Four-horned wasps, description of some remarkable Dirhinus (Hymenoptera, Chalcididae) from Kenya, with a discussion of their taxonomic placement Delvare, Gérard Copeland, Robert S. Zootaxa 2018 2018-01-19 4374 3 301 349 Delvare & Copeland 2018 [207,347,789,815] Insecta Chalcididae Dirhinus Animalia Hymenoptera 8 309 Arthropoda species quadrhinus   Diagnosis, females(character number in parentheses). Body with long setae on mandibles, labrum and procoxa (1) ( Fig. 25); clypeus with long setae on both sides, oriented forwards (2) (Figs 119, 148); mandibles about as long as procoxa (3) ( Fig. 28); apices of mesal mandibular tooth broadly rounded (4) ( Fig. 143); inner tooth of mandible absent (5); outer mandibular tooth hardly projecting, not visible in frontal view (6) ( Fig. 143), head with four horns (8) ( Fig. 31); inner horns quite broad (9); depression between inner horns very narrow, forming an incision (10); inner horns without apical notch (11); ocellar triangle not raised (12) ( Fig. 39); lateral margin of scrobal depression without tooth (13) ( Fig. 39; what could be mistaken here for an additional tooth is actually the profile of the outer horn, compare Fig. 42); flagellomeres short, transverse (14) ( Fig. 43); clava narrowly rounded apically, with very small area of micropilosity (15); pro- and most of mesonotum with lanceolate, thick and adpressed setae, except on apical half of scutellum (16) ( Fig. 49); mesonotum with extensive smooth surfaces, especially on mid lobe of mesoscutum anteriorly, disc of lateral lobes and oblong surface on centre of mesoscutellum (17) ( Fig. 49); anteromedian areola of propodeum short, at most as long as wide (19) (Figs 123, 135, 144); lateral areolae on propodeum with setation similar to the rest of surface (20); medioventral tooth of prepectus emarginate apically (21) ( Fig. 61); epicnemium virtually flat (22) ( Fig. 66); mesodiscrimenal groove wide, appearing as an oblong fovea, truncate ventrally, tapering dorsally and without dorsal fovea (23) ( Fig. 66); ventral section of epicnemial carina without median notch (24); subcoxal tooth on posterior margin of metepimeron projecting and narrowly rounded (25); posterior projection of metepisternum vestigial (26); metacoxa short and faintly strigose dorsally (27) ( Fig. 76); metafemur with sparse setation (28) ( Figs 76, 124); metafemur without engraved network (29) ( Fig. 76); metatibia with evident outer groove in addition to tarsal sulcus, the outer edge of which has a row of short and adpressed setae (30) ( Fig. 84); fore wing hyaline (31) ( Fig. 85); setation on upper side of wing moderately sparse (32); under side of wing with microtrichiae only (33); Rs fold setose (34) ( Fig. 87); very few admarginal setae on fore wing (35), setation on underside of marginal cell reduced (36); only two sensilla placodea at apex of submarginal vein (37) ( Fig. 86); four aligned sensilla placodea on stigma (38); dorsal surface of petiole sloping backwards (39) (Fig. 139); petiolar surface strongly emarginate on anterior margin (40) ( Fig. 118); median sulcus of petiole present, delimited by submedian carina (sulcus secondarily lost in some species) (41) ( Fig. 118); dorsal surface of petiole bare (42); petiole with vestigial basoventral tooth (43) (Fig. 139); strigose surface on first gastral tergite well expanded, about as long as wide (44) ( Figs 112, 140, 147); lateral ridges of first gastral tergite that delimit strigose surface somewhat curved outwards (45); hypopygium with long and diverging apical submedian setae (46).   