Still counting: new records, nomenclatural notes, and three new species of Phaeogenini (Hymenoptera, Ichneumonidae, Ichneumoninae) from the Afrotropical region

A synthesis of the Phaeogenini occurring in the Afrotropical region is provided. Three species are newly described: Centeterichneumon nambi Dal Pos, Diller & Di Giovanni sp. nov. from Uganda, Chauvinia ganota Claridge sp. nov. from Kenya, and Kibalus nonnaritae Dal Pos & Di Giovanni sp. nov. from Uganda. Heterischnus mfongosi Rousse & van Noort, 2013 is newly recorded for Kenya and Tanzania and the male of the species is diagnosed for the first time. Also, the female of Arearia oxymoron Rousse & van Noort, 2013 is diagnosed for the first time from one of the paratype localities. Lusius tenuissimus (Heinrich, 1938) and Chauvinia nyanga Rousse & van Noort, 2013 are recorded for the first time for Uganda and Kenya, respectively. In addition, new localities are given for Chauvinia nitida (Heinrich, 1938), Heterischnus olsoufieffi (Heinrich, 1938) and Hoplophaeogenes curticornis Heinrich, 1938. A new combination, Nesostenodontus mkomazi (Rousse & van Noort, 2013) comb. nov., is proposed to accommodate Heterischnus mkomazi. An updated key to the Afrotropical genera of Phaeogenini and keys to the Afrotropical species of the genera Arearia Seyrig, Centeterichneumon Heinrich, Chauvinia Heinrich, Heterischnus Heinrich, Hoplophaeogenes Heinrich, Kibalus Rousse, van 1 European Journal of Taxonomy 868: 1–71 ISSN 2118-9773 https://doi.org/10.5852/ejt.2023.868.2105 www.europeanjournaloftaxonomy.eu 2023 · Dal Pos D. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0). M o n o g r a p h urn:lsid:zoobank.org:pub:A09D1E2F-E2A8-44C6-B838-6BB1DCB3F657 Noort & Diller, and Lusius Tosquinet are provided. Updated online Lucid keys to genera and species are available from http://www.waspweb.org.

The biology of Phaeogenini is fairly homogeneous, with the wasps usually attacking various microlepidopteran pupae or prepupae (e.g., Gelechiidae Stainton, 1854, Plutellidae Guenée, 1845) (Selfa & Diller 1994;Diller & Shaw 2014;Yu et al. 2016;Broad et al. 2018). However, Shaw & Bennett (2001) reported an exception in the genus Colpognatus Wesmael, 1845, which attacks Crambidae (Lepidoptera Linnaeus, 1758 at the larval stage, subsequently inducing mummification of the hosts before emerging from the prepupa. As noted by Broad et al. (2018), even though some host associations are known and corroborated, the biology of most of the species is poorly understood.
nonnaritae Dal Pos & Di Giovanni sp. nov. from Uganda. New and interesting records are presented, accompanied by distributional maps. The first male of Heterischnus mfongosi Rousse & van Noort, 2013 and the first female of Arearia oxymoron Rousse & van Noort, 2013 are illustrated and diagnosed. The generic placement of Heterischnus mkomazi Rousse & van Noort, 2013 is discussed, and the new combination Nesostenodontus mkomazi (Rousse & van Noort, 2013) comb. nov. is proposed. Taxonomic and nomenclatural remarks are provided where necessary. A key to the Afrotropical genera is also provided and, for each genus treated, a key to the species is presented to aid identification. Images and online interactive Lucid identification keys to the Afrotropical Phaeogenini are available at WaspWeb (http://www.waspweb.org) (van Noort 2023).

Photographs
An OPTIKA SZM-2 dissecting stereo microscope was used for observation and study. All photographs besides those of Chauvinia ganota Claridge sp. nov. and Chauvinia nyanga Rousse & van Noort, 2013 were taken with a Canon Eos 7D, lens Canon MP-E 65 mm f/2.8 1-5 × Macro and Canon Macro Lens EF 100 mm, using Zerene Stacker software ver. 1.04 for the stacking. Images were enhanced using Photoshop ver. 23.2.2.
Photographs of Chauvinia ganota sp. nov. and Chauvinia nyanga were taken with a Canon 1200D body, a Canon EF-S 60 mm macro lens for habitus images and a Venus Optics Laowa 25 mm Ulta-Macro lens for higher magnification images. Image stacking was performed with Helicon Focus 7.
All images included in this paper, as well as additional images and online interactive keys to species are available on WaspWeb (http://www.waspweb.org) (van Noort 2023).

Identification keys
Lucid pathway and Lucid matrix keys were developed using Lucid Builder ver. 4.0.23. Character matrices were generated and edited using Microsoft Excel; matrices were then used as input into Lucid matrix key production ). The online interactive keys were produced using Lucid, meeting the requirements of publishing both static and dynamic interactive keys under an open access model ). These keys were illustrated using high quality annotated images, highlighting diagnostic characters. Online identification keys are presented in two different formats on WaspWeb: traditional static dichotomous keys where a choice needs to be made at each key couplet to continue, which are also presented as an interactive Lucid pathway (dichotomous) key; and Lucid matrix keys where relevant states from multiple character features can be selected independently until identification is achieved. For more information concerning Lucid keys visit http://www.lucidcentral.org.
The LIF3 file for the online Lucid matrix key to all Afrotropical species of Phaeogenini is provided as a supplementary file (Supp. file 1). Lucid Interchange Format ver. 3 (LIF3) files are XML based files that store all the Lucid3 key data, allowing exchange of the key with other key developers such as Intkey (DELTA) or MX. The provision of this LIF3 data set allows future workers to edit the key and to add newly described taxa. The data file for the published key that is stored on the publisher's website and in e-archives has the rights of 'first publication' identified by its bibliographical data, location, and citation ). The concept of publication, citation, preservation, and re-use of data files to interactive keys under the open access model is detailed in Penev et al. (2009Penev et al. ( , 2012.

Data of examined material
Label information for the type specimens is reported verbatim, using the following conventions: / = different lines; // = different labels; italic = handwriting. For non-type specimens, names of collecting localities have been standardized.

