Apronopa haeselbarthi van Achterberg, 1980, by original designation (Figs 2, 3).

Figure 2. 

Apronopa haeselbarthi van Achterberg, 1980 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 3. 

Apronopa haeselbarthi van Achterberg, 1980 (holotype, female) A mesonotum, dorsal view B propodeum C legs, metasoma and ovipositor, lateral view D metasomal tergites, dorsal view D fore wing.

Holotype (Apronopa haeselbarthi) Germany: • ♀, Dransfeld, B/L 2.vi.1966 (Haeselbarth leg.) (ZSSM). Paratypes (Apronopa haeselbarthi) Germany: • 1 ♀, 1 ♂, Schotten, Hessen, Fi., Streu, v.1967 (Haeselbarth leg.) (♀ in RMNH, ♂ in ZSSM).

Mandible small, simple, robust, tridentate. Paraclypeal fovea short, remaining far from inner margin of eyes. Mesoscutum without medio-posterior pit; notauli absent in posterior half of mesoscutum; precoxal sulcus always present; propodeum smooth or with different types of sculpture and sometimes with longitudinal or transverse carinae. Marginal cell of fore wing never shortened; vein r originating approximately from basal quarter of pterostigma; vein 2-SR always present and distinctly sclerotized; veins m-cu and cu-a distinctly postfurcal; first subdiscal cell always closed postero-apically by CU1a vein. Metasoma of ♀ more or less distinctly compressed laterally. First metasomal tergite without dorsope; second tergite often longitudinally striate medially. Ovipositor sheath not longer than metasoma.

This is a small genus with only three described species exclusively from the Palaearctic region (two of these species have an East Palaearctic distribution). Unfortunately, there is no data about its biology. Apronopa is characterised by three distinct diagnostic characters (van Achterberg 1980; Peris-Felipo and Belokobylskij 2018b): the dorsope of the first metasomal tergite are absent, the ovipositor has a distinct dorsal nodus subapically and the second metasomal tergite is sculptured basally (except in A. levis Papp, 2007). The combination of these features is unknown in other Aspilotina and supports well the separate generic status of this taxon.

Alysia ruficornis Nees von Esenbeck, 1834, by monotypy.

Mandible small, simple, tridentate, often with upper (first) tooth diminished with respect to lower (third) tooth. Paraclypeal fovea large, reaching inner margin of eyes. Mesoscutum with or without medio-posterior pit; notauli present only in anterior part of mesoscutum; precoxal sulcus almost always present; propodeum smooth or more common with different types of sculpture and sometimes with longitudinal and/or transverse carinae, rarely forming areas. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR often present and usually distinctly sclerotised but absent in subgenus Eusynaldis; veins m-cu and cu-a postfurcal; first subdiscal cell always closed postero-apically by vein CU1a. In hind wing, subbasal cell usually closed. Metasoma of ♀ more or less distinctly compressed laterally; second tergite always smooth. Ovipositor sheath usually not longer than metasoma.

Members of the genus Aspilota are frequently encountered as they are one of the most common genera among Alysiini wasps. It is mainly distributed in forested and humid areas of the Holarctic region and only a few species have been already recorded from other zoogeographic regions. This genus is undersampled in the tropics where their main hosts (Phoridae) have the greatest diversity.

Aspilota species are koinobiont endoparasitoids of larvae, mainly of the family Phoridae (Diptera). Previous reports established Aspilota as parasitoid of the families Anthomyiidae, Lonchaeidae, Muscidae, Platypezidae, Sarcophagidae, Syrphidae, and Tephritidae. However, these hosts need to be especially reconfirmed. The records of lepidopterous and hymenopterous larvae as hosts (families Erebidae, Bucculatricidae, Lasiocampidae, and Tortricidae, and family Tenthredinidae, respectively) are extremely doubtful because the known biology and perhaps were concerned to the Phoridae living in dead larvae of the species from these families.

The genus Aspilota contains three subgenera, Aspilota sensu stricto, Eusynaldis Zaykov & Fischer, 1982, and Grandilota Fischer, 2002.

Alysia ruficornis Nees von Esenbeck, 1834, by monotypy.

Numerous species from the Palaearctic, Nearctic, and Neotropical regions were reviewed (e.g., Peris-Felipo and Belokobylskj 2014; Peris-Felipo et al. 2016a, 2016b, 2016c, 2016d).

Figure 4. 

Aspilota (Aspilota) ajara Peris-Felipo, 2016 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

This largest and easily recognised subgenus includes most of Aspilota species. Pterusa ruficollis (Stelfox & Graham, 1950) is returned to Aspilota as A. ruficollis Stelfox & Graham, 1950, comb. nov. after the revision of type because the paraclypeal fovea are wide and reaching the inner margin of the eyes. Its new generic position is also supported by Fischer’s re-description of the species (Fischer 1972: 436).

Figure 5. 

Aspilota (Aspilota) ajara Peris-Felipo, 2016 (holotype, female) A propodeum B first metasomal tergite, dorsal view C legs, metasoma and ovipositor, lateral view D fore and hind wings.

Eusynaldis varinervis Zaykov & Fischer, 1972, by monotypy (Figs 6, 7).

Figure 6. 

Aspilota (Eusynaldis) varinervis (Zaykov & Fischer, 1972) (holotype, male) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 7. 

Aspilota (Eusynaldis) varinervis (Zaykov & Fischer, 1972) (holotype, male) A propodeum B first metasomal tergite, dorsal view C hind leg, lateral view D fore wing.

Holotype Regetus balcanicus [= Aspilota (Eusynaldis) globipes] Former Yugoslavia: • ♀, Kosovo, Mts. Sar, Brezovica, 900–1200 m, 20–23.v.1971 (Papp & Hortatovich leg.) (HNHM) [Hym. Typ. No. Mus. Budapest 7878].

Subgenus Eusynaldis shares all diagnostic characters of Aspilota sensu stricto, except the absent vein 2-SR of the fore wing.

