7th metasomal tergites lateral 215 id. dorsal 216 basal antennal segments. https://binary.pensoft.net/fig/39007810.3897/zookeys.919.39642.figures217-220A9F2DD8D-79C855F9-BFD6-3977FBCC6C38Figures 217–220. Aleiodes desertus (Telenga), ♀, paralectotype 217 habitus lateral 218 antenna lateral 219 detail of fore wing 220 ovipositor sheath lateral. https://binary.pensoft.net/fig/39007910.3897/zookeys.919.39642.figures221-23350B12841-BF885E90-B1DE-4330FDF445C8Figures 221–233. Aleiodes desertus (Telenga), ♀, paralectotype, but 224 and 230 of lectotype 221 fore wing 222 hind wing 223 mesosoma lateral 224 mesosoma dorsal 225 metasoma dorsal 226 fore femur lateral 227

hind femur lateral 228 apex of antenna 229 head anterior 230 head dorsal 231 head lateral 232 outer hind tarsal claw 233 base of antenna. https://binary.pensoft.net/fig/39008010.3897/zookeys.919.39642.figures234-239FCCA41AD-56CD5956-81E0-E2E97F9A43E4Figures 234–239. Aleiodes dissector (Nees), U.K., Scotland (in culture) parasitising Orthosia incerta (Hufnagel) 234 pre-mummy, removed from its hideaway, with unparasitised control from the same egg batch (below) 235 pre-mummy 236 early mummification 237 mummy with ventral ooze 238 three fully hard mummies 239 emerged mummy, cut open to expose silken lining. https://binary.pensoft.net/fig/39008110.3897/zookeys.919.39642.figures240-242A06B8684-54995502-94C4-7783632397FDFigures 240–242. Aleiodes dissector (Nees), ♀, Switzerland, Tessin, but 242 from Scotland (culture) 240 habitus lateral 241 ovipositor sheath lateral 242 mummy of Orthosia incerta (Hufnagel). https://binary.pensoft.net/fig/39008210.3897/zookeys.919.39642.figures243-25596EDCBD0-E8975243-9ED2-DB847C7BA91BFigures 243–255. Aleiodes dissector (Nees), ♀, Switzerland, Tessin 243 fore wing 244 hind wing 245 mesosoma lateral 246 mesosoma dorsal 247 propodeum and 1st –4th metasomal tergites dorsal 248 fore femur lateral 249 hind femur lateral 250 outer hind tarsal claw 251 head anterior 252 head dorsal 253 head lateral 254 base of antenna 255 apex of antenna. https://binary.pensoft.net/fig/39008310.3897/zookeys.919.39642.figures256-2630C522D70-D50853F0-9482-B9A05895953EFigures 256–263. Aleiodes dissector (Nees), ♂, Austria, Kärnten 256 habitus lateral 257 head dorsal 258 3rd–7th tergites lateral 259 mesosoma dorsal 260 1st–6th metasomal tergites dorsal 261 fore femur lateral 262 head anterior 263 hind femur lateral. https://binary.pensoft.net/fig/39008410.3897/zookeys.919.39642.figures264-2650DB13A2A-D8B15BE7-9F86-A455F19B96C4Figures 264, 265. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 264 habitus lateral 265 ovipositor sheath lateral. https://binary.pensoft.net/fig/39008510.3897/zookeys.919.39642.figures266-2775C60A2CD-BA745F54-B86B-AAEDBEB6D781Figures 266–277. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 266 wings 267 mesosoma lateral 268 mesosoma dorsal 269 propodeum and metasoma dorsal 270 fore femur lateral 271 hind femur lateral 272 outer hind tarsal claw 273 head anterior 274 head dorsal 275 head lateral 276 base of antenna 277 apex of antenna. https://binary.pensoft.net/fig/39008610.3897/zookeys.919.39642.figures278-288447739B7-76B95CD8-89E5-715011AEF82AFigures 278–288. Aleiodes diversus (Szépligeti), ♂, Italy, Sicily 278 habitus lateral 279 3rd–7th tergites lateral 280 wings 281 mesosoma lateral 282 metasoma dorsal 283 outer hind claw 284 hind tibial spurs lateral 285 head anterior 286 head dorsal 287 base of antenna 288 apex of antenna. https://binary.pensoft.net/fig/39008710.3897/zookeys.919.39642.figures289-291B35C50E5-1C72516F-87E7-1AF88006BC52Figures 289–291. Aleiodes eurinus (Telenga), ♀, Mongolia, but 291 Russia, Chelyabinskoi Obl. 289 habitus lateral 290 ovipositor sheath lateral 291 apex of antenna. https://binary.pensoft.net/fig/39008810.3897/zookeys.919.39642.figures292-30214384170-9C0F50F2-AEC9-1635D7CEC291Figures 292–302. Aleiodes eurinus (Telenga), ♀, Mongolia 292 wings 293 mesosoma lateral 294 mesosoma dorsal 295 1st–3rd metasomal tergite dorsal 296 fore femur lateral 297 hind femur lateral 298 head anterior 299 head dorsal 300 head lateral 301 base of antenna 302 outer hind tarsal claw. https://binary.pensoft.net/fig/39008910.3897/zookeys.919.39642.figures303-3060C5A40D0-4F6053E8-B8A9-05C48B115355Figures 303–306. Aleiodes eurinus (Telenga), ♂, Mongolia 303 habitus lateral 304 head anterior 305 apical half of metasoma lateral 306 head dorsal. https://binary.pensoft.net/fig/39009010.3897/zookeys.919.39642.figures307-309675D22D4-B1D55BD6-99A3-5BF8C3947159Figures 307–309. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 307 habitus lateral 308 ovipositor sheath lateral 309 detail of fore wing. https://binary.pensoft.net/fig/39009110.3897/zookeys.919.39642.figures310-3210B4B9462-8FF15305-969A-53AB1B89A7CBFigures 310–321. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 310 wings 311 mesosoma lateral 312 mesosoma dorsal 313 propodeum and 1st–3rd metasomal tergites dorsal 314 fore femur lateral 315 hind femur lateral 316 base of antenna 317 apex of antenna 318 head anterior 319 head dorsal 320 head lateral 321 inner hind tarsal claw https://binary.pensoft.net/fig/39009210.3897/zookeys.919.39642.figures322-324D4F8A347-2AA65BC0-8AE2-D3D499D8D54FFigures 322–324. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 322 habitus lateral 323 ovipositor sheath lateral 324 mummy of Idaea sp. (either I. aversata (Linnaeus) or I. straminata (Borkhausen)), Poland, Dybki. https://binary.pensoft.net/fig/39009310.3897/zookeys.919.39642.figures325-338B8113AE8-13F95169-9489-132776F01013Figures 325–338. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 325 fore wing 326 hind wing 327 mesosoma lateral 328 mesosoma dorsal 329 1st –3rd metasomal tergites dorsal 330 fore femur lateral 331 hind femur lateral 332 head anterior 333 head dorsal 334 head lateral 335 base of antenna 336 apex of antenna 337 antenna 338 inner hind tarsal claw. https://binary.pensoft.net/fig/39009410.3897/zookeys.919.39642.figures339-342D9FBADDA-4F615BDC-80C2-8EC0C691093DFigures 339–342. Aleiodes fortipes (Reinhard), ♂, England, Santon Downham 339 habitus lateral 340 3rd–7th metasomal tergites dorsal 341 id. lateral 342 metasoma dorsal. Arrows indicating setose depressions or pores of 4th–6th tergites. https://binary.pensoft.net/fig/39009510.3897/zookeys.919.39642.figures343-34612E3CF58-F8FC5280-A93F-7BBCDC981CDCFigures 343–346. Aleiodes gasterator (Jurine), ♀, France, Les Constants, but 345 and 346 from Cyprus 343 habitus lateral 344 ovipositor sheath lateral 345 mummy of Agrotis sp. 346 mummy of Agrotis sp. or Spodoptera littoralis (Boisduval). https://binary.pensoft.net/fig/39009610.3897/zookeys.919.39642.figures347-359A0670B61-D775580C-AFC1-0F049202709CFigures 347–359. Aleiodes gasterator (Jurine), ♀, France, Les Constants 347 fore wing 348 hind wing 349 mesosoma lateral 350 mesosoma dorsal 351 metasoma dorsal 352 fore femur lateral 353 hind femur lateral 354 base of antenna 355 head anterior 356 head dorsal 357 head lateral 358 outer hind tarsal claw 359 apex of antenna. https://binary.pensoft.net/fig/39009710.3897/zookeys.919.39642.figures360-3645EE78E7B-70FF5FBF-99FD-10AD78AE6098Figures 360–364. Aleiodes gasterator (Jurine), ♂, Italy, Livorno, but 364 of ♀, France 360 habitus lateral 361 antenna 362 base of antenna 363 apex of antenna 364 3rd metasomal tergite dorsal. https://binary.pensoft.net/fig/39009810.3897/zookeys.919.39642.figures365-36763534165-56B55939-84DE-4D1E6D226929Figures 365–367. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 365 habitus lateral 366 antenna 367 ovipositor sheath lateral. https://binary.pensoft.net/fig/39009910.3897/zookeys.919.39642.figures368-3805A59FEEF-114E55DA-901F-68B72A70D581Figures 368–380. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 368 fore wing 369 hind wing 370 mesosoma lateral 371 mesosoma dorsal 372 metasoma dorsal 373 fore femur lateral 374 hind femur lateral 375 head anterior 376 head dorsal 377 head lateral 378 base of antenna 379 apex of antenna 380 outer hind tarsal claw. https://binary.pensoft.net/fig/39010010.3897/zookeys.919.39642.figures381-384A320B5B1-162C5A53-B93A-7EB6DB9C648CFigures 381–384. Aleiodes grassator (Thunberg), ♂, Scotland, Isle of Coll 381 habitus lateral 382 base of antenna 383 apex of antenna 384 mummy of ?Cerapteryx graminis (Linnaeus). https://binary.pensoft.net/fig/39010110.3897/zookeys.919.39642.figures385-3895C964117-B7AE5CEC-94EF-2B0AA8C8FB1BFigures 385–389. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 385 habitus lateral 386 antenna 387 apex of antenna 388 base of antenna 389 ovipositor sheath lateral. https://binary.pensoft.net/fig/39010210.3897/zookeys.919.39642.figures390-3996F7E68A7-8F39543C-84C4-CAE338A06F69Figures 390–399. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 390 wings 391 mesosoma lateral 392 mesosoma dorsal 393 metasoma dorsal 394 fore femur lateral 395 hind femur lateral 396 head anterior 397 head dorsal 398 head lateral 399 outer hind tarsal claw. https://binary.pensoft.net/fig/39010310.3897/zookeys.919.39642.figures400-408D1B53275-75FE54C4-BFF9-9020217DADA3Figures 400–408. Aleiodes hemipterus (Marshall), ♂, Tunisia, Teboursouk 400 habitus lateral 401 fore femur lateral 402 fore wing 403 hind wing 404 mesosoma dorsal 405 metasoma dorsal 406 hind femur lateral 407 head anterior 408 head dorsal. https://binary.pensoft.net/fig/39010410.3897/zookeys.919.39642.figures409-410F0E870D7-8F265AC6-A26A-EE876EE0668EFigures 409, 410.

Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 409 habitus lateral 410 ovipositor sheath lateral. https://binary.pensoft.net/fig/39010510.3897/zookeys.919.39642.figures411-42406C9A64C-18EA5F44-AA67-AAC4F9621E02Figures 411–424. Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 411 fore wing 412 hind wing 413 mesosoma lateral 414 mesosoma dorsal 415 metasoma dorsal 416 fore femur lateral 417 hind femur lateral 418 antenna 419 head anterior 420 head dorsal 421 head lateral 422 base of antenna 423 apex of antenna 424 inner hind tarsal claw. https://binary.pensoft.net/fig/39010610.3897/zookeys.919.39642.figures425-42788C46B20-505F5D88-97D8-3EF0D8A85AF0Figures 425–427. Aleiodes hirtus (Thomson), ♂, England, Chippenham Fen 425 habitus lateral 426 head and anterior part of mesosoma lateral 427 propodeum and 1st –3rd metasomal segments lateral. https://binary.pensoft.net/fig/39010710.3897/zookeys.919.39642.figures428-431E9026BF6-FCE75E98-AE08-34587C523785Figures 428–431. Aleiodes improvisus sp. nov., ♀, holotype 428 habitus lateral 429 ovipositor sheath lateral 430 right fore femur lateral 431 right hind femur lateral. https://binary.pensoft.net/fig/39010810.3897/zookeys.919.39642.figures432-444971CFB00-77C85C26-B9A6-D434A855ED2FFigures 432–444. Aleiodes improvisus sp. nov., ♀, holotype 432 fore wing 433 hind wing 434 mesosoma lateral 435 mesosoma dorsal 436 propodeum and 1st –3rd metasomal tergites dorsal 437 left fore femur lateral 438 left hind femur lateral 439 head anterior 440 head dorsal 441 head lateral 442 inner hind tarsal claw 443 base of antenna 444 apex of antenna. https://binary.pensoft.net/fig/39010910.3897/zookeys.919.39642.figures445-452D8EDEF9E-A0745F67-AA9E-AAFDD0186669Figures 445–452. Aleiodes improvisus sp. nov., ♂, paratype, Switzerland (Arolla) 445 habitus lateral 446 fore femur lateral 447 hind femur lateral 448 base of antenna 449 head dorsal 450 4th–7th metasomal tergites dorsal 451 apex of antenna 452 4th–7th metasomal tergites lateral. https://binary.pensoft.net/fig/39011010.3897/zookeys.919.39642.figures453-4541F78D44C-E9E15669-88DD-191C40331D78Figures 453, 454. Aleiodes krulikowskii (Kokujev), ♀, lectotype 453 habitus lateral 454 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011110.3897/zookeys.919.39642.figures455-466F435B958-BEE45E37-B728-26A94FCA0EA3Figures 455–466. Aleiodes krulikowskii (Kokujev), ♀, lectotype 455 fore wing 456 hind wing 457 mesosoma lateral 458 mesosoma dorsal 459 metasoma dorsal 460 fore femur lateral 461 hind femur lateral 462 base of antenna 463 head anterior 464 head dorsal 465 head lateral 466 outer middle tarsal claw. https://binary.pensoft.net/fig/39011210.3897/zookeys.919.39642.figures467-468B69F968C-201E5DCA-B4AD-DCC6DBB65AE9Figures 467, 468. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 467 habitus lateral 468 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011310.3897/zookeys.919.39642.figures469-48112BE9EBC-14EC5ED6-B474-154B8040C7B8Figures 469–481. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 469 fore wing 470 hind wing 471 mesosoma lateral 472 mesosoma dorsal 473 metasoma dorsal 474 fore femur lateral 475 hind femur lateral 476 head anterior 477 head dorsal 478 head lateral 479 base of antenna 480 apex of antenna 481 inner hind tarsal claw. https://binary.pensoft.net/fig/39011410.3897/zookeys.919.39642.figures482-483719368AD-B6D051BF-BD6C-AA524F5F9B58Figures 482, 483. Aleiodes morio (Reinhard), ♀, Hungary, Budapest 482 habitus lateral 483 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011510.3897/zookeys.919.39642.figures484-4950CD0D1F6-47205E21-8A5D-5D8DFB044606Figures 484–495. Aleiodes morio (Reinhard), ♀, Hungary, Budapest, but 490 of ♂ from Nadap 484 fore wing 485 hind wing 486 mesosoma lateral 487 mesosoma dorsal 488 1st–3rd metasomal tergites dorsal 489 fore femur lateral 490 hind femur lateral 491 head anterior 492 head dorsal 493 head lateral 494 outer middle tarsal claw 495 base of antenna. https://binary.pensoft.net/fig/39011610.3897/zookeys.919.39642.figure49645A4FBA6-B8A1-5B8FACD9-3D00389B582EFigure 496. Aleiodes morio (Reinhard), ♂, Hungary, Nadap, habitus lateral. https://binary.pensoft.net/fig/39011710.3897/zookeys.919.39642.figures497-50300E0EC37-09075628-B1CE-A04F977731DFFigures 497–503. Aleiodes morio (Reinhard), ♂, Hungary, Nadap 497 wings 498 apex of metasoma lateral 499 apex of metasoma dorsal 500 base of antenna 501 head anterior 502 head dorsal 503 inner hind tarsal claw. https://binary.pensoft.net/fig/39011810.3897/zookeys.919.39642.figures504-505ED9C75DB-F56D5A94-AD8D-E777BD02E2A3Figures 504, 505. Aleiodes nigrifemur sp. nov. ♀, holotype 504 habitus lateral 505 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011910.3897/zookeys.919.39642.figures506-518677289DC-BD215788-81C9-FB618A370D45Figures 506–518. Aleiodes nigrifemur sp. nov. ♀, holotype 506 wings 507 mesosoma lateral 508 mesosoma dorsal 509 1st–3rd metasomal tergites dorsal 510 fore femur lateral 511 hind femur lateral 512 base of antenna 513 head anterior 514 head dorsal 515 head lateral 516 apex of antenna 517 outer hind tarsal claw 518 antennae. https://binary.pensoft.net/fig/39012010.3897/zookeys.919.39642.figures519-521F8F63618-568157ED-B55E-15D28791425BFigures 519–521. Aleiodes nobilis (Haliday), ♀, neotype 519 habitus lateral 520 ovipositor sheath lateral 521 mummy of Autographa gamma Linnaeus (Germany, Lindenhayn). https://binary.pensoft.net/fig/39012110.3897/zookeys.919.39642.figures522-5355A75452D-61EA5EA4-90FF-37AA665EBD76Figures 522–535. Aleiodes nobilis (Haliday), ♀, neotype 522 wings 523 mesosoma lateral 524 mesosoma dorsal 525 1st–4th metasomal tergites dorsal 526 fore femur lateral 527 hind femur lateral 528 hind tibia lateral 529 head anterior 530 head dorsal 531 head lateral 532 base of antenna 533 apex of antenna 534 outer hind tarsal claw 535 outer fore tarsal claw. https://binary.pensoft.net/fig/39012210.3897/zookeys.919.39642.figure536BC54482C-1073-5A8A9C4F-56DE9E909C57Figure 536. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen, habitus lateral. https://binary.pensoft.net/fig/39012310.3897/zookeys.919.39642.figures537-54296B0B60C-355C5555-B8BF-D9826036FF20Figures 537–542. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen 537 apex of metasoma dorsal 538 apex of metasoma lateral 539 apex of antenna 540 head anterior 541 head dorsal 542 base of antenna. https://binary.pensoft.net/fig/39012410.3897/zookeys.919.39642.figures543-544D134C7D6-B72A58DA-81BB-DBE6AFC61F2BFigures 543, 544. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 543 habitus lateral 544 ovipositor sheath lateral. https://binary.pensoft.net/fig/39012510.3897/zookeys.919.39642.figures545-55772F3170B-E74E530E-9B3B-85E813F12FC6Figures 545–557. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 545 fore wing 546 hind wing 547 mesosoma lateral 548 mesosoma dorsal 549 1st–3rd metasomal tergites dorsal 550 fore femur lateral 551 hind femur lateral 552 head anterior 553 head dorsal 554 head lateral 555 base of antenna 556 apex of antenna 557 inner hind tarsal claw. https://binary.pensoft.net/fig/39012610.3897/zookeys.919.39642.figures558-55942663AF1-09C850CC-89B5-15D17A31DB04Figures 558, 559. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 558 habitus lateral 559 ovipositor sheath lateral. https://binary.pensoft.net/fig/39012710.3897/zookeys.919.39642.figures560-57281DECFA2-65355228-8678-F741198A7730Figures 560–572. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 560 fore wing 561 hind wing 562 mesosoma lateral 563 mesosoma dorsal 564 1st–3rd metasomal tergites dorsal 565 fore femur lateral 566 hind femur lateral 567 apex of antenna 568 head anterior 569 head dorsal 570 head lateral 571 outer hind tarsal claw 572 base of antenna. https://binary.pensoft.net/fig/39012810.3897/zookeys.919.39642.figures573-5766BE0F66C-89AD5079-9229-E7B04609AF8DFigures 573–576. Aleiodes periscelis (Reinhard), ♀, 573, 574 Russia (holotype R. jaroslawensis Kokujev) and 575, 576 Czech Republic, Pisek 573 head anterior 574, 575 habitus lateral 576 ovipositor sheath lateral. Photographs 573, 574 by K. Samartsev. https://binary.pensoft.net/fig/39012910.3897/zookeys.919.39642.figures577-5896D41C33E-DB8B5356-A8A0-2EFA2AEC1DAFFigures 577–589. Aleiodes periscelis (Reinhard), ♀, Czech Republic, Pisek 577 fore wing 578 hind wing 579 mesosoma lateral 580 mesosoma dorsal 581 metasoma dorsal 582 fore femur lateral 583 hind femur lateral 584 head anterior 585 head dorsal 586 head lateral 587 base of antenna 588 apex of antenna 589 outer middle tarsal claw. https://binary.pensoft.net/fig/39013010.3897/zookeys.919.39642.figure59013547743-5004-59EFB602-617196095EE7Figure 590. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov, habitus lateral. https://binary.pensoft.net/fig/39013110.3897/zookeys.919.39642.figures591-6032B48EACD-704C-

52D4-AC60-E77539DD4B55Figures 591–603. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov 591 wings 592 mesosoma lateral 593 mesosoma dorsal 594 1st –5th metasomal tergites dorsal 595 fore femur lateral 596 hind femur lateral 597 hind tarsus lateral 598 head anterior 599 head dorsal 600 head lateral 601 base of antenna 602 apex of antenna 603 inner hind tarsal claw. https://binary.pensoft.net/fig/39013210.3897/zookeys.919.39642.figures604-6079537E7F1-FFB05D78-BF12-CD5E808804DFFigures 604–607. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 604 habitus lateral 605 mummy of Acronicta psi (Linnaeus) in winter 606 ovipositor sheath lateral, 607 mummy of Acronicta psi (Linnaeus) after emergence of parasitoid. https://binary.pensoft.net/fig/39013310.3897/zookeys.919.39642.figures608-621B8F477C8-D2B15F9C-9F6C-875817AB1D36Figures 608–621. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 608 fore wing 609 hind wing 610 mesosoma lateral 611 mesosoma dorsal 612 1st–3rd metasomal tergites dorsal 613 fore femur lateral 614 hind femur lateral 615 hind tarsus lateral 616 head anterior 617 head dorsal 618 head lateral 619 base of antenna 620 apex of antenna 621 outer hind tarsal claw. https://binary.pensoft.net/fig/39013410.3897/zookeys.919.39642.figures622-6268F16822F-43435AEB-A126-BEB3F9F6DE2EFigures 622–626. Aleiodes pulchripes Wesmael, ♂, England (ex Acronicta culture) 622 habitus lateral 623 outer hind claw lateral 624 hind tibial spurs and basitarsus lateral 625 metasoma dorsal 626 4th–7th metasomal tergites lateral. https://binary.pensoft.net/fig/39013510.3897/zookeys.919.39642.figures627-62857583F6D-4E6B5DCB-9270-AFF7110AC143Figures 627, 628. Aleiodes quadrum (Tobias), ♀, France, Accons 627 habitus lateral 628 ovipositor sheath lateral. https://binary.pensoft.net/fig/39013610.3897/zookeys.919.39642.figures629-64198FD5499-CFC95FEB-9444-C0F118BB3D57Figures 629–641. Aleiodes quadrum (Tobias), ♀, France, Accons 629 wings 630 mesosoma lateral 631 mesosoma dorsal 632 metasoma dorsal 633 fore femur lateral 634 hind femur lateral 635 outer hind tarsal claw 636 head anterior 637 head dorsal 638 head lateral 639 base of antenna 640 apex of antenna 641 antenna. https://binary.pensoft.net/fig/39013710.3897/zookeys.919.39642.figure642511FE234-8F0F-5CF18E5E-35FD5BE1957CFigure 642. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo, habitus lateral. https://binary.pensoft.net/fig/39013810.3897/zookeys.919.39642.figures643-65116E990E9-A70B555D-9335-502C10F3FAF2Figures 643–651. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo 643 wings 644 1st–3rd metasomal tergites dorsal 645 3rd–7th metasomal tergites lateral 646 id. dorsal 647 basal antennal segments 648 head anterior 649 head dorsal 650 head lateral 651 inner hind tarsal claw. https://binary.pensoft.net/fig/39013910.3897/zookeys.919.39642.figures652-653DE32596F-2DB15EB0-B66C-444B6902D4D9Figures 652, 653. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 652 habitus lateral 653 ovipositor sheath lateral. https://binary.pensoft.net/fig/39014010.3897/zookeys.919.39642.figures654-6662A1C084A-F6ED55F6-8EE1-6A74CA3E3D94Figures 654–666. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 654 fore wing 655 hind wing 656 mesosoma lateral 657 mesosoma dorsal 658 1st –3rd metasomal tergites dorsal 659 fore femur lateral 660 hind femur lateral 661 head anterior 662 head dorsal 663 head lateral 664 base of antenna 665 apex of antenna 666 inner hind tarsal claw. https://binary.pensoft.net/fig/39014110.3897/zookeys.919.39642.figures667-6696F715591-21295BE5-925F-6BD56577E9D1Figures 667–669. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 667 habitus lateral 668 ovipositor sheath lateral 669 mummy of Hoplodrina blanda (Denis & Schiffermüller). https://binary.pensoft.net/fig/39014210.3897/zookeys.919.39642.figures670-68159011668-ED535A41-A4D5-911A3B72DECEFigures 670–681. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 670 wings 671 mesosoma lateral 672 mesosoma dorsal 673 metasoma dorsal 674 fore femur lateral 675 hind femur lateral 676 head anterior 677 head dorsal 678 head lateral 679 base of antenna 680 apex of antenna 681 outer hind tarsal claw. https://binary.pensoft.net/fig/39014310.3897/zookeys.919.39642.figures682-685D1D0931E-B1F45F94-987C-FF7F38A7FA7AFigures 682–685. Aleiodes ruficornis (Herrich-Schäffer), ♂, U.K. (culture) 682 habitus lateral 683 apex of antenna 684 base of antenna 685 mummy of Hoplodrina octogenaria (Goeze). https://binary.pensoft.net/fig/39014410.3897/zookeys.919.39642.figures686-687171BD0B1-855A54F7-AF64-48254BEB2FB0Figures 686, 687. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 686 habitus lateral 687 ovipositor sheath lateral. https://binary.pensoft.net/fig/39014510.3897/zookeys.919.39642.figures688-700ED43702D-6573513A-AD98-E335A908BD3EFigures 688–700. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 688 fore wing 689 hind wing 690 mesosoma lateral 691 mesosoma dorsal 692 1st–3rd metasomal tergites dorsal 693 fore femur lateral 694 hind femur lateral 695 head anterior 696 head dorsal 697 head lateral 698 outer hind tarsal claw 699 base of antenna 700 apex of antenna. https://binary.pensoft.net/fig/39014710.3897/zookeys.919.39642.figures701-703199CB048-CA905EB9-92BE-3B3ED43EB59CFigures 701–703. Aleiodes rugulosus (Nees), ♀, U.K., culture 701 habitus lateral 702 ovipositor sheath lateral 703 mummy of Acronicta rumicis (Linnaeus). https://binary.pensoft.net/fig/39014810.3897/zookeys.919.39642.figures704-7167D86EEFB-350A5B10-AC56-D894121A288BFigures 704–716. Aleiodes rugulosus (Nees), ♀, U.K., culture 704 wings 705 mesosoma lateral 706 mesosoma dorsal 707 propodeum and 1st–3rd metasomal tergites dorsal 708 fore femur lateral 709 hind femur lateral 710 antenna 711 head anterior 712 head dorsal 713 head lateral 714 base of antenna 715 apex of antenna 716 outer hind tarsal claw. https://binary.pensoft.net/fig/39014910.3897/zookeys.919.39642.figures717-718FE6550A4-E2F6548E-950E-461004EA85F4Figures 717, 718. Aleiodes rugulosus (Nees), dark form, ♀, U.K., Meathop Moss 717 habitus lateral 718 mummy of Acronicta menyanthidis (Esper). https://binary.pensoft.net/fig/39015010.3897/zookeys.919.39642.figure719CA2CA434-9154-5F36B825-181DBFE6D317Figure 719. Aleiodes rugulosus (Nees), Netherlands, Ede, mummy of Acronicta auricoma (Denis & Schiffermüller). Photograph: J. Voogd. https://binary.pensoft.net/fig/39015110.3897/zookeys.919.39642.figures720-727235DC654-7A6E55B8-94B2-5EA701FC2C96Figures 720–727. Aleiodes rugulosus (Nees), ♂, Hungary (Halastó) 720 habitus lateral 721 metasoma lateral 722 hind tibial spurs and basitarsus lateral 723 base of antenna 724 metasoma dorsal 725 fore femur lateral 726 head dorsal 727 hind femur lateral. https://binary.pensoft.net/fig/39015210.3897/zookeys.919.39642.figures728-729604DBFE4-70245A1A-AA0D-6FC00C223E2BFigures 728, 729. Aleiodes schewyrewi (Kokujev) , ♀, holotype var. zaydamensis (Kokujev) 728 habitus lateral 729 ovipositor sheath lateral. https://binary.pensoft.net/fig/39015310.3897/zookeys.919.39642.figures730-7421B6BBA14-89C855F2-96AF-779159DFD13EFigures 730–742. Aleiodes schewyrewi (Kokujev) , ♀, holotype var. zaydamensis (Kokujev) 730 fore wing 731 hind wing 732 mesosoma lateral 733 mesosoma dorsal 734 metasoma dorsal 735 fore femur lateral 736 hind femur lateral 737 head anterior 738 head dorsal 739 head lateral 740 outer hind tarsal claw 741 base of antenna 742 apex of antenna. https://binary.pensoft.net/fig/39015410.3897/zookeys.919.39642.figure7434BD9BC03-28D8-549A82D8-E91877305D8AFigure 743. Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, habitus lateral. https://binary.pensoft.net/fig/39015510.3897/zookeys.919.39642.figures744-757247A8626-09A05325-8C7F-C86C2780979BFigures 744–757.

Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, but 756 ♀, Ukraine 744 fore wing 745 hind wing 746 mesosoma lateral 747 mesosoma dorsal 748 metasoma dorsal 749 fore femur lateral 750 hind femur lateral 751 head anterior 752 head dorsal 753 head lateral 754 base of antenna 755 apex of antenna 756 outer hind tarsal claw 757 antenna. https://binary.pensoft.net/fig/39015610.3897/zookeys.919.39642.figures758-759590006CE-F27D5D9E-965F-E61357DFF390Figures 758, 759. Aleiodes sibiricus (Kokujev), ♀, lectotype 758 habitus lateral 759 head anterior. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39015710.3897/zookeys.919.39642.figures760-762C0C37212-54365166-931C-FA09E12DE0EAFigures 760–762. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves, but 762 Germany, Freiburg 760 habitus lateral 761 ovipositor sheath lateral 762 mummy of Noctua comes Hübner. https://binary.pensoft.net/fig/39015810.3897/zookeys.919.39642.figures763-776C50BCA38-1D445B02-911D-506AD23D5856Figures 763–776. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves 763 fore wing 764 hind wing 765 mesosoma lateral 766 mesosoma dorsal 767 metasoma dorsal 768 fore femur lateral 769 hind femur lateral 770 head anterior 771 head dorsal 772 head lateral 773 base of antenna 774 apex of antenna 775 outer hind tarsal claw 776 antenna. https://binary.pensoft.net/fig/39015910.3897/zookeys.919.39642.figures777-7788ED3635B-E3CF5FF0-AA73-BF600FA0FBFEFigures 777, 778. Aleiodes turcicus sp. nov., ♀, holotype 777 habitus lateral 778 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016010.3897/zookeys.919.39642.figures779-791D9C94A3B-2D755D96-A9CE-74C2BD6EED69Figures 779–791. Aleiodes turcicus sp. nov., ♀, holotype 779 fore wing 780 hind wing 781 mesosoma lateral 782 mesosoma dorsal 783 metasoma dorsal 784 fore femur lateral 785 hind femur lateral 786 head anterior 787 head dorsal 788 head lateral 789 base of antenna 790 apex of antenna 791 outer hind tarsal claw. https://binary.pensoft.net/fig/39016110.3897/zookeys.919.39642.figures792-794BA2D35E3-C8735AD5-95B2-CE577D30A360Figures 792–794. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 792 habitus lateral 793 ovipositor sheath lateral 794 mummy of Apamea unanimis Hübner. https://binary.pensoft.net/fig/39016210.3897/zookeys.919.39642.figures795-80791FB4AA9-CBFA5C83-81A5-87C52817F0C0Figures 795–807. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 795 wings 796 mesosoma lateral 797 mesosoma dorsal 798 metasoma dorsal 799 fore femur lateral 800 hind femur lateral 801 antenna 802 head anterior 803 head dorsal 804 head lateral 805 base of antenna 806 apex of antenna 807 inner hind tarsal claw lateral. https://binary.pensoft.net/fig/39016310.3897/zookeys.919.39642.figures808-81211FC92CF-95685FE1-B32F-5BA260E837E8Figures 808–812. Aleiodes unipunctator (Thunberg), ♂, Scotland, Edinburgh, but 808 Sweden, Särö-Hamra 808 habitus lateral 809 apex of metasoma dorsal 810 apex of metasoma lateral 811 head dorsal 812 base of antenna. https://binary.pensoft.net/fig/39016410.3897/zookeys.919.39642.figures813-8148142EA8E-5E255EDB-A8C9-3B9CCCA957A0Figures 813, 814. Aleiodes venustulus (Kokujev), ♀, holotype 813 habitus lateral 814 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016510.3897/zookeys.919.39642.figures815-82628ABB556-32445A75-A7A0-7CC856C3A26EFigures 815–826. Aleiodes venustulus (Kokujev), ♀, holotype 815 fore wing 816 hind wing 817 mesosoma lateral 818 mesosoma dorsal 819 metasoma dorsal 820 fore femur lateral 821 hind femur lateral 822 head anterior 823 head dorsal 824 head lateral 825 inner hind tarsal claw lateral 826 base of antenna. https://binary.pensoft.net/fig/39016610.3897/zookeys.919.39642.figures827-829A061B54B-32E25484-A351-D78AD200A589Figures 827–829. Aleiodes zwakhalsi sp. nov., ♀, holotype 827 habitus lateral 828 detail of fore wing 829 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016710.3897/zookeys.919.39642.figures830-8404733616C-9C135269-9486-8C6D9C5DD7EBFigures 830–840. Aleiodes zwakhalsi sp. nov., ♀, holotype 830 wings 831 mesosoma lateral 832 mesosoma dorsal 833 metasoma dorsal 834 fore femur lateral 835 hind femur lateral 836 base of antenna 837 head anterior 838 head dorsal 839 head lateral 840 outer hind tarsal claw lateral. https://binary.pensoft.net/fig/390168ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.MarshPM (1979) Braconidae. Aphidiidae. Hybrizontidae. In: KrombeinKVHurdJr PDSmithDRBurksBD (Eds) Catalog of Hymenoptera in America north of Mexico.Smithsonian Institution Press Washington, 144– 313.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1985b) Braconidae (Hymenoptera) from Korea. VII.Acta Zoologica Hungarica31: 341–365.ShawMRHuddlestonT (1991) Classification and biology of Braconid wasps (Hymenoptera: Braconidae).Handbooks for the Identification of British Insects7(11): 1–126.van AchterbergC (1991) Revision of the genera of the Afrotropical and W. Palaearctic Rogadinae Foerster (Hymenoptera: Braconidae).Zoolologische Verhandelingen, Leiden273: 1–102.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xZaldívarRiverónAShawMRSáezAGMoriMBelokobylskijSAShawSRQuickeDLJ (2008) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Biology 8: 329. https://doi.org/10.1186/1471-2148-8-329van AchterbergCShawMR (2016) 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.ZooKeys639: 1–164.TobiasVI (1971) Review of the Braconidae (Hymenoptera) of the U.S.S.R.Trudy Vsesoyuznogo Entomologicheskogo Obshchestva54: 156–268. [in Russian] [Translation (1975). 164 pp, New Delhi, India.]TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]FortierJShermanC (2008) A new Aleiodes (= Tetrasphaeropyx) (= A. pilosus group), with a host record from Missouri, U.S.A., biological remarks, and previous type depository corrections.Entomological News119(5): 445–453. https://doi.org/10.3157/0013-872X-119.5.445ShawSR (1993) Systematic status of Eucystomastax Brues and characterization of the Neotropical species (Hymenoptera: Braconidae, Rogadinae).Journal of Hymenoptera Research2: 1–11.ButcherBAQuickeDLJ (2011) Revision of Aleiodes (Hemigyroneuron) parasitic wasps (Hymenoptera: Braconidae: Rogadinae) with reappraisal of subgeneric limits, descriptions of new species and phylogenetic analysis.Journal of Natural History45: 1403–1476. https://doi.org/10.1080/00222933.2011.557557ButcherBAQuickeDLJ (2015) First record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) from the Arabian Peninsula: description of new species with remarkable wing venation convergence to Gyroneuron and Gyroneuronella.Zootaxa4033(2): 275–279. https://doi.org/10.11646/zootaxa.4033.2.7ShawSRMarshPMFortierJC (1998) Revision of North American Aleiodes Wesmael (Part 2): the apicalis (Brullé) species-group in the New World (Hymenoptera: Braconidae, Rogadinae) Journal of Hymenoptera Research 7: 62– 73.QuickeDLJShawMR (2005) First host record for the rogadine genus Pholichora van Achterberg (Hymenoptera: Braconidae) with description of a new species and notes on convergent wing venation features.Journal of Natural History39(7): 531–537. https://doi.org/10.1080/00222930410001708678ButcherBASmithMASharkeyMJQuickeDLJ (2012) A turbo-taxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae: Rogadinae) based largely on COI barcoded specimens, with rapid descriptions of 179 new species.Zootaxa3457: 1–232.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.

2. XML treatment for Aleiodes apicalis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642D8E9E2A4-D2C7-5C97-B629DB1DC2567902 Aleiodes apicalis group Apical half of marginal cell of hind wing distinctly widened, its maximum width 1.6 × its width near hamuli or wider (Fig. 27) and vein r of fore wing shorter than vein 3-SR (Figs 180, 608), if marginal cell largely parallel-sided (Figs 506, 609, 704) then tarsal claws comparatively robust and with often blackish pecten (Figs 517, 621, 716) or brachypterous (Fig. 390); occipital carina usually reduced ventrally, not reaching hypostomal carina (Figs 600, 663, 713, 788); mesopleuron partly

smooth (at least between punctures), but largely densely sculptured in both sexes of A. hemipterus and A. krulikowskii, as well in some males of A. ruficornis and allies; lateral carina of scutellum absent or if present then weakly developed and lunula wide (Fig. 508); 2nd metasomal tergite with distinct and smooth triangular area medio-basally (Fig. 509); ovipositor sheath distinctly setose apically (Fig. 483); males are often darker than females, most extremely so in A. arnoldii, A. carbonarius and A. carbonaroides; brachypterous specimens of Aleiodes are included in this group. All species of the A. apicalis group for which host data exist are parasitoids of Noctuidae. However, the putatively more basal A. fortipes belonging to the Hemigyroneuron clade (see below) is a parasitoid of Geometridae. Also, only A. fortipes and A. sibiricus are known to parasitise hosts only in spring although these hosts would have been available in autumn of the previous year. Possibly

others in the A. apicalis group will be found to do this too, and we consider the habit putatively as ancestral, in contrast with the more derived A. circumscriptus and A. bicolor groups in which species using hosts that overwinter as larvae invariably (as far as known) parasitise the host in the autumn and overwinter as a young larva inside it. While we have no host data for a disappointingly large number of species of the A. apicalis group, the form of the clypeus may give important clues as to the site at which host mummification occurs, as those species in which mummification is known to take place in open situations (e.g., on a twig or in a leaf curl) invariably have a relatively small hypoclypeal depression and the clypeal margin blunt (A. apicalis, A. aterrimus, A. fortipes, A. nobilis, A. pulchripes, A. rugulosus) while species known to cause their hosts to mummify in concealed situations tend to have the hypoclypeal opening wider and the margin sharper (e.g., A. cruentus, A. dissector, A. ruficornis, A. sibiricus, A. unipunctator). According to the 28S + COI analysis by ZaldÃvar-Riverón et al. (2008) the following former subgenera or genera belong to this group: Chelonorhogas Enderlein, [1st Sept.] 1912 (worldwide), Eucystomastax Brues, [(end of?) Sept.] 1912 (Neotropical group with 2nd and 3rd maxillary palp segments enlarged), Hemigyroneuron Baker, 1917 (Old World group with distal half of subbasal cell of fore wing modified and glabrous), and Dimorphomastax Shenefelt, 1979 (males of this monotypic Neotropical group have a large curved tooth near the base of the mandible (an outgrowth of the condylar carina) and the hind tibial spurs are blunt apically; females have the tooth smaller and triangular, and the hind tibial spurs are acute). Butcher et al. (2012) indicate in their cladogram based on the analysis of COI sequences that A. fortipes (Reinhard) forms together with Hemigyroneuron Baker and Arcaleiodes Chen & He the most basal clade of Aleiodes Wesmael, and it is noteworthy that all known hosts of this clade are Geometridae (see species entry for A. fortipes). According to the same analysis the A. apicalis group consists of three clades: (i) the Hemigyroneuron clade (see above; likely also includes A. caucasicus (Tobias)), (ii) the A. rugulosus clade (including the Asian group with modified pronotum), and (iii) the A. gasterator clade. Since we do not have the COI sequences of all species, we unite these three clades in the A. apicalis group to allow identification based on their morphology. 10.3897/zookeys.919.39642.figures26-3779D0915E-1C6A-5272-A38E-E5E71FA2DAC4Figures 26–37. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River 26 fore wing 27 hind wing 28 mesosoma lateral 29 mesosoma dorsal 30 metasoma dorsal 31 fore femur lateral 32 hind femur lateral 33 base of antenna 34 head anterior 35 head dorsal 36 head lateral 37 inner hind tarsal claw. https://binary.pensoft.net/fig/39005110.3897/zookeys.919.39642.figures180-1915548F088-03BA5D78-BB9B-82B3B3E62130Figures 180–191. Aleiodes coriaceus sp. nov., ♀, holotype 180 wings 181 mesosoma lateral 182 mesosoma dorsal 183 propodeum and 1st–3rd metasomal tergites dorsal 184 fore femur lateral 185 hind femur lateral 186 head anterior 187 head dorsal 188 head lateral 189 outer hind tarsal claw 190 base of antenna 191 apex of antenna. https://binary.pensoft.net/fig/39007410.3897/zookeys.919.39642.figures608-621B8F477C8-D2B15F9C-9F6C-875817AB1D36Figures 608–621. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 608 fore wing 609 hind wing 610 mesosoma lateral 611 mesosoma dorsal 612 1st–3rd metasomal tergites dorsal 613 fore femur lateral 614 hind femur lateral 615 hind tarsus lateral 616 head anterior 617 head dorsal 618 head lateral 619 base of antenna 620 apex of antenna 621 outer hind tarsal claw. https://binary.pensoft.net/fig/39013410.3897/zookeys.919.39642.figures506-518677289DC-BD215788-81C9-FB618A370D45Figures 506–518.

Aleiodes nigrifemur sp. nov. ♀, holotype 506 wings 507 mesosoma lateral 508 mesosoma dorsal 509 1st–3rd metasomal tergites dorsal 510 fore femur lateral 511 hind femur lateral 512 base of antenna 513 head anterior 514 head dorsal 515 head lateral 516 apex of antenna 517 outer hind tarsal claw 518 antennae. https://binary.pensoft.net/fig/39012010.3897/zookeys.919.39642.figures704-7167D86EEFB-350A5B10-AC56-D894121A288BFigures 704–716. Aleiodes rugulosus (Nees), ♀, U.K., culture 704 wings 705 mesosoma lateral 706 mesosoma dorsal 707 propodeum and 1st–3rd metasomal tergites dorsal 708 fore femur lateral 709 hind femur lateral 710 antenna 711 head anterior 712 head dorsal 713 head lateral 714 base of antenna 715 apex of antenna 716 outer hind tarsal claw. https://binary.pensoft.net/fig/39014910.3897/zookeys.919.39642.figures390-3996F7E68A7-8F39543C-84C4-CAE338A06F69Figures 390–399. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 390 wings 391 mesosoma lateral 392 mesosoma dorsal 393 metasoma dorsal 394 fore femur lateral 395 hind femur lateral 396 head anterior 397 head dorsal 398 head lateral 399 outer hind tarsal claw. https://binary.pensoft.net/fig/39010310.3897/zookeys.919.39642.figures591-6032B48EACD-704C52D4-AC60-E77539DD4B55Figures 591–603. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov 591 wings 592 mesosoma lateral 593 mesosoma dorsal 594 1st –5th metasomal tergites dorsal 595 fore femur lateral 596 hind femur lateral 597 hind tarsus lateral 598 head anterior 599 head dorsal 600 head lateral 601 base of antenna 602 apex of antenna 603 inner hind tarsal claw. https://binary.pensoft.net/fig/39013210.3897/zookeys.919.39642.figures654-6662A1C084A-F6ED55F6-8EE1-6A74CA3E3D94Figures 654–666. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 654 fore wing 655 hind wing 656 mesosoma lateral 657 mesosoma dorsal 658 1st –3rd metasomal tergites dorsal 659 fore femur lateral 660 hind femur lateral 661 head anterior 662 head dorsal 663 head lateral 664 base of antenna 665 apex of antenna 666 inner hind tarsal claw. https://binary.pensoft.net/fig/39014110.3897/zookeys.919.39642.figures779-791D9C94A3B-2D755D96-A9CE-74C2BD6EED69Figures 779–791. Aleiodes turcicus sp. nov., ♀, holotype 779 fore wing 780 hind wing 781 mesosoma lateral 782 mesosoma dorsal 783 metasoma dorsal 784 fore femur lateral 785 hind femur lateral 786 head anterior 787 head dorsal 788 head lateral 789 base of antenna 790 apex of antenna 791 outer hind tarsal claw. https://binary.pensoft.net/fig/39016110.3897/zookeys.919.39642.figures482-483719368AD-B6D051BF-BD6C-AA524F5F9B58Figures 482, 483. Aleiodes morio (Reinhard), ♀, Hungary, Budapest 482 habitus lateral 483 ovipositor sheath lateral. https://binary.pensoft.net/fig/390115ZaldÃvarRiverónAShawMRSáezAGMoriMBelokobylskijSAShawSRQuickeDLJ (2008) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Biology 8: 329. https://doi.org/10.1186/1471-2148-8-329ButcherBASmithMASharkeyMJQuickeDLJ (2012) A turbo-taxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae: Rogadinae) based largely on COI barcoded specimens, with rapid descriptions of 179 new species.Zootaxa3457: 1–232.

3. XML treatment for Aleiodes aestuosus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396425C91497F-3402-56BB-944919348F7F57ED Aleiodes aestuosus (Reinhard, 1863)Figs 22–25, 26–37 Rogas aestuosusReinhard, 1863: 265; Shenefelt 1975: 1216–1217; Zaykov 1980a: 112; Tobias 1976: 84, 1986: 78 (transl.: 129); Kotenko 1992: 96 [examined]. Rhogas aestuosus; Fahringer 1931: 232–234. Aleiodes (Neorhogas) aestuosus; Papp 1985a: 152, 1989: 52, 1990: 90, 1991a: 67–68. Aleiodes (Chelonorhogas) aestuosus; Chen and He 1997: 38; He et al. 2000: 665; Belokobylskij 2000: 26; Chen et al. 2003: 211; Papp 2012: 187; Farahani et al. 2015: 238–240. Aleiodes aestuosus; Fortier and Shaw 1999: 230; Zaldívar-Riverón et al. 2004: 234. Rhogas (Rhogas) aestuosus var. desertusTelenga, 1941: 152–153, 404 (not Rhogas (R.) desertus Telenga, 1941). Holotype, ♀ (MNHN), “Cipro [= Cyprus]”, “Muséum Paris, 1867, coll. O. Sichel”, “Rogas aestuosus Rhd.”. Albania, Bulgaria, Cyprus, Greece, Russia, Turkey, Tunisia, [Azerbaijan, Georgia, Iran, Iraq, Israel, Jordan, Syria, Turkmenistan, Uzbekistan]. Specimens in ZJUH, BZL, CNC, HSC, MRC, MTMA, NMS, RMNH, ZSSM, ZISP. Distributed principally in Asia Minor, extending to Cyprus where it has been collected plentifully. Only single specimens examined from Albania (MTMA) and mainland Greece (BZL), but in North Africa it apparently extends westwards to Tunisia (one specimen in BZL). MRS004 (Turkey). Collected March–July, often at light, but it is not clear how many generations are represented nor how the winter is passed. Reared from Noctuidae: Heliothis peltigera (Denis & Schiffermüller) (4 [1 CNC/Iraq, 1 ZISP (with mummy)/Uzbekistan, 2 MTMA/Iraq]), Sesamia sp. (2 [ZJUH/Iran]). This indicates a host range of both endophagous and exophagous larvae, but the individuals purporting to be from Sesamia are labelled [no doubt incorrectly] “ex pupa” and lack mummies, suggesting that they may have resulted from substrate rearings (presumably from stems of crop species of Poaceae, inside which Sesamia larvae feed and pupate) rather than from isolated hosts, with a consequent reduction in the reliability of the host determination and suspicion that mummies of other hosts could have been overlooked on the stems (see also remark under A. apicalis). On the other hand, the large hypoclypeal depression and somewhat protruding clypeus does indicate that A. aestuosus adults are equipped to chew their way out of mummies made in concealed sites. The hosts given above are regular crop pests, but the paucity of reared material examined may suggest that A. aestuosus is not especially associated with cultivated habitats. The single mummy seen (Fig. 25) is rather elongate, scarcely arched, and the cocoon occupies most of the host abdomen. It has the appearance of not being securely stuck to the substrate. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 34); clypeus rather protruding anteriorly and rather thick ventrally (Fig. 36); head brownish yellow; vertex finely punctate; lateral lobes of mesoscutum sparsely and finely punctate, with wide smooth interspaces; precoxal sulcus absent, area only sparsely finely punctate or smooth; 1-CU1 of fore wing subequal to vein 2-CU1 (Fig. 26); hind tarsal claws with brownish pecten (Fig. 37); only apex of hind tibia dark brown; metasoma of ♀ completely yellowish and distinctly depressed subapically, 1st tergite partly and 4th–6th tergites of ♂ often blackish. Sometimes entire body (including propodeum and 1st metasomal tergite) yellowish (“var. desertus”). Redescribed ♀ (RMNH) from Turkey (Icil). Length of fore wing 6.8 mm, of body 8.3 mm. Head. Antennal segments of ♀ 52, length of antenna 1.1 × fore wing, its subapical segments approx. as long as wide; frons with irregular curved rugae, shiny, and rugose behind antennal sockets; OOL 2.4 × diameter of posterior ocellus, and finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate, shiny; clypeus short, coarsely and densely punctate; ventral margin of clypeus thick and rather protruding forwards (Fig. 36); width of hypoclypeal depression 0.65 × minimum width of face (Fig. 34); length of eye 0.8 × temple in dorsal view (Fig. 35); vertex behind stemmaticum sparsely punctate; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely smooth, shiny, sparsely and finely punctate; prepectal carina medium-sized, reaching anterior border; precoxal area of mesopleuron and metapleuron remotely punctate, interspaces much wider than diameter of punctures, shiny; mesopleuron above precoxal area (except speculum) sparsely punctate; scutellum slightly convex, remotely punctate and evenly rounded laterally, no carina; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete, but irregular posteriorly, without tubercles. Wings. Fore wing: r 0.4 × 3-SR (Fig. 26); 1-CU1 horizontal, nearly as long as (0.9 x) 2-CU1; r-m 0.9 × 2-SR, and 0.7 × 3-SR; second submarginal cell medium-sized (Fig. 26); cu-a vertical, not parallel with CU1b, straight; 1-M rather curved posteriorly. Hind wing: marginal cell gradually and evenly widened, its apical width 2.3 × width at level of hamuli (Fig. 27); 2-SC+R shortly longitudinal; m-cu distinct; M+CU:1-M = 23:19; 1r-m 0.7 × 1-M.

