J Au\/ l’tr/ Soc . 1985. 24 261-274 26 I THE AUSTRALIAN SPECIES OF MONOMACHIDAE (HY MENOPTERA: PROCTOTRUPOIDEA), WITH A REVISED DIAGNOSIS OF THE FAMILY I. D. NAUMANN Dii,isioti uj’En/oniologv. CSIRO. G.P.O. Box 1700, Cunherra. A.(‘.T.2601. Abstract The family Monomachidae is considered to comprise Mot1ornuc.hu.s Klug (Irom Australia and South America) and Terruconu.sSzepligeti (South America). A family diagnosis and key to genera are presented, and the generic placement of wing-reduced South American females is discussed. Three Australian species of Monomuchus are recognised: M.untipodu1i.s Westwood ( = M.unripotlulis heirdoru Riek = M. usmondi Riek synn. n.). M. auslrulicusGirault and M.hesprriussp. n. Thedipterous host of M.trntipodali.~ hasGondwanan affinities. Introduction The family Monomachidae is a small group with an austral-disjunct distribution: 3 species occur in southern and eastern Australia and at least 10 in South America. No fossil monomachids are known, but the family exhibits many character states plesiomorphic within the Proctotrupoidea and is probably a very ancient taxon. Adult monomachids favour cool, moist, forest habitats. It is reasonable to assume that the family is Gondwanan in origin. The Australian species Monomachus anripodalis Westwood has been reared from a species of Boreoiiles Hardy (Diptera: Stratiomyidae) (Riek 1970), a fly remarkable in having apterous females. The subfamily Chiromy- zinae, to which Boreoides belongs, also has a largely austral-disjunct distribution, and almost certainly has a Gondwanan origin. The Monomachidae were catalogued by Dalla Torre (1902) and reviewed by Schletterer (1889) and Schulz (191 1). The more recent literature is scanty: descriptions 4 FIGSI-4-Monomachus. Tetraconus spp.: (1-2) right mandible, anterodorsal view: (I) M.unripor/u/i.s female; (2) M. uusfrulicus male. (3) T. mocsur?? holotype female. head. dorsal view. (4) M. anripol1ali.s female. (ore wing venation (in part). See text lor explanation of symbols. Scale line = 0.5 mm. 1. D. NAUMANN FIGS5-8-M. unripodulis female, head: (5) dorsal view; (6)frontal view; (7)dorsal view; (8) clypeal margin. See text for explanation of symbols. Scale line = 0.2 mm. of new species of Monomachus Klug from Australia (Girault 1925) and Chile (Brethes 1928); a revision of the Australian species of Monornachus (Riek 1955); and synoptic notes on fami!y placement and biology (Riek 1970; Townes 1977; Johnson 1982). Riek's (1955) specific and subspecific diagnoses of Australian Monomachus were based on an examination of just 17 specimens and have proved impractical. The present study, based on over 240 specimens, does away with subspecific categories within the Australian Monomuchus and provides a new key to Australian species. Descriptions provided below of Australian species adhere to a new framework which will be applicable to a revision of the more numerous South American species. Morphologicul ierrns, measurernen/s-BL, length of body (excluding antenna); CL, length of pronotal collar (Fig. 10); FW, minimum distance between compound eyes (Fig. 3); FWL, length of fore wing; FIL. FIW, length, width of first flagellar segment; F13L. F13W. length, width of apical flagellar segment; HL. H W, length,width of head (Fig. 3); MAE, maximum diameter ofcompound eye (Fig. 6);mf, subbasal fold of mandible (Figs I, 6); MI, median index = A/B (Fig. 4); ML, minimum distance between compound eye and mouth (Fig. 6); NL, length of pronotal neck (Fig. 10); oc, occipital carina (Figs 9, 16); OD, maximum diameter of lateral ocellus (Fig. 3); OOL. distance between lateral ocellus and compound eye (Fig. 3); PH, PL, height, length of petiole; POL, distance between lateral ocelli (Fig. 3); PW, width of petiole; RI, radial index = C/D (Fig. 4); SCW, width of scutellum (Fig. 11); SI. stigma1 index = E/F (Fig. 4); T2L. length of second metasomal tergite. CoNection.7-AM, Australian Museum, Sydney; ANIC, Australian National Insect Collection, CSIRO, Canberra; BCRI, Biological and Chemical Research Institute, Rydalmere; BMNH, British Museum (Natural History), London; HNHM, Hungarian Natural History Museum, Budapest; MCZ, Museum of Comparative Zoology, Harvard; MV, Museum of Victoria, Melbourne; OUM, Oxford University Museum, Oxford; QDPI, Entomology Branch, Department of Primary Industries, Brisbane; QM. Queensland Museum, Brisbane; WAM. Western Australian Museum, Perth; ZMHU, Zoological Museum, Humboldt University, Berlin. AUSTRALIAN MONOMACHIDAE 263 FIGS9-ISM. antipodah: (9) male. head, posterior view; (10) female, pronotum, dorsal view; (1 1) female. mesosoma. dorsal view; (12-13) female, propodeum. dorsal view. See text for explanation of symbols. Scale line = 0.2 mm. Family MONOMACHIDAE Monomachinae Ashmead. 