FIGURES 21–30. Characters of Dirhininae(continued). 21, 23, 25, 27, 30. Mandible in frontal view. 22, 24, 26, 29. Mandible in lateral view. 28. Head in anterolateral view. 21, 22.  D. aff. hesperidum, ♀. 23, 24.  D. anthracia, ♀. 25.  Dirhinus( Dirhinus) claviger, paratype ♀. 26, 27.  D. aff. ehrhorni, ♂. 28.  D. quadrhinus, holotype ♀. 29.  Dirhinus( Pareniaca) aff. schwarzi, ♀. 30.  Dirhinus( Pareniaca)'Patawa', ♀.   FIGURES 31–42. Characters of Dirhininae(continued). 31–38and 39–42. Head respectively in dorsal view and lateral views. 31.  Dirhinus( Dirhinus) kambae, holotype ♀ (F02). 32.  Dirhinus( Dirhinus) himalayanus, ♀. 33.  D. claviger, paratype ♀. 34.  D. anthracia, ♀. 35.  D. aff. ehrhorni, ♀. 36.  D. inflexus, ♀. 37.  D. aff. schwarzi, ♀. 38.  Dirhinus( Hontalia)aff. caeruleus, ♀. 39.  Dirhinus( Dirhinus) leakeyorum, holotype ♀ (F08). 40.  D. aff. hesperidum, ♀. 41.  D. aff. ehrhorni, ♀. 42. D. aff. caeruleus, ♀.   FIGURES 43–52. Characters of Dirhininae(continued). 43–48. Antenna. 49–52. Mesonotum. 43.  D. gigasetosus, paratype ♀ (F07). 44.  D. claviger, paratype ♀. 45, 51.  D. aff. hesperidum, ♀. 46.  D. aff. ehrhorni, ♀. 47.  D. aff. schwarzi, ♀. 48. D. 'Patawa', ♀. 49.  D. maasaii, holotype ♀ (F05). 50.  D. anthracia, ♀. 52.  D. inflexus, ♀.   FIGURES 53–60. Characters of Dirhininae(continued). 53–54. Mesonotum. 55–60. Propodeum. 53. D. 'Patawa', ♀. 54. D. aff. caeruleus, ♀. 55.  D. clavigerparatype ♀. 56.  D. anthracia, ♀. 57.  D. aff. hesperidum, ♀. 58.  D. aff. schwarzi, ♀. 59. D. 'Patawa', ♀. 60. D. aff. caeruleus, ♀.   FIGURES 61–68. Characters of Dirhininae(continued). 61–65, 67and 66, 68. Mesopleuron respectively in lateral, lateroventral and ventral views. 61.  D. gigasetosus, holotype ♀. 62.  D. anthracia, ♀. 63.  D. aff. hesperidum, ♀. 64.  D. aff. ehrhorni, ♂. 65. D. 'Patawa', ♀. 66.  D. gigasetosus, paratype ♂ (M02). 67.  D. anthracia, ♀. 68.  D. aff. hesperidum, ♀.   FIGURES 69–75. Characters of Dirhininae(continued). 69and 70–72. Mesopleuron respectively in ventral and anteroventral views. 73–75. Metepisternum and petiole in lateral view. 69.  D. aff. ehrhorni, ♀. 70.  D. inflexus, ♀. 71. D. 'Patawa', ♀. 72. D. aff. caeruleus, ♀. 73.  D. aff. hesperidum, ♀. 74.  D. anthracia, ♀. 75.  D. inflexus, ♀. Abbreviation. dml-gr, discrimenal groove.   FIGURES 76–84. Characters of Dirhininae(continued). 76–83. Metacoxa and metafemur with sculpture of metafemur enlarged. 76.  D. maasaii, holotype ♀ (F05). 77.  D. anthracia, ♀. 78.  D. aff. hesperidum, ♀. 79.  D. aff. ehrhorni, ♂. 80.  D. aff. schwarzi, ♀. 81. D. 'Patawa', ♀. 82. D. aff. caeruleus, ♀. 83.  D. anthracia, ♀. Hind leg in posterior view. 84.  D. leakeyorum, holotype #f (F08). Metatibia in laterodorsal view. Abbreviations. tb3-ogr, outer groove; tb3-ogrm, margin of the outer groove; tb3-ts, tarsal sulcus; tb3-tsm, margin of the tarsal sulcus.   FIGURES 85–98. Characters of Dirhininae(continued). 85, 88, 90, 92, 95. Fore wing with setation enlarged. 87, 91, 93, 94, 96–98. Marginal cell. 86. Submarginal vein. 89. Stigmal vein. 85.  D. quadrhinus, holotype ♀ (F01). 86, 87.  D. kambae, holotype ♀ (F02). 88, 89.  D. himalayanus, ♀. 90, 91.  D. anthracia, ♀. 92, 93.  D. aff. ehrhorni, ♀. 94.  D. inflexus, ♀. 95, 96.  