Legs. Tarsal claws subpectinate, with four brown medium-sized pectinal bristles (Fig. 37); hind coxa remotely punctate, shiny; hind trochantellus robust; length of hind femur and basitarsus 3.0 and 3.5 × their width, respectively; length of inner hind spur 0.55 × hind basitarsus; hind tibia slender (Fig. 23). Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites coarsely and densely rugose, robust, with distinct median carina; medio-basal area of 2nd tergite wide and short; 2nd suture deep medially and shallow laterally; basal half of 3rd tergite finely rugose, remainder of metasoma largely smooth, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with medium-sized setae and apically rounded (Fig. 22). Colour. Brownish yellow; antenna, mesosternum (except anteriorly) and mesopleuron (except anteriorly and dorsally), metapleuron, propodeum, ovipositor sheath and stemmaticum black; hind tibia (except apically) pale yellowish; apices of femora (dorsally) and tibiae, palpi, tarsi (except basally), veins and pterostigma dark brown; wing membrane rather infuscate. Variation. Size of eyes and ocelli rather variable. Mesopleuron, mesosternum, metapleuron and propodeum brownish yellow or black; 1st tergite entirely brownish yellow or with dark brown patch basally; in desert areas body can be wholly orange. Antennal segments: ♀ 49 (1), 50 (3), 51 (9), 52 (13), 53 (10), 54 (3), 55 (5), 56 (2); ♂ 51 (10), 52 (11), 53 (5), 54 (4), 55 (3), 56 (1). The two sexes have comparable numbers of antennal segments. Apical tergites of ♂ type 3 and fringe moderately strong; inner hind tibial spur 0.50 × as long as hind basitarsus. Albania, Azerbaijan, Bulgaria, Cyprus, Georgia, *Greece, Iran, *Iraq, Israel, *Jordan, Russia, Syria, Turkey, Tunisia, *Turkmenistan, Uzbekistan. 10.3897/zookeys.919.39642.figures22-2591C13739-EA90-5B88-BA5E-C363154C28E1Figures 22– 25. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River, but 25 from Uzbekistan, Qamashi 22 ovipositor sheath lateral 23 habitus lateral 24 apex of antenna 25 mummy of Heliothis peltigera (Denis & Schiffermüller). https://binary.pensoft.net/fig/39005010.3897/zookeys.919.39642.figures26-3779D0915E-1C6A5272-A38E-E5E71FA2DAC4Figures 26–37. Aleiodes aestuosus (Reinhard), ♀, Cyprus, Yermasoyja River 26 fore wing 27 hind wing 28 mesosoma lateral 29 mesosoma dorsal 30 metasoma dorsal 31 fore femur lateral 32 hind femur lateral 33 base of antenna 34 head anterior 35 head dorsal 36 head lateral 37 inner hind tarsal claw. https://binary.pensoft.net/fig/390051ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980a) Unknown genera and species of Braconidae for the fauna of Bulgaria. Travaux scientifiques d’Université de Plovdiv “Paissi Hilendarski”, Biologie 15(4) [1977]: 111–115. [in Bulgarian]TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]KotenkoAG (1992) A contribution to the fauna of Braconidae (Hymenoptera) of Dauria. In: Amirkhanov AM (Ed.) Insects of Dauria and adjacent territories: 94–107.Izdatelstvo Tsentralnoi Nauchno issledovatelskoi laboratorii okhotnichego khozyaistva i zapovednikov, Moskva, 141 pp.FahringerJ (1931) Opuscula braconologica. Band 3. Palaearktischen Region. Opuscula braconologica 3: 161–240. Wagner, Wien.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1990) Braconidae (Hymenoptera) from Tunisia, 3.Folia Entomologica Hungarica51: 89–96.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.ChenX-XHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.ChenXXPiaoM-HWhitfieldJBHeJ-H (2003) A molecular phylogeny of the subfamily Rogadinae (Hymenoptera: Braconidae) based on the D2 variable region of 28S ribosomal RNA.Acta Entomologica Sinica46(2): 209–217.PappJ (2012) A contribution to the Braconid fauna of Israel (Hymenoptera: Braconidae), 3. Israel Journal of Entomology 41–42: 165– 219.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.x

4. XML treatment for Aleiodes agilis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis

groupZookeys1632020919125910.3897/zookeys.919.39642AE4E13C8-7737-5FFB-BC339181B93B6C14 Aleiodes agilis (Telenga, 1941)Figs 38–39, 40–49 Rhogas (Rhogas) agilisTelenga, 1941: 165–166, 417. Rogas agilis; Shenefelt 1975: 1217. Rogas (Rogas) agilis; Tobias, 1976: 83,

1986: 76 (transl. 122, 124) (lectotype designation). Aleiodes agilis; Fortier and Shaw 1999: 230. Aleiodes (Chelonorhogas) agilis; Ma et al. 2002: 98; Farahani et al. 2015: 240. Rhogas desertus var. armenicaTelenga, 1959: 85; Tobias, 1976: 83 (as synonym of A. agilis (Telenga, 1941)), 1986: 76 (transl. 122, 124; id.). Lectotype, ♀ (ZISP; examined via photos), “Persiya [= Iran], Tavriz, 21.iii.[19]14, Andrievskij”, “Rhogas agilis sp. n., N. Telenga det.”, “Syntypus agilis Tel.”, “Lectotypus Rogas agilis Tel., design. Tobias, 1980”; paralectotype, ♀ (ZISP; id.), “Armenia, pr. Eriwan [= Yerevan], A. Schelkovnikow / Ragakag, 19.iii.[19]25”, “Paralectotypus Rhogas agilis Telenga, design. Tobias, 1986”. In the original description the latter date is incorrectly cited as 24.vii.1925. None. Unknown. It appears to fly very early in the year (March). Maximum width of hypoclypeal depression approx. 0.8 × minimum width of face; anterior part of clypeus very narrow (Fig. 46); OOL 1.0–1.3 × diameter of posterior ocellus and coarsely remotely punctate with some weak rugulosity; head and mesosoma (except pronotal side partly and mesoscutum medio-posteriorly and laterally) blackish; mandible massive triangular, coarsely punctate and with thick ventral lamella (Fig. 46); face largely transversely rugose and conspicuously whitish setose; frons rugose and shiny; vertex and temple coarsely remotely punctate and shiny; area of precoxal sulcus (but posteriorly superficially) distinctly rugose; lateral lobes of mesoscutum largely smooth (anteriorly becoming densely punctate and somewhat rugose), whitish setose and with satin sheen, middle lobe distinctly punctate; basal half of wings (except anteriorly) largely glabrous and remainder of wing inconspicuously setose; vein r of fore wing approx. 0.6 × vein 3-SR (Fig. 40); vein 1-CU1 0.2–0.3 × as long as 2-CU1, narrow and subhorizontal; tarsal claws long, slender, hardly bent and simple (Fig. 44); 1st and base of 2nd tergite weakly longitudinally rugulose with some superficial punctures; metasoma dark brown but with yellow patches (Fig. 38), clypeus and antenna (except yellow scapus and pedicellus) yellowish brown; legs and palpi pale yellowish, but hind coxa and most of middle coxa dark brown. Paralectotype, ♀, length of fore wing 6.6 mm, of body 7.0 mm. Head. Antennal segments of ♀ 47, antenna as long as body and its subapical segments moderately slender; frons rugose, shiny; OOL 1.3 × diameter of posterior ocellus; OOL and vertex remotely punctate, with satin sheen, OOL also with some rugulae; anterior part of clypeus 9 × wider than high, coarsely punctate and rather convex; clypeus above lower level of eyes; ventral margin of clypeus thick and not protruding forwards (Fig. 48); width of hypoclypeal depression 0.8 × minimum width of face (Fig. 46); length of eye 1.8 × temple in dorsal view; vertex behind stemmaticum convex and sparsely punctate; length of malar space 0.19 × length of eye in lateral view; occipital carina nearly complete, fine and ventrally strongly curved; mandible massive triangular, coarsely punctate and with thick ventral lamella (Fig. 46). Mesosoma. Lateral lobes of mesoscutum largely smooth, with satin sheen and whitish setose, middle lobe distinctly punctate and setose; prepectal carina complete and lamelliform; precoxal area of mesopleuron widely rugose, but posterior 0.2 narrowly striate; mesopleuron largely weakly and sparsely punctate, shiny, but anteriorly becoming densely punctate and somewhat rugulose; scutellum largely smooth, with some punctures; propodeum evenly convex, finely rugose and with

medio-longitudinal carina, without tubercles. Wings. Fore wing: basal half largely glabrous; r 0.6 × 3-SR (Fig. 40); 1-CU1 subhorizontal, 0.25 × as long as 2-CU1; r-m 0.7 × as long as 3-SR; 2ndsubmarginal cell robust (Fig. 40); cu-a distinctly inclivous; 1-M weakly curved posteriorly. Hind wing: basal 0.4 of marginal cell slightly widened and distally strongly widened, its apical width approx. twice width at level of hamuli; 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 24:19; 1r-m 0.7 × 1-M. Legs. Tarsal claws slender, slightly curved and only setose (Fig. 44); hind coxa partly obliquely striate dorsally; tarsi slender, segments (except telotarsus) with long apical spiny bristles; length of hind femur and basitarsus 5.0 and 6.0 × their width, respectively; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite robust, as long as wide apically, distinctly narrowed anteriorly and rather flat posteriorly; 1st and 2nd tergites finely longitudinally striate-rugulose; medio-longitudinal carina of 1st and 2nd tergites indistinct; 2nd tergite 0.7 × longer than its basal width; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture shallow and narrow; 3rd tergite mainly smooth and with satin sheen; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath rather slender, with short setae and apically truncate (Fig. 39). Colour. Black; pronotal side largely yellowish brown; mesoscutum medio-posteriorly and posterolaterally partly chestnut brown; tegulae, clypeus and antenna (except yellow scapus and pedicellus) yellowish brown; mandible, legs and palpi pale yellowish, but hind coxa and most of middle coxa dark brown; metasoma dark brown but with yellow patches (Fig. 38); pterostigma brown medially and dark brown laterally; ovipositor sheath dark brown; veins of fore wing (but pale yellow in basal 0.2 of fore wing) brown; wing membrane hyaline. Armenia, Iran. Included in this revision, because it may occur in Turkey. 10.3897/zookeys.919.39642.figures38-3959BD7BC5-62D7-5AC8-B75B-1B987E9AD7A4Figures 38, 39. Aleiodes agilis (Telenga), ♀, paralectotype 38 habitus lateral 39 ovipositor sheath lateral. Photographs by K. Samartsev. https://binary.pensoft.net/fig/39005210.3897/zookeys.919.39642.figures40-49BD25508C-E43B52D7-84C7-A56A662B24A1Figures 40–49. Aleiodes agilis (Telenga), ♀, paralectotype, but 47 of lectotype 40 wings 41 mesosoma lateral 42 fore femur lateral 43 hind femur lateral 44 outer hind tarsal claw 45 fore leg 46 head anterior 47 head dorso-lateral 48 head lateral 49 base of antenna. Photographs by K. Samartsev. https://binary.pensoft.net/fig/390053ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.MaYChenX-XHeJ-H (2002) Newly recorded species of genus Aleiodes Wesmael (Hymenoptera: Braconidae) in China.Entomotaxonomia24(2): 98. [in Chinese]FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2

5. XML treatment for Aleiodes apicalis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642C39FE455-96C0-50F4-9FD184B0711DC24F Aleiodes apicalis (Brullé, 1832)Figs 50–53, 54–65, 66–71 Bracon apicalisBrullé, 1832: 381 [examined]. Rhogas apicalis; Fahringer 1932: 317–318. Rogas apicalis; Shenefelt 1975: 1218. Aleiodes apicalis; Shaw et al., 1998: 63; Fortier and Shaw 1999: 227; Zaldívar-Riverón et al. 2004: 234, 2008: figs 2–6. Aleiodes (Chelonorhogas) apicalis; Falco et al. 1997: 60; Farahani et al. 2015: 240–242; Abdolalizadeh et al. 2017: 36. Rogas reticulatorNees, 1834: 211; Shenefelt 1975 (as synonym of A. ductor); Papp 2005: 176 (id.). Syn. nov. Aleiodes reticulator; Papp, 1991a: 70 (as synonym of A. ductor). Rogas bicolorLucas, 1849: 336–337 (not Spinola 1808); Shenefelt 1975: 1219; Papp 1985a: 157 (lectotype designation), 2005: 176 (as synonym of A. ductor). Syn. nov. Rogas rufo-aterWollaston, 1858: 24; Shenefelt 1975: 1247; Papp 1990: 90 (as synonym of A. ductor) [examined]. Syn. nov. Rhogas rufoater; Fahringer 1934: 321. Rhogas bicolorinusFahringer, 1932: 318 (replacement name for Rogas bicolor Lucas). Syn. nov. Rhogas reticulator var. atripesCosta, 1884: 13; Papp 1990: 90 (as synonym of R. rufoater). Syn. nov. Rhogas ductor var. atripes; Fahringer 1932: 244. Aleiodes (Neorhogas) ductor var. atripes; Papp 1985a: 157. Rhogas similisSzépligeti, 1903: 114 (not Curtis 1834); Papp 1985a: 157–158 (lectotype designation and as synonym of A. ductor), 2005: 176 (id.). Syn. nov. Rhogas ductor var. similis; Fahringer 1932: 245. Rogas ductorauct. p.p.; Shenefelt, 1975: 1226–1227; Zaykov 1980a: 112; Tobias 1976: 85, 1986: 80 (transl.: 133); Samartsev and Belokobylskij 2013: 765. Aleiodes ductorauct. p.p.; Bergamasco et al. 1995: 5. Holotype of B. apicalis, ♂ (MNHN) “[Greece], Morée, Muséum Paris, Brullé 4187-33”, “Type”, “Bracon apicalis Brullé, Type”. Lectotype of R. similis, ♂ (MTMA) “[Hungary], Kecskemét, Szépligeti”, “Hym. Typ. No. 7021, Mus. Budapest”, “Lectotypus”, “Rhogas similis Szépl. 1903 ♂, Papp, 1984”, “Rhogas reticulator var. similis Sz., det. Szépligeti, 1906”, “Aleiodes ductor Thunbg., det. Papp, J., 1983”. Holotype of R. rufo-ater, ♂ (ZJUH) “[Portugal], Madeira, Wollaston, 55.7”, “Rogas rufo-ater, W.”, Type, H.T.”, “B.M. Type Hym., 3.c.241”. Albania, Austria, Bosnia & Herzegovina, Bulgaria, Croatia, Cyprus, Czech Republic, France (including Corsica), Germany, Greece (including Chios, Corfu, Crete, Lesbos, Rhodes), Hungary, Italy (including Sardinia, Sicily), Malta, Moldova, Montenegro, Morocco, North Macedonia, Portugal (including Madeira), Romania, Russia (including Dagestan), Serbia, Slovakia, Spain (including Mallorca and Canary Islands: Tenerife, Fuerteventura), Switzerland, Tunisia, Turkey, Ukraine, [Georgia, Kazakhstan, Oman, Iran, Iraq, Israel, Syria, Turkmenistan]. Specimens in ZJUH, BZL, CMIM, CNC, HSC, MRC, MSC, MTMA, NMS, RMNH, SDEI, UNS, ZISP, ZSSM. This is a mainly Mediterranean species, extending into Central Europe and West Asia, and one of the commonest species of the group in the Mediterranean area. One surprising female from Sweden (Skåne, Käseberga, MV light 17-vii-14.ix.2013, N. Ryrholm & C. Källander, in NMS) is presumed, like two British specimens (England, V.C. 3, S. Devon, Slapton Ley 7–14.vi.1932, H.St.J. Donisthorpe, in ZJUH; V.C. 22, Berkshire, Beale Park, 25–27.vii.2018, Rothamsted trap, in coll. A.C. Galsworthy destined for ZJUH) and one specimen from Netherlands (Lexmond, ZH, 10.viii.2004, C. Gielis in RMNH) to have been deposited there by winds from southern Europe or N. Africa rather than representing an established breeding population. Whether or not A. apicalis can eventually establish permanent populations, i.e., with winter survival, in these relatively northerly parts of Europe may depend on whether its host can do likewise. MRS008 (Turkey), MRS111 (Turkey), MRS112 (Turkey), MRS181 (Russia), MRS869 (Sweden). Time of flight varies according to harshness of summer. In its more temperate sites plurivoltine April-September(October), overwintering in the mummy, but in Cyprus (and presumably other places with extremely hot dry summers) it appears to be most active from autumn to spring (October–May), with a prolonged summer diapause (June–October or later) in the mummy (reared series ex “Plusia” in ZJUH and NMS, W.R. Ingram, six with mummification dates recorded in May or June and adult emergence in the following October–December, further specimens in the series have only one date, which is ambiguous). Reared from Noctuidae: Autographa gamma (Linnaeus) (6 [4 ZISP/Moldova, 1 HSC/Germany, 1 NMS/Malta]; J.L. Gregory, H. Schnee), indet. Plusiinae (14). There is no reason to suppose that the hosts recorded as indet. Plusiinae are anything except A. gamma. A further specimen labelled as ex Peribroma [sic] saucia is accompanied by a clearly Plusiinae mummy (Sicily, NMS). Also, one labelled as from Anarsia lineatella Zeller (Gelechiidae) (Ukraine, ZISP), but without a mummy and clearly in error on grounds of size alone. Another specimen labelled as “ex Sesamia pupa” (Iran, ZJUH) lacks its mummy but accompanies two individuals of A. aestuosus (q. v.) from the same source, and the remarks made under that species apply also to this record – but with the added objection that the small hypoclypeal opening and flat clypeus of A. apicalis strongly suggest that its hosts do not mummify in deep concealment. The mummy (Fig. 52) is of a pale chalky buff colour, and the cocoon occupies approx. abdominal segments 4–7 of the host larva. Several of the mummies examined, all of which seem to be penultimate instar, have been formed in a more or less curled leaf beneath a web (Fig. 53) that the host had been induced to spin before being mummified, and were weakly stuck to the substrate. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 60); antennal segments of ♀ 44–51 and flagellar segments moderately robust (Figs 64, 65); ventral margin of clypeus thick and obtuse apically and clypeus not protruding outwards (Fig. 62); vertex, mesoscutum, metapleuron and scutellum normally shiny and without dense granulation, at most with some superficial micro-sculpture; frons (and more or less vertex) with striae (Fig. 61) or rugae; scutellum largely smooth and shiny; mesopleuron largely smooth; vein 2-CU1 of fore wing approx. as long as vein 1-CU1 (Fig. 54); vein M+CU of hind wing distinctly longer than vein 1-M (Fig. 54); hind tarsal claws of ♀ with rather slender and brownish pecten (Fig. 63); basal half of hind tibia (largely) pale yellowish, or if black (var. rufoater) then also fore femur black; 3rd tergite (except basally) largely smooth; medially 4th–6th tergites of ♂ slightly concave and with dense band of

medium-sized setae (Figs 68, 69); head, mesoscutum and scutellum black; 2nd tergite yellowish or reddish. Redescribed ♀ (RMNH) from Hungary (Budapest), length of fore wing 5.1 mm, of body 6.7 mm. Head. Antennal segments of ♀ more than 40, but apical segments missing (length of antenna of ♀ from Lesbos 1.4 × fore wing and its subapical segments robust); frons with coarse curved rugae, shiny; OOL 1.5 × diameter of posterior ocellus, and distinctly striate; vertex transversely striate, rather weak; clypeus normal, punctulate and convex; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.3 × minimum width of face (Fig. 60); length of eye 1.6 × temple in dorsal view (Fig. 61); vertex behind stemmaticum transversely striate; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view; occipital carina complete, fine. Mesosoma. Mesoscutal lobes largely smooth, punctulate, shiny; prepectal carina complete, rather strong; precoxal area of mesopleuron largely smooth; mesopleuron above precoxal area weakly and sparsely punctate, especially posteriorly; scutellum largely smooth, with striae laterally; propodeum evenly convex, coarsely vermiculate-rugose, only anteriorly with median carina, without tubercles. Wings. Fore wing: r 0.6 × 3-SR (Fig. 54); 1-CU1 horizontal, equal to or slightly longer than 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell comparatively short (Fig. 54); cu-a vertical, slightly curved posteriorly; 1-M straight posteriorly. Hind wing: marginal cell basally slightly and distally strongly widened, its apical width 2.6 × width at level of hamuli (Fig. 54); 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 5:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather slender and medium-sized brownish pecten (Fig. 63); hind coxa largely densely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 5.1 and 6.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite robust, evenly convex; 1st and 2nd tergites rather coarsely obliquely rugose; 1st tergite and basal half of 2nd tergite with median carina; 2nd tergite robust and with striae diverging posteriorly; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture rather deep medially; 3rd tergite largely smooth, except anteriorly with some striae; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with rather long setae and apically rounded (Fig. 51). Colour. Black; scapus, pedicellus, tegulae (but humeral plate brownish yellow), base of hind tibia narrowly, apical half of hind tibia, telotarsi, hind tarsus largely, ventral apical half of metasoma, pterostigma and veins (except C+SC+R of fore wing) dark brown; remainder of basal half of antenna and palpi yellowish brown; remainder of legs (but apical two-fifths of hind femur black), 1st and 2nd tergites, 3rd tergite basally and laterally orange brown; remainder of hind tibia pale yellowish; apex of middle femur and wing membrane somewhat infuscate. Variation. A. apicalis is very variable in colour and the colour patterns are not restricted to certain areas, but in general southern Palaearctic specimens are darker than northern ones (or specimens from high altitudes). The tegula is dark brown or black, and the humeral plate usually paler than the tegula or equally black, but both usually yellowish in southern specimens; the hind tarsus is dark brown or black, but sometimes 3rd and 4th segments yellowish; the hind tibia variably reddish to black, but palest at extreme base; the pronotum is very occasionally reddish. The extent of black colouration of the legs is especially variable, and sometimes all legs are entirely black (var. rufoater (Wollaston, 1858)). Antenna, especially in females, can be more or less light reddish brown, especially basally, or dark brown/black throughout. Antennal segments: ♀ 44(1), 46(3), 47(11), 48(20), 49(31), 50(41), 51(19), 52(10), 54(3), 55(1), 57(1); ♂ 46(3), 47(7), 48(17), 49(29), 50(30), 51(32), 52(11), 53(5), 54(1). Males have on average approx. one antennal segment more than females. Apical tergites of ♂ type 4, setosity dense (making the tergites appear concave; Figs 68, 69) and fringe weak. *Albania, Austria, *Bosnia & Herzegovina, *Bulgaria, *Croatia, Cyprus, *Czech Republic, *France (including Corsica), *Georgia, *Germany, Greece (including Chios, Corfu, Crete, Lesbos, Rhodes), *Hungary, Iran, *Iraq, *Israel, *Italy (including Sardinia, Sicily), *Kazakhstan, *Malta, *Moldova, *Montenegro, *Morocco, *North Macedonia, *Oman, *Portugal (including Madeira), *Romania, *Russia (including Dagestan), *Serbia, *Slovakia, Spain (including Mallorca and Canary Islands: Tenerife, Fuerteventura), *Syria, Switzerland, *Tunisia, Turkey, *Turkmenistan. The synonymy of Rogas rufo-ater Wollaston, 1858, and Rhogas similis Szépligeti, 1903, are based on examination of the types listed above. The lectotype of Rogas bicolor Lucas, 1849 (not Spinola, 1808) and of Rhogas bicolorinus Fahringer, 1932, has been examined by Dr Jenö Papp and we agree with his opinion that it is a synonym of A. ductor auct. (= A. apicalis). The types of Rogas reticulator Nees, 1834, and Rhogas reticulator var. atripes Costa, 1884, are lost or unavailable and their synonymy is based on the original description and the interpretation by later authors. 10.3897/zookeys.919.39642.figures50-5325D7F282-B3A7-5EF0-A169-0A257A48DF74Figures 50– 53. Aleiodes apicalis (Brullé), ♀, Greece, Thimiana Chios, but 52 mummies of Autographa gamma (Linnaeus) from Malta and 53 of undetermined plusiine host from Cyprus 50 habitus lateral 51 ovipositor sheath lateral 52 mummy dorsal 53 mummy covered by silk of host. https://binary.pensoft.net/fig/39005410.3897/zookeys.919.39642.figures54-65FB7FA51D-63DD5751-B715-879670F8A4E0Figures 54–65. Aleiodes apicalis (Brullé), ♀, Bulgaria, Rodopi 54 wings 55 mesosoma lateral 56 mesosoma dorsal 57 1st –3rd metasomal tergites dorsal 58 fore femur lateral 59 hind femur lateral 60 head anterior 61 head dorsal 62 head lateral 63 outer hind tarsal claw 64 base of antenna 65 apex of antenna. https://binary.pensoft.net/fig/39005510.3897/zookeys.919.39642.figures66-71143A68FD-09935A13-ACDF-DC57416186CBFigures 66–71. Aleiodes apicalis (Brullé), ♂, Turkey, Sivas 66 habitus lateral 67 head dorsal 68 3rd –7th metasomal tergites dorsal 69 3rd –7th metasomal tergites lateral 70 head anterior 71 outer hind tarsal claw. https://binary.pensoft.net/fig/390056FahringerJ (1932) Opuscula braconologica. Band 3. Palaearktischen Region. Opuscula braconologica 4: 241–320. Wagner, Wien.ShenefeltRD (1975)

Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115– 1262.ShawSRMarshPMFortierJC (1998) Revision of North American Aleiodes Wesmael (Part 2): the apicalis (Brullé) species-group in the New World (Hymenoptera: Braconidae, Rogadinae) Journal of Hymenoptera Research 7: 62–73.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xZaldívarRiverónAShawMRSáezAGMoriMBelokobylskijSAShawSRQuickeDLJ (2008) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Biology 8: 329. https://doi.org/10.1186/1471-2148-8-329FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2AbdolalizadehFMadjdzadehSMFarahaniSAskari HesniM (2017) A survey of braconid wasps (Hymenoptera: Braconidae: Euphorinae, Homolobinae, Macrocentrinae, Rogadinae) in Kerman province, southeastern Iran.Journal of Insect Biodiversity and Systematics3(1): 33–40.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera).

Folia Entomologica Hungarica.66: 137–194.SpinolaM (1808) Insectorum Liguriae species novae aut rariores, quas in agro Ligustico nuper detexit, descripsit, et iconibus illustravit (Hymenoptera) 2: 1– 262. Genua.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1990) Braconidae (Hymenoptera) from Tunisia, 3.Folia Entomologica Hungarica51: 89–96.FahringerJ (1934) Opuscula braconologica. Band 3. Palaearktischen Region. Opuscula braconologica 5–8: 321–594. Wagner, Wien.CurtisJ (1834) British Entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland 11: 512.ZaykovAN (1980a) Unknown genera and species of Braconidae for the fauna of Bulgaria. Travaux scientifiques d’Université de Plovdiv “Paissi Hilendarski”, Biologie 15(4) [1977]: 111–115. [in Bulgarian]TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]SamartsevKGBelokobylskijSA (2013) On the fauna of the true cyclostome braconid wasps (Hymenoptera, Braconidae) of Astrakhan Province. Entomologicheskoe Obozrenie 92(2): 319–341.Entomological Review93(6): 755– 774.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.

6. XML treatment for Aleiodes arnoldii. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396421E255C34-5930-595A-9BD524D03A2D81BF Aleiodes arnoldii (Tobias, 1976)Figs 72–73, 74–79, 80, 81–92 Rogas (Rogas) arnoldiiTobias, 1976: 84, 222, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) arnoldi[sic!]; Papp 1985a: 152. Aleiodes (Neorhogas) arnoldii; Papp 1991a: 87. Holotype, ♀ (ZISP) “[Azerbaijan], Kosmoljan, Zuvan, 19.v.[1]936, Arnoldi”, “Holotypus Rogas arnoldii Tobias”. 1 ♂ (RMNH), “Turkey, Hakkâri, Tanin Tanin Pass, 25.vi.1985, 2200 m, C.J. Zwakhals”. Male is provisionally associated with this species; it may belong to a related species. None. Unknown. The holotype was collected in May. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 78); clypeus obtuse apically and not protruding in lateral view (Fig. 77); length of malar space of ♀ 0.5–0.6 × height of eye in lateral view; antennal segments of ♀ 35–37 and length of antenna of ♀ 0.8–0.9 × fore wing; OOL sparsely punctate; lateral lobes of mesoscutum largely smooth; posterior half of notauli shallow; precoxal area coarsely vermiculate-rugose medially; head, palpi and part of mesosoma of ♀ yellowish brown; pterostigma dark brown; apex of hind tibia of ♀ yellowish; hind tarsal claws yellowish or brownish setose (Fig. 72); 4th–6th tergites of ♂ flat and normally setose, but setae slightly longer than on basal tergites (Fig. 92). Holotype, ♀, length of fore wing 4.4 mm, of body 5.7 mm. Head. Antennal segments of ♀ 37, length of antenna 0.85 × fore wing, its subapical segments quadrate; frons with rather coarse curved rugae, shiny, and rugose behind antennal sockets; OOL 2.0 × diameter of posterior ocellus, and finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate, shiny; face transversely rugose; clypeus finely rugulose and with long setae; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.45 × minimum width of face; length of eye 1.1 × temple in dorsal view (Fig. 79); vertex behind stemmaticum rugulose; clypeus near lower level of eyes; length of malar space 0.55 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely smooth, shiny, sparsely and finely punctate; precoxal area of mesopleuron coarsely rugose, but absent posteriorly; metapleuron remotely punctate, interspaces much wider than diameter of punctures, shiny; mesopleuron above precoxal area (except speculum) punctate and dorsally rugose; scutellum sparsely punctate or punctulate, medio-posteriorly rugulose and with some striae laterally, no carina; propodeum evenly convex and coarsely vermiculaterugose, medio-longitudinal carina strong in basal 0.6, and without tubercles. Wings. Fore wing: just reaching apex of metasoma; r 0.35 × 3-SR (Fig. 74); 1-CU1 horizontal, 0.45 × 2-CU1; r-m unsclerotized; 2nd submarginal cell medium-sized (Fig. 74); cu-a vertical, straight; 1M nearly straight posteriorly; 1-SR wide. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 72); 2-SC+R subquadrate; m-cu distinct, but unsclerotized and as long as cu-a; M+CU:1-M = 15:9; 1r-m 0.7 × 1-M. Legs. Tarsal claws subpectinate, with six yellowish medium-sized pectinal bristles; hind coxa obliquely striated dorsally, punctulate laterally; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 4.6 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus.

Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites coarsely longitudinally and densely rugose, robust and posterior corners of 1st protruding outside base of 2nd tergite, with distinct median carina; medio-basal area of 2nd tergite wide and short; 2nd suture moderately deep and crenulate; basal half of 3rd tergite longitudinally striate, remainder of metasoma largely smooth, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, setose and apically truncate (Fig. 73). Colour. Yellowish brown; mesosoma (except mesoscutum, scutellum medially, pronotum anteriorly and dorsally), ovipositor sheath, 3rd tergite (except antero-lateral corners) and following segments black; apical half of antenna, pedicellus, palpi, hind femur apico-dorsally, telotarsi, veins, parastigma basally and pterostigma dark brown; wing membrane rather brownish infuscate. Variation. Antennal segments of ♀ 37(1). Male is largely black, except for 2nd tergite and anterior half of 3rd tergite (Fig. 80). Azerbaijan, *Turkey. Easily confused with A. ruficornis (Herrich-Schäffer); the relative size of the clypeus (wider and somewhat shorter in A. arnoldii than in A. ruficornis) seems to be the main difference in both sexes. In addition, the female of A. arnoldii has the temple ventrally and the malar space yellowish brown (dark brown in A. ruficornis). The male has darker legs and 1st metasomal tergite than the female (the sexes more similar in A. ruficornis). Also reported from Uzbekistan (Yuldashev, 2006); the record from Poland (Huflejt, 1997) most likely concerns A. ruficornis (Herrich-Schäffer). Aleiodes arnoldiisensuFarahani et al. (2015) concerns a species closely related to A. gasterator (Jurine) but has basal half of 3rd tergite coarsely longitudinally rugose, antenna of ♀ with 30–35 segments (of ♂ 36), head linearly narrowed ventrally and subbasal antennal segments of ♀ slightly slenderer. 10.3897/zookeys.919.39642.figures72-73D8339728-0BB7-5DE2-AF96-7F6763CD34ECFigures 72, 73. Aleiodes arnoldii (Tobias), ♀, holotype 72 habitus lateral 73 ovipositor sheath lateral. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39005710.3897/zookeys.919.39642.figures74-791EB71D01-9D245985-8EC6-CB0EAF5EA6DAFigures 74–79. Aleiodes arnoldii (Tobias), ♀, holotype 74 wings 75 mesosoma lateral 76 antenna 77 head lateral 78 head anterior 79 head dorsal. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39005810.3897/zookeys.919.39642.figure809FE171F2-0F5F-5F6F90BE-4550C5CAD1C1Figure 80. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass, habitus lateral. https://binary.pensoft.net/fig/39005910.3897/zookeys.919.39642.figures81-92D7B63730-B1C95937-9156-0402627E8E5FFigures 81–92. Aleiodes arnoldii (Tobias), ♂, Turkey, Tanin Pass 81 wings 82 mesosoma lateral 83 mesosoma dorsal 84 metasoma dorsal 85 fore femur lateral 86 hind femur lateral 87 inner hind tarsal claw 88 head anterior 89 head dorsal 90 head lateral 91 base of antenna 92 apex of metasoma lateral. https://binary.pensoft.net/fig/390060PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2

7. XML treatment for Aleiodes aterrimus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642D5B255F1-2788-5659-BE7BE2FCF928FF9E Aleiodes aterrimus (Ratzeburg, 1852)Figs 93–97, 98–100, 101, 102–115 Bracon aterrimusRatzeburg, 1852: 35; Shenefelt 1978: 1467. Aleiodes aterrimus; Belokobylskij et al. 2003: 398; Zaldívar-Riverón et al. 2004: 234. Aleiodes grandisGiraud, 1857: 178; Papp 1991a: 77; Bergamasco et al. 1995: 5; Belokobylskij et al. 2003: 398; Papp 2005: 176 (as valid species) [examined]. Aleiodes (Neorhogas) grandis; Papp 1985a: 159 (lectotype designation and as synonym of A. aterrimus); Riedel et al. 2002: 106. Aleiodes (Chelonorhogas) aterrimus; Falco et al. 1997: 60. Rogas grandis; Shenefelt 1975: 1232. Rogas (Rogas) grandis; Tobias 1976: 87, 1986: 81 (transl.: 134). Rhogas malaiseiShestakov, 1940: 7. Rogas malaisei; Shenefelt 1975: 1237. Aleiodes malaisei; Shaw et al. 1998: 63 (as synonym of A. grandis Giraud); Belokobylskij et al. 2003: 398 (as synonym of A. aterrimus (Ratzeburg)); Papp 2005: 176 (as synonym of A. grandis Giraud). Rogas (Rogas) vicinusPapp, 1977a: 114, 115 [examined]. Syn. nov. Aleiodes (Neorhogas) vicinis; Papp 1991a: 78. Lectotype of A. grandis, ♂ (MNHN), “[Austria:] environs de Vienne”. Holotype of R. vicinus (MTMA), ♀, “Yugoslavia, [Serbia:] Vojvonida, Fruška Gora Mts., Sremska Kamenica, 1–2.v.1972, Papp & Horvatovich”, “Holotypus ♀ Rogas vicinus sp. n., Papp, 1977”, “Hym. Typ. No. 2375, Mus. Budapest”; paratype of R. vicinus, ♀ (MTMA), “[Romania:] Transylvania, Szászkezd%, Silbernagel”, “Paratypus ♀ Rogas vicinus sp. n., Papp, 1977”, “Hym. Typ. No. 2376, Mus. Budapest”; 1 ♂ (MTMA), id., but No. 2375. Austria, Belgium, British Isles (England V.C.s 8, 9, 10, 11, 12, 14, 15, 20, 22, 28, 29, 39), Czech Republic, Finland, Germany, Hungary, Netherlands (GE: Brummen (Leuvenheim); LI: Epen; ZH: Schoonrewoerd), Poland, Romania, Russia, Slovakia, Spain, Switzerland. Specimens in ZJUH, BZL, CNC, FMNH, HSC, MRC, MSC, MTMA, NMS, OUM, RMNH, SDEI, ZSSM. MRS024 (UK), MRS147 (UK). Univoltine, spending ca ten months of the year in the exposed mummy on an aerial twig. Collected from April–June, among broadleaved trees (but see paragraph below). Reared from arboreal Amphipyra spp.: A. pyramidea (Linnaeus) (29; M.G. Bloxham, C. Bystrowski, J. Connell, A.P. Fowles, G.M. Haggett, B.T. Parsons, D.L.J. Quicke, M.R. Shaw); A. berbera (Rungs) (5:1 [5 OUM]; G.C. Varley); Amphipyra sp. (8). Some of the forgoing specimens were reared and labelled in the period before it was known that there are two closely related and sympatric arboreal species of Amphipyra in Britain, and it is possible that British records from A. pyramidea (especially when collected on Quercus; cf. Shaw, 1981) have been overstated at the expense of A. berbera; however, both certainly serve as host. An account of frequency at one site is given by Shaw (1981). Before becoming mummified the host moves to a narrow twig, to which the mummy will be very strongly glued. In the early stage of the mummification process (Fig. 99), in which the anterior end of the host is particularly contracted, the parasitoid larva strongly protrudes anteriorly to spread the necessary glue (Fig. 101). The resultant almost semi-circularly domed and hard mummy (Fig. 100), in which the parasitoid occupies approximately abdominal segments 4–7 of the host, forms in ca May–June and persists through the remainder of the summer and the following winter until the adult emerges in ca April–May. (The univoltine hosts overwinter in the egg stage.) The swollen part of the mummy, which is moderately densely lined with silk, is externally usually matt chalky buff in colour, but dark brown diamond-shaped patches centred dorsally on intersegmental areas tend to remain (Fig. 97), and sometimes (perhaps especially when the mummy is unable to dry as it forms) these are coalesced to leave a single shiny dark brown patch covering most of the dorsal surface. Some of the mummies examined might be of somewhat stunted final instar hosts, but others are more clearly penultimate instar. The outcomes of an experiment involving six females and cultured A. pyramidea larvae were unfortunately marred by unavoidably high temperatures and then disease overcoming the cultures so that no mummies resulted, but the following observations were made: (i) 2nd to 5th instar hosts were potentially attractive; (ii) 2nd instar hosts were, however, often ignored

or else tended to be abandoned after being paralysed with a single jab (i.e., without oviposition subsequently taking place); (iii) 3rd instar hosts were often ignored, but when attacked seemed the most smoothly parasitised, sometimes with a single paralysing jab being followed, after a short pause, by a single insertion of the ovipositor for presumed oviposition, although the pattern observed for 4th and 5th instars also occurred with 3rd instars; (iv) 4th and (2 only) 5th instar hosts were embraced the most enthusiastically, but it required several (3–5) injections to subdue them, and then there were usually several (3–4) separate sequential and lengthy (often as long as 80 seconds) insertions of the ovipositor (which may or may not all have been actual ovipositions), the parasitoid turning between insertions and always grasping the host with all six legs during the insertion; (v) antennation of the host was minimal, and there was no post-oviposition association; (vi) all temporarily paralysed hosts hung from the substrate by one or usually more prolegs until they recovered, presumably preventing their falling from their pabulum; (vii) there is no long-term physiological venom effect. The behaviour of adults observed toward the different instars is intriguing, and the experiment would be well worth repeating under better circumstances. Although the above is a consistent pattern for this species, it does not account for a small number of specimens (14 ♀, 4♂ in BZL, MRC, MTMA, NMS, SDEI, ZSSM) examined from various localities in central Europe (Czech Republic, Germany, Hungary) and S. Russia. These specimens share small but rather consistent morphological differences from the usual form, in particular tendencies towards: more intense sculpture on the metasomal tergites (T3 being more or less strongly punctate or even rugose-punctate); the hind wing marginal cell parallel-sided in basal three fifths; shorter 3SR in fore wing; basal cell of fore wing with more, and more evenly distributed, setae; fewer antennal segments; wing membrane slightly brownish. These differences are not absolutely consistent and would be easy to let pass without comment were it not for the fact that they are correlated with an apparently different phenology, as (of the ten specimens with dates recorded) five ♀ were collected in July and one in August, with only three ♀ in May and one in June (none in April). This is in marked contrast with the earlier flight time of the usual form, and the usual hosts (arboreal Amphipyra spp.) are not available after early June. A further ♀ specimen (MTMA) examined and returned in 1997 by MRS but apparently no longer in the main MTMA collection was labelled “Hungaria, Fót, Somlyó-hegy, 30.vii.1958, Ehik”; “Ex Panchrysia deaurata Esp [J. Papp’s handscript]”; “ex Pytometra deautate [sic]”. Unfortunately, no mummy had been preserved, but this plusiine noctuid feeds on Thalictrum (a low plant, not a tree) and it is unlikely for an arboreal Amphipyra, even if fallen from a tree above, to have been mistaken for it. The date, whether referring to collection of the host larva or emergence of the adult parasitoid, is also out of step with arboreal Amphipyra species. We considered but rejected the possibility that these specimens belong to a separate species, and instead conclude that under certain circumstances A. aterrimus can have a partial 2nd brood (in the southern part of its range) which uses different hosts, and that the morphological variation is merely seasonal. The material (which does not conform to A. sapporensis (Watanabe), see below) is being returned to holding institutions determined as A. aterrimus but with “var: T3 sculpture etc.” appended to facilitate recall if necessary. It should be added that this form has (on account of its heavy metasomal sculpture and extensively parallel-sided marginal cell in the hind wing) sometimes been misidentified as A. rugulosus, but the two species are always easily separated by the sculpture of the mesoscutum and scutellum, as well as by leg colour. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 109); ventral margin of clypeus obtuse apically and not protruding outwards (Fig. 111); OOL of ♀ distinctly longer than diameter of posterior ocellus; mesoscutal lobes densely and finely punctate-coriaceous, rather matt; scutellum densely and finely coriaceous; mesosternal sulcus shallow, obsolescent or absent; vein 1-CU1 of fore wing 0.2 × vein 2-CU1 (Fig. 102); vein 2-SC+R of hind wing subquadrate or vertical (Fig. 102); hind tarsal claws with conspicuous and robust blackish pecten (Fig. 113); head black; hind tibia largely to completely black; metasoma of both sexes black; 4th–6th tergites of ♂ flat and densely short setose, except a narrow glabrous strip centrally. Dr K. Samartsev (in litt.) kindly brought to the first author’s attention that the East Palaearctic A.

sapporensis (Watanabe, 1937) occurs in southern European Russia (Middle and Lower Volga territories). Aleiodes aterrimus and A. sapporensis differ only slightly, mainly by the colour of the extreme base of the hind tibia (completely dark brown in A. sapporensis and usually narrowly pale yellowish in A. aterrimus) and by the shape of temple in dorsal view (roundly narrowed in A. sapporensis and rather linearly narrowed in A. aterrimus). There is also a slight difference in the proportions of the face (A. sapporensis has facial width 1.50–1.60 × medial height including clypeus and A. aterrimus 1.65–1.75 ×). A. sapporensis seems to have the lateral carinae of propodeum more protruding and has 58–66 antennal segments. Redescribed ♀ (RMNH) from England (Pamber Forest). Length of fore wing 7.3 mm, of body 8.6 mm. Head. Antennal segments of ♀ 59, length of antenna 1.1 × fore wing, its subapical segments rather robust; frons largely superficially granulate; OOL 1.8 × diameter of posterior ocellus, and superficially rugulose-granulate and shiny; vertex superficially rugulose-granulate, rather shiny; clypeus with some punctures; ventral margin of clypeus thick and not protruding forwards (Fig. 111); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 109); length of eye 1.3 × temple in dorsal view (Fig. 110); vertex behind stemmaticum superficially granulate-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate-coriaceous, rather matt; precoxal area of mesopleuron largely smooth medially, densely punctate anteriorly and posteriorly; metapleuron densely punctate; metanotum with nearly complete median carina; scutellum punctate-coriaceous; propodeum rather convex and coarsely reticulate-rugose, medio-longitudinal carina nearly complete, and with slightly protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 102); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 102); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 103); 2-SC+R short and vertical; m-cu absent; M+CU:1-M = 12:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws with conspicuous and robust blackish pecten (Fig. 113); hind coxa largely densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st and 2nd tergites with mediolongitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 106); 2nd suture deep and narrow; basal half of 3rd tergite finely punctate-rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 95). Colour. Black; antenna (except scapus and pedicellus), palpi, tegulae, fore and middle telotarsi, veins and pterostigma dark brown; coxae, trochanters and trochantelli, apical third of hind femur (ventrally extended to its apical two-thirds), hind tibia (except pale yellowish basal ring) and hind tarsus black, remainder of legs yellowish brown; wing membrane subhyaline. Variation. Hind femur usually only apically dark brown, but sometimes entirely dark brown; coxae black or sometimes largely yellowish brown. Two females (both NMS, from different localities) have vein r-m of fore wing absent. Males are very similar, apical tergites type 3, with fringe very weak to negligible; hind femur often only apically blackish, but sometimes up to apical 0.6 darkened. Antennal segments: ♀ 57(3), 58(1), 59(5), 60(7), 61(3), 62(6), 63(5), 64(1); ♂ 51(1), 52(1), 53(3), 54(1), 55(5), 56(7), 57(6), 58(5), 59(3), 60(1), 62(2); females have on average ca four more antennal segments than males. The antennal segments for the specimens of the abnormal series (see above) are scored separately here: ♀ 54(2), 55(1), 56(2), 57(3), 58(3), 60(1); ♂ 54(1). Austria, *Belgium, British Isles (England), Czech Republic, *Finland, Germany, Hungary, *Netherlands, Poland, *Romania, Russia, Serbia, Slovakia, Spain, *Switzerland. The synonymy of Rogas vicinus Papp, 1977, with Aleiodes aterrimus (Ratzeburg, 1852) is based on

the examination of the types listed above. The differences between R. vicinus and R. grandis (= A. aterrimus) listed in the original description (head less constricted posteriorly, apical antennal segments more robust, 1st metasomal tergite less robust and 2nd tergite somewhat longer) fall within the normal variation of A. aterrimus. 10.3897/zookeys.919.39642.figures93-977E59AE8D-3A52-583D-BBEE-A55931BCD1A8Figures 93–97. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 93 habitus lateral 94 detail of fore wing with arrow indicating lost vein r-m 95 ovipositor sheath lateral 96, 97 mummies of Amphipyra sp. showing variation in markings. https://binary.pensoft.net/fig/39006110.3897/zookeys.919.39642.figures98-100051EF52B-1F9E5B88-BA41-4A1940BB491FFigures 98–100. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest 98 parasitised caterpillar of Amphipyra sp. 99 early stage of mummy 100 later stage of mummy. https://binary.pensoft.net/fig/39006210.3897/zookeys.919.39642.figure1019D618773-4828-506AA8FF-0BAEB8C42B2AFigure 101. Larva of Aleiodes aterrimus (Ratzeburg) mummifying Amphipyra pyramidea (Linnaeus), with its anterior (indicated by the arrow) projecting from the ventral opening in the host to spread adhesive over a wide area. https://binary.pensoft.net/fig/39006310.3897/zookeys.919.39642.figures102-115A19714FD-6AB9522E-9BF7-1EADD8FAF96EFigures 102–115. Aleiodes aterrimus (Ratzeburg), ♀, England, Pamber Forest, but 102 from Austria, Wien 102 fore wing 103 hind wing 104 mesosoma lateral 105 mesosoma dorsal 106 metasoma dorsal 107 fore femur lateral 108 hind femur lateral 109 head anterior 110 head dorsal 111 head lateral 112 hind tibia and tarsus lateral 113 outer hind tarsal claw 114 base of antenna 115 apex of antenna. https://binary.pensoft.net/fig/390064ShenefeltRD (1978) Braconidae 10. Braconinae, Gnathobraconinae, Mesostoinae, Pseudodicrogeniinae, Telengainae, Ypsistocerinae, plus Braconidae in general, major groups, unplaced genera and species.Hymenopterorum Catalogus (nova editio)15: 1425–1872.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.ZaldívarRiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xPappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1– 122.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97– 108.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72– 85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]ShawSRMarshPMFortierJC (1998) Revision of North American Aleiodes Wesmael (Part 2): the apicalis (Brullé) species-group in the New World (Hymenoptera: Braconidae, Rogadinae) Journal of Hymenoptera Research 7: 62–73.ShawMR (1981) Possible foodplant differences of Amphipyra pyramidea (L.) and A. berbera svenssoni Fletcher (Lepidoptera: Noctuidae), and a note on their parasitoids (Hymenoptera).Entomologist’s Gazette32: 165–167.

8. XML treatment for Aleiodes carbonarius. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642218FBB39-B50D-5B6F-8A6462568C765B70 Aleiodes carbonarius Giraud, 1857Figs 116–118, 119–131, 132–137 Aleiodes carbonariusGiraud, 1857: 177–178 [examined]. Aleiodes (Neorhogas) carbonarius; Papp 1985a: 156 (lectotype designation), 1991a: 88. Aleiodes carbonarius; Papp 2005: 176. Rogas carbonarius; Shenefelt 1975: 1220–1221. Rhogas (Rhogas) carbonarius ab. giraudiFahringer, 1931: 236; Shenefelt 1975: 1221 (invalid name). Lectotype of A. carbonarius, ♂ (MNHN), “Hunga[ry]”, “2”, “Hungaria”, “Neusiedlersee/teste Papp J., 1979”, “Lectotypus”, “Aleiodes carbonarius Gir., 1857, ♂, Papp, 1979”. Paralectotype ♂ (MNHN) from Austria (near Vienna). 3 ♀ (NMS), “Hungary: Veszprém, nr Tótvázsony, larva coll. 21.v.2001, Tholera decimalis, mum. c. 12.vi.[20]01, em. 19.v., 24.v. and 25.v.[20]02, M.R. Shaw”; 1 ♂ (MSC), “A[ustria], Oberösterreich, Wels, Flughaven, 48°10'N, 14°2'E, 30.iv.2012, M. & J. Schwarz”; 1 ♂ (MTMA), “Hungaria, Csákvár”, “Vértes Hgs., Hajduvágás”, “12.v.1961, Sólymosné”, “Rogas carbonarius Gir. ♂, det. Papp, 1979 / compared with lectotype ♂”; 1 ♂ (NMS), “[Hungaria,] P. Szt. Lelek, Ujhelyi”, “Rogas morio Reinh. ♂, det. Szépligeti”, “Rogas carbonarius Gir. ♂, det. Papp, 1979”; 1 ♂ (MTMA), id., but Budapest, Svabhegy; 2 ♂ (MRC) “Russia, E. Siberia Lake Baikal, Biakalo-Lenskiy res. 20.vi. and 19.vii. [19]05, leg. Berlov”; 1 ♂ (BZL), “CSR [Czech Rep.], envir. Prague, 1968, Dr. Pádr”. This species appears to be sporadic in central and eastern Europe. The specimens from which Morley (1937) recorded this species as new to Britain have been examined and belong to A. carbonaroides sp. nov. MRS162 (Hungary), MRS163 (Hungary), MRS 164 (Hungary). Adults of this lowland species have been collected from the very end of April to July (see also Papp, 1999), and it is found in grassland habitats. Reared from the noctuid Tholera decimalis (Poda) (3:1; M.R. Shaw/Hungary). The decidedly large mummy is very similar to that of A. grassator and forms underground (Fig. 118). Univoltine, overwintering in the mummy. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 126); OOL of ♀ ca 2.6 × as long as diameter of posterior ocellus (Fig. 127) and distinctly rugose; length of 4th antennal segment of ♀ ca 0.9 × its width (Fig. 129; in ♂ 0.9–1.0 times; Fig. 135); clypeus thick apically and not protruding anteriorly (Fig. 128); lobes of mesoscutum densely punctate, interspaces superficially granulate and with satin sheen; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending near level of apex of vein 3-M (Fig. 119); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 119); vein 3-SR of ♀ 1.7–2.0 × as long as vein 2-SR; vein 3-SR ca 0.7 × vein SR1 (Fig. 119; of ♂ ca 0.5×); hind tarsal claws yellowish or brownish bristly setose (Fig. 131); inner side of hind tibia of ♀ yellowish; tegulae yellowish brown; 4th and 5th tergites black. Probably a lowland species in C. Europe. Redescribed ♀ (NMS) from Hungary (Veszprém). Length of fore wing 4.1 mm, of body ca 6.0 mm. Head. Antennal segments of ♀ 46, 4th segment 0.9 × longer than wide (Fig. 129); length of antenna 1.15 × fore wing, its subapical segments robust (Fig. 130); frons with coarse curved rugae and rather shiny; OOL 2.6 × diameter of posterior ocellus and rugulose; vertex rugose and shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 128); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 126); length of eye 1.4 × temple in dorsal view (Fig. 127); vertex behind stemmaticum rugose; clypeus distinctly below lower level of eyes; length of malar space 0.7 × length of eye in lateral view (Fig. 128). Mesosoma. Mesoscutal lobes densely punctate, interspaces superficially granulate and with satin sheen; precoxal area of mesopleuron coarsely rugose medially and punctate posteriorly; remainder of mesopleuron mainly coarsely punctate; scutellum flat, sparsely finely punctate and with lateral carina; propodeum coarsely rugose, medio-longitudinal carina indistinct, rounded posteriorly and dorsal part rather short. Wings. Fore wing: r 0.4 × 3-SR; marginal cell ends near level of apex of 3-M (Fig. 119); 1-CU1 horizontal and slightly widened, 0.45 × 2-CU1; r-m 0.3 × 3-SR; 2nd submarginal cell elongate (Fig. 119), 3-SR twice as long as 2-SR; cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR

slender and medium-sized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 120); 2-SC+R slightly longer than wide; m-cu short, postfurcal; M+CU:1-M = 61:36; 1r-m 0.75 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 131); hind coxa largely rather densely punctate; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 4.5 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 0.9 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and short (Fig. 123); 2nd suture deep and crenulate; basal third of 3rd tergite finely longitudinally striate, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 117). Colour. Dark orange brown; apical two-thirds of antenna, patch on hind femur dorso-apically, and telotarsi, dark brown; temple ventrally, malar space, mesosternum, mesopleuron, metapleuron, propodeum, pair of patches on 2nd tergite and most of apical 0.4 of tergite, and 3rd–7th tergites black; palpi (especially labial palp), veins and pterostigma dark brown, basal third of antenna (but scapus dorsally blackish) rather pale yellowish brown; tegulae and remainder of legs; yellowish brown; wings strongly infuscate. Variation. Antennal segments: ♀ 46(2), 49(1); ♂ 47(1), 50(1), 52(1), 54(1), 56(1), 57(1); length of fore wing of ♀ ca two-thirds of body length (0.8 × in ♂); males always darker than females; mainly black with legs mainly dark brown or blackish, but male from Austria has basal half of metasoma orange brown and legs partly yellowish brown. Males have 2nd submarginal cell distinctly shorter than in females (as in A. grassator), antenna 0.9 × length of body and slightly less robust subapically, temple and face long setose and malar space 0.5–0.7 × length of eye in lateral view; metasoma black or 1–2 basal tergites reddish and apical tergites type 1, fringe not observed (Fig. 132). Austria, Czech Republic, Hungary, *Russia (Lake Baikal). Very similar to A. grassator (Thunberg), and especially A. carbonaroides; males of A. carbonarius and carbonaroides are normally black but males with partly orange brown metasoma occur. The three species exhibit sexual dimorphism of the 2nd submarginal cell (less robust (and also longer in A. carbonarius) in female than in male). Giraud (1857) gave an incomplete description of the only two males he possessed, but clearly indicated that the antenna is slightly shorter than the body. The female of this species is reported for the first time. 10.3897/zookeys.919.39642.figures116-1187A65E536-5B45-58A6-A395-B61CA19A0A9DFigures 116–118. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 116 habitus lateral 117 ovipositor sheath lateral 118 mummy of Tholera decimalis (Poda). https://binary.pensoft.net/fig/39006510.3897/zookeys.919.39642.figures119-131C42B5EB2-8FBC5583-A967-300046F11C24Figures 119–131. Aleiodes carbonarius Giraud, ♀, Hungary, Veszprém 119 fore wing 120 hind wing 121 mesosoma lateral 122 mesosoma dorsal 123 metasoma dorsal 124 fore femur lateral 125 hind femur lateral 126 head anterior 127 head dorsal 128 head lateral 129 base of antenna 130 apex of antenna 131 inner hind tarsal claw. https://binary.pensoft.net/fig/39006610.3897/zookeys.919.39642.figures132-13714702ABD-1CCA5F18-87A4-7B99B412D4F1Figures 132–137. Aleiodes carbonarius Giraud, ♂, Hungary, Csákvár 132 habitus lateral 133 head dorsal 134 wings 135 base of antenna 136 apex of antenna 137 head anterior. https://binary.pensoft.net/fig/390067PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.MorleyC (1937) The Hymenoptera of Suffolk, portio tertio et ultimo.Transactions of the Suffolk Naturalists’ Society3: 233–248.GiraudJ (1857) Description de quelques hyménoptères nouveaux ou rares.Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien7: 163–184.