Treatment of taxa
The overall morphological terminology follows the Hymenoptera Anatomy Ontology (HAO, Yoder et al. 2010) and Broad et al. (2018).
For each genus, a comparative diagnosis is compiled based on the relevant literature, namely Heinrich (1938) and Rousse et al. (2013).
For each species, type information, material examined, and relevant comments are provided. Type localities are reported as they appeared in the original publication with the addition of the country of origin. Moreover, a complete list of synonyms is listed below the valid species-level name, together with the original combination as well as subsequent combinations. Unavailable names are identified in square brackets (as in Sforzi & Sommaggio 2021). For each name, an exhaustive list of the known references is provided with indications of their contribution.
The distribution for Madagascar follows the official division, which recognizes 23 regions ("faritra") instead of the former six provinces (INSTAT 2010).

Diagnosis
One of the key features that separates Arearia from all the other Afrotropical Phaeogenini is the presence of a deeply impressed sternaulus, extending beyond the mid-length of the mesopleuron. Moreover, the genus can be further distinguished by the combination of the following characters: (1) gastrocoeli and thyridia absent; (2) mandible bidentate; (3) malar space with subocular sulcus present and distinct; (4) areolet open (3rs-m absent). Seyrig (1952: 17) placed Arearia within the subfamily Cryptinae (tribe Stilpnini) due to the very long, deeply impressed sternaulus. Later on, it was transferred to Ichneumoninae by Townes (1971: 233), after examination of the type. So far, the genus occurs only in the Afrotropical region with two species already described: Arearia oxymoron Rousse & van Noort, 2013, in South Africa, and Arearia paradoxa Seyrig, 1952, in Madagascar (Rousse et al. 2013Yu et al. 2016). Rousse & van Noort, 2013 Fig. 1 Arearia oxymoron Rousse & van Noort in Rousse et al., 2013: 17-20 (original description, key).

Diagnosis of female
The female of the species is hereby diagnosed for the first time bearing the same label data as one of the male paratypes reported in the original description. Compared to the male, no major differences are Genus Centeterichneumon Heinrich, 1938Centeterichneumon Heinrich, 1938. Type species: Centeterichneumon denticoxatus Heinrich, 1938, by original designation and monotypy.

Diagnosis
According to Heinrich (1938: 22) the genus Centeterichneumon is characterized by having "Clypéus largement arrondi. Spiracules du propodéum ovales chez les grands exemplaires, ronds chez les petits. Hanches III avec une petite dent chez la ♀ [= Clypeus broadly rounded. Propodeum spiracles oval in large specimens, round in small ones. Coxae III with a small tooth in ♀]." However, Heinrich's (1938) diagnosis is insufficient to separate Centeterichneumon from the other Afrotropical genera. We hereby attempt a more in-depth diagnosis of the genus. Centeterichneumon can be distinguished from all the other genera by the following combination of characters: (1) bidentate mandibles (that sets Centeterichneumon apart from Lusius and Heterischnus); (2) presence of distinct gastrocoeli, even though superficial (that allows the separation from Arearia, Chauvinia, Dicaelotus, and Hoplophaeogenes); (3) non-modified hypostomal carina, meeting occipital carina at the base of mandible (modified in Aethecerus and meeting occipital carina above mandibular base in Tycherus); (4) 2 nd metasomal tergite roughly square (and not elongated as in Kibalus); (5) clypeus well-separated from face (epistomal sulcus present) (that allows a clear separation from Diadromus); (6) malar space less than 0.5 × as long as the mandible (different from Diadromus); (7) hind coxae of female with a ventral tooth or small carina.
Except for the original type series, no other records have been reported for the genus. Townes & Townes (1973) provided a catalogue to the species with no new information on the distribution, while Rousse et al. (2013) failed to include the taxon into their review of the Afrotropical Phaeogenini. The species hereby newly described represents the first record of the genus after its original conception and expands its distribution for the first time to mainland Africa (Figs 5A-B, 8A-B).
A more in-depth study of the MZPW collection will be required prior to the designation of any lectotypes; therefore, for the moment, the type specimens should be referred to as syntypes (ICZN 1999: article 73.2).

Type locality
Madagascar: Rogez (denticoxatus denticoxatus); Ankaratra, Kalambatritra, Ampandrandava (denticoxatus obscutatus). Heinrich, 1938, subspecies (MZPW). A-B. Centeterichneumon denticoxatus denticoxatus Heinrich, 1938 (Heinrich 1938) (Fig. 6A-B). Heinrich (1938: 130) differentiated the two subspecies based only on the different coloration of the metasoma. Centeterichneumon denticoxatus denticoxatus has the first three tergites completely reddishbrown, the 4 th progressively infuscating with an apical narrow white band, the 5 th anteriorly black and posteriorly white-banded, while the remaining tergites are completely white (   European Journal of Taxonomy 868: 1-71 (2023) Centeterichneumon denticoxatus obscutatus has the 1 st tergite and the anterior part of the 2 nd reddish brown, the posterior part of the 2 nd tergite infuscate and the 3 rd and 4 th tergites entirely black, the 5 th anteriorly black and apically white, the 6 th and 7 th completely white (Figs 2C-D, 4B). We deemed these different characters as mere colour variation that does not allow an unequivocal differentiation of the two subspecies either based on consistent morphological traits or on clear distributional patterns. Moreover, also Heinrich (1938: 130) was unsure on the exact position of the two groups of specimens (see Type series) and, therefore, because of the above reasons, the two subspecies are hereby synonymized: Centeterichneumon denticoxatus denticoxatus Heinrich, 1938 = Centeterichneumon denticoxatus obscutatus Heinrich, 1938 syn. nov. Heinrich, 1938, subspecies. A. Known occurrence records: C. denticoxatus denticoxatus (blue dot) and C. denticoxatus obscutatus (yellow dots). B. Regional distribution: C. denticoxatus denticoxatus (in blue) and C. denticoxatus obscutatus (in yellow).

