Revision of the Afrotropical species of Pristomerus (Ichneumonidae: Cremastinae), with descriptions of 31 new species

We revised the species of the genus Pristomerus Curtis, 1836 in the Afrotropical Region. Fortynine species are recognized, of which 31 are newly described. The following new species are described: P. afrikaner, P. aka, P. babinga, P. bemba, P. dikidiki, P. herero, P. hutu, P. kagga, P. khoikhoi, P. kuku, P. masai, P. mbaka, P. mbama, P. mboum, P. nzakara, P. protea, P. restio, P. san, P. sara, P. sotho, P. swahili, P. teke, P. tutsi, P. venda, P. wolof, P. xhosa, P. yakoma, P. yangere, P. yoccolo, P. zande and P. zulu spp. nov. New synonyms: P. africator, P. cunctator and P. luteolus are new junior synonyms of P. pallidus. New records: new host records are reported for the widespread P. pallidus; new country-level distribution records are added for P. bullis, P. keyka, P. kelikely and P. pallidus; and reports of Trathala concolor and P. veloma in South Africa are shown to have been erroneous. An illustrated dichotomous key to females is provided; an online Lucid interactive matrix key is also available at www.waspweb.org. Finally, the ecological and geographical correlates of colour patterns exhibited by Pristomerus in the Afrotropical region are discussed.


Introduction
Most of the biodiversity in the tree of life evolved in the tropics and diversified later into temperate latitudes (Jansson et al. 2013). The genus Pristomerus Curtis, 1836 is no exception, with Townes (1971) already stating that its diversity peaks in the tropics. Paradoxically only eight of the approximately 100 described Pristomerus species are recorded from the Afrotropical region (Yu et al. 2012). This gap between expected and documented species richness pinpoints the poor knowledge we still have of the entomofauna in this region. This general rule concerning entomology in Subsaharan Africa is even more evident when considering the huge family of Ichneumonidae, especially when dealing with diverse genera such as Pristomerus.  (Morley, 1912) 24. Malar line exceptionally long, longer than mandible base (ML > 1) ( -Female femoral tooth weak to moderate, at least distinct as a subapical ventral protuberance (a) and distinctly shorter than basally wide, sometimes followed by secondary denticles (FFT = 1) (b) ... 34 -Female femoral tooth strong, about as high as wide (aa) to strongly higher than basally wide (bb), always followed by secondary denticles (  Pristomeridia Ashmead, 1900: 100. Pristocelus Szépligeti, 1905: 48. Neopristomerus Viereck, 1912: 592. Nesanomalon Morley, 1913.

Diagnosis (updated from Gauld 2000)
Female Small to moderate sized species, fore wing length 2-8 mm; main background colour yellow to black; head with mandible not twisted, without obvious ventral flange, teeth subequal to upper tooth slightly longer than lower tooth; palpi formula 5:4; frons biconcave, sometimes with a mid-longitudinal ridge that may be strongly raised; occipital carina mid-dorsally complete to narrowly obsolescent, ventrally joining hypostomal carina at to distinctly above mandible base; antenna with flagellum finely setose; inner margins of eyes from slightly converging to distinctly diverging ventrally; mesosoma punctate with smooth or coriaceous parts to extensively punctate; notaulus weakly to strongly impressed; pronotum unspecialized with epomia usually discernible and quite short; scutellum flat to moderately convex, without lateral carina; epicnemial and postpectal carinae complete; propodeum with carination usually complete, sometimes longitudinal carinae partially obsolescent; tarsal claws unspecialized, pectinate; mid tibia with two apical spurs; hind femur often with a ventral subapical tooth, often followed by auxiliary denticles; fore wing with areolet open (vein 3 rs-m absent); pterostigma stout, usually as broad or broader than 1 st subdiscal cell; distal abscissa of M from virtually absent to complete to wing margin; abscissa of M between rs-m and 2m-cu usually as long to far longer than rs-m, rarely distinctly shorter; hind wing with distal abscissae of Rs, M, Cu1 and 1A usually absent, almost invariably without a trace of the distal abscissa of Cu1 basally so that Cu&cu-a is not broken but smoothly curved; metasoma with tergites 1-2 usually aciculate and following ones coriaceous or granulate; first segment moderately stout, ventral margins of tergite 1 widely separated so that sternite 1 is exposed for its full length, glymma distinct in front of spiracle; tergite 2 slender to moderately stout, slightly depressed, with welldefined thyridium subcircular to elongate, close to anterior margin, at most separated from it by its own maximum diameter; laterotergite membranous and folded up; ovipositor 0.8-3.2× as long as hind tibia, straight to slightly down-curved but sometimes evenly bi-curved, its apex usually weakly to strongly sinuous but sometimes straight.

Male
Ocelli usually strongly enlarged, inner margins of eyes sometimes distinctly more convergent, mesoscutum sometimes strongly smoother, area superomedia sometimes more slender, hind femur stouter and femoral tooth stronger, nearly always present; gonosquama apically rounded, without distal lobe. Otherwise similar to female.

Differential diagnosis
Pristomerus is usually an easily identifiable ichneumonid genus, mostly defined by the presence of thyridia close to the anterior margin of tergite 2 and the separation of the ventral margins of tergite 1. Apart from a few exceptions, the apex of the ovipositor is also distinctly sinuous. This derived character is present in other ichneumonid and braconid subfamilies (Tersilochinae, Doryctinae, Braconinae…), and is probably an adaptation enabling the female to actively bend the ovipositor tip and thus to forage more efficiently within the host's hideaway (Quicke 1991). The only other cremastine genus with comparable features is the Neotropical Xiphosomella Szépligeti,1905, in which the thyridia are distinctly removed from the anterior margin of tergite 2, and the ovipositor tip is rarely sinuous. Some extra-limital Xiphosomella cannot, however, be unambiguously separated from Pristomerus (Gauld 2000). When present, the femoral tooth is also a reliable feature, though many Pristomerus females lack it. The only other cremastine genus in the Afrotropical region with a ventral tooth on the femur is the Malagasy genus Fafana: in the only known Fafana Rousse, Villemant & Seyrig, 2011 species, the sculpture is almost entirely smooth, the distal abscissa of Cu1 reaches the hind wing margin and the ovipositor tip is straight.

Phylogeny
No comprehensive phylogenetic analysis of the subfamily Cremastinae has been published. The generic identification of Pristomerus is usually easy, but not one of the diagnostic morphological features may be unambiguously considered as a synapomorphy. The presence of thyridia (absent in most other cremastine genera), is considered as a plesiomorphic feature within Ichneumonidae (Gauld 1985). The sinuous ovipositor tip and the femoral tooth cannot be considered as derived characters, because they are not restricted to Pristomerus and are totally absent in some Pristomerus females. It is therefore impossible to ascertain whether the genus is monophyletic or not. Hence, Pristomerus is either a widespread and rather variable clade or part of a paraphyletic clade including Xiphosomella and other smaller cremastine genera.