9. XML treatment for Aleiodes carbonaroides. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642F9211028-9578-55D5-BA051033F2974422 Aleiodes carbonaroides http://zoobank.org/0BE2C69B-E310-4DFB-BE5C07218AC6F018 van Achterberg & Shawsp. nov.Figs 138–141, 142–153, 154–160 Holotype, ♀ (NMS), “[Netherlands: Friesland], Holland [sic!], Schiermonnikoog, em. 20.v. [19]82”, “ex Cerapteryx graminis larva”. Paratypes: 2 ♀ (NMS, RMNH), 3 ♂ (NMS,

RMNH), topotypic and from same host, em. 19 or 20.v.1982; 1 ♂ (ZSSM) “[Germany], Münehey, 26.iv.[18]85 R7”, “1-653”; 2 ♂ (CMIM) “[England] 25.v.[19]22, Bdn. [= Brandon, Suffolk] HF”, “Named by Claude Morley 2 Rhogas carbonarius Giraud. NEW TO BRIT. CM V.22”; 1 ♂ (ZJUH) “[England], Totternhoe, [Bedfordshire], 30.v.[19]64 [V.H. Chambers]”. Sporadic in western Europe. None. Adults of this lowland species have been collected in April and May. The two paratypes from Suffolk were swept from Breck grassland (Morley, 1937, misidentified as A. carbonarius). Reared

from the grass-feeding noctuid Cerapteryx graminis (Linnaeus) (6 [2 are RMNH]; K.P. Carl/Netherlands). If it is a specialist, it is presumably univoltine and overwinters in the mummy (the univoltine known host overwinters in the egg stage). Mummy similar to that of the closely related A. carbonarius and A. grassator, but slightly smaller. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 149); OOL of ♀ 1.8–2.0 × as long as diameter of posterior ocellus (Fig. 150) and distinctly rugose or rugulose; length of 4th antennal segment of ♀ 0.7–0.9 × its width (Fig. 152; in ♂ up to 1.0 times); clypeus thick apically and not protruding anteriorly (Fig. 151); lobes of mesoscutum punctate, interspaces largely coriaceous and superficially coriaceous; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 142); vein 1-CU1 of fore wing 0.5–0.6 × as long as vein 2-CU1 (Fig. 142); 2nd submarginal cell of fore wing medium-sized (Fig. 142); hind tarsal claws slender and yellowish or brownish bristly setose; hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; head and mesoscutum of ♀ reddish; palpi and tegulae of ♀ brownish yellow; males entirely black, with palpi, tegulae and antenna dark brown or blackish. Holotype, ♀, length of fore wing 4.2 mm, of body 7.1 mm. Head. Antennal segments of ♀ 45, 4th segment 0.9 × longer than wide (Fig. 152); length of antenna 1.1 × fore wing, its subapical segments robust (Fig. 153) and scapus oblique apically; frons with coarse curved rugae and shiny; OOL 1.8 × diameter of posterior ocellus and rugulose; vertex rugose and shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 151); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 149); length of eye 1.2 × temple in dorsal view (Fig. 150); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view. Mesosoma. Mesoscutal lobes moderately punctate, interspaces superficially granulate-coriaceous and with satin sheen; precoxal area of mesopleuron coarsely rugose medially, but largely smooth posteriorly; remainder of mesopleuron mainly punctate; scutellum flat, sparsely finely punctate and with irregular lateral carina; propodeum coarsely rugose, medio-longitudinal carina complete, rounded posteriorly and dorsal part approx. as long as posterior part. Wings. Fore wing: r 0.4 × 3-SR (Fig. 142); marginal cell ends basad of level of apex of 3-M; 1CU1 horizontal, 0.5 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell robust (Fig. 142), 3-SR 1.4 × as long as 2-SR; cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR similar to 1-M and medium-sized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 1.7 × width at level of hamuli (Fig. 143); 2-SC+R subquadrate; m-cu short; M+CU:1-M = 27:15; 1r-m 0.7 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 140); hind coxa largely rugulose dorsally; hind trochantellus robust; length of hind femur and basitarsus 3.2 and 4.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus.

Metasoma. First tergite rather flattened, 0.7 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite without medio-longitudinal carina; medio-basal area of 2nd tergite triangular and short; 2nd suture deep and crenulate; basal half of 3rd tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured; 4th and apical third of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 139). Colour. Dark orange brown; apical half of antenna, patch on hind femur dorso-apically, and telotarsi apically, dark brown; mesosternum, mesopleuron (except dorsally and postero-ventrally), metapleuron (except medio-dorsally), propodeum (except pair of posterior patches), 3rd–7th tergites (except antero-lateral corners of 3rd tergite) black; palpi, basal half of antenna, tegulae and remainder of legs rather pale yellowish brown; veins and pterostigma dark brown; wings strongly infuscate but hind wing less than fore wing. Variation. Basal third or half of antenna of ♀ pale yellowish brown; vein 3-SR 1.4–1.6 × as long as vein 2-SR; hind femur of ♀ 3.2–3.5 × longer than wide; 1st metasomal tergite 0.7–0.8 × its apical width; temple and occiput ventrally, and malar space ventrally orange brown or black. Antennal segments: ♀ 43(1), 45(1); ♂ 48(1), 49(2), 51(1), 50(1), 53(2); males clearly have many more antennal segments than females. Males are much darker than females; body black with palpi and legs mainly dark brown or blackish (Fig. 154). Males have 2nd submarginal cell slightly smaller than females (Fig. 158), temple and face long setose, malar space 0.5–0.7 × length of eye in lateral view, and apical tergites type 1 and fringe not observed (Fig. 154); sometimes superficial granulosity of 3rd tergite and of mesoscutum are absent. Germany, Netherlands, U.K. The suffix “-oides” indicates similar to; in this case the high similarity to A. carbonarius Giraud. 10.3897/zookeys.919.39642.figures138-141A0138053-F634-5848-AFE7-BB186A4EAA4FFigures 138–141. Aleiodes carbonaroides sp. nov., ♀, holotype 138 habitus lateral 139 ovipositor sheath lateral 140 outer hind tarsal claw lateral 141 mummy of Cerapteryx graminis (Linnaeus). https://binary.pensoft.net/fig/39006810.3897/zookeys.919.39642.figures142-153E65C2ACC-84B65071-AB21-E6A3AE3F6212Figures 142–153. Aleiodes carbonaroides sp. nov., ♀, holotype 142 fore wing 143 hind wing 144 mesosoma lateral 145 mesosoma dorsal 146 metasoma dorsal 147 fore femur lateral 148 hind femur lateral 149 head anterior 150 head dorsal 151 head lateral 152 base of antenna 153 apex of antenna. https://binary.pensoft.net/fig/39006910.3897/zookeys.919.39642.figures154-16086D778D8-38175FEF-9743-028C28D1C1ECFigures 154–160. Aleiodes carbonaroides sp. nov., ♂, paratype 154 habitus lateral 155 apex of antenna 156 antenna 157 base of antenna 158 wings lateral 159 head anterior 160 head dorsal. https://binary.pensoft.net/fig/390070

10. XML treatment for Aleiodes caucasicus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642F81E320E-FEC2-5292-ACFC800C6F9C8DEE Aleiodes caucasicus (Tobias, 1976)Figs 161–163, 164–177 Rogas (Rogas) caucasicusTobias, 1976: 86, 222, 1986: 81 (transl.: 133) [examined]. Aleiodes (Neorhogas) caucasicus; Papp 1985a: 152. Aleiodes caucasicus; Papp 1991a: 75 (as synonym of A. fortipes), 2005: 176 (id.); Fortier and Shaw 1999: 227; Žikić et al. 2002: 108; Aydogdu and Beyarslan 2005: 191. Holotype, ♀ (ZISP), “[Russia], Sotchi, Lazarevskoe [terras], 26.iv.[1]973, V. Tobias”, “Holotypus Rogas caucasicus Tobias”; 2 ♀, paratype (MTMA), id., but 29.iv.1973. Figured ♀ (NMS), “[Russia], Sotchi, Lazarevskoe terras. Sklony, les [= forest], 25.iv.1988, V. Tobias”, “Rogas caucasicus Tob.”, “Aleiodes caucasicus (Tobias), det. Belokobylskij, 2005. ♀ Ant. 40”; 2 ♀ (ALC, RMNH), id., but 7.v.1975; 1 ♀ (MTMA), “Bulgaria”, “Rhodopi, St[ara] Zagora, 17.iv.1977, J. Kolarov”, “Rogas sp. n.?, det. Zaykov, 1983”, “Aleiodes fortipes Rh. ♀, det. Papp J., 1985”. None. Unknown. Specimens collected in April-May and flight time probably April–May. We have not seen reared material. Probably, like A. fortipes, it will be found to be univoltine, overwintering in the mummy, but direct evidence is lacking. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 171); antenna of ♀ with 38–41 segments and 2nd – 10th antennal segments yellowish, contrasting with remaining segments; OOL coarsely transversely striate; clypeus obtuse apically and not protruding in lateral view (Fig. 173); precoxal area finely striate (Fig. 166); tegulae yellow; lobes of mesoscutum finely coriaceous-granulate and rather dull, with satin sheen; vein 1-CU1 of fore wing much shorter than vein 2-CU1 (Fig. 164); posteriorly vein m-cu of fore wing diverging from anterior half of vein 1-M; length of hind femur 3.6–3.8 × its maximum width (Fig. 170); hind tarsal claws brownish setose (Fig. 177); length of fore wing 3.7–5.0 mm. Very similar to A. fortipes (Reinhard) and differs mainly by its body colour and sculpture of mesopleuron. Holotype, ♀, length of fore wing 3.7 mm, of body 4.6 mm. Head. Antennal segments of ♀ 41, length of antenna 1.3 × fore wing, its subapical segments rather robust; frons largely finely rugulose medially; OOL 2.2 × diameter of posterior ocellus, and coarsely transversely striate; vertex transversely striate and rather shiny; clypeus rugulose, but ventrally depressed and smooth; ventral margin of clypeus thick and not protruding forwards (Fig. 173); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 171); length of eye twice temple in dorsal view (Fig. 172); vertex behind stemmaticum rugulose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view; occipital carina largely absent dorsally and weakly developed ventrally. Mesosoma. Mesoscutal lobes largely rugulose-granulate, rather matt; precoxal area of mesopleuron transversely striate medially, distinctly rugose antero-dorsally and remainder largely punctulate; pleural sulcus moderately crenulate (Fig. 166); ventral half of metapleuron rugose; metanotum with nearly complete median carina; scutellum coriaceous; propodeum densely and finely granulaterugose and medio-longitudinal carina medium-sized. Wings. Fore wing: r 0.6 × 3-SR; 1-CU1 horizontal, 0.5 × 2-CU1; r-m unsclerotized and 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 164); cu-a vertical, straight and rather short; 1-M slightly curved posteriorly; posteriorly vein m-cu diverging from anterior half of vein 1-M. Hind wing:

marginal cell linearly widened, its apical width 2.0 × width at level of hamuli (Fig. 165); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 5:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws robust and with brownish bristles (Fig. 177); hind coxa densely rugulose and rather dull; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 5.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, 0.9 × longer than wide apically; 1st and 2nd tergites with indistinct medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 168); 2nd suture rather shallow and crenulate; medio-basally 3rd tergite striate, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 162). Colour. Orange brown; head, 3rd tergite (except antero-laterally) and subsequent tergites black; scapus, pedicellus basally, 11th and following antennal segments, palpi, veins, parastigma, pterostigma and femora apico-dorsally, tibia and tarsal segments apically, ventral half of metasoma and ovipositor sheath dark brown; tegulae, 3rd–10th antennal segments brownish yellow; wing membrane subhyaline. Variation. Head black or mainly dark brown, specimen from Bulgaria also anterior half of mesosoma; antenna of ♀ with 38 or 41 segments according to the original description; 11th and 12th antennal segments of ♀ dark brown or brownish yellow; hind femur 3.6–3.8 × as long as wide. The male is unknown, or possibly has not been distinguished from that of A. fortipes. *Bulgaria, Russia (SW). It remains unclear whether this predominantly rather yellowish orange species is distinct from A. fortipes, which in its more western localities is a much darker insect. Females intermediate in colour (and included in A. fortipes) seem to predominate in eastern Europe. More material (preferably with biological data) is needed to clarify the status of A. caucasicus. 10.3897/zookeys.919.39642.figures161-1633948DC74-1B56-5A6F-A324-8F80387803A4Figures 161–163. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 161 habitus lateral 162 ovipositor sheath lateral 163 apex of antenna (of paratype). https://binary.pensoft.net/fig/39007110.3897/zookeys.919.39642.figures164-17726C7F91D-55A352EE-A04C-89C66E6AAC9FFigures 164–177. Aleiodes caucasicus (Tobias), ♀, Russia, Sotchi 164 fore wing 165 hind wing 166 mesosoma lateral 167 mesosoma dorsal 168 metasoma dorsal 169 fore femur lateral 170 hind femur lateral 171 head anterior 172 head dorsal 173 head lateral 174 base of antenna 175 antenna 176 hind tibia and tarsus lateral 177 outer hind tarsal claw. https://binary.pensoft.net/fig/390072PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.ŽikićVBrajkovićMTomanovićZ (2002) Preliminary results of Braconid fauna research (Hymenoptera: Braconidae) found in Sicevo Gorge, Serbia. Acta Entomologica Serbica 5 (1–2) (2000): 95–110.AydogduMBeyarslanA (2005) The first records of Aleiodes Wesmael 1838 (Hymenoptera: Braconidae: Rogadinae). The fauna of Thrace region of Turkey. Linzer biologische Beiträge 37/1: 185–193.

11. XML treatment for Aleiodes coriaceus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964229D9CDFF-ED7A-537C-845299B316FC6122 Aleiodes coriaceus http://zoobank.org/EA99A74A-AA7C-460F-85F5AC7405FE67B9 van Achterberg & Shawsp. nov.Figs 178–179, 180–191, 192–195 Holotype, ♀ (NMS), “Sweden: Hr, Sveg, Duybergshammaren, 17.vii.2004, N. Ryholm, NMSZ 2004.167”, “MRS Aleiodes DNA 377”, “COI worked”. Paratypes: 1 ♂ (NMS), same label data as holotype; 1 ♀ (RMNH), “Sweden: Ås. Lilla, Vammasj. Window trap on Betula F2, 8, vii.2003, J. Hilszczanski”, “MRS Aleiodes DNA 311”, “COI worked”. MRS311 (Sweden), MRS377 (Sweden). Unknown. The available specimens were collected in July, and it is almost certainly univoltine, but we have not seen reared material. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Fig. 186); OOL of ♀ 0.9–1.1 × as long as diameter of posterior ocellus (Fig. 187), and rugulosecoriaceous or only coriaceous; ventral margin of clypeus rather thin or blunt and not protruding forwards (Fig. 188); vertex mainly coriaceous and rather dull; mesoscutal lobes coriaceous and largely matt; scutellum remotely punctate; area of precoxal sulcus largely smooth, with some punctulation; length of vein 1-CU1 of fore wing 0.3–0.4 × vein 2-CU1 and 0.4–0.5 × vein m-cu; marginal and 2nd submarginal cells of fore wing elongate (Fig. 180); tarsal claws with robust apical tooth and with medium-sized dark brown pecten (Fig. 190); hind femur and basitarsus slender (Figs 178, 185); 1st metasomal tergite comparatively steep anteriorly (Fig. 178); basal half of 3rd tergite with posteriorly diverging rugulae; head black; dorsal half of hind femur largely black dorsally; basal half of hind tibia largely dark brown; fore and middle trochanters and trochantelli infuscate or dark brown; 2nd tergite yellowish or reddish and rather slender (Fig. 183); 5th–7th tergites of ♂ medially glabrous and convex, and laterally with long setae (Figs 194, 195). Closely related to A. rufipes (Thomson) and differs mainly by the sculpture of the mesoscutum (matt instead of rather shiny), darker colour of legs, different COI and less robust 2nd and 3rd metasomal tergites. Holotype, ♀, length of fore wing 6.1 mm, of body 6.7 mm.

Head. Antennal segments of ♀ 54, antenna 1.1 × as long as fore wing, its basal segments robust, subapical segments medium-sized and apical segment with spine; frons largely smooth, except for some micro-sculpture; OOL 0.9 × diameter of posterior ocellus, rugulose-coriaceous and rather dull, groove beside posterior ocellus deep and smooth; vertex coriaceous with some rugulae, rather dull; face transversely rugose; clypeus densely rugulose; ventral margin of clypeus thin and not protruding forwards (Fig. 188); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 186); length of eye 2.1 × temple in dorsal view (Fig. 187); vertex behind stemmaticum coriaceous; clypeus partly above lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely coriaceous and matt; precoxal area of mesopleuron partly remotely punctulate and superficially micro-sculptured; medio-longitudinal carina of metanotum distinct posteriorly; scutellum punctate and with lateral carina; propodeum convex and rugose, medio-longitudinal carina absent posteriorly, and without protruding carinae laterally. Wings. Fore wing: r 0.35 × 3-SR (Fig. 180); 1-CU1 slightly oblique, 0.35 × 2-CU1; r-m 0.4 à — 3-SR; 2nd submarginal cell long (Fig. 180); cu-a slightly inclivous, straight but posteriorly slightly curved; 1-M nearly straight posteriorly; 1-SR widened; surroundings of M+CU1, 1-M and 1CU1 densely setose. Hind wing: marginal cell linearly widened, its apical width 2.3 × width at level of hamuli (Fig. 180); 2-SC+R slightly longer than wide; m-cu absent; M+CU:1-M = 50:46; 1rm 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized dark brown pecten (Fig. 189); hind coxa (except depression) coriaceous and with some rugulae dorsally; hind trochantellus robust and with long setae; length of hind femur and basitarsus 4.5 and 5.8 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex and basally rather steep, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and longitudinally rugose; maximum width of 2nd tergite 1.5 × its median length; medio-basal area of 2nd tergite medium-sized triangular and rather short (Fig. 183); 2nd suture distinct and moderately crenulate; basal half of 3rd tergite finely rugulose and rugulae diverging posteriorly, remainder of metasoma nearly smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 179).

Colour. Black; mesoscutum posteriorly, legs (but fore and middle telotarsi, fore and middle femora basally and apically, fore and middle trochanters and trochantelli, hind tarsus dark brown or infuscate, posterior half of hind femur dorsally and hind tibia largely blackish), propodeum and 1st –3rd metasomal tergites (but posterior half of 3rd tergite blackish posteriorly) reddish brown; tegulae brownish yellow, but humeral plate largely dark brown; palpi, pterostigma and veins dark brown; wing membrane slightly infuscate. Variation. Antennal segments: ♀ 52(1), 54(1); ♂ 53(1). Length of fore wing 5.3–6.1 mm. Male is very similar to female (Figs 195–195). Apical tergites of male type 1–2, and fringe scarcely visible in the single male seen. Sweden. Coriaceus is Latin for leathery, because of the coriaceous sculpture of vertex and mesoscutum. 10.3897/zookeys.919.39642.figures178-179A24DA8CB-1766-5059-A4FE-A315916CCE31Figures 178, 179.

Aleiodes coriaceus sp. nov., ♀, holotype 178 habitus lateral 179 ovipositor sheath lateral. https://binary.pensoft.net/fig/39007310.3897/zookeys.919.39642.figures180-1915548F088-03BA5D78-BB9B-82B3B3E62130Figures 180–191. Aleiodes coriaceus sp. nov., ♀, holotype 180 wings 181 mesosoma lateral 182 mesosoma dorsal 183 propodeum and 1st–3rd metasomal tergites dorsal 184 fore femur lateral 185 hind femur lateral 186 head anterior 187 head dorsal 188 head lateral 189 outer hind tarsal claw 190 base of antenna 191 apex of antenna. https://binary.pensoft.net/fig/39007410.3897/zookeys.919.39642.figures192-195212724B8-BD435195-A895-702B10123BABFigures 192–195. Aleiodes coriaceus sp. nov., ♂, paratype 192 habitus lateral 193 inner hind claw lateral 194 3rd–7th metasomal tergites dorsal 195 3rd –7th metasomal tergites lateral. https://binary.pensoft.net/fig/390075

12. XML treatment for Aleiodes cruentus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964229199599-262E-5DC0-AB00C26D2EC2466D Aleiodes cruentus (Nees, 1834)Figs 196–198, 199–211, 212–216 Rogas cruentusNees, 1834: 212; Shenefelt 1975: 1222; Zaykov 1980a: 112; Kotenko 1992: 96. Rogas (Rogas) cruentus; Tobias 1976: 85, 1986: 80 (transl.: 130). Aleiodes (Neorhogas) cruentus; Papp 1985a: 156–157 (neotype designation), 1987b: 35, 1991a: 83; Belokobylskij 1996: 6; Riedel et al. 2002: 106. Aleiodes (Chelonorhogas) cruentus; Chen and He 1997: 39; Belokobylskij 2000: 32. Aleiodes cruentus; Bergamasco et al. 1995: 5; Belokobylskij et al. 2003: 398; Papp 2005: 176. Rhogas cruentus ab. nigricansFahringer, 1932: 238; Papp 1991a: 83 (invalid name). Rhogas cruentus ab. basalisHellén, 1927: 22 (invalid name). Rhogas cruentus ab. nigromaculataHellén, 1927: 22 (invalid name). Rhogas cruentus ab. rufofasciataHellén, 1927: 22 (invalid name). Rogas dorsalisHerrich-Schäffer, 1838: 154; Shenefelt 1975: 1222 (as synonym of A. cruentus); Papp 2005: 176 (id.). Rogas affinisHerrich-Schäffer, 1838: 124 (key only); Shenefelt 1975: 1174–1175 [neotype designated below]. Syn. nov. Aleiodes affinis; Belokobylskij et al. 2003: 398. Neotype of A. affinis here designated, ♀ (RMNH), “Museum Leiden, Nederland, Melissant (ZH), [at light], 10.viii.1980, K.J. Huisman”. It is important for nomenclatorial stability to fix our interpretation of A. affinis because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; the first author could not find any specimen in ZMB), the original description is rudimentary and there are very similar species in Europe. The specimen from Netherlands is selected because it fits best the original description, Netherlands is relatively close to the probable German (but unknown) type location and it is in good condition. Another complication is that the neotype of A. cruentus by Papp (1985) is an old male from uncertain origin in the Gravenhorst Collection (Wroclaw). Austria, Bulgaria, Croatia, Czech Republic, Finland, France, Germany, Greece, Italy (including Sicily), Moldova, Netherlands (FR: Ried, GE: Beusichem; Heerde; Voorst (Twello), LI: Thorn, NB: Eindhoven; Tilburg (Kaaistoep), OV: Buurse; Hasselt, ZH: Lexmond; Melissant; Middelharnis; Oostvoorne, ZL: Oostkapelle), Norway, Romania, Slovakia, Slovenia, Spain, Sweden, Ukraine, [Mongolia]. Specimens in ZJUH, BZL, FMNH, HSC, IKC, MSC, MTMA, NMS, NRS, RMNH, SDEI, ZSSM. Widespread in the region but rather sporadic. The specimen (CMIM) from which Morley (1915) recorded this species as new to Britain has been examined and proves to be A. alternator (Nees). A further specimen in CMIM recorded by Lyle (1919) as A. cruentus has been examined and belongs to A. diversus (Szépligeti), q. v., as do another three British specimens in ZJUH and one in NMS, and there is no evidence that A. cruentus has ever occurred in Britain. Probably univoltine, certainly overwintering as a mummy. Collected June-August, often at light and including around Dianthus barbatus harbouring larvae of the noctuid Hadena confusa (Hufnagel) (H. Schnee/Germany). In Austria it has been collected up to 2000 m. Only one reared specimen seen, from H. confusa [FMNH], the adult emerging in June in the year following host mummification. Extensive rearings of this host in various parts of Britain in recent years by one of us (MRS) has not produced A. cruentus, strengthening the view that it does not occur in Britain. The predominantly dark mummy seen (Fig. 198) is stout, rather short and weakly swollen dorsally, and has a paler and moderately strong lateral keel. The cocoon is substantially silk-lined and occupies most of the host’s abdomen (approx. 2nd–7th abdominal segments). The mummy probably forms underground, albeit from penultimate instar hosts, and the somewhat reflexed and sideways twisted head suggests that it is not or scarcely stuck down; the caudal segments are also somewhat recurved ventrally. Although oviposition has not been witnessed, the somewhat laterally compressed apex of the female’s metasoma appears to be an adaptation for attacking the host at rest or feeding within the seed capsules of its food plants (Dianthus, Silene, etc.). MRS558 (France), MRS624 (Germany), MRS625 (Germany). Maximum width of hypoclypeal depression (0.5–)0.6–0.7 × minimum width of face (Fig. 206); OOL of ♀ coarsely punctate and 0.5–0.8(–1.0) × diameter of posterior ocellus; ventral margin of clypeus (rather) obtuse apically and not protruding (Fig. 208), but sometimes intermediate; length of eye 1.5– 1.9 × temple in dorsal view; lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures; precoxal area with some rugae medially; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein r of fore wing 0.3–0.4 × vein 3-SR (Fig. 199); vein 1-CU1 of fore wing 0.8–1.1 × vein 2-CU1 (Fig. 199), rarely shorter; hind tarsal claws with conspicuous dark brown pecten (Fig. 205); 1st tergite widened apically; 2nd tergite 0.7–0.9 × as long as wide (Fig. 202), its colour variable, often reddish; head black; vein 1-M of fore wing brownish; wing membrane subhyaline; 4th–6th tergites of ♂ with long setae, but flattened and narrowly glabrous medially. Neotype of A. affinis, ♀, length of fore wing 7.3 mm, of body 10.2 mm. Head. Antennal segments of ♀ 61, length of antenna 1.2 × fore wing, its subapical segments rather robust; frons largely smooth and shiny, but rugulose near stemmaticum; OOL 0.6 × diameter of posterior ocellus, and coarsely punctate, interspaces approx. equal to diameter of punctures; vertex mainly densely punctate, shiny; clypeus coarsely punctate-rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 208); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 206); length of eye 1.9 × temple in dorsal view and temple rather long and densely setose (Fig. 207); vertex behind stemmaticum punctate-rugose; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view (Fig. 208). Mesosoma. Mesoscutal lobes densely and finely punctate, with satin sheen; precoxal area of mesopleuron with some rugae medially, rather densely punctate anteriorly and posteriorly; metapleuron mainly sparsely punctate, shiny; scutellum rather weakly punctate and slightly convex; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete and straight. Wings. Fore wing: r 0.4 × 3-SR (Fig. 199); 1-CU1 horizontal, as long as 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell rather short (Fig. 199); cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR wide; anterior half of subbasal and of subdiscal cells largely glabrous. Hind wing: basal half of marginal cell slightly widened, but apical half wide, apical width of cell 2.5 × width at level of hamuli (Fig. 199); 2-SC+R subquadrate; m-cu short and obsolescent; surroundings of M+CU and 1-

M glabrous; M+CU:1-M = 75:47; 1r-m 0.8 × 1-M. Legs. Tarsal claws with conspicuous and robust dark brown pecten (Fig. 205); hind coxa largely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.3 and 5.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with mediolongitudinal carina and largely coarsely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 202); 2nd suture deep medially, shallow laterally and crenulate; 2nd tergite 0.7 × as long as wide (Fig. 202); anterior 0.7 of 3rd tergite densely and finely punctate, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically rather rounded (Fig. 197). Colour. Black; posterior half of mesoscutum, scutellum largely, apical rim of 1st tergite and basal rim of 2nd tergite reddish brown; fore coxa, bases of middle and hind coxae blackish; apex of hind tibia, telotarsi, hind tarsus, palpi, veins and pterostigma dark brown; tegulae and remainder of hind tibia pale yellowish; remainder of legs reddish brown; wing membrane subhyaline. Variation. Vein 1-CU1 of fore wing 0.8–1.1 × as long as 2-CU1; mesoscutum, scutellum, metanotum, 1st and 2nd metasomal tergites are most often entirely reddish or orange brown but variably partly blackish, in particular 1st tergite sometimes with dark medial patch; pronotum and mesopleuron black or reddish dorsally; parastigma narrowly dark brown or yellowish brown; coxae entirely reddish to entirely dark brown. Antennal segments: ♀ 53(1), 55(1), 56(3), 57(5), 58(9), 59(9), 60(10), 61(9), 62(3), 63(1), 65(2), 67(1). ♂ 60(6), 61(7), 62(2), 63(5), 64(3), 65(1), 66(1), 67(5), 69(1). The males have on average approx. three more antennal segments than females. Males are very similar but often darker than females, 2nd tergite 0.9–1.0 × as long as basal width of tergite and apical tergites type 1 and (usually) type 2, with fringe present in the latter (Fig. 215); hind femur at most apically blackish, and hind tibial spurs sometimes blunt. *Austria, Bulgaria, Croatia, Czech Republic, Finland, France, Germany, *Greece, Italy, *Moldova, Mongolia, *Netherlands, Norway, *Romania, Slovakia, *Slovenia, Spain, Sweden, Ukraine. An examined female (NMS) from Albania (Mt Mali me Gropa, above Shengiergi, 1400 m, 13.viii.2019, MV light, C.W. Plant) has a CO1 sequence (MRS940) 3 % different from A. cruentus (19 differences in 626 bp of overlap) and although superficially similar in colour is clearly distinct in having OOL shorter (0.5 × lateral ocellus), a smaller hypoclypeal depression (0.5 × width of face), slenderer hind femur (5 × as long as wide), and several other differences. It may be A. parvicauda (Tobias, 1985) described from Afghanistan, but it has more (64; 58–60 in type series) and somewhat more elongate antennal segments than described for A. parvicauda, as well as other small deviations. Additional material as well as comparison with the type series of A. parvicauda are needed to settle the status of the Albanian species. 10.3897/zookeys.919.39642.figures196-198FD1C577A-699A-5351-9934-D041B273800AFigures 196–198. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg, but 198 from Finland, Mäntyharju 196 habitus lateral 197 ovipositor sheath lateral 198 mummy of Hadena confusa (Hufnagel). https://binary.pensoft.net/fig/39007610.3897/zookeys.919.39642.figures199-21167A8AF3F-595956F5-96CE-E198218BF417Figures 199–211. Aleiodes cruentus (Nees), ♀, Germany, Markkleeberg 199 wings 200 mesosoma lateral 201 mesosoma dorsal 202 metasoma dorsal 203 fore femur lateral 204 hind femur lateral 205 outer hind tarsal claw 206 head anterior 207 head dorsal 208 head lateral 209 base of antenna 210 apex of antenna 211 hind tarsus lateral. https://binary.pensoft.net/fig/39007710.3897/zookeys.919.39642.figures212-2160F129638-7BE65A59-9DF4-74DA62458E82Figures 212–216. Aleiodes cruentus (Nees), ♂, Hungary, Hársbokorhegy, but 213–215 from Germany, Markkleeberg 212 habitus lateral 213 1st–3rd metasomal tergites dorsal 214 4th–7th metasomal tergites lateral 215 id. dorsal 216 basal antennal segments. https://binary.pensoft.net/fig/390078ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980a) Unknown genera and species of Braconidae for the fauna of Bulgaria. Travaux scientifiques d’Université de Plovdiv “Paissi Hilendarski”, Biologie 15(4) [1977]: 111–115. [in Bulgarian]KotenkoAG (1992) A contribution to the fauna of Braconidae (Hymenoptera) of Dauria. In: Amirkhanov AM (Ed.) Insects of Dauria and adjacent territories: 94–107.Izdatelstvo Tsentralnoi Nauchno issledovatelskoi laboratorii okhotnichego khozyaistva i zapovednikov, Moskva, 141 pp.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1–12.RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97–108.ChenX-XHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2– 29.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.LyleGT (1919) Contributions to our knowledge of British Braconidae. No. 4 – Rhogadinae. Entomologist 49: 134– 136, 149–155, 178–181.

13. XML treatment for Aleiodes desertus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642737A772F-5088-5899-810503925C5ACB00 Aleiodes desertus (Telenga, 1941)Figs 217–220, 221–233 Rhogas (Rhogas) desertusTelenga, 1941: 184–185, 423 (not R. aestuosus var. desertus Telenga, 1941, from China) [examined]. Rogas desertus; Shenefelt 1975: 1223. Rogas (Rogas) desertus; Tobias 1986: 76 (transl. 124) (lectotype designation). Aleiodes desertus; Fortier and Shaw 1999: 230. Lectotype, ♀ (ZISP), “[Uzbekistan:] Khiva, 30.iv.[1]927, V. Gussakovskij/ S.Kh.Op.Ot., at light”, “Lectotypus Rogas desertus Tel., design. [V.I.] Tobias, 1980”. Paralectotypes: 1 ♀ (ZISP), “[Turkmenistan:] Ashkhabad [= Ashgabat], 25.iii.[1]905, S. Ahnger”, “Paralectotypus Rogas desertus Tel., design. [V.I.] Tobias, 1980”; 1 ♀ (ZJUH, figured), “Khiva, Rabat, 3.v.[1]927, V. Gussakovskij/collected at light”, “Paratypus Rogas desertus Telenga”, “Rec[eived] in exchange Academy of Science, Leningrad, B.M.1963-211”. None. Unknown. It seems to fly in spring (March–May) and may be univoltine. Maximum width of hypoclypeal depression 0.9–1.0 × minimum width of face; anterior part of clypeus very narrow, most of clypeus depressed (Fig. 229); OOL approx. 0.9 × diameter of posterior ocellus and remotely punctate; mandible massive triangular, coarsely punctate and with thick ventral lamella (Figs 229, 231); face largely transversely rugose; malar space 0.15 × as long as height of eye and 0.27 × basal width of mandible; area of precoxal sulcus (but posteriorly superficially) and

anteriorly area above it distinctly rugose; lateral lobes of mesoscutum largely smooth, strongly shiny and glabrous, middle lobe remotely punctulate and with satin sheen; basal half of wings (except anteriorly) largely glabrous and remainder of wing inconspicuously setose; vein r of fore wing 0.7– 0.8 × vein 3-SR (Fig. 221) vein 1-CU1 0.1 × as long as 2-CU1, narrow and oblique; tarsal claws long, slender, hardly bent and simple (Fig. 232); tarsal segments (except telotarsus) with four apical spines; 1st and base of 2nd tergite aciculate-rugulose, 3rd tergite micro-sculptured and matt, remainder of metasoma shiny and rather smooth; head and mesosoma (except prothorax anteriorly and mesoscutum posteriorly) black; pterostigma dark brown; legs and palpi pale yellowish. According to original description antenna of ♀ with 50–52 segments, but ZJUH paralectotype has 63 segments. Lectotype, ♀, length of fore wing 7.5 mm, of body 8.2 mm. Head. Antennal segments of ♀ more than 45, but apical segments missing, length of antenna of paralectotype 1.1 × body and its subapical segments moderately slender; frons rugose, shiny; OOL 0.9 × diameter of posterior ocellus; OOL and vertex remotely punctate, shiny; anterior part of clypeus 9 × wider than high, coarsely punctate and rather convex; clypeus above lower level of eyes; ventral margin of clypeus thick and not protruding forwards; width of hypoclypeal depression 0.9 × minimum width of face (Fig. 229); length of eye 1.7 × temple in dorsal view (Fig. 230); vertex behind stemmaticum convex and sparsely punctate; length of malar space 0.15 × length of eye in lateral view; mandible massive triangular, coarsely punctate and with thick ventral lamella (Figs 229, 231); occipital carina nearly complete, fine and ventrally strongly curved. Mesosoma. Lateral lobes of mesoscutum largely smooth, strongly shiny and glabrous, middle lobe

remotely punctulate and with satin sheen; prepectal carina complete and lamelliform; precoxal area of mesopleuron widely rugose, but posterior 0.2 narrowly striate; mesopleuron above precoxal area anteriorly rugose and remainder weakly and sparsely punctate, shiny; axilla crenulate but posteriorly densely and coarsely rugose; scutellum largely smooth, with some punctures; propodeum evenly convex, finely rugose and with strong medio-longitudinal carina, without tubercles. Wings. Fore wing: basal half largely glabrous; r 0.7 × 3-SR (Fig. 219); 1-CU1 oblique, 0.1 × as long as 2-CU1; r-m nearly as long as 3-SR; 2nd submarginal cell comparatively short (Fig. 221); cu-a inclivous; 1-M nearly straight posteriorly. Hind wing: basal 0.4 of marginal cell slightly widened and distally strongly widened, its apical width 2.7 × width at level of hamuli (Fig. 222); 2-SC+R subquadrate; m-cu indistinct; M+CU:1-M = 3:2; 1r-m 0.8 × 1-M. Legs. Tarsal claws slender, slightly curved and only setose (Fig. 232); hind coxa partly obliquely striate dorsally; tarsi slender, segments (except telotarsus) with long apical spines; length of hind femur and basitarsus 5.0 and 6.8 × their width, respectively; length of inner hind spur 0.3 × hind basitarsus. Metasoma. First tergite robust, 0.9 × longer than wide apically, strongly narrowed anteriorly (Fig. 225) and rather flat posteriorly; 1st and 2nd tergites finely longitudinally striate-rugulose; mediolongitudinal carina of 1st and 2nd tergites indistinct; 2nd tergite 0.6 × longer than its basal width; medio-basal area of 2nd tergite wide triangular, rather short; 2nd suture shallow and narrow; 3rd tergite matt and micro-sculptured, anteriorly finely striate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath with rather short setae and apically truncate (Fig. 220). Colour. Black; mesoscutum posteriorly partly chestnut brown; antenna, clypeus, malar space ventrally, mandible, pronotum and propleuron anteriorly and metasoma, brownish yellow; tegulae, legs and palpi pale yellowish; pterostigma and ovipositor sheath dark brown; veins of fore wing (but pale in basal 0.3 of fore wing) brown; wing membrane hyaline. Variation. Length of body 7.0–8.2 mm, of fore wing 7.5–7.9 mm; temple punctate to smooth; precoxal sulcus area finely to rather coarsely rugose; pronotal side largely black (except ventrally)

black or brownish yellow; lateral lobes of mesoscutum entirely dark chestnut brown or only posteriorly so, or mesoscutum largely yellowish brown posteriorly and prolonged to base of notauli; first tergite usually entirely brownish yellow, but sometimes dark brown and only posteriorly and laterally yellowish; pterostigma dark brown or brown. Antennal segments: ♀ 63(1). Turkmenistan, Uzbekistan. We have included this extralimital species from Central Asia because we suspect it may occur in Turkey. It should not be confused with Rogas aestuosus var. desertus Telenga, 1941, described from China in the same paper. The latter is an unavailable name (a primary homonym) and most likely a colour variety of R. aestuosus. 10.3897/zookeys.919.39642.figures217-220A9F2DD8D-79C8-55F9-BFD6-3977FBCC6C38Figures 217–220. Aleiodes desertus (Telenga), ♀, paralectotype 217 habitus lateral 218 antenna lateral 219 detail of fore wing 220 ovipositor sheath lateral. https://binary.pensoft.net/fig/39007910.3897/zookeys.919.39642.figures221-23350B12841-BF885E90-B1DE-4330FDF445C8Figures 221–233. Aleiodes desertus (Telenga), ♀, paralectotype, but 224 and 230 of lectotype 221 fore wing 222 hind wing 223 mesosoma lateral 224 mesosoma dorsal 225 metasoma dorsal 226 fore femur lateral 227 hind femur lateral 228 apex of antenna 229 head anterior 230 head dorsal 231 head lateral 232 outer hind tarsal claw 233 base of antenna. https://binary.pensoft.net/fig/390080ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.

14. XML treatment for Aleiodes dissector. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642446B0A23-445D-52A8-905F648AEDEF5326 Aleiodes dissector (Nees, 1834)Figs 234–239, 240–242, 243–255, 256–263 Rogas dissectorNees, 1834: 208; Shenefelt 1975: 1225–1226; Papp 1977a: 110. Rogas (Rogas) dissector; Tobias 1976: 81, 1986: 75 (transl.: 121). Aleiodes (Neorhogas) dissector; Papp, 1985a: 145, 1987b: 35, 1991a: 74, 1991d: 5, 1999: 550; Belokobylskij, 1996: 9; Riedel et al., 2002: 106. Aleiodes (Chelonorhogas) dissector; Belokobylskij 2000: 34; Ku et al. 2001: 234, 235. Aleiodes dissector; Fortier and Shaw 1999: 230; Belokobylskij et al. 2003: 398; Zaldívar-Riverón et al. 2004: 234, 2008: 392; Papp 2005: 176; Belokobylskij et al. 2008: 136–137. Phylax aestivalisSnellen van Vollenhoven, 1858: 282; Shenefelt, 1975: 1226 (as synonym of A. dissector); van Achterberg 1992: 363 (id.); Papp 2005: 176 (id.) [examined]. Holotype of A. aestivalis, ♀ (RMNH), “[Netherlands], Haag [= near The Hague], 6 [= June], v.Voll.”. According to the original description the ♂ holotype of R. dissector from Germany should be in the Gravenhorst collection (Museum of Natural History, University of Wrocław, Wrocław), but so far it has not been found. Austria, British Isles (England: V.C.s 15, 17, 20, 22, 23, 24, 30, 31, 34, 37, 58; Scotland: V.C.s 73, 88, 89, 95, 96, 97, 107), Croatia, Czech Republic, Finland, France, Germany, Greece, Hungary, Montenegro, Italy, Netherlands (FL: Lelystad, GE: Barneveld, OV: Raalte (Heino), ZH: Wassenaar), Norway, Russia, Serbia, Slovakia, Switzerland, Ukraine, [Armenia]. Specimens in ZJUH, BZL, CNC, IKC, MRC, MSC, MSNV, MTMA, NMS, OUM, RMNH, SDEI, UNS, UWIM, ZSSM. MRS007 (Turkey), MRS025 (Turkey), MRS145 (UK), MRS146 (UK). Univoltine, collected in May and June in deciduous scrub and woodland. In Britain it is widespread but particularly common in birch-dominated woodland in upland Scotland. Reared from the noctuids Orthosia incerta (Hufnagel) (17, M.R. Shaw), O. gothica (Linnaeus) (1, J.L. Yela) and Orthosia sp. (3), overwintering in the concealed mummy. An additional specimen, lacking a mummy but labelled as reared doubtfully from the sesiid Paranthrene tabaniformis (Rottemburg) ( RMNH), which normally feeds under Populus bark at ground level or below, can be discounted as a probable substrate rearing in which the mummy of the true host was overlooked. Parasitised host larvae in their penultimate instar leave their feeding sites and enter the soil or other site of moderate concealment (including below loose bark), where they prepare a chamber as though to pupate. At this time the parasitoid larva within the strongly retarded host (Fig. 234) is around half its final length, and the host lies quiescent for approx. a week until the parasitoid has completed its feeding (Fig. 235). During mummification (Figs 236, 237) the caudal end of the host recurves ventrally as the host’s body becomes weakly retracted. A ventral opening at the head end is made, but the head (as with the caudal segments) is usually tucked downwards rather than becoming raised, and so the resulting expelled fluid (Fig. 237) usually dries without the mummy becoming stuck down. The eventual outcome is a rather distinctive (Fig. 238) elongate and curved dark brown structure with a paler and weakly raised lateral keel. The parasitoid’s pupation chamber occupies ca 2nd–8th abdominal segments of the host, which are moderately strongly lined with silk (Fig. 239). The moderately large hypoclypeal opening and protruding sharp-rimmed clypeus of A. dissector is seen in some other species (e.g., A. modestus (Reinhard), treated in part 1 of this work) whose hosts also pupate in shallow soil. In culture experiments A. dissector was found to prefer hosts in the early to middle part of the 3rd instar, although late 2nd instar host were often also acceptable. Oviposition into suitable hosts was rapid (1–2 seconds) and accomplished with a single insertion of the ovipositor, following only brief antennation and no use of the legs. There was no clear temporary paralysis. Experimental rearings from O. incerta (6:107\85\\75+10) and O. gothica (6:61\49\\34+15) were comparable (given that some insertions of less than a full second might have been scored as ovipositions incorrectly; and furthermore that some failures to oviposit into these hosts might be ascribed to temporary egg depletion, as the protocol of normally ceasing to offer hosts to a particular female after four apparent ovipositions on the day had not been developed until after the experiments were undertaken), and clearly demonstrated the suitability of both species as hosts. In contrast, no parasitoids developed (and indeed possibly no ovipositions occurred) in the other species of Orthosia tested, which were all found to be clearly outside the host range: O. cerasi (Fabricius) (3:32\?3\\0+3); O. cruda (Denis & Schiffermüller) (2:12\0\\-); O. munda (Denis & Schiffermüller) (3:10\0\\-); O. gracilis (Denis & Schiffermüller) (2:11\?1\\0+1). Of these four, only O. gracilis is not fully arboreal. There is no adverse venom effect on host development. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 251); OOL of ♀ 0.6–0.7 × diameter of posterior ocellus (Fig. 252) and sparsely punctate; ventral margin of anterior part of clypeus comparatively sharp and more or less protruding outwards (Fig. 253); length of malar space 0.2 × length of eye in lateral view (Fig. 253); head transverse in dorsal view and eye 1.5–2.0 × as long as temple in dorsal view (Fig. 252); lobes of mesoscutum punctulate, with interspaces smooth to superficially micro-sculptured; precoxal area completely smooth or nearly so; vein 1-CU1 of fore wing 0.2–0.3 × vein 2-CU1 and horizontal (Fig. 243); hind tarsal claws with conspicuous dark brown pecten close to apical tooth (Fig. 250); 1st tergite rounded antero-laterally and 1.0–1.1 × as long as wide apically; basal half of metasoma black and weakly sculptured; 3rd tergite smooth; head black; palpi yellowish; basal half of hind tibia pale yellowish, but in some males almost uniformly dark; 4th–6th tergites of males depressed medially and conspicuously setose (Fig. 258). Redescribed ♀ (RMNH) from Austria (Burgenland, Winden am See). Length of fore wing 8.5 mm, of body 9.0 mm. Head. Antennal segments of ♀ 60, antenna as long as fore wing, its subapical segments rather slender, slightly longer than wide; frons largely smooth; OOL 0.7× diameter of posterior ocellus, sparsely punctate, shiny and with deep groove near posterior ocellus (Fig. 252); vertex sparsely punctate, rather shiny; clypeus coarsely punctate; ventral margin of clypeus rather thin and forward protruding (Fig. 253); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 251); length of eye 1.5 × temple in dorsal view (Fig. 252); vertex behind stemmaticum superficially rugose-punctate; clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Mesoscutal lobes punctulate with interspaces superficially micro-sculptured and shiny; precoxal area of mesopleuron smooth except some punctulation, mesopleuron punctulate anteriorly and posteriorly; metapleuron densely punctate; metanotum with nearly complete median carina;

scutellum flat (but with rugulose depression medio-posteriorly), remainder punctulate and with weak lateral carinae; propodeum evenly convex and coarsely rugose, and medio-longitudinal carina absent posteriorly. Wings. Fore wing: r 0.4 × 3-SR; 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.3 × 3-SR; 2nd submarginal cell medium-sized (Fig. 243); cu-a inclivous, straight; 1-M straight posteriorly; 1-SR medium-sized; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell rather narrow basally, apical half gradually widened, its apical width 3.1 × width at level of hamuli (Fig. 244); 2SC+R subquadrate; m-cu absent; M+CU:1-M = 35:33; 1r-m 0.7 × 1-M. Legs. Tarsal claws with conspicuous and robust dark brown pecten (Fig. 250); hind coxa distinctly punctate and with some oblique striae postero-dorsally; hind trochantellus robust; length of hind femur and basitarsus 3.8 and 5.3 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite flattened, basally narrowed, as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and largely finely punctate-rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite wide and triangular, distinct (Fig. 247); 2nd suture rather deep and micro-sculptured; 3rd and subsequent tergites largely smooth; apical half of 3rd and 4th tergites without sharp lateral crease; ovipositor sheath wide, with long and medium-sized setae and apically truncate (Fig. 241). Colour. Black; apical half of hind tibia and hind tarsus blackish; basal half of hind tibia pale yellowish; remainder of legs, palpi and tegulae yellowish brown; most veins and pterostigma dark brown; wing membrane slightly yellowish basally and remainder slightly infuscate. Variation. Interspaces between punctulation of mesoscutum smooth to superficially microsculptured; medio-longitudinal carina of propodeum complete or absent posteriorly; 3rd metasomal tergite largely finely sculptured (except posteriorly) to largely smooth; mesopleuron black or with brownish longitudinal stripe; hind tibia usually ivory or pale yellowish basally. Antennal segments: ♀ 51(2), 55(2), 56(7), 57(4), 58(7), 59(12), 60(14), 61(18), 62(6), 63(4); ♂ 51(1), 53(2), 54(2), 55(4), 56(8), 57(29), 58(29), 59(27), 60(15), 61(4), 62(4), 63(2), 64(2). Females have on average ca one to two more antennal segments than males. Males are very similar but hind femur more or less blackish and, in some males, hind tibia almost uniformly dark, OOL approx. as long as diameter of posterior ocellus (Fig. 257) and apical tergites type 3–4 with fringe long and strong (Figs 258, 260). *Armenia, *Austria, British Isles (England, Scotland), Croatia, Czech Republic, Finland, France, Germany, *Greece, Hungary, *Montenegro, *Italy, Netherlands, Norway, Russia, *Serbia, Switzerland, Ukraine. 10.3897/zookeys.919.39642.figures234-239FCCA41AD-56CD-5956-81E0-E2E97F9A43E4Figures 234–239. Aleiodes dissector (Nees), U.K., Scotland (in culture) parasitising Orthosia incerta (Hufnagel) 234 pre-mummy, removed from its hideaway, with unparasitised control from the same egg batch (below) 235 pre-mummy 236 early mummification 237 mummy with ventral ooze 238 three fully hard mummies 239 emerged mummy, cut open to expose silken lining. https://binary.pensoft.net/fig/39008110.3897/zookeys.919.39642.figures240-242A06B8684-54995502-94C4-7783632397FDFigures 240–242. Aleiodes dissector (Nees), ♀, Switzerland, Tessin, but 242 from Scotland (culture) 240 habitus lateral 241 ovipositor sheath lateral 242 mummy of Orthosia incerta (Hufnagel). https://binary.pensoft.net/fig/39008210.3897/zookeys.919.39642.figures243-25596EDCBD0-E8975243-9ED2-DB847C7BA91BFigures 243–255. Aleiodes dissector (Nees), ♀, Switzerland, Tessin 243 fore wing 244 hind wing 245 mesosoma lateral 246 mesosoma dorsal 247 propodeum and 1st –4th metasomal tergites dorsal 248 fore femur lateral 249 hind femur lateral 250 outer hind tarsal claw 251 head anterior 252 head dorsal 253 head lateral 254 base of antenna 255 apex of antenna. https://binary.pensoft.net/fig/39008310.3897/zookeys.919.39642.figures256-2630C522D70-D50853F0-9482-B9A05895953EFigures 256–263. Aleiodes dissector (Nees), ♂, Austria, Kärnten 256 habitus lateral 257 head dorsal 258 3rd–7th tergites lateral 259 mesosoma dorsal 260 1st–6th metasomal tergites dorsal 261 fore femur lateral 262 head anterior 263 hind femur lateral. https://binary.pensoft.net/fig/390084ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1977a) Contributions to the Braconid fauna of Yugoslavia. III.Folia Entomologica Hungarica30: 105–117.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97–108.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.KuDSBelokobylskijSAChaJY (2001) Hymenoptera (Braconidae). Economic Insects of Korea 16. Insecta Koreana, Suppl.23: 1– 283.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.ZaldívarRiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xZaldívarRiverónAShawMRSáezAGMoriMBelokobylskijSAShawSRQuickeDLJ (2008) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Biology 8: 329. https://doi.org/10.1186/1471-2148-8-329PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.BelokobylskijSAZaldívarRiverónAMaetôKSaezAG (2008) Asian Betylobraconinae (Hymenoptera, Braconidae), with description of a new genus and phylogenetic affinities of the tribe Facitorini.Insect Systematics and Evolution39(2): 133–154. https://doi.org/10.1163/187631208788784110van AchterbergC (1992) Revisionary notes on the subfamily Homolobinae (Hymenoptera: Braconidae).Zoologische Mededelingen, Leiden66(25): 359–368.

15. XML treatment for Aleiodes diversus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642CF7B3762-3684-53C6-8D8CC0F3B0486369 Aleiodes diversus (Szépligeti, 1903)Figs 264–265, 266–277, 278–288 Rhogas

diversusSzépligeti, 1903: 114; Papp 2004: 216 (as synonym of A. dissector) [examined]. Rogas dissector var. diversus; Shenefelt 1975: 1226 (lectotype designation). Aleiodes (Neorhogas) diversus; Papp 1977a: 110–112 (re-instated), 1985: 145, 1991a: 81. Aleiodes diversus; Belokobylskij et al., 2003: 398 (as synonym of A. dissector); Papp 2005: 176; Merz and Pasche 2012: 244; van Achterberg 2014: 209.

