Trentino-South Tyrol: Schwangart 1913, 1918; Stellwaag 1921.

Species of temperate-cold zones of the Palearctic region, widespread in Europe (excluding the Balkan Peninsula), Russia, Turkey, eastwards to Korea and Japan (Yu et al. 2012). From Europe it has been introduced in Canada from 1956 to 1958 and from 1979 to 1981 to control the winter moth Opheroptera brumata (Linnaeus, 1758) (Lepidoptera, Geometridae) and has been reported as being established there (Carlson 1979, Barron 1989).

Yu et al. (2012) list 58 species, mostly belonging to the Lepidopteran family Geometridae (27 species). Further records include Tortricidae (8 species), Yponomeutidae (7), Noctuidae (5), Lasiocampidae, Nolidae (2 species each), Pyralidae, Thyatiridae, Notodontidae, Lycaenidae, Erebidae (Lymantria dispar (Linnaeus, 1758)), and Diprionidae (HymenopteraSymphyta) (1 species each).

This larval-pupal koinobiont endoparasitoid emerged from overwintering pupae of L. botrana and Eupoecilia ambiguella (Hübner, 1796) (Stellwaag 1928, Thompson 1957) in Italy (South Tyrol), France and Germany and Sparganothis pilleriana (Denis & Schiffermüller, 1775) (LepidopteraTortricidae) in France (Stellwaag 1928). These hosts are not included in the list of Yu et al. (2012). Another species of the same genus, Agrypon anxium (Wesmael, 1849), is reported on the first generation larvae of EGVM in Switzerland and France (Moreau et al. 2010), while A. minutum (Bridgman and Fitch, 1894) has been obtained by E. ambiguella in France and Switzerland and by S. pilleriana in Germany (Villemant et al. 2011).

After its introduction in Canada, A. flaveolatum was confused by some American authors (Carlson 1979, Dasch 1984) with similar Nearctic species belonging to the same genus (Barron 1989).

Trentino-South Tyrol: Silvestri 1912.

This species is widespread over most of the temperate Holarctic region. It is quite common in the Nearctics (Yu et al. 2012), while in the Palaearctics its distribution is limited to the Western part only: Europe, Central Russia and Turkey (Yu et al. 2012, Zwakhals and van Achterberg 2017).

Yu et al. (2012) list 38 hosts for this species, 21 of which belonging to the family Tortricidae, the remaining, in order of importance, to the families Pyralidae (4 species), Gelechiidae (3), Noctuidae (2), Choreutidae, Galacticidae, Geometridae, Lycaenidae, Nolidae, Sesiidae and Psychidae (all represented by a single species). Some economically important species are included, as the Oriental fruit moth Grapholita molesta (Busck, 1916) and the codling moth Cydia pomonella (Linnaeus, 1758), Argyrotaenia spp., Rhyacionia spp., Choristoneura spp. (Tortricidae), and the European corn borer, Ostrinia nubilalis (Hübner, 1796) (Crambidae). The report of Andricus kollari (Hartig, 1843) (HymenopteraCynipidae) of Fulmek (1968) as host of P. geniculata is not reliable.

In Italy, the species is reported on Paranthrene tabaniformis (Rottemburg, 1775) (LepidopteraSesiidae) (Dasch 1984, without indication of locality), on Gypsonoma aceriana (Duponchel, 1843) (LepidopteraTortricidae) (Haeselbarth 1989) and L. botrana (Silvestri 1912). The latter is not included in the list of Yu et al. (2012) and represents the only record for this species on EGVM.

Silvestri (1912) reared a single male of this species from overwintering pupae of EGVM in San Michele (Trentino) in May.

Parania geniculata is one of the smallest European species of the subfamily Anomaloninae. It parasitizes mostly Tortricidae (Schnee 2008). The species was originally described by Holmgren (1857) as Anomalon geniculatum and subsequently transferred to the genus Atrometus Förster, 1869 by Thomson (1892). Silvestri (1912) has erroneously attributed the authorship of the species to Förster, who just described the genus Atrometus (Förster 1869). Then Townes (1971) transferred the species to Parania Morley, 1913, with P. geniculata as the only European species of this small, but widely distributed genus. Probably because of this nomenclatural inexactness, Hoffmann and Michl (2003) have misinterpreted the specimen obtained by Silvestri and put it in synonymy with Sinophorus geniculatus Gravenhorst, 1829, which belongs to the subfamily Campopleginae. The specimen figured by Silvestri (1912, fig. XXXIX, Figure 1A) clearly belongs to Anomaloninae. After comparing the figure by Silvestri with specimens of Atrometus insignis Förster, 1878 (a South European species that could be confused with Parania), and Parania geniculata in his collection, Heinz Schnee recognized the specimens depicted by Silvestri as P. geniculata, for the following reasons: “…small number of flagellomeres, small brachial cell, mesoscutum and scutellum somewhat longer, and slender hind tarsi”; on the contrary, “…Atrometus insignis is therefore out of the question, because in the drawing the characteristic transverse furrow on posterior part of the mesoscutum is absent and the brachial cell is too small. Also the hind tarsi of P. geniculata are much thinned, while they are strongly thickened in males of A. insignis. Moreover, the hosts of A. insignis are Zygaena spp. (LepidopteraZygaenidae) and other host assignments are very likely wrong (Schnee in litteris)”. We have searched for the specimen identified by Silvestri without finding it.