Regetus Papp, 1999 and Adelphenaldis Fischer, 2003 are junior synonyms of Eusynaldis Zaykov & Fischer, 1982 because both taxa are characterised by the same diagnostic features (Zhu et al. 2017). Moreover, the study of the holotype of Regetus balcanicus Papp, 1999 (the type species of Regetus) showed this species to be a junior synonym of Aspilota (Eusynaldis) globipes (Fischer, 1962) (Peris-Felipo and Belokobylskij 2019).

Grandilota kenyaensis Fischer, 2002, by original designation (Figs 8, 9).

Figure 8. 

Aspilota (Grandilota) kenyaensis (Fischer, 2002), comb. nov. (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 9. 

Aspilota (Grandilota) kenyaensis (Fischer, 2002), comb. nov. (holotype, female) A mesonotum, dorsal view B propodeum. C legs, metasoma and ovipositor, lateral view D first metasomal tergite, dorsal view E fore and hind wings.

Holotype (Grandilota kenyaensis). Kenya: • ♀, Mt. Elgon Natural Park, bamboo (Arundinaria alpine) thicket, 2740 m; swept. No. 496, 22.i.1992 (G. Varkonyl leg.) [Hym. Typ. No. Mus. Budapest 11673] (HNHM). Paratype (Grandilota kenyaensis) Kenya: • ♀, same label as holotype but [Hym. Typ. No. Mus. Budapest 11674] (HNHM).

Mandible well developed, tridentate, with upper (first) tooth diminished to respect to lower (third) tooth. Paraclypeal fovea long, reaching inner margin of eyes. Mesoscutum without mesoscutal pit; notauli present only in anterior half of mesoscutum; precoxal sulcus present; propodeum with pentagonal areola, delineated by carinae. In fore wing, marginal cell reaching apex of wing; vein r originating from basal quarter of pterostigma; vein 2-SR present and sclerotised; veins m-cu and cu-a postfurcal; first subdiscal cell closed postero-apically by CU1a vein. In hind wing, subbasal cell open. Metasoma of ♀ more or less distinctly compressed laterally. Ovipositor sheath not longer than metasoma.

This subgenus has only one known species, Aspilota (Grandilota) kenyaensis Fischer, 2002, from Kenya and shares the main characters with Aspilota sensu stricto, however, the subbasal cell of the hind wing open distally (absent vein cu-a) and the wing membrane is pigmented, which distinguishes it at the subgeneric level.

Dinostigma muesebecki Fischer, 1966, by monotypy (Figs 10, 11).

Figure 10. 

Dinostigma muesebecki Fischer, 1966 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 11. 

Dinostigma muesebecki Fischer, 1966 (holotype, male) A propodeum B legs, metasoma and ovipositor, lateral view C first metasomal tergite, dorsal view D fore and hind wings.

Holotype (Dinostigma muesebecki). United States Of America: • ♀, North East, Pa. [= Pennsylvania], No 9019, 6.vii.1912 (F. Johnson leg.) (NMNH).

Mandible small, simple, tridentate. Paraclypeal fovea short, far from reaching inner margin of eyes. Mesoscutum without mesoscutal pit; notauli present only in anterior part of mesoscutum; precoxal sulcus absent; propodeum always smooth; spiracles of propodeum large. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR absent; vein cu-a postfurcal; first subdiscal cell open distally (without vein 2-1A). Hind wing with all cells open. Metasoma of ♀ more or less distinctly compressed laterally. Ovipositor sheath shorter than metasoma.

After careful revision of former Dinostigma and Eudinostigma (as subgenus of Dinostigma) species, only the type species of this genus, Dinostigma muesebecki Fischer, 1966, is retained in Dinostigma. The species D. stenosoma (van Achterberg, 1988) is transferred to the genus Dinotrema as a type species of the new subgenus, Pseudoprosapha subgen. nov. (see below), because this species has the first subdiscal cell of fore wing closed, the pterostigma broad and wider than vein r length, and all cells of the hind wing closed (Dinotrema (P.) stenosoma (van Achterberg, 1988), comb. nov.).

This genus is very close to the Oriental-Afrotropical Lysodinotrema Fischer, 1995, because both of them share, among others, such main diagnostic cha­racters as simple tridentate mandible, short paraclypeal fovea, and mesoscutum without medio-posterior pit. However, the lack of closed cells in the hind wing in Dinostigma (present in Lysodinotrema), absence of vein 2-SR (present in Lysodinotrema), and absence of the precoxal sulcus (present in Lysodinotrema) are sufficient to separate both as different genera.

Dinotrema erythropa Foerster, 1863, by monotypy.

Mandible small, simple, tridentate, often with upper (first) tooth diminished with respect to lower (third) tooth. Paraclypeal fovea short, not reaching more than half distance between clypeus and inner margin of eyes. Mesoscutum with or without mesoscutal pit; notauli usually present only in anterior part of mesoscutum, although in some species of the subgenus Alitha it is rather well developed and reaching or almost reaching mesoscutal pit; precoxal sulcus usually present, propodeum with different types of sculpture and sometimes with longitudinal and/or transverse carinae, rarely entirely smooth. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR usually present and distinctly sclerotised or sometimes (subgenus Synaldis) absent or weakly developed and vein r not angled with vein 3-SR (van Achterberg 1988); veins m-cu and cu-a postfurcal; first subdiscal cell always closed postero-apically by CU1a vein. Venation of hind wing more or less reduced, sometimes without closed cells (Zhu et al. 2017). Metasoma of ♀ more or less distinctly compressed laterally. Ovipositor sheath usually not longer than metasoma.

Dinotrema is the most complicated and largest genus within the tribe Alysiini with more than 440 known species, predominantly occurring in the temperate climatic regions (Peris-Felipo and Belokobyslkij 2018a). However, after studying a large amount of type material from different regions it should be possible to present a new generic classification, including the following subgenera: Alitha Cameron, 1906, stat. nov. (with Carinthilota Fischer as a new synonym), Dinotrema sensu stricto, Prosapha Foerster, 1863, Pseudoprosapha Peris-Felipo subgen. nov. and Synaldis Foerster, 1863 (with Eudinostigma Tobias as a new synonym).