Lectotype, ♀ (MTMA), “Croatia, Buccari [= Bakar], 1893, Pavel”, “Lectotypus, ♀”, Rogas (s. str.) diversus Szépligeti, 1906 [sic!], Papp, 1968”, “Hym. Typ. No. 1011, Mus. Budapest”. Austria, British Isles (England: V.C.s 8, 25, 70), Bulgaria, Croatia, Hungary, Italy (Sicily), Norway, Switzerland. Specimens in ZJUH, BZL, CMIM, MHNG, MRC, MTMA, NMS, RMNH, SDEI, ZSSM. The most recent of the five English specimens seen is dated 1931, and it seems likely that this rather large and showy insect is extinct in Britain. None. Unknown. Female specimens have been collected in (May–)June, and also September, suggesting that it may be plurivoltine. This is reinforced by the date of capture of the two available males (which would not have hibernated as an adult) in Sicily on 30.iv.1965 (ZJUH) and 1.v.1994 (RMNH). There is no indication of habitat on data labels and we have not seen reared material. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 273); OOL of ♀ coarsely punctate and 1.0–1.2 × diameter of posterior ocellus; ventral margin of clypeus (rather) obtuse apically and clypeus not protruding outwards (Fig. 275), but sometimes intermediate; length of eye 1.0–1.2 × temple in dorsal view; lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures, shiny and smooth; precoxal area with some rugae medially; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein 1-CU1 of fore wing 0.7–1.1 × vein 2-CU1 and approx. as long as vein m-cu (Fig. 266), rarely shorter; hind femur 3.0–3.3 × longer than wide; hind tarsal claws with medium-sized dark brownish pecten up to apical tooth (Fig. 272); 1st tergite widened apically and moderately wide basally (Fig. 269); 2nd tergite 0.7–0.8 × as long as wide (Fig. 269) and black; 4th–7th tergites of males flat and with long yellowish setae (Figs 279, 282); head black; vein 1-M of fore wing brownish; wing membrane subhyaline. Lectotype, ♀, length of fore wing 7.0 mm, of body 10.0 mm. Head. Antennal segments of ♀ 56, antenna as long as fore wing, its subapical segments robust; frons largely smooth behind antennal sockets; OOL 1.2 × diameter of posterior ocellus, and coarsely punctate, interspaces less than diameter of puncture; vertex coarsely punctate; clypeus rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 275); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 273); length of eye 1.2 × temple in dorsal view (Fig. 274); vertex behind stemmaticum superficially punctate-rugose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, interspaces largely smooth, shiny; precoxal area of mesopleuron coarsely punctate and without rugae medially, mesopleuron coarsely punctate anteriorly and posteriorly; metapleuron moderately punctate; scutellum remotely punctate; propodeum rather convex and coarsely rugose. Wings. Fore wing: r 0.5 × 3-SR (Fig. 266); 1-CU1 horizontal, 0.7 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell rather long (Fig. 266); cu-a vertical, straight; 1-M rather curved posteriorly; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened, its apical width 2.3 × width at level of hamuli (Fig. 266); 2-SC+R transverse; m-cu largely absent, only as short antefurcal remnant (Fig. 266); M+CU:1-M = 35:23; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather conspicuous, medium-sized dark brown pecten up to apical tooth (Fig. 272); hind coxa largely punctate; hind trochantellus robust; length of hind femur and basitarsus 3.1 and 4.4 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with mediolongitudinal carina and coarsely vermiculate-rugose; medio-basal area of 2nd tergite triangular and distinct (Fig. 269); 2nd suture deep; 2nd tergite 0.7–0.8 × as long as wide (Fig. 269); 3rd tergite

densely punctate and interspaces largely smooth, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; apical third of metasoma rather compressed; ovipositor sheath wide, with rather short setae and apically truncate (Fig. 265). Colour. Black; mesoscutum (except anterior third), scutellum, clypeus ventrally, mandible, tegulae and legs largely brownish red; palpi, fore coxa largely, telotarsi, hind tarsus and apex of hind tibia (excluding spurs) dark brown; pterostigma blackish brown; veins dark brown, but near wing base yellowish; wing membrane slightly infuscate. Variation. OOL 1.0–1.2 × diameter of posterior ocellus; mesoscutum of ♀ entirely brownish red or yellowish brown, or anteriorly black; 1st tergite 1.0–1.1 × longer than wide apically; metasoma rarely partly obscurely reddish dark brown; mesopleuron may be just punctate or may have some rugae in lower half. Antennal segments ♀: 55(3), 56(3), 57(3), 58(1), 59(1); ♂ 58(1). Males have mesosoma black (Fig. 278), 2nd tergite 0.8–0.9 × as long as basal width of tergite (Fig. 282) and apical tergites type 2, setae rather long, fringe long and strong (Fig. 279). *Austria, *British Isles (England; probably extinct), *Bulgaria, Croatia, Hungary, *Italy (Sicily), *Norway, *Switzerland. Close to A. cruentus which, however, almost always has much or all of 1st and 2nd metasomal tergites orange-red (usually wholly black or dark brown in A. diversus). In addition to characters given in the key A. diversus is a more robust insect, and females have broader antennal segments (distinctly transverse near middle of flagellum) and on average they are fewer in number (although with overlap). 10.3897/zookeys.919.39642.figures264-2650DB13A2A-D8B1-5BE7-9F86-A455F19B96C4Figures 264, 265. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 264 habitus lateral 265 ovipositor sheath lateral. https://binary.pensoft.net/fig/39008510.3897/zookeys.919.39642.figures266-2775C60A2CD-BA745F54-B86B-AAEDBEB6D781Figures 266–277. Aleiodes diversus (Szépligeti), ♀, Italy, Sicily 266 wings 267 mesosoma lateral 268 mesosoma dorsal 269 propodeum and metasoma dorsal 270 fore femur lateral 271 hind femur lateral 272 outer hind tarsal claw 273 head anterior 274 head dorsal 275 head lateral 276 base of antenna 277 apex of antenna. https://binary.pensoft.net/fig/39008610.3897/zookeys.919.39642.figures278-288447739B7-76B95CD8-89E5-715011AEF82AFigures 278–288. Aleiodes diversus (Szépligeti), ♂, Italy, Sicily 278 habitus lateral 279 3rd–7th tergites lateral 280 wings 281 mesosoma lateral 282 metasoma dorsal 283 outer hind claw 284 hind tibial spurs lateral 285 head anterior 286 head dorsal 287 base of antenna 288 apex of antenna. https://binary.pensoft.net/fig/390087PappJ (2004) Type specimens of the braconid species by Gy. Szépligeti deposited in the Hungarian Natural History Museum (Hymenoptera: Braconidae).Annales Historico-Naturales Musei Nationalis Hungarici96: 153–223.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1977a) Contributions to the Braconid fauna of Yugoslavia. III.Folia Entomologica Hungarica30: 105– 117.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.MerzBPascheA (2012) 8.31.10. Superfamily Ichneumonoidea (Braconidae): 241–244. In: MerzB (Ed.) Liste annotée des insectes (Insecta) du canton de Genève.Instrumenta Biodiversitatis8: 1–532. Genève.van AchterbergC (2014) Notes on the checklist of Braconidae (Hymenoptera) from Switzerland.Mitteilungen der Schweizerischen Entomologischen Gesellschaft87: 191–213.

16. XML treatment for Aleiodes eurinus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642596C7289-B8C1-5E50-AA277E3F46A5D296 Aleiodes eurinus (Telenga, 1941)Figs 289–291, 292–302, 303–306 Rhogas (Rhogas) eurinusTelenga, 1941: 422. Rogas eurinus; Shenefelt 1975: 1228; Papp 1971: 359. Rogas (Rogas) eurinus; Tobias 1976: 85, 1986: 80 (transl.: 130; lectotype designation). Aleiodes (Neorhogas) eurinus; Papp 1985a: 145; 1991a: 94. Aleiodes (Chelonorhogas) eurinus; Chen and He, 1997: 39; He et al. 2000: 667; Belokobylskij 2000: 49; Ku et al. 2001: 235; Farahani et al. 2015: 242–243; Beyarslan et al. 2017: 330. Aleiodes eurinus; Fortier and Shaw 1999: 223, 230; Belokobylskij et al. 2003: 398; Papp 2005: 176. Rogas eurinus ab. nigratusPapp, 1967: 223 (invalid name). Rogas eurinus ab. nigrimaculatusPapp, 1967: 223 (invalid name). Rogas eurinus ab. nigripesPapp, 1967: 223 (invalid name). None seen. Italy, Russia (Siberia and Far East), Spain, Turkey, [China, Mongolia]. Specimens in ZJUH, BZL, MRC, MSNV, MTMA, NMS, RMNH, SDEI, ZISP. None. Specimens have been collected from April to August, and the presence of males in both April and July clearly demonstrates that it is plurivoltine and overwinters in the mummy. We have not seen reared material, but specimen labelling indicates that it occurs among Ammophila and Schoenus in the Venice Lido and Triticum (presumably cultivated wheat) in Turkey, suggesting that its hosts will occur in open grassland habitats. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 298); OOL of ♀ approx. as long as diameter of posterior ocellus (Fig. 299) and densely rugose; clypeus rather thin apically and rather protruding anteriorly (Fig. 300); eyes prominent (Fig. 299); lobes of mesoscutum distinctly punctate-granulate and rather matt; precoxal area more or less rugose and comparatively wide medially, and posteriorly punctate; vein 1-CU1 of fore wing 0.3 × vein 2-CU1 (Fig. 292); hind tarsal claws slender, brownish setose and without pecten (Fig. 302); basal half of 3rd tergite striate; 3rd antennal segment of ♀ dark brown; basal half of hind tibia pale yellowish or ivory, at least inner side contrasting with reddish or dark brown colour of basal half of hind femur (usually less pronounced in ♂). Redescribed ♀ (RMNH) from Turkey (Ankara). Length of fore wing 7.0 mm, of body 8.0 mm. Head. Antennal segments of ♀ 51 remaining, but apical segments missing, length of antenna 1.2 × fore wing; frons with coarse curved rugae and dorsally coarsely rugose; OOL equal to diameter of posterior ocellus, and densely rugose; vertex spaced rugose, rather dull; clypeus medium-sized and coarsely rugose (as face); ventral margin of clypeus rather thin and rather protruding forwards (Fig. 300); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 298); length of malar space in anterior view 0.7–1.0 × maximum width of hypoclypeal depression (Fig. 298); head in anterior view trapezoid; length of eye 1.3 × temple in dorsal view and temples directly narrowed behind eyes (Fig. 299); vertex behind stemmaticum densely rugose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes distinctly punctate-granulate, and with satin sheen; precoxal area of mesopleuron coarsely rugose, rather wide medially and posteriorly coarsely punctate and some short rugae, densely punctate; remainder of mesopleuron mainly sparsely and finely punctate; metapleuron densely punctate; metanotum with nearly complete median carina; scutellum punctulate and weakly granulate; propodeum coarsely vermiculate-rugose, medio-longitudinal carina irregular. Wings. Fore wing: r 0.4 × 3-SR (Fig. 292); m-cu far antefurcal; 1-CU1 horizontal, slightly widened, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 292); cu-a inclivous, somewhat curved posteriorly; 1-M rather curved posteriorly; 1-SR wide; surroundings of M+CU1, 1M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened, its apical width 2.6 × width at level of hamuli (Fig. 292); 2-SC+R subquadrate; m-cu medium-sized and only pigmented; M+CU:1-M = 50:43; 1r-m 0.7 × 1-M. Legs. Tarsal claws slender and brownish setose (Fig. 302); hind coxa rather finely and densely punctate; hind trochantellus robust; length of hind femur and basitarsus 5.0 and 6.5 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus.

Metasoma. First tergite rather flattened, 1.1 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and rather regularly longitudinally rugose; medio-basal area of 2nd tergite narrow triangular (Fig. 295); 2nd suture deep and crenulate; basal half of 3rd tergite longitudinally striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 290). Colour. Black; palpi and basal half of antenna (except scapus and pedicellus) brown; scapus, pedicellus, clypeus largely, apex of hind femur (but ventrally reddish), apex of hind tibia, hind tarsus, all telotarsi, pterostigma (but basally narrowly pale) and veins (except yellowish veins of basal quarter of wings) dark brown; remainder of legs and 1st–3rd tergites orange brown; tegulae and hind tibia (except apically) pale yellowish; wing membrane subhyaline. Variation. Coxae and hind femur (except its basal third) largely dark brown, black or orange brown; apical half of hind tibia dark brown or only apically so; 1st tergite largely dark brown (except posteriorly), with pair of dark brown spots or entirely orange or reddish brown; apical half of 3rd tergite orange brown or largely black. Antennal segments: ♀ 54(1), 55(2), 57(2), 58(3), 59(1), 60(2); ♂ 52(1), 60(1). Male is very similar and has apical tergites type 1–2, setae moderately dense, glabrous stripe only rarely evident and fringe very short, negligible (Figs 303, 305). China, *Italy, Mongolia, Russia (Siberia and Far East), Spain, *Turkey. 10.3897/zookeys.919.39642.figures289-291B35C50E5-1C72-516F-87E7-1AF88006BC52Figures 289–291. Aleiodes eurinus (Telenga), ♀, Mongolia, but 291 Russia, Chelyabinskoi Obl. 289 habitus lateral 290 ovipositor sheath lateral 291 apex of antenna. https://binary.pensoft.net/fig/39008810.3897/zookeys.919.39642.figures292-30214384170-9C0F50F2-AEC9-1635D7CEC291Figures 292–302. Aleiodes eurinus (Telenga), ♀, Mongolia 292 wings 293 mesosoma lateral 294 mesosoma dorsal 295 1st–3rd metasomal tergite dorsal 296 fore femur lateral 297 hind femur lateral 298 head anterior 299 head dorsal 300 head lateral 301 base of antenna 302 outer hind tarsal claw. https://binary.pensoft.net/fig/39008910.3897/zookeys.919.39642.figures303-3060C5A40D0-4F6053E8-B8A9-05C48B115355Figures 303–306. Aleiodes eurinus (Telenga), ♂, Mongolia 303 habitus lateral 304 head anterior 305 apical half of metasoma lateral 306 head dorsal. https://binary.pensoft.net/fig/390090ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1971) Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 215. Braconidae (Hym.) 3.Annales Historico-Naturales Musei Nationalis Hungarici63: 307–363.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512– 588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.KuDSBelokobylskijSAChaJY (2001) Hymenoptera (Braconidae). Economic Insects of Korea 16. Insecta Koreana, Suppl.23: 1– 283.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2BeyarslanAGözüaçikCGüllüMKonuksalA (2017) Taxonomical investigation on Braconidae (Hymenoptera: Ichneumonoidea) fauna in northern Cyprus, with twenty six new records for the country.Journal of Insect Biodiversity and Systematics3(4): 319–334.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.

17. XML treatment for Aleiodes fahringeri. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964269C3C3F4-C019-54A9-BCE3C4C0073CBB92 Aleiodes fahringeri (Telenga, 1941)Figs 307–309, 310–321 Rhogas (Rhogas) fahringeriTelenga, 1941: 173. Rogas fahringeri; Shenefelt 1975: 1228. Rogas (Rogas) fahringeri; Papp 1977b: 113. Aleiodes (Chelonorhogas) fahringeri; Chen and He 1997: 40; He et al. 2000: 666; Belokobylskij 2000: 39 (lectotype designation); Papp 2009: 149. Aleiodes fahringeri; Chen and He, 1992: 125; Fortier and Shaw 1999: 230. Rhogas (Rhogas) flavipennisTelenga, 1941: 174, 419. Rogas flavipennis; Shenefelt 1975: 1229. Aleiodes (Chelonorhogas) flavipennis; Belokobylskij 2000: 39 (lectotype designation and synonymised with A. fahringeri (Telenga, 1941)). None examined.

3 ♀ (MTMA, NMS, RMNH), “Mongolia: Südgobi aimak, Somon Bulgan, Talyn bulag, 1350 m, Exp. Dr. Z. Kaszab, 1967”, “Nr. 889, 5.vii.1967”, “Rogas fahringeri Tel., ♀, det. Papp, 1976”; 1 ♀ (RMNH, ZJUH), “China: Ningxia, Yinchuan, 6.vii.1983, no. 840994, Xu Wenzhong, RMNH’99”. None. Unknown. Specimens have been collected in June–August. Presumed to be univoltine, but we have not seen reared material and the means of overwintering is unclear. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face; OOL 0.9 × diameter of posterior ocellus, largely smooth with spaced punctures; ventral margin of clypeus thin, anterior part shiny and distinctly protruding anteriorly (Fig. 320); mesoscutum shiny and moderately punctulate; precoxal area with only some rugulae medially; vein r of fore wing 0.5–0.6 × as long as vein 3-SR; tarsal claws rather slender and with yellowish or brown pecten, pecten remains removed from apical tooth (Fig. 321); hind tarsus fairly elongate and segment with medium-sized apical

spines (Figs 307, 321); pterostigma brownish yellow; wings subhyaline; head and mesosoma laterally and dorsally (except more or less dark brown propodeum) yellowish brown; fore wing longer than 5 mm. Redescribed ♀ (RMNH) from Mongolia (Somon Bulgan). Length of fore wing 6.9 mm, of body 7.7 mm. Head. Antennal segments of ♀ 58, length of antenna 1.1 × fore wing, its basal and subapical segments slender (Figs 316, 317); frons largely smooth anteriorly and rugulose posteriorly; OOL 0.9 × diameter of posterior ocellus, largely smooth with spaced punctures, (but superficially coriaceous near eye) and with satin sheen; vertex largely smooth, but superficially rugulose behind ocelli; anterior part of clypeus nearly 5 × wider than long, medially distinctly wider than laterally, largely smooth, punctulate, its ventral margin thin and protruding forwards (Fig. 320); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 318); length of eye 1.6 × temple in dorsal view (Fig. 319); clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Pronotum medio-dorsally flat, shiny and largely smooth; mesoscutal lobes largely smooth except for punctulation, shiny and densely setose; precoxal area of mesopleuron largely smooth medially, with only some superficial rugulae; remainder of mesopleuron finely punctate and antero-dorsally rugose; metapleuron remotely punctate and largely smooth medially; scutellum remotely punctulate; metanotum with fine complete median carina; propodeum weakly convex and densely rugose, its medio-longitudinal carina complete and fine. Wings. Fore wing: r 0.5 × 3-SR (Fig. 310); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 310); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR somewhat widened; surroundings of M+CU1, 1-M and 1-CU1 evenly setose but setae pale and easily overlooked. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 310); 2-SC+R short and longitudinal; m-cu vaguely indicated; M+CU:1-M = 10:7; 1r-m 0.8 × 1-M. Legs. Tarsal claws with rather inconspicuous and pale brownish pecten remaining far removed from apical tooth (Fig. 321); hind coxa largely superficially finely punctate, but dorso-anteriorly densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.4 and 6.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flat, 1.1 × as long as wide apically; 1st and 2nd tergites with fine medio-longitudinal carina and finely longitudinally (1st) or irregularly (2nd) densely rugose; mediobasal area of 2nd tergite triangular and medium-sized (Fig. 313); 2nd suture rather deep, finely crenulate and narrow; basal half of 3rd tergite finely rugulose, remainder of metasoma superficially micro-sculptured or nearly smooth; 4th and apical half of 3rd tergite without sharp lateral crease;

ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 308). Colour. Yellowish brown; antenna (except dark brown scapus and pedicellus), stemmaticum and ovipositor sheath black; tarsi, medio-posterior patch of propodeum, basal patch of 1st tergite and apex of hind tibia dark brown; veins rather dark brown at medial third of fore wing, remainder of veins pale brown or yellowish; pterostigma brownish yellow; wing membrane subhyaline. Variation. Scapus entirely dark brown or largely yellowish brown; dark patches of propodeum and 1st tergite sometimes absent (♀ RMNH from Ningxia). Antennal segments: ♀ 56(2), 58(2), 59(1); ♂ 57(1), 59(1). Male is very similar with apical tergites type ?1–2, setae short, sparse and hard to see, with fringe very short and negligible. China (Ningxia), Mongolia. This Asian species is included here because it was reported from Poland (Huflejt, 1997). The record needs confirmation to rule out confusion with a similar European species. 10.3897/zookeys.919.39642.figures307-309675D22D4-B1D5-5BD6-99A3-5BF8C3947159Figures 307–309. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 307 habitus lateral 308 ovipositor sheath lateral 309 detail of fore wing. https://binary.pensoft.net/fig/39009110.3897/zookeys.919.39642.figures310-3210B4B9462-8FF15305-969A-53AB1B89A7CBFigures 310–321. Aleiodes fahringeri (Telenga), ♀, Mongolia, Somon Bulgan 310 wings 311 mesosoma lateral 312 mesosoma dorsal 313 propodeum and 1st–3rd metasomal tergites dorsal 314 fore femur lateral 315 hind femur lateral 316 base of antenna 317 apex of antenna 318 head anterior 319 head dorsal 320 head lateral 321 inner hind tarsal claw https://binary.pensoft.net/fig/390092ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1977b) Braconidae (Hymenoptera) from Mongolia, VI.Folia Entomologica Hungarica30(1): 109–118.ChenX-XHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.PappJ (2009) Braconidae (Hymenoptera) from Mongolia, XVII. Eleven subfamilies.Acta Zoologica Academiae Scientiarum Hungaricae55(2): 139– 173.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.

18. XML treatment for Aleiodes fortipes. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642543DE8ED-DC0F-5A62-8241D34B299DFD30 Aleiodes fortipes (Reinhard, 1863)Figs 322–324, 325–338, 339–342 Rogas fortipesReinhard, 1863: 272; Shenefelt 1975: 1229 [examined]. Aleiodes (Neorhogas) fortipes; Papp 1985a: 158, 1987a: 333, 1987b: 35, 1991a: 75. Aleiodes fortipes; Papp 2005: 176; Lozan et al. 2010: 17; Butcher et al. 2012: 14. Rhogas freyiHellén, 1927: 25–26; Papp 1985a: 158 (unnecessary lectotype designation and as synonym of A. fortipes), 2005: 176 [examined]. Rogas freyi; Shenefelt 1975: 1229–1230; Tobias 1986: 75 (transl.: 121). Holotype of A. fortipes, ♂ (ZMB), “Gallia [France]”, “Type”, “Coll. H. Rhd.”, “26723”, “fortipes Rhd.”, “Holotypus”, “Rogas fortipes Reinh., 1863, ♂, Papp, 1983.”. Holotype of A. freyi, ♂ (ZMH), “[Finland], Nagu”, “R. Frey”, “Freyi n. sp., Hellén det.”, “Mus. Zool. H:fors, sp. typ. No. 5363, Rhogas Freyi Hellén”, “Lectotypus Rogas freyi Hellén, design. Tobias”, “Aleiodes % ♂ fortipes Rh., det. Papp J., 1983/ compared with ♂ holotype of A. fortipes”. The lectotype designation is superfluous because it is evident from the description that the author had only one male. Austria, British Isles (England: V.C.s 16, 26, 28), Bulgaria, Czech Republic, Finland, France, Germany, Hungary, Netherlands (GE: ‘t Harde, Nunspeet), Poland, Spain, Sweden, Turkey. Specimens in ZJUH, BZL, CMIM, FC, MTMA, NMS, RMNH, SDEI. It has been collected in open or understory habitats, including (but not exclusively) growths dominated by Vaccinium and/or Calluna below sparse conifers. Generally, found on sandy well-drained soils in England (Breck heaths of East Anglia) and the Netherlands (Veluwe). MRS650 (France), MRS807 (Poland). The flight time of this univoltine species is (April)May–June, and ca 10 months of the year is spent as an exposed mummy. The only mummy seen (Fig. 324) formed in captivity firmly attached beneath a thin stem and would have been positioned low down in the vegetation, but probably aerially. It is light brown, moderately slender, and the parasitoid occupied approximately abdominal segments 3–8. The host was Idaea sp. (Geometridae), either I. aversata (Linnaeus) or I. straminata (Borkhausen), and the rearing arose when a few larvae of the foregoing were collected (MRS) along with an adult female of A. fortipes at the same site in Poland (22.v.2016) and offered to the parasitoid, which had been fed honey water, on 24.v.2016. Although two of the caterpillars were well-grown, in their final instars and at least twice as long as the parasitoid, one was accepted avidly. This host was first pricked several times, at intervals. Paralysis was rather slow to take effect and not complete until after the host was revisited for oviposition: a single insertion of ca 30 seconds duration, with no post-oviposition association (the parasitoid simply walked away after oviposition). The host mummified on 9.vi.2016 and an adult female emerged on 22.v.2017. The other host was rejected after being pricked just once, and later died. A penultimate instar caterpillar of the same host aggregate was also parasitised but died after an ecdysis. Subsequent barcoding (through the kindness of Axel Hausmann, ZSSM) of the dead caterpillars revealed one specimen each of I. aversata and I. straminata, leaving the precise determination of the successful host unclear. It is possible that the parasitised host had already been attacked before it was collected, but the rather long time before mummification occurred suggests not. In any case, at least one Idaea species in the aversata/ straminata group clearly serves as host. Some individuals of the long and slender, morphologically very different, larvae of Idaea muricata (Hufnagel) were also offered. Although possibly of less interest to the parasitoid, one penultimate instar larva (1.7 times the length of the female parasitoid) was immediately parasitised (a single prick for eventual paralysis, followed after an interval by a single insertion for oviposition lasting just more than a minute), but this larva later produced a moth. Final instars of this very elongate species of caterpillar were generally ignored, but one did elicit a downwards curl of the metasoma without, however, being stung. There are two particularly significant aspects to the successful rearing. The first is that these Idaea species overwinter as quite well-grown larvae, so during the flight period of the parasitoid they are in late instars, and attacking hosts at this stage is an unusual strategy for Aleiodes (but see A. aterrimus and A. sibiricus). The second is that we know of no other Aleiodes species apart from A. sibiricus (q. v.) among those whose host overwinters as a larva that fails to take advantage of that to overwinter as an early instar larva within it. The apparently riskier strategy taken by A. fortipes, in both respects, may be plesiomorphic. Aleiodes fortipes is the only known West Palaearctic species in which males have small, subapical setose pore (probably associated with pheromone release) situated mid-dorsally on each of the 4th– 6th metasomal tergites (Fig. 340). We also expect these pores to be present in A. caucasicus, which is only doubtfully distinct from A. fortipes, but we have not seen the male of A. caucasicus. Similar, probably homologous, pores are also a feature of males of Aleiodes (Hemigyroneuron) species which are found in the near East, Oriental and Afrotropical regions (Butcher & Quicke, 2015). Outside of Hemigyroneuron, metasomal pores are also found the New World Aleiodes cameronii (Dalla Torre) species complex and in a number of undescribed Madagascan Aleiodes. In Hemigyroneuron the pores have been shown to connect with large sub-tergal glands (Butcher & Quicke, 2011). Collectively these taxa form a basal clade in our molecular phylogeny (Fig. 1). Aleiodes fortipes is the only species among those treated in this part of our revision with known hosts outside the Noctuidae and, although no host is known for rather a lot of these species, the apparently basal position of A. fortipes in the group is noteworthy and using geometrid hosts may also be plesiomorphic. The rather slender ovipositor sheath (Fig. 322) is another indication for its basal position. It is interesting that the known hosts of both A. fortipes and of the subgenus Hemigyroneuron are all Geometridae (two species of Hemigyroneuron with examined mummies, India and S. Africa, cited by Butcher & Quicke, 2011 [a label record indicating a pierid host of a 3rd species is also cited in that paper but is discounted here because no mummy was present]). An Australian species described under Hemigyroneuron with examined mummy reported to be that of a geometrid by Butcher & Quicke (2016) is probably (a) actually not a member of A. (Hemigyroneuron) and (b) may be from a lasiocampid (W. Moore in litt.). The hosts of members of the Aleiodes cameronii complex, based on multiple rearings in both North America and Costa Rica include both Geometridae and Erebidae (Eiseman & Charney, 2010; http://v4.boldsystems.org). Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 332); 2nd– 10th antennal segments yellowish, contrasting with remaining darker segments; clypeus obtuse apically and not protruding in lateral view (Fig. 334); precoxal area largely smooth, at most with some aciculae or punctures medially (Fig. 327); tegulae brown; lobes of mesoscutum finely

coriaceous-granulate and rather dull, with satin sheen; vein 1-CU1 of fore wing 0.4–0.6 × vein 2CU1 (Fig. 332); length of hind femur 3.5–3.8 × its maximum width (Fig. 331); hind tarsal claws small and only yellowish or brownish setose (Fig. 338); body of ♂ completely black, antenna completely blackish, dark brown or with some segments yellowish subbasally and 4th–6th tergites with a setose medio-dorsal depression; length of fore wing 3.7–5.0 mm. Holotype of A. freyi, ♂, length of fore wing 4.5 mm, of body 5.3 mm. Head. Antenna incomplete, (length of antenna of ♀ from Santon Downham 1.2 × fore wing, its subapical segments rather robust: Fig. 336); frons smooth anteriorly and with coarse curved striae posteriorly; OOL 2.7 × diameter of posterior ocellus, and rather regularly and rather coarsely striate; vertex transversely striate, rather shiny; clypeus narrow, strongly curved dorsal margin, rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 334); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 332); length of eye 1.4 × temple in dorsal view (Fig. 333); vertex behind stemmaticum transversely rugose; clypeus near lower level of eyes; face coarsely transversely rugose; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes very densely coriaceous-granulate, with vague micro-reticulate sculpture, matt; precoxal area of mesopleuron largely smooth (except some micro-sculpture) medially, rather depressed; remainder of mesopleuron largely smooth, except some punctures and antero-dorsally coarsely rugose; scutellum superficially granulate and with some punctures; propodeum coarsely rugose-reticulate and medio-longitudinal carina nearly complete. Wings. Fore wing: r 0.6 × 3-SR (Fig. 325); 1-CU1 horizontal and somewhat widened, 0.45 × 2-CU1; r-m 0.65 × 3-SR; 2nd submarginal cell rather short (Fig. 325); cu-a vertical, straight; 1-M rather curved posteriorly; 1-SR short and narrow; surroundings of M+CU1, 1-M and 1-CU1 evenly setose; m-cu subvertical, slightly diverging from 1-M posteriorly. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 326); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 80:57; 1r-m 0.6 × 1-M; 1-M straight. Legs. Tarsal claws small but robust and only yellowish setose (Fig. 338); hind coxa largely rugulosegranulate; hind trochantellus medium-sized; length of hind femur and basitarsus 3.8 and 7.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus; fore femur 4.8 × as long as wide. Metasoma. First tergite evenly convex, 1.3 × as long as wide apically; 1st and 2nd tergites with weak medio-longitudinal carina and together with basal half of 3rd tergite densely and finely longitudinally rugose; medio-basal area of 2nd tergite narrow but rather distinct (Fig. 329); 2nd suture rather shallow; remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; 4th–6th tergites with a setose medio-dorsal depression. Colour. Dark brown or blackish; palpi dark brown; mesopleuron with reddish brown streak; legs yellowish brown but tarsi, apex of hind femur (and indistinctly apices of fore and middle femora, and of tibiae) and base of hind coxa infuscate; tegulae and pterostigma brown; wing membrane slightly infuscate. Variation. Maximum width of marginal cell of hind wing 2.0–2.6 × its width near hamuli (Fig. 326); vein 1-CU1 of fore wing 0.4–0.6 × vein 2-CU1; length of hind femur 3.5–3.8 × its maximum width; length of 1st tergite 1.0 (♀)–1.3 (♂) × its apical width. Propodeum and metapleuron posteriorly, 1st and 2nd tergites and base of 3rd tergite of ♀ more or less brown, basal third of antenna (except scapus and base of pedicellus) brownish yellow or yellowish brown and ovipositor sheath rather slender, with long setae and apically narrowed (Fig. 323). Antennal segments: ♀ 38(1), 39(1), 41(1), 43(1), 44(1), 45(1); ♂ 36(1), 39(3), 40(6), 41(3), 42(7), 43(2), 44(2), 45(1). The number of antennal segments appear to be comparable between the sexes. Males have 1st–3rd metasomal tergites completely black, and basal third of antenna completely blackish, dark brown or with some subbasal segments yellowish. The male apical tergites (besides pores, see above) are type 1, setae rather dense, no fringe observed and probably absent (Fig. 341). *Austria, *British Isles (England), Bulgaria, Czech Republic, Finland, France, *Germany, Hungary, *Netherlands, *Poland, *Spain, *Sweden, *Turkey. 10.3897/zookeys.919.39642.figures322-324D4F8A347-2AA6-5BC0-8AE2D3D499D8D54FFigures 322–324. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 322 habitus lateral 323 ovipositor sheath lateral 324 mummy of Idaea sp. (either I. aversata (Linnaeus) or I. straminata (Borkhausen)), Poland, Dybki. https://binary.pensoft.net/fig/39009310.3897/zookeys.919.39642.figures325-338B8113AE8-13F95169-9489-132776F01013Figures 325–338. Aleiodes fortipes (Reinhard), ♀, England, Santon Downham 325 fore wing 326 hind wing 327 mesosoma lateral 328 mesosoma dorsal 329 1st –3rd metasomal tergites dorsal 330 fore femur lateral 331 hind femur lateral 332 head anterior 333 head dorsal 334 head lateral 335 base of antenna 336 apex of antenna 337 antenna 338 inner hind tarsal claw. https://binary.pensoft.net/fig/39009410.3897/zookeys.919.39642.figures339-342D9FBADDA-4F615BDC-80C2-8EC0C691093DFigures 339–342. Aleiodes fortipes (Reinhard), ♂, England, Santon Downham 339 habitus lateral 340 3rd–7th metasomal tergites dorsal 341 id. lateral 342 metasoma dorsal. Arrows indicating setose depressions or pores of 4th–6th tergites. https://binary.pensoft.net/fig/39009510.3897/zookeys.919.39642.figures1-6A5EF9B58-048B-5FD2A778-8CC686DC3F21Figures 1–6. Terminology and measurements used in this paper 1 wing venation: pa = parastigma, pt = pterostigma, 1 = marginal cell, 2a, b, c = 1st, 2nd and 3rd submarginal cell, respectively, 3a, b = 1st and 2nd discal cell, respectively, 4a = 1st subdiscal cell, 5 = basal cell, 6 = subbasal cell 2 head, dorsal aspect: a = length of eye, b = length of temple 3 head, lateral aspect: c = width of temple, d = width of eye, e = height of eye, f = width of malar space (measured as actual true distance in its own plane) 4 head, anterior aspect: g = width of face, h = width of hypoclypeal depression 5 fore femur, lateral aspect: i = length, j = width 6 1st metasomal tergite, dorsal aspect: k = length of tergite (measured from adductor), l = apical width of tergite. https://binary.pensoft.net/fig/390045ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1987a) First survey of the braconid fauna of the Kiskunság National Park, Hungary (Hymenoptera, Braconidae). In: MahunkaS (Ed.) The Fauna of the Kiskunság National Park II.Akademiai Kiado, Budapest, 314–334.PappJ (1987b) First outline of the braconid fauna of Southern Transdanubia, Hungary (Hymenoptera, Braconidae), 3. Helconinae, Homolobinae,

Macrocentrinae, Blacinae, Doryctinae and Rogadinae. A Janus Pannonius Muzeum Evkonyve 30–31 (1985–1986): 33–36.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.LozanABelokobylskijSvan AchterbergCMonaghanM (2010) Diversity and distribution of Braconidae, a family of parasitoid wasps in the Central European peatbogs of South Bohemia, Czech Republic Journal of Insect Science 10: 16(1–21).

https://doi.org/10.1673/031.010.1601ButcherBASmithMASharkeyMJQuickeDLJ (2012) A turbotaxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae: Rogadinae) based largely on COI barcoded specimens, with rapid descriptions of 179 new species.Zootaxa3457: 1–232.TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]

19. XML treatment for Aleiodes gasterator. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396420E8476B0-62DA-5838-80F9B17814F7587A Aleiodes gasterator (Jurine, 1807)Figs 343–346, 347–359, 360–364 Bracon gasterator Jurine, 1807: 118, pl. 8. [examined]. Rogas gasterator; Shenefelt 1975: 1230–1231; Zaykov 1980a: 112. Rogas (Rogas) gasterator; Tobias 1976: 86, 1986: 81 (transl.: 133) p.p. Aleiodes (Neorhogas) gasterator; Papp 1991a: 91 p.p. Aleiodes (Chelonorhogas) gasterator; Falco et al. 1997: 60; Ghahari et al. 2011: 267; Rastegar et al. 2012: 3; Farahani et al. 2015: 243. Aleiodes gasterator; Bergamasco et al. 1995: 5; Zaldívar-Riverón et al. 2004: 234; Papp 2005: 176. Bracon dimidiatusSpinola 1808: 123–124. Syn. nov. Aleiodes dimidiatus; Bergamasco et al. 1995: 5. Rogas (Rogas) dimidiatus: Tobias 1976: 86; 1986: 81 (transl.: 134) p.p. Rogas dimidiatus; Zaykov 1980a: 112. Aleiodes (Neorhogas) dimidiatus; Papp 1991a: 90 p.p. Aleiodes (Chelonorhogas) dimidiatus; Samartsev and Belokobylskij 2013: 765; Farahani et al. 2015: 242. Rhogas (Rhogas) dimidiatus var. turkestanicusTelenga, 1941: 184, 409; Shenefelt 1975: 1225 [examined]. Syn. nov. Holotype of A. gasterator, ♀ (Museum Genève), “[? Switzerland], gasterator J.”, “Typus”, “Bracon gasterator Jur., Type”, “Type du g. Rogas [= incorrect]”, “vu par [R.D.] Shenefelt, U.S.A., 1967” (metasoma on separate card and pin). Lectotype of A. turkestanicus here designated, ♀ (ZISP), “[Turkmenistan], Transcaspia, Bajram-aly, 17.viii.1930, T. Boguj/311, i.s.”, “Rhogas dimidiatus Spin. var. turkestanica [sic!] nov., N. Telenga det.”. Albania, Cyprus, France (including Corsica), Greece (including Crete), Italy (including Sardinia, Sicily), North Macedonia, Portugal (including Madeira), Spain (including Mallorca, Menorca, Tenerife), Tunisia, Turkey, [Iraq, Jordan, Syria]. Specimens in ZJUH, BZL, CNC, MSC, MTMA, NMS, RMNH, ZSSM. Widespread in the Mediterranean region, where it tends to replace A. ruficornis. MRS046 (France), MRS048 (France), MRS892 (Spain). Collected chiefly in May–July and September–November, but specimens have occurred in every month of the year. Plurivoltine; there is no indication of a unique overwintering mode in the material seen. Reared from low-feeding Noctuidae: Agrotis segetum (Denis & Schiffermüller) (6 [6 ZJUH], Spain), Agrotis sp. (1 [ZJUH], Cyprus; W.R. Ingram), mixed Agrotis and Spodoptera littoralis (Boisduval) (1 [ZJUH], Cyprus; W.R. Ingram). The two mummies seen are rather different (Figs 345, 346), though it may be that neither overwintered; a small, pale and relatively slender one from Agrotis sp. produced a small male, while a more normal sized individual emerged from the larger, dark and stout mummy whose host was (even) less certain. Even in the latter case, the mummy is less keeled, less lined with silk and much more in relation to the size of the emerging adult than is the situation with the overwintering mummies of the A. grassator/ carbonarius/ carbonaroides/ ruficornis complex. The appearance of both mummies suggest that they would normally form below ground. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 355); OOL of ♀ 1.2–1.6 × as long as diameter of posterior ocellus (Fig. 356) and distinctly rugose or rugulose; antennal segments of ♀ 29–39, of ♂ 36–46(–51) (usually 39–43); antenna of ♀ 0.8–0.9 × fore wing; length of malar space of ♀ 0.5–0.6 × height of eye in lateral view (Fig. 357); clypeus thick apically and not protruding anteriorly (Fig. 357); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; posterior half of notauli deep; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 347); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 347); hind tarsal claws yellowish or brownish bristly setose; third tergite with (faint) curved or antero-medially transverse rugulae or striae (Fig. 364) or largely smooth (sometimes with only longitudinal striae baso-laterally), often with distinct punctures laterally; hind femur at least apico-dorsally dark brown or black; inner and/or dorsal side of hind tibia (largely) yellowish or red; tegulae usually (partly) dark brown; pale males have nearly always frons medially and stemmaticum black; palpi usually brownish or yellowish, sometimes dark brown; 3rd metasomal tergite frequently partly or completely reddish or yellowish; 4th and 5th tergites black. Redescribed ♀ (RMNH) from France (Isle sur le Sorque). Length of fore wing 4.9 mm, of body 6.1 mm. Head. Antennal segments of ♀ 35, length of antenna 0.9 × fore wing, its subapical segments robust (Fig. 359), 4th segment 1.2 × longer than wide; frons with coarse curved rugae; OOL 1.6 × diameter of posterior ocellus, and densely rugose or rugulose; vertex densely rugose, shiny; clypeus densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 357); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 355); length of eye 1.3 × temple in dorsal view (Fig. 356); vertex behind stemmaticum punctate-rugose; clypeus just below lower level of eyes; length of malar space 0.5 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, interspaces largely smooth with superficial granulation, shiny; precoxal area of mesopleuron evenly vermiculate-rugose medially, but only sparsely punctate posteriorly; metanotum without median carina; scutellum rather flat, sparsely punctate, but rugose laterally; propodeum coarsely vermiculate-rugose, medio-longitudinal carina nearly complete, and angulate latero-posteriorly. Wings. Fore wing: r 0.35 × 3-SR (Fig. 347); marginal cell distinctly ending basad of level of apex of vein 3-M; 1-CU1 horizontal, slender, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell mediumsized (Fig. 347); cu-a inclivous, posteriorly curved; 1-M rather curved posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened, but slightly basally, its apical width 2.3 × width at level of hamuli (Fig. 348); 2-SC+R subquadrate; m-cu largely absent; M+CU:1-M = 28:19; 1r-m 0.6 × 1-M. Legs. Tarsal claws robust and with only bristly brownish setae (Fig. 358); hind coxa largely densely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.2 and 5.8 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex medially and 0.9 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose; medio-basal area of 2nd tergite short and rather distinct (Fig. 351); 2nd suture deep; subbasally 3rd tergite with faint curved striae and medially transverse (Fig. 364); remainder of metasoma superficially micro-sculptured; 4th and apical

half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically oblique, dorsally longer than ventrally (Fig. 344). Colour. Black; face (except medio-dorsally), malar space, dorsal half of temple, frons largely laterally, notauli, mesoscutum laterally, scutellum, pronotum postero-dorsally, mesopleuron dorsally and posteriorly, metapleuron largely, 1st and 2nd metasomal tergites and base of 3rd tergite orange brown; palpi and humeral plate and veins of hind wing yellowish brown; tegula rather dark brownish; ventral half of temple largely, dorso-apical patch of hind femur, pterostigma and veins of fore wing dark brown; fore wing membrane slightly infuscate, of hind wing subhyaline. Variation. A very colour-variable species; head and mesoscutum of female may be largely black (nominate form) or reddish (= “A. dimidiatus / var. turkestanicus”, but especially the mesoscutum may be intermediate). Especially males may have the hind coxa black and most of hind tibia dark brown, sometimes the entire leg is nearly completely black or dark brown. Antennal segments: ♀ 29(1), 31(2), 32(3), 33(13), 34(13), 35(15), 36(9), 37(9), 38(3), 39(2); ♂ 36(2), 37(3), 38(4), 39(8), 40(13), 41(9), 42(5), 43(11), 44(7), 45(2), 46(2). Additionally, an exceptionally large male with 50 segmented antennae from Cyprus (ZJUH) appears to belong to this species, as does a female from Spain (RMNH) with 41 antennal segments, basal half of 3rd tergite largely obliquely rugose and blackish scapus. On average males have ca 7 more antennal segments than females. Males are very similar with apical tergites type 1–2, setae rather sparse and with evident but short fringe (Fig. 360), hind tibial spurs often blunt apically and 3rd tergite remotely punctate basally. *Albania, *Cyprus, France (including Corsica), Greece (including Crete), *Iraq, Italy (including Sardinia, Sicily), *Jordan, *North Macedonia, *Portugal (including Madeira), Spain (including Mallorca, Menorca and Tenerife), *Syria, *Tunisia, *Turkmenistan, Turkey. The new synonymy of Rhogas dimidiatus var. turkestanicus Telenga, 1941, is based on direct comparison of the types of both taxa. The identity of Bracon dimidiatus Spinola, 1808, is problematic because the holotype from Italy (Genoa) is lost and the original description is far too incomplete for an easy identification. Its colour pattern (head completely yellowish, hind tibia and 3rd tergite reddish) does not fit with A. ruficornis (Herrich-Schäffer); if the head is largely reddish brown then the temple ventrally and malar space remain blackish. This pattern agrees better with that of pale specimens of A. gasterator (named as A. dimidiatus var. turkestanicus Telenga, 1941). Aleiodes ruficornis occurs also in Italy, but its females have the head partly black ventrally, the apex of the hind tibia dark brown and most of the 3rd metasomal tergite black. Therefore, we synonymise Bracon dimidiatus with A. gasterator (syn. nov.). The holotype of Bracon gasterator Jurine, 1807, has the 3rd metasomal tergite finely curved (nearly circular) aciculate or striate basally, palpi (as far as present) pale brownish, maximum with of hypoclypeal depression 0.45 × minimum width of face, vein 1-CU1 of fore wing half as long as vein 2-CU1 and 4th antennal segment 1.3 × as long as wide. Aleiodes arnoldiisensuFarahani et al. (2015) concerns a species closely related to A. gasterator (Jurine) having basal half of 3rd tergite coarsely longitudinally rugose, antenna of ♀ with 30–35 segments (of ♂ 36), head linearly narrowed ventrally in anterior view and subbasal antennal segments of ♀ slightly slenderer. 10.3897/zookeys.919.39642.figures343-34612E3CF58-F8FC-5280-A93F-7BBCDC981CDCFigures 343–346. Aleiodes gasterator (Jurine), ♀, France, Les Constants, but 345 and 346 from Cyprus 343 habitus lateral 344 ovipositor sheath lateral 345 mummy of Agrotis sp. 346 mummy of Agrotis sp. or Spodoptera littoralis (Boisduval). https://binary.pensoft.net/fig/39009610.3897/zookeys.919.39642.figures347-359A0670B61-D775580C-AFC1-0F049202709CFigures 347–359. Aleiodes gasterator (Jurine), ♀, France, Les Constants 347 fore wing 348 hind wing 349 mesosoma lateral 350 mesosoma dorsal 351 metasoma dorsal 352 fore femur lateral 353 hind femur lateral 354 base of antenna 355 head anterior 356 head dorsal 357 head lateral 358 outer hind tarsal claw 359 apex of antenna. https://binary.pensoft.net/fig/39009710.3897/zookeys.919.39642.figures360-3645EE78E7B-70FF5FBF-99FD-10AD78AE6098Figures 360–364. Aleiodes gasterator (Jurine), ♂, Italy, Livorno, but 364 of ♀, France 360 habitus lateral 361 antenna 362 base of antenna 363 apex of antenna 364 3rd metasomal tergite dorsal. https://binary.pensoft.net/fig/390098ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980a) Unknown genera and species of Braconidae for the fauna of Bulgaria. Travaux scientifiques d’Université de Plovdiv “Paissi Hilendarski”, Biologie 15(4) [1977]: 111–115. [in Bulgarian]TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.GhahariHFischerMPappJ (2011) A study on the Braconidae (Hymenoptera: Ichneumonoidea) from Ilam province, Iran.Calodema160: 1– 5.RastegarJSakeninHKhodaparastSHavaskaryM (2012) On a collection of Braconidae (Hymenoptera) from East Azarbaijan and vicinity, Iran.Calodema226: 1–4.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.ZaldívarRiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xPappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.SpinolaM (1808) Insectorum Liguriae species novae aut rariores, quas in agro Ligustico nuper detexit, descripsit, et iconibus illustravit (Hymenoptera) 2: 1–262. Genua.SamartsevKGBelokobylskijSA (2013) On the fauna of the true cyclostome braconid wasps (Hymenoptera, Braconidae) of Astrakhan Province. Entomologicheskoe Obozrenie 92(2): 319– 341.Entomological Review93(6): 755–774.

20. XML treatment for Aleiodes grassator. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642B81F3B1B-63AE-54A5-B6FBEFC49244C7EC Aleiodes grassator (Thunberg, 1822)Figs 365–367, 368–380, 381–384 Ichneumon grassatorThunberg, 1822: 256 [examined]. Rogas grassator; Shenefelt 1975: 1232. Aleiodes (Neorhogas) grassator; Papp 1991a: 86. Aleiodes grassator; Zaldívar-Riverón et al. 2004: 234; Quicke et al. 2014: 240; Butcher et al. 2014: 458. Rhogas grassator ab. thoracicusHellén, 1927: 24; Shenefelt 1975: 1232 (unavailable name). Rogas (Rogas) flavipalpisThomson, 1892: 1672 [examined]. Aleiodes flavipalpis; Papp 1991a: 86 (as synonym of A. grassator). Rogas alpinusThomson, 1892: 1671; Shenefelt 1975: 1217 [examined]. Syn. nov. Aleiodes alpinus; Papp 1991a: 90 (as synonym of A. dimidiatus). Holotype of A. grassator, ♀ (ZMUU), unlabelled. Lectotype of A. flavipalpis, ♀ (ZIL), “åre”, “Sverige, Ǻreskutan I Jemtland/teste Papp, 1983”, “Lectotypus Rogas flavipalpis Thomson, 1899 [sic!], Papp, 1983”, “Aleiodes grassator Thb., det. Papp J., 1983”. Lectotype of A. alpinus, ♀ (ZIL), “[Norway:] Dovre”, “alpinus m.”, “Lectotypus Rogas alpinus Thoms., 1891, ♀. Papp, 1983”, “Aleiodes dimidiatus var. alpinus Th., ♀, det. Papp J., 1983”. Austria, British Isles (England: V.C. 70; Scotland: V.C.s 83, 85, 88, 89, 97, 103; Finland, France (both Alps and Pyrenees), Italy, Germany, Norway, Romania, Sweden, Switzerland. Specimens in ZJUH, BZL, MRC, MTMA, NMS, RMNH, SDEI, ZIL, ZMUU, ZSSM. This is essentially a montane grassland species, though occurring at low altitudes in northern Europe. MRS215 (UK), MRS721 (UK), MRS725 (UK). Collected in (April)May–July. Univoltine, overwintering in the mummy. Reared from the noctuid Cerapteryx graminis (Linnaeus) (9: K.P. Bland, M.J.W. Cock, M.R. Shaw) and from mummies compatible with that (3), and it may be strictly monophagous. The known host overwinters in the egg stage, and feeds on Poaceae near ground level. The tough dark brown mummy is formed on or

below the soil surface and seems spectacularly too large for the adult that will emerge from it (Fig. 384). It is more or less cylindrical, though with a pronounced lateral keel, and well-lined with silk. The cocoon chamber occupies most of the abdominal segments. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 375); OOL of ♀ ca twice as long as diameter of posterior ocellus (Fig. 376) and distinctly rugose or rugulose; length of 4th antennal segment of ♀ 0.7–0.9 × its width (Fig. 378; in ♂ 0.9–1.0 ×); clypeus thick apically and not protruding anteriorly (Fig. 377); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 368); vein 1-CU1 of fore wing 0.5–0.6 × as long as vein 2-CU1 (Fig. 368); hind tarsal claws robust (Fig. 380) and yellowish or brownish bristly setose; hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; pale males have whole frons and stemmaticum yellowish; palpi dark brown or blackish, rarely brown; 3rd metasomal tergite only antero-laterally reddish or yellowish; 4th and 5th tergites black. Redescribed ♀ (RMNH) from Finland (Sb: Leppävirta). Length of fore wing 4.6 mm, of body 5.7 mm. Head. Antennal segments of ♀ 39, 4th segment 0.8 × longer than wide (Fig. 378); antenna as long as fore wing, its subapical segments robust (Fig. 379); frons with coarse curved rugae and shiny; OOL 1.9 × diameter of posterior ocellus and rugose; vertex rugose and shiny; face rugose-punctate; clypeus rugose; ventral margin of clypeus rather thick and not protruding forwards (Fig. 377); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 375); length of eye 1.1 × temple in dorsal view (Fig. 376); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.6 × length of eye in lateral view. Mesosoma. Mesoscutal lobes moderately punctate, laterally interspaces mainly smooth, medially superficially granulate and rather shiny; precoxal area of mesopleuron coarsely rugose medially and largely smooth posteriorly; remainder of mesopleuron mainly punctate, but dorsally coarsely rugose;

scutellum flat, sparsely finely punctate and only anteriorly with lateral carina; propodeum coarsely rugose, medio-longitudinal carina present on anterior half, rounded posteriorly and dorsal part approx. as long as posterior part. Wings. Fore wing: r 0.3 × 3-SR (Fig. 368); marginal cell ends basad of level of apex of 3-M; 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell robust (Fig. 368), 3-SR 1.3 × as long as 2-SR; cu-a vertical, straight; 1-M slightly curved posteriorly; 1-SR similar to 1-M and mediumsized; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width twice width at level of hamuli (Fig. 369); 2-SC+R subquadrate; m-cu short; M+CU:1-M = 27:18; 1r-m 0.7 × 1-M. Legs. Tarsal claws robust and with only brownish bristly setae (Fig. 380); hind coxa largely densely punctate, but dorsally with some rugae; hind trochantellus robust; length of hind femur and basitarsus 3.1 and 3.9 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus. Metasoma. First tergite rather flattened, 0.8 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose; medio-basal area of 2nd tergite wide triangular and short (Fig. 372); 2nd suture deep and crenulate; basal half of 3rd tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured; 4th tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 367). Colour. Orange brown; apical two thirds of antenna, labial palp, patch on hind femur dorso-apically, posterior patch of 2nd tergite and telotarsi, dark brown; head, mesosoma (except side of pronotum postero-dorsally and pair of latero-posterior patches of propodeum), 3rd–7th tergites (except anterolateral corners of 3rd tergite) black; maxillary palp, basal third of antenna, tegulae and remainder of legs rather pale yellowish brown; veins and pterostigma dark brown; wings distinctly infuscate but hind wing less than fore wing. Variation. Basal third or half of antenna of ♀ pale yellowish brown; head partly and mesosoma anteriorly of ♀ dark orange brown or both entirely black; 3rd tergite longitudinally striate or rugulose basally (sometimes narrowly so), without curved sculptural elements (Fig. 372), except sometimes some weak transverse striae occasionally present at extreme apex. Males are always darker than females; mainly black with legs mainly dark brown or blackish (Fig. 381). Antennal segments: ♀ 36(2), 37(4), 38(6), 39(10), 40(6), 41(1); ♂ 47(1), 48(2), 49(2), 50(3), 51(2), 52(4), 53(1), 60(1). On average males have ca 12 more antennal segments than females. Males have 2nd submarginal cell slightly shorter than of females, temple and face long setose, malar space 0.5–0.7 × length of eye in lateral view, apical tergite type 1, rarely type 2, setae rather dense, fringe not observed and probably absent (Fig. 381). The superficial granulosity of 3rd tergite and mesoscutum may be absent. *Austria, British Isles (England, Scotland), Finland, *France, *Ireland, *Italy, *Germany, Norway, *Romania, Sweden, *Switzerland.

