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Hyposoter didymator (Thunberg, 1824) Ichneumon didymator Thunberg, 1824: holotype ♀ in University Museum, Uppsala, Sweden, examined by Horstmann. syn. Ichneumon rotundator Thunberg, 1824: Roman (1912) syn. Campoplex ruficincta Gravenhorst, 1829: Roman (1912) syn. Anilastus schmiedeknechti Smits van Burgst, 1913: Aubert (1966a) syn. Trophocampa nigrata Kiss, 1926: Horstmann (2009) Roman (1912) referred to a ‘single well preserved specimen’ in the Thunberg collection. Townes et al. (1965) designated what is presumably the same specimen as ‘type’. This is a common and well-known species. Horstmann placed it in the key in his section covering species with predominantly or completely black metasomata as well as in that covering species with red on the metasoma. Specimens from most of Europe which we have seen always have a considerable amount of red on the metasoma, usually involving the whole of at least one tergite. However, a long series from Norway is much darker, with some specimens showing no red at all. This may perhaps be a boreal form of the species. The most likely species with which it might be confused is Hyposoter barrettii: while didymator always has a smooth or only faintly sculptured speculum, barrettii has some fine striae extending across it. In the female the broad emarginate notch in the seventh sternite of barrettii is entirely lacking in didymator. The dark banding on the hind tibia of didymator is also more sharply defined and broader than in barrettii. Rearing records are very numerous from a wide variety of lepidopteran hosts, including Geometridae, Lasiocampidae, Noctuidae, Notodontidae, Papilionidae and Pieridae. Shaw et al. (2016) provide a long list of species. In addition to these, we have found among the previously unidentified NMS material 2♀ reared from Lithomoia solidaginis (Ḩbner) in Poland, 1♀ from Noctua pronuba (Linnaeus) in Austria, 1♀ marked as reared from imported Spodoptera litura (Fabricius) in the UK, and 1♀ from Trachea atriplicis (Linnaeus) in France. In NHMUK there are the following reared specimens, all from Noctuidae: ex Anarta myrtilli (Linnaeus) (1♁), “ H. neglecta ” probably = Xestia castanea (Esper) (1♁), Xestia agathina (Duponchel) (2♁), Plusia on potatoes (2♁ 1♀), on lucerne (1♁ 3♀), and on beans (1♁), Spodoptera littoralis (Boisduval) on lucerne (3♁ 3♀), Helicoverpa armigera (Ḩbner) on lucerne (3♀) and on tomato (1♁), Heliothis peltigera (Denis & Schiffermüller) on weeds (2♁ 1♀), Spodoptera exigua (Hübner) on alfalfa (2♀), on maize (1♁ 1♀), and on lucerne (1♁ 1♀), and Cucullia gnaphthalii (Hübner) (1♁, 1♀). There are many more records, though of uncertain authenticity, in the literature. We have seen specimens from the UK, the Netherlands, France, Belgium, Norway, Sweden, Austria, Spain, Hungary, Poland, Greece and Cyprus. The species has been recorded widely in the literature across Eurasia from Ireland to Russia and Tajikistan. It was introduced to Australia and New Zealand. We have seen specimens taken in every month from May to October, with numbers fairly evenly spread through June to September. This no doubt reflects the diversity of available hosts for this plurivoltine species. Three specimens from the third author’s collection were barcoded (BOLD Sample IDs CollHH1222, 1223 and 1224)., Published as part of Galsworthy, Anthony, Shaw, Mark R. & Haraldseide, Håkon, 2023, A key to European species of Hyposoter Förster, 1869 (Ichneumonidae: Campopleginae) with descriptions of 18 new species, and notes on all included species, pp. 1-73 in Zootaxa 5290 (1) on page 40, DOI: 10.11646/zootaxa.5290.1.1, http://zenodo.org/record/7959256, {"references":["Thunberg, C. P. (1824) Ichneumonidea, Insecta Hymenoptera illustrata. Memoires de l'Academie Imperiale des Sciences de Saint Petersbourg, 9, 285 - 368.","Roman, A. (1912) Die Ichneumonidentypen C. P. Thunbergs. Zoologiska Bidrag fran Uppsala, 1, 229 - 293.","Gravenhorst, J. L. C. (1829) Ichneumonologia Europaea. Pars III. Sumtibus auctoris, Vratislaviae, 1097 pp. https: // doi. org / 10.5962 / bhl. title. 65750","Smits van Burgst, C. A. L. (1913) Tunesien Hymenoptera. Privatdruk, den Haag, 37 pp.","Aubert, J. - F. (1966 a) Les Ichneumonides du rivage mediterraneen francais (9 e serie: Ouest de l'Herault et Aude) (Hym.). (suite et fin). Bulletin de la Societe Entomologique de France, 71, 166 - 176. https: // doi. org / 10.3406 / bsef. 1966.20879","Kiss, von Zilah A. (1926) Zweiter Beitrag zur Kenntnis der ungarischen und siebenb ¸ rgischen Ichneumoniden- (Schlupfwespen-) Fauna. Verhandlungen und Mitteilungen des Siebenburgischen Vereins fur Naturwissenschaften in Hermannstadt, 75 / 76, 74 - 120.","Horstmann, K. (2009) Typenrevisionen der von Kiss beschriebenen Taxa der Ichneumonidae III. Verschiedene Unterfamilien (Hymenoptera, Ichneumonidae). Linzer Biologische Beitrage, 41 (1), 673 - 689.","Townes, H. K., Momoi, S. & Townes, M. (1965) A catalogue and reclassification of the eastern Palearctic Ichneumonidae. Memoirs of the American Entomological Institute, 5, 1 - 661.","Shaw, M. R., Horstmann, K. & Whiffin, A. L. (2016) Two hundred and twenty-five species of reared western Palaearctic Campopleginae (Hymenoptera: Ichneumonidae) in the National Museums of Scotland, with descriptions of new species of Campoplex and Diadegma, and records of fifty-five species new to Britain. Entomologist's Gazette, 67, 177 - 222."]}