A revision of Eudinostigma Tobias species was carried out for this reclassification. After careful study of the type species of Eudinostigma we consider it a synonym of Dinotrema. However, depending on the presence or absence of vein 2-SR of the fore wing, its species are divided between the subgenera Dinotrema and Synaldis. The main diagnostic characters of Eudinostigma are as follows: distinctly depressed head (resulting in antennal sockets situated at the upper level of eye and maximum width of head in dorsal view 1.6–2.4× width of mesoscutum), compressed mesosoma, and vein 2-SR of fore wing often absent (Tobias 1986; van Achterberg 1988). These characters also occur sometimes in Dinotrema species, e.g., among others, in Dinotrema brevissimicorne (Stelfox et Graham, 1948), D. compressum (Haliday, 1838), D. parapunctatum (Fischer, 1976), and D. robertoi Peris-Felipo, 2013.

The following species previously belonging to Eudinostigma are transfered to the subgenus Dinotrema sensu stricto: D. (D.) alox (van Achterberg, 1988), comb. nov.; D. (D.) entabeniense (Fischer, 2009), comb. nov.; D. (D.) latum (Chen & Wu, 1994), comb. nov.; D. (D.) planiceps (Fischer, Tormos & Pardo, 2006), comb. nov. and D. (D.) subpulvinatum (Fischer, 2009), comb. nov.. Moreover, four other Eudinostigma species are transferred to the subgenus Synaldis: D. (S.) bienesae (Fischer, Tormos & Pardo, 2006), comb. nov., D. (S.) fischeri (Tobias, 1986), comb. nov. (type species of Eudinostigma), D. (S.) latistigma (Fischer, 1962), comb. nov., and D. (S.) planiceps (Fischer, Tormos & Pardo, 2006), comb. nov.

Furthermore, after studying the types of Dinostigma and Eudinostigma species, we consider the features of Eudinostigma stenosoma van Achterberg, 1988 (see below) enough different to transfer it to a new subgenus Pseudoprosapha subgen. nov.: Dinotrema (Pseudoprosapha) stenosoma (van Achterberg, 1988), comb. nov.

In summary, five subgenera of the genus Dinotrema are recognised, Alitha Cameron, 1906, stat. nov., Dinotrema sensu stricto, Prosapha Foerster, 1863, Pseudoprosapha Peris-Felipo subgen. nov. and Synaldis Foerster, 1863.

Alitha longipennis Cameron, 1906, by monotypy (lost).

Holotype (Carinthilota parapsidalis) (Figs 12, 13) Austria: • ♀, Kärnten (88), 1 km O. Heft b(ei) Hüttenberg, 1000–1100 m, 25.viii.1973 (Fischer leg.) (NHMW). Holotype (Carinthilota vechti) The Netherlands: • ♀, Putten (Gld.), Malaise trap, 24–28.ix.1970 (J.v.d. Vecht leg.) (RMNH).

Figure 12. 

Dinotrema (Alitha) parapsidalis (Fischer, 1975), comb. nov. (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 13. 

Dinotrema (Alitha) parapsidalis (Fischer, 1975), comb. nov. (holotype, female) A propodeum B first metasomal tergite, dorsal view C fore and hind wings.

This subgenus has all main characters of Dinotrema sensu stricto but differs from it by having the notauli more or less complete posteriorly, reaching or almost reaching the mesoscutal pit.

Despite the loss of the type material of Alitha Cameron, 1906 described from South Africa (van Achterberg 1988) and thanks to the relative complete description of this genus given by Cameron (1906) and the additional comments by van Achterberg (1988), Alitha is considered a subgenus of Dinotrema (stat. nov.). Moreover, the genus Carinthilota Fischer, 1975 is considered a junior synonym of the subgenus Alitha (syn. nov.) because both share identical diagnostic characters. Unfortunately, so far only four Palaearctic and Oriental species are known and no studied specimens from the Afrotropical region.

The development of the notauli in Alitha species is highly variable: usually they are developed as rows of closely circular grooves more or less reaching the medio-posterior mesoscutal pit [Dinotrema (A.) longipennis (Cameron, 1906), comb. nov., D. (A.) parapsidalis (Fischer, 1975), comb. nov. and D. (A.) vechti (van Achterberg, 1988), comb. nov.] while the distal part of the notauli is more or less reduced in the two Eastern Palaearctic species [D. (A.) lada (Belokobylskij, 1998), comb. nov. and D. (A.) mavka (Belokobylskij, 1998), comb. nov.]. The variable development of the notauli supports our opinion that the presence of nearly complete notauli in several genera of Aspilota group (Dinotrema and Orthostigma) is an unsuitable generic character; at most it may be used provisionally at subgeneric level.

Dinotrema erythropum Foerster, 1863, by original designation (Figs 14, 15).

Figure 14. 

Dinotrema (Dinotrema) erythropum Foerster, 1863 (female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antennae E head, front view F head and mesonotum, dorsal view.

Figure 15. 

Dinotrema (Dinotrema) erythropum Foerster, 1863 (female) A propodeum, dorsal view B hind leg, lateral view C first metasomal tergite, dorsal view D metasoma and ovipositor, lateral view E fore and hind wings.

(Dinotrema (Dinotrema) erythropum): England: • 1 female (paratype of Aspilota praecipua) and 1 male (paratype id.), Coll. Marshall Cornwall, Botusfleming (HNHM). Denmark: • 1 female, E-Jylland, Frisenborg, 28.vii.1986 (Munk leg.) (NMA). Finland: • 1 female, Sa. Valkeala, 6772:483, 28.vii.1977 (Koponen leg.) (NMA); • 1 female, same locality but, 29.vii.1977 (NMA). Hungary: • 1 female, Ugod, Somberek Hubertlak-Kórnyéle, 26–29.vi.1967 (Papp leg.) (HNHM). Luxembourg: • 2 females, Tratten, b. Murau Stmk. Coll Fulmek, 14.viii.1942 and 13.x.1954 (NHMW). Netherlands: • 1 female, Wijster (Dr.) opposite Biological Station, 22–30.ix.1975 (van Achterberg leg.) (RMNH). Spain: • 1 female, Valencia, 16.vii.1942 (NHMW).

