Revision of the world species of Zambion (Hymenoptera, Ichneumonidae, Tryphoninae)

Abstract The world species of Zambion Kasparyan (Hymenoptera: Ichneumonidae: Tryphoninae) are revised including re-descriptions of Zambion monodon Kasparyan and Zambion hirtum Delobel. Five new species are described: Zambion kasparyani sp. n., Zambion rogeri sp. n., Zambion eileenae sp. n., Zambion wahli sp. n. andZambion broadi sp. n. A key to species is provided. The genus is endemic to Africa (Angola to Kenya) and is one of only three genera of the tribe Tryphonini recorded from the Afrotropical region.


Introduction
The tribe Tryphonini is poorly represented in the Afrotropical region with only 3 of 21 extant genera recorded (Yu et al. 2009;Bennett 2002): Boethus Förster (Scaramozzino 1991) and two endemic genera: Ibornia Seyrig (Seyrig 1935) and Zambion Kasparyan (Kasparyan 1993). In total, there are only eight described species of Tryphonini recorded from the Afrotropical region: two species each of Boethus and Zambion and four species of Ibornia (Yu et al. 2009). It was speculated by Gauld (1987) that a reason for the lack of ctenopelmatine ichneumonids from tropical regions is because their hosts (sawflies) are rare in the tropics in terms of species and biomass. The same hypothesis could explain the low number of species of Afrotropical Tryphonini because almost all known host records for the tribe are from sawflies (Gauld et al. 1997). The hosts of Ibornia and Zambion are not known, but they likely parasitize tenthredinid sawflies based on the known host range of related genera (e.g., Dyspetes Förster and Otoblastus Förster) (Gupta 1983;Kasparyan 1973). Of the 10 genera of Tryphonini for which hosts are known, only two have been reared from hosts other than Tenthredinidae: Boethus species have been consistently reared from Argidae (Hymenoptera) (Gauld et al. 1997) and some species of Grypocentrus Ruthe have been reared from Eriocraniidae (Lepidoptera) (Jordan 1998) (most rearing records of Grypocentrus are from Tenthredinidae) (Kasparyan 1973).
Apart from the presumed lower number of potential hosts, another factor that definitely contributes to the small number of species of Afrotropical Tryphonini (and Ichneumonidae in general) is the relatively little amount of taxonomic research on the ichneumonids of the region. In other regions such as the Neotropical, recent active research programmes have greatly elucidated the ichneumonid fauna (e.g., Gauld 1991;Gauld et al. 1997Gauld et al. , 2000Gauld et al. , 2002. In the Afrotropical region, some excellent work has been done in the past, for example, Seyrig (1932Seyrig ( , 1934Seyrig ( , 1935Seyrig ( , 1952 on various subfamilies, Heinrich (1967aHeinrich ( , b, c, 1968a on Ichneumoninae, and Gauld and Mitchell (1978) on Ophioninae, but recently, taxonomic studies including Afrotropical species have been uncommon. For example, Yu et al. (2009) listed only 8 taxonomic papers that included Afrotropical species published between 2000 and 2008 (compared to 65 for the Neotropical region and 46 for the Oriental). In terms of described species, the Afrotropical ichneumonid fauna is not as speciose as the Neotropical and Oriental regions (1,946 described species in the Afrotropical compared to 3,803 in the Neotropical and 3,595 in the Oriental) (Yu et al. 2009). But based on examination of unidentified material at the American Entomological Institute, Canadian National Collection of Insects and the Natural History Museum, London, there are large numbers of undescribed species in the Afrotropical region, especially from the tropics and South Africa. Without additional taxonomic studies of the fauna, well-founded assessments of species diversity and biological patterns for sub-Saharan Africa will be difficult to achieve. The purpose of this paper is to revise the genus Zambion including description of five new species. It is hoped that the revision of such a beautiful and morphologically distinct genus will encourage other researchers to spend much-needed time working on this interesting, but under-studied fauna.