En los últimos años, el creciente interés por buscar nuevos agentes de control biológico y el fácil acceso a las tecnologías de secuenciación masiva ha facilitado el descubrimiento de un gran número de virus presentes en insectos. Muchos de estos virus infectan de forma encubierta sin causar síntomas evidentes. Spodoptera exigua iflavirus 1 (SeIV1) es un virus de ARN que causa infecciones encubiertas tanto en las poblaciones de laboratorio como en las de campo de la rosquilla verde, Spodoptera exigua (Hübner). Su abundancia en el campo sugiere que podría causar un efecto sobre la bioecología del insecto, ya sea de forma directa afectando a su fisiología o de forma indirecta a través de la interacción con otros niveles tróficos.

Hyposoter didymator (THUNBERG,1822) M a t e r i a l e x a m i n e d: Markazisite 1: 1♀ 11.vi.-20.vii.2018, leg. M. Parachami-Araghi; Khuzestan site 1: 2♀♀ 4♂♂ 11.iii.-10.v.2015, leg. E. Gilasian. D i s t r i b u t i o n: Palaearctic region, known from Iran (BARAHOEI et al. 2012)., Published as part of Riedel, Matthias, Ameri, Ali, Talebi, Ali Asghar & Ebrahimi, Ebrahim, 2019, Contribution to the Ɨchneumonidae (Hymenoptera) oI Ɨran, with descriptions oI seven new species, pp. 361-390 in Linzer biologische Beiträge 51 (1) on page 367, DOI: 10.5281/zenodo.3758345, {"references":["BARAHOEI H., RAKHSHANI E. & M. RIEDEL (2012): A checklist of Ichneumonidae (Hymenoptera: Ichneumonoidea) from Iran. 2 Iranian Journal of A nimal Biosystematics 8: 83 - 132."]}

Hyposoter didymator (Hymenoptera, Ichneumonidae) is a generalist solitary endoparasitoid of noctuid larvae. In the present work, we tested whether populations of H. didymator were divided in several genetically distinct taxa as described for many other generalist parasitoid species, and whether differences in H. didymator parasitism rates were explained by the insect host species and/or by the plant on which these hosts were feeding on. The genetic analysis of natural populations collected in different regions in France and Spain on seven different insect hosts and seven different host plants (775 individuals) showed that H. didymator populations belong to a unique single taxon. However, H. didymator seems to be somewhat specialized. Indeed, in the fields it more often parasitized Helicoverpa armigera compared to the other host species collected in the present work. Also, H. didymator parasitism rates in field conditions and semi-field experimental studies were dependent on the host plants on which H. armigera larvae are feeding. Still, H. didymator can occur occasionally on non-preferred noctuid species. One hypothesis explaining the ability of H. didymator to switch hosts in natura could be related to fluctuating densities of the preferred host over the year; this strategy would allow the parasitoid to avoid seasonal population collapses.