The synonymy of Rogas alpinus Thomson, 1892, with Aleiodes grassator (Thunberg, 1822) is based on direct comparison of the types listed above. Although males of A. carbonaroides are generally easily distinguished from A. grassator through being black, it is possible that lighter forms occur which would be difficult to recognise. Also, females of A. carbonaroides are similar in colour to those of A. grassator. Therefore, specimens collected at low altitude away from northern areas that appear, on other characters, to be A. grassator might well really be A. carbonariodes. See also remarks under A. carbonarius and A. ruficornis. 10.3897/zookeys.919.39642.figures365-36763534165-56B5-5939-84DE-4D1E6D226929Figures 365–367. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 365 habitus lateral 366 antenna 367 ovipositor sheath lateral. https://binary.pensoft.net/fig/39009910.3897/zookeys.919.39642.figures368-3805A59FEEF-114E55DA-901F-68B72A70D581Figures 368–380. Aleiodes grassator (Thunberg), ♀, Scotland, Beinn Ghlas 368 fore wing 369 hind wing 370 mesosoma lateral 371 mesosoma dorsal 372 metasoma dorsal 373 fore femur lateral 374 hind femur lateral 375 head anterior 376 head dorsal 377 head lateral 378 base of antenna 379 apex of antenna 380 outer hind tarsal claw. https://binary.pensoft.net/fig/39010010.3897/zookeys.919.39642.figures381-384A320B5B1-162C5A53-B93A-7EB6DB9C648CFigures 381–384. Aleiodes grassator (Thunberg), ♂, Scotland, Isle of Coll 381 habitus lateral 382 base of antenna 383 apex of antenna 384 mummy of ?Cerapteryx graminis (Linnaeus). https://binary.pensoft.net/fig/390101ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xQuickeDLJShawMRvan AchterbergCBlandKPButcherBALyszkowskiRZhangYM (2014) A new Australian genus and five new species of Rogadinae (Hymenoptera: Braconidae), one reared as a gregarious endoparasitoid of an unidentified limacodid (Lepidoptera).Zootaxa3881(3): 237–257. https://doi.org/10.11646/zootaxa.3881.3.3ButcherBAZaldívar-RiverónAvan de KampTRoloTDSBaumbachTQuickeDLJ (2014) Extension of historical range of Betylobraconinae (Hymenoptera: Braconidae) into Palaearctic Region based on a Baltic amber fossil, and description of a new species of Mesocentrus Szepligeti from Papua New Guinea.Zootaxa3860(5): 449–463. https://doi.org/10.11646/zootaxa.3860.2

21. XML treatment for Aleiodes hemipterus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964272D68C85-7514-5203-B410C4FFFB93F0DA Aleiodes hemipterus (Marshall, 1897)Figs 385–389, 390–399, 400–408 Rhogas hemipterusMarshall, 1897: 137. Rogas hemipterus; Shenefelt 1975: 1233. Aleiodes hemipterus; Papp 1990: 90, 2003: 138 (lectotype listed). Aleiodes (Chelonorhogas) hemipterus; Belokobylskij and Kula 2012: 35–38. Lectotype here designated, ♀ (ZJUH), “Tunisie, Sicard”, “Type, H.T.”, “B.M. Type Hym. 3.c.243”, “B.M. Type Hym., Rhogas hemipterus Marshall, 1896”, “hemipterus Marsh.”, “Marshall coll. 1904120”. Paralectotype: 1 ♀ (MTMA), “Tunisie”, “hemipterus M. coll. Marshall”, “Paratypus Rhogas hemipterusMarshall 1897 sp. n. % des Papp J. 1986”, “Hym. Type No 10582 Museum Budapest”, “Aleiodes ♀ hemipterus Mshl. Det. Papp J. 1991”. 1 ♀ + 1 ♂ (MNHN), “[N. Tunisia:] Teboursouk”, “Rhogas hemipterus Mrsh.”, “Muséum Paris, Coll. J. de Gaulle, 1919” [figured specimens]; 1 ♂ [but metasoma missing] (ZJUH) “Rabat, Maroc [= Morocco], coll. Thery”. None. Biology. Unknown. The specimens seen do not have phenological data, and we have not seen reared material. As the female is brachypterous it is likely that the host will be found near the ground. Maximum width of hypoclypeal depression approx. 0.6 × minimum width of face (Fig. 396); OOL of ♀ 1.2–1.3 × as long as diameter of posterior ocellus, of ♂ 0.9 × (Figs 397, 408) and finely reticulate-rugose; stemmaticum protruding dorsally; antennal segments of ♀ 46–50 (of ♂ unknown), penultimate segments approx. as long as wide and antenna 0.8 × as long as body; length of malar space 0.3–0.4 × height of eye; mesoscutal lobes densely rugose or rugulose and rather matt, middle lobe with medio-longitudinal ridge or carina, of ♀ surrounded by shallow grooves (Fig. 392); propodeum angulate posteriorly (Fig. 392); ♀ brachypterous and ♂ macropterous; marginal cell of hind wing of ♀ hardly widened (Fig. 390) and of ♂ distinctly widened apically (Fig. 403); hind tarsal claws with rather conspicuous pale brown pecten (Fig. 399); 1st–3rd metasomal tergites very densely and finely longitudinally rugose; 1st tergite of ♀ 1.0–1.1 × its apical width, of ♂ approx. 1.4 ×; 2nd metasomal suture of ♀ hardly impressed but densely costate, of ♂ medium-sized. Lectotype, ♀, length of hind wing 1.7 mm (fore wing missing, but in other specimens ca one-third longer than hind wing and 2.2 mm, brachypterous), of body 7.8 mm. Head. Antenna incomplete, segments robust; frons largely striate-rugose (but transversely costate in figured ♀); OOL 1.2 × diameter of posterior ocellus, (as vertex) rather finely and densely reticulaterugose and rather dull; clypeus rugose; ventral margin of clypeus rather thick ventrally and rather forward protruding (Fig. 398); width of hypoclypeal depression 0.6 × minimum width of face and long (Fig. 396); length of eye 1.2 × temple in dorsal view (Fig. 397); vertex behind stemmaticum rather coarsely reticulate-rugose; clypeus near lower level of eyes; length of malar space 0.35 × length of eye in lateral view. Mesosoma. Antescutal depression distinct; mesoscutal lobes coarsely rugose-punctate (but superficial in figured ♀) and rather matt, middle lobe of pair of submedian grooves (Fig. 392); nearly entire mesopleuron (except minute smooth speculum) densely and coarsely reticulate-rugose; scutellum coarsely rugose and without lateral carinae; propodeum coarsely vermiculate-rugose, dorsal face long and rectangularly angulate postero-laterally (Fig. 392). Wings. Fore wing brachypterous, hardly surpassing propodeum (Marshall, 1897): (of ♀ from Tunisia r 0.2 × 3-SR; 1-CU1 distinctly widened and oblique, 0.4 × 2-CU1; r-m 0.8 × 3-SR; 2nd submarginal cell medium-sized (Fig. 390); cu-a short, vertical, straight; 1-M straight posteriorly; 1SR widened; 1st subdiscal cell open apically and posteriorly; surroundings of M+CU1, 1-M and 1-

CU1 setose; 2m-cu present as curved and only pigmented vein). Hind wing brachypterous: marginal cell reduced, sinuate and apically narrowed (Fig. 390); 2-SC+R quadrate and widened; m-cu absent; M+CU:1-M = 27:13; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous pale brown pecten, remaining far from apical tooth and much shorter (Fig. 399); hind coxa finely and densely reticulate-rugose; hind trochantellus robust; length of hind femur and basitarsus 3.6 and 6.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st–3rd tergites regularly finely and very densely longitudinally rugose, rather matt and medio-longitudinal carina rather weak; mediobasal area of 2nd tergite triangular and short (Fig. 393); 2nd suture shallow; 4th and subsequent tergites superficially punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 389). Colour. Brown; stemmaticum and ovipositor sheath black; frons, vertex medially, occiput, femora, propodeum, 1st and 2nd tergites somewhat infuscate; wing membrane subhyaline. Variation. Length of body 7.8–8.8 mm. Antennal segments: ♀ 46(1), 50(1); ♂ unknown. Male is normally winged (vein 3-SR of fore wing 1.5 × vein 2-SR, vein r 0.3 × 3-SR, vein cu-a oblique, vein 1-CU1 narrow and 0.3 × vein 2-CU1) and has marginal cell of hind wing 2.2 × wider than width at level of hamuli (with vein m-cu present anteriorly, 2-SC+R quadrate and M+CU:1-M:1r-m = 40:30:26). Apical metasomal segments of ♂ type 1 and sparsely setose. Morocco, Tunisia. Marshall (1897) based his description on three females from Tunisia. Papp (2003) listed a female in ZJUH as lectotype, but this was not accepted as a designation by Belokobylskij & Kula (2012). Therefore, the redescribed female above is here designated formally as lectotype and is the same specimen intended to become lectotype by Papp (2003). 10.3897/zookeys.919.39642.figures385-3895C964117-B7AE-5CEC-94EF-2B0AA8C8FB1BFigures 385–389. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 385 habitus lateral 386 antenna 387 apex of antenna 388 base of antenna 389 ovipositor sheath lateral. https://binary.pensoft.net/fig/39010210.3897/zookeys.919.39642.figures390-3996F7E68A7-8F39543C-84C4-CAE338A06F69Figures 390–399. Aleiodes hemipterus (Marshall), ♀, Tunisia, Teboursouk 390 wings 391 mesosoma lateral 392 mesosoma dorsal 393 metasoma dorsal 394 fore femur lateral 395 hind femur lateral 396 head anterior 397 head dorsal 398 head lateral 399 outer hind tarsal claw. https://binary.pensoft.net/fig/39010310.3897/zookeys.919.39642.figures400-408D1B53275-75FE54C4-BFF9-9020217DADA3Figures 400–408. Aleiodes hemipterus (Marshall), ♂, Tunisia, Teboursouk 400 habitus lateral 401 fore femur lateral 402 fore wing 403 hind wing 404 mesosoma dorsal 405 metasoma dorsal 406 hind femur lateral 407 head anterior 408 head dorsal. https://binary.pensoft.net/fig/390104ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1990) Braconidae (Hymenoptera) from Tunisia, 3.Folia Entomologica Hungarica51: 89–96.PappJ (2003) Braconid wasps from the Cape Verde Islands (Insecta: Hymenoptera: Braconidae) 2. Doryctinae, Braconinae, Hormiinae, Rogadinae, Gnamptodontinae, Homolobinae, Opiinae, Alysiinae, Cheloninae, Adeliinae and Microgastrinae.Faunistische Abhandlungen (Dresden)24: 137–167.BelokobylskijSAKulaRR (2012) Review of the brachypterous, micropterous, and apterous Braconidae of the cyclostome lineage (Hymenoptera: Ichneumonoidea) from the Palearctic Region.Zootaxa3240: 1–62. https://doi.org/10.11646/zootaxa.3240.1.1MarshallTA (1897) Les Braconides (Supplément): 1–144. In: AndréE (Ed.) 1897–1900.Species des Hyménoptères d’Europe et d’Algérie 5 bis, 1–369.

22. XML treatment for Aleiodes hirtus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396425B1094D6-D8D0-5EC9-9D98A133001D9CD2 Aleiodes hirtus (Thomson, 1892)Figs 409–410, 411–424, 425–427 Rogas hirtusThomson, 1892: 1672; Shenefelt 1975: 1233. Aleiodes (Neorhogas) hirtus; Papp 1985a: 153, 155, 161 (lectotype designation and as synonym of A. pallidicornis), 1991a: 75 (id.). Aleiodes hirtus; Papp 2005: 177 (as synonym of A. pallidicornis). Rhogas hirtus ab. coloratusHellén, 1927: 23; Shenefelt 1975: 1233; Papp 2005: 177 (as synonym of A. pallidicornis) (unavailable name; not Rogas coloratus Motschulsky, 1863). Lectotype of A. hirtus, ♂ (ZIL), “[Norway], Norl.”, “hirtus m.”, “Funne I Norrland, teste Papp J., 1983”, “Lectotypus”, “Rogas hirtus Thms. 1891, ♂, Papp J., 1983”, “Aleiodes pallidicornis HS ♂, det. Papp J., 1983”. Austria, Belgium, British Isles (England: V.C.s 26, 29, 32, 62; Scotland: V.C. 78; Ireland: V.C. H12), Bulgaria, Czech Republic, Finland, France, Germany, Hungary, Netherlands (DR: Borger), Norway, Romania, Russia, Serbia, Slovakia, Switzerland, Ukraine, [? Mongolia]. Specimens in ZJUH, BZL, CMIM, CNC, FMNH, MRC, MSC, MTMA, NMS, RMNH, SDEI, UNS, USNM, UWIM, ZIL, ZSSM. MRS619 (UK), MRS882 (Romania), MRS883 (Romania). Unknown. Collected in June–August, presumably univoltine but the mode of overwintering is unclear. Most British sites are more or less damp and calcareous grasslands, approaching fens. We have not seen reared material, but the clypeus suggests that the mummy will form in the soil. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 419); OOL of ♀ approx. 1.3 × as long as diameter of posterior ocellus (Fig. 420; in ♂ approx.1.6 ×) and punctaterugose; ventral margin of clypeus rather thick but rather strongly protruding forwards (Fig. 421; stronger in in ♂: Fig. 426); mesoscutal lobes largely smooth, only indistinctly punctulate and shiny; precoxal area finely punctate and often with some rugulae (Fig. 413); vein 1-CU1 0.3–0.6 × vein 2CU1 of fore wing (Fig. 411); hind tarsal claws with rather conspicuous brownish pecten (Fig. 424); length of inner spur of hind tibia 0.5–0.7 × hind basitarsus; palpi dark brown; basal half of metasoma

at least partly reddish or orange and 1st tergite rather coarsely sculptured; setae of body of ♂ (but of ♀ mainly its head) conspicuous and dense (Fig. 426); hind coxa black; hind femur largely or completely reddish or brownish; basal half of hind tibia usually (pale) yellowish or yellowish brown, but sometimes uniformly reddish and of ♂ ivory. In the past this species has been frequently misidentified as “Rogas dimidiator” or “Rogas gasterator”. Redescribed ♂ (RMNH) from Germany (Graswang). Length of fore wing 6.2 mm, of body 8.0 mm. Entire body with long whitish setae. Head. Antennal segments of ♂ 60, length of antenna 1.3 × fore wing, its subapical segments somewhat longer than wide; frons medially largely smooth, laterally with some fine curved rugae; OOL 1.6 × diameter of posterior ocellus, and punctate-rugose, POL approx. half as long as diameter of ocellus; vertex spaced rugose, shiny; clypeus punctate; ventral margin of clypeus rather thick but distinctly protruding forwards (Fig. 421); width of hypoclypeal depression 0.55 × minimum width of face (Fig. 419); length of eye 1.1 × temple in dorsal view (Fig. 420), temples conspicuously setose (Figs 419, 426); vertex behind stemmaticum rugose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Pronotum rugose and anteriorly oblique, without antescutal depression; mesoscutal lobes large smooth and shiny, only indistinctly punctulate and densely setose; precoxal area of mesopleuron punctulate and medially with some superficial rugulae; remainder of mesopleuron sparsely punctate; scutellum sparsely punctate and largely smooth, posteriorly with lateral rugae; propodeum rather convex and coarsely rugose, its medio-longitudinal carina only in anterior half of propodeum. Wings. Fore wing: r 0.3 × 3-SR (Fig. 411); 1-CU1 horizontal, 0.45 × 2-CU1; r-m 0.7 × 3-SR and as long as 2-SR; 2nd submarginal cell medium-sized (Fig. 411); cu-a vertical, straight; 1-M straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell gradually widened (but less so basally: Fig. 412), its apical width 2.5 × width at level of hamuli; 2-SC+R short longitudinal; m-cu narrowly indicated; M+CU:1-M = 9:7; 1r-m 0.7 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized brownish pecten (Fig. 424); hind coxa pimply punctate and shiny; hind trochantellus robust; length of hind femur and basitarsus 3.8 and 6.0 × their width, respectively; length of inner hind spur 0.65 × hind basitarsus (0.6 × in ♀).

Metasoma. First tergite evenly convex, approx. as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and densely longitudinally rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite minute (Fig. 415); 2nd suture deep and moderately crenulate; basal half of 3rd tergite finely rugose, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; 4th – 6th tergites with long setae and flat. Colour. Black; legs (except black coxae, trochanters and trochantelli, 1st and 2nd metasomal tergites (but base of 1st tergite partly infuscate) and base of 3rd tergite orange brown; vaguely near base of femora, telotarsi, apex of hind femur, apical half of hind tibia, hind tarsus largely black or blackish; basal half of hind tibia pale yellow; palpi, tegulae, veins and pterostigma dark brown; wing membrane slightly infuscate. Variation. Hind femur varies from apically black to entirely orange. Propodeum can be partly orangish in posterior part. Usually both sexes have hind trochanter (often also trochantellus) more or less infuscate and darker than the orange part of the hind femur, but this is scarcely evident in a series from S. Russia (MRC, NMS). Hind coxa varies from orange to black. Female is similar to the more distinctive male but is less conspicuously setose (Figs 419–421) and its ovipositor sheath is wide, with long setae and apically truncate (Fig. 410). Precoxal sulcus smooth to superficially rugulose medially. A female and a male from Romania (NMS) are slightly different from British ones; ocelli approx. 1/5 larger and frons coarsely rugose posteriorly. This appears to be reflected by a small divergence in CO1 (2.75 %), but for the moment we treat them as belonging to A. hirtus. Antennal segments of ♀ 54(1), 55(2), 56(6), 57(6), 59(2), 60(2), 61(1), of ♂ 56(1), 58(1), 59(3), 60(3), 61(1). Apical tergites of male type (1–)2, setae rather sparse but long and glabrous stripe consequently not always evident and fringe present but poorly differentiated. A female from

Mongolia (BZL) with completely black hind femur and base of hind tibia pale yellowish and with dark basal ring may be another very similar species. *Austria, *Belgium, *British Isles (England, Scotland, Ireland), *Bulgaria, *Czech Republic, Finland, *France, *Germany, *Hungary, *Netherlands, Norway, *Romania, *Russia, *Serbia, *Slovakia, *Switzerland, *Ukraine. 10.3897/zookeys.919.39642.figures409-410F0E870D7-8F26-5AC6-A26A-EE876EE0668EFigures 409, 410. Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 409 habitus lateral 410 ovipositor sheath lateral. https://binary.pensoft.net/fig/39010510.3897/zookeys.919.39642.figures411-42406C9A64C-18EA5F44-AA67-AAC4F9621E02Figures 411–424. Aleiodes hirtus (Thomson), ♀, Scotland, Peebles 411 fore wing 412 hind wing 413 mesosoma lateral 414 mesosoma dorsal 415 metasoma dorsal 416 fore femur lateral 417 hind femur lateral 418 antenna 419 head anterior 420 head dorsal 421 head lateral 422 base of antenna 423 apex of antenna 424 inner hind tarsal claw. https://binary.pensoft.net/fig/39010610.3897/zookeys.919.39642.figures425-42788C46B20-505F5D88-97D8-3EF0D8A85AF0Figures 425–427. Aleiodes hirtus (Thomson), ♂, England, Chippenham Fen 425 habitus lateral 426 head and anterior part of mesosoma lateral 427 propodeum and 1st –3rd metasomal segments lateral. https://binary.pensoft.net/fig/390107ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.

23. XML treatment for Aleiodes improvisus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396423EAD392B-2586-5259-9C5BB2EEDA721EB6 Aleiodes improvisus http://zoobank.org/BA464FC8-D95C-4CF8-A27F788BB7D2D3C4 van Achterberg & Shawsp. nov.Figs 428–431, 432–444, 445–452 Holotype, ♀ (ZJUH), “Austria: Tirol, Obergurgl, 2000 m, vii.[19]81, Day & Fitton”. Paratypes: 1 ♂ (NMS), “Switzerland: Valais, Aletschwald, 6000–7000 ft, 7–17.vi.1959, J.E. & R.B. Benson”; 1 ♂ (ZJUH), “Switzerland: Valais, J.E. & R.B. Benson, B.M. 1935581”, “Arolla, 6500 ft, 12.vi.1935”; 3 ♂ (ZJUH, NMS), “Austria: Tirol Vent., 1860 m, vii.1981, Fitton & Day”; 1 ♂ (ZJUH), “Austria: Tirol Vent (Winterstallen), 1750 m. vii.1981, Fitton & Day”. None. Unknown. Collected above the tree line in the Alps in June–July, and presumably univoltine. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 439); OOL of ♀ 1.8 × as long as diameter of posterior ocellus and densely rugose (Fig. 440); antenna of ♀ as long as fore wing; clypeus thick apically and not protruding anteriorly in lateral view (Fig. 441); lobes of mesoscutum densely punctate, interspaces largely finely coriaceous and with satin sheen; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 432); vein 1-CU1 of fore wing 0.3–0.6 × as long as vein 2-CU1; fore femur subparallel-sided (Fig. 437); hind tarsal claws yellowish or brownish bristly setose and with few dark brown or brown pectinal teeth submedially (Fig. 442); 3rd tergite longitudinally rugulose basally, without curved sculptural elements (Fig. 436); head of ♀ black; inner side of hind tibia of ♀ dark brown ventrally; palpi dark brown or blackish; hind trochanter and trochantellus largely dark brown; 2nd metasomal tergite of both sexes orange or dark reddish brown; 4th and 5th tergites black. Holotype, ♀, length of fore wing 5.5 mm, of body 7.7 mm. Head. Antennal segments of ♀ 41, antenna as long as fore wing, its subbasal and subapical segments rather robust (Fig. 444); frons with curved rugae; OOL 1.8 × diameter of posterior ocellus, densely rugose and with satin sheen; vertex densely rugose (also behind stemmaticum), with satin sheen; clypeus transversely rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 441); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 439); eye as long as temple in dorsal view (Fig. 440); clypeus below lower level of eyes; length of malar space 0.55 × length of eye in lateral view; temple striate near eye, and remainder rugose; head with long setae. Mesosoma. Mesoscutal lobes densely punctate, interspaces largely finely coriaceous and with satin sheen; precoxal area of mesopleuron coarsely vermiculate-rugose medially, but posteriorly rugose; mesopleuron remotely punctate and shiny medially; metapleuron densely rugose and rather dull; scutellum largely smooth (except for spaced punctures), shiny and nearly flat, with lateral carina; propodeum coarsely rugose but antero-laterally rugulose, laterally dorsal face longer than posterior one, somewhat angulate laterally but without tubercles, and with complete medio-longitudinal carina. Wings. Fore wing: r 0.4 × 3-SR; marginal cell fairly short (Fig. 432); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.8 × 3-SR; 2nd submarginal cell medium-sized (Fig. 432); cu-a vertical, straight; 1M slightly curved posteriorly; 1-SR wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell linearly widened, its apical width 2.1 × width at level of hamuli (Fig. 433); 2-SC+R quadrate; m-cu narrowly pigmented; M+CU:1-M = 30:21; 1r-m 0.7 × 1-M. Legs. Tarsal claws mainly setose but submedially with four rather short and dark brown pectinal teeth (Fig. 442); fore femur largely parallel-sided and rather slender (Fig. 437); hind coxa punctate and shiny, but dorsally mainly rugose; hind trochantellus rather robust; length of hind femur and basitarsus 3.3 and 5.7 × their width, respectively; length of inner hind spur 0.4 × hind

basitarsus. Metasoma. First tergite distinctly convex medially, its length 0.8 × apical width, robust and irregularly longitudinally rugose as 2nd tergite; both tergites with medio-longitudinal carina; mediobasal area of 2nd tergite triangular and small (Fig. 436); 2nd suture moderately deep and crenulate; basal half of 3rd tergite largely longitudinally striate, remainder of metasoma superficially microsculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide medially, with long setae and apically truncate (Fig. 429). Colour. Black; antenna (but only scapus partly yellowish), right fore coxa, trochanter, trochantellus, and femur (but left all yellowish brown except dark base of coxa and infuscated apex of femur), middle femur dorso-apically, middle coxa basally, hind trochanter, trochantellus and femur (but dorso-basally yellowish and left femur also ventrally), apical third of hind tibia (but left tibia yellowish ventrally), tegulae, pterostigma, veins largely, and metasoma ventrally largely dark brown; dorsal part of scutellum, 1st tergite laterally and narrowly medially and posteriorly, 2nd tergite and antero-laterally 3rd tergite orange brown; right fore tibia (except basally and left one yellowish brown) and tarsi more or less infuscate (but left fore tarsus only telotarsus dark brown); fore wing membrane somewhat infuscate, but hind wing nearly subhyaline. Variation. Eye of ♀ as long as temple in dorsal view (of ♂ 1.0–1.4 ×); length of malar space 0.5–0.6 × length of eye in lateral view; palpi black or largely dark brown; 1-CU1 0.3–0.6 × 2-CU1; length of fore wing 4.0–6.5 mm. Antennal segments: ♀ 41(1); ♂ 44(2), 49(2), 51(2). Male often has much darker legs (largely dark brown with coxae black as right legs of holotype, but legs are more extensively orange, including basal half of hind femur, in two paratypes) than female and scutellum black; metasoma similarly sculptured and coloured or also basal half of 3rd tergite orange brown or 1st tergite only posteriorly orange or only 2nd and 3rd tergites (except posteriorly) dark reddish brown; in the largest male paratype (Winterstallen) traces of inwardly curved sculpture are discernible posteriorly on the almost completely longitudinally rugose 3rd tergite; marginal cell of fore wing similar to ♀, with apical tergites type 1 and fringe not observed (Figs 450, 452). Austria, Switzerland. Improvisus is Latin for unexpected, unforeseen, because at first sight the specimens were expected to belong to A. gasterator or A. ruficornis. As suggested by its name this species can be easily confused with A. gasterator or A. ruficornis. It differs from A. gasterator mainly by being darker (subbasal antennal segments of ♀, hind trochanter and trochantellus, inner and dorsal side of hind tibia, parastigma) and somewhat higher number of antennal segments of ♀ (41 vs 29–39). Aleiodes ruficornis has an inflated fore femur (hardly or not inflated in A. improvisus), antenna of ♀ medium-sized (1.0–1.2 × fore wing vs 0.8–0.9 ×) and head of ♀ at least partly reddish brown. 10.3897/zookeys.919.39642.figures428-431E9026BF6-FCE7-5E98-AE08-34587C523785Figures 428–431. Aleiodes improvisus sp. nov., ♀, holotype 428 habitus lateral 429 ovipositor sheath lateral 430 right fore femur lateral 431 right hind femur lateral. https://binary.pensoft.net/fig/39010810.3897/zookeys.919.39642.figures432-444971CFB00-77C85C26-B9A6-D434A855ED2FFigures 432–444. Aleiodes improvisus sp. nov., ♀, holotype 432 fore wing 433 hind wing 434 mesosoma lateral 435 mesosoma dorsal 436 propodeum and 1st –3rd metasomal tergites dorsal 437 left fore femur lateral 438 left hind femur lateral 439 head anterior 440 head dorsal 441 head lateral 442 inner hind tarsal claw 443 base of antenna 444 apex of antenna. https://binary.pensoft.net/fig/39010910.3897/zookeys.919.39642.figures445-452D8EDEF9E-A0745F67-AA9E-AAFDD0186669Figures 445–452. Aleiodes improvisus sp. nov., ♂, paratype, Switzerland (Arolla) 445 habitus lateral 446 fore femur lateral 447 hind femur lateral 448 base of antenna 449 head dorsal 450 4th–7th metasomal tergites dorsal 451 apex of antenna 452 4th–7th metasomal tergites lateral. https://binary.pensoft.net/fig/390110

24. XML treatment for Aleiodes krulikowskii. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642795E8253-12E4-5D60-9CBBCA9878D5443B Aleiodes krulikowskii (Kokujev, 1898)Figs 453–454, 455–466 Rhogas (Rhogas) krulikowskiiKokujev, 1898: 302; Telenga 1941: 157–158. Rogas krulikowskii; Papp 1971: 360; Shenefelt 1975: 1235. Rogas (Rogas) krulikovskii; Tobias 1976: 83. Rogas (Rogas) krulikowskii; Tobias 1986: 77 (transl. 125) (lectotype designation). Aleiodes (Neorhogas) krulikowskii; Papp 1985a: 153, 1991a: 84; Belokobylskij 1996: 9. Aleiodes (Chelonorhogas) krulikowskii; Belokobylskij 1996: 2000: 31; Chen and He, 1997: 40; He et al. 2000: 665. Aleiodes krulikowskii; Fortier and Shaw 1999: 230; Papp 2004: 153. Rhogas csikiiSzépligeti, 1901 150. Rogas csikii; Shenefelt 1975: 1223. Aleiodes csikii; Papp 1991: 84 (as synonym of A. jaroslawensis); 2004: 216 (as synonym of A. krulikowskii). Lectotype of A. krulikowskii, ♀ (ZISP), “[Russia:] Kirovsk ts., Malmyzh, L.K. Krulikovsk, N. 1906”, “Rh. Krulikowskii Kokw., No. 1906”, “Lectotypus Rogas krulikowskii Kok., design. [V.I.] Tobias, 1980”. Holotype of A. csikii, ♀ (MTMA), “[Russia:] Siberia, Minusinsk, 30.vii.[18]98, Exp. Zichy, leg. Csiki”, “Holotypus ♀ Rhogas csikii sp. n. Szépl., 1901/ des. Papp J, 1967”, “Hym. Typ. No. 403, Museum Budsapest”, “Aleiodes ♀ krulikowskii Kok., det. Papp J., 1983”. 1 ♀ (MTMA), “[Romania: N Siebenburgen,] Radnai havas, Páváy V.F/ 26.vii.1906, 1400”; 1 ♂ (MTMA), “[Hungary:] Jaruer, 20.vi.”, “Rhogas carbonarius Gir. var. det. Szépligeti”; 1 ♀ (MTMA), “[Russia:] Ussuri, Kasakewitsch, 1907, Korb”; 1 ♀ (MTMA), “Mongolia: Suchebaator aimak, 44 km SSW von Baruun urt, 1050 m, Exp. Dr. Z. Kaszab, 1965, nr. 349, 2–3.viii.1965”; 1 ♂ (MTMA), “Mongolia: Cojbalsan aimak, Somon Chalchingol, 600 m, Exp. Dr. Z. Kaszab, 1965, nr. 409, 13.viii.1965”. None. Unknown. The collection dates (June–August) suggest that it is univoltine, but there is nothing to suggest how it overwinters. Maximum width of hypoclypeal depression 0.7–0.8 × minimum width of face (Fig. 463); OOL of ♀ approx. 1.3 × longer than diameter of posterior ocellus and coarsely rugose (Fig. 464); ventral margin of anterior part of clypeus thin, clypeus approx. 5 × wider than long medially (Fig. 463) and more or less protruding in lateral view (Fig. 465); head robust in anterior view (Fig. 463); lateral mesoscutal lobes densely and coarsely punctate, with interspaces narrower than punctures but interspaces becoming wider posteriorly, middle lobe coriaceous, but punctate near narrow and distinctly impressed notauli; mesopleuron densely and coarsely punctate, interspaces approx. equal to diameter of punctures or narrower; vein 1-CU1 of fore wing 0.8 × vein 2-CU1, widened and 1.1 × longer than vein m-cu; hind tarsal claws robust and with inconspicuous fine subbasal brownish pecten (Fig. 466); 1st and 2nd metasomal tergites comparatively slender and 1st tergite moderately widened (Fig. 459); 2nd tergite basally and 3rd tergite apically distinctly convex in lateral view (Fig.

453); 3rd tergite coarsely punctate, with complete lamelliform lateral margin (Fig. 453); hind coxa and femur completely dark brown or blackish; hind tibia usually ivory or pale yellowish basally; first and 2nd metasomal tergites reddish or orange. Lectotype, ♀, length of fore wing 6.9 mm, of body 9.6 mm. Head. Antenna incomplete, 32 segments remaining; frons mainly with curved or oblique rugae; OOL 1.3 × diameter of posterior ocellus, coarsely rugose and rather matt; vertex densely and rather finely rugose, hardly shiny; anterior part of clypeus densely punctate and flat; ventral margin of clypeus thin and rather forward protruding (Fig. 465); clypeus approx. 5 × wider than long medially; width of hypoclypeal depression 0.8 × minimum width of face (Fig. 463); length of eye 1.4 × temple in dorsal view (Fig. 464); vertex behind stemmaticum coarsely rugose; clypeus largely above lower level of eyes; length of malar space 0.2 × length of eye in lateral view; mandible punctate-rugose and with long setae (Fig. 465). Mesosoma. Lateral mesoscutal lobes densely and coarsely punctate, with interspaces narrower than punctures but interspaces becoming wider posteriorly, middle lobe coriaceous, but punctate near narrow and distinctly impressed notauli; precoxal area of mesopleuron and metapleuron coarsely and densely rugose punctate; remainder of mesopleuron densely punctate; metanotum with incomplete median carina; scutellum coarsely punctate; axilla largely densely rugose; propodeum rather flat and coarsely reticulate-rugose, medio-longitudinal carina on only anterior half. Wings. Fore wing: r 0.6 × 3-SR (Fig. 455); 1-CU1 horizontal, 0.8 × 2-CU1 and widened; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 455); cu-a vertical, straight; 1-M nearly straight posteriorly; 1-SR widened; surroundings of M+CU1, 1-M and 1-CU1 evenly setose; M+CU1 curved distally. Hind wing: marginal cell linearly widened, its apical width 2.8 × width at level of hamuli (Fig. 456); 2-SC+R quadrate; m-cu absent; M+CU:1-M = 40:21; 1r-m 1.1 × 1-M. Legs. Hind tarsal claws robust and with inconspicuous fine subbasal brownish pecten (Fig. 466); hind coxa largely densely finely punctate, dorso-posteriorly with oblique rugae; hind trochantellus slender; length of hind femur and basitarsus 4.3 and 5.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather convex and moderately widened (Fig. 459), 1.1 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugosepunctate; medio-basal area of 2nd tergite triangular and minute (Fig. 459); 2nd suture deep and rather wide; 1st tergite; 2nd tergite basally and 3rd tergite apically distinctly convex in lateral view (Fig. 453); 3rd tergite coarsely punctate, interspaces approx. equal to diameter of punctures, with complete lamelliform lateral margin; remainder of metasoma smooth and shiny; ovipositor sheath rather slender, with long setae and apically truncate (Fig. 454). Colour. Black; orbit near ocelli reddish brown; fore and middle legs (except blackish or dark brown coxae, trochanters and trochantelli), apex of hind trochantellus and basal third of hind tibia brownish yellow; tarsi darkened and remainder of legs dark brown; palpi (except basally) pale yellowish; mandible yellowish but basally largely dark brown; propleuron and tegula anteriorly dark brown and

tegula posteriorly brown; 1st and 2nd metasomal tergites and metasoma ventrally (except apically) orange brown; pterostigma dark brown; veins brown; wing membrane subhyaline. Variation. Orbit near hind ocellus sometimes only very slightly lighter in colour. Antennal segments: ♂ 68(1); according to the original descriptions of A. krulikowskii and A. csikii, the female types have 60 and 62 antennal segments, respectively. Apical tergites of ♂ type 2 and no fringe observed. *Hungary, Mongolia, *Romania, Russia (Central and Far East). 10.3897/zookeys.919.39642.figures453-4541F78D44C-E9E1-5669-88DD-191C40331D78Figures 453, 454. Aleiodes krulikowskii (Kokujev), ♀, lectotype 453 habitus lateral 454 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011110.3897/zookeys.919.39642.figures455-466F435B958-BEE45E37-B728-26A94FCA0EA3Figures 455–466. Aleiodes krulikowskii (Kokujev), ♀, lectotype 455 fore wing 456 hind wing 457 mesosoma lateral 458 mesosoma dorsal 459 metasoma dorsal 460 fore femur lateral 461 hind femur lateral 462 base of antenna 463 head anterior 464 head dorsal 465 head lateral 466 outer middle tarsal claw. https://binary.pensoft.net/fig/390112TelengaNA (1941) Family Braconidae, subfamily Braconinae (continuation) and Sigalphinae. Fauna USSR.Hymenoptera5(3): 1–466.PappJ (1971) Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 215. Braconidae (Hym.) 3.Annales Historico-Naturales Musei Nationalis Hungarici63: 307–363.ShenefeltRD (1975) Braconidae 8.

Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1– 12.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.HeJ-HChenXXMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.PappJ (2004) Type specimens of the braconid species by Gy. Szépligeti deposited in the Hungarian Natural History Museum (Hymenoptera: Braconidae).Annales Historico-Naturales Musei Nationalis Hungarici96: 153–223.

25. XML treatment for Aleiodes miniatus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396426706941D-AB04-5B9C-89F11F6D2D90A343 Aleiodes miniatus (Herrich-Schäffer, 1838)Figs 467–468, 469–481 Rogas miniatusHerrich-Schäffer, 1838: 156; Shenefelt 1975: 1238–1239 (type series lost). Rogas (Rogas) miniatus; Tobias 1976: 81, 1986: 75–76 (transl.: 122). Aleiodes (Neorhogas) miniatus; Papp 1987b: 36, 1991a: 88. Aleiodes (Chelonorhogas) miniatus; Belokobylskij et al. 2003: 398. Aleiodes miniatus; Bergamasco et al. 1995: 5; Papp 2005: 177. Rogas bicoloratusBoheman, 1853: 180; Shenefelt 1975: 1239 (as synonym of A. miniatus); Papp 2005: 177 (id.). Aleiodes formosusGiraud, 1857: 177; Shenefelt 1975: 1239; (as synonym of A. miniatus); Papp 1985a: 159 (lectotype designation and as synonym of A. miniatus), 2005: 177 [examined]. Lectotype of A. formosus, ♀ (MNHN), “[Austria, Wien,] Prata 16 juin”, “Austria, Vienne, Prater, 16 juin/Papp 1979”, “Lectotypus Aleiodes formosus Gir., 1857, ♀, Papp, 1979”, “Rogas miniatus HS ♀, det. Papp J., 1979”. Austria, Czech Republic, France, Finland, Germany, Hungary, Romania, Russia, Sweden, Ukraine, [Kazakhstan, Kyrgyzstan]. Specimens in ZJUH, BZL, SDEI, MNHN, MTMA, NMS, OUM, RMNH, ZSSM. The OUM specimen is labelled “Litchfield L.A. Carr 23” but there are very evidently numerous non-British specimens in the (now somewhat dispersed) Carr collection labelled Litchfield, and good reasons for discounting them as British are given by Perkins (1953). Such labelling may have been a means of identifying ownership of specimens at a time of considerable exchange and identification by others, and there is no evidence that this species has ever been collected in the British Isles. Material examined from central Europe (often labelled “Germany” or “Bohemia”) is mostly much more than 100 years old, when it seems to have been quite readily collected. Three recent specimens (NMS) from different sites in Sweden (Öland: Halltorp, 2015, 2017 and Skåne: Ravlunda, 2018, all N. Johansson) were swept from herb-rich sandy or gravelly grasslands overlying calcareous bedrock, with outstanding biodiversity partly maintained by grazing (Niklas Johansson, pers. comm.). The evident decline of A. miniatus in central Europe, as evidenced by specimen data showing a declining number of specimens collected in that region through time, probably reflects the loss of similar steppe habitat and, although a fairly recent (1994) specimen from Romania is in MTMA, it may now be extinct in large parts of central Europe. MRS950 (Sweden), MRS951 (Sweden). Unknown, but it seems to inhabit herb-rich calcareous steppe grasslands. Collected in (May)June– August; presumably univoltine, but we have not examined reared material of this large and distinctive species and there is no indication of how it may overwinter. A series in BZL (one now in NMS) is labelled “Wien D. Au” which can be interpreted as [? wet] woodland near the Danube (M. Schwarz, pers. comm.), which would probably be well under 200 m a.s.l. In contrast, a recent specimen (also in BZL) from Kyrgyzstan was collected higher at 2550 m. Maximum width of hypoclypeal depression approx. 0.5 × minimum width of face (Fig. 469); OOL of ♀ approx. twice as long as diameter of posterior ocellus and punctate (Fig. 477); ventral margin of clypeus thin and distinctly protruding in lateral view; length of malar space approx. equal to height of eye in lateral view (Fig. 478); mesoscutal lobes densely punctate; area of precoxal sulcus wide and coarsely rugose; length of vein 1-CU1 of fore wing 0.4 × vein 2-CU1; 2nd submarginal cell of fore wing short and square (Fig. 469); vein 1r-m of hind wing longer than vein 1-M; vein 2-SC+R of hind wing subquadrate; 3rd tergite densely punctate (Fig. 473); head and mesoscutum orange or brownish yellow; basal half of hind tibia (largely) pale yellowish; metasoma (except part of 1st tergite) orange or brownish yellow. Redescribed ♀ (RMNH) from Russia (Yaaseni). Length of fore wing 6.5 mm, of body 7.9 mm. Head. Antennal segments of ♀ 65, length of antenna 1.1 × fore wing, its subapical segments somewhat longer than wide; frons with coarse curved rugae; OOL 2.3 × diameter of posterior ocellus, and punctate; vertex densely punctate and shiny; clypeus densely punctate; ventral margin of clypeus thin and distinctly protruding forwards (Fig. 478); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 476); length of eye as long as temple in dorsal view (Fig. 477); vertex behind stemmaticum densely punctate; clypeus just below lower level of eyes; malar space 0.5 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, with minute interspaces and rather shiny; precoxal area of mesopleuron wide and coarsely rugose medially, mesopleuron above it coarsely and densely punctate, even speculum with some punctures; scutellum convex and punctate; propodeum evenly

convex and coarsely reticulate-rugose, medio-longitudinal carina incomplete. Wings. Fore wing: r 0.7 × 3-SR (Fig. 469); 1-CU1 horizontal, 0.4 × 2-CU1; r-m 1.2 × 3-SR; 2nd submarginal cell short (Fig. 469); cu-a inclivous, straight and rather short; 1-M rather curved posteriorly; 1-SR slender and short; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell evenly widened, its apical width 2.7 × width at level of hamuli (Fig. 470); 2SC+R subquadrate; m-cu absent; M+CU:1-M = 35:16; 1r-m 1.5 × 1-M. Legs. Tarsal claws with only three conspicuous brownish and widened bristles basally (Fig. 481); hind coxa densely and rather finely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 4.0 and 4.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite evenly convex, 0.9 × as long as wide apically; 1st tergite coarsely reticulaterugose, 2nd tergite coarsely and densely rugose-punctate, without median carina; medio-basal area of 2nd tergite short triangular (Fig. 473); 2nd suture deep and finely crenulate; basal half of 3rd tergite densely punctate, remainder of metasoma superficially micro-sculptured; apical half of 3rd tergite with sharp lateral crease; ovipositor sheath moderately wide, with long setae and apically rounded (Fig. 468). Colour. Brownish yellow; antenna, mesosternum, mesopleuron (except antero-dorsally), metapleuron, propodeum, 1st tergite, and ovipositor sheath black; propleuron, small patch on middle mesoscutal lobe anteriorly, apices of femora, fore and middle tibiae, tarsi, apical half of hind tibia, veins, and pterostigma dark brown; wing membrane subhyaline; basal half of hind tibia pale yellowish. Variation. Second submarginal cell square or somewhat narrower; propleuron dark brown or yellowish; mesopleuron black or yellowish anteriorly and dorsally; medio-longitudinal carina of posterior half of propodeum absent, obsolescent or incomplete. Antennal segments: ♀ 64(5), 65(3), 66(2), 67(3), 68(2), 70(1); ♂ 61(1), 64(2), 66(1), 67(2), 68(1), 69(1), 70(1). On this limited evidence there seems to be little, if any, difference in the number of antennal segments between the sexes. Males are very similar but have the metasoma infuscated apically and the apical tergites are type 3, setae short and dense, glabrous stripe rather narrow and fringe not observed. Austria, Czech Republic, Finland, *France, Germany, Hungary, Kazakhstan, *Kyrgyzstan, *Romania, Russia, Sweden, *Ukraine. 10.3897/zookeys.919.39642.figures467-468B69F968C-201E-5DCA-B4ADDCC6DBB65AE9Figures 467, 468. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 467 habitus lateral 468 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011310.3897/zookeys.919.39642.figures469-48112BE9EBC-14EC5ED6-B474-154B8040C7B8Figures 469–481. Aleiodes miniatus (Herrich-Schäffer)), ♀, Russia, Stavropolskij kraj 469 fore wing 470 hind wing 471 mesosoma lateral 472 mesosoma dorsal 473 metasoma dorsal 474 fore femur lateral 475 hind femur lateral 476 head anterior 477 head dorsal 478 head lateral 479 base of antenna 480 apex of antenna 481 inner hind tarsal claw. https://binary.pensoft.net/fig/390114ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1987b) First outline of the braconid fauna of Southern Transdanubia, Hungary (Hymenoptera, Braconidae), 3. Helconinae, Homolobinae, Macrocentrinae, Blacinae, Doryctinae and Rogadinae. A Janus Pannonius Muzeum Evkonyve 30–31 (1985–1986): 33–36.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1– 122.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341– 435.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PerkinsJF (1953) Note on British Ichneumonidae with descriptions of new species (Hym., Ichneumonidae).Bulletin of the British Museum (Natural History) (Entomology)3: 103–176. https://doi.org/10.5962/bhl.part.1050

26. XML treatment for Aleiodes morio. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964213CC2154-7801-57A1-BC88A29E700F37D9 Aleiodes morio (Reinhard, 1863)Figs 482–483, 484–495, 496, 497–503 Rogas morioReinhard, 1863: 255; Shenefelt 1975: 1239. Rogas (Rogas) morio; Tobias 1976: 83, 1986: 76 (transl.: 124). Aleiodes (Neorhogas) morio; Papp 1985a: 160 (♀ type lost, designation as lectotype), 1991a: 92. Aleiodes (Chelonorhogas) morio; Belokobylskij et al. 2003: 398. Aleiodes morio; Papp 2005: 177. Lectotype ♀ from southern Germany most probably lost (Papp 1985a); the only paralectotype (♂, ZMB) is a melanistic male of A. coxalis (Spinola, 1808) which does not fit the original description. The paralectotype was listed as A. tristis Wesmael by Papp (1985a). 1 ♀ (MTMA), “[Hungary:] Budapest, Rákospalita, 4.iv.%”, “Rhogas morio Reinh. ♀, det. Szépligeti”, “Aleiodes morio Reinh., ♀, det. Papp J., 1984”; 1 ♂ (MTMA), “[Hungary:] Nadap, 19– 21.iv.1951, Móczár Bajári”; 1 ♂ (ZSSM), “[? Germany:] Rogas n. sp.?, /: Ruthe, Berlin[?]/”; 1 ♀ (FMNH), “Suomi [= Finland:] U: Nurmijärvi, 6712:373, 13.v.1984, M. Koponen”; 2 ♀ + 1 ♂ (FMNH, RMNH), id., but 15.v.1984. None. Unknown. Specimens of both sexes have been collected in April and May, from which from which it is safe to surmise that the winter is passed in the mummy as the male would not hibernate. But we have seen no reared material nor any indication of habitat for this central European species. Its early flight time might be one reason why it is seldom collected and apparently rare. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face; OOL of ♀ 0.8 × (of ♂ 0.9 ×) diameter of posterior ocellus and rugose; ventral margin of anterior part of clypeus comparatively sharp and more or less protruding in lateral view (Fig. 493); head rather transverse (Fig. 492); mesoscutal lobes coriaceous; precoxal area of mesopleuron rugose medially; vein 1-CU1 0.3–0.4 × vein 2-CU1 and 0.3 × vein m-cu; hind tarsal claws with conspicuous and robust brownish pecten (Figs 494, 503); posterior orbit black; pterostigma of ♀ pale brown medially, of ♂ dark brown; coxae and femora completely black or dark brown; hind tibia usually ivory or pale yellowish basally; 1st and 2nd metasomal tergites of both sexes black. According to Papp (1985) most closely related to A. sibiricus (Kokujev), but that species does not have all black females and has the shape of the clypeus different. According to the original description the pterostigma is yellowish and laterally darkened, 1st subdiscal cell of the fore wing rather short, because vein cu-a distinctly more postfurcal than its own length (ca twice its own length) and meaning vein 1-CU1 of intermediate [approx. 0.6 ×] length of 2-CU1 and 0.8 × vein m-cu] [= “discoidali posterior brevior” as indicated for A. pallidicornis], precoxal sulcus area rugose medially, 1st tergite twice wider posteriorly than basally, hind leg black, except pale yellowish dorso-basal area of hind tibia and palpi dark brown.

Here we accept the interpretation of the first reviser (Szépligeti 1906) despite the difference in the shape of the 1st subdiscal cell, because it may be part of intraspecific variation. Redescribed ♀ (MTMA) from Hungary (Budapest). Length of fore wing 7.5 mm, of body 9.4 mm. Head. Antenna incomplete, 47 segments remaining (54 in lectotype), length of complete antenna approx. 0.9 × fore wing, its subbasal and subapical segments short; frons largely rugose; OOL 0.8 × diameter of posterior ocellus, and mainly rugulose and with satin sheen; depression near posterior ocellus smooth; vertex densely rugulose and with satin sheen; clypeus with some punctures; ventral margin of clypeus rather thin and protruding forwards (Fig. 493); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 491); length of eye 1.7 × temple in dorsal view (Fig. 492); vertex behind stemmaticum punctate-rugulose; clypeus largely above lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, with satin sheen; precoxal area of mesopleuron widely and densely rugose, but densely punctate posteriorly; middle of mesopleuron densely rugulose and dorsally coarsely rugose; metapleuron largely rugose; scutellum punctatecoriaceous; propodeum rather flat and densely rugose or rugulose, medio-longitudinal carina complete, and without protruding carinae laterally. Wings. Fore wing: r 0.5 × 3-SR (Fig. 484); 1-CU1 slightly oblique, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell robust and posteriorly somewhat diverging (Fig. 484); cu-a inclivous, straight; 1-M slightly curved posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 densely but inconspicuously setose. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 485); 2-SC+R short and subquadrate; m-cu present anteriorly; M+CU:1-M = 4:3; 1r-m 0.6 × 1-M. Legs. Tarsal claws bristly setose, medium-sized, and with robust pecten basally (cf. Fig. 494); hind leg missing in redescribed specimen. Metasoma. First tergite evenly convex, 0.9 × as long as wide apically; 1st tergite with mediolongitudinal carina; 1st and 2nd tergites and basal half of 3rd tergite finely and densely longitudinally rugulose; medio-basal area of 2nd tergite short triangular (Fig. 488); 2nd suture distinct and finely crenulate; remainder of metasoma superficially micro-sculptured; 4th tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Fig. 483). Colour. Black; palpi brownish yellow, but basally dark brown; tegulae pale yellowish; legs (except pale base of tibiae), metasoma ventrally and veins dark brown; pterostigma brown and medially yellowish brown; wing membrane subhyaline. Variation. Clypeus distinctly to moderately protruding and ventrally rather thin to thick. Antennal segments of ♀ 51(1), 52(1), 54(1), of ♂ 55(1). Males are very similar to the redescribed female (including the wing venation: Fig. 497), apical tergites type 1 with fringe not observed and probably absent (Fig. 498), mesopleuron rugulose or punctate medially and pterostigma entirely dark brown.

*Finland, Germany, Hungary. The lost lectotype from Germany had hyaline wings (which separates it from the A. carbonarius/ grassator/ carbonaroides complex), the pterostigma paler medially than laterally (entirely dark brown), base of the hind tibia pale yellow (black in ♂) and the body of ♀ entirely black (more or less yellowish or reddish). 10.3897/zookeys.919.39642.figures482-483719368AD-B6D0-51BF-BD6C-AA524F5F9B58Figures 482, 483. Aleiodes morio (Reinhard), ♀, Hungary, Budapest 482 habitus lateral 483 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011510.3897/zookeys.919.39642.figures484-4950CD0D1F6-47205E21-8A5D-5D8DFB044606Figures 484–495. Aleiodes morio (Reinhard), ♀, Hungary, Budapest, but 490 of ♂ from Nadap 484 fore wing 485 hind wing 486 mesosoma lateral 487 mesosoma dorsal 488 1st–3rd metasomal tergites dorsal 489 fore femur lateral 490 hind femur lateral 491 head anterior 492 head dorsal 493 head lateral 494 outer middle tarsal claw 495 base of antenna. https://binary.pensoft.net/fig/39011610.3897/zookeys.919.39642.figure49645A4FBA6-B8A1-5B8FACD9-3D00389B582EFigure 496. Aleiodes morio (Reinhard), ♂, Hungary, Nadap, habitus lateral.

https://binary.pensoft.net/fig/39011710.3897/zookeys.919.39642.figures497-50300E0EC37-09075628-B1CE-A04F977731DFFigures 497–503. Aleiodes morio (Reinhard), ♂, Hungary, Nadap 497 wings 498 apex of metasoma lateral 499 apex of metasoma dorsal 500 base of antenna 501 head anterior 502 head dorsal 503 inner hind tarsal claw. https://binary.pensoft.net/fig/390118ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.SzépligetiG (1906) Braconiden aus der Sammlung des ungarischen National-Museums, 1.Annales Historico-Naturales Musei Nationalis Hungarici4: 547–618.

27. XML treatment for Aleiodes nigrifemur. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642BC5C8E9C-EA96-55BC-B052089A49510904 Aleiodes nigrifemur http://zoobank.org/2535423C-36E7-4ECC-9FC3362039FC4928 van Achterberg & Shawsp. nov.Figs 504–505, 506–518 Holotype, ♀, (RMNH), “Greece, Peloponn[esus], Chelmos, 1700 m, 29.v.1987, H. Teunissen”. None. Unknown; the only known specimen was collected at the end of May which gives no clue of voltinism or how the winter is passed. Maximum width of hypoclypeal depression 0.6 × minimum width of face (Fig. 513); OOL of ♀ 1.6 × longer than diameter of posterior ocellus and rugulose (Fig. 514); width of clypeus intermediate apically, but strongly protruding in lateral view (Fig. 515); lobes of mesoscutum densely punctate, coriaceous between punctures; precoxal area widely rugose, and posteriorly punctate; vein 1-CU1 of fore wing 0.2 × vein 2-CU1 (Fig. 506); hind tarsal claws rather robust and with few yellowish pectinal teeth (Fig. 517); 3rd antennal segment of ♀ and basal third of hind femur black; basal third of hind tibia pale yellowish, contrasting with black basal half of hind femur. Similar to A. morio (Reinhard), but has pterostigma black (pale brown in A. morio), fore wing darkened apically (subhyaline), vein 1-M of hind wing linear with M+CU (angled); metasoma largely yellowish brown (entirely blackish) and eye in lateral view comparatively small (eye larger). Holotype, ♀, length of fore wing 7.2 mm, of body 8.2 mm. Head. Antennal segments of ♀ 62, length of antenna 1.1 × fore wing, its subapical segments medium-sized (Fig. 516); frons largely rugose; OOL 1.6 × diameter of posterior ocellus, rugulose and shiny; depression near posterior ocellus rugose; vertex largely rugose, rather shiny; clypeus rugulose; ventral margin of clypeus intermediate and distinctly protruding forwards (Fig. 515; as face dorsally); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 513); length of eye 1.3 × temple in dorsal view (Fig. 514); vertex behind stemmaticum rugulose; clypeus largely above lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate, rather shiny and interspaces coriaceous; precoxal area of mesopleuron widely rugose but posteriorly punctate, and area above it densely punctate or rugulose; metapleuron densely punctate dorsally and rugose ventrally; metanotum with short median carina anteriorly; scutellum remotely punctate, with some lateral rugae; propodeum rather short and flat, coarsely reticulate-rugose, medio-longitudinal carina complete, and without protruding carinae laterally.