The main diagnostic characters of this subgenus are the short paraclypeal fovea which remain far from the eye margins, the pterostigma very narrow (linear) and vein 2-SR of fore wing present and more or less completely sclerotised.

This is the largest subgenus including the main part of Dinotrema species with approx. 440 species described worldwide (Peris-Felipo and Belokobyslkij 2018a). As shown by van Achterberg and Vikberg (2014), Pterusa Fischer, 1958 is a synonym of Dinotrema sensu stricto, because any differences are absent in females, and it is based only on the brachyptery of the males.

Alysia speculum Haliday, 1838, by original designation (Figs 16, 17).

Figure 16. 

Dinotrema (Prosapha) speculum (Haliday, 1838) (A, C, E: female; B, D, F: male) A, B habitus, lateral view C, D antenna E, F fore and hind wings.

Figure 17. 

Dinotrema (Prosapha) speculum (Haliday, 1838) (female) A head and mesosoma, lateral view B mandible C head and mesonotum, dorsal view D propodeum E first metasomal tergite, dorsal view F legs, metasoma and ovipositor, lateral view.

(Dinotrema (Prosapha) speculum): Austria: • ♂, Spitzzicken, Burgenland, 24.viii.1959 (Fischer leg.) (NHMW). Russia: • 8 ♂, Leningradskaya Province, Tolmachevo, 23.viii.1960 (Tobias leg.) (ZISP); • 2 ♂, Novgorod Province, 20 km NW of Pestovo, 6.vii.1986 (Tobias leg.) (ZISP); • ♀, ibid, 15.viii.1990 (ZISP); • ♀, ibid, 27.vii.1999 (ZISP); • ♀, ibid, 1.viii.1999 (ZISP); • ♀, ibid, 5.viii.2001 (ZISP); • ♀, Volgograd Province, 10 km S of Novokhopersk, 10.vii.1977 (Tobias leg.) (ZISP); • ♂, Krasnodar Territory, Sochi, Lazarevskoe, 30.v.1988 (Tobias leg.) (ZISP); • ♂, Chelyabinsk Province, Ilmenskiy Nature Reserve, 17.vii.1958 (Tobias leg.) (ZISP).

This subgenus shares the main characters of Dinotrema sensu stricto but differs by having, in the fore wing, the maximum width of pterostigma wider than vein r (especially in males) and vein 2-SR of fore wing always present.

This subgenus includes five Palaearctic species: D. (P.) comptum Tobias, 2003, D. (P.) pachysemoides Tobias, 2003, D. (P.) speculum (Haliday, 1838), D. (P.) tobiasi (Fischer, 1994) and D. (P.) ussuricum Tobias, 2007. The status of Prosapha has been variable for a long time. Foerster (1863) described this genus based on the distinctive large, cuneiform and heavily sclerotised pterostigma of the male. Van Achterberg (1988) and Tobias (2003b) considered Prosapha species inside of Dinotrema based on their morphological similarity and because Prosapha females possess a narrower pterostigma weakly se­parated from the metacarp (1-R1). Prosapha species can be differentiated from Pseudoprosapha subgen. nov. by the presence of vein 2-SR (which is absent in Pseudoprosapha).

Eudinostigma stenosoma van Achterberg, 1988 (Figs 18, 19).

Figure 18. 

Dinotrema (Pseudoprosapha) stenosoma (van Achterberg, 1988), comb. nov. (holotype, male) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 19. 

Dinotrema (Pseudoprosapha) stenosoma (van Achterberg, 1988), comb. nov. (holotype, male) A head and mesonotum, dorsal view B propodeum C legs and metasoma, lateral view D first metasomal tergite, dorsal view E fore and hind wings.

Holotype (Eudinostigma stenosoma) Hungary: • ♂, Budapest, Biró, 21.ix.1927, “226” (RMNH).

Mandible small, simple, tridentate, with upper (first) tooth diminished with respect to lower (third) tooth. Paraclypeal fovea short, length not more than half distance between clypeus and inner margin of eyes. Mesoscutum without medio-posterior pit; notauli present only in anterior part of mesoscutum; precoxal sulcus present; propodeum completely smooth. Marginal cell of fore wing never shortened; vein r originating from basal quarter of wide pterostigma; vein 2-SR absent; vein cu-a postfurcal; first subdiscal cell always closed postero-apically by vein CU1a. Hind wing with all cells closed. Metasoma of ♀ more or less distinctly compressed posteriorly. Ovipositor sheath not longer than metasoma.

This new subgenus includes only a single species, Dinotrema (Pseudoprosapha) stenosoma (van Achterberg, 1988), comb. nov.. This subgenus shares with Prosapha the comparatively broad pterostigma (viz., wider than the length of vein r) and in female vein r + 3-SR forming a (nearly) straight line but differs by the loss of vein 2-SR (present in Prosapha), the depressed head (antennal sockets situated near the upper level of the eyes), the strongly compressed mesosoma and the very narrow clypeus. These differences make it worth to name a different subgenus for it.

Bassus concolor Nees, 1812, by original designation (lost) (Figs 20, 21).

Figure 20. 

Dinotrema (Synaldis) concolor (Nees, 1812) (female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 21. 

Dinotrema (Synaldis) concolor (Nees, 1812) (female) A propodeum B legs, metasoma and ovipositor, lateral view C first metasomal tergite, dorsal view D fore and hind wings.

Dinotrema (Synaldis) concolor: Austria: • ♀, Mischerdorf, Burgenland, 5.ix.1956 (Fischer leg.) (NHMW); • ♀, Mischerdorf, Burgenland, 6.viii.1958 (Fischer leg.) (NHMW); • ♀, Spitzzicken, Burgenland, 5.ix.1956 (Fischer leg.) (NHMW). Hungary: • ♀, Nagyrákos, 6.vi.1985 (Rozner leg.) (HNHM). Slovakia: • ♀, Orosva Polhora, 25.vii.1988 (Podlussány leg.) (HNHM). Dinotrema (Synaldis) cracipes [= Pterusa cracipes]: Holotype: Austria: • ♂, Wimpassing, Nieder-Österreich (Leitha-Gebirge), 2.v.1915 (Fischer leg.) (NHMW).