D. aff. schwarzi, ♀. 97. D. 'Patawa', ♀. 98. D. aff. caeruleus, ♀. Abbreviations. fw-Mt, macrotrichia; fw-mt, microtrichia.   FIGURES 99–111. Characters of Dirhininae(end). 99–101, 105, 106. Petiole in dorsal view. 102–104. Petiole and GT1 in dorsal view. 107. Metasoma in dorsal view. 108. Metasoma in lateral view with hypopygium enlarged. 109. Metasomal apex in lateral view. 110,111. Hypopygium. 99.  D. himalayanus, ♀. 100, 107.  D. claviger, paratype ♀. 101, 102, 110.  D. anthracia, ♀. 103, 111.  D. aff. hesperidum, ♀. 104.  D. aff. ehrhorni, ♀. 105. D. 'Patawa', ♀. 106. D. aff. caeruleus, ♀. 108.  D. aff. schwarzi, ♀. 109. D. 'Patawa', ♀. Abbreviations. GT1-cr1, primary crest; GT1-cr2, secondary crest.   TABLE 1. List of the selected species used for the character analysis of the genus  Dirhinusand origin of the specimens. The names of the undescribed species mostly refer to already described ones being morphologically similar, the primary types of which were examined by GD, except for  D. ehrhorni. Depositories.  D. claviger(BMNH),  D. aff. claviger(NKM), other specimens (CIRAD).    Selected species Code or status Country Province Locality Coordinates Sampling notes Date Collector     Dirhinus( Dirhinus) himalayanusWestwood 13021 Union of Arab Emirates Wadi Shawkah 25.06° N 56.01° E water trap 01-15/03/2011 van Harten A.    Dirhinus( Dirhinus) clavigerBouček & Narendran paratype India Utah Pradesh Aligarh 08-10/11/1979 Noyes J.S.    Dirhinus( Dirhinus) aff. claviger 0 8670 Kenya Rift Valley LakeNakuru National Park 0.47203° S 36.05388° E Malaise trap 17/02- 03/03/2006 Copeland R.   Dirhinus( Dirhinus) anthraciaWalkerGDEL0154 Cameroon Centre Messamendongo, (surrounds of Yaoundé) 03°57'07" N 011°29'27" E Malaise trap 31/03- 14/04/2003 Tussac M.  Dirhinus( Dirhinus) aff. hesperidumMorocco Fez - Meknès Moulay Idriss, Volubilis antic city 34°04'34" N 05°04'14" W 17/05/1995 Tussac H.      Dirhinus( Dirhinus) aff. hesperidum Morocco Gharb - Chrarda - Béni Messen 18 km N Souk-el-Arab-du- Rharb 34°50'34" N 05°57'36" W on flower of Apiaceae 14/05/2000 Delvare G.     Dirhinus( Pareniaca) aff. ehrhorni GDEL0463 Kenya Central Nairobi, ICIPE Campus 1.22165° S 36.89579° E ex  Hermetia illucens 15/12/2015 Copeland R.    Dirhinus( Pareniaca) inflexusWaterston Benin Plateau Pobé on extrafloral nectary 17/11/1993 Delvare G.    Dirhinus( Pareniaca) aff. schwarzi Dominican Rep. Pedernales Parque Nat. Jaragua, 4km W Oviedo 28/11- 04/12/1991 Masner L. & Peck J.    Dirhinus( Pareniaca)'Patawa' GDEL0254 French Guiana  Regina Roura, Patawa camp, Montagne de Kaw 4°33'N 52°10' W   Dirhinus( Hontalia)aff. caeruleusGDEL0261 French Guiana Regina Roura, Patawacamp, Montagne de Kaw 4°33'N 52°10' W   TABLE 2. Character states of  Dirhinusspecies (females) from the specimens listed in Table 1.    Characters   Dirhinus( Dirhinus) quadrhinus   Dirhinus( Dirhinus) himalayanus   Dirhinus( Dirhinus) claviger   Dirhinus( Dirhinus) aff. claviger   Dirhinus( Dirhinus) anthracia   Dirhinus( Dirhinus) aff. hesperidum  (1) Setation on mandible, labrum and procoxa long (as in Figs 23̅24) long (as in Fig. 23) long (Fig. 25) long (as in Figs 23̅24) long (Figs 23̅24) moderately long (Fig. 21)    (6) Mandible, outer tooth moderately projecting, hardly visible in front view (as in Figs 144, 152)  well expanded, visible in front view (as in Fig. 23)  well expanded, visible in front view (Fig. 25) well expanded, visible