Methods
All terms of ichneumonid morphology follow Townes (1969) with the following modifications: hypostomal carina for 'oral carina', supra-antennal area for 'frons', supracl-ypeal area for 'face', gena for 'temple', occiput for 'postocciput', malar space for 'cheek', epicnemial carina for 'prepectal carina', laterotergites for 'epipleura', gonoforceps for 'claspers', and hypopygium for 'subgenital plate'. The term 'mesosoma' is used for the body region that includes the thorax and first abdominal segment (the propodeum). The term 'metasoma' is used for the apparent abdomen, with MS1, MS2, etc. referring to metasomal segments 1, 2, etc., T1, T2, etc. referring to the tergites of metasomal segments 1, 2, etc. and S1 referring to the sternite of metasomal segment 1. The term T2+ refers to tergite 2 and all tergites posterior to T2. Terms of relative position of the body follow Goulet and Huber (1993). Wing venation terms follow the Comstock-Needham system as updated by Ross (1936) and incorporate the recommendations of Goulet and Huber (1993) except for naming of the vein that forms the distal edge of fore wing cell 1+2Rs (the 'areolet' of Townes 1969). This vein is of uncertain origin and is here referred to as 'vein 3rs-m' in conformity with Wahl and Gauld (1998). The following terms for specialized structures are defined: epomia: a raised ridge (carina) on pronotum (Figs 30-31); glymma: lateral depression sub-basally on T1; notaulus: longitudinal groove sublaterally on mesoscutum (Fig. 29); sternaulus: longitudinal groove subventrally on mesopleuron; ventral transverse carina of propodeum: a carina that extends from the metasomal foramen of the propodeum to the pleural carina (Fig. 35).
Measurements for particular structures were made as follows: supraclypeal width was measured at midheight compared to height in the middle (laterally); horn length measured in dorsal view versus pedicel width at middle in lateral view; gena width measured at lower (narrowest) point versus transverse width of eye at midheight (both in lateral view); length of scutellar carina measured from base of scuto-scutellar groove to apex of scutellum; point of union of posterior transverse carina of propodeum (ptc) and pleural carina (pc) was estimated by measuring the distances shown in Figs 35-37: from the propodeal spiracle to the ptc and from the ptc to the ventral transverse carina (vtc) (or in Z. monodon and Z. kasparyani, where the vtc would be, if present); hind femur dimensions were calculated by dividing the maximum length including the distal trochantellus by the height at midpoint; length of T1 measured in lateral view compared to width of T1 measured in dorsal view at posterior edge. Measurements of holotypes are shown in parentheses after range. Characters listed in the generic description are those deemed relevant to distinguish Zambion from all other genera of Tryphonini as defined by Kasparyan (1973), Kasparyan and Tolkanitz (1999) and Bennett (2002).
Specimens are deposited at the following institutions (acronyms according to Evenhuis 2011): Canadian National Collection of Insects (CNC); The Natural History Museum, London (NHM); Muséum national d'Histoire naturelle, Paris (MNHN), Finnish Museum of Natural History, Helsinki (MZH), California Academy of Sciences, San Francisco (CAS). Label data for holotypes is reported verbatim from the labels, with our comments about the labels indicated in square brackets. Locality data for other specimens are provided in a standardized format.
Digital photos were taken using a Leica MZ16 stereomicroscope with motorized focus drive attached to a Leica DFC420 digital camera. Photos were combined and edited Diagnosis. Differentiated from other genera of Tryphoninae by combination of both of the following characters: 1) unidentate mandibles (Fig. 22); 2) supra-antennal area with a medial horn (not a vertical lamella) slightly ventral to medial ocellus (24)(25)(26)(27). Zambion would most easily be mistaken for Ibornia Seyrig, but Ibornia has a supra-antennal vertical lamella instead of a horn and T1 and T2 are fused (not fused in Zambion).
Description. Adult. Fore wing length 4.9 to 7.2 mm. Clypeus separated from supraclypeal area by a strong groove (most species) (Fig. 11) or weakly separated in Z. hirtum Delobel (Fig. 10); strongly convex or moderately flat in profile; not divided into dorsal and ventral faces by a transverse line (most species) (Fig. 11) or divided (Fig. 12). Apical edge of clypeus not projecting ventroanteriorly in lateral view, truncate to slightly convex medially in anterior view, without a pair of medial denticles. Mandible unidentate (Fig. 22), outer face (anterior surface) basally flat and coarsely punctate (not inflated and impunctate as in Ctenochira). Glossa slightly to strongly elongated (Fig. 15). Posterior mandibular condyles separated by greater width than inner margins of eyes at level of clypeal foveae. Malar space 0.4 to 0.7 times basal width of mandible. Occipital carina present only dorsally (between inner margins of eyes), therefore not joining hypostomal carina, the latter not directed posteriorly into a tooth. Supra-antennal area with or without a glabrous depression, if present then just as a deep dimple directly ventral to medial ocellus, not an elongate groove as in Cosmoconus Förster. Supra-antennal area with a medial horn (24)(25)(26)(27). Vertex posterior to medial ocellus without a rounded, conical protuberance. Distance between inner edges of toruli 0.1 to 0.4 times width of one torulus.