Figure 1. 

Parania geniculata (Holmgren, 1857), male (IT, Udine, Aug. 28, 1985, leg. Allegro ex Gypsonoma aceriana; ZSM) A drawing of male in a dorsal vision by Silvestri (1912)B habitus, lateral view C head and anterior part of thorax, dorsal view D head, frontal view E wings F head and mesosoma lateral view.

Piedmont: Colombera et al. 2001.

Tuscany: Scaramozzino et al. 2017b.

Palaearctic species occurring throughout Europe, Canary Islands, Near East (Turkey and Syria), Northern and Central Russia, Kazakhstan, Russian Far East, Korea and Japan (Yu et al. 2012; Zwakhals and van Achterberg 2017). In 1931, it was released in North America (New Jersey, USA) to control the Oriental fruit moth, without becoming established (Carlson 1979).

Females lay eggs on young larvae that live hidden in the vegetation. Yu et al. (2012) list 28 host species, many of which belong to the family Tortricidae, including the fruit crop pests Cydia pomonella (Linnaeus, 1758), Archips rosana (Linnaeus, 1758) and Grapholita molesta (Busck, 1916). The following species need to be added to the list: Aphelia (Zelotherses) peramplana (Hübner, 1825) (= A. amplana Hübner, 1813) (Vas et al. 2015), Archips crataegana (Hübner, 1799) (Lungu-Constantineanu 2009), Cacoecimorpha pronubana (Hübner, 1799) (Scaramozzino et al. 2017b), Cydia pactolana (Zeller, 1840) (Jansons 2013), Eudemis profundana (Denis & Schiffermüller, 1775) (Zaemdzhikova 2017), Sparganothis pilleriana (Denis & Schiffermüller, 1775) (Habermehl 1922), Spilonota ocellana (Denis & Schiffermüller, 1775) (Nuzhna 2010) among the Tortricidae; Anacampsis populella (Clerck, 1759) (Hassanein 1978) among Gelechiidae; Macrothylacia rubi (Linnaeus, 1758) (Nuzhna 2010) among Lasiocampidae; Acrobasis suavella (Zincken, 1818) (Morley 1915) among Pyralidae; and Prismosticta fenestrata Butler, 1880 among Endromidae (Morley 1915). Habermehl (1922) reports as host also Gelechia boticella, but the identity of this species still remains unclear.

Ultimately, 40 host species of Trichomma enecator have been reported: 25 species belonging to Tortricidae, three each to Gelechiidae and Pyralidae, and one each to Elachistidae, Endromidae, Erebidae (Lymantria dispar (Linnaeus, 1758)), Gracillariidae, Lasiocampidae, Noctuidae, Nolidae and Psychidae. The record by Starke (1956 in Yu et al. 2012) of Plioreocepta poeciloptera (Schrank, 1776) (DipteraTephritidae) seems unlikely and is probably wrong.

In Italy, T. enecator is reported on: Earias clorana (Linnaeus, 1761) (LepidopteraNolidae) on goat willow (Salix caprea Linnaeus, 1753) (Leonardi 1925); Cydia pomonella in Campania (Sciarra 1915) and Emilia Romagna (Faggioli 1938); Tortrix viridana Linnaeus, 1758 and Aleimma loeflingiana (Linnaeus, 1758) (LepidopteraTortricidae) on oak in Calabria and Campania, respectively (Silvestri 1923); Grapholita molesta in Emilia Romagna (Grandi 1937); Rhyacionia buoliana (Denis & Schiffermüller, 1775) in Tuscany (Zocchi 1952); Cacoecimorpha pronubana feeding on spurge flax (Daphne gnidium Linnaeus, 1753) in Tuscany (Scaramozzino et al. 2017b).

Trichomma enecator is a solitary, koinobiont, larval-pupal endoparasitoid on fruit-mining or other concealed lepidopterous larvae. It is one of the most common parasitoids of the codling moth in Europe (Franck et al. 2017). Although quite common, its control action on the codling moth is limited, with parasitization rates rarely exceeding 5% (Table 4), being inexplicably absent in some apple orchards (Maalouly et al. 2013).

What we know about its biology is mainly due to Rosenberg (1934), who studied the codling moth in French apple orchards. This parasitoid attacks the host larvae, hibernating in the larval stage inside the host; the adult emerges from the pupa, some weeks before the host; the emergence period, in outdoor insectary, lasts 10–24 days, from the second half of May to the beginning of June. In captivity both genders may live approximately a month. Females start to oviposit one or two days after emergence, and their eggs hatch in approximately eight days (Rosenberg 1934). Despite its presence in most of the areas in Rosenberg’s survey, the parasitism rate never surpassed 3.11%.