The main characters of this subgenus are shared with Dinotrema sensu stricto, but it has vein 2-SR of fore wing absent.

The status of Synaldis has been uncertain for a long time. Van Achterberg (1988) revised the genera of the Aspilota group and first sy­nonymised this genus with the re-established Dinotrema based on the not enlarged paraclypeal fovea (the plesiomorphic state). As a result, the former Synaldis species were distributed among the genera Aspilota and Dinotrema according to the new used diagnostic feature, the size and position of the paraclypeal fovea. For some time, the synonymy of Synaldis was not accep­ted by several experts working on alysiine taxa (Fischer 1993a, 1993b; Papp 2001; Belokobylskij 2002; Peris-Felipo et al. 2014a). It is necessary to underline that the apomorphic feature of the subgenus, the complete reduction of vein 2-SR of the fore wing, is a peculiar evolutionary event which also resulted in the disappearance of the distinct break (corner) between veins r and 3-SR and this part is only gently and relatively widely curved. Such an apomorphic state of the wing venation might represent an important qualitative transformation and could support at least a subgeneric status of Synaldis (Belokobylskij 2002; Peris-Felipo and Belokobylskij 2014, 2017). However, the intermediate state is also known, both with the presence of non-sclerotised vein 2-SR and vein r not angled with vein 3-SR (e.g., D. (D.) pulvinatum (Stelfox & Graham) as depicted by van Achterberg 1988) or vein 2-SR entirely absent (e.g., D. (S.) cespitator (Belokobylskij, 2004), comb. nov.), D. (S.) perfidum (Fischer, 1970), comb. nov. (as depicted by Fischer 1970) and D. (S.) trematosum (Fischer, 1967), comb. nov. (as depicted by Fischer 1967) with vein r weakly angled with 3-SR). The variation of vein 2-SR from entirely absent to entirely present and non-sclerotised vein is aptly shown in D. (D.) concinnum (Haliday, 1838). Therefore, we agree with Zhu et al. (2017) to recognise Synaldis as a subgenus for convenience. Its position as separate genus likely will render the genus Dinotrema paraphyletic, and the loss of vein 2-SR occurred probably more than once in Dinotrema and is variable within some taxa as illustrated by D. concinnum (König 1972) and the type species of the genus Synaldotrema (Belokobylskij and Tobias 2002).

Aspilota dentifemur Stelfox, 1943, by original designation (Figs 22, 23).

Figure 22. 

Leptotrema dentifemur (Stelfox, 1943) (female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 23. 

Leptotrema dentifemur (Stelfox, 1943) (female) A propodeum B fore femur, lateral view C legs, metasoma and ovipositor, lateral view D first metasomal tergite, dorsal view E fore and hind wings.

(Leptotrema dentifemur) Denmark: • ♀, Stegelykke VG, 8–15.vii.1991 (Munk leg.) (PFEC). The Netherlands: • ♀, Putten (Gld.), Malaise trap, 26.ix–2.x.1973 (J.v.d. Vecht leg.) (RMNH).

Mandible small, simple, tridentate, often with upper (first) tooth diminished with respect to lower (third) tooth. Paraclypeal fovea short, not reaching more than half distance between clypeus and inner margin of eyes. Mesoscutum with or without mesoscutal pit; notauli usually present only in anterior part of mesoscutum; precoxal sulcus usually present, propodeum with different types of sculpture and sometimes with longitudinal and/or transverse carinae, rarely entirely smooth. Fore femur has a distinct apomorphic character, viz., the presence of a large obtuse tooth (flange) or two or three small teeth. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR usually present and distinctly sclerotised; veins m-cu and cu-a postfurcal; first subdiscal cell always closed postero-apically by vein CU1a. Metasoma of ♀ more or less distinctly compressed laterally. Ovipositor sheath usually not longer than metasoma.

Only three rare Leptotrema species are known from the Palaearctic, Oriental and Australasian regions, L. bovefemora (Bhat, 1979), L. dentifemur (Stelfox, 1943) and L. wilhelmense Braet & van Achterberg, 2014. Wharton (2002) treated Leptotrema only as subgenus of Dinotrema, but some other experts (Fischer 2002; Belokobylskij and Tobias 2007; Braet and van Achterberg 2014) preferred to consider it as a valid genus based on its unique apomorphic character: the presence of the ventral tooth or teeth of the fore tibia. The study based on the main morphological characters show that Leptotrema deserves generic status.

Lysodinotrema madli Fischer, 1995, by original designation (Figs 24, 25).

Figure 24. 

Lysodinotrema madli Fischer, 1995 (holotype, female) A habitus, lateral view B head, lateral view C mandible D antenna E head, front view F mesosoma, lateral view.

Figure 25. 

Lysodinotrema madli Fischer, 1995 (holotype, female) A mesonotum, dorsal view B propodeum C hind leg, lateral view D metasoma and ovipositor, lateral view E first metasomal tergite, dorsal view E fore and hind wings.

Holotype (Lysodinotrema madli) Madagascar: • ♀, Ste. Marie Flues Manandriana, 14–25.xi.1994 (Fischer leg.) (NHMW).

Mandible small, simple, tridentate. Paraclypeal fovea short, remaining far from the inner margin of eyes. Mesoscutum without mesoscutal pit; notauli present only in anterior half of mesoscutum; precoxal sulcus present; propodeum mainly sculptured, without areola. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR present and sclerotised; veins m-cu and cu-a always postfurcal; first subdiscal cell completely open posteriorly and without vein 2-1A. Hind wing without closed cells. Metasoma of ♀ more or less distinctly compressed. Ovipositor sheath shorter than metasoma.