Dorsal edge of pronotum (seen in dorsal view) strongly thickened laterally (Fig.  28). Epomia present only as a slight sharpening of ventroanterior ridge of pronotum (not crossing pronotal groove) (Figs 30-31). Notaulus weak (Fig. 29) or absent. Subtegular ridge slightly protruding laterally, not extending dorsally as a vertical lamina that nearly touches tegula, without a posterior longitudinal slot. Epicnemial carina not joined by an auxiliary carina that extends from anterior edge of mesopleuron. Vertical portion of epicnemial carina close to anterior edge of mesopleuron and dorsally curving anteriorly to meet anterior edge of mesopleuron . Longitudinal carinae of mesopleuron and mesoscutum (as in Ibornia) absent. Sternaulus weak and wide to 0.2 to 0.3 length of mesopleuron. Angular flange on posterolateral edges of mesoscutum and associated 'axillary tongue' both present and strongly angulate. Scutellum completely black (most species), completely yellow (Z. hirtum and Z. kasparyani) or black with yellow apically (Z. monodon), lateral carinae present only basally or up to 0.5. Propodeum with all carinae present except anterior transverse carina completely absent in all species (Fig. 34), medial longitudinal carinae absent anteriorly in Z. wahli and Z. broadi and posterior transverse carina interrupted medially in the males of Z. wahli and Z. broadi. Posterior transverse carina angulate where it intercepts medial longitudinal carinae (Fig. 34). Foretibia with a strong tooth on apex of anterior side or tooth weak to absent. Trochantelli not fused on middle and hind legs. Hind tarsal claws stoutly pectinate in basal 0.4 to 0.6. Fore wing vein 3rs-m present. Cell 1 + 2Rs subrectangular or subrhombic, width not greater than 1.4 times height, petiolate anteriorly. Fore wing vein 2m-cu with an angulation (zig-zag) that is strong in most species  or weak (Fig. 7) and with one wide or two narrowly spaced bullae. Fore wing hyaline basally and infumate apically (Figs 38-39) or completely moderately infumate (Figs 7-8).
T1 gradually widening from anterior to posterior, 1.2 to 1.8 times as long as posterior width, with moderate anterolateral projections (as seen in dorsal view). Glymma present as a depression ventral to anterolateral projections. Dorsal longitudinal carinae of T1 present basally or extending up to 0.7 of segment. Dorsolateral longitudinal carinae of T1 present (most species) or absent; if present, not extending posterior to spiracle (Figs 40,41) or extending up to 0.7 of segment, not bifurcated into two branches above and below spiracle. Spiracle of T1 just anterior to middle. T1 and T2 not fused. T2 without a postmedial transverse groove or grooves that delineate anterolateral corners. T2 to T4 completely yellow/orange, completely black, or black with yellow at posterior of each segment. Laterotergites wide (Fig. 42), of MS3 divided from tergite by a crease, of MS4 not divided. T7 and T8 not turned anteriorly under metasoma. Ovipositor straight, about equal to apical height of metasoma or a little shorter, carrying one egg (Figs 5,45) or no eggs, dorsal valve tapered to a sharp point (Fig. 44). Ovipositor sheath in profile, widest basally, gradually tapering from base to apex, flexible from 0.3 to 0.6. Body polished, face and supraclypeal area coarsely punctate (Figs 10-15); pronotum, mesopleuron and metapleuron with fine, sparse, but distinct punctures ; all other body regions almost impunctate (40)(41)(42)(43). Body covered in dense setae, particularly long on supraclypeal area, shorter on pleura of mesosoma and metasomal tergites.

Species included. Z. monodon, Z. hirtum Delobel and five new species described below.