Trichomma enecator females parasitize all the larval instars of the codling moth inside the fruits. The females are attracted by exudates that accumulate on the surface of the fruits infested by the codling moth larvae; in the absence of these exudates, the parasitization behavior is disrupted (Mills and Dixon 1993). To breed this species in insectarium is very difficult (Mills 2005), even if Russ and Faber (1978) were able to rear it until F6 generation by the same method used to rear Ascogaster quadridentatus Wesmael, 1845 (Hymenoptera, Braconidae).

At our latitude, T. enecator is a multivoltine species, while in Central-Northern Europe (Gauld and Mitchell 1977, Sedivy 2001) and Spain (Miñarro and Dapena 2004) it is presumably bivoltine. According to our personal observations, in Italy it shows three generations per year, and can attack all three generations of EGVM.

Trichomma enecator has a secondary importance on EGVM; Telenga (1934) obtained it in early June in Crimea. In Piedmont (Italy), a single specimen emerged from pupae of the overwintering generation of L. botrana (Colombera et al. 2001). In the Natural Reserve of San Rossore (Pisa, Tuscany), we obtained 13 specimens of T. enecator from EGVM pupae in July 2012 and from EGVM and C. pronubana pupae in May and July 2014. Pupae of both tortricids were collected into the nests formed by the larva on the apical buds of the spurge flax Daphne gnidium (Malvales, Thymelaeaceae) (Scaramozzino et al. 2017b).

Figure 2. 

Trichomma enecator (Rossi, 1790) (IT, San Rossore Pisa, ex Lobesia botrana) A female habitus lateral view B male habitus lateral view C female head frontal view D female head and mesosoma dorsal view E female head and mesosoma lateral view.

Piedmont: Pinna et al. 1989; Lozzia and Rigamonti 1991.

The cosmopolitan genus Campoplex Gravenhorst, 1829 includes ca. 210 species of koinobiont endoparasitoids of microlepidopteran larvae (mainly of the Tortricidae family, but also of Coleophoridae, Gelechiidae, Pyralidae etc.), and to a lesser extent of macrolepidoptera, HymenopteraSymphyta and few ColeopteraCurculionidae (see Yu et al. 2012 for a more comprehensive review). In Europe approximately 90 species are recorded (Zwakhals and van Achterberg 2017).

Pinna et al. (1989) obtained an unidentified species of Campoplex from overwintering specimens of L. botrana, with parasitism rates variable from 4.7 to 36.9%. Lozzia and Rigamonti (1991) found an unidentified species from overwintering generation of the EGVM, with a parasitism rate close to 4%.

Apulia: Nuzzaci and Triggiani 1982 [on Daphne gnidium L.].

The species is widely spread throughout Europe and Northern Russia (Yu et al. 2012; Zwakhals and van Achterberg 2017). In 1937, 96 adults of C. borealis were released in U.S.A. (Connecticut) from Europe, to control the European pine shoot moth, Rhyacionia buoliana (Denis & Schiffermüller, 1775) (LepidopteraTortricidae), but the species has not become established (Dowden 1962, Bartlett et al. 1978).

Yu et al. (2012) report 19 host species associated with C. borealis, belonging to the families Coleophoridae (8 species), Tortricidae (4 species, L. botrana is not into the list), Psychidae, Yponomeutidae, Eriocraniidae, Gelechiidae, Gracillariidae, Simaethidae (one species each) and on the weevil Anthonomus pomorum (Linnaeus, 1758) (ColeopteraCurculionidae).

In three years of sampling on Daphne gnidium, Nuzzaci and Triggiani (1982) obtained three specimens of this parasitoid from larvae of L. botrana (identified by K. Horstmann). This is the only record on EGVM so far.

Campoplex borealis is the species that gives its name to a “borealis” species-group of the genus Campoplex (Horstmann 2012). With the name of C. borealis were indicated at least six different species (Horstmann 2012), morphologically very similar and mainly characterized by their host preferences. Right now, eight species are included in this species-group: C. borealis (Zetterstedt, 1838), C. jaeckhi (Bauer, 1936), C. psammae (Morley, 1915), C. punctipleuris Horstmann, 1980, C. serratellae Horstmann, 2012, C. caloptiliae Horstmann, 2013, C. tussilaginis Horstmann, 2013 and Campoplex linosyridellae Horstmann, 2016. They are mainly related to Coleophoridae and Gelechiidae; one species, C. caloptiliae, lives on Gracillariidae, while a second species close to C. psammae lives on Psychidae (Horstmann 1980, 2012, 2013, Shaw et al. 2016). Horstmann (1985) does not mention EGVM among the hosts of C. borealis, despite a male and a female collected by Nuzzaci and Triggiani are in his collection in ZSM.

Figure 3. 

Campoplex borealis (Zetterstedt, 1838) (female: SP, Barcelona, March 28, 1964; male: CH, Sustenpaß, September 10, 1989, ex Acleris variegana; ZSM) A female habitus lateral view B male habitus lateral view C female head lateral view D female head frontal view E female head dorsal view F female propodeum dorsal view G female metasoma dorsal view H opened cocoon.

Trentino-South Tyrol: Catoni 1910, 1914; Silvestri 1912; Schwangart 1913, 1918; Ruschka and Fulmek 1915.