in front view (as in Fig. 25)  well expanded, visible in front view (Fig. 23)  well expanded, visible in front view (Figs 21̅ 22)    (2) Clypeus, long lateral setae oriented forwards at right angle with clypeal surface present and long (as in Figs 39, 152) present and long (as in Fig. 24) present and long (as in Fig. 24) present and long (as in Fig. 24) present and long (Fig. 24) present and moderately long (Fig. 40)  (3) Mandible, relative length as long as procoxa (as in Fig. 8) longer than procoxa (Fig. 23) longer than procoxa (Fig. 25) longer than procoxa (as in Fig. 25) longer than procoxa (Fig. 23) as long as procoxa (Fig. 22)  (4) Mandible, apex of mesal tooth  widely rounded (as in Figs 144, 152)  widely rounded (as in Fig. 23)  widely rounded (Fig. 25) widely rounded (as in Fig. 25)  widely rounded (Fig. 23)  widely rounded (Fig. 21)  (5) Mandible, inner tooth  absent  absent  absent absent  present (Fig. 23)  present (Fig. 21)   (7) Mandible, outer tooth apex rounded (as in Fig. 152) rounded (as in Fig. 23) rounded (Fig. 25) rounded (as in Fig. 25) rounded (Fig. 23) rounded (Fig. 22)  (12) Ocellar triangle not raised (as in Fig. 39) not raised (as in Fig. 39) not raised (as in Fig. 39) not raised (as in Fig. 39) not raised (as in Fig. 39) slightly raised (Fig. 40)  (13) Margin of scrobal depression, secondary tooth absent but outer horn present (as in Fig. 39) absent (as in Fig. 40) absent (as in Fig. 40) absent (as in Fig. 40) absent (as in Fig. 40) absent (Fig. 40)  (14) Flagellomeres, relative length short, transverse (as in Fig. 43) short, strongly transverse (as in Fig. 44) short, strongly transverse (Fig. 44) short, strongly transverse (as in Fig. 44) short, strongly transverse (as in Fig. 44) short, transverse except F1 and F2 (Fig. 45)    (8) Horns on head, number four (Fig. 113) two (Fig. 32) two (Fig. 33) two (as in Fig. 33) two (Fig. 34) two (as in Fig. 34)  (9) Horns on head, habitus wide (Fig. 113) very wide (Fig. 32) moderately wide (Fig. 33) moderately wide (as in Fig. 33) moderately wide (Fig. 34) moderately wide (as in Fig. 34)  (10) Depression between horns quite narrow (Fig. 113) quite narrow (Fig. 32) relatively narrow (Fig. 33) relatively narrow (as in Fig. 33) relatively narrow (Fig. 34) relatively narrow (as in Fig. 34)  (11) Horns on head, apex without notch or incision, narrowly rounded (Fig. 113) without notch or incision, widely rounded (Fig. 32) with sinuate margin near apex (Fig. 33) with sinuate margin with subapical notch near apex (as in Fig. 33) (Fig. 34) with subapical notch (as in Fig. 34)   (15) Clava, apex narrowly rounded (as in Fig. 43) narrowly rounded truncate (Fig. 44) truncate (as in Fig. 44) narrowly rounded (as in Fig. 43) moderately rounded (Fig. 45)  …….continued on the next page ……continued on the next page  Dirhinus( Dirhinus)  Dirhinus( Dirhinus)  Dirhinus( Dirhinus)  Dirhinus( Dirhinus)aff.  Dirhinus( Dirhinus)  Dirhinus( Dirhinus)aff.  