Comments. Zambion, Ibornia and Thibetoides are all closely related based on seven synapomorphies, especially the thickened dorsoposterior corner of the pronotum and the supra-antennal prominence (Bennett 2002). Kasparyan (1993) speculated that Zambion and Ibornia must be sister genera because of the unidentate mandibles and because they are both Afrotropical, but Bennett (2002) found that Ibornia and Thibetoides are sister taxa based on seven characters including the fusion of T1 and T2 and the ventral curving of T7 and T8 under the metasoma which makes Zambion the sister of these two genera. Fore and mid legs orange, hind coxa, trochanter and femur orange, hind tibia and tarsus dark brown, except yellow at base of hind tibia (Fig. 6). Tergites 3 to 7 with medioposterior ivory bands (Fig. 6 (Fig. 36). The only other species of Zambion that lacks this carina is Z. kasparyani, but the latter species has a mesopleuron that is completely yellow to orange (Fig. 46). Description. Adult. Female (based on holotype only -see comments below). Fore wing length 7.2 mm. Clypeus separated from supraclypeal area by a strong groove (Fig. 11). Malar space 0.5 times basal width of mandible. Supraclypeal area 2.0 times as wide as high (Fig. 11), dorsomedially without a short, narrow, longitudinal lamella between the antenna. Supra-antennal horn moderately long and triangular in dorsal view (Fig. 17), about equal in length to width of pedicel at midheight in lateral view (Fig. 25). Ocello-ocular distance 1.4 times ocellar diameter. Gena 0.6 times transverse diameter of the eye. Antennal flagellum with 37 segments.
Notaulus present, but weak. Epicnemial carina medially curving slightly away from anterior edge of mesopleuron near ventral corner of pronotum (intermediate between Z. rogeri shown in Fig. 32 and Z. wahli shown in Fig. 33). Scutellum with lateral carinae 0.4 length of scutellum. Medial longitudinal carinae of propodeum complete and strong anterior to posterior transverse carina (Fig. 3). Carina extending from pleural carina to spiracle complete (Fig. 36). Posterior transverse carina of propodeum with lateral abscissa roughly straight, joining pleural carina at about 0.4 distance from posterior end of pleural carina to propodeal spiracle (Fig. 36). Ventral transverse carina extending from metasomal foramen to pleural carina absent (Fig. 36). Fore tibia with a strong apical point on dorsal surface. Hind femur 3.4 times as long as medial width (Fig. 1). Hind tarsal claw pectinate to about 0.5. Angulation of fore wing vein 2m-cu strong (Fig. 1). Tergite 1 of metasoma 1.7 times as long as apical width. Dorsal longitudinal carinae of T1 extending posterior to spiracle (0.6 length of tergite), dorsolateral longitudinal carinae extending about half way to spiracle (0.3 distance of T1) (Fig. 41 of male, similar to female).
Yellow. Legs and metasoma become slightly darker yellow apically/ posteriorly. Scape, pedicel except apically, supra-antennal area except orbits ventral to medial ocellus, vertex, gena dorsal to ventral edge of eye, occiput, apex of mandibles, pronotum dorsal to furrow, mesoscutum, dorsal half of mesopleuron except subtegular ridge and scutellum except basolaterally black/ dark brown. Apex of pedicel, flagellum and distal tarsomere of hind leg brown. Entire membrane of fore wing moderately infumate (Fig. 5). Body covered with dense golden setae.
Male (based only on single male paratype noted by Kasparyan 1993 Comments. Z. monodon appears related to Z. kasparyani and Z. hirtum on the basis of the similar moderately long, triangular-shaped supra-antennal horn (Figs 16,17,47). Within this grouping, Z. monodon and Z. kasparyani both lack the ventral transverse carina of the propodeum (Figs 36, 48). Outgroup comparison with Ibornia and Thibetoides spp., reveals that this character varies between species within these two genera as well, therefore its polarity (and phylogenetic utility) in Zambion is unclear. Zambion monodon is the only species of Zambion for which the egg is known. Kasparyan (1993) stated that the dermestid damaged paratype was a female. Because the metasoma posterior to T3 is missing, it is difficult to confirm the sex; however, the length of T1 (1.8 times posterior width) and ocellar-ocular distance (1.2 times ocellar diameter) suggest that this specimen may be a male based on comparison with the dimensions of these structures in the female holotype and male paratype.  Male: Fore wing length 5.8 mm. Clypeus separated from supraclypeal area by a strong groove. Malar space 0.4 times basal width of mandible. Supraclypeal area 1.7 times as wide as high, dorsomedially without a short, narrow, longitudinal lamella between the antenna. Supra-antennal horn moderately long and triangular in dorsal view (Fig. 47), about equal in length to width of pedicel at midheight in lateral view. Ocello-ocular distance 1.2 times ocellar diameter. Gena 0.8 times transverse diameter of the eye. Antennal flagellum with 32 segments.