This rare genus, with only three described species from the tro­pical areas (L. madli Fischer, 1995, L. minimum Fischer, 2004, L. sarawakense Fischer, 1995), is considered to be related with Dinostigma Fischer, 1966, because of sharing the open first subdiscal cell in the fore wing. However, the combination of closed cells in the hind wing (absent in Dinostigma), presence of vein 2-SR of fore wing (absent in Dinostigma) and of the precoxal sulcus (absent in Dinostigma) makes it possible to maintain Lysodinotrema as a valid genus.

Panerema inops Foerster, 1863, by original designation (lost) (Figs 26, 27).

Figure 26. 

Panerema inops Foerster, 1863, comb. nov. (female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 27. 

Panerema inops Foerster, 1863, comb. nov. (female) A propodeum B first metasomal tergite, dorsal view C legs, metasoma and ovipositor, lateral view D fore wing.

(Panerema inops): Germany: • ♀, Zaarensee, Seggenwiese, 29.vi.1998 (v. Broen leg.) (PFEC). The Netherlands: • ♀, Cadier, 5.v.1975 (B. v. Aartsen leg.) (RMNH).

Mandible small, simple, tridentate, often with upper (first) tooth diminished with respect to lower (third) tooth. Paraclypeal fovea short, not reaching more than half distance between clypeus and inner margin of eyes. Third antennal segment distinctly elongated. Mesoscutum with or without mesoscutal pit; notauli usually present only in anterior part of mesoscutum. Scutellum with a transverse crenulate depression subposteriorly. Females are brachypterous with strongly reduced wings (commonly in this group of genera males are brachypterous but females macropterous). The preserved distal anterior veins in such wing are distinctly thickened, with veins r and 2-SR of the fore wing absent but hind wing with closed cells (van Achterberg 1988; Belokobylskij and Kula 2012). Metasoma of ♀ more or less distinctly compressed laterally. Ovipositor sheath usually not longer than metasoma.

During many years, Panerema was considered as a valid genus (van Achterberg 1988; Fischer 2002; Belokobylskij and Tobias 2007; Belokobylskij and Kula 2012). As shown by van Achterberg (1988) despite its uncertain position of this taxon it has two synapomorphies, viz., the scutellum has a transverse crenulate depression subposteriorly and the third antennal segment is at least 1.5× longer than the fourth segment. The value of both characters is uncertain (although perhaps apomorphic), but the diagnostic character study carried out shows that Panerema deserves the status of genus due to its distance from other genera.

Dinotrema (Synaldotrema) speciosum Belokobylskij & Tobias, 2002, by original designation (Figs 28, 29).

Figure 28. 

Synaldotrema speciosum Belokobylskij & Tobias, 2002, comb. nov. (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 29. 

Synaldotrema speciosum Belokobylskij & Tobias, 2002, comb. nov. (holotype, female) A propodeum B legs, metasoma and ovipositor, lateral view C first metasomal tergite, dorsal view D fore and hind wings.

Holotype (Dinotrema (Synaldotrema) speciosum) Russia: • ♀, Primorskiy kray, 10 km SSW of Partizansk, border of forest, 12–13.vii.1996 (S. Belokobylskij) (ZISP). Paratypes (Dinotrema (Synaldotrema) speciosum) Russia: • ♀, Primorskiy kray, 50 km N of Olga, mixed forest, 29.vii.1979 (S. Belokobylskij) (ZISP); • ♀, Primorskiy kray, Pogranichnyi District, Barabash-Levada, forest, 3–6.vi.1980 (S. Belokobylskij) (ZISP); • ♀, Primorskiy kray, 42 km S of Plastun, forest, 24.vi.1979 (S. Belokobylskij) (ZISP); • ♀, Republic of Tuva, 14 km E of Kyzyl, lowland of Ka Khem River, 31.v.1975 (D. Kasparyan) (ZISP).

Synaldotrema Belokobylskij & Tobias shares the main characters of Dinotrema sensu stricto but differs by having the hypopygium of the female strongly retracted under the posterior tergites of metasoma and the fourth tergite strongly elongated, ~ 2.5× longer than fifth tergite (vs approximately of equal length in Dinotrema s. str.).

The type species of Synaldotrema (D. speciosum Belokobylskij & Tobias, 2002) has a variable vein 2-SR of the fore wing. This vein is usually present, but sometimes, mostly discoloured and its posterior half or sometimes entire vein 2-SR is absent (Belokobylskij and Tobias 2002). Previous studies showed that the value of the reduction of this vein illustrates well the subgeneric division (Belokobylskij and Tobias 2002), however the current diagnostic character study proved that retraction of hypopygium has enough value to consider Synaldotrema as a valid genus.

Cubitalostigma reichli Fischer, 1998, by monotypy (Figs 30, 31).

Figure 30. 

Cubitalostigma reichli Fischer, 1998 (holotype, female) A habitus, lateral view B head, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 31. 

Cubitalostigma reichli Fischer, 1998 (holotype, female) A mesosoma, lateral view B mesonotum, dorsal view C propodeum D legs, metasoma and ovipositor, lateral view E first metasomal tergite, dorsal view D fore and hind wings.

Holotype (Cubitalostigma reichli) Indonesia: • ♀, Sumatra, Aceh, Gunung Leuser Nat. Pk., Ketambe Res. Sta., 1° rainforest, Mature forest, Terrace 4 closed canopy, 400 m, 3°41'N, 97°29'E, Malaise trap W/pans, 1–30.xi.1989 (D.C. Darling leg.) (NHMW).

Mandible small, simple, tridentate, with upper (first) tooth diminished with respect to lower (third) tooth, with complete submedial transverse curved carina. Paraclypeal fovea short, remaining far removed from edge of eyes. Mesoscutum without mesoscutal pit; notauli present only in anterior part of mesoscutum; precoxal sulcus always absent; propodeum smooth. In fore wing, marginal cell never shortened; vein r originating from almost middle of pterostigma; first submarginal cell very reduced; second submarginal cell widened; vein 2-SR present and well sclerotised; first subdiscal cell closed postero-apically by CU1a vein. Subbasal cell of hind wing minute but closed. Metasoma of ♀ distinctly compressed laterally. Ovipositor sheath shorter than metasoma.