Veneto: Marchesini and Dalla Montà 1992, 1994, 1998; Dalla Montà et al. 1993; Marchesini et al. 2006; Marchesini 2007.

Piedmont: Colombera et al. 2001.

Tuscany: Dalla Montà et al. 1993; Bagnoli and Lucchi 2006; Lucchi and Santini 2011; Loni et al. 2016; Scaramozzino et al. 2017b.

Campania: Silvestri 1912.

Sicily: Alcamo (TP), ex Lobesia botrana on grapes (ZSM new record).

Campoplex capitator is a Mediterranean species, occurring in the Iberian Peninsula, France, Corsica, Italy, Switzerland and Turkey (Yu et al. 2012; Zwakhals and van Achterberg 2017). It is widespread in most of the southern European wine-growing areas (Bagnoli and Lucchi 2006), although its presence on EGVM in Southern Italy was not definitely ascertained. Nuzzaci and Triggiani (1982), in Apulia, underline the presence of C. difformis on EGVM feeding on Daphne gnidium and the absence of C. capitator, as already stated by Silvestri (1912). When checking his collection in Naples, we found two series of specimens, both reported as C. difformis from L. botrana. Actually, the two series are composed of at least three different species:

1. Campoplex capitator from Portici (Naples), 5 females and 3 males, and from San Michele all’Adige (Trento), 3 females, 3 males and 1 individual without metasoma. The specimens from San Michele all’Adige have the same origin of those studied and published by Catoni (1910) with the name of C. difformis and identified by O. von Schmiedeknecht.

2. Diadegma stigmatellae Horstmann, 1980 (Campopleginae), 6 males and 4 females from Portici, a parasitoid of Gracillariidae (Shaw and Horstmann 1997).

3. Pristomerus vulnerator (Panzer, 1799) (Cremastinae) 1 male and 1 female from Portici.

We are not sure if the two series of specimens correspond to those actually studied by Silvestri but we think that the Campoplex specimens he had attributed to Omorgus difformis belong to C. capitator.

In the Horstmann collection, as well as in the general collection of ZSM, we found 7 females and 6 males of C. capitator from Sicily (Alcamo, TP), emerged from larvae of L. botrana feeding on grapes in July 2007, August 2009 and late May-June 2010. Also in the Horstmann collection we examined a male and a female of C. capitator from Piacenza (Northern Italy), obtained from E. ambiguella.

Campoplex capitator seems to have an extremely limited host range. It was discovered on EGVM for the first time by Coscollá (1980) in Spain. Yu et al. (2012) list only two host species, L. botrana and Ancylis mitterbacheriana (Denis & Schiffermüller, 1775) (LepidopteraTortricidae). According to Villemant et al. (2011), in French vineyards C. capitator lives mainly at the expenses of L. botrana and E. ambiguella, though it has been obtained occasionally also from S. pilleriana. All the mentioned hosts live mainly on the grapevine, with the exception of A. mitterbacheriana, an univoltine leaf folder which lives on deciduous woodlands and whose larvae feed on the leaves of beech, common hornbeam, oaks, and sweet chestnut (Alford 2012, Brown et al. 2008).

Campoplex capitator is a solitary koinobiont larval endoparasitoid. Its development is strongly synchronized with L. botrana: both species overwinter in the same places, and live in close association, the first at the expense of the larvae of all the moth generations. The female oviposits into the body of EGVM larvae of 2nd-4th instar (Thiéry 2008, Villemant et al. 2011). Endophagous larva kills the host after spinning its own cocoon inside the moth cocoon. The larva of C. capitator builds a delicate elongated semi-transparent cocoon characterized by rounded poles, white color and a thin median opaque transverse line (Figure 4D).

Figure 4. 

Campoplex capitator Aubert, 1960 (female and male: IT, San Rossore Pisa ex Lobesia botrana) A female habitus lateral view B male habitus lateral view C propodeum female, dorsal view D opened cocoon inside a EGVM cocoon, in a dried nest on Daphne gnidium E distal 24 articles of female antenna.

The parasitization rates recorded in Europe (Italy excluded) are shown in Table 5, while those recorded in Italy are shown in Table 6.

Silvestri (1912) frequently found C. capitator on EGVM, both in Trentino (Northern Italy), in spring, and in Portici (Naples), from July to September. In Veneto (Northern Italy) it attacks all the generations of EGVM, with irregular and not particularly high rate of parasitism, often less than 1%, sometimes close to 12% in the first generation and 14% in the second and slightly more than 8% in the third generation (Marchesini and Dalla Montà 1994, Marchesini et al. 2006); sometimes it is absent. In Piedmont, where EGVM developed 2 generations per year, Colombera et al. (2001) recorded parasitization rates of 7.4% and 5.9%, respectively. In Tuscany (Central Italy), on grapevine, C. capitator is the most frequent species among larval parasitoids, showing a good parasitic activity throughout the region, mostly on larvae of the first two generations of the year (Bagnoli and Lucchi 2006). In the Natural Reserve of San Rossore (Pisa, Tuscany), it is very frequent on Daphne gnidium, where it represents the dominant species in the parasitoid community of EGVM; attacking larvae of all three generations, it contributed for more than 58% of the total number of parasitoids found in 2014 and more than 73% in 2015, with an overall annual parasitization rate for 2014 next to 10% (Loni et al. 2016).