Notaulus absent. Epicnemial carina mostly straight medially, not curving away from anterior edge of mesopleuron near ventral corner of pronotum (similar to Z. rogeri shown in Fig. 32). Scutellum with lateral carinae extending 0.4 length of scutellum. Medial longitudinal carinae of propodeum complete and strong anterior to posterior transverse carina (as in Fig. 3). Carina extending from pleural carina to spiracle complete (Fig. 48). Posterior transverse carina of propodeum with lateral abscissa roughly straight, joining pleural carina at about 0.4 distance from posterior end of pleural carina to propodeal spiracle (Fig. 48). Ventral transverse carina extending from metasomal foramen to pleural carina absent (Fig. 48). Fore tibia with a moderately strong apical point on dorsal surface. Hind femur 3.6 times as long as medial width. Hind tarsal claw pectinate to about 0.5. Angulation of fore wing vein 2m-cu strong.
Tergite 1 of metasoma 1.6 times as long as apical width. Dorsal longitudinal carinae of T1 extending to spiracle (0.4 length of tergite), dorsolateral longitudinal carinae extending just over half way to spiracle (0.3 distance of T1).
Yellow. Legs and metasoma slightly darker yellow apically/ posteriorly. Vertex and supra-antennal area medially, scape and mesoscutum orange. Pedicel and apex of mandible dark brown. Flagellum medium brown except base of first flagellomere orange. Entire membrane of fore wing moderately infumate (Fig. 46 Diagnosis. Zambion hirtum can be distinguished from all other Zambion spp. in that the supraclypeal area dorsomedially has a short, longitudinal lamella between the an-tennae (Fig. 23) (lamella absent in all other species). In addition, Z. hirtum is the only species that has the following colour combination: mesosoma and metasoma completely light coloured (yellow or orange) (Fig. 4) AND head yellow with black markings dorsally (Fig. 16). Description. Adult. Female (only holotype known). Fore wing length 7.1 mm. Clypeus separated from supraclypeal area by a weak groove (Fig. 10). Malar space 0.5 times basal width of mandible. Supraclypeal area 1.7 times as wide as high (Fig. 10), dorsomedially with a short, narrow, longitudinal lamella between the antenna (Fig. 23). Supraantennal horn triangular in dorsal view (Fig. 16), 1.0 times as long as width of pedicel at midheight in lateral view (Fig. 24). Ocello-ocular distance 1.6 times ocellar diameter. Gena 0.7 times transverse diameter of the eye. Antennal flagellum with 40 segments.
Notaulus absent. Epicnemial carina mostly straight medially (similar to Z. rogeri shown in Fig. 32), not curving away from anterior edge of mesopleuron near ventral corner of pronotum. Scutellum with lateral carinae at base only. Medial longitudinal carinae of propodeum strong and complete anterior to posterior transverse carina (Fig.  34). Carina extending from pleural carina to spiracle absent. Posterior transverse carina of propodeum with lateral abscissa roughly straight (Fig. 34), point of union with pleural carina at about 0.2 distance from posterior end of pleural carina to propodeal spiracle. Ventral transverse carina extending from metasomal foramen to pleural carina present (Fig. 35). Fore tibia with a weak apical point on dorsal surface. Hind femur 3.2 times as long as medial width (original description = 3.8). Hind tarsal claw pectinate to 0.6 times length of claw. Angulation of fore wing vein 2m-cu strong (Fig. 38).
Tergite 1 of metasoma 1.6 times as long as apical width. Dorsal longitudinal carinae of T1 extending close to spiracles (about 0.4 times length of tergite), dorsolateral longitudinal carina extending just over half way to spiracles (similar to Fig. 40 of male Z. hirtum).
Yellow. Pedicel, medial 0.5 of supra-antennal area except dorsomedial part of horn (Fig. 16) and medial 0.3 of occiput black to dark brown. Flagellum and dorsomedial area of supra-antennal horn brown. Medial 0.5 of vertex posterior to ocelli, mesoscutum and apical 2 tarsomeres of fore and mid legs and apical 3 tarsomeres of hind leg brownish yellow. Membrane of fore wing hyaline, except slightly infumate in apical 0.3 (Fig. 38). Body covered with dense golden setae.