Etymology
Named after the goddess Nambi, daughter of the sky god Gulu; she married Kintu, the first king of Uganda in the Baganda mythology (Lynch & Roberts 2010). Noun in apposition. Female, in very good condition, missing only the 2-5 tarsomeres and claws on the right hind leg.
Head (Figs 7C,8a). Strongly transverse in dorsal and frontal view, overall shining; face distinctly transverse, more than 3.5 × as wide as medially high, densely punctate, medio-apically protruding in a very distinct blunt tubercule right below the antennal sockets, median field present and delimited by carinae on the ventral part of the face; clypeus well separated from the rest of the face (epistomal sulcus distinct); clypeus sparsely punctuate, shining and lenticular; malar space short, about 0.3 × basal width of mandible; mandible departing from the horizontal plane, slightly bending downward, with sparse setiferous punctures in the central area and on the ventral margin, teeth rather stout with the ventral tooth shorter (about 0.5 ×) than the upper tooth and slightly bent downward; antennal socket slightly projecting into a small flange, connected laterally to the internal orbit by means of a blunt, almost indistinct carina; frons and vertex almost completely smooth, with sparse and superficial punctuation, and with short and sparse hairs; ocellar triangle wider than high, slightly elevated; occipital carina distinct and complete, meeting hypostomal carina at the base of the mandible; temples slightly rounded in dorsal view; gena, in lateral view, medially strongly inflated, with indistinct punctures; antenna filiform, not tapering, flagellum with 28 flagellomeres, the 2 nd flagellomere about 1.5 × as long as 1 st , preapical flagellomeres slightly longer than wide. . Dorsoventrally flattened, overall shining; pronotum irregularly striate on ventral margin, smooth centrally, and densely punctate dorsally, its ventral corner pointed; pronotal neck aciculate; pronotal collar centrally smooth and from densely to irregularly punctate on the side; epomia strong; propleuron finely punctate, with sparse hairs, projected into a blunt flange ventroapically; mesoscutum finely and moderately punctate, notauli short and distinct only anteriorly; scutellum flat, with only a few, very superficial punctures, carinated only at the base; mesopleuron from densely punctate to puncto-striate, speculum smooth; epicnemial carina slightly raised between fore coxae, laterally ending at anterior edge of mesopleuron; sternaulus distinct on anterior third, crenulate; posterior transverse carina of mesosternum complete, slightly raised between mid coxae; metapleuron punctate anteriorly and punctate-rugose apically and ventrally, juxtacoxal carina strong; propodeum rather long in lateral view, not gently sloping; anterior transverse carina absent, with area basalis and area superomedia not separated and transversely striated, area externa and area dentipara not separated, punctate basally, and transversely striated apically; area petiolaris and area postero-externa transversely striated; area spiracularis moderately punctate; area lateralis transversely striated.

MetasoMa (
CoLoration (Figs 7a-C, 8a). Head with face, clypeus, mandibles (except the black apical teeth), orbits all around the eyes, and genae pale yellow; black are: mandibular teeth, ocellar triangle, vertex, and temples; central part of the frons and the lower part of the temples orange-brown. Antenna with scape and pedicel entirely orange; flagellum with the first three basal segments orange-brown, the 4 th gradually infuscating, and the rest black with the exception of the brown ventral side of the apical segments (from the 12 th /13 th flagellomeres); white annulus present only on the dorsal side of the flagellum, from the 8 th to 14 th /15 th flagellomeres. Mesosoma entirely orange-yellow with preaxilla black and tegulae pale yellow, the overall ventral side is pale yellow while the dorsal sclerites are mostly orange. All legs are orange yellow, with fore and mid coxae and fore and mid trochanters pale yellow; basal part of hind femur, hind tibia, and hind tarsus infuscate. Wing entirely hyaline. Metasoma entirely orange, except for the 7 th tergite, which is dorsally brown with a white apical margin; ovipositor sheath deeply infuscate.
Variation (based on the two paratypes). White annulus on the flagellum starting from the 7 th flagellar segment in one paratype; mesoscutal suprahumeral sulcus black in both paratypes.

Male
Unknown.

Key to the species of Centerichneumon Heinrich, 1938
1. Head strongly transverse, face about 3.7 × as wide as medially high (width measured as the maximum distance of the inner orbits, height measured in the center from the edge of the clypeus to the level of the antennal sockets) (A); area superomedial not clearly separated from area basalis, forming an elongated continuous area (shaded area) (B); area externa and area dentipara not separate ( -Head not strongly transverse, face about 3.0 × as wide as medially high (a); area superomedia inverted heart-shaped (shaded area), weakly separated from area basalis (red arrow), and not European Journal of Taxonomy 868: 1-71 (2023) forming an elongated continuous area (b); area externa and area dentipara distinctly separated (anterior transverse carina clearly present) (b); mesoscutum convex in lateral view (Figs 2B, D, 5A, D); clypeus, face and frons pale white (a); metasoma with white posterior band on apical tergites ( (3) propodeal apophyses absent (present in Hoplophaeogenes); (4) notauli indistinct (deep in Arearia); (5) propodeum smooth, with area superomedia not separated from area basalis and area petiolaris, forming a single mid-longitudinal area (area superomedia clearly distinct in Dicaelotus). The genus can further be distinguished by: clypeus strongly transverse, its ventral margin sharp and more or less regularly rounded; flagellum of female enlarged from middle; temples moderately swollen behind eyes; occipital and hypostomal carinae joining above mandibular base; epomia present, moderately marked; notauli indistinct; propodeum elongate, in profile slightly and regularly rounded to uniformly sloping backwards in a single plane; posterior transverse carina interrupted in front of mid coxae; fore wing with areolet pentagonal, closed; hind wing with distal abscissa of CU present, unpigmented; tarsal claws simple; metasoma of female elongate to strongly elongate, ventral margins of apical tergites overlapping, hiding sternites; metasoma of male not so unusually modified; ovipositor sheath wide, barely extending beyond metasomal apex (Heinrich 1938;Rousse et al. 2013).