Wings. Fore wing: r 0.5 × 3-SR (Fig. 506); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 506); cu-a inclivous, straight; 1-M curved posteriorly; 1-SR wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 1.9 × width at level of hamuli (Fig. 506); 2SC+R short and longitudinal; m-cu present anteriorly; vein 2-1A comparatively long (Fig. 506); M+CU:1-M = 24:37; 1r-m 0.65 × 1-M. Legs. Tarsal claws rather robust, bristly setose and few small yellowish teeth (Fig. 517); hind coxa largely punctate and with some oblique striae dorsally; hind trochantellus rather robust; length of hind femur and basitarsus 4.5 and 5.1 × their width, respectively; length of inner hind spur 0.45 à — hind basitarsus. Metasoma. First tergite rather flat medially, 0.8 × as long as wide apically; 1st tergite and anterior half of 2nd tergite with medio-longitudinal carina; 1st–2nd tergites densely longitudinally rugose; 3rd tergite (except posterior third) mainly rugulose; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 509); 2nd suture rather deep and crenulate; remainder of metasoma superficially micro-sculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 505). Colour. Black; maxillary palp apically, basal 0.4 of hind tibia and tegulae pale yellowish; mandible (but with dark brown patch), side of pronotum dorso-posteriorly, fore and middle tibiae, hind basitarsus basally, 1st tergite apically, 2nd–5th tergites orange brown; remainder of legs dark brown; remainder of palp, veins and pterostigma dark brown; lateral lobes of mesoscutum (except anteriorly and medially) dark reddish brown; wing membrane subhyaline, but apically infuscated (Fig. 506). Greece (main). The species is named after its black femur; niger is Latin for black, dark, dusky. 10.3897/zookeys.919.39642.figures504-505ED9C75DB-F56D-5A94-AD8DE777BD02E2A3Figures 504, 505. Aleiodes nigrifemur sp. nov. ♀, holotype 504 habitus lateral 505 ovipositor sheath lateral. https://binary.pensoft.net/fig/39011910.3897/zookeys.919.39642.figures506-518677289DC-BD215788-81C9-FB618A370D45Figures 506–518. Aleiodes nigrifemur sp. nov. ♀, holotype 506 wings 507 mesosoma lateral 508 mesosoma dorsal 509 1st–3rd metasomal tergites dorsal 510 fore femur lateral 511 hind femur lateral 512 base of antenna 513 head anterior 514 head dorsal 515 head lateral 516 apex of antenna 517 outer hind tarsal claw 518 antennae. https://binary.pensoft.net/fig/390120

28. XML treatment for Aleiodes nobilis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396421EFAF609-9B50-5562-98338120CCCF5138 Aleiodes nobilis (Haliday [in Curtis], 1834)Figs 519–521, 522–535, 536, 537–542 Rogas nobilisHaliday [in Curtis], 1834: 512; Papp 2005: 176 (as syn. of A. ductor). Rogas ductor var. nobilis; Shenefelt 1975: 1227. Aleiodes (Neorhogas) nobilis; Papp 1991a: 70 (as synonym of A. ductor). Aleiodes (Chelonorhogas) nobilis; van Achterberg 1997: 62 (both syntypes lost); Belokobylskij et al. 2003: 398. Aleiodes nobilis; Bergamasco et al. 1995: 5; O’Connor et al. 1999: 91–92; Papp 2005: 177. Rogas medianusThomson, 1892: 1668; Shenefelt 1975: 1237; van Achterberg 1997: 62 (as synonym of A. nobilis); Belokobylskij et al. 2003: 398 (id.); Papp 2005: 177 (id.) [examined]. Rogas (Rogas) medianus; Tobias 1976: 85, 1986: 80 (transl.: 133). Aleiodes (Neorhogas) medianus; Papp 1991a: 69; Belokobylskij 1996: 13. Aleiodes medianus; Papp and Vas 2016: 152. Aleiodes ductor; auct. p.p. Neotype of A. nobilis here designated: ♀ (NMS), “[Scotland:] W. Ross, Coppachy, Letterewe Estate, ix.2007, Mal. trap, P. Tinsley-Marshall”, “BCLDQR _00123”. Lectotype of A. medianus, ♀ (ZIL), “[Sweden:] Scan”, “medianus m.”, “Funnen vid Esperöd I Skåne, teste Papp J., 1983”, “Lectotypus Rogas medianus Thoms., 1891, ♀, Papp, 1983”, “Aleiodes medianus Th., ♀, det. Papp J., 1983”. The lectotype designation for A. nobilis is necessary for nomenclatural stability, because the type series is lost (van Achterberg, 1997) and the species has been confused with similar species in the past. The specimen from Scotland is selected neotype because it fits well the original description, Scotland is relatively close to both type localities (near Holywood in Ireland and Monk’s Wood in England) and it is in good condition. Austria, British Isles (Scotland: V.C.s 72, 88, 105; Ireland: V.C. H29), Bulgaria, Croatia, Czech Republic, Finland, Germany, Hungary, Italy, Moldova, Netherlands (LI: Gulpen; St. Pietersberg; Geulle (Bunderbos); NB: Udenhout (“de Brand”), OV: Voorst (Twello), ZH: Lexmond), Poland, Romania, Russia, Serbia, Slovakia, Sweden. Specimens in ALC, ZJUH, BZL, HSC, MTMA, NMS, NRS, RMNH, SDEI, Tullie House Museum Carlisle, USNM, ZSSM. MRS401 (Finland), MRS880 (Russia), MRS881 (UK). Collected predominantly in grassy places, June–October. Reared from the noctuid Autographa gamma (Linnaeus) (4 [1 NRS, 2 HSC], Germany, Sweden; H. Schnee) but, in view of its moderately northern areas of occurrence, it seems very likely that other plusiine noctuids would play an important part in its host range. The rearing data indicate that it is plurivoltine, and adult emergence in November from mummies forming in October suggests that it overwinters in the host larva. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 529); OOL of ♀ approx. as long as diameter of posterior ocellus and granulate (Fig. 530); ventral margin of clypeus thick and not protruding in lateral view (Fig. 531); mesoscutal lobes (as vertex) very finely and densely granulate, with satin sheen; precoxal area smooth; vein 1-CU1 0.7–1.3 × vein 2-CU1 and vein 1-CU1 wider than 2-CU1 (Fig. 522); tarsal claws with distinct dark brown pecten (Figs 534, 535); hind femur and basitarsus slender (Figs 519, 527); 1st metasomal tergite comparatively slender (Fig. 525); at least basal half of 4th–6th tergites of ♂ usually with long dense setosity (Figs 537, 538); head black; pronotum usually (partly) orange brown; both tegula and humeral plate equally yellowish; base of hind tibia pale yellowish; hind basitarsus brownish yellow, strongly contrasting with dark brown telotarsus; 2nd tergite yellowish or reddish. Redescribed ♀ (RMNH) from Slovakia (Kubrica). Length of fore wing 5.1 mm, of body 5.9 mm. Head. Antennal segments of ♀ 48, length of antenna 1.25 × fore wing, its subapical segments slender (Fig. 533); frons matt and granulate; OOL equal to diameter of posterior ocellus, and coriaceousgranulate; vertex coriaceous-granulate and rather dull; clypeus punctate-coriaceous; ventral margin of clypeus thick and not protruding forwards (Fig. 531); width of hypoclypeal depression 0.3 × minimum width of face (Fig. 529); length of eye 2.5 × temple in dorsal view (Fig. 530); vertex behind stemmaticum granulate; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely granulate, rather shiny near tegulae; precoxal area of mesopleuron smooth, surroundings sparsely punctulate; metapleuron mostly granulate; metanotum without median carina; scutellum granulate and with lateral carina; propodeum slightly convex, granulate with spaced rugosity, medio-longitudinal carina only anteriorly present, and no protruding carinae laterally. Wings. Fore wing: r 0.6 × 3-SR (Fig. 522); 1-CU1 straight, 1.2 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell short (Fig. 522); cu-a vertical, nearly straight; 1-M slightly curved posteriorly; 1-SR rather narrow; surroundings of M+CU1, 1-M and 1-CU1 evenly setose. Hind wing: marginal cell linearly widened, its apical width 2.4 × width at level of hamuli (Fig. 522); 2-SC+R subquadrate; m-

cu absent; M+CU:1-M = 14:13; 1r-m 0.6 × 1-M. Legs. Tarsal claws with conspicuous and robust dark brown pecten (Figs 534, 535); hind coxa sparsely finely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.7 and 8.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites rather regularly sublongitudinally striate, without medio-longitudinal carina on 2nd tergite; medio-basal area of 2nd tergite wide triangular and rather distinct (Fig. 525); 2nd suture rather deep and narrow; basal quarter of 3rd tergite finely striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath rather long and slender, with long setae and apically rounded (Fig. 520). Colour. Black; pterostigma (except yellowish extreme base and apex), veins (except brown vein C+SC+R), clypeus, apical third of hind tibia and telotarsus dark brown; palpi, tegulae, remainder of tibiae and tarsi, pale yellowish; apex of middle femur and apical half hind femur, black; remainder of legs, antenna (but apical segments and to some degree scapus infuscate) yellowish brown; 1st–3rd metasomal tergites (except black medial patch of 1st tergite), propleuron and pronotum orange; wing membrane subhyaline. Variation. 1-CU1 0.7–1.2 × 2-CU1; striae of 2nd tergite regularly sublongitudinal or somewhat diverging posteriorly (Fig. 525), but in male sometimes only granulate; basal third or half of 3rd tergite finely striate, rarely completely smooth; fore and middle femora black or dark brown apically or brownish yellow; pronotal side orange to dark brown dorsally; dark patch of 1st tergite absent (e.g., lectotype of A. medianus), small, large or occupying most of tergite; posterior half of 3rd tergite orange or black. Antennal segments: ♀ 46(3), 47(8), 48(3), 49(5), 50(5); ♂ 45(2), 46(2), 47(6), 48(5), 49(6); with little difference in the number of antennal segments between the sexes. Males are very similar, but apical tergites type 4, dense setae (making the tergites appear concave) and fringe strong (Figs 537, 538). *Austria, British Isles (Scotland, Ireland), Bulgaria, *Croatia, Czech Republic, Finland, Germany, Hungary, Italy, Moldova, Netherlands, *Poland, *Romania, Russia, *Serbia, *Slovakia, Sweden. 10.3897/zookeys.919.39642.figures519-521F8F63618-5681-57ED-B55E-15D28791425BFigures 519–521. Aleiodes nobilis (Haliday), ♀, neotype 519 habitus lateral 520 ovipositor sheath lateral 521 mummy of Autographa gamma Linnaeus (Germany, Lindenhayn). https://binary.pensoft.net/fig/39012110.3897/zookeys.919.39642.figures522-5355A75452D-61EA5EA4-90FF-37AA665EBD76Figures 522–535. Aleiodes nobilis (Haliday), ♀, neotype 522 wings 523 mesosoma lateral 524 mesosoma dorsal 525 1st–4th metasomal tergites dorsal 526 fore femur lateral 527 hind femur lateral 528 hind tibia lateral 529 head anterior 530 head dorsal 531 head lateral 532 base of antenna 533 apex of antenna 534 outer hind tarsal claw 535 outer fore tarsal claw. https://binary.pensoft.net/fig/39012210.3897/zookeys.919.39642.figure536BC54482C-1073-5A8A9C4F-56DE9E909C57Figure 536. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen, habitus lateral. https://binary.pensoft.net/fig/39012310.3897/zookeys.919.39642.figures537-54296B0B60C-355C5555-B8BF-D9826036FF20Figures 537–542. Aleiodes nobilis (Haliday), ♂, Netherlands, Gulpen 537 apex of metasoma dorsal 538 apex of metasoma lateral 539 apex of antenna 540 head anterior 541 head dorsal 542 base of antenna. https://binary.pensoft.net/fig/390124PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.van AchterbergC (1997) Revision of the Haliday collection of Braconidae (Hymenoptera).Zoologische Verhandelingen, Leiden314: 1–115.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341– 435.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.O’ConnorJPNashRvan AchterbergC (1999) A catalogue of the Irish Braconidae (Hymenoptera: Ichneumonoidea).Occasional Publication of the Irish Biogeographical Society4: 1–123.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1– 286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far

East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1–12.PappJVasZ (2016) Contributions to the knowledge of Braconidae fauna of Sălaj County, Romania

(Hymenoptera).Studia Universitatis “Vasile Goldiş”, Seria Ştiinţele Vieţii26(1): 141–152.

29. XML treatment for Aleiodes pallidicornis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964246C8126E-8FD3-54D4-B32E95E448D882E1 Aleiodes pallidicornis (Herrich-Schäffer, 1838)Figs 543–544, 545–557 Rogas pallidicornisHerrich-Schäffer, 1838: 156; Shenefelt 1975: 1241; Zaykov 1980b: 87. Rhogas pallidicornis; Fahringer 1932: 266. Rogas (Rogas) pallidicornis; Tobias 1976: 84, 1986: 80 (transl.: 130). Aleiodes (Neorhogas) pallidicornis; Papp 1987b: 36, 1991a: 70 (as senior synonym of A. hirtus). Aleiodes (Chelonorhogas) pallidicornis; Belokobylskij 2000: 42; Ku et al. 2001: 236; Belokobylskij et al. 2003: 398. Aleiodes pallidicornis; Papp 2005: 177. Rhogas pallidipennisDalla Torre, 1898: 221 [invalid emendation]. Rogas ductorauctt. p.p. [North & Central Europe, e.g., Lozan et al. 2010: 17]. Neotype of A. pallidicornis here designated, ♀ (RMNH), “[Netherlands], [Zuid-]Holland, Asperen, 6.viii.1972, C.J. Zwakhals”. The neotype designation is necessary for nomenclatorial stability, because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; no specimens could be found by the first author in ZMB), and the species has been confused with similar species in the past. The specimen is selected because it fits well the original description, Netherlands is relatively close to the probable type location in Germany and it is in excellent condition. Austria, Belarus, British Isles (Scotland: V.C. ?92), Bulgaria, Germany, Hungary, Italy, Montenegro, Netherlands (ZH: Asperen; Schoonrewoerd; Waarder), Romania, Russia, Slovakia, Switzerland, Turkey [Iran, North Korea]. Specimens in ZJUH, BZL, MRC, MSC, MTMA, NMS, RMNH, UNS, ZSSM. MRS885 (Russia). Very little is known. Specimens collected in (May) June–August (September), the great majority in June-July strongly suggesting that it is at least largely univoltine. The Dutch specimens were collected in fairly humid coppice woods. The single British specimen (ZJUH; G.T. Lyle) was reared (emergence 20.vi.1926) from a “noctua” caterpillar collected by E.A. Cockayne in Aberdeenshire. The mummy is lost. At that time, the term “noctua” was used generally for Noctuidae rather than in the restricted sense of the genus of that name, and it would appear (as Cockayne was by then a distinguished amateur lepidopterist) that the host larva did not belong to an obviously identifiable species. Otherwise we have not seen reared material, and there is no indication of how the winter is passed. Maximum width of hypoclypeal depression approx. 0.3 × minimum width of face (Fig. 552); OOL of ♀ approx. as long as diameter of posterior ocellus and remotely punctate with interspaces superficially granulate (Fig. 553); ventral margin of clypeus thick and not protruding in lateral view (Fig. 554); mesoscutal lobes and vertex very finely and densely granulate, with satin sheen; precoxal area smooth medially, but sometimes some rugae below it; vein 1-CU1 0.4–0.6 × vein 2-CU1 and equally slender (Fig. 545); tarsal claws with distinct dark brown pecten (Fig. 557); hind femur and basitarsus slender (Fig. 543); basal quarter of 3rd tergite largely finely striate; at least basal half of 4th–6th tergites of ♂ usually with long dense setosity; head and pronotum black; both tegula and humeral plate equally yellowish; base of hind tibia with narrow dark brown band; hind femur and tibia at least partly black or dark brown; 2nd tergite yellowish or reddish. Description. Neotype, ♀, length of fore wing 5.9 mm, of body 6.6 mm. Head. Antennal segments of ♀ 54, length of antenna 1.3 × fore wing, its subapical segments rather robust (Fig. 556); frons largely superficially granulate, anteriorly with some weak striae; OOL 1.4 × diameter of posterior ocellus, and punctate, interspaces granulate; vertex spaced punctate, shiny; clypeus densely and coarsely punctate, with granulate interspaces; ventral margin of clypeus thick and not protruding forwards (Fig. 554); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 552); length of eye 2.1 × temple in dorsal view (Fig. 553); vertex behind stemmaticum granulate with some transverse rugae; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate with largely granulate interspaces, with satin sheen; precoxal area of mesopleuron distinctly remotely punctate, interspaces larger than punctures; metapleuron densely punctate-granulate; metanotum with median carina; scutellum punctategranulate; propodeum evenly convex and coarsely rugose, its medio-longitudinal carina complete. Wings. Fore wing: r 0.4 × 3-SR (Fig. 545); 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 545); cu-a vertical, straight; 1-M nearly straight posteriorly; 1SR wide; surroundings of M+CU1, 1-M and 1-CU1 evenly setose. Hind wing: marginal cell gradually widened, its apical width 2.6 × width at level of hamuli (Fig. 546); 2-SC+R short and longitudinal; m-cu present basally; M+CU:1-M = 15:14; 1r-m 0.6 × 1-M.

Legs. Tarsal claws with rather small dark brownish pecten, absent near apical tooth (Fig. 557); hind coxa largely densely and finely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 5.0 and 7.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites with mediolongitudinal carina and coarsely irregularly rugose, but posteriorly 2nd tergite largely smooth and no median carina; medio-basal area of 2nd tergite triangular and rather large (Fig. 549); 2nd suture rather deep and finely crenulate; basal half of 3rd tergite smooth (except for punctuation) and shiny as remainder of metasoma; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath moderately wide, with long setae and apically truncate (Fig. 544). Colour. Black; hind tarsus largely infuscate, but 3rd and 4th segments paler than other segments; apices of fore and middle tibiae slightly infuscate, base of middle and hind tibiae and telotarsi dark brown; apical two-fifths of hind femur and hind tibia (except a pale yellowish band subbasally) black; remainder of legs, 1st and 2nd tergites, and 3rd tergite antero-laterally orange brown; palpi and tegulae brownish yellow; most of veins and pterostigma dark brown; wing membrane subhyaline. Variation. Antennal segments: ♀ 49(2), 50(2), 51(2), 52(6), 53(6), 54(1), 56(1); ♂ 50(1), 51(3), 52(1), 53(2), 54(2), 55(1), 56(3). On average males have ca two more antennal segments than females. Males are similar but have a large dark brown patch on 1st tergite, hind tarsus largely dark brown and apical tergites type 3, positioned rather posteriorly, setae long and fringe not observed. Austria, *Belarus, *British Isles (Scotland), Bulgaria, Hungary, *Iran, *Italy, *Montenegro, *Netherlands, North Korea, *Romania, Russia, *Slovakia, Switzerland, *Turkey. The type of Rogas pallidicornis Herrich-Schäffer, 1838, has been lost. Traditionally, it has been considered to belong to Aleiodes ductor (Thunberg, 1822), but the latter species is a synonym (see under A. unipunctator). The inadequate original description indicates that the 2nd tergite has diverging rugae, which excludes part of A. ductor auctt. Female specimens with yellowish or brownish palpi, basal half of the antenna yellowish and blackish hind tibia (except its pale yellowish base) fit well the original description of A. pallidicornis. 10.3897/zookeys.919.39642.figures543-544D134C7D6-B72A-58DA-81BBDBE6AFC61F2BFigures 543, 544. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 543 habitus lateral 544 ovipositor sheath lateral. https://binary.pensoft.net/fig/39012510.3897/zookeys.919.39642.figures545-55772F3170B-E74E530E-9B3B-85E813F12FC6Figures 545–557. Aleiodes pallidicornis (Herrich-Schäffer), ♀, neotype 545 fore wing 546 hind wing 547 mesosoma lateral 548 mesosoma dorsal 549 1st–3rd metasomal tergites dorsal 550 fore femur lateral 551 hind femur lateral 552 head anterior 553 head dorsal 554 head lateral 555 base of antenna 556 apex of antenna 557 inner hind tarsal claw. https://binary.pensoft.net/fig/390126ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980b) New species of the family Braconidae (Hymenoptera) for the Bulgarian fauna.Acta Zoologica Bulgarica16: 86–89. [in Bulgarian with English summary]FahringerJ (1932) Opuscula braconologica. Band 3. Palaearktischen Region. Opuscula braconologica 4: 241–320. Wagner, Wien.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1987b) First outline of the braconid fauna of Southern Transdanubia, Hungary (Hymenoptera, Braconidae), 3. Helconinae, Homolobinae, Macrocentrinae, Blacinae, Doryctinae and Rogadinae. A Janus Pannonius Muzeum Evkonyve 30–31 (1985–1986): 33–36.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.KuDSBelokobylskijSAChaJY (2001) Hymenoptera (Braconidae). Economic Insects of Korea 16. Insecta Koreana, Suppl.23: 1–283.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.LozanABelokobylskijSvan AchterbergCMonaghanM (2010) Diversity and distribution of Braconidae, a family of parasitoid wasps in the Central European peatbogs of South Bohemia, Czech Republic Journal of Insect Science 10: 16(1–21). https://doi.org/10.1673/031.010.1601

30. XML treatment for Aleiodes pallidistigmus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396422DDCBC74-9D5C-5AA5-9C5B4B625E197404 Aleiodes pallidistigmus (Telenga, 1941)Figs 558–559, 560–572 Rhogas (Rhogas) pallidistigmusTelenga, 1941: xii, 143, 177 (but also as palidistigma (p. 409) and pallidistigma (p. 420)) [examined]. Rogas pallidistigmus; Shenefelt 1975: 1241. Aleiodes (Neorhogas) pallidistigmus; Papp 1985b: 348. Aleiodes (Chelonorhogas) pallidistigmus; Chen and He 1997: 41; Fortier and Shaw 1999: 228; Belokobylskij 2000: 47; Ku et al. 2001: 237. Aleiodes pallidistigma; He et al., 2000: 667. Rogas heterostigmaStelfox, 1953: 149; Shenefelt 1975: 1233 [examined]. Syn. nov. Aleiodes (Neorhogas) heterostigma; Papp 1985a: 143, 146–147, 153, 1991a: 95. Aleiodes heterostigma; O’Connor et al. 1999: 91; Papp 2005: 177. Paratypes of A. heterostigma, 4 ♀ + 1 ♂ (ZJUH, USNM), “[Ireland], Rye Water, Co. KD, 5/9.vii. [19]42, AWS[telfox]”; 1 ♀ (RMNH), id., but 5.vii.1942; 1 ♀ (NMI), id., but 8.vii.1948; 1 ♂ (NMI), “[Ireland], Woodbrook, OC, 26.vi.[19]38”. Holotype of A. pallidistigmus (♀, ZISP) from Far East Russia (Primorsky Krai, Ussuri area, Vinogradovka, 10.viii.1929, Kiritshenko). 1 ♀ (RMNH), “Belgium: Liège, Mt. Rigi, 650 m, 2.viii.1986, at light, C. v. Achterberg, RMNH”; 1 ♀ (MTMA), “Dania [= Denmark]: S-Jutland, Kragelund Mose, near Baekke, 11.viii.1973, [T.] Munk”; 1 ♀ (NMS), “Wales: Cereigion, Rhôs Rhydd, SN572738, Molinia bog, 30.vii.1987, NCC Welsh Peatland Survey, P. Holmes, NMSZ 1996.023”; 1 ♂ (NMS) “Wales; Ceredigion, Comin Esgair Maen, SN652649, Equisetum bog, 23.vii.1987, NCC Welsh Peatland Survey, P. Holmes, NMSZ 1996.023”; 1 ♂ (FMNH), “Fennia [= Finland]: Helsinki, 27.vii.1978, O. Ranin”; 1 ♀ (NMS) Far East Russia, Anismovka v. Shkotova/S Primorje reg., 11–13.viii.2003, leg. Osipov”; 2 ♀ (MRC) “Far East Russia, S. Primorje reg., Lazo distr. Valentin 17–18.vii.2003 leg. Osipov”: 1 ♀ (NMS) [Russian, Far East]; 2 ♀ (RMNH), “China: Jilin, Gomngzhuling, 43°5'N, 124°8'E, viii–ix.1983, Wang Chenghun”. None. Unknown for West Palaearctic populations. Specimens have been collected in (June)July-August in open boggy areas, certainly at least sometimes over limestone. Presumably univoltine, but we have not seen reared material from West Palaearctic and the overwintering mode is unclear. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 568); OOL of ♀ approx. 1.3 × as long as diameter of posterior ocellus and densely granulate (Fig. 569); penultimate segments rather slender and antenna 1.5 × as long as fore wing; ventral margin of clypeus thick, not protruding in lateral view; mesoscutal lobes finely granulate-punctulate and matt; precoxal area coarsely rugose; marginal cell of fore wing of ♀ ending rather close to wing apex (Fig. 560); vein 1CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 560); hind tarsal claws rather robust and only brownish setose (Fig. 571); 3rd tergite and basal half of 4th tergite coriaceous and dull; labial palp yellowish brown or brown; basal half of hind tibia reddish or yellowish, slightly paler than basal half of hind femur, and its apex reddish or yellowish; 4th and 5th tergites black. Redescribed ♀ paratype of A. heterostigma (RMNH) from Ireland (Rye Water). Length of fore wing 4.9 mm, of body 6.3 mm. Head. Antennal segments of ♀ 58, length of antenna 1.1 × fore wing, its subapical segments rather robust (Fig. 567); frons largely superficially granulate; OOL 1.8 × diameter of posterior ocellus, and superficially rugulose-granulate and shiny; vertex superficially rugulose-granulate, rather shiny; clypeus with some punctures; ventral margin of clypeus thick and not protruding forwards (Fig. 570); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 568); length of eye 1.3 × temple in dorsal view (Fig. 569); vertex behind stemmaticum superficially granulate-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate-granulate, matt; precoxal area of mesopleuron coarsely rugose medially; remainder of mesopleuron coarsely punctate, with some

rugae near speculum and interspaces superficially granulate; scutellum rather flat, punctulategranulate and with weak lateral carinae; propodeum rather convex, shiny and coarsely rugose, medio-longitudinal carina distinct only on its anterior half. Wings. Fore wing: r 0.3 × 3-SR (Fig. 560); 1-CU1 horizontal, 0.45 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell short (Fig. 560); cu-a inclivous, straight; 1-M slightly curved posteriorly; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell gradually widened with basal half rather narrow (Fig. 561), its apical width 2.6 × width at level of hamuli; 2-SC+R short longitudinal; m-cu weakly developed; M+CU:1-M = 35:26; 1r-m 0.5 × 1-M. Legs. Tarsal claws rather robust and only brownish setose (Fig. 572); hind coxa coarsely punctate, dorsally with oblique striae; hind trochantellus robust; length of hind femur and basitarsus 4.1 and 4.6 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically; 1st and 2nd tergites and base of 3rd tergite finely and irregularly longitudinally rugose, with medio-longitudinal carina weak; mediobasal area of 2nd tergite triangular and short (Fig. 564); 2nd suture rather deep and crenulate; apical half of 3rd tergite punctate-granulate, remainder of metasoma smooth except for some superficial micro-sculpture; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 559). Colour. Black; telotarsi largely and basal quarter of antenna dark brown; palpi, tegulae and pterostigma pale yellow; remainder of legs, 1st and 2nd tergites, basal half of 3rd tergite largely and pronotum orange brown; veins brown; wing membrane subhyaline. Variation. Pronotum anteriorly and basal half of antenna orange brown to dark brown, pterostigma is yellowish to largely (except base) rather dark brown; length of malar space 1.0–1.4 × basal width of mandible; OOL 0.7–1.8 × diameter of ocellus and metapleuron medially more or less punctate, rugulose-coriaceous or rugose. Antennal segments: ♀ 54(1), 58(1), 59(4), 60(2), 62(1), 64(1); ♂ 59(1), 60(1), 62(1). Apical tergites of ♂ type 1, fringe absent. *Belgium, British Isles (Ireland, Wales), China, *Denmark, *Finland, Russia (Far East). We tried to separate the East Palaearctic A. pallidistigmus from the West Palaearctic A. heterostigma, but efforts were in vain. The differences such as the colour of the basal half of the antenna (dark brown in A. heterostigma and usually yellowish or brown in A. pallidistigmus), the eyes and ocelli often smaller, OOL 1.1–1.8 × diameter of ocellus (0.7–1.4 ×), malar space 1.2–1.3 × basal width of mandible (1.0–1.4 ×) and metapleuron with a shiny and more or less punctate area (less shiny and rugulose-coriaceous or rugose) are too variable to justify separation of A. heterostigma. Therefore, we synonymise A. heterostigma with A. pallidistigmus (syn. nov.). 10.3897/zookeys.919.39642.figures558-55942663AF1-09C8-50CC-89B5-15D17A31DB04Figures 558, 559. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 558 habitus lateral 559 ovipositor sheath lateral. https://binary.pensoft.net/fig/39012710.3897/zookeys.919.39642.figures560-57281DECFA2-65355228-8678-F741198A7730Figures 560–572. Aleiodes pallidistigmus (Telenga), ♀, Denmark, Kragelund Mose 560 fore wing 561 hind wing 562 mesosoma lateral 563 mesosoma dorsal 564 1st–3rd metasomal tergites dorsal 565 fore femur lateral 566 hind femur lateral 567 apex of antenna 568 head anterior 569 head dorsal 570 head lateral 571 outer hind tarsal claw 572 base of antenna. https://binary.pensoft.net/fig/390128ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1985b) Braconidae (Hymenoptera) from Korea. VII.Acta Zoologica Hungarica31: 341–365.ChenX-XHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far

East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.KuDSBelokobylskijSAChaJY (2001) Hymenoptera (Braconidae). Economic Insects of Korea 16. Insecta Koreana, Suppl.23: 1–283.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18.

Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1– 122.O’ConnorJPNashRvan AchterbergC (1999) A catalogue of the Irish Braconidae (Hymenoptera: Ichneumonoidea).Occasional Publication of the Irish Biogeographical Society4: 1–123.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137– 194.

31. XML treatment for Aleiodes periscelis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396421A94464D-518E-5653-88100820C4462E09 Aleiodes periscelis (Reinhard, 1863)Figs 573–576, 577–589, 590, 591–603 Rogas periscelisReinhard, 1863: 254; Shenefelt 1975: 1242; Kotenko 1992: 96 [examined]. Rhogas (Rhogas) periscelis var. charkowensisKokujev, 1898: 297. Rhogas (Rhogas) periscelis var. charkoviensis[sic!]; Telenga 1941: 163, 164. Rogas (Rogas) periscelis; Tobias 1976: 86, 1986: 81 (transl. 133); Papp 1983: 330. Aleiodes (Neorhogas) periscelis; Papp 1985a: 161 (lectotype designation), 1985b: 348; 1991a: 79, 1991c: 641, 1994: 307. Aleiodes (Chelonorhogas) periscelis; Belokobylskij 2000: 36. Aleiodes periscelis; Fortier and Shaw 1999: 230; Belokobylskij and Taeger 2001: 115; Belokobylskij et al. 2003: 400 (excluded from German checklist); Papp, 2005: 177. Rhogas jaroslawensisKokujev, 1898: 302. Syn. nov. Rhogas (Rhogas) jaroslavensis; Telenga 1941: 176, 408 (invalid emendation). Rogas jaroslawensis; Shenefelt 1975: 1235. Rogas (Rogas) jaroslavensis; Tobias 1976: 85. Rogas (Rogas) jaroslawensis; Tobias 1986: 81 (transl.: 133; as synonym of A. rufipes (Thomson)). Aleiodes (Neorhogas) jaroslawensis; Papp 1985a: 153, 1991a: 80. Aleiodes jaroslawensis; Papp 2005: 177 (as valid species). Lectotype of R. periscelis, ♂ (ZMB), “[Austria:] Neusiedler See”, “Type”, “Coll. H. Rhd.”, “26696”, “periscelis Gir. [= from Giraud]”, “Lectotypus Rogas periscelis Rhd., 1863, ♂, Papp, 1982”, “Aleiodes periscelis Rhd., ♂, det. Papp J., 1983”; 2 ♂ paralectotypes (MNHN), one with “[Austria:] Prater, Mai”, “ex coll. Giraud”. Holotype of R. jaroslawensis, ♀ (ZISP; examined photos made by K. Samartsev), “[S. Russia:] Berditsino [Yaroslavskiy rayon, 57.454N, 40.108E], 22.vi.1892, A.M. Yakovlev, 1909”, “Rh. jaroslawensis Kokw., No. 1909” and with a round golden label. Czech Republic, Germany, Hungary, Russia. Specimens in ALC, BZL, MTMA, NMS, SDEI, ZISP, ZMB, ZSSM. None. Unknown but presumably univoltine. Specimens of both sexes collected in April and May suggest that the winter is passed in the mummy. We have not seen reared material, but several Hungarian specimens appear to have been collected in Quercus-dominated woodland, but without indication of any association with Quercus as such. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Figs 573, 584); OOL of ♀ 1.5 × as long as diameter of posterior ocellus (Fig. 585; of ♂ 1.4 ×; Fig. 599), rugulose or rugose and with satin sheen; ventral margin of clypeus thick and not protruding anteriorly (Fig. 586); mesoscutum remotely punctulate and with satin sheen, interspaces of lateral lobes largely smooth, of middle lobe superficially coriaceous; area of precoxal sulcus smooth and shiny; length of vein 1CU1 of fore wing 0.3–0.4 × vein 2-CU1 and 0.5 × vein m-cu; vein 2-SC+R of hind wing subquadrate; hind basitarsus robust; head (including basal half of mandible) black; antenna of ♀ (except scapus and pedicellus, and apically darkened) brownish yellow; apex of hind femur usually largely black dorsally; basal half of hind tibia (largely) pale yellowish; fore coxa dark brown; 2nd tergite of ♀ orange or dark reddish brown, of ♂ largely black; 4th–6th tergites of males flat, and with long dense setosity (Fig. 590). Redescribed ♀ (BZL), Czech Republic (Pisek); length of fore wing 5.8 mm, of body 8.1 mm. Head. Antennal segments 45 (holotype ♀ of A. jaroslawensis: 42), length of antenna approx. as long as fore wing, its subbasal and subapical segments robust (Figs 587, 588); frons largely smooth anteriorly and with curved rugae posteriorly; OOL 1.5 × diameter of posterior ocellus, rugulose and shiny; vertex finely rugose and with satin sheen; clypeus slightly convex and mainly transversely aciculate; ventral margin of clypeus thick and not protruding anteriorly (Fig. 586); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 584); length of eye 1.5 × temple in dorsal view (Fig. 585); vertex behind stemmaticum rugulose; clypeus below lower level of eyes; length of malar space 0.45 × length of eye in lateral view and 1.3 × basal width of mandible. Mesosoma. Mesoscutum remotely punctulate and with satin sheen, interspaces of lateral lobes largely smooth, and of middle lobe superficially coriaceous; scutellum superficially punctate, laterally rugose; precoxal area of mesopleuron smooth and shiny; metapleuron largely densely punctate, but ventrally coarsely rugose; metanotum with distinct median carina anteriorly; propodeum rather flat and coarsely vermiculate rugose, medio-longitudinal carina complete, and slightly tuberculate laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 577); 1-CU1 horizontal, 0.3 × as long as 2-CU1; r-m 0.6 × 3SR; 2nd submarginal cell medium-sized (Fig. 577); cu-a vertical, straight; 1-M nearly straight posteriorly and subparallel; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 densely setose. Hind wing: marginal cell gradually widened (but less so in its basal third) and apical width 2.2 × width at level of hamuli (Fig. 578); 2-SC+R quadrate; m-cu short; M+CU:1-M = 40:33; 1r-m 0.5 × 1-M. Legs. Tarsal claws with conspicuous and medium-sized brownish pecten, remaining removed from tarsal tooth (Fig. 589); hind coxa largely coriaceous-punctate, but dorsal besides smooth depression rugose; hind trochantellus rather robust; length of hind femur 3.7 × its width. Metasoma. First tergite rather flat posteriorly, wide subbasally and 0.9 × longer than wide apically; 1st and 2nd tergites with coarse medio-longitudinal carina and coarsely longitudinally rugose, but posterior quarter of 2nd tergite rather finely rugose; medio-basal area of 2nd tergite triangular and wide (Fig. 581); 2nd suture moderately deep, finely crenulate and narrow; basal two-thirds of 3rd

tergite finely longitudinally rugose, remainder of metasoma superficially micro-sculptured and with satin sheen; 4th and apical half of 3rd tergites without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Figs 574, 576). Colour. Black; palpi dark brown basally and remainder pale brown; antenna (except dark brown scapus and pedicellus), tegulae (but anteriorly dark brown), middle and hind trochanters and trochantelli brownish yellow; fore coxa, trochanter and femur dark brown; basal 0.4 of hind tibia ivory and remainder black; remainder of legs (but hind femur with a blackish patch dorso-apically), 1st and 2nd tergites and basal two thirds of 3rd tergite, largely dark reddish brown; pterostigma dark brown; veins mainly yellowish brown, but medially brown (Figs 577, 578); wing membrane subhyaline. Variation. Holotype of A. jaroslawensis has apex of hind femur yellowish brown (Fig. 574). Antennal segments: ♀ 42(1), 45(1); ♂ 50(1), 53(1), 54(2), 56(1). Males appear to have ca ten more antennal segments than females. Male has apical tergites type 1, setae moderately long and fringe not observed, probably absent (Fig. 590); antenna rather dark brown, but scapus largely blackish brown and antennal segments slightly slenderer than of female and 1.1–1.2 × as long as fore wing; metasoma black, but extreme apex of 1st tergite yellowish brown; OOL approx. 1.4 × width of posterior ocellus. Austria, Czech Republic, Germany, Hungary, Russia. The ♀ holotype of Rhogas jaroslawensis lacks the antennae, but according to the original description the antenna was 42-segmented, distinctly shorter than the body, reddish brown, except for the darkened apex and the black scapus. This and the other characters still visible agree well with our interpretation of A. periscelis (except that the hind femur is yellowish brown apically); therefore, we synonymise R. jaroslawensis with A. periscelis (syn. nov.). 10.3897/zookeys.919.39642.figures573-5766BE0F66C-89AD-5079-9229-E7B04609AF8DFigures 573–576. Aleiodes periscelis (Reinhard), ♀, 573, 574 Russia (holotype R. jaroslawensis Kokujev) and 575, 576 Czech Republic, Pisek 573 head anterior 574, 575 habitus lateral 576 ovipositor sheath lateral. Photographs 573, 574 by K. Samartsev. https://binary.pensoft.net/fig/39012910.3897/zookeys.919.39642.figures577-5896D41C33E-DB8B5356-A8A0-2EFA2AEC1DAFFigures 577–589. Aleiodes periscelis (Reinhard), ♀, Czech Republic, Pisek 577 fore wing 578 hind wing 579 mesosoma lateral 580 mesosoma dorsal 581 metasoma dorsal 582 fore femur lateral 583 hind femur lateral 584 head anterior 585 head dorsal 586 head lateral 587 base of antenna 588 apex of antenna 589 outer middle tarsal claw. https://binary.pensoft.net/fig/39013010.3897/zookeys.919.39642.figure59013547743-5004-59EFB602-617196095EE7Figure 590. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov, habitus lateral. https://binary.pensoft.net/fig/39013110.3897/zookeys.919.39642.figures591-6032B48EACD-704C52D4-AC60-E77539DD4B55Figures 591–603. Aleiodes periscelis (Reinhard), ♂, Russia, Serpukhov 591 wings 592 mesosoma lateral 593 mesosoma dorsal 594 1st –5th metasomal tergites dorsal 595 fore femur lateral 596 hind femur lateral 597 hind tarsus lateral 598 head anterior 599 head dorsal 600 head lateral 601 base of antenna 602 apex of antenna 603 inner hind tarsal claw. https://binary.pensoft.net/fig/390132ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.KotenkoAG (1992) A contribution to the fauna of Braconidae (Hymenoptera) of Dauria. In: Amirkhanov AM (Ed.) Insects of Dauria and adjacent territories: 94–107.Izdatelstvo Tsentralnoi Nauchno issledovatelskoi laboratorii okhotnichego khozyaistva i zapovednikov, Moskva, 141 pp.TelengaNA (1941) Family Braconidae, subfamily Braconinae (continuation) and Sigalphinae. Fauna USSR.Hymenoptera5(3): 1–466.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1983) A survey of the Braconid fauna of the Hortobágy National Park (Hymenoptera, Braconidae), II: 315–337. In: Mahunka S (Ed.) The Fauna of the Hortobágy National Park II. Budapest.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.BelokobylskijSATaegerA (2001) Braconidae: 103–115. In: DatheHHTaegerABlankSM (Eds) Verzeichnis der Hautflügler Deutschlands (Entomofauna Germanica 4).Entomologische Nachrichten und Berichte (Dresden), Beiheft7: 1–178.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.

32. XML treatment for Aleiodes pulchripes. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642252D41F9-7D7C-5A97-BB696CD6F339C89E Aleiodes pulchripes Wesmael, 1838Figs 604–607, 608–621, 622–626 Aleiodes pulchripesWesmael. 1838: 102; Čapek and Lukás 1989: 31; Papp 1991a: 73, 2005: 177; Fortier and Shaw 1999: 224; O’Connor et al. 1999: 92; Belokobylskij et al. 2003: 398 [examined]. Rogas pulchripes; Shenefelt 1975: 1245. Rogas (Rogas) pulchripes; Tobias 1976: 83, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) pulchripes; Papp 1985a: 149, 153, 161, 1991a: 73. Aleiodes pulchricornisKolubajiv, 1962: 27; Shenefelt 1975: 1245 (invalid emendation; not A. pulchricornis (Szépligeti, 1902)); Papp 2005: 177 (as synonym of A. pulchripes). Holotype of A. pulchripes, ♂ (KBIN), “A. pulchripes ♂ mihi”, “A. pulchripes mihi, dét. C. Wesmael”, “Coll. Wesmael”, “Belgique, Charleroi/ teste Papp J., 1983”, “Holotypus”, “Aleiodes pulchripes Wesm., 1838, ♂, Papp, 1983”. Austria, British Isles (England: V.C. 59; Isle of Man: V.C. 71: Ireland: V.C. H21), Czech Republic, Finland, Germany, Hungary, Netherlands (GE: Vierhouten; ZH: Leiden; NH: Amsterdam; Sloten), Russia, Sweden. Specimens in ZJUH, CNC, IKC, KBIN, MSC, MTMA, NMS, NRS, RMNH, USNM, UWIM, ZISP, ZSSM. MRS847 (Sweden), MRS873 (Sweden). Collected in (June)July and August. Univoltine, overwintering in an exposed mummy. Reared from the following arboreal acronictine Noctuidae: Acronicta aceris (Linnaeus) (2 [CNC, MSC], Austria, Germany; J. Schwarz), Acronicta psi (Linnaeus) (22 [1 NRS, 2 ZISP]; M.R. Shaw), Acronicta leporina (Linnaeus) (1 [IKC], Finland; M.J. Pellinen), Acronicta tridens (Dennis & Schiffermüller) (4:2; M.R. Shaw), Acronicta psi or tridens (2), indet. Acronictinae (1). A quantitative account of rearing this species at its only known English site, comprising old hedges rich in Sorbus aucuparia bordering a largely reclaimed peat bog, is given by Shaw (1979). Experimental rearings were unfortunately limited to unobserved exposures of multiple hosts in closed boxes; extremely hot weather marred the results, but from one box containing 15 of each of Subacronicta megacephala (Dennis & Schiffermüller) and A. tridens, the surviving 13 S. megacephala were dissected after three days of exposure and contained no hosts, while at least eight of the A. tridens contained parasitoids (two found by dissection + six mummies formed; of the other seven, one contained no parasitoid on dissection + six resulted in moths). This suggests that S. megacephala is outside the host range. Similar but less well quantified experiments also excluded the low-feeding Acronicta rumicis (Linnaeus) and the arboreal lymantriine ErebidaeEuproctis similis (Fuessly). It is worth adding that the rather frequent citation of lymantriine hosts in the literature can undoubtedly be explained by misidentification of the setose and rather colourful larvae of most species of arboreal acronictine noctuids. The mummy is dark grey in colour, leaving only little evidence of the patches of bright colour that had been a feature of the host larva. It forms in the caudal part of the host, the anterior

segments of which strongly contract towards the extensive point of attachment, and the cocoon occupies approx. 4th–7th abdominal segments. As mummification approaches, the host aligns itself on a narrow aerial twig to which the mummy becomes ventrally adpressed, thus leaving a weakly arched dorsal profile bearing a strong resemblance to an overwintering lateral bud (e.g., of Sorbus aucuparia: Fig. 605). Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 616); OOL distinctly less than diameter of posterior ocellus, largely smooth but micro-sculptured near eyes; ventral margin of clypeus thick and not protruding in lateral view (Fig. 618); mesoscutal lobes coriaceous; mesopleuron (including precoxal sulcus area) nearly or completely smooth; propodeum with pair of crest-like protuberances laterally; vein 1-CU1 of fore wing much shorter than vein 2CU1; basal half of marginal cell of hind wing parallel-sided (Fig. 609); tarsal claws with large dark brown pecten up to apical tooth of claw (Fig. 621); hind spurs dark brown; hind tibial spurs of ♂ obtuse apically (Fig. 624); head black; pterostigma pale yellowish or light brown; mesopleuron, mesosternum and scutellum brownish yellow; apex of hind femur yellowish or reddish; basal half of hind tibia pale yellowish. Redescribed ♀ (RMNH) from England (Chat Moss). Length of fore wing 5.3 mm, of body 6.5 mm. Head. Antennal segments of ♀ 56, length of antenna 1.3 × fore wing, its subapical segments slender (Fig. 620); frons largely smooth; OOL 0.3 × diameter of posterior ocellus, and smooth near ocelli, but micro-sculptured near eye, shiny; vertex largely smooth, with few punctures, shiny; face crestlike protruding medio-dorsally; clypeus densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 618); width of hypoclypeal depression 0.35 × minimum width of face (Fig. 616); length of eye 5.6 × temple in dorsal view (Fig. 617); vertex behind stemmaticum mainly smooth but partly rugulose; clypeus above lower level of eyes; length of malar space 0.15 × length of eye in lateral view. Mesosoma. Mesoscutal lobes coriaceous, rather shiny; precoxal area of mesopleuron smooth as most of mesopleuron; metanotum with medio-longitudinal carina anteriorly; scutellum finely punctate, interspaces smooth, but posteriorly coriaceous; propodeum rather flat medially and rather remote rugose, medio-longitudinal carina nearly complete, and with slightly protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 608); 1-CU1 slightly oblique, 0.4 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell rather robust (Fig. 608); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 sparsely setose. Hind wing: basal half of marginal cell parallel-sided, apical half linearly widened, its apical width twice width at level of hamuli (Fig. 609); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 31:26; 1r-m 0.7 × 1-M and 1-M oblique. Legs. Tarsal claws with conspicuous and robust dark brown pecten up to apical tooth of claw (Fig. 621); hind coxa dorsally largely smooth and remainder remotely punctate; hind trochantellus robust; length of hind femur and basitarsus 4.3 and 5.0 × their width, respectively; length of inner hind spur 0.45 × hind basitarsus.

Metasoma. First tergite evenly convex medially, 0.9 × longer than wide apically, wider than base of 2nd tergite; 1st and 2nd tergites with medio-longitudinal carina and coarsely irregularly sublongitudinally rugose; medio-basal area of 2nd tergite triangular and rather large (Fig. 612); 2nd suture deep and coarsely crenulate; basal half of 3rd tergite rugulose, remainder of metasoma largely smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 606). Colour. Blackish or dark brown; telotarsi, apical 0.4 of hind tibia, hind tibial spurs and hind tarsus dark brown; remainder of hind tibia and palpi yellowish; remainder of legs, pterostigma and tegulae pale brownish yellow; veins brown; mesoscutum medio-dorsally, scutellum, metanotum, mesopleuron (except partly antero-dorsally), mesosternum and metapleuron orange yellow; wing membrane subhyaline. Variation. Scutellum largely finely punctate, coriaceous medio-posteriorly, but may be striate. Specimens from Sweden are appreciably darker than those from Britain. Vein m-cu of hind wing absent or faintly indicated. Antennal segments: ♀ 51(1), 52(2), 53(2), 54(3), 55(4), 56(6), 57(7), 58(7), 59(2), 60(1), 62(1); ♂ 49(1), 51(1), 53(2), 54(9), 55(6), 56(6), 57(2). Females have on average ca three more antennal segments than males. Males have obtuse hind tibial spurs and the tarsal pecten less developed than in females, propleuron and pronotum yellowish or blackish posteriorly; posterior half of mesoscutum largely yellowish or blackish; apical tergites type 2, somewhat sparse setose, glabrous stripe broad but with some setae directed into it and fringe rather weak (Figs 625, 626). *Austria, British Isles (England, Isle of Man, Ireland), Czech Republic, Finland, Germany, Hungary, *Netherlands, Russia, Sweden. 10.3897/zookeys.919.39642.figures604-6079537E7F1-FFB0-5D78-BF12-CD5E808804DFFigures 604–607. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 604 habitus lateral 605 mummy of Acronicta psi (Linnaeus) in winter 606 ovipositor sheath lateral, 607 mummy of Acronicta psi (Linnaeus) after emergence of parasitoid. https://binary.pensoft.net/fig/39013310.3897/zookeys.919.39642.figures608-621B8F477C8-D2B15F9C-9F6C-875817AB1D36Figures 608–621. Aleiodes pulchripes Wesmael, ♀, England, Chat Moss 608 fore wing 609 hind wing 610 mesosoma

lateral 611 mesosoma dorsal 612 1st–3rd metasomal tergites dorsal 613 fore femur lateral 614 hind femur lateral 615 hind tarsus lateral 616 head anterior 617 head dorsal 618 head lateral 619 base of antenna 620 apex of antenna 621 outer hind tarsal claw. https://binary.pensoft.net/fig/39013410.3897/zookeys.919.39642.figures622-6268F16822F-43435AEB-A126-BEB3F9F6DE2EFigures 622–626. Aleiodes pulchripes Wesmael, ♂, England (ex Acronicta culture) 622 habitus lateral 623 outer hind claw lateral 624 hind tibial spurs and basitarsus lateral 625 metasoma dorsal 626 4th–7th metasomal tergites lateral. https://binary.pensoft.net/fig/390135WesmaelC (1838) Monographie des Braconides de Belgique 4.Nouveaux Mémoires de l’Academie Royale des Sciences et Belles-lettres de Bruxelles11: 1– 166.ČapekMLukásJ (1989) ApocritaParasitica, Ichneumoidea, Braconidae.Acta Faunistica Entomologica Musei Nationalis Pragae19: 27–44.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.O’ConnorJPNashRvan AchterbergC (1999) A catalogue of the Irish Braconidae (Hymenoptera: Ichneumonoidea).Occasional Publication of the Irish Biogeographical Society4: 1– 123.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115– 1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.ShawMR (1979) Rogas pulchripes (Wesmael) (Hymenoptera: Braconidae) and other parasites of arboreal Acronicta species (Lepidoptera: Noctuidae) at Chat Moss, Manchester.Entomologist’s Gazette30: 291–294.

33. XML treatment for Aleiodes quadrum. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.3964233A49D24-B0A2-5511-B3FE383C9E346F4C Aleiodes quadrum (Tobias, 1976)Figs 627–628, 629–641, 642, 643–651 Rogas (Rogas) quadrumTobias, 1976: 83, 221, 1986: 76 (transl.: 125). Aleiodes (Neorhogas) quadrum; Papp 1985a: 162, 1991a: 83. Aleiodes quadrum; Papp 2005: 177. Rogas (Rogas) illustrisPapp, 1977a: 112, 1985a: 162 (as synonym of A. quadrum), 1991a: 83 (id.), 2005: 176 (id.) [examined]. Holotype of A. illustris, ♀ (MTMA), “Yugoslavia, [Croatia:] Kostrena, Rijeka, 12.viii.1966, Uremović”, “Holotypus ♀ Rogas illustris sp. n., Papp, J., 1977”, “Hym. Typ. No. 2378, Mus. Budapest”; paratype, ♀ (MTMA), “[Hungary], Hársbokorhegy, Nagykovacsi”, “1.viii.1952, Bajári”, “Paratypus ♀ Rogas illustris sp. n., Papp, J., 1977”, “Hym. Typ. No. 2380, Mus. Budapest”. 1 ♀ (NMS), “France: Ardèche, Accons, UV light, 24.vi.2013, M.R. Shaw”, “MRS Aleiodes DNA 796”; 1 ♀ (NMS), “France: Savoie, Queige, Le Villaret, 700m., 19.vi. 2019, C.W. Plant”; 1 ♀ (BZL), “Turkey, 15 km W Refahye, W of Erzincan, 1600 m, 7.vii.2000, M. Halada”; 1 ♀ ( BZL), “GRC [= Greece], Westmakadonien, Florina, Aussichtsplatz SE Karies, 40°45'2"N, 21°10'39"E, 1080 m msl, 27.vi.2016, 2016/31, LF, H. u. R. Rausch”; 1 ♀ (MTMA), “[North] Macedonia, Skopje Prov., Mt. Vodno, 16.vii.1997, Gy. Rozner”; 1 ♀ (NMS), “N. Macedonia, Vardar river valley, above Demir Kapiya, N41°22'58", E22°11'45", 244m, 13.vii.2019 S. Beshkov & A. Nahirnic”; 1 ♂ (NMS), “Bulgaria: Haskovo, E. Rhodopes, SW Mezek, 450 m, MV light, 17.vii.2015, C.W. Plant”. MRS796 (France), additionally MRS824 (Bulgaria) likely to be a male of this species. Unknown. Collected in June–July, likely to be univoltine, but there is nothing to suggest how the winter is passed. We have not seen reared material, but the elongate and strongly apically compressed metasoma suggests that the host would be concealed, perhaps between spun leaves, in a leaf sheath, in a seed capsule or in some similar situation. Maximum width of hypoclypeal depression approx. 0.6 × minimum width of face (Fig. 636); OOL of ♀ 0.6–0.7 × diameter of posterior ocellus and rugose; ventral margin of clypeus obtuse apically and clypeus hardly protruding anteriorly (Fig. 638); lobes of mesoscutum densely finely punctate, with interspaces approx. equal to diameter of punctures, smooth and shiny; precoxal area distinctly rugose, but posteriorly only punctate; vein cu-a of fore wing vertical; surroundings of veins M+CU1 and 1-+2-CU1 largely glabrous; vein 1-CU1 of fore wing approx. 0.8 × vein 2-CU1 and as long as m-cu (Fig. 629); surroundings of veins M+CU and 1-M of hind wing largely glabrous; hind tarsal claws with conspicuous dark brown pecten close to apical tooth (Fig. 635); 1st tergite parallel-sided and longer than wide apically (Fig. 632); 2nd tergite of ♀1.0–1.2 × as long as wide basally and black; head black; vein 1-M of fore wing dark brown; wing membrane slightly infuscate. ♀ (NMS) from France (Accons). Length of fore wing 6.9 mm, of body 9.6 mm. Head. Antennal segments of ♀ 53, antenna as long as fore wing, its subapical segments rather robust (Fig. 640); frons with curved striae but medially largely smooth; OOL 0.7 × diameter of posterior ocellus, finely rugose and shiny; vertex coarsely punctate but behind ocelli rugose, rather shiny; clypeus nearly flat and coarsely rugose-punctate; ventral margin of clypeus thick and hardly protruding anteriorly (Fig. 638); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 636); length of eye 1.6 × temple in dorsal view (Fig. 637); clypeus near lower level of eyes; length of malar space 0.2 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely and finely punctate, with interspaces approx. equal to diameter of punctures, smooth and shiny; scutellum finely and densely punctate; precoxal sulcus area of mesopleuron distinctly rugose but posteriorly only punctate, remainder of mesopleuron distinctly but remotely punctate; metapleuron remotely punctate, but ventrally rugose; propodeum evenly convex and coarsely vermiculate-rugose and medio-longitudinal carina nearly complete. Wings. Fore wing: r 0.4 × 3-SR (Fig. 629); 1-CU1 horizontal, 0.8 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 629); cu-a nearly vertical and straight; 1-M curved posteriorly; 1-SR rather slender; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 629); 2-SC+R short longitudinal; m-cu narrowly present; M+CU:1-M = 50:33; 1r-m 0.8 × 1-M.