Male: as female except fore wing length 4.9 to 7.1 mm. Malar space 0.4 times basal width of mandible. Width of supraclypeal area 1.7 to 2.0 times height. Ocelloocular distance 1.7 to 1.9 times ocellar diameter. Gena 0.6 times transverse diameter of eye. Flagellum with 36 to 40 segments. Notaulus weak anteriorly in some specimens. Lateral abscissa of posterior transverse carina of propodeum slightly sinuous (Fig. 35) or completely straight (Fig. 34), joining pleural carina at 0.2 to 0.4 distance from posterior end of pleural carina. Hind femur length 3.0 to 3.4 times medial width. Hind tarsal claw pectinate in basal 0.4 to 0.5 of claw. Tergite 1 1.3 to 1.7 times as long as apical width. Dorsolateral longitudinal carina of T1 extending less than half way to spiracle (Fig. 40) or a little bit more than half. Colour as female except, in some specimens medial 0.5 of vertex light brown to dark brown and occiput more extensively brown (up to medial 0.5). Supra-antennal horn completely black to dark brown in some specimens.
Comments. Z. hirtum is a mostly pale-coloured species, most similar in colour to Z. kasparyani, although the latter species is completely yellow to orange dorsomedially on the head (Fig. 47), whereas Z. hirtum is extensively black in this area (Fig. 16). Zambion hirtum is the species with the most known specimens (eight) and is the only species known from two countries. Note that only six of the eight specimens were examined in this study. Also note that the supraclypeal longitudinal lamella of Z. hirtum is not homologous with the supra-antennal lamella of Ibornia. The latter is positioned much more dorsally on the head (extending nearly to the medial ocellus) and its dorsal margin is strongly thickened with a longitudinal groove (Townes, 1969).
Notaulus absent. Epicnemial carina mostly straight medially, not curving away from anterior edge of mesopleuron near ventral corner of pronotum (Fig. 32). Scutellum with lateral carinae 0.4 to 0.5 (0.5) length of scutellum. Medial longitudinal carinae of propodeum complete and strong anterior to posterior transverse carina (as in Z. hirtum shown in Fig. 34). Carina extending from pleural carina to propodeal spiracle absent. Posterior transverse carina of propodeum with lateral abscissa roughly straight, point of union with pleural carina at about 0.4 distance from posterior end of pleural carina to propodeal spiracle (similar to Z. monodon shown in Fig. 36). Ventral transverse carina extending from metasomal foramen to pleural carina present (as in Fig. 35). Fore tibia with a weak apical point on dorsal surface. Hind femur 3.5 to 3.7 (3.6) times as long as medial width. Hind tarsal claw pectinate to 0.4 times length of claw. Angulation of fore wing vein 2m-cu moderate.
Tergite 1 of metasoma 1.6 to 1.8 (1.6) times as long as apical width. Dorsal longitudinal carinae of T1 extending to about the level of spiracle or a bit beyond, 0.4 to 0.6 (0.6) times length of T1, dorsolateral longitudinal carinae extending to  spiracle, or in holotype to at least 0.7 times length of T1 (posterior to this, present as a rounded ridge).
Black to dark brown. Extreme apex of hind femur, basal 0.2 of hind tibia and posterior 0.2+ of T3-T7 ivory (the ivory bands longer medially than laterally and increasing in length from T3 to T7) (Fig. 6). Scape, pedicel and flagellum ventrally, mandibles except at apex, palpi, pronotal collar medially, tegula, posterior of subtegular ridge, fore leg, mid leg, hind coxa, trochanter and femur orange (middle and hind trochanters and femora brownish orange, especially medial surface of hind trochanter and femur). Flagellum dorsally, apical 0.8 of hind tibia and all of hind tarsus brown (flagellum darkening from base to apex). Membrane of fore wing slightly infumate except moderately infumate in apical 0.3 (Fig. 6). Body covered with dense silver setae. Etymology. This species is named in honour of the senior author's father, Dr. Roger Bennett, for his support and encouragement during the long (and sometimes dark) journey towards becoming a systematic entomologist.