Distribution, species richness and ecology
Including the three new synonyms and 31 descriptions in the present revision, the genus now comprises 140 species. This is a worldwide genus, most diversified in the tropics, which contain about two thirds of the currently known species. All known hosts are small lepidopterans (including several major pests) living in leaf rolls, tunnels, silk shelters or other concealed substrates. The biology of some widespread Pristomerus parasitizing agricultural pests has been documented: for example see Smith & Johnson (1986) and Cave (1995) for P. spinator (Fabricius, 1804), or Rosenberg (1934) and Athanassov et al. (1997)

Differential diagnosis
Large yellow to reddish-testaceous species, differentiated from all other Afrotropical species by the combination of the strong and narrow femoral tooth in the female, the blunt transverse rugosities on the propodeum, the long malar line, the long antenna and the extensively punctate mesonotum.

Differential diagnosis
Small and mostly yellowish species with large black dorsal markings; differentiated from all other Afrotropical species by the combination of the straight ovipositor tip, the narrow clypeus and the converging inner margins of eyes.

Differential diagnosis
Large and mostly yellow to orange species; differentiated from all other Afrotropical species by the combination of the rather strong femoral tooth, the ventrally diverging inner margins of eyes, the strongly transverse clypeus, the numerous flagellomeres and the long ovipositor. The only other Afrotropical species with such a transverse clypeus is P. bullis, in which the femoral tooth and the ovipositor are shorter.

Male
Unknown.

Distribution
Central African Republic.

Differential diagnosis
Small testaceous and black species with inner orbits lighter, pale yellow, and hind legs noticeably mottled; differentiated from all other Afrotropical species by the combination of this unique colour pattern, the largely coriaceous mesoscutum with scutellum punctate, the strong femoral tooth and the antenna with few flagellomeres.

Differential diagnosis
Moderately sized and yellow to orange overall; differentiated from most other Afrotropical species by the very transverse clypeus. It is thus apparently closely related to P. babinga sp. nov. but P. bullis has a significantly shorter ovipositor.

Differential diagnosis
Rather small species from Madagascar, mostly yellowish to testaceous-orange with basal tergites blackened and whitish head, though dorsally and often medially darkened. Differentiated from most other Afrotropical species by the absence of the femoral tooth in females, the whitish face, the mostly yellowish-orange metasoma and the moderately long ovipositor. It is hence morphologically related to P. vahaza, which has a narrower clypeus, a more slender mesosoma and a rather flat scutellum, and to P. ranomafana, which is darker with a significantly longer ovipositor.

Distribution
Madagascar. Comments (from Rousse et al. 2013) The large series of specimens available in the MNHN collection shows considerable variation with respect to the original description of P. caris. The propodeum carination is highly variable, the area basalis being petiolate to truncate, and the area superomedia greatly varies in elongation. Colour also varies from lighter specimens with reduced brownish markings to darker ones with larger black spots on face, vertex, mesoscutum and propodeum. We furthermore observed two male groups with differently sized ocelli. Such variability is, however, not correlated with any other morphological or geographical difference. These specimens might represent a species complex whose study needs more comprehensive molecular and morphometric investigations.

Diagnosis
Large; mostly black with legs mostly testaceous and trochanters and apex of femora pale dotted; anterior edge of pterostigma whitish; face densely punctate, clypeus almost totally punctate, sculpture smoother on remainder of head; clypeus transverse; malar line moderately short; inner margins of eyes parallel; antenna with 38 flagellomeres, penultimate flagellomere quadrate; mesosoma densely and deeply punctate with a smoother dorsal area on pronotum, ventral half of speculum smooth; area superomedia rather stout, weakly carinate laterally; female femoral tooth very strong; ovipositor moderately long, strongly sinuous. Male unknown.

Differential diagnosis
Large and mostly black species, differentiated from most other Afrotropical species by the combination of the colour pattern, the strong femoral tooth and the extensively punctate mesonotum. It is morphologically closely related to P. nzakara sp. nov., whose ovipositor is distinctly stouter with a straight apex.
Colour. Head black with frontal orbit orange, mandible and palpi yellow; mesosoma black; metasoma black with apical and lateral margins of tergites 3 and following testaceous-orange; fore and mid legs testaceous, hind leg mostly dark brown; trochanters and apex of femora pale dotted; wings apically infuscate, venation pale brown, anterior edge of pterostigma whitish.
Head. Face deeply and densely punctate, mid-longitudinally weakly bulging; inner margins of eyes parallel; clypeus transverse, entirely punctate; malar line short; frons finely and densely punctate with a weak mid-longitudinal ridge from upper face to median ocellus; vertex and temple coriaceous; occipital carina joining hypostomal carina distinctly above mandible base; antenna with 38 flagellomeres, penultimate flagellomere quadrate.

Male
Unknown.

Differential diagnosis
Rather large Madagascan species, differentiated from all other Afrotropical species by the atypical colour pattern: head mostly white, mesosoma testaceous orange and metasoma mostly black with apical margins of tergites white. The only other Afrotropical species with a somewhat similar colour pattern is P. zulu sp. nov., whose face is medially reddish-testaceous and the malar line is distinctly shorter.

Differential diagnosis
White faced species from Madagascar, chiefly characterized by the very long ovipositor, much longer than in any other Afrotropical species of Pristomerus.

Distribution
Madagascar.

Diagnosis
Large; mottled yellow to reddish-orange with numerous variable black markings on entire body; face deeply and densely punctate, punctation slightly sparser on clypeus; clypeus weakly transverse; malar line long; remainder of head coriaceous except inter-ocellar area deeply and densely punctate; antenna with 31-33 flagellomeres, penultimate flagellomere shorter than wide; mesosoma coarsely and densely punctate but pronotum mid-dorsally and speculum ventrally smooth; area superomedia elongate and not carinate postero-laterally; female femoral tooth strongly reduced, reduced to a subapical point followed by denticles; ovipositor very long, its apex sinuous. Male with mesosoma more shallowly punctate, inner margins of eyes distinctly diverging ventrally, ocelli enlarged, femur and femoral tooth strongly stouter, and area superomedia fully carinate.

Differential diagnosis
Large and mostly yellow and reddish-orange; differentiated from all other Afrotropical species by the combination of the coarse punctation on face and mesosoma, the near absence of the femoral tooth in the female and the very long ovipositor. Probably related to P. roberti in Madagascar, which is, however, far smaller with a stronger femoral tooth and a fully delimited area superomedia in both sexes.
Colour. Background colour mottled yellow and reddish-orange with face brown and variable black markings (sometimes strongly reduced) on: frons, ocellar area, occiput, anterior margin of mesosternum and mesopleuron, ventral margin of metapleuron, axillar trough, metanotum and basal margin of propodeum, and basal half of tergites 1-3; legs yellowish orange, hind tibia somewhat darker and hind tarsus strongly infuscate; flagellum and ovipositor sheath strongly infuscate; wings hyaline, venation brown.
Head. Face very densely and deeply punctate, punctation slightly sparser on median bulge; inner margins of eyes weakly diverging ventrally; clypeus weakly transverse, long, punctation sparser and shallower than on face; malar line long; frons, vertex and temple coriaceous except inter-ocellar area deeply and densely punctate; occipital carina joining hypostomal carina at mandible base; antenna with 32-33 flagellomeres, rather thick, penultimate flagellomere shorter than wide.