Legs. Tarsal claws with conspicuous and robust blackish pecten, close to level of apical tooth (Fig. 635); hind coxa largely densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.3 and 5.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, 1.3 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and finely longitudinally rugose, but posterior quarter of 2nd tergite smooth and no median carina; medio-basal area of 2nd tergite triangular and wide (Fig. 632); 2nd tergite as long as wide basally and with shallow transverse impression; 2nd suture shallow and narrowly crenulate; 3rd and subsequent tergites finely punctulate and strongly shiny; apical half of 3rd and 4th tergites without sharp lateral crease; ovipositor sheath widened apically, with mediumsized setae and apically truncate (Fig. 628). Colour. Black; hind tibia dark brown apically and remainder pale yellowish; middle and hind tarsi, and fore telotarsus blackish or dark brown; mandible, remainder of legs, mesoscutum, pronotum postero-dorsally, mesopleuron dorsally, scutellum, metanotum, ovipositor sheath and 1st tergite (except pair of dark patches apically) orange; tegulae yellowish; palpi and pterostigma blackish; vein dark brown, but vein at base of wings yellowish; wing membrane largely slightly infuscate. Variation. OOL of ♀ 0.6–0.7 × diameter of posterior ocellus. The female from Turkey is very similar but has 1st tergite 1.4 × as long as wide apically and 2nd tergite 1.2 × longer than wide basally. Antennal segments: ♀ 53(1), 56(1), 57(1), 58(1). Apical tergites of male type 2 with fringe rather strong (Figs 645, 646). The figured male from Bulgaria has OOL rugose anteriorly, 1st tergite 1.2 × as long as wide posteriorly, 2nd tergite completely parallel-sided and 0.9 × as long as wide, 3rd tergite rugose-striate in anterior half and parallel-sided 0.7 × as long as wide, fore femur 4.5 × longer than wide and hind femur 4.1 × longer than wide. Azerbaijan, *Bulgaria, Croatia, *France, *Greece, Hungary, *North Macedonia, *Turkey. The holotype of A. quadrum is a male and it is less reliable to identify this species from it than from the holotype female of A. illustris; nevertheless, we accept the synonymy proposed by Papp (1985a). The figured male from Bulgaria (NMS; Figs 642–651) is considered to be this species (initially through its CO1 sequence); it is morphologically very similar to A. cruentus and there is a possibility that some similar males have been returned to depositories determined as A. cruentus with no recognition that they might belong to A. quadrum. However, the matter remains unresolved until more males of A. quadrum become available. 10.3897/zookeys.919.39642.figures627-62857583F6D-4E6B-5DCB-9270-AFF7110AC143Figures 627, 628. Aleiodes quadrum (Tobias), ♀, France, Accons 627 habitus lateral 628 ovipositor sheath lateral. https://binary.pensoft.net/fig/39013610.3897/zookeys.919.39642.figures629-64198FD5499-CFC95FEB-9444-C0F118BB3D57Figures 629–641. Aleiodes quadrum (Tobias), ♀, France, Accons 629 wings 630 mesosoma lateral 631 mesosoma dorsal 632 metasoma dorsal 633 fore femur lateral 634 hind femur lateral 635 outer hind tarsal claw 636 head anterior 637 head dorsal 638 head lateral 639 base of antenna 640 apex of antenna 641 antenna. https://binary.pensoft.net/fig/39013710.3897/zookeys.919.39642.figure642511FE234-8F0F-5CF18E5E-35FD5BE1957CFigure 642. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo, habitus lateral. https://binary.pensoft.net/fig/39013810.3897/zookeys.919.39642.figures643-65116E990E9-A70B555D-9335-502C10F3FAF2Figures 643–651. Aleiodes quadrum (Tobias), ♂, Bulgaria, Haskovo 643 wings 644 1st–3rd metasomal tergites dorsal 645 3rd–7th metasomal tergites lateral 646 id. dorsal 647 basal antennal segments 648 head anterior 649 head dorsal 650 head lateral 651 inner hind tarsal claw. https://binary.pensoft.net/fig/390139PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.

34. XML treatment for Aleiodes ruficeps. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642E86D9554-FF63-508C-B1D8901E801532D8 Aleiodes ruficeps (Telenga, 1941)Figs 652–653, 654–666 Rhogas (Rhogas) ruficepsTelenga, 1941: 179, 421, Fig. [examined]. Rogas ruficeps; Shenefelt 1975: 1146; Zaykov 1980b: 87. Rogas (Rogas) ruficeps; Tobias 1976: 81, 1986: 76 (transl.: 122; lectotype designation). Aleiodes ruficeps; Papp 1991a: 88; Fortier and Shaw 1999: 230. Rogas gasteratorauctt. p.p. Lectotype of A. ruficeps, ♀ (ZISP), “[Russia: Crimea,] Eupatoria [= Eupatoriya], Tavrits, gub, 7.v.1907, V.E. Jakovlev”, “Rhogas ruficeps sp. n., Telenga det.”, “Lectotype Rogas ruficeps Tl., design. Tobias, 1980”. Bulgaria, Russia, Turkey, [Armenia, Iran]. Specimens in ZJUH, BZL, NMS, RMNH, ZISP, ZSSM. None. Unknown. Specimens collected in April-May; presumably univoltine, but there is nothing to suggest how it overwinters. We have not seen reared material. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 661); OOL ca twice diameter of posterior ocellus and moderately punctate (Fig. 662); 4th–10th antennal segments approx.as long as wide (Figs 652, 664); ventral margin of clypeus thick to rather sharp and distinctly protruding in lateral view (Fig. 663); mesoscutal lobes punctate and interspaces largely smooth and shiny, lobes rather convex; scutellum sparsely punctate; precoxal sulcus coarsely vermiculaterugose; marginal cell of fore wing of ♀ ending rather removed from wing apex (Fig. 654); length of vein r of fore wing 0.3 × vein 3-SR (Fig. 654); vein 1-CU1 of fore wing 0.3–0.5 × vein 2-CU1 (Fig. 654); hind tarsal claws rather slender, hardly curved and only brownish setose (Fig. 666); head completely or largely orange or yellowish; palp dark brown basally; hind femur apico-dorsally dark brown or black; hind tibia pale yellowish but apically darkened; 4th and 5th tergites black; wing membrane nearly entirely infuscate. Resembles A. grassator because of the robust antennal segments and dark wings, but A. ruficeps has frons, OOL, vertex, malar space, and third tergite less sculptured, apex of hind tibia and palpi dark brown, basal antennal segments of ♀ somewhat less robust, hypoclypeal depression wider, marginal cell of fore wing slenderer and vein r of fore wing shorter. Differs from the similar A. ruficornis by

having hypoclypeal depression wider, clypeus wider and lower, apical antennal segments of ♀ slenderer, OOL less sculptured and more antennal segments (♀: 45–47 vs 35–39(–41) of A. ruficornis). Lectotype, ♀, length of fore wing 7.3 mm, of body 8.6 mm. Head. Antennal segments of ♀ 45, length of antenna 1.1 × fore wing, its subapical segments moderately robust (Fig. 665); frons with few rugae, remainder smooth; OOL 2.0 × diameter of posterior ocellus, and moderately densely punctate; vertex spaced punctate and shiny; clypeus punctate; ventral margin of clypeus thick and distinctly protruding forwards (Fig. 663); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 661); length of eye 1.1 × temple in dorsal view (Fig. 662), temples subparallel-sized behind eyes; vertex behind stemmaticum punctate; clypeus distinctly below lower level of eyes; occipital carina widely reduced ventrally (Fig. 663); length of malar space 0.7 × length of eye in lateral view. Mesosoma. Mesoscutal lobes punctate and interspaces smooth, shiny; precoxal area of mesopleuron coarsely vermiculate-rugose, near precoxal area mesopleuron distinctly punctate; scutellum rather flattened, sparsely punctate; propodeum coarsely vermiculate-rugose, medio-longitudinal carina incomplete, absent posteriorly and propodeum rounded laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 654); 1-CU1 horizontal, 0.3 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell rather long (Fig. 654); cu-a slightly inclivous, straight; 1-M slightly curved posteriorly; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell linearly widened, its apical width 2.5 × width at level of hamuli (Fig. 655); 2-SC+R longitudinal; m-cu pigmented only basally; M+CU:1-M = 7:5; 1r-m 0.7 × 1-M. Legs. Tarsal claws rather slender, hardly curved and with six brownish bristles (Fig. 666); hind coxa rather weakly punctate; hind trochantellus robust; hind femur distinctly punctate; fore femur 3.3 × longer than wide; length of hind femur and basitarsus 3.2 and 5.2 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 0.9 × as long as wide apically; 1st and 2nd tergites with medio-longitudinal carina and regularly longitudinally rugose; medio-basal area of 2nd tergite triangular and short (Fig. 658); 2nd suture rather deep and finely crenulate; basal half of 3rd tergite finely striate, remainder of metasoma smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 653).

Colour. Orange brown; antenna, palpi, apices of femora, telotarsi, bases of fore and middle coxae, pterostigma, and veins (but 1-SR much paler than 1-M) dark brown; mesosoma (except for mesoscutum and scutellum), ovipositor sheath, 3rd tergite (except basally) and subsequent tergites black or blackish; apex of hind tibia only narrowly dark brown; wing membrane nearly entirely infuscate. Variation. Vein 1-CU1 of fore wing 0.3–0.5 × vein 2-CU1. Females may have the flagellum extensively pale basally, or entirely dark. Antennal segments: ♀ 45(1), 46(1), 47(1); ♂ 56(1), 58(1). Males appear to have ca ten more antennal segments than females. Male has apical tergites type 1, setae rather long and sparse, and fringe not observed. *Armenia, Bulgaria, *Iran, Russia, *Turkey. 10.3897/zookeys.919.39642.figures652-653DE32596F-2DB1-5EB0-B66C-444B6902D4D9Figures 652, 653. Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 652 habitus lateral 653 ovipositor sheath lateral. https://binary.pensoft.net/fig/39014010.3897/zookeys.919.39642.figures654-6662A1C084A-F6ED55F6-8EE1-6A74CA3E3D94Figures 654–666.

Aleiodes ruficeps (Telenga), ♀, Turkey, Konya 654 fore wing 655 hind wing 656 mesosoma lateral 657 mesosoma dorsal 658 1st –3rd metasomal tergites dorsal 659 fore femur lateral 660 hind femur lateral 661 head anterior 662 head dorsal 663 head lateral 664 base of antenna 665 apex of antenna 666 inner hind tarsal claw. https://binary.pensoft.net/fig/390141ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980b) New species of the family Braconidae (Hymenoptera) for the Bulgarian fauna.Acta Zoologica Bulgarica16: 86–89. [in Bulgarian with English summary]TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512– 588. Lebanon, U.S.A.]PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1– 122.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.

35. XML treatment for Aleiodes ruficornis. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642F561F7BF-1CC5-5CBC-B33CE3BA444A17B9 Aleiodes ruficornis (Herrich-Schäffer, 1838)Figs 667–669, 670–681, 682–685 Rogas ruficornisHerrich-Schäffer, 1838: 156, fig.; Shenefelt 1975: 1224 (as synonym of A. dimidiatus) [neotype designated below]. Aleiodes (Neorhogas) ruficornis; Papp 1985a: 152 (as synonym of A. dimidiatus), 1991a: 90 (id.). Aleiodes (Chelonorhogas) ruficornis; Belokobylskij et al. 2003: 398; van Achterberg 2014: 209; Abdolalizadeh et al. 2017: 37. Aleiodes ruficornis; Bergamasco et al. 1995: 5; Zaldívar-Riverón et al. 2004: 234; Papp 2005: 176 (as synonym of A. dimidiatus). Aleiodes brevicornisWesmael, 1838: 98; Shenefelt 1975: 1224; Papp 1985a: 152, 157 (as synonym of A. dimidiatus), 2005: 176 (id.); Belokobylskij et al. 2003 (as synonym of A. ruficornis) [examined]. Aleiodes (Neorhogas) brevicornis; Papp 1991a: 90 (as synonym of A. dimidiatus). Aleiodes nigripalpisWesmael, 1838: 97; Shenefelt 1975: 1224; Papp 1985a: 152, 157 (lectotype designation; as synonym of A. dimidiatus), 1991a: 90, 2005: 176 (id.); Belokobylskij et al. 2003 (as synonym of A. ruficornis) [examined]. Rhogas dimidiatus ab. nigrobasalisHellén, 1927: 24 (invalid name). Rhogas dimidiatus ab. ruficollisHellén, 1927: 24 (invalid name). Rhogas carbonarius ab. giraudiFahringer, 1931: 236 [unavailable name for melanistic males]. Rhogas carbonarius var. giraudiTelenga, 1941: 168. Syn. nov. Rogas dimidiatus ab. infuscatusHellén, 1957: 49 (invalid name). Rogas dimidiatus ab. nigripesHellén, 1957: 49 (invalid name). Aleiodes (Aleiodes) arnoldii; Farahani et al. 2015: 232–233 (but see note under A. arnoldii). Rogas gasteratorauctt. p.p. Rogas dimidiatusauctt. p.p. Neotype of A. ruficornis here designated, ♀ (NMS), “[Germany,] Einbeck, L. 31.v.[19]85, [R. Hinz]”, “ex: Hoplodrina blanda Schiff. (Lep.)”. Holotype of A. brevicornis, ♀ ( KBIN), “A. brevicornis ♀ mihi. 3.”, “Coll. Wesmael”, “A. brevicornis mihi, dét. C. Wesmael”, “Belgique, Liège, leg. Robert/ teste Papp J., 1983”, “Holotypus Aleiodes brevicornis Wesm., 1838 ♀, det. Papp, 1983”, “Aleiodes dimidiatus Spin. ♀, det. Papp J., 1983”. Holotype of A. nigripalpis, ♂ (KBIN), “A. nigripalpis ♂ mihi. 2.”, “Coll. Wesmael”, “A. nigripalpis mihi, dét. C. Wesmael”, “Belgique, Liège/ teste Papp J., 1983”, “Lectotypus Aleiodes nigripalpis Wesm., 1838 ♂, det. Papp, 1983”, “Aleiodes dimidiatus Spin. ♂, det. Papp J., 1983”. The neotype designation for A. ruficornis is necessary for nomenclatural stability, because the types of Braconidae described by Herrich-Schäffer are lost (Horn and Kahle 1935–37; the first author could not find any specimen in ZMB) and the species has been confused with similar species in the past. The specimen is selected because it fits well the original description, the probable type location was in Germany, it has been reared and it is in good condition. Andorra, Austria, British Isles (England (V.C. 5, 11, 13, 15, 17, 19, 20, 22, 23, 24, 28, 29, 31, 33, 38, 39, 60, 63); Wales (V.C. 52) [no specimens seen from Scotland]), Bulgaria, Croatia, Czech Republic, Finland, France, Germany, Greece, Hungary, Italy (including Sicily), Moldova, Montenegro, Netherlands (DR: Borger; LI: Venlo, ZH: Oostkapelle), North Macedonia, Norway, Romania, Russia (including Far East), Serbia, Slovakia, Sweden, Switzerland, Turkey, Ukraine, [Afghanistan, Dagestan, Iran, Kazakhstan, Kyrgyzstan]. Specimens in ALC, ZJUH, BZL, FMNH, MMUM, MRC, MSC, MSNV, MTMA, NMS, NRS, OUM, RMNH, SDEI, UNS, UWIM, ZISP, ZSSM. This is a widespread species, generally common, but partly replaced by A. gasterator in the Mediterranean region. MRS140 (UK), MRS877 (Sweden), MRS888 (UK), MRS890 (UK), MRS891 (UK). Specimens collected from April–September; probably plurivoltine in the southern part of its range but in Britain univoltine, flying from June–August with a varied means of overwintering (see below). Reared from the noctuids Agrotis clavis (Hufnagel) (4 [4 ZISP]/ Russia), Agrotis segetum (Dennis & Schiffermüller) (1 [FMNH]/Finland), Agrotis sp. (3), Euxoa nigricans (Linnaeus) (1 [FMNH]/Finland), Euxoa sp. (3:1 [3 FMNH/Finland], Hoplodrina blanda (Dennis & Schiffermüller) (4:1, Germany; R. Hinz), Hoplodrina octogenaria (Goeze) (1; W.A. Watson), Mythimna impura (Hübner) (1 [ZSSM]/Germany; E. Haeselbarth). A further mummy from H. blanda failed to emerge (M.R. Shaw), but was no doubt of this species. The above hosts belong to three different subfamilies of Noctuidae, all feeding and resting close to ground level. In addition, we have seen a specimen labelled as ex the nymphalid Brenthis ino (Rottemburg) but accompanied by a mummy of a noctuid, probably Hoplodrina sp. (det. M.R. Shaw). In experiments a range of hosts recorded for this species in the literature (several arctiine and lymantriine Erebidae and the lasiocampid Lasiocampa quercus (Linnaeus)) that are actually hosts of superficially similar species such as A. alternator (Nees) were offered to the female reared ex H. octogenaria but, as expected, they were firmly rejected. However, this female readily accepted late 1st instar larvae of Agrotis exclamationis (Linnaeus), from which adult progeny resulted very smoothly (1:16\13\\12\12+0. The few failures to oviposit were almost certainly due to egg depletion). Searching in the vicinity of hosts included antennal drumming (the tips curled downwards) and indeed the antennae seemed to be the only proximal means of locating and assessing the host. Once the host was found it was immediately accepted, rapidly jabbed and stood over or often withdrawn from (1.0–1.5 cm) while the venom took affect (20–40 secs), then relocated via antennal searching (when it had been withdrawn from this might take up to a minute, but it was always eventually successful) scooped in with the fore legs (the antennae only slightly involved), positioned and held between the mid legs for the duration of oviposition (20–30 secs). Frequently the host larva was kicked free of the ovipositor by the parasitoid’s hind leg(s) and the parasitoid rapidly left without any period of post-oviposition association. Recovery from the venom was rather protracted (up to 20 mins), during which time hosts were rejected if rediscovered. Towards the end of successful oviposition runs it was evident that venom depletion ran ahead of egg depletion, resulting in erratic (but nevertheless successful) oviposition sequences. No host feeding took place. In this experimental series oviposition took place in mid-July with mummification at the end of August and adult emergence in late May of the following year. However, although the winter was passed in the mummy in this entire series (and probably also the case for the other, natural, Agrotis hosts), it is clear that the rearings from Hoplodrina and probably also Mythimna involved overwintering in the host larva with adult emergence in the year of mummification. This host-related difference in overwintering is not inconsistent with univoltinism in Britain (where the experiments and other observations were done) but it is certainly an interesting quirk of its host range and might be of significance in suggesting one way in which a temporal isolating mechanism could potentially arise as a forerunner to speciation (cf. Shaw, 2003). The mummy is formed in the soil and is not strongly (if at all) glued to the substrate. It is predominantly dark brown in colour, very large in relation to the size of the insect that will emerge and, although basically cylindrical, somewhat flattened in appearance owing to a pronounced but blunt lateral keel (Fig. 669). It is more or less strongly contracted at the anterior end, markedly less so caudally, and the copiously silken lining typically occupies 3rd–8th abdominal segments. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 676); OOL of ♀ 1.4–1.6 × as long as diameter of posterior ocellus and distinctly rugose or rugulose (Fig. 677); length of 4th antennal segment of ♀ 1.0–1.4 (of ♂ 1.1–1.4) × its width (Fig. 679); ventral margin of clypeus thick and not protruding anteriorly (Fig. 678); lobes of mesoscutum densely punctate, interspaces largely smooth and shiny; precoxal area coarsely vermiculate-rugose medially; marginal cell of fore wing of ♀ usually ending rather removed from wing apex (Fig. 670); vein 1-CU1 of fore wing 0.4– 0.6 × as long as vein 2-CU1; hind tarsal claws yellowish or brownish bristly setose and with few yellowish pectinal teeth (Fig. 681); hind femur at least apico-dorsally dark brown or black; inner side of hind tibia of ♀ yellowish; pale males have whole frons and stemmaticum yellowish; palpi dark brown or blackish, rarely brown; 3rd metasomal tergite only antero-laterally reddish or yellowish; 4th and 5th tergites black. Specimens from high altitude have the head conspicuously long setose and the tarsal claws brownish pectinate basally. In this respect males are similar to A. hirtus, but A. hirtus has precoxal area and mesoscutum largely smooth and clypeus distinctly protruding in lateral view.

Neotype, ♀, length of fore wing 3.9 mm, of body 6.5 mm. Head. Antennal segments of ♀ 35, antenna as long as fore wing, its subbasal and subapical segments robust (Figs 679, 680); frons with curved rugae; OOL 1.2 × diameter of posterior ocellus, rugose and moderately shiny; vertex rugose, rather shiny; clypeus rugose; ventral margin of clypeus thick and not protruding forwards (Fig. 678); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 676); length of eye twice temple in dorsal view (Fig. 677); vertex behind stemmaticum rugose; clypeus below lower level of eyes; length of malar space 0.5 × length of eye in lateral view; temple punctate and shiny, but rugulose near occipital carina. Mesosoma. Mesoscutal lobes distinctly punctate, interspaces of lateral lobes smooth and shiny; precoxal area of mesopleuron coarsely vermiculate-rugose medially, but posteriorly punctate; mesopleuron punctate medially; metapleuron distinctly rugose ventrally and dorsally punctate; scutellum largely smooth (except for punctulation), rather shiny and nearly flat, with lateral carina; propodeum coarsely reticulate-rugose, laterally dorsal face longer than posterior one, somewhat angulate laterally but without tubercles, and with complete medio-longitudinal carina. Wings. Fore wing: r 0.3 × 3-SR; marginal cell short (Fig. 670); 1-CU1 horizontal, 0.5 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 670); cu-a inclivous, straight; 1-M rather curved posteriorly; 1-SR slightly wider than 1-M; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 670); 2-SC+R subquadrate; m-cu narrowly pigmented; M+CU:1-M = 30:19; 1r-m 0.6 × 1-M. Legs. Tarsal claws mainly setose and medially with 4 yellowish rather short pectinal teeth (Fig. 681); hind coxa punctate and shiny; hind trochantellus robust; length of hind femur and basitarsus 3.3 and 4.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather convex medially, 0.9 × longer than wide apically, robust and coarsely irregularly longitudinally rugose as 2nd tergite; 1st tergite and basal half of 2nd tergite with mediolongitudinal carina; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 673); 2nd suture deep and crenulate; basal half of 3rd tergite largely longitudinally striate, remainder of metasoma superficially micro-sculptured or smooth; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with medium-sized setae and apically truncate (Fig. 668). Colour. Reddish or orange-brown; stemmaticum medially, malar space largely, temple and occiput ventrally, mesosternum, mesopleuron ventrally, 3rd tergite (except antero-laterally), 4th–7th tergites black; palpi (only labial palp darkened basally) brown; basal half of antenna, tegulae, parastigma, and base of pterostigma pale yellowish; apical half of antenna, pedicellus dorsally, propodeum dorsally and medially, middle and hind femora apico-dorsally, and telotarsi dark brown; remainder of pterostigma and veins dark brown or brown (Fig. 670); fore wing membrane rather infuscate, but hind wing nearly subhyaline. Variation. Female: mesosoma occasionally wholly black. Male face and mesosoma usually black but can be variably marked with red; scape and pedicel usually (partly) reddish in central and southern populations but most often entirely black in more north-western ones (e.g., British Isles, Sweden); hind coxa varies from black to red. Length of malar space 0.5–0.6 × length of eye in lateral view; head black or largely reddish brown (except temple ventrally and malar space); interspaces of mesoscutal lobes smooth to micro-sculptured; 1-CU1 0.4–0.6 × 2-CU1; 3rd tergite longitudinally striate or rugulose basally (sometimes narrowly so), without curved sculptural elements (Fig. 673), except sometimes some weak transverse striae occasionally present at extreme apex; males from montane habitats are generally darker than lowland males. Antennal segments: ♀ 34(6), 35(12), 36(15), 37(22), 38(17), 39(13), 40(2), 41(3), 42(2), 43(1); ♂ 43(1), 44(1), 45(1), 46(1), 47(5), 48(13), 49(32), 50(19), 51(36), 52(36), 53(28), 54(11), 55(4), 56(4), 57(2). On average males have ca 14 more antennal segments than females. Male has marginal cell of fore wing less robust than in ♀, with apical tergites type 1–2, density of setae rather variable and fringe evident but sparse (Fig. 682). *Afghanistan, *Andorra, Austria, British Isles (England, Wales), *Bulgaria, *Croatia, *Czech Republic, *Finland, *France, Germany, *Iran, *Kazakhstan, *Kyrgyzstan, *Montenegro, *Netherlands, *North Macedonia, *Norway, *Romania, *Russia (including Dagestan and Far East), *Serbia, *Slovakia, *Sweden, Switzerland, *Turkey, *Ukraine. An examined female (NMS) from Hungary, Borzsony Mts., 140 m altitude, 20–30.vii.2005 (unfortunately, too damaged for description) represents a very similar but new species. The 4th–10th antennal segments are not moniliform, slenderer than in typical A. ruficornis, the fore femur is more robust than in A. ruficornis, and the COI sequence (MRS886) is different (2.1 %). Aleiodes ruficornis is the commonest and most widespread of a small group of related species parasitising grassland and “cutworm” hosts, exhibiting strong sexual dimorphism with unremarkable males but the more extensively orange females having a stronger build and much shorter antennae. The least extreme in these respects is A. gasterator, which largely (but not completely) replaces A. ruficornis in the Mediterranean region. Aleiodes grassator is similar to A. ruficornis, but it appears to be restricted to montane and northern habitats where it might be thought to replace A. ruficornis. However, some males that morphologically agree best with A. ruficornis have been collected at high altitude in the Alps (up to 2550 m), where A. improvisus also occurs, but whether these high-altitude A. ruficornis males are parts of breeding populations or have simply been carried up in thermals is impossible to say. The females in this group (excluding A. gasterator) are scarcer in collections than males, as they fly very little and rarely enter Malaise traps. 10.3897/zookeys.919.39642.figures667-6696F715591-2129-5BE5-925F-6BD56577E9D1Figures 667–669. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 667 habitus lateral 668 ovipositor sheath lateral 669 mummy of Hoplodrina blanda (Denis & Schiffermüller). https://binary.pensoft.net/fig/39014210.3897/zookeys.919.39642.figures670-68159011668-ED535A41-A4D5-911A3B72DECEFigures 670–681. Aleiodes ruficornis (Herrich-Schäffer), ♀, neotype 670 wings 671 mesosoma lateral 672 mesosoma dorsal 673 metasoma dorsal 674 fore femur lateral 675 hind femur lateral 676 head anterior 677 head dorsal 678 head lateral 679 base of antenna 680 apex of antenna 681 outer hind tarsal claw. https://binary.pensoft.net/fig/39014310.3897/zookeys.919.39642.figures682-685D1D0931E-B1F45F94-987C-FF7F38A7FA7AFigures 682–685. Aleiodes ruficornis (Herrich-Schäffer), ♂, U.K. (culture) 682 habitus lateral 683 apex of antenna 684 base of antenna 685 mummy of Hoplodrina octogenaria (Goeze). https://binary.pensoft.net/fig/390144ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341– 435.van AchterbergC (2014) Notes on the checklist of Braconidae (Hymenoptera) from Switzerland.Mitteilungen der Schweizerischen Entomologischen Gesellschaft87: 191– 213.AbdolalizadehFMadjdzadehSMFarahaniSAskari HesniM (2017) A survey of braconid wasps (Hymenoptera: Braconidae: Euphorinae, Homolobinae, Macrocentrinae, Rogadinae) in Kerman province, southeastern Iran.Journal of Insect Biodiversity and Systematics3(1): 33– 40.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xPappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227– 250. https://doi.org/10.11646/zootaxa.3973.2.2

36. XML treatment for Aleiodes rufipes. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642AEFA9612-4F10-5BCC-8892DBC764BE5C09 Aleiodes rufipes (Thomson, 1892)Figs 686–687, 688–700 Rogas rufipesThomson, 1892: 1669; Shenefelt 1975: 1224; Kotenko 1992: 96 [examined]. Rogas (Rogas) rufipes; Tobias 1986: 81 (transl.: 133). Aleiodes (Neorhogas) rufipes; Papp 1985a: 162, 1987b: 36, 1991a: 88; Belokobylskij 1996: 14; Riedel et al. 2002: 106. Aleiodes (Chelonorhogas) rufipes; Chen et al. 1992: 496; Belokobylskij 2000: 40; Chen and He 1997: 42; He et al. 2000: 665; Ghahari et al. 2011: 4; Farahani et al. 2015: 229, 244. Aleiodes rufipes; Fortier and Shaw 1999: 228; Papp 2002: 562, 2005: 177; Aydogdu and Beyarslan 2005: 191, 192. Holotype, ♀ (ZIL), “Lap”, “rufipes m”, “Sverige [= Sweden], Lappland, teste Papp J., 1983”, “Holotypus”, “Rogas rufipes Thoms., 1891, ♀, Papp, 1983”. Finland, Norway, Sweden. Specimens in FMNH, NMS, MTMA, RMNH, ZIL. MRS294 (Sweden), MRS312 (Sweden), MRS314 (Sweden), MRS673 (Finland), MRS674 (Finland), MRS676 (Finland), MRS680 (Finland). Unknown. Collected from July–August; presumably univoltine. We have not seen reared material and there is no indication of how the winter may be passed. Maximum width of hypoclypeal depression approx. 0.4 × minimum width of face (Fig. 695); OOL of ♀ 1.1–1.5 × as long as diameter of posterior ocellus and punctate-rugulose to coriaceous-rugose (Fig. 696); length of antenna of ♀ 1.0–1.1 × length of fore wing; ventral margin of clypeus thin and not protruding in lateral view (Fig. 697); mesoscutal lobes remotely punctulate and with satin sheen; area of precoxal sulcus smooth; length of vein 1-CU1 of fore wing 0.2–0.3 × vein 2-CU1 and 0.4 × vein m-cu; vein 2-SC+R of hind wing subquadrate; tarsal claws with robust apical tooth and with medium-sized yellowish brown pecten (Fig. 698); hind femur and basitarsus slender (Fig. 686); 1st metasomal tergite comparatively steep anteriorly (Fig. 686); head (largely) black; apex of hind femur usually largely black dorsally; basal half of hind tibia (largely) pale yellowish; 2nd tergite yellowish or reddish; males usually with dense and long setosity on at least basal half of 4th–6th tergites. Redescribed ♀ (RMNH) from Finland (Enntekiö). Length of fore wing 5.3 mm, of body 5.8 mm. Head. Antennal segments of ♀ 51, antenna as long as fore wing, its subapical segments rather robust and apical segment with short spine (Fig. 700); frons largely smooth, except for some microsculpture; OOL 1.2 × diameter of posterior ocellus, coriaceous-rugose and slightly shiny, groove beside posterior ocellus rather shallow and crenulate; vertex coriaceous with some rugulae, rather dull; clypeus rugose; ventral margin of clypeus thin and not protruding forwards (Fig. 697); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 695); length of eye 1.7 × temple in dorsal view (Fig. 696); vertex behind stemmaticum coriaceous-rugulose; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes largely punctate-coriaceous, with satin sheen; precoxal area of mesopleuron partly remotely punctate as its surroundings; medio-longitudinal carina of metanotum distinct posteriorly; scutellum punctate; propodeum convex and coarsely rugose, medio-longitudinal carina absent posteriorly, and without protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 688); 1-CU1 slightly oblique, 0.3 × 2-CU1; r-m 0.5 × 3-SR; 2nd submarginal cell long (Fig. 688); cu-a slightly inclivous, straight but posteriorly slightly curved; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 densely setose. Hind wing: marginal cell linearly widened, its apical width 1.7 × width at level of hamuli (Fig. 689); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 26:25; 1r-m 0.6 × 1-M. Legs. Tarsal claws with rather conspicuous and medium-sized brownish pecten (Fig. 698); hind coxa densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.8 and 6.4 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite convex and basally rather steep, 0.9 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and longitudinally rugose; maximum width of 2nd tergite 1.6 × its median length; medio-basal area of 2nd tergite medium-sized triangular and rather short (Fig. 692); 2nd suture deep and finely crenulate; basal half of 3rd tergite rugulose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 687). Colour. Black; mesoscutum largely, legs, and 1st–3rd metasomal tergites (but 3rd tergite narrowly

infuscate posteriorly) reddish brown; tegulae brownish yellow; ovipositor sheath, palpi, pterostigma and veins dark brown; telotarsi partly infuscate; wing membrane subhyaline. Variation. Legs usually largely reddish, but telotarsi, apices of hind femur and tibia frequently dark brown and sometimes most of hind tibia and apical half of hind femur black; clypeus blunt to rather acute ventrally; depression near posterior ocelli smooth or finely crenulate; mesoscutum of ♀ usually partly reddish brown, but sometimes largely or entirely black; ventral third of mesopleuron regularly and finely punctate. Antennal segments: ♀ 57(2), 59(1); ♂ 57(1), 59(1), 60(5), 61(1), 62(1). Male is very similar with mesoscutum black (rarely partly reddish) and at least 2nd tergite orange brown (sometimes with pair of dark brown patches), apical tergites type 1–2 with fringe rather strong when visible, and also often evident on tergites following the third. Finland, Norway, Sweden. 10.3897/zookeys.919.39642.figures686-687171BD0B1-855A-54F7-AF64-48254BEB2FB0Figures 686, 687. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 686 habitus lateral 687 ovipositor sheath lateral. https://binary.pensoft.net/fig/39014510.3897/zookeys.919.39642.figures688-700ED43702D-6573513A-AD98-E335A908BD3EFigures 688–700. Aleiodes rufipes (Thomson), ♀, Sweden, Lillav 688 fore wing 689 hind wing 690 mesosoma lateral 691 mesosoma dorsal 692 1st–3rd metasomal tergites dorsal 693 fore femur lateral 694 hind femur lateral 695 head anterior 696 head dorsal 697 head lateral 698 outer hind tarsal claw 699 base of antenna 700 apex of antenna. https://binary.pensoft.net/fig/390147ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.KotenkoAG (1992) A contribution to the fauna of Braconidae (Hymenoptera) of Dauria. In: Amirkhanov AM (Ed.) Insects of Dauria and adjacent territories: 94–107.Izdatelstvo Tsentralnoi Nauchno issledovatelskoi laboratorii okhotnichego khozyaistva i zapovednikov, Moskva, 141 pp.TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143–164.PappJ (1987b) First outline of the braconid fauna of Southern Transdanubia, Hungary (Hymenoptera, Braconidae), 3. Helconinae, Homolobinae, Macrocentrinae, Blacinae, Doryctinae and Rogadinae. A Janus Pannonius Muzeum Evkonyve 30–31 (1985–1986): 33–36.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1–12.RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97–108.ChenX-XHeJHMaY (1992) New records of Aleiodes Wesmael from China 2 (Hymenoptera: Braconidae: Rogadinae).Acta Zootaxonomica Sinica17(4): 495–496. [in Chinese with English summary]BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.ChenXXHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.GhahariHFischerMPappJ (2011) A study on the Braconidae (Hymenoptera: Ichneumonoidea) from Ilam province, Iran.Calodema160: 1–5.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–237.PappJ (2002) The Braconid wasps (Hymenoptera: Braconidae) of the Ferto-Hansag National Park (NW Hungary). In: Mahunka S (Ed.) The Fauna of the Ferto-Hansag National Park, Hungarian Natural History Museum, Budapest, 557– 581.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.AydogduMBeyarslanA (2005) The first records of Aleiodes Wesmael 1838 (Hymenoptera: Braconidae: Rogadinae). The fauna of Thrace region of Turkey. Linzer biologische Beiträge 37/1: 185–193.

37. XML treatment for Aleiodes rugulosus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642BC048A17-A771-52A8-AE01FD716951E65F Aleiodes rugulosus (Nees, 1811)Figs 701–703, 704–716, 717, 718, 719, 720–727 Bracon rugulosusNees, 1811: 32; Papp 1985a: 162 (neotype designation). Rogas rugulosus; Shenefelt 1975: 1247–1248. Rogas (Rogas) rugulosus; Tobias 1976: 84, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) rugulosus; Papp 1987b: 36, 1991a: 79; Riedel et al. 2002: 106. Aleiodes (Chelonorhogas) rugulosus; Belokobylskij et al. 2003: 398. Aleiodes rugulosus; Bergamasco et al. 1995: 5; Zaldívar-Riverón et al. 2004: 234; Papp 2005: 177. Rhogas rugulosus var. pictusKokujev, 1898: 296; Shenefelt 1975: 1247–1248 (not Herrich-Schäffer 1838). Neotype, ♀ (KBIN), “A. rugulosus”, “dét. C. Wesmael”, “Coll. Wesmael”, “Belgique, Bruxelles”/ teste Papp J., 1983”, “Neotypus, Bracon rugulosus Nees, 1812 [sic!], ♀, Papp 1983”, “Aleiodes rugulosus Ns. ♀, det. Papp J., 1983”. Albania, Austria, Belgium, Bulgaria, Czech Republic, British Isles (England: V.C.s 1, 3, 4, 11, 17, 25, 27, 29, 69; Wales: V.C.s 41, 48; Scotland: V.C.s 78, 81, 88, 98; Ireland: V.C. H30), Finland, France, Germany, Hungary, Moldova, Netherlands (DR: Borger; GE: ‘t Harde; Heerde; Otterlo, NB: Kampina; NH: Crailo; OV: Buurserzand), North Macedonia, Norway, Poland, Romania, Russia, Slovakia, Spain, Sweden. Specimens in ALC, ZJUH, BZL, HHC, IKC, MMUM, MRC, MSC, MSNV, MTMA, NMS, NRS, OUM, RMNH, SDEI, UWIM, ZSSM. MRS191 (Hungary), MRS217 (UK), MRS398 (France), MRS884 (Poland). Collected in (May)July–August(September), usually in open habitats such as mosses, heaths, herbrich grasslands and fens. Partly plurivoltine, at least in the southern part of its range, but largely univoltine in the north (in a UK culture only one female out of 20 reared, and two males out of 69, emerged in the same year as mummy formation). Reared from Noctuidae, Acronictinae: Acronicta auricoma (Denis & Schiffermüller) (7 [1 BZL, 1 MSC, 1 ZMUO]; J. Voogd/Netherlands, M & J. Schwarz/Austria, Finland), Acronicta euphorbiae (Dennis & Schiffermüller)/cinerea (Hufnagel) (5 [2 IKC, 1 BZL, 1 FMNH]; M.J. Pellinen, D. & J. Steedan), Acronicta menyanthidis (11; R.P. KnillJones, W.A. Watson), Acronicta rumicis (Linnaeus) (5 [1 ZJUH, 1 IKC, 1 MTMA]; R.J. Heckford, M.J. Pellinen, M.R. Shaw), Oxicestra geographica (Fabricius) (17 [12 BZL, 5 MTMA]; Hungary), Acronicta sp. on low plants (3), Simyra albovenosa (Goeze) (14 [5 ZJUH, 4 FMNH, 2 UMZC, 1 NRS]; M.R. Shaw); A. Lozan, Romania), Acronicta euphorbiae (Dennis & Schiffermüller) on Euphorbia sanguinea Hochst (5 [2 RMNH, 3 ALC]). These species all feed on low plants. The mummy is moderately arched, very strongly glued down (usually to a narrow twig or stem low in the vegetation), and persists through the winter. The pupation chamber, occupying approximately abdominal segments 3–7 of the host, is rather densely lined with silk which is laid down after the mummy has hardened suggesting that the larva within can turn easily. Rearing experiments, undertaken using stock originally reared from A. menyanthidis, suggested that this host and A. rumicis were equally suitable, but most experiments were not conducted in a way to provide clear data in this respect. The behaviour of the adult females towards these hosts indicated some adaptation to use of highly aggregated species (i.e., that lay large batches of eggs) as, firstly there was a habituation process whereby repeated contacts with hosts generally preceded oviposition, and secondly there was only weak displacement following oviposition (resulting in rather frequent superparasitism). The antennae were used to locate hosts with wide sweeping motions, and usually the host curled up and was manipulated backwards against the hind tarsi before the ovipositor was inserted and the egg was laid. Generally, there was no pre-oviposition sting and post-oviposition association with the only slightly subdued host was minimal, but the oviposition process was variable and occasionally there was a brief jab, but no subsequent waiting period, before oviposition. Less enthusiasm for sub-active hosts, such as those oviposited into a few seconds or minutes earlier, provided a short-lived impediment to super-parasitism, although sometimes two (on one occasion four, confirmed by dissection) eggs were laid into a single host in separate consecutive bouts without the parasitoid really relinquishing the host. First instar hosts were less easy than 2nd or 3rd instars for the parasitoid to deal with, and although oviposition into 2nd instar hosts was somewhat more successful than into 3rd instars, occasionally successful oviposition into early 4th instar hosts occurred. Mean development times from oviposition to mummification in different instar hosts (A. rumicis) under the same ambient conditions (Reading, S. England, July) varied as might be expected given that mummification was always at essentially the same (penultimate instar) stage of the host’s larval life: for 1st (N = 23), 2nd (N = 40) and 3rd (N = 7) instars, 27.0, 25.5 and 20.7 days,

respectively. There is no venom effect to influence successful host development. Opportunities to offer other hosts were limited but it was clear that, although oviposition into larvae of the closely related arboreal species Subacronicta megacephala (Dennis & Schiffermüller) was fairly readily obtained (N = 15), though slightly inhibited by the host’s adherence to its silken pad rather than curling up, the parasitoid was always encapsulated (as a 1st instar larva in observed cases) and no progeny resulted. No rearings of A. rugulosus from arboreal Acronictinae have been seen, although these conspicuous larvae are often collected and reared. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 711); OOL approx. equal to diameter of posterior ocellus and coarsely punctate (Fig. 712); vertex flattened behind ocelli; ventral margin of clypeus thick and not protruding in lateral view (Fig. 713); mesoscutal lobes coriaceous; mesopleuron (including precoxal sulcus area) nearly or completely smooth; propodeum with pair of crest-like protuberances laterally; vein 1-CU1 of fore wing much shorter than vein 2-CU1; basal half of marginal cell of hind wing parallel-sided and subapically widened (Fig. 704); tarsal claws with large dark brown pecten up to apical tooth of claw (Fig. 716); hind spurs (dark) reddish brown; hind tibial spurs of ♂ acute apically (Fig. 722); head black; dorsal 0.4 of mesopleuron, mesosternum and scutellum black; metasoma entirely black (typical) or 1st and 2nd tergites orange or yellowish brown; apex of hind femur yellowish or reddish; basal half of hind tibia pale yellowish. Redescribed ♀ (RMNH) from Netherlands (Buurserzand). Length of fore wing 6.3 mm, of body 7.7 mm. Head. Antennal segments of ♀ 65, length of antenna 1.3 × fore wing, its subapical segments rather slender (Fig. 716); frons largely smooth except few striae; OOL equal to diameter of posterior ocellus, coarsely punctate and shiny; vertex coarsely punctate with some rugae, shiny; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 713); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 711); length of eye twice temple in dorsal view (Fig. 712); vertex behind stemmaticum flattened and punctate-rugose; clypeus between eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate with very finely granulate interspaces, rather matt; precoxal area of mesopleuron smooth, mesopleuron densely punctate posteriorly; scutellum coarsely punctate and rather flat; propodeum rather flattened medially, very coarsely reticulate-rugose, medio-longitudinal carina nearly complete, and with small crest-like protuberances laterally. Wings. Fore wing: r 0.6 × 3-SR (Fig. 704); 1-CU1 horizontal, 0.4 × 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell short (Fig. 704); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: basal 0.6 of marginal cell subparallel-sided and remainder linearly widened, its apical width 1.8 × width at level of hamuli (Fig. 704); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 37:28; 1r-m 0.8 × 1-M. Legs. Tarsal claws with conspicuous and robust blackish pecten (Fig. 716); hind coxa largely densely and coarsely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 3.9 and 5.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite moderately convex, 0.9 × longer than wide apically; 1st and 2nd tergites with coarse medio-longitudinal carina and very coarsely and irregularly longitudinally rugose; mediobasal area of 2nd tergite large and distinct (Fig. 707); 2nd suture deep, rather wide and coarsely crenulate; basal half of 3rd tergite punctate-rugose, remainder of metasoma finely punctate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically narrowed and rounded (Fig. 702). Colour. Black (including fore and middle telotarsi, apical half of hind tibia and hind tarsus); basal half of hind tibia pale yellowish; palpi (but basally somewhat infuscate) and remainder of legs reddish brown; tegulae yellowish brown; hind tibial spurs and pterostigma dark brown; veins brown; mesopleuron with broad dark reddish longitudinal band; wing membrane slightly infuscate. Variation. Micro-sculpture of lateral lobes of mesoscutum very finely granulate or absent and resulting in a largely smooth surface; maximum width of marginal cell of hind wing 1.5–2.3 × its width near hamuli; body entirely black (both sexes) or 1st and 2nd metasomal tergites and mesosoma partly dark reddish (both sexes, but 2nd and 3rd tergites of males more or less darkened); hind tibial spurs dark brown or reddish brown. Antennal segments: ♀ 60(4), 61(7), 62(10), 63(18), 64(14), 65(2), 66(8), 67(4), 68(3), 69(1); ♂ 53(1), 56(2), 57(1), 58(7), 59(6), 60(15), 61(15), 62(19), 63(28), 64(12), 65(10), 66(4), 67(2), 68(1). Females have on average ca one more antennal segment than males. Male is similar to the dark female form, with acute hind tibial spurs (Fig. 722), 3rd tergite convex in lateral view (Fig. 720), with dense and long setosity and apical tergites type 1 and no

fringe observed (Figs 721, 724). Aleiodes rugulosus is a very colour-variable species; the 1st and 2nd metasomal tergites are quite frequently mostly or entirely dark red or orange brown (Fig. 701) in populations in which entirely black females (Fig. 717) also often occur. The variation is not geographical, since most populations definitely have both forms. In rearing experiments, a strong genetic basis for this feature became evident: from a virgin female with completely black metasoma, all five males reared were black, and a cross between one of them and a (wild-reared) black female produced eleven female progeny, all black. In contrast, a lineage from a pairing between a weakly red male and a red female comprised three red males and eight red females, separate individuals of which produced a single red and a single black male as progeny. When sufficient material from single sites is available it is usual to see a clear predominance of one form or the other. Sometimes the lower part of mesopleuron is pale (often looking unpigmented there, but narrowly), and in extreme reddish specimens the scutellum, much of the mesopleuron (but usually the mesosternum remains darkish), the metanotum dorsally, the metapleuron in part and much of the propodeum are also reddish. Albania, Austria, Belgium, Bulgaria, Czech Republic, British Isles (England, Wales, Scotland, Ireland), Finland, France, Germany, Hungary, *Netherlands, *North Macedonia, *Moldova. Norway, Poland, *Romania, Russia, *Slovakia, *Spain, Sweden. 10.3897/zookeys.919.39642.figures701-703199CB048-CA90-5EB9-92BE-3B3ED43EB59CFigures 701–703. Aleiodes rugulosus (Nees), ♀, U.K., culture 701 habitus lateral 702 ovipositor sheath lateral 703 mummy of Acronicta rumicis (Linnaeus). https://binary.pensoft.net/fig/39014810.3897/zookeys.919.39642.figures704-7167D86EEFB-350A5B10-AC56-D894121A288BFigures 704–716. Aleiodes rugulosus (Nees), ♀, U.K., culture 704 wings 705 mesosoma lateral 706 mesosoma dorsal 707 propodeum and 1st–3rd metasomal tergites dorsal 708 fore femur lateral 709 hind femur lateral 710 antenna 711 head anterior 712 head dorsal 713 head lateral 714 base of antenna 715 apex of antenna 716 outer hind tarsal claw. https://binary.pensoft.net/fig/39014910.3897/zookeys.919.39642.figures717-718FE6550A4-E2F6548E-950E-461004EA85F4Figures 717, 718. Aleiodes rugulosus (Nees), dark form, ♀, U.K., Meathop Moss 717 habitus lateral 718 mummy of Acronicta menyanthidis (Esper). https://binary.pensoft.net/fig/39015010.3897/zookeys.919.39642.figure719CA2CA434-9154-5F36B825-181DBFE6D317Figure 719.

Aleiodes rugulosus (Nees), Netherlands, Ede, mummy of Acronicta auricoma (Denis & Schiffermüller). Photograph: J. Voogd. https://binary.pensoft.net/fig/39015110.3897/zookeys.919.39642.figures720-727235DC654-7A6E55B8-94B2-5EA701FC2C96Figures 720–727. Aleiodes rugulosus (Nees), ♂, Hungary (Halastó) 720 habitus lateral 721 metasoma lateral 722 hind tibial spurs and basitarsus lateral 723 base of antenna 724 metasoma dorsal 725 fore femur lateral 726 head dorsal 727 hind femur lateral. https://binary.pensoft.net/fig/390152PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72– 85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1987b) First outline of the braconid fauna of Southern Transdanubia, Hungary (Hymenoptera, Braconidae), 3. Helconinae, Homolobinae, Macrocentrinae, Blacinae, Doryctinae and Rogadinae. A Janus Pannonius Muzeum Evkonyve 30–31 (1985–1986): 33– 36.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97– 108.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341– 435.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.ZaldívarRiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.0300-3256.2004.00144.xPappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.Herrich-SchäfferGAW (1838) Faunae Insectorum Germaniae initiae oder Deutschlands Insecten. Hymenoptera. Heft 124 & 156. Regensburg.PappJ (1983) A survey of the Braconid fauna of the Hortobágy National Park (Hymenoptera, Braconidae), II: 315–337. In: Mahunka S (Ed.) The Fauna of the Hortobágy National Park II. Budapest.

38. XML treatment for Aleiodes schewyrewi. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642D9F506D0-8EC9-55FD-89A7EA36438530AA Aleiodes schewyrewi (Kokujev, 1898)Figs 728–729, 730–742 Rhogas (Rhogas) schewyrewiKokujev, 1898: 304. Rhogas (Rhogas) schevyrevi[sic!]; Telenga 1941: 186. Rogas schewyrewi; Shenefelt 1975: 1248 Rogas (Rogas) schewyrewi; Papp 1977b: 116. Aleiodes (Chelonorhogas) schewyrewi; Belokobylskij 2000: 46. Rhogas (Rhogas) schewyrewi var. zaydamensisKokujev, 1898: 305; Telenga 1941: 186 [examined]. Rogas schewyrewi var. zaydamensis; Shenefelt 1975: 1248–1249. Rogas (Aleiodes) schewyrewi var. zaydamensis; Papp 1977b: 116, 117. Holotype of A. schewyrewi zaydamensis, ♀ (ZISP), “[Mongolia], Kerijsk Kr., Ruio Zaydam, Przewalski”, “1910a”, [illegible handwritten label], “Rh. Schewyrewi Kokw. var. zaydamensis Kokw., No.1910a”, “♀ Rhogas schewyrewi var. zaydamensis Kok., C. van Achterberg, 1992, holotype”. Holotype of A. s. schewyrewi not found, according to the original description with same label data and with a larger part of the body blackish. 1 ♀ (BZL), “S. Russia, [Volgograd obl.], Elton Lake env., 20.v.2001, J. Miatleuski”; 1 ♂ (MTMA), “Mongolia, Gobi Altay aimak, Mongol els, 10 km SO von Somon Chechmort, 1600 m, Exp. Dr. Z. Kaszab, 1966”, “Nr. 684, 13.vii.1966”, “Rogas schewyrewi Kok., det. Papp J., 1977/ compared with ♀ det. Kokujev, Papp, 1983, 57”. None. Unknown. A female collected in May and a male in July may suggest that it is plurivoltine (or, less probably, that the female overwinters as an adult). Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 737); OOL of ♀ approx. as long as diameter of posterior ocellus and densely rugose (Fig. 738); head in anterior view rather robust (Fig. 737); clypeus distinctly protruding anteriorly in lateral view, thick apically and with long setae on medium-sized anterior part (Fig. 739); lobes of mesoscutum largely superficially punctate, interspaces finely granulate or smooth and with satin sheen; precoxal area densely rugose, but posterior third only finely punctate; vein 1-CU1 of fore wing 0.3 × vein 2-CU1 and 0.5 × vein mcu (Fig. 730); hind tarsal claws long and slender, nearly straight and only brownish bristly setose (Fig. 740); tarsal segments (except telotarsus) with long apical spiny bristles (Fig. 728); basal half of hind tibia pale yellowish, contrasting with dark brown colour of basal half of hind femur. Holotype, ♀, length of fore wing 5.8 mm, of body 7.5 mm. Head. Antennal segments of ♀ 48, length of antenna 1.1 × fore wing, its subapical segments moderately slender (Fig. 742); frons largely with fine curved rugae; OOL equal to diameter of posterior ocellus, and densely rugose; vertex superficially rugose-punctate, rather shiny; clypeus convex and densely punctate; ventral margin of clypeus thick and protruding forwards (Fig. 740); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 737); length of eye 1.3 × temple in dorsal view (Fig. 738); vertex behind stemmaticum finely rugose-punctate and with long setae; clypeus largely above lower level of eyes; length of malar space 0.3 × height of eye in lateral view. Mesosoma. Mesoscutal lobes largely superficially punctate, interspaces finely granulate and with satin sheen; precoxal area of mesopleuron largely smooth medially, densely punctate anteriorly and posteriorly densely rugose, but posterior 0.3 only finely punctate; metapleuron spaced coarsely punctate; metanotum with fine and nearly complete median carina; scutellum sparsely punctate,

shiny; propodeum rather convex and rather coarsely rugose, medio-longitudinal carina on anterior 0.4 of propodeum. Wings. Fore wing: r curved and 0.4 × 3-SR (Fig. 730); 1-CU1 horizontal and slightly widened, 0.3 × 2-CU1; r-m 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 730); cu-a slightly inclivous, straight; 1-M slightly curved; 1-SR slender; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: basal third of marginal cell subparallel-sided and remainder linearly widened; 2-SC+R short and longitudinal; m-cu slightly indicated; M+CU:1-M = 10:7; 1r-m 0.6 × 1-M. Legs. Tarsal claws long and slender, nearly straight and only brownish bristly setose (Fig. 740); tarsal segments (except telotarsus) with long apical spiny bristles (Figs 728, 740); hind coxa largely punctate, but dorsally punctate-rugose; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.6 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex and strongly widened posteriorly, 0.9 × longer than wide apically; 1st and 2nd tergites with weak medio-longitudinal carina (absent posteriorly) and finely longitudinally rugose, but 2nd tergite smooth medio-posteriorly; medio-basal area of 2nd tergite wide triangular and rather distinct (Fig. 734); 2nd suture deep and narrow; basal half of 3rd tergite aciculate, remainder of metasoma smooth and shiny; 2nd and 3rd tergites with sharp lateral crease; ovipositor sheath moderately widened, with medium-sized setae and apically truncate (Fig. 729). Colour. Yellowish brown (including basal half of antenna); apical half of antenna, frons largely, stemmaticum, occiput dorso-laterally, pronotal side medially, axilla, mesopleuron (except anterodorsally), mesosternum, metapleuron, propodeum, hind femur and apical third of hind tibia, 5th–7th tergites, last two posterior sternites, ovipositor sheath and pterostigma dark brown or blackish brown; veins brown; wing membrane subhyaline. Variation. Length of 4th hind tarsal segment 1.8–2.0 × longer than wide; malar space and temple ventrally largely dark brown or yellowish brown. Antennal segments: ♀ 45(1); ♂ 54(1); according to original description ♀ type has 58 segments. Male has clypeus yellowish and contrasting with black face, apical tergites type 1, and no fringe observed. *Iran, Mongolia, *Russia (European part). 10.3897/zookeys.919.39642.figures728-729604DBFE4-7024-5A1A-AA0D-6FC00C223E2BFigures 728, 729. Aleiodes schewyrewi (Kokujev) , ♀, holotype var. zaydamensis (Kokujev) 728 habitus lateral 729 ovipositor sheath lateral. https://binary.pensoft.net/fig/39015310.3897/zookeys.919.39642.figures730-7421B6BBA14-89C855F2-96AF-779159DFD13EFigures 730–742. Aleiodes schewyrewi (Kokujev) , ♀, holotype var. zaydamensis (Kokujev) 730 fore wing 731 hind wing 732 mesosoma lateral 733 mesosoma dorsal 734 metasoma dorsal 735 fore femur lateral 736 hind femur lateral 737 head anterior 738 head dorsal 739 head lateral 740 outer hind tarsal claw 741 base of antenna 742 apex of antenna. https://binary.pensoft.net/fig/390154TelengaNA (1941) Family Braconidae, subfamily Braconinae (continuation) and Sigalphinae. Fauna USSR.Hymenoptera5(3): 1–466.ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115– 1262.PappJ (1977b) Braconidae (Hymenoptera) from Mongolia, VI.Folia Entomologica Hungarica30(1): 109–118.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.