Remarks
The genus Chauvinia occurs only in the Afrotropical region with 4 species, one of which is newly described here: C. ganota Claridge sp. nov., C. nitida Heinrich, 1938, C. nyanga Rousse & van Noort, 2013, C. pelecinoides Heinrich, 1938. Claridge (2021a, commenting on the biology of Jethsura Cameron, 1902, noticed that the needle-like terebra is similar to that of other amblypygous species and is enclosed within the 7 th metasomal tergite. These observations led the author to suggest that its morphology could potentially be indicative of a larva oviposition behaviour. This could also be the case for Chauvinia, since it has a similar sclerite organization, but a better understanding of the entire ovipositor morphology and its functionality are required to fully associate a certain structure to a given host association. In fact, in many Atrophini (Ichneumonidae, Banchinae) the enlarged hypopygium is associated with big muscles that run from the first or second valvifers to the lateral part of the abdominal sternum (DDP, personal obs.), suggesting a potentially higher maneuverability of the terebra associated with the type of substrate, while in other groups (e.g., Platygastroidea), the mechanism can be even more complex, involving a telescoping conjunctiva (Talamas et al. 2017). Because for the majority of Ichneumonidae with a short terebra the ovipositor morphology is almost completely unstudied, any conclusion at this stage can only be considered a speculation.

Differential diagnosis
Chauvinia ganota sp. nov. can be distinguished from the other species of Chauvinia by the following combination of characters: (1) second tergite approximately 3 × as long as apically wide (approximately 2 × in C. nitida and C. nyanga) (Fig. 10C); (2) female predominantly brownish-orange, with dorsal margin of pronotum predominantly yellowish-white (female predominantly black in C. nyanga, dorsal margin of pronotum orange to yellowish-orange and concolorous in C. nitida and C. pelecinoides) (Figs 10B, 11A); (3) female antenna with incomplete yellowish-white annulus from 9 th to the 11 th flagellomeres (annulus starting at the 7 th /8 th flagellomeres in C. nitida and C. pelecinoides) (Fig. 10B); (4) male overall lighter in colour than males of C. nyanga and with mesopleuron predominantly yellowish-white (mesopleuron predominantly black in C. nyanga) (Fig. 13A); (5) male face with less distinct and less impressed punctures (more distinct and deeply impressed in C. nyanga) (Fig. 12A).

Etymology
From the Greek 'ganotos' meaning 'polished' or 'brightened' as this species is overall lighter in colour compared to the species to which it is most similar, Chauvinia nyanga Rousse & van Noort, 2013.
Head (Figs 11a, 12a). Subquadrate in frontal view (slightly higher than wide), overall shining; face transverse, more than 3 × as wide as medially high, very finely, sparsely punctate with punctation denser laterally, medio-apically protruding in a very distinct blunt tubercule immediately ventral to antennal sockets, median field present, weakly convex, and delimited by indistinct carinae on the ventral part of the face; antennal socket slightly projecting into a small flange; dorsal tentorial arm pit distant from antennal sockets by less than 2 × the diameter of the pit; frons and vertex almost completely smooth, with extremely sparse and superficial punctuation; ocellar triangle indistinct; occipital carina distinct and complete, meeting hypostomal carina before the base of the mandible; temples slightly rounded in dorsal view; gena, in lateral view, weakly inflated and nearly impunctate except for a few scattered, fine, superficial punctures; clypeus moderately convex, shining, and nearly impunctate except for a few scattered, fine, superficial punctures, particularly near ventral and dorsal margins; malar space short, about 0.6 × basal width of mandible; mandible with sparse setiferous punctures in the central area, teeth rather stout with the ventral tooth shorter (about 0.7 ×) than the upper tooth; antenna subclavate with apical flagellomeres short and wide; flagellum with 23 flagellomeres, the 2 nd flagellomere about 0.8 × length of 1 st , preapical flagellomeres slightly wider than long.  (Fig. 11A-B). Overall shining; pronotum smooth and impunctate; epomia small but welldeveloped; propleuron finely punctate, with sparse hairs, projected into a blunt flange ventro-apically; mesoscutum smooth and impunctate except for a few scattered, fine, superficial punctures, notauli obsolete; scutellum flat and impunctate, carinated only at the base; mesopleuron smooth and impunctate except for a few scattered, fine, superficial punctures, speculum smooth; epicnemial carina slightly raised between fore coxae, laterally ending at anterior edge of mesopleuron; sternaulus distinct on anterior fourth; posterior transverse carina of mesosternum widely obsolete in front of mid coxae; metapleuron smooth and impunctate, juxtacoxal carina strong; propodeum long in lateral view, gently sloping posteriorly; propodeal carinae well-developed except for the posterior transverse carinae obsolete between median longitudinal carinae and subobsolete between median and lateral longitudinal carinae; propodeum overall smooth and sparsely punctate laterally, medially with confluent area consisting of area petiolaris, and area superomedia transversely rugulose, becoming coarser and denser posteriorly. European Journal of Taxonomy 868: 1-71 (2023) Legs (Fig. 10B). All coxae finely, densely punctate; hind coxa without tooth-like projection or carina. Hind femur about 3.1-3.2 × as long as medially high. Tarsal claws without pecten.
CoLoration (Figs 10B, 12a). Head with face, clypeus, mandibles (except for black mandibular apex) pale yellow; inner eye orbit and postero-dorsal section of eye orbit yellow; remainder of head light brownish-orange ventrally and brownish-orange dorsally. Antenna with scape brownish-yellow ventrally, becoming dark orange-brown dorsally; pedicel brown; flagellum dark brown, becoming darker apically, except for 9 th to 11 th flagellomeres yellowish-white medio-dorsally. Mesosoma with pronotum light brownish-orange except for posterior 0.7 of dorsal margin yellowish-white; prosternum brownish-yellow; mesonotum brown medially and dark brown laterally and medially at anterior margin; scutellum orange-brown; mesopleuron light brownish-orange except for longitudinal yellowish-white area immediately dorsal to sternaulus; propodeum brownish-orange except for anteriorly dark brownish. Fore and mid legs with coxae, trochanters and trochantelli yellowish-white; femora light brownishorange; tibiae light brownish-orange ventrally and dark orange-brown dorsally; tarsi dark orange-brown except for 4 th -5 th tarsomeres dark brown. Hind leg with coxa light brownish-orange ventrally and brown dorsally; tarsus dark brown except for 1 st -3 rd tarsomeres orange-brown basally. Metasoma brownishorange with anterior and posterior areas of 1 st -4 th tergites lighter and 6 th -7 th tergites dark orange-brown; ovipositor sheath dark reddish-brown.
Variation (based on the 24 paratypes). Posterior transverse carina between median longitudinal carinae varying from obsolete (20 specimens) to subobsolete (1 specimen) such that area superomedia partially distinct; incomplete white banding on flagellum ranging from 8 th /9 th to 10 th /11 th flagellomeres. European Journal of Taxonomy 868: 1-71 (2023) Male (Figs 12B, 13B) As in female, except for: head transverse, slightly wider than high; face finely, sparsely punctate; 2 nd -7 th tergites densely, finely punctate with punctae becoming progressively finer and sparser at posterior tergites; metasoma shorter than in female. Coloration: head with face, clypeus, mandibles (except for black mandibular apex), ventral 0.3 of gena, inner eye orbits, postero-dorsal section of eye orbit yellowish-white, remainder of head varying from brown to black. Antenna with scape yellowish-white ventrally and remainder dark brown; scape and flagellum dark brown. Mesosoma with pronotum varying from brown to dark brown except for entirety of ventral margin and 0.7-0.8 of dorsal margin yellowishwhite; prosternum yellowish-white; mesonotum varying from entirely black to black with irregular brown submedial longitudinal areas; scutellum predominantly yellowish-white with variable light to dark brown area medially; mesopleuron with large yellowish-white area approximately spanning area ventral to speculum and dorsal to sternaulus except for small light to dark brown area at anterior margin along sternaulus, remainder varying from brownish-orange to dark brown, propodeum usually entirely dark brown to black, occasionally lighter and with small yellowish to yellowish-white mark immediately lateral to propodeal apophyses. Fore and mid legs with coxae, trochanters and trochantelli yellowishwhite; tarsi light yellowish-brown except for 5 th tarsomere dark brown. Hind leg with coxa having large yellowish to dark brown spot on basal and ventral area, remainder yellowish-white; trochanter dark brown except yellowish-white apex; trochantellus yellowish-white; femur brown to dark brown except for basal 0.2 light brownish dorsally; tibia brown to dark brown dorsally and brown ventrally except for basal 0.2 dark brown to black; tarsus dark brown except 1 st tarsomere lighter ventrally. Metasoma with 1 st tergite dark brown; 2 nd -7 th tergites brown to dark brown with extreme apical margins yellowish-white.