Diagnosis
Most of body ventrally and laterally mostly pale yellow to yellowish-orange, dorsally nearly entirely black; face except orbits dark brown; fore and mid legs yellowish-orange, hind legs mostly dark brown to black, femora and trochanters apically white-dotted; anterior edge of pterostigma whitish; face laterally densely punctate, clypeus and middle of face sparsely and finely punctate; clypeus weakly transverse, convex in profile; malar line short; remainder of head coriaceous; antenna with 28-31 flagellomeres; mesosoma moderately to densely punctate but pronotum almost entirely smooth, speculum ventrally smooth, and area superomedia usually transversely striate; area superomedia elongate, usually hardly delimited laterally beyond anterior transverse carina; female femoral tooth weak but distinct; ovipositor moderately long, apically moderately sinuous. Male with ocelli and hind femur enlarged, femoral tooth long and acute, otherwise similar to female.

Differential diagnosis
Moderately sized, ventrally yellow to orange and dorsally black. Differentiated from all other Afrotropical Pristomerus species by the combination of the light facial orbits, the always black scutellum, the very short malar line, and the weak though distinct female femoral tooth.
Colour. Background colour of head yellowish-orange with face often medially dark brown, mandible, malar space, clypeus and frontal orbit sometimes pale yellow, and frons, inter-ocellar area, and occiput black; background colour of mesosoma orange, dorsally entirely black except sometimes notaulus and lateral margin of mesoscutum, metanotum and a variable basal part of propodeum; tergites 1-2 nearly entirely black, following tergites pale yellow to orange with a large median black spot of variable extent; fore and mid legs yellowish-orange, hind leg mostly dark brown to black, all trochanters and femora apically white; wings hyaline, venation brown, pterostigma more or less whitish anteriorly.

Distribution
Central African Republic, Uganda.

Differential diagnosis
Moderately sized species with a characteristic colour pattern, rufo-testaceous with large black markings; differentiated from all other Afrotropical species by the combination of the very long malar line and the strong femoral tooth in females. The reduced ocelli and the dense punctation over the entire mesosoma are also helpful for identification.

Etymology
Kagga is old Norse for a keg or casket and is taken from the name of the type locality.
MesosoMa. Moderately elongate; pronotum almost smooth with some sparse punctures along dorsal, posterior and ventral margins; mesopleuron and metapleuron densely, deeply and evenly punctate, speculum smooth except some punctures dorsally; mesoscutum and scutellum densely, deeply and evenly punctate, apical third of mesoscutum with punctures somewhat confluent into weak transverse rugosities, notaulus relatively strong; propodeum densely, deeply and evenly punctate with area superomedia smoother centrally, area superomedia elongate, weakly delimited laterally beyond anterior transverse carina. Legs. Femoral tooth very strong, obviously higher than basally wide, followed by strong denticles.

Male
Unknown.

Distribution
South Africa (Western Cape).

Differential diagnosis
Small and entirely yellow to yellowish orange species; differentiated from most other Afrotropical species by the absence of the femoral tooth in females, the short malar line, the strongly transverse clypeus and the rather long ovipositor. It is closely related to P. moramora and P. venda sp. nov., from which it may be differentiated by the colour and antennal length characters given in the key (see comments below). It is also structurally very similar to P. sara sp. nov.; however, this species exhibits white-marked femora and tibiae, a colour pattern that appears to have ecological relevance (see general discussion).  48E 25.xii.1995-29.i.1996, S. van Noort, Malaise trap, Acacia / Commiphora / Combretum bushland, SAM-HYM-P048059" (SAMC); 1 ♀, same locality, "Kikolo plot, 04°08.72'S 38°01.37'E, 16 April-2 May 1996, S. van Noort, Commiphora woodland, Malaise trap, SAM-HYM-P016192" (SAMC).

Distribution
Madagascar. New record: Tanzania.

Comments
The three species P. kelikely, P. moramora and P. venda can only be keyed by rather weak though constant differentiating features. Pristomerus moramora is mostly separated from the other two by the extensive dark colouration of the mesosoma and metasoma. Furthermore, the males of P. kelikely and P. moramora have ocelli that are not enlarged, which is very unusual in Pristomerus and separate both from P. venda. However, several specimens in the SAMC and MNHN collections could not unambiguously be assigned to one of these three species: this species-complex needs further examination before being fully resolved. Rousse, Villemant & Seyrig, 2013 Fig. 15 Diagnosis (updated from Rousse et al. 2013) Small; pale yellow to testaceous orange with very variable dark markings on inter-ocellar area, around scutellum and on metasoma; flagellum dark brown, basally yellow; face shallowly punctate; inner margins of eyes subparallel; clypeus smooth, strongly transverse; malar line moderately long; occipital carina joining hypostomal carina shortly above mandible base; antenna short with 23-27 flagellomeres, penultimate flagellomere subquadrate; mesosoma moderately elongate, laterally densely and shallowly punctate-granulate including most of speculum but pronotum dorsally smooth, mesoscutum coriaceous with some punctures along notaulus, scutellum almost smooth; propodeum with area superomedia often weakly delimited laterally beyond anterior transverse carina; female femoral tooth small, followed by minute denticles; ovipositor long, its apex weakly sinuous. B 4.5-4.9; A 2.3-2.5; F 2.4-2.8; CT 2.2; ML 0.6; POL 1.0; OOL 1.3; Fl n-1 1.1; ASM 1.8; OT 1.8-2.0; FFT 1. Male with inner margins of eyes diverging ventrally, ocelli strongly enlarged, hind femur and femoral tooth stouter, scutellum quite smooth. POL 0.6; OOL 0.3.

Differential diagnosis
Small and mostly yellowish with variable dark dorsal markings; differentiated from all other Afrotropical species by the combination of the short antenna (basally yellow fading to dark brown), the mesoscutum coriaceous with only few punctures along the notaulus, the strongly transverse clypeus and the short female femoral tooth.

Differential diagnosis
Small and mostly black, apparently restricted to the south of South Africa. Differentiated from all other species of Pristomerus by the combination of the coriaceous mesoscutum, the smooth scutellum, the short antenna and the very long malar line.

Diagnosis
Moderately large; mostly black with fore and mid legs and metasoma from tergite 3 yellowish-orange; trochanters and femora apically white-dotted; pterostigma anteriorly white; face coarsely and densely punctate, sculpture sparser and smoother on remainder of head; malar line moderately long; frons with a strong median protuberance; antenna with 35 flagellomeres; mesosoma densely and coarsely sculptured, punctate to rugose-punctate, including pronotum and almost entire speculum; female femoral tooth nearly absent; ovipositor short, weakly sinuous apically. Male with ocelli slightly enlarged, femoral tooth strong, tergites 3 and following medially black; otherwise similar to female.

Differential diagnosis
Large black species, differentiated from all other Afrotropical Pristomerus by the absence of a femoral tooth in females, the short ovipositor and the strong protuberance on the frons. It is related to P. yangere sp. nov., which lacks the frontal protuberance and has a distinctly longer ovipositor.
Colour. Mostly black with metasoma orange from tergite 3 and pale yellow parts on head: a thin margin along frontal orbit, malar space, mandible and palpi; legs testaceous-brown but hind femur black and all trochanters and femora apically white-dotted; wings slightly infuscate, venation brown, pterostigma anteriorly white.
Head. Face mid-longitudinally bulging, coarsely punctate, punctures somewhat confluent into transverse rugosities; clypeus transverse, more sparsely punctate, distinctly convex in profile view, its ventral margin strongly convex; malar line moderately long; frons finely and densely punctate with a strong median protuberance: laterally compressed, about triangular, its tip rounded, about as high as basally wide; vertex and temple coriaceous; occipital carina joining hypostomal carina distinctly above mandible base; antenna with 35 flagellomeres, penultimate one slightly elongate.