Characters  quadrhinus himalayanus claviger claviger anthracia hesperidum16) Pronotum and setae thick, lanceolate and setae thick, lanceolate and setae thick, lanceolate and setae thick, lanceolate and setae thick, lanceolate setae slender, adpressed mesonotum, setation adpressed (as in Fig. 49) adpressed (as in Fig. 49) adpressed (as in Fig. 49) adpressed (as in Fig. 49) adpressed ( Fig. 50) ( Fig. 51) with smooth surfaces on with smooth surfaces on with smooth surfaces on with smooth surfaces on With small smooth surfaces with reduced smooth 17) Mesonotum, lateral lobes of mesoscutum lateral lobes of mesoscutum lateral lobes of mesoscutum lateral lobes of mesoscutum on lateral lobes of surface on center of pattern of puncturation and mesoscutellum (as in and mesoscutellum (as in and mesoscutellum (as in and mesoscutellum (as in mesoscutum and mesoscutellum ( Fig. 51) Fig. 49) Fig. 49) Fig. 49) Fig. 49) mesoscutellum ( Fig. 50) 18) Propodeum, present (as in Fig. 123) present present ( Fig. 55) present (as in Fig. 55) present ( Fig. 56) present ( Fig. 57) spiracular tooth 19) Propodeum, as long as wide (as in Fig. wider than long (as in Fig. wider than long (as in Fig. as long as wide (Fig. as long as wide ( Fig. 55) as long as wide ( Fig. 56) anteromedian areola 135) 134) 135) 57) 20) Propodeum, setae thin as on rest of setae thin as on rest of setae thin as on rest of setae lanceolate and thick in setae thin as on rest of setae thin as on rest of setation of lateral propodeal surface (as in propodeal surface (as in propodeal surface (as in contrast with rest of propodeal surface (Fig. propodeal surface ( Fig. 55) areola Fig. 135) Fig. 55) Fig. 55) propodeal surface ( Fig. 56) 57)    (21) Prepectus, apical margin of medioventral tooth distinctly emarginate (as in Fig. 61) distinctly emarginate (as in Fig. 62) distinctly emarginate (as in Fig. 62) distinctly emarginate (as in Fig. 62) distinctly emarginate (Fig. 62) distinctly emarginate (Fig. 63)  (22) Mesepisternum, mesal elevation of epicnemium absent, epicnemium virtually flat (as in Fig. 66) epicnemium only very slightly raised mesally (as in Fig. 67) epicnemium only very slightly raised mesally (as in Fig. 67) epicnemium only very slightly raised mesally (as in Fig. 67) epicnemium only very slightly raised mesally (Fig. 67) epicnemium only very slightly raised mesally (Fig. 68)  (23) Mesodiscrimenal groove wide and superficial (as in Fig. 66) wide and deep (as in Fig. 67) wide and deep (as in Fig. 67) wide and deep (as in Fig. 67) wide and deep (Fig. 67) moderately wide, deep (Fig. 68)  (27) Metacoxa, habitus short (as in Fig. 76) short (as in Fig. 76) short (as in Fig. 76) short (as in Fig. 76) moderately long (Fig. 77) moderately long (Fig. 78)  (28) Metafemur, setation sparse (Fig. 112) very sparse sparse (as in Fig. 76) very sparse dense (Fig. 77) dense (Fig. 78)    (24) Epicnemial carina, ventral section continuous (as in Fig. 66) with mesal notch (as in Fig. 67) with mesal notch (as in Fig. 67) with mesal notch (as in Fig. 67) with mesal notch (Fig. 67) continuous (Fig. 68)  (25) Posterior margin  of metepimeron, present present present vestigial present present  subcoxal tooth  (26) metepisternum, posterior projection vestigial present (as in Fig. 74) [not visible on specimen] present present (Fig. 74) present (Fig. 73) piliferous points only, piliferous points only, piliferous points only, piliferous points only, piliferous points only, piliferous points only, 29) Metafemur, without engraved network without engraved network without engraved network without engraved network without engraved network without engraved sculpture (as in Fig. 76) (as in Fig. 76) (as in Fig. 76) (as in Fig. 76) ( Fig. 77) network ( Fig. 78) present but setose, visible 30) Metatibia, outer present and evident (as in present and evident (as in present and evident (as in present and evident (as in through outer ridge only absent groove Fig. 84) Fig. 126, 127) Fig. 126, 127) Fig. 126, 127) ( Fig. 83) ……continued on the next page  Dirhinus( Pareniaca)aff.  