Host
Unknown.
Heinrich (1938: 125) described the species based on "Nombreux exemplaires [= numerous specimens]" without clearly designating one of them as the name-bearing type. Initially, Rousse et al. (2013: 21) correctly referred to the types as "syntypes", without designating a lectotype. However, the same authors refer to the specimen depicted in figs 5-6 of their work (Rousse et al. 2013: 22-23) as the "holotype". The employment of the term "holotype" does not constitute a valid lectotype designation (ICZN 1999: article 74.5, 74.7). Moreover, one of the specimens belonging to the type series bears a red label that reads verbatim: "aes Lectotypus Sawoniewicz '77" (Rousse et al. 2013: 23, fig. 6f). Unfortunately, there is no record of a published work clearly designating a lectotype for this species, and therefore the label does not constitute a valid lectotype designation (ICZN 1999: article 74.5). A more in-depth study of the

Remarks
The three specimens examined here represent the first record of the species since the original description, as well as an expansion of its range to the Diana region ( Fig. 16A-B). (Heinrich, 1938). A. Previous (blue dots) and new occurrence records (yellow star). B. Previous (blue) and new (yellow) regional records.

Remarks
In the original description, Rousse et al. (2013: 23) reported that the mesosoma is mostly black, with "a longitudinal stripe on mesopleuron sometimes lightening to pale yellow" (see Rousse et al. 2013: fig. 8e). However, the female specimens from Kenya here examined display a larger yellow area reaching the middle of the mesopleuron and extending to the mesosternum, at least partially ( Fig. 17A-B).
The 67 specimens recorded from Kenya represent the first record for the country (Fig. 18A-B). -Apex of propodeum hardly elongated between hind coxae (a); clypeus extremely transverse, more than 3 × as wide as long (b); metasoma of female shorter than or as long as hind leg, with ventral margins of 4 th and following tergites overlapping (c  Ashmead (1906: 171). Synonymized by Perkins (1959: 77).

Diagnosis
The unidentate and falcate mandibles set Heterischnus Wesmael, 1859 apart from all the other Afrotropical genera of Phaeogenini, except for Lusius, from which it can be distinguished by the presence of vein 3rsm (areolet closed) in the fore wing and the abscissa of CU in the hind wing. Additionally, the genus can be separated from other genera by the following combination of characters: (1) clypeus separated from face by clupeal suture; (2) basal flagellar segments slender and long; (3) vertex long and slightly convex behind ocelli; (4) hypostomal carina joining occipital carina distinctly at or above mandibular base; (5) epicnemial carina strongly raised ventrally and flexed over the base of the fore coxae; (6) notauli distinct anteriorly, deep and long; (7) tarsal claws simple; (8) gastrocoeli nearly indistinct to deep, thyridia wide; European Journal of Taxonomy 868: 1-71 (2023) (9) ovipositor extending relatively strongly beyond apex of metasoma (Perkins 1959;Selfa & Diller 1994;Rousse et al. 2013;Claridge 2021b) .