Distribution
Central African Republic, Uganda.

Differential diagnosis
Moderately sized species from tropical mainland Africa; differentiated from all other Afrotropical species by the combination of the black dorsal markings (including most of the occiput in females), the rather enlarged ocelli with an unusually reduced POL, the short malar line, the small female femoral tooth, the long and narrow thyridium and the long ovipositor. It is apparently very closely related to other undescribed species (see comments) and to P. wolof sp. nov., from which it is separated by the size of the ocelli and femoral tooth as given in the key.

Distribution
Democratic Republic of the Congo, Guinea-Bissau, Ivory Coast, Kenya.

Comments
Nine morphologically related specimens in the SAMC collection (listed under material examined as excluded from type series), from Tanzania and Gabon, could not be definitively associated with P. masai sp. nov., but their morphological differences are also not strong enough to warrant description of a distinct species. Pristomerus masai sp. nov. is probably included within a widespread tropical species-complex; resolution of species delimitation within this complex requires thorough examination using molecular and morphometric tools.

Differential diagnosis
Moderately small and mostly black; differentiated from all other Afrotropical species by the very short and entirely straight ovipositor. Also characterized by the very short malar line, the coriaceous scutellum and the entirely punctate speculum.

Distribution
Central African Republic, Ivory Coast.

Differential diagnosis
Moderately small and mostly black; differentiated from all other Pristomerus species by the combination of the very short malar line, the entirely punctate speculum and almost entirely sculptured pronotum, and the very weak female femoral tooth. It appears closely related to P. mbaka sp. nov. in which the scutellum is impunctate and the ovipositor is significantly shorter, apically straight.

Male
Unknown.

Diagnosis
Small; yellow to yellowish-orange with dorsal black markings on head and metasoma; femora and trochanters apically white-dotted; anterior edge of pterostigma whitish; face densely punctate; clypeus smooth, moderately transverse; malar line very short; remainder of head coriaceous; mesosoma densely punctate to punctate-granulate, with speculum ventrally, most of pronotum and scutellum quite smooth; area superomedia long and narrow, hardly carinate laterally; femoral tooth small; ovipositor moderately short, apically strongly sinuous. Male unknown.

Differential diagnosis
Small, mostly yellowish-orange; differentiated from all other Afrotropical Pristomerus by the combination of the quite smooth scutellum, the short malar line, the small female femoral tooth and the rather enlarged ocelli. The apical white dot on the femora and the black occiput are also reliable identification cues, making P. mboum sp. nov. morphologically close to P. masai sp. nov. and related species.
Colour. Yellow to yellowish-orange overall, with face fading to dark brown medially and numerous black markings: frons, inter-ocellar area, occiput, tergite 2 medially and a basal spot on following tergites; femora and trochanters apically white-dotted; wings slightly infuscate apically, venation pale brown, anterior edge of pterostigma whitish.

Male
Not known with certainty (see comments).

Comments
The probable male from Uganda shares the colour pattern and the quite smooth scutellum and pronotum. It was, however, not included as a paratype because its head is missing and it therefore could not be unambiguously determined.

Differential diagnosis
Small species from Madagascar; differentiated from most other Afrotropical species by the absence of the femoral tooth in females, the very short malar line, the strongly transverse clypeus and the long ovipositor. It is closely related to P. kelikely and P. venda sp. nov., from which it may differentiated by the colour and antenna length characters given in the key (see comments on P. kelikely).

Madagascar.
Pristomerus nzakara sp. nov. urn:lsid:zoobank.org:act:719C9EFB-B4A0-4848-8B83-217E77DD93B6 Fig. 24 Diagnosis Large; black with lighter parts on head and metasoma fading to yellowish-orange from tergite 3; trochanters and apex of femora pale dotted; pterostigma anteriorly white; face moderately punctate, sculpture smoother on remainder of head; clypeus transverse; malar line moderately short; inner margins of eyes parallel; antenna with 36 flagellomeres, penultimate flagellomere quadrate; mesosoma densely and deeply punctate with a large smooth dorsal area on pronotum, ventral half of speculum and apical half of lateral lobe on mesoscutum almost smooth; notaulus noticeably deep and rugose; area superomedia stout, hardly carinate laterally; female femoral tooth very strong; ovipositor moderately long, exceptionally stout and entirely straight. Male unknown.

Differential diagnosis
Large, black species from tropical mainland Africa; chiefly differentiated from all other Afrotropical Pristomerus species by the very stout and apically straight ovipositor, also characterized by the long antenna, the deep and rugose notaulus, and the posteriorly smooth lateral mesoscutal lobe.
Colour. Head black with frontal orbit testaceous, mandible, palpi and malar space yellow; mesosoma black; metasoma black with tergite 3 and following fading laterally to testaceous-orange; fore and mid legs yellowish-orange, hind leg mostly black, with tibia and tarsus strongly infuscate; trochanters and apex of femora white dotted; wings apically infuscate, venation pale brown, pterostigma anteriorly white. Head. Face deeply and moderately punctate, mid-longitudinally bulging; inner margins of eyes parallel; clypeus transverse, smooth with some punctures dorsally, distinctly convex in profile view; malar line moderately short; frons finely and densely punctate with a weak mid-longitudinal ridge from upper face to median ocellus; vertex and temple coriaceous; occipital carina joining hypostomal carina distinctly above mandible base; antenna with 36 flagellomeres, penultimate flagellomere quadrate.

Male
Unknown.

Distribution
Central African Republic.

Diagnosis
Moderately small to moderately sized; yellow to reddish overall with variable dark markings on first tergites and often around scutellum, dark markings sometimes also present on propodeum and apical tergites; frons darker than face, mesosoma varying from almost uniformly yellowish to strikingly mottled yellowish and reddish, with notaulus and scutellum almost always lighter than remainder of mesonotum; face densely punctate; inner margins of eyes barely diverging ventrally; clypeus strongly transverse, densely but very shallowly punctate; malar line moderately long to long; remainder of head coriaceous to punctategranulate; occipital carina joining hypostomal carina shortly above mandible base; antenna long with 28-35 flagellomeres, penultimate flagellomere subquadrate; mesosoma moderately elongate and very densely punctate but pronotum dorsally and ventral half of speculum smooth; propodeum with area superomedia moderately stout to elongate; female femoral tooth moderate to very small, sometimes reduced to a barely distinct subapical tubercle; ovipositor moderately short, its apex strongly sinuous. B 7.4-12.1; A 4.7-6.8; F 4.2-6.4; CT 1.7-2.4; ML 0.6-0.7; POL 0.9; OOL 1.1; Fl n-1 1.1; ASM 1.8-2.3; OT 1.3-1.6; FFT 1. Male with inner margins of eyes more strongly diverging ventrally, mesoscutum most often weakly sculptured, mostly coriaceous with dense punctation on anterior edge of median mesoscutal lobe, punctation occasionally more extended; ocelli strongly enlarged, hind femur and femoral tooth stouter. POL 0.6; OOL 0.2.