In France, the rates of parasitism can be very high, especially in the EGVM first generation (Villemant et al. 2011). In the vineyards of Valencia (Spain), C. capitator is the only larval parasitoid that plays a significant role in the control of EGVM, even if the total parasitism levels found in that region were low across all the three generations (Coscollá 1997). In Douro Wine Region (Portugal), C. capitator is the second most abundant parasitoid of EGVM (Carlos et al. 2013), representing the 11.8% of parasitoids obtained in 10-year surveys. In Turkey, the species is mostly widespread in the Aegean vineyards (Koclu et al. 2005, Özsemerci et al. 2016).

Despite being considered one of the possible candidates for use in the biological control of EGVM, the knowledge about its behavior and its development are too limited and still some difficulties have to be overcome to develop an efficient mass rearing in bio-factory (Bagnoli and Lucchi 2006). Nevertheless, a recent cooperation between Italian and Chilean entomologists seems very promising (Lucchi et al. 2017).

Horstmann (1985) divided the Western Palaearctic species of the genus Campoplex in four species-groups: melanostictus (including the spurius-group), continuus, discrepans and difformis species-groups; later, the C. borealis species-group was added (Horstmann 2012). Both Campoplex capitator and C. difformis belong to “difformis” species-group, which is characterized by occipital carina joining hypostomal carina at a right angle at the base of the mandibles (i.e., occipital carina turned outwards ventrally); slender body with apically compressed metasoma; hind tibia with the median outer part from yellowish red to reddish brown, and basal part not clearly brightened; ovipositor sheath relatively long (at least as long as the hind tibia). Horstmann (1985) describes the female of C. capitator as follows in the key: body size approx. 5 mm, face wider than long, temples behind the eyes narrow, the lines (as seen in profile from above) touching the outside of the eyes and temples usually intersecting in or behind the scuto-scutellar groove, antennal segments, in the last quarter, at least as long as wide, prepectal carina medially not significantly broader than ventrolaterally and not clearly notched, hind coxae black with hind femora predominantly red, ovipositor sheath 1.4–1.7 times as long as the hind tibia, second metasomal segment not more than 1.6 times as long as wide.

The females of C. difformis, which are very similar to those of C. capitator, in Horstmann’s keys are distinguished by: body size approx. 8 mm, face longer than wide, antennal segments, in the last quarter, much wider than long, and the area superomedia of propodeum wide, and not clearly separated from the area petiolaris; both areas are clearly depressed.

Villemant et al. (2011) pointed out that C. capitator has often been confused with C. difformis in the past and many reports of this species on EGVM should probably be related to C. capitator. The identifications of C. difformis made by Silvestri as well as those of Catoni are to be referred to C. capitator (see above). The record of Nuzzaci and Triggiani (see C. difformis), whose specimens were identified by Horstmann, has to be considered correct.

Molecular-based studies indicate that C. capitator could be conspecific of C. formosanae Horstmann, 2012, a species reared on Enarmonia formosana in Germany (Hunt and Kuhlmann 2007, Hunt et al. 2008). The species was identified as C. dubitator at first, but then recognized as a valid species by Horstmann (2012). Despite molecular differences between the two species were not significant (Hunt and Kuhlmann 2007, Hunt et al. 2008), laboratory tests showed that C. formosanae was unable to develop on EGVM larvae and small but constant morphological characters have been found that lead to consider C. formosanae as a distinct species from C. capitator (Hunt et al. 2008, Jenner et al. 2013).

Figure 5. 

Campoplex capitator Aubert, 1960 (female: IT, San Rossore Pisa ex Lobesia botrana) A wings B head frontal view C head and mesosoma dorsal view D mesopleuron with epicnemial and postpectal carinae, ventral view.

Apulia: Nuzzaci and Triggiani 1982 [on Daphne gnidium L.].

The species is present throughout Europe up to the Caucasus and Uzbekistan, the Canary Islands and Madeira, Tunisia and Greenland (Yu et al. 2012; Zwakhals and van Achterberg 2017).

Yu et al. (2012) list 64 host species belonging to 18 different families (15 of Lepidoptera and 3 of Hymenoptera). This long list has to be verified, because in the past the specific interpretation of C. difformis was rather uncertain (see taxonomic notes under C. capitator and Horstmann 1985). The most represented family is that of Tortricidae, with 35 species (including L. botrana and E. ambiguella). Tortricids could be actually the only hosts of C. difformis, because all known hosts of the “difformis” species-group belong to this family (Horstmann 1985). Archips podana (Scopoli, 1763) was the only host ascertained for this species in the work of Horstmann (1985). In Evenhuis and Vlug (1983), a hypothetical Campoplex difformis, so identified by Horstmann, is reported attacking three other tortricid species, Pandemis cerasana (Hübner, 1786), Adoxophyes orana (Fischer v. Röslerstamm, 1834) and Acleris rhombana (Denis & Schiffermüller, 1775).