Dirhinus( Pareniaca)aff.  Dirhinus( Hontalia)aff.   TABLE 2 (continued). Character states of  Dirhinusspecies. Characters  Dirhinus( Pareniaca) inflexus  Dirhinus( Pareniaca)'Patawa'  ehrhorni schwarzicaeruleusnarrow, on top of mesal quite narrow, on top of mesal quite narrow, on top of mesal quite narrow, on top of mesal quite narrow, on top of mesal    (16) Pronotum and mesonotum, setation setae slender, adpressed (as in Fig. 52) setae slender, adpressed (Fig. 52) setae lanceolate, adpressed setae lanceolate, adpressed (Fig. 53) setae very short and thin (Fig. 54)  (17) Mesonotum, pattern of puncturation without smooth surfaces, punctured overall (as in Fig. 52) without smooth surfaces, punctured overall (Fig. 52) without smooth surfaces, punctured overall (as in Fig. 52) without smooth surfaces, punctured overall (Fig. 53) with reduced smooth surfaces, mostly (Fig. 54)   (18) Propodeum, spiracular tooth present present present (Fig. 58) present present (Fig. 60)  (19) Propodeum, anteromedian areola as long as wide as long as wide longer than wide (Fig. 58) much longer than wide (Fig. 59) much longer than wide (Fig. 60) (20) Propodeum, setation of lateral areola setae thin as on rest of propodeal surface (as in Fig. 57) setae thin as on rest of propodeal surface (as in Fig. 57) setae lanceolate, thick (Fig. 58) setae lanceolate, thick (Fig. 59) setae thin, of medium length (Fig. 60) (21) Prepectus, apical margin of medioventral tooth convex (Fig. 64) strongly convex (as in Fig. 65) strongly convex (as in Fig. 65) strongly convex (Fig. 65) strongly convex (as in Fig. 65) (22) Mesepisternum, mesal elevation of epicnemium epicnemium moderately raised mesally (Fig. 69) epicnemium strongly raised (Fig. 70) epicnemium strongly raised (as in Fig. 70) epicnemium strongly raised (Fig. 71) epicnemium strongly raised (Fig. 72) ) Mesodiscrimenal groove elevation of epicnemium (Fig. elevation of epicnemium (Fig. elevation of epicnemium (as in elevation of epicnemium (Fig. elevation of epicnemium ( Fig. 69) 70) Fig. 70) 71) 72)    (24) Epicnemial carina, ventral section continuous (Fig. 69) continuous (Fig. 70) continuous (as in Fig. 70) continuous (Fig. 71) continuous (Fig. 72)  (25) Posterior margin of metepimeron, subcoxal tooth present present present, hardly projecting, vestigial absent  (26) metepisternum, posterior projection present present (Fig. 75) absent absent absent ) Metacoxa, habitus moderately long ( Fig. 79) moderately long (as in Fig. 79) moderately long ( Fig. 80) long ( Fig. 81) very long ( Fig. 82) ) Metafemur, setation dense ( Fig. 79) dense (as in Fig. 79) dense ( Fig. 80) dense ( Fig. 81) dense ( Fig. 82) with engraved network in with engraved network in with engraved network in with engraved network in without engraved network, only ) Metafemur, sculpture addition to piliferous points addition to piliferous points (as addition to piliferous points addition to piliferous points piliferous points ( Fig. 82) ( Fig. 79) in Fig. 79) ( Fig. 80) ( Fig. 81) ) Metatibia, outer groove absent absent absent absent absent  ……continued on the next page ……continued on the next page) Hypopygium, pair of apical absent absent absent ( Fig. 108) absent ( Fig. 109) absent