Differential diagnosis
Mostly yellow to yellowish-orange with notaulus and scutellum most often usually distinctly lighter than remainder of mesoscutum; otherwise differentiated from all other Afrotropical species by the combination of the rather long malar line, the strongly transverse clypeus, the somewhat reduced ocellus, the weak female femoral tooth and the rather short ovipositor. Confusion may occur with P. moramora, P. kelikely or P. venda sp. nov., these three species having a shorter malar line and no trace of a femoral tooth in females; P. pallidus may also be confused with P. bullis in which the clypeus is significantly more transverse and the ovipositor shorter.

Comments
Pristomerus pallidus is distributed throughout the whole Afrotropical region and is very common in collections. It exhibits a noticeable intra-specific variation in the size of the femoral tooth in the female, although the tooth is always distinct and shorter than wide. The extent of the dorsal dark markings and the intensity of the reddish markings on the mesosoma are also variable. Finally, some males from Western Africa also have a mostly punctate mesoscutum. The holotype of P. africator (from Ivory Coast) is in the private collection of Jacques F. Aubert and was not available for examination. According to Horstmann (1990), the record from Egypt is doubtful. We could not examine this specimen, but we consider the record as probable because of the species' wide distribution.  Rousse, Villemant & Seyrig, 2013 Fig. 26 Diagnosis (range of variation from Rousse et al. 2013) Background colour yellowish-orange with some median dark spots on metasomal tergites; face densely punctate; inner margins of eyes subparallel; clypeus transverse, very sparsely punctate, inter-puncture spaces smooth; malar line long; remainder of head coriaceous; occipital carina joining hypostomal carina shortly above mandible base; antenna very long with more than 37-38 flagellomeres, penultimate flagellomere elongate; mesosoma moderately elongate, moderately to densely punctate, but pronotum and speculum almost entirely smooth and scutellum distinctly more sparsely punctate than mesoscutum; area superomedia weakly delimited laterally beyond anterior transverse carina; female femoral tooth small, followed by some minute denticles; ovipositor long, slightly sinuous apically. B 7.8-10.1; A 5.3-6.9; F 5.9-7.5; CT 1.8; ML 0.7; POL 0.7; OOL 1.1; Fl n-1 1.2; ASM 2.0; OT 1.8-1.9; FFT 1. Male with inner margins of eyes diverging ventrally, and ocelli, hind femur and femoral tooth enlarged. B 8.9-9.8; A 5.8-7.1; F 6.3-6.7: POL 0.6; OOL 0.4.

Differential diagnosis
Large yellowish-orange species from Madagascar; differentiated from all other Pristomerus species in the region by the combination of the large size, the antenna with numerous flagellomeres, the long malar line, the largely smooth speculum, the punctation on the scutellum, which is often distinctly sparser than on the mesoscutum, the area superomedia hardly delimited postero-laterally, the small female femoral tooth and the long ovipositor.

Diagnosis
Small; head and mesosoma black, metasoma mostly testaceous-brown with tergites 1-2 black; face and clypeus deeply and sparsely to densely punctate without granulate background sculpture; clypeus transverse; malar line very long; upper head punctate-granulate; ocelli exceptionally reduced; antenna with 26 flagellomeres, penultimate one shorter than wide; mesosoma entirely deeply and densely punctate except speculum ventrally smooth; propodeum with area superomedia shorter than wide and not laterally defined beyond anterior transverse carina; female femoral tooth absent; ovipositor moderately short, evenly up-curved and apically down-curved. Male unknown.

Differential diagnosis
Small and mostly black, highly atypical among Pristomerus because of the exceptionally reduced ocelli, the fore wing venation with the abscissa of M between rs-m and 2m-cu very short, and the ovipositor shape, evenly curved without the usual sinuous apex of most Pristomerus. It is only close to P. restio sp. nov., in which the face is lighter and more roughly punctate, and the area superomedia more slender.

Etymology
Refers to one of the most emblematic flowers of the Cape Floral Kingdom. Noun in apposition.
Colour. Head and mesosoma black with mandible and a thin margin along posterior orbit yellowishtestaceous; metasoma testaceous-brown with tergites 1-2 mostly black and posterior margins of tergites yellow; antenna blackish; fore and mid legs yellow; hind leg testaceous with coxa and trochanter black; wings hyaline, venation pale yellowish with pterostigma basally whitish.
MesosoMa. Mesosoma stout; entirely deeply and very densely punctate except ventral half of speculum smooth; area superomedia short, subquadrate, laterally not delimited beyond anterior transverse carina. Wings. Fore wing with 2m-cu subopposite 2rs-m, both separated by less than their own width. Legs. Female femoral tooth absent.

Male
Unknown.

Distribution
South Africa.

Differential diagnosis
Moderately sized species from Madagascar, mostly reddish-testaceous with basal tergites black, and head mostly white, posteriorly and mid-longitudinally black. Differentiated from most of other Afrotropical species by the absence of a femoral tooth in the female, the white orbits, and the very long ovipositor.
Morphologically related to P. vahaza, whose clypeus is narrower and mesosoma more slender, and to P. caris, whose ovipositor is significantly shorter.

Differential diagnosis
See P. protea sp. nov.

Etymology
Refers to another one of the most emblematic flowers of the Cape Floral Kingdom. Noun in apposition.

Male
Unknown.

Distribution
South Africa.

Differential diagnosis
Rather small, overall yellow to yellowish orange with metasoma basally black; differentiated from all other Afrotropical species by the combination of the rather strongly transverse clypeus, the long malar line, the small female femoral tooth and the markedly long ovipositor. Morphologically close to other Pristomerus species in Madagascar: P. kelikely and related species, which have a shorter malar line and no femoral tooth in the female; P. patator, which is distinctly larger with longer antennae and an incompletely carinate area superomedia; and the widespread P. pallidus, which has a shorter ovipositor and nearly always a differently coloured mesosoma with the notaulus and the scutellum paler than the remainder of the mesonotum. Pristomerus roberti is otherwise probably closely related to P. herero sp. nov. (see differential diagnosis of P. herero sp. nov.).  (Morley, 1912). Holotype, ♀. A. Habitus, lateral view. B. Head, mesosoma, dorsal view. C. Head, anterior view. D. Propodeum, dorsal view. E. Mesosoma, lateral view. F. Hind tibia, lateral view (inset: data labels). (Morley, 1912) Fig. 32

Differential diagnosis
Rather small, yellowish-orange and black species from the Seychelles Islands; readily differentiated from all other Afrotropical species by the colour pattern with the mesosoma dorsally black and ventrally yellowish-orange, the coriaceous mesoscutum, the strong femoral tooth and the short ovipositor.

Differential diagnosis
Moderately sized and yellowish orange with no defined dorsal dark markings; differentiated from all other Afrotropical species by the combination of the long malar line, the almost smooth pronotum, the deep and dense punctation, the absence of a femoral tooth in females and the rather short ovipositor. Morphologically close to P. moramora and related species which are also present in Southern Africa, but the longer malar line, the smooth pronotum and the deep dense punctation of P. san sp. nov. are reliable identification cues.
Colour. Yellowish-orange to orange overall with legs, mandible and clypeus somewhat paler, with sometimes reduced infuscate markings around scutellum and on basal tergites; flagellum and ovipositor sheath dark brown; wings hyaline, venation pale brown.