Hyposoter didymator (THUNBERG, 1822) M a t e r i a l: H 1: 1♀ 4-14. VI; H 3: 1♀ 15-25. V. D i s t r i b u t i o n: Palaearctic region, known from Azerbaijan (ABDINBEKOVA 1963), new for Georgia., Published as part of Riedel, Matthias, Diller, Erich & Japoshvili, George, 2018, The Ichneumonid fauna (Hymenoptera: Ichneumonidae) of Lagodekhi Reserve, Sakartvelo (Georgia), with descriptions of four new species, pp. 1447-1507 in Linzer biologische Beiträge 50 (2) on page 1454, DOI: 10.5281/zenodo.5275090, {"references":["ABDINBEKOVA A. A. (1963): [The Ichneumonid fauna of the Kuba Khachmassk zone of Azerbaidzhan.] (in Russian). - Izvestiya Akademii Nauk Azerbaid zhanskoi SSR 1963 (5): 43 - 51."]}

The wasp Hyposoter didymator (Hymenoptera, Ichneumonidae) parasitizes several agricultural pest moths and could therefore be used in biological control. The 454 FLX Titanium pyrosequencing technology was used to define two distinct sets of multiplex combining 14 polymorphic microsatellite loci: 10 (referred to as HD) located within the genome of H. didymator and 4 (referred to as HdIV) located within the Ichnovirus genome which is integrated into the wasp genome. Genotyping of two populations collected in France on Helicoverpa armigera revealed that most of the loci are independent and at Hardy-Weinberg equilibrium.

Ichnoviruses are large dsDNA viruses that belong to the Polydnaviridae family. They are specifically associated with endoparasitic wasps of the family Ichneumonidae and essential for host parasitization by these wasps. We sequenced the Hyposoter didymator Ichnovirus (HdIV) encapsidated genome for further analysis of the transcription pattern of the entire set of HdIV genes following the parasitization of four different lepidopteran host species. The HdIV genome was found to consist of at least 50 circular dsDNA molecules, carrying 135 genes, 98 of which formed 18 gene families. The HdIV genome had general features typical of Ichnovirus (IV) genomes and closely resembled that of the IV carried by Hyposoter fugitivus. Subsequent transcriptomic analysis with Illumina technology during the course of Spodoptera frugiperda parasitization led to the identification of a small subset of less than 30 genes with high RPKM values in permissive hosts, consisting with these genes encoding crucial virulence proteins. Comparisons of HdIV expression profiles between host species revealed differences in transcript levels for given HdIV genes between two permissive hosts, S. frugiperda and Pseudoplusia includens. However, we found no evident intrafamily gene-specific transcription pattern consistent with the presence of multigenic families within IV genomes reflecting an ability of the wasps concerned to exploit different host species. Interestingly, in two non-permissive hosts, Mamestra brassiccae and Anticarsia gemmatalis (most of the parasitoid eggs were eliminated by the host cellular immune response), HdIV genes were generally less strongly transcribed than in permissive hosts. This suggests that successful parasitism is dependent on the expression of given HdIV genes exceeding a particular threshold value. These results raise questions about the mecanisms involved in regulating IV gene expression according to the nature of the lepidopteran host species encountered.

Corrigendum to “Venom gland extract is not required for successful parasitism in the polydnavirus-associated endoparasitoid Hyposoter didymator (Hym. Ichneumonidae) despite the presence of numerous novel and conserved venom proteins” [Insect Biochem. Mol. Biol. 43 (2013), 292–307] http://dx.doi.org/10.1016/j.ibmb.2013.07.001(UT:000329772300006); The venom gland is a conserved organ in Hymenoptera that shows adaptations associated with life-style diversification. Few studies have investigated venom components and function in the highly diverse parasitic wasps and all suggest that the venom regulates host physiology. We explored the venom of the endoparasitoid Hyposoter didymator (Campopleginae), a species with an associated polydnavirus produced in the ovarian tissue. We investigated the effects of the H. didymator venom on two physiological traits of the host Spodoptera frugiperda (Noctuidae): encapsulation response and growth rate. We found that H. didymator venom had no significant effect on host cellular immunity or development, suggesting that it does not contribute to parasitism success. The host physiology seemed to be modified essentially by the ovarian fluid containing the symbiotic polydnaviruses. Proteomic analyses indicated that the H. didymator venom gland produces a large variety of proteins, consistent with the classical hymenopteran venom protein signature, including: reprolysin-like, dipeptidyl peptidase IV, hyaluronidase, arginine kinase or allergen proteins. The venom extracts also contained novel proteins, encoded by venom genes conserved in Campopleginae ichneumonids, and proteins with similarities to active molecules identified in other parasitoid species, such as calreticulin, reprolysin, superoxide dismutase and serpin. However, some of these proteins appear to be produced only in small amounts or to not be secreted. Possibly, in Campopleginae carrying polydnaviruses, the host-modifying activities of venom became. redundant following the acquisition of polydnaviruses by the lineage.