This is a peculiar monotypic genus with only the type species known from Indonesia. Cubitalostigma is characterised by the very aberrant venation of the fore wing, with vein r arising almost from the middle of the very narrow pterostigma, very far from its basal part. This is a unique character within the subtribe Aspilotina.

Neorthostigma eoum Belokobylskij, 1998 (= Aspilota macrops Stelfox & Graham, 1951), by original designation (Figs 32, 33) [synonymised by Peris-Felipo et al. 2020].

Figure 32. 

Neorthostigma macrops (Stelfox & Graham, 1951) (A, B: female, holotype of A. macrops; C–F: female, holotype of Neorthostigma eoum) A, C habitus, lateral view B habitus, dorsal view D head and mesosoma, lateral view E mandible F antenna.

Figure 33. 

Neorthostigma macrops (Stelfox & Graham, 1951) (female, holotype of Neorthostigma eoum) A head, front view B head and mesonotum, dorsal view C propodeum, dorsal view D first metasomal tergite, dorsal view E hind leg, metasoma and ovipositor, lateral view F fore and hind wings.

Holotype (Aspilota macrops) Ireland: • ♀, Sligo, S. shore of Lough Gill near Doonee Rock, 15.x.1937 (AWS leg.) [USNM #76022; USNMENT01569377] (NMNH). Holotype (Neorthostigma eoum (= macrops)) Russia: • ♀, Primorskiy kray, Anisimovka, forest, glades, 16.viii.1979 (S. Belokobylskij leg.) (ZISP). Paratypes (Neorthostigma eoum (= macrops)) Russia: • 2 ♀, Primorskiy kray, Spassk-Dal’niy, forest, glades, 16 and 22–23.viii.1995 (S. Belokobylskij leg.) (ZISP); • 1 ♀, Sakhalin Island, 10 km W of Aniva, mixed forest, 15.viii.1981 (S. Belokobylskij leg.) (ZISP). Japan: • 1 ♂, Fukuoka, Nogochi, Fukuoka-shi, 28.viii.1992 (V. Makarkin leg.) (ZISP).

Norway: • 1 ♀, Oslo [AK], Maridalen, Dausjøen, Spruce forest, 5.vi–16.x.2010, 60.01234 N 10.787665 E, 160 m, Malaise trap, river outlet (Lars Ove Hansen leg.) (NHMO). Russia. Leningradskaya Province: • 1 ♀, Tolmachevo, mixed forest, 22.viii.1960 (V. Tobias leg), “Aspilota macrops Stelf., Tobias det. 1961” (ZISP). Primorskiy kray: • 1 ♀, 30 km E of Spassk-Dal’niy, forest, glades, 4.vi.1984 (S. Belokobylskij leg.) (ZISP); • 1 ♀, Nadezhdinskiy District, 15 km SSW of Nezhino, forest, 16–18.vii.1993 (S. Belokobylskij leg.) (ZISP); • 1 ♀, 30 km SE of Ussuriysk, forest, border of forest, 12–17.vii.2001 (S. Belokobylskij leg.) (ZISP); • 1 ♀, Vladivostok, Okeanskaya, forest, 25.vii.2001 (S. Belokobylskij leg.) (ZISP); • 1 ♀, Vladivostok, Sedanka, forest, border of forest, 30.vii.2001 (S. Belokobylskij leg.) (ZISP).

Mandible small, tridentate, with very small and screwed upper tooth, with complete transverse and curved submedian carina. Paraclypeal fovea long, reaching or almost reaching inner margin of eyes. Mesoscutum always without mesoscutal pit; notauli present only in anterior half of mesoscutum; precoxal sulcus always developed; propodeum with wide and rather distinctly delineated by carina areola and with different types of sculpture but sometimes almost smooth. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR always developed; veins m-cu and cu-a postfurcal; first subdiscal cell always closed postero-apically by vein CU1a. Metasoma of ♀ more or less distinctly compressed. Ovipo­sitor sheath shorter than metasoma.

Five described species are known: one from the Palaearctic region (widely distributed from Ireland to Japan), two from Papua New Guinea (Peris-Felipo et al. 2020) and two from Brazil (Dias de Oliveira and Penteado-Dias 2023). This genus is closely related to Orthostigma according to its specialised mandibles. Wharton (2002) treated Neorthostigma as a subgenus of Orthostigma. However, the combination of such important diagnostic characters such as the absence of eye-antennal socket sulcus, the large paraclypeal fovea reaching or almost reaching inner margin of eye, the usual absence of mesoscutal pit, the face and sometimes the mesoscutum entirely covered by dense setosity distinctly indicate a separate position of this taxon at the genus level (Belokobylskij et al. 2019; Peris-Felipo et al. 2020). Its hosts are still unknown.

Aphidius flavipes Ratzeburg, 1844: 71, by monotypy.

Delocarpa Foerster, 1863; Ischnocarpa Foerster, 1863; Patrisaspilota Fischer, 1995; Africostigma Fischer, 1995; Whartonstigma Peris-Felipo, 2020.

Mandible small, tridentate and with a wide ventral lobe as third tooth, with complete submedial transverse curved carina. Paraclypeal fovea short, far distant from inner margin of eyes. First flagellar segment usually longer or sometimes as long as second flagellar segment (slightly shorter in subgenus Africostigma). Mesoscutum usually with mesoscutal pit; notauli often present only in anterior part of mesoscutum, but in subgenus Patrisaspilota notauli almost reaching mesoscutal pit; precoxal sulcus always present; propodeum usually with different types of sculpture and sometimes with longitudinal or transverse carinae, rarely almost smooth. In fore wing, marginal cell never shortened; vein r originating from basal quarter of pterostigma; vein 2-SR usually distinctly sclerotised (but absent in subgenus Whartonstigma); veins m-cu and cu-a postfurcal; first subdiscal cell always closed postero-apically by vein CU1a. Metasoma of ♀ more or less distinctly laterally compressed. Ovipositor sheath usually not longer than metasoma.