Distribution
South Africa.

Differential diagnosis
Small and mostly yellowish-orange species, differentiated from most other Afrotropical species by the combination of the strongly transverse clypeus, the rather short malar line, the shallow sculpture of face and pronotum, the absence of a femoral tooth in females and the rather long ovipositor. Morphologically, it is therefore very close to P. kelikely, from which it differs only by having the apices of the femora and the anterior half of the pterostigma strikingly white (see general discussion). Pristomerus sara sp. nov. seems also closely related to P. wolof sp. nov. (see differential diagnosis of P. wolof sp. nov.). Paratype CENTRAL AFRICAN REPUBLIC: ♀, same label data.

Male
Unknown.

Comments
This species is mostly differentiated from P. kelikely by the colour pattern, the femora being apically white-dotted and the anterior half of the pterostigma being white. We consider this pattern to be a significant ecological indicator (see general discussion).

Differential diagnosis
Moderately sized, black and reddish-orange species; readily differentiated from every other Afrotropical species by the unique colour pattern, with most of the head and mesosoma ventrally black, the remainder of the mesosoma being reddish-orange; also characteristic because of the combination of the densely hairy face and mandibles, the short ovipositor, the strong female femoral tooth and the punctate scutellum.
Colour. Head including antenna black with clypeus reddish-orange and mandible yellow; mesosoma dorsally reddish-orange, ventrally black; metasoma with tergites 1-2 black, following tergites reddishorange, medially blackened; legs orange with base and apex of hind tibia and hind femur blackened; wings hyaline, venation brown. Head. Face and clypeus densely punctate with very dense silver hairs obscuring most of sculpture; mandible also unusually hairy; inner margins of eyes parallel; clypeus transverse, punctation ventrally smoother; malar line long; frons and vertex sparsely punctate-granulate; temple coarsely coriaceous; occipital carina joining hypostomal carina distinctly above mandible base; both antennae broken.

Male
Unknown.

Differential diagnosis
Moderately sized, mostly yellowish to orange with numerous black markings dorsally; differentiated from all other Pristomerus species in the Afrotropical region by the combination of the quite smooth scutellum, the almost entirely smooth pronotum, the mostly coriaceous mesoscutum and the strong female femoral tooth.
Colour. Yellowish-orange to reddish-orange, with yellow and black markings; yellow to pale yellow: clypeus, mandible, facial orbits, lower gena, and sometimes pronotum, fore and mid legs; black: inter- ocellar area, occiput more or less extensively, mesosoma postero-dorsally, most of tergites 1-2 and a large basal part of following tergites; base and apex of hind tibia strongly infuscate; trochanters and femora apically white-dotted; wings hyaline, venation brownish, anterior edge of pterostigma whitish.
MesosoMa. Pronotum nearly smooth with anterior margin weakly crenulate and some punctures along posterior and dorsal margins; mesopleuron moderately punctate, with an oblique striate furrow below speculum, ventral half of speculum smooth; punctation slightly denser and coarser on metapleuron; mesoscutum alutaceous, somewhat longitudinally rugose posteriorly, with punctures along notaulus; notaulus moderate; scutellum quite smooth; propodeum moderately and deeply punctate, area superomedia centrally smoother and relatively short, area petiolaris transversely strigose. Legs. Femoral tooth strong, higher than basally wide, followed by a row of denticles.

Male
Unknown.

Differential diagnosis
Rather small and mostly black; readily differentiated among Afrotropical Pristomerus by the flat and coriaceous scutellum and the elongate penultimate flagellomere. Also characterized by the combination of the colour pattern, the elongate mesosoma, the moderate female femoral tooth, the short ovipositor, and the enlarged ocelli in females. It might be related to P. aka sp. nov., the latter having a mostly coriaceous mesoscutum, a punctate scutellum, and the ocelli not enlarged. Pristomerus teke sp. nov. is also strongly morphologically similar to P. mbaka sp. nov., the latter having a shorter and apically straight ovipositor, and not having the elongate mesosoma nor the flat scutellum of P. teke sp. nov..
Colour. Head mostly black with clypeus, malar space, mandible, frontal and temporal orbits yellow to orange; mesosoma black with some lighter orange markings along anterior margin of pronotum and reddish glints on mesopleuron; metasomal tergites black with apical and lateral margins of tergites yellow from tergite 3; fore and mid legs yellow, hind leg darker brown, with apices of all femora and trochanters white-marked; wings hyaline, venation brown with anterior half of pterostigma slightly paler; flagellum and ovipositor sheath dark brown.

Male
Unknown.

Differential diagnosis
Moderately large and mostly black; differentiated from all other Afrotropical species by the combination of the mostly smooth pronotum, the coarsely punctate-granulate mesoscutum, the short post-ocellar line, the absence of a femoral tooth in females and the long ovipositor.

Differential diagnosis
Moderately large, mostly yellowish orange and white-faced; differentiated from all other species by the combination of the white orbits, the narrow clypeus, the transverse head, the absence of a femoral tooth and the short ovipositor. Pristomerus vahaza appears to be related to P. albescens with which it shares the unusually transverse head and narrow clypeus, but these species cannot be confused because P. albescens has an apically straight ovipositor, a strongly reduced but distinct femoral tooth and a distinctly darker head.

Differential diagnosis
Large, reddish-orange with metasoma black; differentiated from all other Afrotropical Pristomerus species by the combination of that colour pattern, the distinctly diverging inner orbits in both sexes, the long antenna and the strong female femoral tooth. Pristomerus zulu sp. nov., in South Africa, shares a similar colour pattern, but is readily differentiated by its white and subparallel inner orbits.

Distribution
Madagascar.

Diagnosis
Moderately small; background colour yellow to testaceous with variable dark brown to black dorsal markings; head sparsely to moderately punctate; inner margins of eyes subparallel; clypeus strongly transverse; malar line short; antenna with 29-30 flagellomeres, penultimate flagellomere subquadrate; mesosoma moderately to densely punctate, pronotum hardly smoother postero-dorsally, and speculum ventrally smoother; propodeum with area superomedia sometimes weakly carinate postero-laterally; female femoral tooth absent; ovipositor moderately long, apically strongly sinuous. Male darker with ocelli, hind femur and femoral tooth strongly enlarged, inner margins of eyes strongly diverging ventrally, and area superomedia slightly more slender.

Differential diagnosis
Rather small and mostly yellowish-orange; differentiated from most other Afrotropical species by the combination of the absence of a femoral tooth in females, the short malar line, the strongly transverse clypeus and the rather long ovipositor. It is closely related to P. moramora and P. kelikely, from which it may be differentiated by the colour and antenna length characters given in the key (see comments).
Colour. Head yellow with face testaceous, and frons, vertex, temples and occiput darker brown, frons sometimes with a mid-longitudinal yellow stripe; mesosoma yellowish-orange with pronotum paler and axillary troughs slightly infuscate; metasoma yellowish-orange with basal tergites variously dark marked; legs yellow to yellowish-orange; wings hyaline, venation brown; ovipositor sheath black.