Remarks
Despite being currently a synonym of Heterischnus, Aethiopischnus requires a focused commentary to resolve some nomenclatural issues. Aethiopischnus firstly appeared in Heinrich (1936: 244) without being accompanied by any formal description. The genus name served only to accommodate the description of a new species, Aethiopischnus africanus. Later on, Heinrich (1938: 127) introduced the name Aethiopischnus again as "gen. nov." providing this time an adequate description and designating Aethiopischnus olsoufieffi as the type species. Townes & Townes (1973: 221) acknowledged the two names (Aethiopischnus Heinrich, 1936 andAethiopischnus Heinrich, 1938) and placed them as a synonym of Heterischnus. Yu & Horstmann (1997: 489) deemed Aethiopischnus Heinrich, 1936 an unavailable name without providing any discussion for their action. According to ICZN (1999: article 13.1.1), to be available a name proposed after 1930 needs to be accompanied "by a description or definition that states in words characters that are purported to differentiate the taxon.". Because of the lack of a proper description, the genus name Aethiopischnus Heinrich, 1936 is deemed a nomen nudum, and therefore unavailable. Aethiopischnus Heinrich, 1938 is the available name (even though invalid) in accordance with Yu & Horstmann (1997: 489). Rousse & van Noort, 2013 Figs 19-21 Heterischnus mfongosi Rousse & van Noort, 2013: 47-51 (original description, key).

Diagnosis (male)
For the first time, a male specimen of the species is hereby diagnosed, bearing the same label data as one of the female paratypes reported in the original description. Compared to the female, no major differences are reported. Body length is approximately 7.0-7.5 mm, fore wing length is 4.2 mm. The coloration is identical, with the exception of: (1) absence of white annuli in the antennae, and (2) a slightly lighter coloration of the anterior part of 6 th tergite (Fig. 19B). Rousse & van Noort, 2013 (NHMUK). A. ♀, habitus, lateral view. B. ♂, habitus, lateral view.

Remarks
In the original description, Rousse et al. (2013: 51) reported that the propodeum of the species is "without carination" and that the metasomal tergites are testaceous "with median black maculae." However, the specimens examined by one of us (DDP) from Zimbabwe and matching the paratype locality, clearly showed the presence of a complete and well-defined posterior transverse carina (Fig. 20A-B), and no presence of black maculae on the metasoma. Based on the examination of the holotype, it seems clear that the reported black maculae are only an artifact due to the poor state of preservation of the specimen. When muscles and body fat tissue break up, the external coloration of the tergites becomes darker, creating the illusion of maculae. Similar situations are not rare in preserved specimens, and this has been briefly discussed for Genaemirum filipazzii Dal Pos & Rousse, 2018(Dal Pos & Rousse 2018. Based on the specimens examined here, the coloration is orangish-yellow for the entire metasoma, with only the apical tergites (6 th and 7 th ) infuscated (Fig. 19A).
Before this contribution, the species was known only from the type series and recorded only from southern Africa. The additional examined material included here extends the distributional range of the species to East Africa ( Fig. 21A-B).

Original type series
Syntypes: 2 ♀♀, 3 ♂♂; MZPW. Heinrich (1938: 128) described the species based on 5 specimens from different locations in Madagascar without designating any of them as the name-bearing type. Rousse et al. (2013: 54), referring to the type series, employed the terms "holotype" and "paratype". This latter usage of the terms is incorrect. Holotypes are designated in the original description or are implied when only one specimen is described (ICZN 1999: 73.1). In this case, Heinrich (1938: 128) mentioned more than one specimen in the type series and did not choose any of them as holotype. Moreover, the employment of the term "holotypus" does not constitute a valid lectotype designation either (ICZN 1999: article 74.5). A more in-depth study of the MZPW collection will be required prior to the designation of a lectotype; therefore, for the moment, the specimens of the original type series should be referred to as syntypes (ICZN 1999: article 73.2).

Remarks
In the description of Aethiopischnus africanus, Heinrich (1936: 244) introduced the name "Ae. Olsoufieffi Hein." without providing any other detail. Aethiopischnus olsoufieffi reappeared later on in Heinrich (1938: 127), but this time it was accompanied by a formal description and was designated as the type species of the genus Aethiopischnus. Townes & Townes (1973: 221) and Diller & Schönitzer (1999: 295) reported "Heinrich, 1938" as the authorship of the species, while Yu & Horstmann (1997: 489) and Rousse et al. (2013: 54) retained "Heinrich, 1936". When Heinrich (1936 introduced olsoufieffi, he did by comparing it with A. africanus. However, according to ICZN (1999: article 13.1. 1 & 13.1.2) a new name after 1930 must be accompanied "by a description or definition that states in words characters that are purported to differentiate the taxon" or "by a bibliographic reference to such a published statement.". Basically, if the original statements allow the construction of a conclusion that can be used as a description or differential diagnosis, then the name should be considered available. However, from Heinrich's (1936: 244) comparison, it is impossible to differentiate olsoufieffi from africanus. The statement "Dem Ae. Olsoufieffi Heinr. nahestehend, aber von dunklerer, unbestimmterer Grundfarbe, Fühlergeissel ohne weisse Zeichnung. Schläfen relativ schmaler. Wengen und Schläfen dichter punktiert. Thyridien kaum noch angedeutet [= Close to the species Ae. Olsoufieffi Heinr., but of darker, more indistinct ground colour, antennal flagella without white markings. Temples relatively narrower. Cheeks and temples more densely punctured. Thryridia hardly indicated]" is too vague and does not allow a clear separation between the two species. Before Heinrich (1936), also africanus (the species with which olsoufieffi was compared) was not known and the comparison could not be anchored to a previous species hypothesis. Therefore, Aethiopischnus olsoufieffi Heinrich, 1936 is hereby deemed a nomen nudum (unavailable) and the available name is Aethiopischnus olsoufieffi Heinrich, 1938. The circumstances that led Heinrich (1936) to introduce the name olsoufieffi are probably due to a publication issue. Reading the introduction, it seems clear that Heirnich's (1938) monograph was meant to be published before Heinrich's (1936) work, but a contingent situation delayed the publication of the Malagasy work.
The three additional CAS specimens represent the first record of the species since the original description, as well as an expansion of its distributional range to the Amoron'i Mania and Vatovavy-Fitovinany regions (Fig. 23A-B). (Heinrich, 1938). A. Previous (blue dots) and new records (yellow stars). B. Previous (blue) and new (yellow) regional records.  (Heinrich, 1936) European Journal of Taxonomy 868: 1-71 (2023) Hoplophaeogenes Heinrich, 1938Hoplophaeogenes Heinrich, 1938. Type species: Hoplophaeogenes amoenus Heinrich, 1938, by original designation.