An endoparasitic life style is widespread among Hymenoptera, and various different strategies allowing parasitoids to escape from the host encapsulation response have been reported. Species carrying polydnaviruses (PDVs), such as the ichneumonid Hyposoter didymator, generally rely on the viral symbionts to evade host immune responses. In this work, we show that H. didymator eggs can evade encapsulation by the host in the absence of calyx fluid (containing the viral particles), whereas protection of the larvae requires the presence of calyx fluid. This evasion by the eggs depends on proteins associated with the exochorion. This type of local passive strategy has been described for a few species carrying PDVs. Immune evasion by braconid eggs appears to be related to PDVs or proteins synthesized in the oviducts being associated with the egg. We report that in H. didymator, by contrast, proteins already present in the ovarian follicles are responsible for the eggs avoiding encapsulation. Mass spectrometry analysis of the egg surface proteins revealed the presence of host immune-related proteins, including one with similarities with apolipophorin-III, and also the presence of three viral proteins encoded by IVSPERs (Ichnovirus Structural Protein Encoding Regions).

The ability to cope with plant defense chemicals differs between specialist and generalist species. In this study, we examined the effects of the concentration of the two main iridoid glycosides (IGs) in Plantago lanceolata, aucubin and catalpol, on the performance of a specialist and two generalist herbivores and their respective endoparasitoids. Development of the specialist herbivore Melitaea cinxia was unaffected by the total leaf IG concentration in its host plant. By contrast, the generalist herbivores Spodoptera exigua and Chrysodeixis chalcites showed delayed larval and pupal development on plant genotypes with high leaf IG concentrations, respectively. This result is in line with the idea that specialist herbivores are better adapted to allelochemicals in host plants on which they are specialized. Melitaea cinxia experienced less post-diapause larval and pupal mortality on its local Finnish P. lanceolata than on Dutch genotypes. This could not be explained by differences in IG profiles, suggesting that M. cinxia has adapted in response to attributes of its local host plants other than to IG chemistry. Development of the specialist parasitoid Cotesia melitaearum was unaffected by IG variation in the diet of its host M. cinxia, a response that was concordant with that of its host. By contrast, the development time responses of the generalist parasitoids Hyposoter didymator and Cotesia marginiventris differed from those of their generalist hosts, S. exigua and C. chalcites. While their hosts developed slowly on high-IG genotypes, development time of H. didymator was unaffected. Cotesia marginiventris actually developed faster on hosts fed high-IG genotypes, although they then had short adult longevity. The faster development of C. marginiventris on hosts that ate high-IG genotypes is in line with the “immunocompromized host” hypothesis, emphasizing the potential negative effects of toxic allelochemicals on the host’s immune response.

Many thousands of endoparasitic wasp species are known to inject polydnavirus (PDV) particles into their caterpillar host during oviposition, causing immune and developmental dysfunctions that benefit the wasp larva. PDVs associated with braconid and ichneumonid wasps, bracoviruses and ichnoviruses respectively, both deliver multiple circular dsDNA molecules to the caterpillar. These molecules contain virulence genes but lack core genes typically involved in particle production. This is not completely unexpected given that no PDV replication takes place in the caterpillar. Particle production is confined to the wasp ovary where viral DNAs are generated from proviral copies maintained within the wasp genome. We recently showed that the genes involved in bracovirus particle production reside within the wasp genome and are related to nudiviruses. In the present work we characterized genes involved in ichnovirus particle production by analyzing the components of purified Hyposoter didymator Ichnovirus particles by LC-MS/MS and studying their organization in the wasp genome. Their products are conserved among ichnovirus-associated wasps and constitute a specific set of proteins in the virosphere. Strikingly, these genes are clustered in specialized regions of the wasp genome which are amplified along with proviral DNA during virus particle replication, but are not packaged in the particles. Clearly our results show that ichnoviruses and bracoviruses particles originated from different viral entities, thus providing an example of convergent evolution where two groups of wasps have independently domesticated viruses to deliver genes into their hosts., Author Summary The polydnaviruses (PDVs) are a unique virus type used by an organism (a parasitic wasp) to manipulate the physiology of another organism (a lepidopteran host) in order to ensure successful parasitism. The evolutionary origin of these unusual viruses, found in ∼17,500 braconid wasps (Bracoviruses) and ∼15,000 ichneumonid wasps (Ichnoviruses), has been a major question for the last decade. We thus undertook an exclusive work aiming at investigating this origin via the characterization of genes encoding structural components for both types of PDVs. The present paper constitutes the first report on the identity and genome organisation of the viral machinery producing Ichnovirus virions. Our results strongly suggest that Ichnoviruses originated from a virus belonging to a group as yet uncharacterized that integrated its genome into that of an ichneumonid wasp ancestor. More importantly, our results demonstrate that the ancestor of Ichnoviruses differs from that of Bracoviruses, which originated from a nudivirus. We have now identified, for the two types of PDVs, the non packaged viral genes and their products involved in producing particles injected into the host during oviposition. Together, these data provide an example of convergent evolution where different groups of wasps have independently domesticated viruses to deliver genes into their hosts.