39. XML treatment for Aleiodes schirjajewi. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642ACE6E3C7-C81F-5A63-AE5AC438590A3E56 Aleiodes schirjajewi (Kokujev, 1898)Figs 743, 744–757 Rhogas reticulator var. schirjajewiKokujev, 1898: 299 [examined]. Rogas schirjajewi; Shenefelt 1975: 1249. Rogas (Rogas) schirjaevi[sic!]; Tobias 1976: 85. Rogas (Rogas) schirjaewi[sic!]; Tobias 1986: 80 (transl.: 132). Aleiodes (Neorhogas) schirjajewi; Papp 1991a: 71, 2002: 562. Aleiodes (Chelonorhogas) schirjajewi; Samartsev and Belokobylskij 2013: 766. Aleiodes schirjajewi; Shaw et al. 1998: 63; Papp 2005: 177. Holotype, ♂ (ZISP), “[Kazakhstan], Kemropavl., Akmolin, 908a”, “K. Kokujeva”, “908a, Rh. reticulator Nees v. schirjajewi Kokw.”, “Holotypus”. Bulgaria, Hungary, Italy, Moldova, Russia, Serbia, Ukraine [Dagestan, Kazakhstan]. Specimens in BZL, ZJUH, MRC, MSC, MTMA, NMS, RMNH, SDEI, ZISP. None. Unknown. Specimens collected throughout April–September, presumably plurivoltine. We have not seen reared material and it is unclear how the winter is passed. Diagnosis. Maximum width of hypoclypeal depression 0.3–0.4 × minimum width of face (Fig. 751); length of antenna of ♀ 1.1–1.4 × fore wing; ventral margin of clypeus thick and obtuse apically and clypeus not protruding in lateral view (Fig. 753); vertex and frons with strong striae or rugae; mesoscutum, metapleuron and scutellum normally shiny and without dense granulation, at most with some superficial micro-sculpture; precoxal area of mesopleuron smooth; vein 2-CU1 of fore wing approx. as long as vein 1-CU1 or shorter (Fig. 744); vein M+CU of hind wing distinctly longer than vein 1-M (Fig. 745); hind tarsal claws with medium-sized dark brown pecten (Fig. 756); head black; mesoscutum and scutellum orange brown; fore and middle femora distinctly black or dark brown apically; basal half of hind tibia dark brown; anterior half of mesosoma, 1st and 2nd metasomal tergites yellowish or orange brown; at least basal half of 4th–6th tergites of ♂ with long and dense setosity. Holotype, ♂, length of fore wing 5.0 mm, of body 5.8 mm. Head. Antennal segments of ♂ 50, length of antenna 1.3 × fore wing, its subapical segments rather robust; 4th segment of maxillary palp slender and cylindrical; frons with rather coarse curved rugae and interspaces smooth; OOL 1.1 × diameter of posterior ocellus, coarsely rugose and shiny; coarsely transversely rugose and shiny; clypeus punctate-rugulose; ventral margin of clypeus thick and not protruding forwards (Fig. 753); width of hypoclypeal depression 0.4 × minimum width of face (Fig. 751); length of eye 3.2 × temple in dorsal view (Fig. 752); vertex behind stemmaticum coarsely rugose; clypeus below lower level of eyes; occipital carina complete; length of malar space

0.4 × length of eye in lateral view. Mesosoma. Mesoscutal lobes finely punctate, shiny, interspaces micro-sculptured; precoxal area of mesopleuron smooth medially except for some crenulations and punctures, its surroundings smooth; scutellum moderately punctate; propodeum rather convex and coarsely reticulate, coarse mediolongitudinal carina present anteriorly, carinae not protruding laterally. Wings. Fore wing: r 0.7 × 3-SR (Fig. 744); 1-CU1 horizontal, as long as 2-CU1; r-m 0.9 × 3-SR; 2nd submarginal cell short (Fig. 744); cu-a vertical, largely straight; 1-M nearly straight posteriorly; 1SR slender; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: marginal cell linearly widened, its apical width 2.2 × width at level of hamuli (Fig. 745); 2-SC+R subquadrate; m-cu absent; M+CU:1-M = 15:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws with medium-sized dark brown pecten (Fig. 756); hind coxa largely densely punctate; hind trochantellus medium-sized; length of hind femur and basitarsus 4.0 and 7.0 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite moderately flattened, 0.9 × longer than wide apically; 1st and 2nd tergites with medio-longitudinal carina and densely vermiculate-rugose; medio-basal area of 2nd tergite wide triangular and distinct (Fig. 748); 2nd suture deep and moderately crenulate; 3rd tergite finely striate basally, remainder of metasoma largely smooth; 4th without sharp lateral crease; basal half of 4th– 6th tergites of ♂ with long and dense setosity. Colour. Orange brownish; basal half of antenna, palpi largely and parastigma (except base) yellowish brown; head, mesosternum (except anteriorly), mesopleuron (except anteriorly and anterodorsally), apical 0.4 of hind femur, 3rd tergite (except antero-laterally) and subsequent tergites black; scapus, pedicellus basally, apical half of antenna, apex of fore and middle femora, apex of middle and hind tibiae, hind basitarsus, 2nd hind tarsal segment apically, telotarsi, pterostigma, parastigma basally and veins dark brown; wing membrane subhyaline. Variation. Vein 1-CU1 of fore wing 1.0–2.3 × vein 2-CU1; maximum width of marginal cell of hind wing 1.6–2.6 × its width near hamuli (Fig. 745). Antennal segments: ♀ 46(1), 48(3), 49(1), 50(2), 51(1); ♂ 47(1), 48(2), 49(2), 50(1), 51(1). The sexes have comparable numbers of antennal segments. Apical tergites of ♂ type 4, dense, making the tergites look concave and fringe not observed. Female is very similar to the redescribed male; ovipositor sheath wide, with long setae and apically truncate. *Bulgaria, Hungary, *Italy, Kazakhstan, Moldova, Russia (including Dagestan and Far East), Serbia, Ukraine. 10.3897/zookeys.919.39642.figure7434BD9BC03-28D8-549A-82D8-E91877305D8AFigure 743. Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, habitus lateral. https://binary.pensoft.net/fig/39015510.3897/zookeys.919.39642.figures744-757247A8626-09A05325-8C7F-C86C2780979BFigures 744–757.

Aleiodes schirjajewi (Kokujev), ♀, Hungary, Budapest, but 756 ♀, Ukraine 744 fore wing 745 hind wing 746 mesosoma lateral 747 mesosoma dorsal 748 metasoma dorsal 749 fore femur lateral 750 hind femur lateral 751 head anterior 752 head dorsal 753 head lateral 754 base of antenna 755 apex of antenna 756 outer hind tarsal claw 757 antenna. https://binary.pensoft.net/fig/390156ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1991a) Parasitic wasps of the superfamily

Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.PappJ (2002) The Braconid wasps (Hymenoptera: Braconidae) of the FertoHansag National Park (NW Hungary). In: Mahunka S (Ed.) The Fauna of the Ferto-Hansag National Park, Hungarian Natural History Museum, Budapest, 557–581.SamartsevKGBelokobylskijSA (2013) On the fauna of the true cyclostome braconid wasps (Hymenoptera, Braconidae) of Astrakhan Province. Entomologicheskoe Obozrenie 92(2): 319–341.Entomological Review93(6): 755– 774.ShawSRMarshPMFortierJC (1998) Revision of North American Aleiodes Wesmael (Part 2): the apicalis (Brullé) species-group in the New World (Hymenoptera: Braconidae, Rogadinae) Journal of Hymenoptera Research 7: 62–73.PappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.

40. XML treatment for Aleiodes sibiricus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642CDFDDC7E-C46C-5695-881ED661D6998B63 Aleiodes sibiricus (Kokujev, 1903)Figs 758–759, 760–762, 763–776 Rhogas sibiricusKokujev, 1903: 286 [examined]. Rogas sibiricus; Shenefelt 1975: 1250. Rogas (Rogas) sibiricus; Tobias 1976: 83, 84, 1986: 76, 78 (transl.: 124, 128; lectotype designation). Aleiodes (Neorhogas) sibiricus; Papp 1985a: 150, 153, 162, 1991a: 92; Belokobylskij 1996: 15. Aleiodes sibiricus; Papp, 2005: 177. Rhogas hungaricusSzépligeti, 1906: 616; Papp 1985a: 150, 153, 162 (as synonym of A. sibiricus; lectotype designation), 2005: 177 (id.); 2004: 216 (id.) [examined]. Rogas hungaricus; Shenefelt 1975: 133. Rhogas reinhardiFahringer, 1931: 221 (description in key only), 1932: 275 (full description; as R. rheinhardi); Papp 1985a: 153, 162 (as synonym of A. sibiricus; holotype examined), 2005: 177 (id.). Paralectotype of A. sibiricus, ♀, (ZJUH), “[Russia], Irkutsk, v., I. Jakovlev”, “K. Kokujeva”, “Paratypus Rogas sibiricus Kokujev”, “Rec. in exchange [from] Academy of Science, Leningrad, BM.1963.211”, “Ant. 69”. Lectotype of A. hungaricus, ♀, (MTMA), “[Hungary], Budapest, Szépligeti”, “Lectotypus Rhogas hungaricus Szépl. 1906, ♀, Papp, 1966”, “Hym. Typ. No. 401, Mus. Budapest”, “Aleiodes sibiricus Kok., ♀, det. Papp J., 1983/compared with ♀ paralectotype”. Albania, Austria, Bulgaria, France, Germany, Greece, Hungary, Italy, North Macedonia, Sweden, Turkey. Specimens in ZJUH, BZL, MTMA, NMS, RMNH, ZSSM. MRS310 (Sweden), MRS313 (Sweden), MRS805 (France). Collected in April and May, and presumably univoltine, but 2 ♀ from Sweden: Ångermanland, Lillavammasjon were collected apparently in July in window traps set on the trunks of Betula and Picea. We have examined four males collected in April which suggests spring emergence from the mummy rather than overwintering as an adult. This is corroborated by the data with the single reared specimen examined (MTMA), from the noctuid Noctua comes Hübner collected 7.iv.1961 and emerging on 3.iv.1962 (Germany; [R.] Hinz). The rearer was widely experienced with caterpillars, and the host determination is unlikely to be wrong (the other caterpillar species with which it might conceivably be confused all have similar biology and phenology in any case). This host initiates its overwintering as a small larva, feeding in mild weather through the winter and normally being wellgrown by April, by then in its penultimate or final instar. The rearing is of great interest because it shows that A. sibiricus, like A. fortipes (q. v.), not only parasitises a host that has overwintered as a larva, but also must habitually attack late instar hosts. The reared specimen is accompanied by a stout mummy, large but not unduly so for the size of the adult that emerged, lacking its anterior portion to leave a partitioned chamber comprising abdominal segments 4 onwards, which is well lined with silk and would presumably normally form in the soil (Fig. 762). Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Figs 759, 770); antenna of ♀ with 65–72 segments and 5th–10th segments wider than long; anterior part of clypeus short and transverse, its height 0.2–0.3 × height of hypoclypeal depression (Fig. 770); ventral margin of clypeus rather thin and slightly protruding in lateral view (Fig. 772); mesoscutal lobes densely punctate and interspaces smooth; precoxal area (rather) coarsely vermiculate-rugose medially; length of vein r of fore wing 0.3–0.5 × vein 3-SR (Fig. 763); vein 1-CU1 horizontal and 0.2–0.3 × vein 2CU1; hind tarsus and claws slender and claws with inconspicuous brownish teeth (Fig. 775); 4th and 5th metasomal tergites more or less yellowish to reddish brown; head, mesoscutum, scutellum, mesopleuron and apex of metasoma black. Paralectotype of A. sibiricus, ♀, length of fore wing 9.1 mm, of body 10.0 mm. Head. Antennal segments of ♀ 69, antenna as long as fore wing, its subapical segments mediumsized; frons smooth; OOL 1.1 × diameter of posterior ocellus, and finely coriaceous-rugulose; vertex rugulose and rather dull; clypeus coriaceous and strongly transverse (4–6 × wider than high; Figs 759, 770); ventral margin of clypeus rather thin and slightly protruding forwards (Fig. 772); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 770); length of eye 1.1 × temple in dorsal view (Fig. 771); vertex behind stemmaticum rugulose; clypeus near lower level of eyes; length of malar space 0.3 × length of eye in lateral view. Mesosoma. Mesoscutal lobes densely punctate and interspaces smooth, rather matt; precoxal area of mesopleuron rugose medially and anteriorly, its surroundings moderately punctate; scutellum sparsely punctate and no lateral carina; propodeum rather convex and densely and finely rugose, medio-longitudinal carina complete and no protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 763); 1-CU1 horizontal, 0.2 × 2-CU1; r-m unsclerotized, 0.7 × 3-SR; 2nd submarginal cell rather short (Fig. 763); cu-a inclivous, straight; 1-M nearly straight

posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 setose. Hind wing: basal half of marginal cell slightly widened, but apical half distinctly linearly widened, its apical width 2.5 × width at level of hamuli (Fig. 764); 2-SC+R subquadrate; m-cu slightly indicated; M+CU:1-M = 51:38; 1r-m 0.7 × 1-M. Legs. Tarsal claws with four inconspicuous brownish pecten-teeth (Fig. 775); hind coxa punctulate; hind trochantellus robust; length of hind femur and basitarsus 4.1 and 6.0 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite rather flattened, 0.8 × longer than wide apically; 1st and 2nd tergites with weak medio-longitudinal carina and densely finely rugose, but posterior quarter of 2nd tergite irregularly rugose and no median carina; medio-basal area of 2nd tergite wide and short (Fig. 767); 2nd suture deep, rather wide medially and finely crenulate; basal half of 3rd tergite finely rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 761). Colour. Black; palpi and tegulae pale yellowish; legs (except black coxae; apex of hind femur dorsally, inner side of hind tibia apically (not outer side!) and telotarsi infuscated), apex of first tergite, 2nd–5th tergites and metasoma ventrally, yellowish brown; ovipositor sheath largely, pterostigma and most veins dark brown; vein 1-R1 of fore wing yellowish brown; wing membrane subhyaline. Variation. Face, clypeus, mesoscutum, propleuron, upper part of mesopleuron, and first tergite partly, or rarely entirely, reddish brown. Usually in males and rarely in females mesoscutum wholly black; vein r of fore wing 0.3–0.5 × vein 3-SR; clypeus flattened and subparallel-sided or convex and ventrally concave; pterostigma medially dark brown or yellowish brown. Antennal segments ♀: 65(4), 66(8), 67(6), 68(4), 69(2), 70(5), 71(2), 72(2); ♂ 66(1), 71(1). Male apical tergites of type 1 and fringe not observed. *Albania, *Austria, *Bulgaria, *France, Germany, *Greece, Hungary, Italy (main), *North Macedonia, Russia (Siberia), *Sweden, *Turkey. The holotype of A. reinhardi (Fahringer, 1931) from Bolzano (N Italy) was examined by Papp (1985a) and directly compared with the lectotype of A. hungaricus. Unfortunately, the holotype could not be located in NHMW, but there is no obvious reason not to follow the synonymy with A. sibiricus (Kokujev) proposed by Papp (1985a). Aleiodes agilis (Telenga, 1941) from China, Iran, and Caucasus is very similar to A. sibiricus, but A. agilis has antenna of ♀ with ca 48 segments (69–72 segments in A. sibiricus), 2nd tergite narrowly smooth posteriorly (finely sculptured), pronotum largely yellow (black), clypeus not protruding in lateral view (somewhat protruding) and is often smaller (body length 7–8 mm vs 7–11 mm). 10.3897/zookeys.919.39642.figures758-759590006CE-F27D-5D9E-965F-E61357DFF390Figures 758, 759. Aleiodes sibiricus (Kokujev), ♀, lectotype 758 habitus lateral 759 head anterior. Photographs: K. Samartsev. https://binary.pensoft.net/fig/39015710.3897/zookeys.919.39642.figures760-762C0C37212-54365166-931C-FA09E12DE0EAFigures 760–762. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves, but 762 Germany, Freiburg 760 habitus lateral 761 ovipositor sheath lateral 762 mummy of Noctua comes Hübner. https://binary.pensoft.net/fig/39015810.3897/zookeys.919.39642.figures763-776C50BCA38-1D445B02-911D-506AD23D5856Figures 763–776. Aleiodes sibiricus (Kokujev), ♀, Hungary, Heves 763 fore wing 764 hind wing 765 mesosoma lateral 766 mesosoma dorsal 767 metasoma dorsal 768 fore femur lateral 769 hind femur lateral 770 head anterior 771 head dorsal 772 head lateral 773 base of antenna 774 apex of antenna 775 outer hind tarsal claw 776 antenna. https://binary.pensoft.net/fig/390159ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1–12.

41. XML treatment for Aleiodes turcicus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396423DCD7753-2BA1-58CB-8965223FBD52EE35 Aleiodes turcicus http://zoobank.org/885C0189-5A7B-4D35-B7D382F1094633AA van Achterberg & Shawsp. nov.Figs 777–778, 779–791 Holotype, ♀ (NMS), “Turkey: Sivas, v.2001, D.L.J. Quicke”, “MRS Aleiodes DNA 126 [one middle leg]”. Paratype: 1 ♂ (RMNH), “Turkey, Hakkâri, [20 km S Siirt, 500 m, 23.vi.1985], C.J. Zwakhals”. MRS126 (Turkey). Unknown. The material examined was collected in the period May–June. It is not clear how many generations occur, or how the winter is passed. Maximum width of hypoclypeal depression 0.4–0.5 × minimum width of face (Fig. 786); OOL approx. 2.3 × diameter of posterior ocellus and sparsely punctate (Fig. 787); ventral margin of clypeus obtuse and not protruding in lateral view (Fig. 788); length of antenna of ♀ 1.1–1.2 à — fore wing; lateral lobes of mesoscutum largely smooth; precoxal area coarsely vermiculate-rugose medially; vein 1-CU1 of fore wing approx. 0.4 × as long as vein 2-CU1; hind tarsal claws yellowish or brownish setose (Fig. 791); head and part of mesosoma black; palpi, pterostigma and apical 0.2–0.3 of hind tibia of ♀ blackish; wing membrane distinctly infuscate. Holotype, ♀, length of fore wing 5.3 mm, of body 7.9 mm. Head. Antennal segments of ♀ 47, length of antenna 1.15 × fore wing, length of 4th segment 1.1 × its width, and its subapical segments 1.2 × as long as wide (Figs 789, 790); frons with regular curved rugae, shiny, and rugose behind antennal sockets; OOL 2.3 × diameter of posterior ocellus, and area mostly finely remotely punctate, interspaces much larger than diameter of punctures; vertex spaced punctate laterally, densely punctate and with transverse rugae medially, shiny; clypeus medium-sized, coarsely and densely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 788); width of hypoclypeal depression 0.5 × minimum width of face (Fig. 786); length of eye 1.3 × temple in dorsal view (Fig. 787); vertex behind stemmaticum sparsely punctate; clypeus near lower level of eyes; length of malar space 0.5 × length of eye in lateral view; eyes medium-sized, elliptical (Fig. 788). Mesosoma. Mesoscutal lobes smooth between rather remote punctures, strongly shiny, more densely punctate on middle lobe; notauli distinct but shallow, especially posterior half; mesoscutum short setose, widely and strongly rugose medio-posteriorly; scutellum slightly convex, remotely punctate and evenly rounded laterally, no carina; prepectal carina strong, reaching anterior border; precoxal area coarsely vermiculate rugose anteriorly and medially, posteriorly absent; mesopleuron above precoxal area (except speculum) remotely punctate, shiny, and antero-dorsally coarsely vermiculate-

rugose; metapleuron densely rugose, but dorsally punctate, interspaces approx. equal to diameter of punctures; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete, but irregular, no tubercles. Wings. Fore wing: r 0.35 × 3-SR (Fig. 779); 1-CU1 horizontal, 0.4 × as long as 2-CU1; r-m 0.9 × 2-SR, and 0.7 × 3-SR; 2nd submarginal cell medium-sized (Fig. 779); cu-a slightly oblique, approx. parallel with CU1b, straight; 1-M rather curved posteriorly. Hind wing: marginal cell gradually and evenly widened, its apical width 1.9 × width at level of hamuli (Fig. 780); 2-SC+R subquadrate; m-cu distinct, shorter than cu-a. Legs. Tarsal claws subpectinate, with four brown medium-sized pectinal bristles and some finer ones basally (Fig. 791); hind coxa moderately coarsely punctate, with several long oblique rugae, shiny; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 4.9 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus; hind tibia slender (Fig. 777). Metasoma. First tergite rather flattened; 1st and 2nd tergites coarsely and densely rugose, robust, with distinct median carina; medio-basal area of 2nd tergite wide and short (Fig. 783); 2nd suture deep medially and shallow laterally; basal 0.4 of 3rd tergite finely striate, remainder of metasoma largely smooth, strongly shiny, punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath rather wide, with long setae and apically rounded (Fig. 778). Colour. Black; palpi, base of middle coxa, apical 0.2 (dorsally)–0.3 (inner side) of hind tibia, apex of hind femur and telotarsi blackish; remainder of tarsi more or less darkened and base of hind tibia dark brown; basal seven segments of antenna (remainder more or less dark brown), 1st and 2nd tergites and antero-lateral corners of 3rd tergite, and remainder of legs orange brown; humeral plate pale yellowish but partly darkened; pterostigma blackish; veins dark brown; wing membrane blackish infuscate. Variation. Apical metasomal tergites of ♂ type 2; inner hind tibial spur 0.50 × hind basitarsus; mesopleuron, metapleuron and propodeum may be largely yellowish. Turkey. 10.3897/zookeys.919.39642.figures777-7788ED3635B-E3CF-5FF0-AA73-BF600FA0FBFEFigures 777, 778. Aleiodes turcicus sp. nov., ♀, holotype 777 habitus lateral 778 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016010.3897/zookeys.919.39642.figures779-791D9C94A3B-2D755D96-A9CE-74C2BD6EED69Figures 779–791. Aleiodes turcicus sp. nov., ♀, holotype 779 fore wing 780 hind wing 781 mesosoma lateral 782 mesosoma dorsal 783 metasoma dorsal 784 fore femur lateral 785 hind femur lateral 786 head anterior 787 head dorsal 788 head lateral 789 base of antenna 790 apex of antenna 791 outer hind tarsal claw. https://binary.pensoft.net/fig/390161

42. XML treatment for Aleiodes unipunctator. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.39642B0E1D2FC-6C1C-54EB-AB9C1858C4DF4BC9 Aleiodes unipunctator (Thunberg, 1822)Figs 792–794, 795–807, 808–812

Ichneumon unipunctatorThunberg, 1822: 267 [examined]. Rogas unipunctator; Shenefelt 1975: 1254–1255; Zaykov 1980c: 229; Jakimavicius 1974: 96. Rogas (Rogas) unipunctator; Tobias 1976: 84, 1986: 78 (transl.: 128). Aleiodes (Neorhogas) unipunctator; Papp 1985a: 151, 163, 1991a: 86, 1996: 456; Belokobylskij 1996: 18; Papp and Rezbanyai-Reser 1996: 71, 73, 95, 96; Riedel et al. 2002: 106. Aleiodes (Chelonorhogas) unipunctator; Belokobylskij 2000: 44, 2003: 399; Chen and

He 1997: 43; He et al. 2000: 667; Rastegar et al. 2012: 3; Farahani et al. 2015: 229, 244. Aleiodes unipunctator; Čapek and Lukás 1989: 31; Bergamasco et al. 1995: 6; O’Connor et al. 1999: 92; Fortier and Shaw 1999: 230; Belokobylskij and Taeger 2001: 115; Marsh and Shaw 2001: 303; Belokobylskij et al. 2003: 399; Zaldívar-Riverón et al. 2004: 234; Papp 2005: 177; Lozan et al. 2010: 17. Ichneumon ductorThunberg, 1822: 269; Papp 1985a: 157 (not auctt.) [examined]. Syn. nov. Aleiodes irregularisWesmael, 1838: 101; Shenefelt 1975: 1255 (as synonym of A. unipunctator); Papp 1985a: 163 (id.) [examined]. Rhogas unipunctator ab. nigrescensHellén, 1927: 23; Shenefelt 1975: 1255 (excluded name). Holotype of A. unipunctator, ♂ (ZMUU) with holotype label by CvA. Holotype of A. ductor, ♂ (ZMUU), “α”, “Rhogas ductor Thbg”, “Uppsala Univ. Zool Mus., Thunbergsaml. Nr. 25332, Ichneumon ductor Sv. Type”. Holotype of A. irregularis, ♂ (KBIN), “[Belgium], 11 Juin, Brig.”, “A. irregularis ♂ mihi 5”, “dét. C. Wesmael”, “Belgique, Bruxelles/teste Papp J., 1983”, “Holotype”, “Aleiodes irregularis Wesm., 1838, ♂, Papp, 1983”, “Aleiodes unipunctator Thb. ♂, det. Papp J., 1984”. Austria, Belgium, British Isles (England: V.C.s 1, 3, 5, 6, 8, 15, 16, 17, 18, 20, 21, 23, 25, 26, 27, 28, 29, 32, 33, 40, 53, 55, 58, 59, 63, 64, 65, 66, 69; Wales: V.C.s 35, 52 ; Scotland: V.C.s 72, 75, 76, 77, 79, 80, 82, 83, 85, 86, 87, 88, 89, 90, 91, 95, 96, 97, 98, 99, 100, 101, 103, 105, 108, 109, 110, 111, 112; Ireland: V.C.s H5, H19, H20, H21, H22, H28, H30), Bulgaria, Czech Republic, Denmark, Finland, Germany, Greece (mainland, Corfu), Hungary, Italy, Montenegro, Netherlands (DR: Wijster; Borger; GE: Nunspeet; Kemperberg, NB: Bergen op Zoom, ZH: Arkel; Melissant; Oostvoorne; ZE: Oostkapelle), Norway, Romania, Russia, Sweden, [Kazakhstan, Tadzhikistan, W. Caucasus]. Specimens in AAC, ALC, ZJUH, BZL, CNC, FMNH, HHC, HSC, IKC, OUM, MMUM, MRC, MSC, MSNV, MTMA, NMI, NMS, RMNH, SDEI, UMZC, UNS, UWIM, ZMUU, ZSSM. MRS211 (UK), MRS221 (Germany), MRS354 (UK), MRS893 (UK). Univoltine, flying from May to August. Reared from the noctuids Apamea unanimis (Hübner) (23 [1 ZJUH, 1 FMNH, 1 NRS, 1 MTMA]; M.R. Shaw), Apamea crenata (Hufnagel) (2:1, ZJUH; G.T. Lyle), Apamea ? sordens (Hufnagel) (3:1; K.P. Bland), and from mummies compatible with Apamea spp. (8). These closely related hosts all live in the shoots or leaf sheaths of Poaceae. The large number reared from A. unanimis may be at least partly the result of a sampling bias, as the larvae of that species are so readily detected and collected when feeding on Phalaris. The winter is passed in the relatively slender brown mummy, which is nearly cylindrical and only weakly keeled laterally (Fig. 794). It probably usually forms at or below soil level and is rather weakly contracted at the head end (which is bent sideways in a high proportion of cases), with the thinly silken cocoon occupying abdominal segments (2–)3–8. This species is widespread and often abundant, especially in rank or damp grassland habitats, in the northern part of its range. Maximum width of hypoclypeal depression 0.5–0.6 × minimum width of face (Fig. 802); OOL of ♀ approx. 0.8 × as long as diameter of posterior ocellus and mainly granulate mixed with some punctures (Fig. 803); ventral margin of clypeus thick, not protruding in lateral view (Fig. 804); mesoscutal lobes finely granulate-punctulate and matt; precoxal area comparatively narrow and moderately rugose medially; marginal cell of fore wing of ♀ ending rather close to wing apex (Fig. 795); vein 1-CU1 of fore wing 0.4–0.5 × as long as vein 2-CU1 (Fig. 795); hind tarsal claws rather robust and only brownish setose (Fig. 807); 1st tergite rather slender basally (Fig. 798); whole 4th and part of 3rd metasomal tergite smooth and very glossy (Fig. 798); labial palp yellowish brown or brown; basal half of hind tibia pale yellowish or ivory, at least inner side contrasting with reddish or dark brown colour of basal half of hind femur (usually less pronounced in ♂), and its apex dark brown or black; 4th and 5th tergites black; wings rather slender and their membrane subhyaline. Redescribed ♀ (RMNH) from Sweden (Storbacken). Length of fore wing 5.8 mm, of body 5.7 mm. Head. Antennal segments of ♀ 50, length of antenna 1.2 × fore wing, its subapical segments rather robust; frons largely smooth anteriorly (except some fine sculpture) and densely rugulose posteriorly; OOL 0.8 × diameter of posterior ocellus, granulate with some punctures and matt; vertex granulate with some rugulosity and rather dull; clypeus coarsely punctate; ventral margin of clypeus thick and not protruding forwards (Fig. 804); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 802); length of eye 1.7 × temple in dorsal view (Fig. 803); vertex behind stemmaticum rugulose-granulate; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view. Mesosoma. Mesoscutal lobes moderately punctate and interspaces distinctly granulate, with satin sheen; precoxal area of mesopleuron moderately rugose medially, sparsely punctulate posteriorly as surroundings of precoxal area; scutellum rather sparsely punctate, but medio-posteriorly rugulose, shiny; propodeum rather convex and moderately rugose, medio-longitudinal carina complete, and no protruding carinae laterally. Wings. Fore wing: r 0.3 × 3-SR (Fig. 795); 1-CU1 narrow and horizontal, 0.4 × 2-CU1; r-m 0.5 × 3SR; 2nd submarginal cell medium-sized and 1st subdiscal cell slender (Fig. 795); cu-a nearly vertical, straight; 1-M slightly curved posteriorly; 1-SR narrow posteriorly and widened anteriorly; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: basal half of marginal cell slightly wider and its apical half distinctly gradually widened, its apical width 2.2 × width at level of hamuli (Fig. 795); 2-SC+R short longitudinal; m-cu indistinct; M+CU:1-M = 4:3; 1r-m 0.7 × 1-M. Legs. Tarsal claws yellowish bristly setose, without distinct pecten (Fig. 807); hind coxa largely distinctly punctate; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 6.3 × their width, respectively; length of inner hind spur 0.5 × hind basitarsus. Metasoma. First tergite rather flattened, 1.2 × as long as wide apically; 1st and 2nd tergites with distinct medio-longitudinal carina and longitudinally striate; medio-basal area of 2nd tergite wide triangular and short (Fig. 798); 2nd suture rather deep and finely crenulate; 3rd tergite nearly entirely smooth and strongly shiny, as remainder of metasoma; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with rather long setae and apically truncate (Fig. 793). Colour. Black; pronotum dorso-posteriorly, telotarsi largely, hind tarsus, apical half of hind tibia, pterostigma (except paler extreme base), most veins and apical fifth of 2nd tergite dark brown; palpi brown; tegulae and basal half of hind tibia pale yellowish; apical third of 1st tergite and 2nd tergite (except apically) and remainder of legs, orange brown; wing membrane subhyaline. Variation. Propodeum and pronotum sometimes weakly marked with orange. One male seen with vein r-m of fore wing absent. Antennal segments: ♀ 47(1), 48(7), 49(12), 50(21), 51(26), 52(40), 53(40), 54(15), 55(5), 56(3), 57(1); ♂ 47(1), 48(1), 49(1), 50(8), 51(7), 52(22), 53(27), 54(34), 55(35), 56(21), 57(6), 58(8), 59(1). On average males have ca one to two more antennal segments

than females. Male is very similar with apical tergites type 2, setae rather sparse and fringe short (Figs 809, 810). Melanistic females and males occur, metasoma entirely blackish or with only apex of 1st tergite narrowly and 2nd tergite medially and antero-laterally dark orange; clypeus reddish brown or blackish; apex of hind femur often somewhat infuscate. Austria, British Isles (England, Wales, Scotland, Ireland), Bulgaria, Czech Republic, *Denmark, Finland, Germany, *Greece (mainland and Corfu), Hungary, Italy, *Kazakhstan, *Montenegro, Netherlands, Norway, *Romania, Russia (including W. Caucasus), Sweden, *Tadzhikistan. Both male holotypes of I. unipunctator and I. ductor are preserved in the Thunberg collection, but are severely damaged. From the holotype of A. ductor only the head, fore coxa, mesoscutum and metasoma remain (Roman, 1912). The holotype of A. unipunctator has the head and the metasoma separately glued on a card and the mesosoma is still attached to the pin. Judging from these remnants (especially the mainly smooth and very shiny 3rd tergite, the wide hypoclypeal depression (0.6 × minimum width of face) and the coriaceous vertex), it is obvious that both belong to the same species. Consequently, A. ductor (Thunberg) is synonymised with A. unipunctator (Thunberg) (syn. nov.). Aleiodes ductor auctt. is divided among A. pallidicornis (Herrich-Schäffer, 1838) (N and C European populations) and A. apicalis (Brullé, 1832) (Mediterranean and southern C European populations). 10.3897/zookeys.919.39642.figures792-794BA2D35E3-C873-5AD5-95B2-CE577D30A360Figures 792–794. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 792 habitus lateral 793 ovipositor sheath lateral 794 mummy of Apamea unanimis Hübner. https://binary.pensoft.net/fig/39016210.3897/zookeys.919.39642.figures795-80791FB4AA9-CBFA5C83-81A5-87C52817F0C0Figures 795–807. Aleiodes unipunctator (Thunberg), ♀, England, Fletcher Moss 795 wings 796 mesosoma lateral 797 mesosoma dorsal 798 metasoma dorsal 799 fore femur lateral 800 hind femur lateral 801 antenna 802 head anterior 803 head dorsal 804 head lateral 805 base of antenna 806 apex of antenna 807 inner hind tarsal claw lateral. https://binary.pensoft.net/fig/39016310.3897/zookeys.919.39642.figures808-81211FC92CF-95685FE1-B32F-5BA260E837E8Figures 808–812. Aleiodes unipunctator (Thunberg), ♂, Scotland, Edinburgh, but 808 Sweden, Särö-Hamra 808 habitus lateral 809 apex of metasoma dorsal 810 apex of metasoma lateral 811 head dorsal 812 base of antenna. https://binary.pensoft.net/fig/390164ShenefeltRD (1975) Braconidae 8. Exothecinae, Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.ZaykovAN (1980c) Contribution to the braconid fauna of Bulgaria (Hymenoptera, Braconidae). Travaux scientifiques d’Université de Plovdiv “Paissi Hilendarski”, Biologie 16(4) [1978]: 227–234. [in Bulgarian with English summary]JakimaviciusAB (1974) Species of braconids new for the fauna of the Lithuanian SSR (Hym., Braconidae).Trudy Akademii Nauk Litovskoi SSR B2(66): 93–102.TobiasVI (1976) Braconids of the Caucasus (Hymenoptera, Braconidae).Opredeliteli po Faune SSSR110: 1–286. [in Russian]TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]PappJ (1985a) Contributions to the Braconid fauna of Hungary, VII. Rogadinae (Hymenoptera: Braconidae).Folia Entomologica Hungarica46: 143– 164.PappJ (1991a) Parasitic wasps of the superfamily Ichneumonoidea. Braconidae. 18. Doryctinae. 19. Rogadinae.Magyarorszag Allatvilaga [Fauna Hungariae]167: 1–122.BelokobylskijSA (1996) Contribution to the knowledge of braconid fauna of the subfamily Rogadinae (Hymenoptera, Braconidae) of Russian Far East and Eastern Siberia. Part 1. Far Eastern Entomologist 27–28: 1– 12.PappJRezbanyai-ReserL (1996) Contributions to the braconid fauna of Monte Generoso, Canton Ticino, southern Switzerland (Hymenoptera: Braconidae).Entomologische Berichte Luzern35: 59– 134.RiedelMHansenLOBergØ (2002) Braconidae (Hymenoptera) of Norway, Part 1.Norwegian Journal of Entomology49: 97–108.BelokobylskijSA (2000) Rogadinae: 15–109. In: LerPA (Ed.) Key to the insects of Russian Far East, vol. 4.Neuropteroidea, Mecoptera, Hymenoptera, pt4: 1–651. Vladivostok.ChenX-XHeJ-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China.Zoologische Verhandelingen, Leiden308: 1–187.HeJ-HChenX-XMaY (2000) HymenopteraBraconidae. Fauna Sinica. Insecta 18: 1–757. Science Press, Beijing.RastegarJSakeninHKhodaparastSHavaskaryM (2012) On a collection of Braconidae (Hymenoptera) from East Azarbaijan and vicinity, Iran.Calodema226: 1–4.FarahaniSTalebiAAvan AchterbergCRakhshaniE (2015) A review of the subfamily Rogadinae (Hymenoptera: Braconidae) from Iran.Zootaxa3973(2): 227–250. https://doi.org/10.11646/zootaxa.3973.2.2ČapekMLukásJ (1989) ApocritaParasitica, Ichneumoidea, Braconidae.Acta Faunistica Entomologica Musei Nationalis Pragae19: 27–44.BergamascoPPennacchioFScaramozzinoPLTremblayE (1995) HymenopteraBraconidae.Checklist della Specie della Fauna Italiana95: 2–29.O’ConnorJPNashRvan AchterbergC (1999) A catalogue of the Irish Braconidae (Hymenoptera: Ichneumonoidea).Occasional Publication of the Irish Biogeographical Society4: 1– 123.FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204– 237.BelokobylskijSATaegerA (2001) Braconidae: 103–115. In: DatheHHTaegerABlankSM (Eds) Verzeichnis der Hautflügler Deutschlands (Entomofauna Germanica 4).Entomologische Nachrichten und Berichte (Dresden), Beiheft7: 1–178.MarshPMShawSR (2001) Revision of North American Aleiodes Wesmael (Part 6): The gasterator (Jurine) and unipunctator (Thunberg) species-groups (Hymenoptera: Braconidae: Rogadinae).Proceedings of the Entomological Society of Washington103(2): 291–307.BelokobylskijSATaegerAvan AchterbergCHaeselbarthERiedelM (2003) Checklist of the Braconidae (Hymenoptera) of Germany.Beiträge zur Entomologie53(2): 341–435.Zaldívar-RiverónAButcherBAShawMRQuickeDLJ (2004) Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa.Zoologica Scripta33(3): 223–237. https://doi.org/10.1111/j.03003256.2004.00144.xPappJ (2005) A checklist of the Braconidae of Hungary (Hymenoptera). Folia Entomologica Hungarica.66: 137–194.LozanABelokobylskijSvan AchterbergCMonaghanM (2010) Diversity and distribution of Braconidae, a family of parasitoid wasps in the Central European peatbogs of South Bohemia, Czech Republic Journal of Insect Science 10: 16(1–21). https://doi.org/10.1673/031.010.1601

43. XML treatment for Aleiodes venustulus. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396424DEAB885-8DBC-588C-B538E5754A94E3FC Aleiodes venustulus (Kokujev, 1905)Figs 813–814, 815–826 Rhogas venustulusKokujev, 1905: 15. Rogas venustulus; Shenefelt 1975: 1255–1256; Tobias 1986: 78 (transl.: 129). Aleiodes venustulus; Fortier and Shaw 1999: 230; Aydogdu and Beyarslan 2005: 192, 2006: 87. Rhogas (Rhogas) robustusTelenga, 1941: 151, 415. Rogas robustus; Tobias 1986: 78 (transl.: 129; as synonym of A. venustulus). Holotype of A. venustulus, ♀ (ZISP), “[Kyrgyzstan], Ushch. Kyzyl-su[u], [= village south of Lake Issyk Kul], 7–9.000’[ft], 5.vii.[19]03, E. Pojarkov”, No. 2273, Rh. venustulus Kok.”, “K. Kokujeva”. 1 ♀ + 1 ♂ (MTMA), “[Kazakhstan:] Turkestan, Almásy, Naryn-kol, 1906, Tekkes”. None. Unknown. Maximum width of hypoclypeal depression 0.6–0.7 × minimum width of face (Fig. 822); OOL twice as long as diameter of posterior ocellus and coarsely rugose; clypeus rather short, rather flat, weakly protruding anteriorly and its ventral margin thick (Fig. 824); mesoscutum coarsely and remotely punctate, with interspaces smooth and wider than punctures; precoxal sulcus area coarsely and densely punctate; vein 1-CU1 of fore wing approx. 0.8 × as long as vein 2-CU1 and as long as vein m-cu (Fig. 815); membrane near veins M+CU1 and 1-CU1 of fore wing sparsely setose; tarsal claws rather robust and with some fine dark brown spiny bristles subbasally (Fig. 825); head brownish yellow; vein 1-M of fore wing brown; apical half of hind tibia dark brown; metasoma of ♀ yellowish, but anterior 0.6 of first tergite black; wing membrane subhyaline. Holotype of A. venustulus, ♀, length of fore wing 6.7 mm, of body 8.6 mm. Head. Antenna incomplete, with eight segments remaining; frons rugose and shiny; OOL twice diameter of posterior ocellus, mainly rugose and shiny; stemmaticum densely punctate; vertex remotely punctate and shiny; clypeus punctate and slightly convex; ventral margin of clypeus thick and anterior part weakly protruding (Fig. 824); width of hypoclypeal depression 0.6 × minimum width of face (Fig. 822); length of eye 1.3 × temple in dorsal view (Fig. 823); vertex behind stemmaticum convex and remotely punctate; clypeus near lower level of eyes; length of malar space 0.4 × length of eye in lateral view and temple as wide as eye. Mesosoma. Mesoscutum coarsely and remotely punctate, with interspaces smooth and wider than punctures; precoxal area coarsely and densely punctate, remainder of mesopleuron remotely punctate and antero-dorsally rugose; metapleuron densely and coarsely punctate; metanotum with coarse medio-longitudinal carina anteriorly; scutellum punctate; propodeum convex and coarsely rugose, its

medio-longitudinal carina present only on anterior third of propodeum. Wings. Fore wing: r 0.4 × 3-SR (Fig. 815); 1-CU1 slightly oblique, 0.2 × 2-CU1; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 815); cu-a inclivous, straight; 1-M nearly straight posteriorly; 1-SR wide; surroundings of M+CU1, 1-M and 1-CU1 largely glabrous. Hind wing: marginal cell linearly widened, its apical width twice width at level of hamuli (Fig. 816); 2-SC+R short and vertical; m-cu absent; M+CU:1-M = 12:11; 1r-m 0.7 × 1-M. Legs. Tarsal claws rather robust and with some fine dark brown spiny bristles subbasally (Fig. 825); hind coxa largely densely punctate; hind trochantellus rather robust; length of hind femur and basitarsus 4.7 and 6.5 × their width, respectively; length of inner hind spur 0.4 × hind basitarsus. Metasoma. First tergite evenly convex, as long as wide apically; 1st and 2nd tergites with mediolongitudinal carina and coarsely rugose-reticulate; medio-basal area of 2nd tergite triangular and rather distinct (Fig. 819); 2nd suture deep and narrow; basal half of 3rd tergite punctate-rugose, remainder of metasoma superficially micro-sculptured; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath wide, with long setae and apically truncate (Fig. 814). Colour. Black; antenna (except scapus and pedicellus), palpi, tegulae, fore and middle telotarsi, veins, and pterostigma dark brown; coxae, trochanters and trochantelli, apical third of hind femur (ventrally extended to its apical two-thirds), hind tibia (except pale yellowish basal ring), fore and middle femora apically, and hind tarsus black; remainder of legs yellowish brown; wing membrane subhyaline. Variation. No specimens with intact antennae examined; 2nd tergite coarsely rugose-reticulate or coarsely longitudinally rugose. Male is very similar and with apical tergites type 3, setae quite dense, glabrous stripe narrow, and fringe very weak. Kazakhstan, Kyrgyzstan. This Central Asian species bears a superficial resemblance to A. miniatus and A. aestuosus. It is included in this revision, because it has been reported twice from Turkey (Aydogdu and Beyarslan 2005, 2006). 10.3897/zookeys.919.39642.figures813-8148142EA8E-5E25-5EDB-A8C93B9CCCA957A0Figures 813, 814. Aleiodes venustulus (Kokujev), ♀, holotype 813 habitus lateral 814 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016510.3897/zookeys.919.39642.figures815-82628ABB556-32445A75-A7A0-7CC856C3A26EFigures 815–826. Aleiodes venustulus (Kokujev), ♀, holotype 815 fore wing 816 hind wing 817 mesosoma lateral 818 mesosoma dorsal 819 metasoma dorsal 820 fore femur lateral 821 hind femur lateral 822 head anterior 823 head dorsal 824 head lateral 825 inner hind tarsal claw lateral 826 base of antenna. https://binary.pensoft.net/fig/390166ShenefeltRD (1975) Braconidae 8. Exothecinae,

Rogadinae.Hymenopterorum Catalogus (nova editio)12: 1115–1262.TobiasVI (1986) Rogadinae: 72–85. In: MedvedevGS (Ed.) Opredelitel Nasekomych Evrospeiskoi Tsasti SSSR 3, Peredpontdatokrylye 4.Opredeliteli po Faune SSSR145: 1–501. [Translation (1995): 512–588. Lebanon, U.S.A.]FortierJCShawSR (1999) Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera: Braconidae).Journal of Hymenoptera Research8(2): 204–

237.AydogduMBeyarslanA (2005) The first records of Aleiodes Wesmael 1838 (Hymenoptera: Braconidae: Rogadinae). The fauna of Thrace region of Turkey. Linzer biologische Beiträge 37/1: 185–193.AydogduMBeyarslanA (2006) First records of Aleiodes Wesmael, 1838 species in East Marmara region of Turkey (Hymenoptera: Braconidae: Rogadinae).Acta Entomologica Slovenica14(1): 81–88.

44. XML treatment for Aleiodes zwakhalsi. Taxon classificationAnimaliaHymenopteraBraconidaevan AchterbergCornelisShawMark R.QuickeDonald L.J.Revision of the western Palaearctic species of Aleiodes Wesmael (Hymenoptera, Braconidae, Rogadinae). Part 2: Revision of the A. apicalis groupZookeys1632020919125910.3897/zookeys.919.396429BE89C08-4F59-52A4-913446994C08F756 Aleiodes zwakhalsi http://zoobank.org/3C42EDAB-B9DB-45DB-884CB23A630B249F van Achterberg & Shawsp. nov.Figs 827–829, 830–840 Holotype, ♀ (RMNH), “Turkey, Ankara, Kizilcahaman, 1100 m, 17.vi.1985, C.[J.] Zwakhals”. Paratypes: 1 ♂ (RMNH), “Turkey, Agri, Mt. Ararat, 1800 m, 3.vii.1985, C.J. Zwakhals”; 1 ♀ (RMNH), “Turkiye, Gümüshane, 40–46 km E [of] Bagburt, 1500 m, 19.vii.1989, J.A.W. Lucas”; 1 ♀ (NMS), “Turkey: Zigana Dagi, 5,000 ft., SW of Tabzon, 10.viii.1959, K.M. Guichard”. None. Unknown. Probably univoltine as all known adults were collected in the period June to August, but there is no indication of its means of overwintering. Maximum width of hypoclypeal depression approx. 0.7 × minimum width of face (Fig. 837); OOL of ♀ finely remotely punctate and 0.8–1.0 × diameter of posterior ocellus; ventral margin of clypeus thick apically and not protruding in lateral view (Fig. 839); lobes of mesoscutum densely finely punctate, with interspaces shiny; precoxal area densely punctate and with some rugae medially; vein cu-a of fore wing oblique, parallel with vein 3-CU1; surroundings of veins M+CU1 and 1-+2-CU1 largely setose; vein r of fore wing 0.4–0.5 × vein 3-SR (Fig. 830); vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1 (Fig. 830); hind tarsal claws with rather conspicuous pale brown pecten (Fig. 840); 1st tergite gradually narrowed basally (Fig. 833); 2nd tergite 0.7 × as long as wide basally and black; 3rd tergite densely punctulate basally, and sparsely so apically; head black; vein 1-M of fore wing brown; wing membrane subhyaline. This new species is similar to A. cruentus (Nees), but it differs by having the surroundings of veins M+CU1 and 1-+2-CU1 partly setose (Fig. 828; largely glabrous in A. cruentus), vein M+CU1 of fore wing with bend near its distal fifth (Fig. 828; straight or slightly curved), vein cu-a of fore wing oblique, parallel with vein 3-CU1 (vertical or nearly so, rarely oblique), vein r of fore wing 0.4–0.5 × vein 3-SR (0.3–0.4 ×), vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1 (0.8–1.1 ×, rarely less), OOL of ♀ 0.8–1.0 × diameter of posterior ocellus (0.5–0.8 à —, rarely longer), length of eye 1.1–1.6 × temple in dorsal view (1.5–1.9 ×), ovipositor sheath comparatively slender and area between ocelli and eyes moderately punctate (coarsely punctate). Also very similar to A. diversus (Szépligeti), it differs from the latter by having vein 1CU1 of fore wing distinctly shorter than vein m-cu, hind femur approx. 4 × as long as wide, vein cu-a inclivous (parallel with vein 3-CU1; vertical and vein 3-CU1 diverging posteriorly in A. diversus), 5th–10th antennal segments of ♀ as long as wide (shorter than wide), vertex and OOL remotely punctate (densely punctate) and ovipositor sheath slender (robust). Holotype, ♀, length of fore wing 7.0 mm, of body 9.0 mm. Head. Antennal segments of ♀ 59, length of antenna 1.1 × fore wing, its subapical segments moderately slender; frons largely smooth; OOL equal to diameter of posterior ocellus, finely remotely punctate and shiny; vertex distinctly punctate and shiny; clypeus punctate-rugose, wide and short; ventral margin of clypeus thick and not protruding forwards (Fig. 839); width of hypoclypeal depression 0.7 × minimum width of face (Fig. 837); length of eye 1.1 × temple in dorsal view (Fig. 838); vertex behind stemmaticum densely punctate; clypeus near lower level of eyes; length of malar space 0.25 × length of eye in lateral view.

Mesosoma. Mesoscutal lobes largely densely and finely punctate, shiny; precoxal area of mesopleuron densely punctate, medially with few rugae; surroundings of precoxal area densely punctate; scutellum sparsely and finely punctate, rather flat, shiny and laterally rugose-punctate; propodeum evenly convex and coarsely rugose, medio-longitudinal carina complete but irregular posteriorly, and no protruding carinae laterally. Wings. Fore wing: r 0.4 × 3-SR (Fig. 830); 1-CU1 horizontal, 0.5 × 2-CU1 and 0.7 × m-cu; r-m 0.6 × 3-SR; 2nd submarginal cell medium-sized (Fig. 830); cu-a inclivous, straight; 1-M slightly curved posteriorly; vein M+CU1 of fore wing with distinct bend near its distal fifth (Fig. 828); 1-SR widened; surroundings of M+CU1, 1-M and 1-CU1 largely setose. Hind wing: marginal cell gradually widened, its apical width 2.1 × width at level of hamuli (Fig. 830); 2-SC+R subquadrate; short m-cu weakly developed; M+CU:1-M = 7:4; 1r-m 0.9 × 1-M. Legs. Tarsal claws with rather conspicuous pale brownish pecten (Fig. 840); hind coxa largely densely punctulate; hind trochantellus robust; length of hind femur and basitarsus 4.0 and 5.0 × their width, respectively; length of inner hind spur 0.55 × hind basitarsus. Metasoma. First tergite rather flattened, as long as wide apically and distinctly narrowed basally (Fig. 833); 1st and 2nd tergites with medio-longitudinal carina and coarsely longitudinally rugose; medio-basal area of 2nd tergite wide triangular and short (Fig. 833); 2nd tergite 0.7 × as long as its basal width; 2nd suture deep and finely reticulate; basally 3rd tergite densely punctulate and apically (as remainder of metasoma) sparsely punctulate; 4th and apical half of 3rd tergite without sharp lateral crease; ovipositor sheath slender, with medium-sized setae and apically rounded (Fig. 829). Colour. Black (including fore coxa anteriorly and basally); apex of hind tibia, telotarsi, hind tarsus, palpi, pterostigma and veins, dark brown; hind tibia (except apex) brownish yellow; clypeus narrowly ventrally, remainder of legs, pronotum (except ventrally), mesopleuron dorsally, mesoscutum, scutellum, and metanotum, orange-brown; tegulae pale brownish yellow; wing membrane subhyaline. Variation. Antennal segments of ♀ 58(1), 59(1), 60(1), of ♂ 62(1); vein r of fore wing 0.4–0.5 × vein 3-SR; vein 1-CU1 of fore wing 0.3–0.6 × vein 2-CU1; OOL of ♀ 0.8–1.0 × diameter of posterior ocellus; length of eye 1.1–1.6 × temple in dorsal view; clypeus ventrally orange brown or black; mesopleuron dorsally or largely orange brown; basal half of third tergite rugose, punctate-rugose or punctulate; entire 1st tergite orange brown, posterior half of 1st tergite brownish and rest of tergite blackish (as base of middle coxa) or entirely black. Male is very similar, apical tergites type 1 and no fringe observed. Turkey (Asian part: 1100–1800 m). 10.3897/zookeys.919.39642.figures827-829A061B54B-32E2-5484-A351-D78AD200A589Figures 827–829. Aleiodes zwakhalsi sp. nov., ♀, holotype 827 habitus lateral 828 detail of fore wing 829 ovipositor sheath lateral. https://binary.pensoft.net/fig/39016710.3897/zookeys.919.39642.figures830-8404733616C-9C135269-9486-8C6D9C5DD7EBFigures 830–840. Aleiodes zwakhalsi sp. nov., ♀, holotype 830 wings 831 mesosoma lateral 832 mesosoma dorsal 833 metasoma dorsal 834 fore femur lateral 835 hind femur lateral 836 base of antenna 837 head anterior 838 head dorsal 839 head lateral 840 outer hind tarsal claw lateral. https://binary.pensoft.net/fig/390168