Original type series
Syntypes: 2 ♀♀ (MZPW). Heinrich (1938: 127) described the species based on two females (one from "Montagne d'Ambre" and the other from "Diégo-Suarez") without clearly designating any of them as the name-bearing type. Initially, Rousse et al. (2013: 59) correctly referred to the types as "syntypes", without designating a lectotype. However, the same authors (Rousse et al. 2013: 60-61, figs 33-34) referred to the specimens as the "holotype". The use of the term "holotype" does not constitute a valid lectotype designation (ICZN 1999: article 74.5, 74.7). A more in-depth study of the MZPW collection will be required prior to designating a lectotype; therefore, for the moment, the type specimens should be referred to as syntypes (ICZN 1999: article 73.2).

Type locality
Madagascar

Remarks
The only two specimens known are those belonging to the type series, from the Diana region (Northern Madagascar). The specimen here recorded represents the first record of the species for the Analamanga region and, more importantly, for the central part of Madagascar ( Fig. 25A-B).  Heinrich, 1938 Genus Kibalus Rousse, van Noort & Diller, 2013 Kibalus Rousse, van Noort & Diller in Rousse et al., 2013: 59.  (4) 2 nd metasomal tergite more than 2 × as long as apically wide (almost square in Centeterichneumon and Diadromus). The genus can further be distinguished by: hemispherical head, coarsely sculptured; upper tooth of mandible much longer than lower tooth; mesoscutum steeply elevated above pronotum; propodeum elongate, regularly rounded without differentiated horizontal anterior part in profile view; propodeal carination weak, more or less complete; fore wing with areolet closed; hind wing with distal abscissa CU absent; ovipositor very shortly projecting beyond metasomal apex .
European Journal of Taxonomy 868: 1-71 (2023) According to Rousse et al. (2013), the genus is also characterized by a complete and strong postpectal carina. However, in the new species described below, Kibalus nonnaritae Dal Pos & Di Giovanni sp. nov., the postpectal carina is absent in front of the mid coxa.

Etymology
The species is dedicated to the first author's grandmother, Rita Zanin, and the specific epithet is the combination of two terms: 'nonna' (Italian word for 'grandmother') and 'ritae' (female noun in the genitive case formed from a modern person name (ICZN 1999: article 31.1.2)). For her constant and indefatigable support to her family over the entire span of her life and for the tacit understanding of DDP during the many summers he spent in grandma's backyard, looking for small creatures. Without that initial support, this new species could not have been described. Female, in perfect condition, micropinned.
Head (Figs 26B, 27a). Overall shining; face subquadrate, as wide as medially high, smooth and almost impunctate, medio-apically protruding in a very distinct blunt tubercule right below the antennal sockets, clear delimitation between clypeus and face absent; frons and vertex almost completely smooth, impunctate; ocellar triangle equilateral and not elevated; ocular-ocellar distance about 1.3 × ocellus diameter, inter-ocellar distance 1 × ocellus diameter; occipital carina distinct and complete, meeting hypostomal carina at base of mandible; temples slightly inflated in dorsal view; gena, in lateral view, European Journal of Taxonomy 868: 1-71 (2023) medially strongly inflated, smooth; clypeus smooth, shining with straight apical margin; malar space short, about 0.5 × basal width of mandible; mandible robust, with sparse setiferous punctures in the central area, teeth rather stout and widely separated with the ventral tooth shorter (about 0.8 ×) than the upper tooth; maxillary palpi long, reaching fore coxae, 5 th segment about 1.5 × as long as 4 th ; antenna filiform, not tapering, flagellum with 28 flagellomeres, the 1 st flagellomere about 1.4 × as long as the 2 nd , preapical flagellomeres distinctly longer than wide.
MesosoMa ( Fig. 26A-C). Overall shining; pronotum smooth, shining and impunctate; epomia absent; propleuron smooth, shining and almost impunctate, projected into a blunt, rounded flange ventro-apically; mesoscutum subquadrate, smooth, finely and sparsely punctate, notauli short and distinct only anteriorly; scutellum slightly elevated over metascutellum, almost impunctate, carinated throughout; mesopleuron shining all over, smooth on the upper ⅔ and sparsely and finely punctate ventrally, speculum smooth, epicnemial carina laterally ending at anterior edge of mesopleuron; sternaulus distinct on ⅔ of the length; posterior transverse carina of mesosternum incomplete, absent in front of mid coxae and slighty raised European Journal of Taxonomy 868: 1-71 (2023) between mid coxae; metapleuron smooth, shining and impunctate, juxtacoxal carina strong; propodeum short in lateral view, sloping gently with almost no horizontal part; anterior transverse carina present with area basalis and area superomedia separated and shagreen, area externa and area dentipara separated and shagreen; area spiracularis and area lateralis shagreen and shining.
CoLoration . Head with face, clypeus, mandibles (except the black apical teeth), orbits all around the eyes (except for a spot on vertex), and genae pale yellow; black are: mandibular teeth, ocellar triangle, frons, vertex, and temples. Antenna with scape and pedicel reddish-brown; flagellum black with white annulus present only on the dorsal side, from the 9 th to 13 th /14 th flagellomeres. Mesosoma yellow, with base and lateral lobes of mesoscutum, metanotal trough, and propodeum (except area petiolaris) infuscated; tegulae pale yellow; the overall ventral side is pale yellow while the dorsal sclerites are mostly orange. Legs overall yellowish-brown, with fore and mid coxae, fore and mid trochanter, and basal portion of fore and mid tibia pale yellow; hind coxae bicolored, pale yellow in the basal half and reddish brown in the apical half; hind leg overall more infuscated than the fore and mid legs. Wing entirely hyaline. Metasoma entirely brownish-black, except for the white basal portion of the 1 st tergite, the white apical bands on 2 nd and 3 rd tergites, the white gastrocoeli and thyridia, and the white central spot on the 7 th tergite; gonoforceps brownish-black with a white apex, penis valvae also pale yellow.

Female
Unknown.