Distribution
Zimbabwe.

Comments
Pristomerus venda sp. nov. females are likely to be very difficult to differentiate from some P. kelikely and P. moramora. However, we keep it as a separate species because it could be unambiguously associated with males. These males exhibit two of the usual features of sexual dimorphism in Pristomerus spp. which are absent in the males of P. kelikely and P. moramora (ventrally diverging eyes and enlarged ocelli). See P. kelikely for further comments about this species-complex.

Diagnosis
Small to moderately small; pale yellow to yellowish with base of tergites 2-3 black; femora and trochanters apically white-dotted; pterostigma anteriorly paler; face and clypeus moderately to densely punctate, remainder of head more finely sculptured; clypeus strongly transverse; inner margins of eyes subparallel; malar line moderately short; ocelli small; antenna long with 32-34 flagellomeres; mesosoma moderately elongate, densely punctate-granulate but upper pronotum and ventral half of speculum smooth; female femoral tooth barely distinct; ovipositor long, apically strongly sinuous. Male with ocelli and femoral tooth hardly enlarged, inner margins of eyes slightly diverging ventrally, area superomedia more slender, and mesoscutum sculpture smoother.

Differential diagnosis
Rather to fairly small, yellow to pale yellow overall; otherwise distinguished from all other Afrotropical species by the strongly transverse clypeus, the weak but distinct female femoral tooth and the long ovipositor. Pristomerus wolof sp. nov. is part of the P. masai sp. nov. species-complex (see comments for P. masai sp. nov.): the reduced femoral tooth and barely enlarged ocelli in both sexes differentiate P. wolof sp. nov. from P. masai sp. nov. It is also somewhat morphologically similar to P. sara sp. nov., whose ovipositor and malar line are shorter and whose femoral tooth is totally absent in the female.

Differential diagnosis
Moderately sized, mottled yellow, black and reddish-testaceous; unmistakably characterized by this unique colour pattern, the extremely long malar line, the strongly reduced ocelli and the long clypeus, all features making P. xhosa sp. nov. highly characteristic within Pristomerus. The only confusion might occur with P. kaggasp. nov. , from which it may otherwise be differentiated by the very weak femoral tooth in P. xhosa sp. nov. females.
Colour. Head mostly reddish with mandible, malar space and frontal orbits pale yellow, and black markings on mid-dorsal face, tentorial pit, frons, inter-ocellar area, occiput and antenna; mesosoma mottled: background colour yellow variously fading to reddish-testaceous, with numerous variable black markings on: mesoscutal lobes, scuto-scutellar groove, scutellar trough, metanotum, propodeum dorsally, mesopleuron dorsally, metapleuron anteriorly and most of mesosoma ventrally; metasomal tergites black with apical half reddish to yellow but tergite 1 basally yellow, and tergite 3 and following sometimes without black marking but one or two postero-lateral dark spots; fore and mid legs reddish with coxae and trochanters somewhat paler, hind leg reddish with black markings of variable extent: coxa ventrally, trochanter ventrally, femur ventrally, tibia basally and apically, and entire tarsus; wings hyaline, venation pale brown.
MesosoMa. Moderately elongate; entire mesosoma deeply, densely and evenly punctate, but pronotum mid-dorsally and ventral half of speculum smooth; notaulus shallow; punctures somewhat confluent in area superomedia and area petiolaris, area superomedia elongate. Legs. Femoral tooth distinct but very weak, reduced to a barely distinct subapical point followed by minute denticles.

Male
Unknown.

Differential diagnosis
Moderately sized, mostly yellowish-orange with basal tergites mostly black; differentiated from all other Afrotropical Pristomerus species by the combination of the transverse head, the hardly sculptured mesonotum and the strong femoral tooth.

Differential diagnosis
Small, entirely yellow with flagellum bicoloured or tricoloured; otherwise differentiated from most Afrotropical species by the combination of the very short malar line, the strongly transverse clypeus, the coriaceous to smooth mesonotum and the strong female femoral tooth. Pristomerus yakoma sp. nov. seems closely related to P. swahili sp. nov., whose flagellum is uniformly dark and clypeus significantly narrower.

Male
Unknown.

Distribution
Central African Republic, Mozambique.

Differential diagnosis
Large and black with metasoma partially orange; differentiated from all other Afrotropical species by the combination of the colour pattern, the large size and the absence of a female femoral tooth; it might be closely related to P. kuku sp. nov., which is, however, characterized by a strong frontal protuberance and a distinctly shorter ovipositor.
Head. Face densely punctate-granulate; inner margins of eyes subparallel; clypeus transverse, sparsely punctate but punctation denser along dorsal margin, distinctly convex in profile; malar line long; frons, vertex and temple finely and densely punctate; frond with a moderate mid-longitudinal ridge from median ocellus to upper face; occipital carina joining hypostomal carina at mandible base; antenna with 35 flagellomeres, penultimate flagellomere quadrate.

Diagnosis
Moderately sized; yellowish orange overall; face moderately punctate; clypeus transverse and almost smooth; inner margins of eyes subparallel; malar line moderately short; upper head finely coriaceous; antenna with 30 flagellomeres, penultimate one slightly shorter than wide; mesosoma moderately to densely punctate but pronotum quite smooth and speculum hardly sculptured; female femoral tooth long and unusually narrow; tergite 2, apical half of tergite 1 and base of tergite 3 longitudinally aciculate; ovipositor long and straight, barely sinuous apically. Male unknown.

Differential diagnosis
Moderately sized and yellowish-orange overall; differentiated from most other Afrotropical species by the long and basally noticeably narrow female femoral tooth. Such a narrowed tooth is only otherwise found in P. afrikaner sp. nov. and P. yakoma sp. nov. The former, from South Africa, has, however, a longer malar line and longer antennae, and the latter has a coriaceous to smooth mesonotum. Pristomerus yoccolo sp. nov. is otherwise somewhat similar to P. babinga sp. nov., whose femoral tooth is distinctly stouter and in which the clypeus is more transverse and the inner orbits diverge ventrally.

Male
Unknown.

Distribution
Ivory Coast.

Diagnosis
Moderately small to small; yellowish-orange with mandible, clypeus and frontal orbits pale yellow, and variable dorsal black markings; trochanters and femora apically white-dotted; fore wing with anterior edge of pterostigma whitish; face densely punctate-granulate, clypeus almost entirely smooth, remainder of head coriaceous; clypeus strongly transverse; malar line short; antenna with 27 flagellomeres, penultimate one quadrate; mesosoma moderately to densely punctate, including sometimes pronotum and almost entire speculum, with punctures confluent into rugosities on notaulus; propodeal carination thick; female femoral tooth absent; ovipositor short, apically nearly straight. Male unknown.

Differential diagnosis
Small to moderately small and yellow to yellowish orange overall with some black markings dorsally; differentiated from all other Afrotropical species by the combination of the short malar line, the absence of a femoral tooth and the short ovipositor. Pristomerus zande sp. nov. is somewhat morphologically similar to P. albescens and P. vahaza, but is easily differentiated because both have a narrower clypeus.

Male
Unknown.

Distribution
Central African Republic.