Campoplex difformis is a koinobiont larval endoparasitoid often reported as a parasitoid of E. ambiguella in France (Voukassovitch 1924). Its larva kills the moth larva when it is ready to pupate, and weaves its own cocoon next to the host spoils (Marchal 1912, Voukassovitch 1924). In Apulia (Southern Italy), this species occurred frequently, showing a parasitism rate of approx. 4% on EGVM larvae feeding on Daphne gnidium (Nuzzaci and Triggiani 1982). The species has been reported on L. botrana in Austria, France, Germany, Russia, Spain and Bulgaria, as well as in Italy (Thompson 1957, Zapryanov and Stoeva 1982, Zapryanov 1985).

Campoplex difformis is itself the victim of Dibrachys microgastri (Bouché, 1834) (HymenopteraPteromalidae) (Faure and Zolotarewsky 1925, Zapryanov and Stoeva 1982), Perilampus tristis Mayr, 1905 (HymenopteraPerilampidae) (Thompson 1958) and Cirrospilus sp. (HymenopteraEulophidae) (Noyes 2017).

Campoplex difformis (Gmelin) was designated as the type species of the genus Campoplex Gravenhorst, 1829 by Westwood (1840). It also gives the name to a complicated group of very similar species, with morphological characteristics insufficient to allow a definitive identification (Jenner et al. 2013). In the past this species was mainly attributed to the genera Limneria Holmgren, 1859 and Omorgus Förster, 1869 (= Omorga Thomson, 1887). Unfortunately, the interpretation of the species Ichneumon difformis Gmelin, until the studies of K. Horstmann (1969, 1985), has been uncertain. The type of Gmelin was destroyed. Then, following the first interpretation of the species given by Gravenhorst (1829), Horstmann fixed the lectotypus of Limneria mutabilis Holmgren, in Holmgren’s collection in Stockholm as a neotypus of C. difformis (Horstmann 1969, 1985). Thus, C. mutabilis of Holmgren became a junior synonym of C. difformis, and C. difformissensu Holmgren (and Thomson) became Campoplex deficiens Gravenhorst, 1829. Therefore, the interpretation of the species given by Gravenhorst (1829) [and hence by Horstmann (1985)] differed from that of other taxonomists (mainly Holmgren, Thomson and Schmiedeknecht), who considered C. difformis and C. mutabilis two distinct species. For this reason Aubert (1971, 1974 and 1981), another leading authority in the ichneumonid taxonomy, rejected the neotypus fixed by Horstmann and created another typus in the collection of Thomson in Lund, in order to keep C. mutabilis as a separate species from C. difformis. Consequently, C. deficiens Gravenhorst became synonym of C. difformis (see Table 7). In this work we follow the interpretation of Gravenhorst (1829), Horstmann (1969, 1985), and Yu and Horstmann (1997). Campoplex difformis has three synonyms: Campoplex lineolatus Ratzeburg, 1844, Limneria mutabilis Holmgren, 1860 and Nepiera algerica Habermehl, 1922, and a variety with dark hind legs (var. obscuripes Greese, 1927).

Campania: Silvestri 1912.

Diadegma armillata is a Palaearctic widespread species. It is found throughout Europe, Middle East, Caucasus, Kazakhstan, China and Korea (Yu et al. 2012; Zwakhals and van Achterberg 2017).

It is an important koinobiont larval endoparasitoid of macro- and microlepidoptera (Arctiidae, Argyresthiidae, Choreutidae, Coleophoridae, Gelechiidae, Geometridae, Gracillariidae, Lymantriidae, Noctuidae, Pieridae, Plutellidae, Psychidae, Pterophoridae, Pyralidae, Simaethidae, Tortricidae, Yponomeutidae). Yu et al. (2012) list 57 host species. Further three species have to be added to the list: Celastrina argiolus (Linnaeus, 1758) (Lycaenidae), Acrobasis marmorea (Haworth, 1811) (Pyralidae) and Swammerdamia caesiella (Hübner, 1796) (Yponomeutidae) (Shaw et al. 2016).

Diadegma armillata is particularly active against various Yponomeuta spp., attacking crop fruits, and it was introduced in 1989–1991 from France to northwestern Washington (USA), to control the apple ermine moth, Yponomeuta malinellus (Zeller, 1838) without becoming established (Unruh et al. 2003).

Diadegma armillata is a multivoltine species. Silvestri (1912) obtained few specimens of this wasp from EGVM cocoons in Portici (Naples). The record of Silvestri remains the only one for this species on EGVM.

The first serious attempt to bring order in the existing confusion for the interpretation of the European species of the genus Diadegma Förster, 1869 is due to the efforts of Horstmann (1969), who revised many types of the described species. The identifications of most species were based on poor morphological characters and are unfortunately unreliable (Horstmann 1969).