Diagnosis
The unidentate and falcate mandibles set Lusius Tosquinet, 1903 apart from all the other genera of Phaeogenini, except for Heterischnus Wesmael, 1859, from which it can be distinguished by the absence of vein 3rs-m (areolet open) in the fore wing and abscissa of CU in the hind wing. Additionally, the genus can be separated from other genera by the following combination of characters: (1) head hemispherical; (2) basal flagellar segments slender and long; (3) vertex long and slightly convex behind ocelli; (4) occipital and hypostomal carinae joining at mandibular base; (5) notaulus complete, ending posteriorly in a median depression; (6) gastrocoeli long with thyridia faint; (7) ovipositor extending beyond apex of metasoma; (8) male genitalia with gonoforceps reduced to rod-like processes (Baltazar 1964;Diller 2006;Rousse et al. 2013).

Remarks
The genus has a tropical distribution, and it occurs in the Afrotropical, Indomalayan and Neotropical regions. In Africa, Lusius is represented only by two species: L. flummox Rousse &van Noort, 2013, andL. tenuissimus Heinrich, 1938 (Table 1). (Heinrich, 1938 Heinrich (1938: 129) described the species based on multiple specimens, males and females, without clearly declaring the number of specimens nor designating any of them as the name-bearing type. Initially, Rousse et al. (2013: 72-73, figs 42-43) incorrectly referred to the type as holotype. The employment of the term "holotype" does not constitute a valid lectotype designation (ICZN 1999: article 74.5, 74.7). A more in-depth study of the MZPW collection will be required prior to the designation of a lectotype; therefore, for the moment, the type specimens should be referred to as syntypes (ICZN 1999: article 73.2).

Fig. 29. Lusius tenuissimus
European Journal of Taxonomy 868: 1-71 (2023) Heinrich (1938: 129) stated that the species is "Largement répandu, mais rare [= widely distributed, but rare]". This statement seems to be largely true as Lusius tenuissimus is probably the most widely spread species of Phaeogenini across Africa. It cannot be considered "rare" as Heinrich (1938) stated, but surely it is not largely abundant. The number of specimens per country from Rousse et al. (2013) and this current contribution seem to corroborate that. Also, the specimen from Uganda represents the first record for the country (Figs 30-31). (Heinrich, 1938). A-B. Previous (blue dots) and new records (yellow star). A. Mainland Africa. B. Madagascar. (Heinrich, 1938 Before this contribution, the genus was recorded only from the Indomalayan region. This new combination marks the first record for the Afrotropical region.

Type locality
Tanzania: Mkomazi Game Reserve.
The above features are instead shared with the genus Nesostenodontus Cushman, 1922, from which it slightly differs by the ventral margin of the clypeus, which is irregularly emarginate in mkomazi as opposed to strongly dentate in bakeri (the type species) (Cushman 1922: 554) (Fig. 32A). However, this difference can be regarded as a simple variation within the genus. Moreover, three other unidentified Afrotropical specimens were located at EMUS, belonging to the genus Nesostenondontus. However, their species identity remains, at the moment, doubtful. Therefore, because of the above shared features of H. mkomazi with Nesostenodontus, and based on an examination of the new material, we feel confident in proposing a new combination for mkomazi: Nesostenodontus mkomazi (Rousse & van Noort, 2013) comb. nov.
Nesostenodontus was previously known only from three described species in the Indo-Malaysian region Heinrich 1934;Gupta 1988), but it is herewith also recorded for the first time from the Afrotropical region, specifically in Tanzania and Senegal (N. mkomazi) , and Sierra Leone and South Africa (undescribed species at EMUS). Because of this generic transfer, the genus Heterischnus is no longer recorded for Senegal (Table 1).

Discussion
Despite the comprehensive revision by Rousse et al. (2013), the tribe Phaeogenini in the Afrotropical region is still severely understudied, with most countries without a single record for the tribe, and others with less than 2 ( Fig. 35; Table 1). Madagascar, Uganda, and South Africa are best represented but these countries still only have a maximum of 7, 8 and 11 species, respectively ( Fig. 35; Table 1). Most of the species are recorded from the eastern part of the continent (see Table 1), a trend reflected across the entire subfamily Ichneumoninae (e.g., Rousse et al. 2016;Dal Pos & Rousse 2018), and also across the family Ichneumonidae on the whole (e.g., Di Giovanni & Varga 2021). This 'biogeographic bias' is most likely not the result of a natural distribution of the taxon, but rather an artifact caused by mainly the combination of three factors: (1) a shortage of taxonomists, which is a problem affecting the entire entomological community (e.g., Engel et al. 2021;Zamani et al. 2022a), but particularly exacerbated in Ichneumonidae (e.g., Riedel et al. 2021); (2) a lack of modern and comprehensive taxonomic treatment (the most recent contribution to the Afrotropical Ichneumoninae fauna is Riedel & Dal Pos 2019); and (3) the reduced availability of specimens from the west of the continent due to different historical and contemporary reasons. Collaborative efforts among the ichneumonoid researchers and a thorough study of other collections, as well as partnerships with local African institutions, will be required to fill these gaps.
From our results, it is self-evident that also the nomenclatural aspect needs substantial improvement. Many types of Ichneumoninae require examination and careful study in order to improve the taxonomic stability of the family (e.g., syntype/lectotype problems). These problems should not come as a surprise. Broad (2021) discovered several issues in a single collection (Morley's) focusing only on a small European Journal of Taxonomy 868: 1-71 (2023) selection of taxa across Ichneumonoidea. The curation of existing names (and not just the description of new species) is a fundamental practice in the compilation of meaningful taxonomic work and, as pointed out by Zamani et al. (2022aZamani et al. ( , 2022b, is one of the goals of taxonomy. This is especially true in Ichneumonidae, given their huge number of species (and therefore names), and the recent discontinuity of resources, such as Taxapad (Yu et al. 2016), which can potentially negatively affect even more the already poor status of knowledge of this family.
Hopefully, this paper will serve the community, providing tools for the future advancement of knowledge of the tribe, and, more broadly, of the entire subfamily. To paraphrase what Heinrich stated at the opening