This genus includes more than 60 described species and is easily separated from other genera in the Aspilota group by the presence of the peculiar structure of mandible with complete transverse and curved submedial carina and usually wide lobe-shaped third tooth.

Currently four subgenera are recognised within this genus, Africostigma Fischer, 1995, Orthostigma sensu stricto, Patrisaspilota Fischer, 1995, and Whartonstigma Peris-Felipo, 2020 (Peris-Felipo and Belokobylskij 2020).

Orthostigma (Africostigma) karkloofense Fischer, 1995, by original designation (Figs 34, 35).

Figure 34. 

Orthostigma (Africostigma) karkloofense Fischer, 1995 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 35. 

Orthostigma (Africostigma) karkloofense Fischer, 1995 (holotype, female) A mesonotum, dorsal view B propodeum C legs, metasoma and ovipositor, lateral view D first metasomal tergite, dorsal view E fore and hind wings.

Holotype (Orthostigma (Africostigma) karkloofense) South Africa: • ♀, Howick, Natal, Karkloof Forest, 19.ix.1963 (Haeselbarth leg.) (ZSSM).

This Afrotropical subgenus includes two species from South Africa and shares the main diagnostic characters with Orthostigma but differs from all other subgenera by having the first flagellar segment of antenna shorter than the second one.

Aphidius flavipes Ratzebrug, 1844, by monotypy.

Several species from Palaearctic region were studied. For example:

– Orthostigma beyarslani Fischer, 1995: Spain: • ♀, Alicante, Torrevieja, Natural Park of Lagunas de la Mata-Torrevieja, 25.v.2004 (ENV).

– Orthostigma laticeps (Thompson, 1895): Spain: • ♀, Alicante, Alcoi, Natural Park of Carrascal de La Font Roja, 20.v.2004 (ENV).

– Orthostigma maculipes (Haliday, 1838): Spain: • ♀, Castellon, Pobla de Benifassà, Natural Park of Tinença de Benifassà, 26.ix.2005 (ENV).

– Orthostigma mandibulare Tobias, 1962: Russia: Holotype: • ♀, Leningradskaya oblast’. Tolmachevo, border of forest near floodplain of Ostrovenka River, 19.viii.1960, Tobias [leg] (ZISP).

– Orthostigma pumilum (Nees, 1834): Spain: • ♀, Castellon, Pobla de Benifassà, Natural Park of Tinença de Benifassà, 17.vi.2004 (ENV).

– Orthostigma sculpturatum (Tobias, 1962): Spain: • ♀, Castellon, Pobla de Benifassà, Natural Park of Tinença de Benifassà, 28.viii.2006 (ENV).

Main characters for the subgenus Orthostigma are the long first flagellar segment (longer than second segment), the notauli only anteriorly present on the mesoscutum and fore wing with vein 2-SR present and more or less distinctly sclerotised.

Figure 36. 

Orthostigma (Orthostigma) mandibulare (Tobias, 1962) (female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and messonotum, dorsal view.

Figure 37. 

Orthostigma (Orthostigma) mandibulare (Tobias, 1962) (female) A propodeum, and metasomal tergites, dorsal view B legs, metasoma and ovipositor, lateral view C fore and hind wings.

This is the largest subgenus with about 60 known species from the Holarctic, Oriental, and Australasian regions.

Patrisaspilota memoranda Fischer, 1995 (= Orthostigma multicarinatum Tobias, 1990) by original designation (Figs 38, 39). Synonymised by Peris-Felipo et al. 2019.

Figure 38. 

Orthostigma (Patrisaspilota) multicarinatum Tobias, 1990 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head, dorsal view.

Figure 39. 

Orthostigma (Patrisaspilota) multicarinatum Tobias, 1990 (holotype, female) A mesonotum, dorsal view B propodeum C first metasomal tergite D legs, metasoma and ovipositor, lateral view E fore and hind wings.

Holotype (Orthostigma (Patrisaspilota) multicarinatum) Vietnam: • ♀, Bathuok, 125 km W of Thanh Hoa, prov. Thanh Hoa, 26.i.1989 (B. Korotyaev leg.) (ZISP). Paratype (Orthostigma (Patrisaspilota) multicarinatum) Vietnam: • ♀, Vietnam, Tram Lap, 20 km N of Buon Luoi, prov. Gia Lai – Con Tum, 6.xii.1988 (Sharkov leg.) (ZISP).

This subgenus shares the main characters of Orthostigma but has the notauli almost reaching the mesoscutal pit.

The four Oriental species of this subgenus share the almost completely developed notauli as in Dinotrema (Alitha). The notauli consist of a row of closely located large points more or less reaching the mesoscutal pit. The presence of such type of notauli in different genera of the subtribe Aspilotina (Orthostigma and Dinotrema) is obviously a parallelism and perhaps indicates the limited value of the character, viz., at least as subgeneric character.

Orthostigma gallowagi Wharton, 2002, by original designation (Figs 40, 41).

Figure 40. 

Orthostigma (Whartonstigma) gallowagi Wharton 2002 (holotype, female) A habitus, lateral view B head and mesosoma, lateral view C mandible D antenna E head, front view F head and mesonotum, dorsal view.

Figure 41. 

Orthostigma (Whartonstigma) gallowagi Wharton 2002 (holotype, female) A propodeum B legs, metasoma and ovipositor, lateral view C first metasomal tergite D fore and hind wings.

Holotype (Orthostigma (Whartonstigma) gallowagi): Australia: • ♀, Queensland, Wongabel S. F., 6 km S of Atherton, 12.xi–1.xii.1983, Storey and Brown. M.T. (QMBA). Paratypes (Orthostigma (Whartonstigma) gallowagi): Australia: • 1 ♀, 1 ♂, same data as holotype [No.111581] (ANIC).

Very similar to the subgenus Orthostigma sensu stricto but differs from it by the absence of vein 2-SR of the fore wing.

This recently described subgenus includes four species from Australia and Papua New Guinea (Peris-Felipo and Belokobylskij 2020).