Diadegma armillata belongs to the subgenus Nythobia Förster, 1869, which includes group species with the seventh metasomal tergite deeply notched medially, the ovipositor sheath longer than the first metasomal tergite and shorter than the hind tibia (Horstmann 1969). The female of D. armillata is distinguished from the related species by a head with strongly narrowed temples; last article of the antennae longer than wide; propodeum with area superomedia wider than long and opened posteriorly, costulae strong; petiolar area slightly sunken and transversely striated; first metasomal segment with slightly protruding spiracles and postpetiole with parallel sides; ovipositor sheath approx. twice the length of the first metasomal segment; front and middle coxae yellow, the middle ones sometimes darkened at the base; femora and tibiae reddish yellow; hind tibia externally dark brown with white-yellow spots at the base and in the middle; metasoma variably stained with red (Horstmann 1969).

Some doubts regarding the distribution and host range of D. armillata arises from the fact that D. semiclausum (Hellen, 1949), a common parasitoid of the diamondback moth Plutella xylostella (Linnaeus, 1758), has been misidentified with D. tibialis (Gravenhorst, 1829), which is currently a synonym of D. armillata (Horstmann 1969, Azidah et al. 2000). Under the name tibialis, D. semiclausum was introduced in 1951 from Italy to Australia to control the diamondback moth (Oatman 1978).

Figure 6. 

Diadegma armillata (Gravenhorst, 1829) (female: D, Ostfriesischen Inseln, Mollum Memmert, July 24–30, 1986, leg. V. Haeseler; male: IT, South Tyrol, Kaltern, July 29, 1959; ZSM) A female habitus lateral view B male habitus lateral view C female head dorsal view D female head frontal view E female tergites 7–9 of metasoma with ovipositor and ovipositor sheath, lateral view F female ovipositor with subapical dorsal notch lateral view.

Trentino-South Tyrol: Catoni 1910, 1914; Schwangart 1913.

A Palaearctic species ranging from Europe up to Mongolia; in Europe it is widely present with the exception of the Iberian Peninsula, Balkans and Greece. In North Africa it is reported only from Tunisia (Yu et al. 2012; Zwakhals and van Achterberg 2017).

Diadegma tenuipes is a solitary koinobiont larval endoparasitoid of a dozen of hosts, which belong to the Lepidoptera families Coleophoridae, Momphidae, Pieridae, Plutellidae, Psychidae, Tortricidae and to the Hymenoptera families Tenthredinidae and Braconidae (Yu et al. 2012). Among these, some hosts of economic importance are indicated as the diamondback moth, the Oriental fruit moth (G. molesta), the European grape berry moth (E. ambiguella) and EGVM.

The only Italian records of D. tenuipes on EGVM are those of Catoni (1910 and 1914). In the vineyards of southern Romania, Bărbuceanu and Jenser (2009) found this species, along with three other species of the same genus, attacking the overwintering generation of EGVM, with a rather low parasitization rates (0.8%). In Romania it is reported by Petcu (1978) living on E. ambiguella as well.

Like the previous species, D. tenuipes has been assigned by Horstmann (1969) to the subgenus Nythobia. It measures approx. 6 mm in length, with the head posteriorly narrowed, propodeum with evident costulae (anterior transverse carinae), and the area superomedia shorter than twice its width; areolet of the fore wing rather large and intercepted by the second recurrent vein (2m-cu) after the middle; mesopleuron with speculum almost smooth, very shiny, the area close to mesopleural suture finely dotted; the seventh metasomal tergite dorsally deeply notched, ovipositor sheath 0.8 times the length of the hind tibia and 1.4 times that of the first metasomal tergite; body black, wings with pterostigma light brown, fore coxae light, femora and tibiae reddish yellow, posterior tibiae dark behind the base and at the apex, sides of the third metasomal tergite stained with red (Horstmann 1969).

Tuscany: Del Guercio 1899.

The limited information provided by Del Guercio (1899) does not allow designation of the two species of Nemeritis to any of the parasitoids associated to EGVM. Though Leonardi (1925) included the work of Del Guercio in his bibliography, he did not quote these species, which are not mentioned by any other author.

The species of Nemeritis have been divided by Horstmann (1975) in four groups: caudatula- and elegans-group, which parasitizes Raphidioptera, macrocentra-group which parasitizes Coleoptera (Cleridae, Malachiidae) and Lepidoptera and lissonotoides-group for which no host records are available (Horstmann 1994). Even though most of the species seem to attack concealed hosts under the bark or in bark crevices (Horstmann 1975), few species of the macrocentra-group have been recorded on moth species of economic importance, like the Mediterranean flour moth (Ephestia kuehniella Zeller, 1879), the European grain moth (Nemapogon granella (Linnaeus, 1758)) or the strawberry fruitworm (Cnephasia longana (Haworth, 1811)) (Horstmann 1994; Yu et al. 2012).

In the past, the genus Nemeritis included species of other campoplegine genera like Campoplex, Cymodusa or Venturia (Thomson 1887, Schmiedeknecht 1909). It is possible that the two species cited as Nemeritis sp. by Del Guercio (1899) are actually Venturia canescens (Gravenhorst, 1829). Schmiedeknecht, who identified ichneumonids obtained by Del Guercio (1899, page 156), refers to Venturia canescens as Nemeritis canescens in his fundamental work on European ichneumonids (Schmiedeknecht 1909, page 1688). At least the general habitus and wing venation in the picture of the second species (Del Guercio 1899, page 156) fit with the general aspect of V. canescens.