Comments. Zambion rogeri is most closely related to Z. eileenae on the basis of the similar short triangular horn of the supra-antennal area and the length of the longitudinal carinae of T1. Even though Z. rogeri is only known from males, and Z. eileenae from a single female, it is highly unlikely that they are conspecific. Zambion rogeri has extensive yellow colouration on the legs and ivory bands on the metasoma, whereas Zambion eileenae is completely dark. In the three species of Zambion for which both males and females are known, there are minimal sex-related colour differences and this is also the case for species in the related genera Ibornia (Seyrig 1935) and Thibetoides (Kasparyan 1973). In addition, the malar space of Z. rogeri is 0.4 to 0.5 times the basal width of the mandible compared to 0.7 for Z. eileenae. Zambion rogeri also lacks the carina extending from the pleural carina to the propodeal spiracle, whereas this carina is complete in Z. eileenae.
Description. Adult. Female. Fore wing length 6.0 mm. Clypeus separated from supraclypeal area by a strong groove (Fig. 13). Malar space 0.7 times basal width of mandible. Supraclypeal area 1.9 times as high as wide (Fig. 13), dorsomedially without a short, narrow, longitudinal lamella between the antenna. Supra-antennal horn short and triangular in dorsal view (Fig. 19), 0.3 times as long as width of pedicel at  midheight in lateral view (Fig. 26). Ocello-ocular distance 1.8 times ocellar diameter. Gena 0.8 times transverse diameter of the eye. Flagellum with 29 segments on left, 28 on right.
Notaulus absent. Epicnemial carina mostly straight medially, not curving away from anterior edge of mesopleuron near ventral corner of pronotum (similar to Z. rogeri shown in Fig. 32). Scutellum with lateral carinae extending 0.5 length of scutellum. Medial longitudinal carinae of propodeum complete and strong anterior to posterior transverse carina (as in Z. hirtum shown in Fig. 34). Carina running from pleural carina to propodeal spiracle complete. Posterior transverse carina of propodeum with lateral abscissa roughly straight, point of union with pleural carina at about 0.4 distance from posterior end of pleural carina to propodeal spiracle (similar to Z. monodon shown in Fig. 36). Ventral transverse carina extending from metasomal foramen to pleural carina present (as in Fig. 35). Fore tibia without an apical point on dorsal surface. Hind femur 3.6 times as long as medial width. Hind tarsal claw pectinate to about 0.5 times length of claw. Angulation of fore wing vein 2m-cu weak.
Tergite 1 of metasoma 1.8 times as long as apical width. Dorsal longitudinal carinae of T1 extending to 0.7 length of T1, dorsolateral longitudinal carinae extending beyond spiracles up to 0.7 length of segment.
Black. Legs, metasomal sternites and ovipositor sheaths at base brown, the legs lightening from base to apex except hind leg with tibia and tarsus darker brown. Flagellomere 1 at extreme base and flagellomeres 2+ on ventral surface yellow brown, gradually lightening towards apex, dorsal surface of flagellum brown, lightening to yellow brown apical to middle. Apical 0.3 of clypeus orange-brown. Membrane of fore wing strongly infumate (Fig. 7), slightly less imfumate basally. Body covered with dense silver setae.
Etymology. This species, collected in the "Mountains of the Moon", is named in honour of the senior author's mother, Mrs. Eileen Bennett, for her support and understanding, especially when finding dead caterpillars and grasshoppers in jars or worse (live ones wandering through the house).
Notaulus present, but weak. Epicnemial carina medially curving away from anterior edge of mesopleuron near ventral corner of pronotum (Fig. 33) (not mostly straight medially as in Fig. 32). Scutellum with lateral carinae to 0.3 length of scutellum. Medial longitudinal carinae of propodeum incomplete anterior to posterior transverse carina (present between posterior transverse carina and the level of the propodeal spiracles, but absent anterior to this point). Carina running from pleural carina to spiracle absent. Posterior transverse carina of propodeum with lateral abscissa roughly straight, point of union with pleural carina at about 0.4 distance from posterior end of pleural carina to propodeal spiracle (similar to Z. monodon shown in Fig. 36). Ventral transverse carina extending from metasomal foramen to pleural carina present (as in Z. hirtum shown in Fig. 35). Fore tibia with a moderately strong apical point on dorsal surface. Hind femur 3.8 times as long as medial width. Hind tarsal claw with pectination not known (both distal tarsomeres missing). Angulation of fore wing vein 2m-cu strong.