1902: 241, 243; Szepligeti, 1903: 387-388. Monomachidae Schulz. 191 1: 405-410; Johnson. 1982: 661. Female Head-Inner orbits not strongly convergent ventrally. Antenna1 socket very narrowly separated from clypeus. Labrum not forming median process. Mandible endodont. bidentate or tridentate. Labial palp 5- segmented. Maxillary palp 3-segmented. Anrenna-15-segmented; scape 2.4-3.3 times longer than wide: anellus absent. Mesosoma-Pronotum without reentrant declivity, posterior margin broadly overlapping meso- scutum. Mesoscutum anteriorly with continuous, transverse. postmarginal carina. Acetabular carina absent. Mesoscutellum posteriorly with transverse row of punctures. Fore wing-Rs not forked. Radial cell posteriorly closed by tracheate vein or colour streak. Rs + M absent. Rs intercept of basalis tracheate. sclerotised. Medial cell absent. Isolated medial sclerite absent. Hind wing-Posterobasally convex. Basal vein (Irm/Rs) and M + Cu, tracheate (Fig. 35). Legs-Trochantellus present on all legs. Hind tibia not grossly clavate: blunt tubercle present (Figs 21-22). Tibia1 spur formula I, 2, 2. Mefasoma-Petiole (Fig. 33) formed by TI and SI only; TI and SI separated by suture. Caster slender, tapering. not laterally compressed, with pair of spiracles. Gastral tergites and sternites broadly overlapping. Apical sternite not medially slit. Cercus plate-like. Ovipositor internal. Apical tergites partially overlapped by apical sternite. Male Antenna- 14-segmented, without modified flagellar segments. Metusoma-Gaster posteriorly blunt. Cercus digitiform. Apical tergites not overlapped by apical sternite. Genitalia as in Figs 32, 34. 264 I. D. NAUMANN FIGS14-17---M. uusirulicusmale. head: (14) posterior view; (15) frontal view;(l6) posterior view; (17)clypeal margin. See text for explanation of symbol. Scale line = 0.2 mm. Discussion Phylogenetic analysis of the proctotrupoid families (1. D. Naumann and L. Masner unpubl.) indicates that the genera Monomachus and Tetraconus Szepligeti are sister groups (synapomorphies: female metasoma slender, hind tibia1 tubercle present). On the other hand the sister group of Monomachus + Tetraconus is uncertain and Riek’s (1955, 1970) conglomerate family Heloridae (comprising Helorus Latreille, Austronia Riek, Monomachus, Tetraconus, Vanhornia Crawford and Roproniu Provancher) is not supported by a single apomorphy. Helorus. Austronia, Mon- omuchus + Tetraconus, Vanhornia and Ropronia are each morphologically and biologically very distinct and phenetically as remote from each other as are the widely recognised, polytypic proctotrupoid families (Diapriidae, Proctotrupidae, Platygas- tridae, Scelionidae and Pelecinidae) (Naumann and Masner unpubl.). In view of the phenetic isolation of Monomachus + Tetraconus and in the absence of a clear sister group relationship, it is more informative and less misleading to regard these 2 genera as comprising a discrete family, Monomachidae. This is a reversion to Schulz’s (191 1) concept of the family Monomachidae, a concept promoted more recently by Townes (1 977) and Johnson (1982). Similarities, for example in the overlapping articulation of the pronotum and mesoscutum, suggest that the Monomachidae (comprising Monomachus and Tetraconus), Roproniidae (comprising only Ropronia) and Austro- niidae (comprising only Austronia) are closely related and may form a monophyletic group. The Australian Monomachidae comprise a small, morphologically uniform group. On the other hand the South American Monomachidae are more hetero- geneous. Manotypic Tetraconus is in most respects a typical Monomachus and several authors (Schulz 191 1; Townes 1977) have questioned whether the presence of genal AUSTRALIAN MONOMACHIDAE 265 FIGS18-22-M. ausfralicusmale: (18) mesosoma. dorsal view; (19) pronotum. dorsal view; (20) propodeum, dorsal view; (21) hind tibia; (22) hind tibia, detail. Scale line = 0.2 mm. tubercles (Fig. 3) and clypeal tubercles justifies generic separation. I have examined the holotype of T. mocsaryi Szepligeti (in HNHM), but hesitate to synonymise Tetraconus with Monomachus until a review of the South American monomachid fauna can reevaluate the diagnostic characters summarised in the key below: 1. Clypeus and gena with strong tubercles (Fig. 3); dorsal mandibular tooth divided; South America . .. Tetraconus Szepligeti 266 I. D. NAUMANN FIGS23-28-M. hesperius paratype female: (23-26) head: (23) dorsal view; (24) frontal view; (25) posterodorsalview; (26) clypeal margin. (27) pronotum, dorsal view. (28)propodeum, dorsal view. Scale line = 0.2 mm. Clypeus and gena without strong tubercles (Fig. 5); dorsal mandibular tooth undivided (Fig. 6); Australia and South America .. .. .. .. .. .. .. Monomachus Klug Wing reduction has not previously been reported in the Monomachidae. A