Trentino-South Tyrol: Schwangart 1913, 1918; Stellwaag 1928.

Transpalaeartic species widespread in Europe, present in Caucasus and Central and Far Eastern Russia as well (Yu et al. 2012; Zwakhals and van Achterberg 2017). The report for India by Morley (1913) needs to be confirmed.

Yu et al. (2012) list 14 host species for S. crassifemur, including 2 Geometridae, 2 Pieridae, 3 Pyralidae, 3 Tortricidae (Lepidoptera) and 4 species of Pamphiliidae (Hymenoptera).

The very little information available on this species derives from Stellwaag (1928), who reports that it has been obtained by Schwangart and Catoni in April. Nevertheless, Catoni (1910, 1914, 1915), Leonardi (1925) and Boselli (1928) do not mention this species in their lists. Thompson (1946) quotes it in Austria based on an article of Schwangart (1918) who obtained it from EGVM and E. ambiguella in Trentino. Hoffmann and Michl (2003) do not cite it. The species was also obtained from EGVM in Bulgaria (Athanassov 1981, Zapryanov 1985).

Very likely the species referred as Eulimneria crassifemur Thomson by Schwangart (1918) and Stellwaag (1928) is Sinophorus turionum (Ratzeburg, 1844). Carlson (1979) writes: “…in literature published before the description of alkae in 1928 [Sinophorus alkae (Ellinger and Sachtleben, 1928) is a junior synonym of S. turionum] and in some literature for more than ten years thereafter, the species was misidentified as crassifemur (Thomson)”. Sanborne (1984), in his review of the world species of the genus Sinophorus Förster, 1869, indicates among the hosts of S. crassifemur, only the web-spinning larvae of Cephalcia sp. and Acantholyda sp. (HymenopteraPamphiliidae) on Pinus spp.

The indication of this species on EGVM is due to Thompson (1946) that found it in a compendium of Hymenoptera parasitoids of European corn borer of Chu and Hsia (1937) that, unfortunately, we were not able to examine. Thompson and Parker (1928), on the basis of a record by Paillot (1924), report this species under the name of Eulimneria crassifemur Thomson, both on L. botrana and on E. ambiguella.

The species is widely distributed throughout the Palearctic region, except North Africa (Yu et al. 2012). It has been introduced several times in the United States and Canada for the biological control of the European corn borer O. nubilalis (Bartlett et al. 1978, Carlson 1979) and the pine shoot borer, R. buoliana (Syme 1971, 1984, Carlson 1979) without being established (Sanborne 1984). Its presence in the Oriental region (India and Sri Lanka) (Townes et al. 1965, Yu and Horstmann 1997) has to be confirmed.

Yu et al. (2012) list 26 host species, belonging to nine families of Lepidoptera and one family of Hymenoptera. Tortricidae and Pyraloidea are the most represented. Five other lepidopteran host species need to be added: Anania hortulata (Linnaeus, 1758), Nascia cilialis (Hübner, 1796), Pyrausta aurata (Scopoli, 1763) (Crambidae), Gymnoscelis rufifasciata (Haworth, 1809) (Geometridae) and Acrobasis advenella (Zincken, 1818) (Pyralidae) (Shaw et al. 2016).

It is reported as one of the main parasitoids of the European corn borer in Europe (Thompson and Parker 1928) under the name Eulimneria crassifemur, and in the Northern part of Far East (Manchuria and North Korea) (Clark 1934) as Eulimneria alkae. In the case of L. botrana, it is certainly an occasional parasitoid of minor importance, perhaps a secondary adaptation to a host different from the usual ones. In the literature we found, besides those mentioned for Italy, scattered reports of his presence on EGVM, mostly under the name crassifemur in its various generic combinations: in Austria (Thompson and Parker 1930), France (Paillot 1924, Thompson and Parker 1928), Bulgaria (Athanassov 1981, Zapryanov 1985), Germany (Thompson 1946) and Spain (Thompson 1946, Coscollá 1997). Another species, S. costalis (Thomson, 1887), has been recorded on EGVM in Moldavian vineyards, Romania (Pisicá and Páişescu-Bărbuceanu 2002, Bărbuceanu and Jenser 2009).

Many authors dealt with this species, especially in relation to its main hosts, O. nubilalis and R. buoliana. The species has been often confused with Limnerium crassifemur (recte Sinophorus crassifemur) (Paillot 1924, Thompson and Parker 1928, Goidanich 1931, Carlson 1979), which, instead, is mainly related to species of the family Pamphiliidae (Hymenoptera) (Sanborne 1984; see notes on previous species). The name crassifemur was later corrected in alkae by Ellinger and Sachtleben (1928), which then turned out to be a synonym of Sinophorus turionum (Sanborne 1984). It is likely that the report of Thompson (1946) for Italy on EGVM, quoted by Schwangart (1918) and Stellwaag (1928), concerns the previous species (see above).