Hyposoter didymator (THUNBERG 1824) Ichneumon didymator THUNBERG 1824. Mem. Acad. Sc. St. Petersbourg 8: 265. Anilasta rufocincta: STROBL 1904: 90, Croatia. Hyposoter didymator: MACELJSKI & BALORIN 1975: 113, ex Autographa gamma L. D i s t r i b u t i o n: Tunisia, Europe, Azerbaijan, Turkey, Cyprus, Israel, Tajikistan, Dagestan, Uzbekistan, Siberia and Australia, introduced into New Zealand., Published as part of Kolarov, J., 2008, A Catalogue of the [former] Yugoslavian Ichneumonidae (Hymenoptera, Insecta), pp. 1585-1739 in Linzer biologische Beiträge 40 (2) on page 1607, DOI: 10.5281/zenodo.5431642, {"references":["THUNBERG C. P. (1824): Ichneumonidea, Insecta Hymenoptera illustrata. - Mem. Acad. Imp. Sci. Sankt Petersbourg 9: 285 - 368.","STROBL G. (1904): Ichneumoniden Steiermarks (und der Nachbarlander). V. Fam. Ophionidae. - Mitt. Naturwiss. Ver. Steiermark, Graz 40 (1903): 43 - 160."]}

Background The abundance and the conservation of the repeated element (rep) genes in Ichnoviruses genomes suggest that this gene family plays an important role in viral cycles. In the Ichnovirus associated with the wasp Hyposoter didymator, named HdIV, 10 rep genes were identified to date. In this work, we report a relative quantitative transcription study of these HdIV rep genes in several tissues of the lepidopteran host Spodoptera frugiperda as well as in the H. didymator wasps. Results The data obtained in this work indicate that, in the early phases of infection (24 hours), HdIV rep genes each display different levels of transcripts in parasitized 2nd instar or HdIV-injected last instar S. frugiperda larvae. Only one, rep1, is significantly transcribed in female wasps. Transcript levels of the HdIV rep genes were found as not correlated to their copy number in HdIV genome. Our results also show that HdIV rep genes display different tissue specificity, and that they are primarily transcribed in S. frugiperda fat body and cuticular epithelium. Conclusion This work is the first quantitative analysis of transcription of the ichnovirus rep gene family, and the first investigation on a correlation between transcript levels and gene copy numbers in Ichnoviruses. Our data indicate that, despite similar gene copy numbers, not all the members of this gene family are significantly transcribed 24 hours after infection in lepidopteran larvae. Additionally, our data show that, as opposed to other described HdIV genes, rep genes are little transcribed in hemocytes, thus suggesting that they are not directly associated with the disruption of the immune response but rather involved in other physiological alterations of the infected lepidopteran larva.

The present study was conducted to determine egg and larval parasitoids of the beet armyworm Spodoptera exigua Hübner (Lepidoptera: Noctuidae), which is an important but sporadic pest in Turkey. High beet armyworm population levels were recently observed in fields of first and second crop maize in the southeast Mediterranean region of Turkey. The parasitoid species complex and its impact on the pest were analyzed in a 4-year study in first and second crop maize. The braconid larval parasitoids Microplitis rufiventris Kokujev, M. tuberculifer Wesmael, Meteorus ictericus Nees, Chelonus obscuratus (Herrich Schäffer) (an egg-larval parasitoid), Apanteles ruficrus (Haliday); the ichneumonid larval parasitoids Hyposoter didymator (Thunberg) and Sinophorus xanthostomus Gravenhorst; and the egg parasitoid Trichogramma evanescens (Westwood) were found to be the natural enemies attacking the pest. Among the parasitoid species the solitary endoparasitoid H. didymator was the most prevalent species, being reared from 40.5% of the parasitized larvae found. Higher parasitism rates were recorded on first crop than on second crop maize in every year. Possible reasons for this difference in larval parasitism between two growing seasons include lower population of the pest and reduced insecticide applications in first crop maize fields which permitted higher parasitism. However, parasitoid activity was insufficient to counterbalance the population growth of the pest on subsequent second crop maize.