Differential diagnosis
Rather to quite large and with an unusual colour pattern: metasoma black, head and mesosoma reddishtestaceous except orbits and lower head white, and legs largely white-maculated. That colour pattern readily distinguishes this species from almost all other Pristomerus species; P. veloma is somewhat similar in colour, but its orbits are not contrastingly pale and diverge ventrally.
Colour. Head mostly reddish-testaceous with face sometimes fading to black medially, and whitish parts: inner orbits, most of clypeus, mandible but teeth, malar space and palpi; mesosoma reddishtestaceous with scutellum weakly lighter orange and sometimes an infuscate marking on metanotum; metasoma black; legs mostly testaceous with anterior margins of fore and mid tibiae, and apex of all femora and trochanters whitish; ovipositor sheath dark brown; antenna dark brown, sometimes fading to testaceous toward apex; wing hyaline, venation pale brown with anterior half of pterostigma whitish.

Distribution
South Africa.

Comments
The three paratypes were previously reported as non-type specimens of P. veloma in Rousse et al. 2013.

Discussion
What else, other than pure taxonomic information, does this revision contribute to our knowledge of the genus? One of the first observations arising from this revision and the previous revision of Madagascan Pristomerus (Rousse et al. 2013) is the definition of colour patterns associated with geography or ecology. Five of the Pristomerus species from Madagascar are uniquely white-faced. No other species in Africa display such a pale face colour but two species from South Africa (P. zulu sp. nov. and another non-described male in the SAMC collections) exhibit whitish orbits on the otherwise testaceous face. We cannot associate that colour pattern with any ecological traits, mostly because Seyrig's labels do not provide information on habitat association. This correlates with a similar situation in Xanthophenax Saussure, 1892, where the white-faced species are restricted to Madagascar (Rousse, Villemant & van Noort, in prep).
A second colour pattern is more informative. This pattern is defined by white-dotted femora and halfwhite pterostigma, usually associated with a mostly black colour overall, although sometimes combined with a lighter overall colour, a pattern that is exhibited by 20 of the 49 species. These 20 species have all been collected in tropical Africa with the noticeable exception of P. zulu sp. nov. and, in part, P. bemba sp. nov.. Furthermore, 15 of these 20 species have been collected in rainforest, and two more in dryer forest environments. We lack ecological data for P. dikidiki sp. nov., but the collection locality strongly suggests a similar rainforest habitat. Moreover, all the 15 species from rainforest habitats have enlarged ocelli in both sexes. By contrast, 22 species of the 29 non white-dotted group have ocelli that are not enlarged in females, and for some of them not even in males. The 20 white-dotted species do not appear to be closely related, since they exhibit much variation in characters which are probably of more critical importance in terms of indicating phylogenetic relationships: the size of the femoral tooth, the length of the malar line, the sculpture of the mesonotum. On the other hand, some clearly related (and possibly sister-species) (e.g. P. sara sp. nov./P. kelikely) can be differentiated only by the presence or absence of this white dot. This colour pattern is thus clearly a phenotypic convergence conditioned by the environmental parameters of a closed and humid forest habitat.
Interestingly, the general black colour of most of these white-dotted species may be compared to one of the patterns displayed by Braconinae, where the darkest species are also reported from the tropical humid forests of Central Africa, putatively for concealment purpose (Quicke 1986). Dark patterns are also often associated with high altitude and low temperatures (Gauld & Mitchell 1978;Quicke 1997), though such a correlation could not be shown in Pristomerus. The comparison with the South American fauna brings further informative data. The predominant colour pattern in the Neotropical region is yellow to brown, black being an exception (Gauld 2000) even in the tropical rainforests (Sääksjaärvi pers. com.). Interestingly, most if not all of the rainforest species exhibit similar white dots on the femora (Sääksjaärvi pers. com.). What function can be hypothesized for the presence of these white marks on species in closed forest environments? To our knowledge, no other sympatric insect exhibits a similar pattern, so we can exclude the hypothesis of a mimicry complex (Quicke 1986). We suggest that the markings are involved in intra-specific communication, as contrasting markings are known to be used for this purpose in other Hymenoptera. For example, social Vespidae use individual markings for kin recognition (Baracchi et al. 2013) or minimization of conflict costs (Tibbets & Lindsay 2008). Even though they are taxonomically distantly related, from a functional perspective many lepidopterans exhibit "white" surfaces reflecting UV flashes, hence favoring mate location (Brunton & Majerus 1995). "Semaphore signaling" is also fairly well documented in Tephritidae (Diptera) (Sivinski et al. 2000), with the noticeable example of Eutreta diana (Osten Sacken), in which the mid femur is black and contrastingly yellow-tipped in both sexes, and is apparently involved in the mating behavior of the male (Goeden 1990). However, none of these examples provide definitive evidence to support any hypothesis for the precise function of white markings in Pristomerus, even though hypotheses of inter or intra-specific communication using body markings have been proposed and tested in Braconidae by Quicke (1984).
Finally, a third colour pattern groups the species which have a mostly yellow to orange overall colour, with various infuscate to black markings. This pattern is exhibited by most of the remaining Afrotropical species, with the exception of some very unusual species such as P. xhosa sp. nov., P. fynbos and P. protea sp. nov., or even P. sotho sp. nov. Some of the species in this colour group have the femur somewhat paler at the apex, but never as strikingly contrasting as in the white-dotted group. Species with this third colour pattern were collected in drier, open habitats. The few species known from the arid areas of the Mediterranean and Middle East regions are similarly coloured (Horstmann 1990). Here again, it makes sense to hypothesize that this colour pattern is well suited for camouflage purposes in lighter habitats with short vegetation, and/or to prevent over-heating in those often hotter environments. The perfect example supporting this dichotomy is P. pallidus, a common species with hundreds of specimens present in collections. Pristomerus pallidus is present from Madagascar to Western Africa and Southern Europe. Interestingly, very few records exist (with no ecological information) from Central Africa: this otherwise very widespread pale species appears to be largely absent from tropical rainforest.
The available data are indeed affected by a strong "agricultural bias", i.e., they largely include wasps reared from pests in cultivated areas, collected by agricultural services. Specimens of Cremastinae in collections are mostly associated with dry and open areas, at least in temperate areas (Fitton & Gauld 1980;Gauld 2000), but the few expeditions led into the Afrotropical forests brought back many new species. In the present study and the previous revision of Madagascan species (Rousse et al. 2013), about 60% of the newly described species emanate from limited and punctuated sampling in the forests of Madagascar and tropical Africa and are described from very few individuals if not from singletons. This suggests that, although our taxonomic knowledge of the genus has been expanded by the present revision, many more species will still be unveiled by further collecting.
Assuming the lack of knowledge we have on the precise biodiversity and phylogeny of the genus, we cannot hypothesize about the geographical location of its original radiation. The pantropical distribution of this genus is a tempting, but not definitive argument to support divergence of Pristomerus before the complete breakup of Gondwana, c. 95 million years ago (Le Gall et al. 2010). Such a distribution may, however, also be the result of a more recent dispersal event through temporary inter-continental connections (Sääksjärvi et al. 2004). As a consequence, we are still in need of taxonomic and phylogenetic data to reconstruct the evolutionary history of Pristomerus and related genera within the tropics.