Tergite 1 of metasoma 1.2 times as long as apical width. Dorsal longitudinal carina of T1 extending to about level of spiracle (0.4 length of segment), dorsolateral longitudinal carina absent (no obvious carina visible between anterior part of dorsal longitudinal carina and spiracle).
Male as female except: Fore wing length 6.2 to 6.3 mm. Malar space 0.4 to 0.5 times basal width of mandible. Supraclypeal area 1.9 to 2.0 times as wide as high. Supra-antennal horn 1.3 times width of pedicel at middle. Ocello-ocular distance 1.8 to 2.0 times ocellar diameter. Gena 0.8 to 0.9 times transverse diameter of eye. Flagellum with 27 segments. Hind femur length 3.7 to 3.8 times medial width. Hind tarsal  Etymology. This species is named in honour of Dr. David Wahl (American Entomological Institute) (AEIC) for his contributions to ichneumonid taxonomy and his hospitality to the senior author during many visits to the AEIC.
Comments. Zambion wahli is most closely related to Z. broadi on the basis of the long, sub-parallel-sided horn (Figs 20, 21), the curved epicnemial carina (Fig. 33) and the lack of a complete medial, longitudinal carina of the propodeum anterior to the posterior transverse carina. See comments under the latter species for distinction of these two species.
Description. Adult. Male. Fore wing length 6.4 mm. Clypeus separated from supraclypeal area by a strong groove. Malar space 0.4 times basal width of mandible. Supraclypeal area 1.9 times as wide as high (Fig. 15), dorsomedially without a short, narrow, longitudinal lamella between the antenna. Supra-antennal horn long and sub-parallel-sided in dorsal view (Fig. 21 Notaulus present, but weak (Fig. 29). Epicnemial carina medially curving away from anterior edge of mesopleuron near ventral corner of pronotum (similar to Z. wahli shown in Fig. 33). Scutellum with lateral carinae at base only. Medial longitudinal carinae of propodeum absent anterior to posterior transverse carina. Posterior transverse carina incomplete medially. Carina extending from pleural carina to propodeal spiracle absent. Posterior transverse carina of propodeum with lateral abscissa strongly curving posteriorly where it joins pleural carina, point of union at about 0.2 distance from posterior end of pleural carina to propodeal spiracle (Fig. 37). Ventral transverse carina extending from metasomal foramen to pleural carina present (Fig.  37). Fore tibia apical point not examined (both fore tibiae absent). Hind femur 3.6 times as long as medial width. Hind tarsal claw pectinate to 0.4 times length of claw. Angulation of fore wing vein 2m-cu strong.
Tergite 1 of metasoma 1.2 times as long as apical width. Dorsal longitudinal carinae of T1 extending to about level of spiracle (0.4 length of tergite), dorsolateral longitudinal carinae present only as a short stub bifurcating near base of dorsal longitudinal carina.
Black. Apical half of clypeus, glossa, apex of scutellum, propodeum, metasoma, except as noted below, orange. Tegula, coxae, apical 0.2 of hind tibia, hind tarsus except basal 0.3 of basal tarsomere, T2+ in spots laterally and sublaterally, T5 to T6 medially and hypopygium brown. Palpi, hind trochanter, femur, basal 0.8 of hind tibia and basal 0.3 of basal tarsomere, sternites and gonoforceps brownish yellow. Note that the distinction between orange and brownish yellow is not always clear. The hind leg is slightly more yellowish than the propodeum, the latter being slightly more yellow (and slightly less orange) than the scutellum and tergites. Membrane of fore wing uniformly, moderately infumate (Fig. 9). Body covered with dense golden setae. Etymology. This species is named in honour of Dr. Gavin Broad (NHM) in recognition of his contributions to ichneumonid taxonomy and for finding the type specimen of this species within the unidentified ichneumonids of the Natural History Museum, London.
Comments. Zambion broadi is most closely related to Z. wahli as noted under the comments for that species. The differences are the colour of the propodeum (compare Figs 8 and 9), the shape and location of the lateral abscissa of the posterior transverse carina of the propodeum, and the complete lack of the medial longitudinal carina dorsal to the posterior transverse carina of the propodeum in Z. broadi (present, but incomplete in Z. wahli). It is possible that Z. broadi is conspecific with Z. wahli, but we think this unlikely given the difference in both colour and propodeal carination. When both of these characters differ in related taxa, it is a good indicator that two species are present (e.g., in Dyspetes Förster) (He and Wan 1987).    Fig. 36).