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: ) (Riek 1970), a 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 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 micropterous female (in MCZ) from San Fabian Province, Chile, has a minute, yellowish fore wing (smaller than the corresponding tegula and devoid of setae or venation), a rather large metanotum, a vestigial hind wing and a slightly globose propodeum. For the present, I prefer to regard this specimen as an aberrant, flightless species of Monomachus. On the other hand Masner (pers. comm.) advises me of 2 AUSTRALIAN MONOMACHIDAE 261

FIGS29-34-Monomachus spp.: (29-31) male S8: (29) M. australicus; (30) M. hesperius paratype; (31) M. antipodalis. (32-34) M. antipodalis: (32) male genitalia, lateral view; (33) female, petiole, lateral view; (34) male genitalia, dorsal view. Scale line = 0.2 mm. additional, undescribed, flightless species (from Peru and Chile) in which the female is apterous, the petiole very short and the radial cell of the male fore wing open. These 2 species may deserve generic separation from Monomachus. Biology Riek’s (1970) rearing of M. antipodalis from mature larvae and puparia of a species of Boreoides (Diptera: Stratiomyiidae: ) is the only known host association for the Monomachidae. The relatively small, male Boreoides hosts each produce 1 M. antipodalis. Several wasps can complete their development successfully in each of the larger, female hosts. Boreoides has a more restricted distribution than does Monomachus (Fig. 39). M. australicus Girault and M. hesperius sp. n. must parasitise a different host or hosts: among the Chiromyzinae larger species of Chiromyza Wiedemann and Inopus Walker are possible candidates. Monomachus spp. are not among the known parasites of (Macquart) despite intensive 268 I. D. NAUMANN

37

FIGS35-38-Monomachus spp., hind wings: (35-36) M. anripodalis; (37-38) M. uu.rrru1icu.s. Scale line = 2 mrn. biological studies of this economically important species (L. N. Robertson pers. comm.), suggesting that Chiromyza spp. are the more likely hosts of M. australirus and M. hesperius. Adults of Australian Monomachus spp. tend to fly during the cooler months. M. antipoddis is almost exclusively an autumn-winter species, and in southern Australia is 1 of the few wasps active when the daily temperature range is 0-9°C. M. australicus, from north-eastern Australia, predominantly during late summer and winter, but records from the Bellenden Ker Range suggest that even in tropical regions at high altitudes (over 1000 m) this species has a longer summer flight period. M. hesperius is active from late summer to early winter. Monomachus spp. occur in rainforest, in wet sclerophyll forest and in moist pockets in otherwise dry sclerophyll forest. M. anripodalis has been recorded from domestic gardens where these are relatively shaded and moist. Adult wasps are collected by Malaise traps and are attracted to light; they are rarely taken by diurnal sweeping. A series of 6 females of M. antipodalis was recovered from the stomach of a trout caught on the Fish R., N.S.W. Biogeography Australian Monomachus spp. are allopatric and restricted to the higher rainfall regions of eastern and southern Australia (Fig. 39). The absence of Tasmanian records may reflect a lack of collecting during cooler months. Their distribution falls almost entirely within the known distribution of the Chiromyzinae, which includes the only known host of a species of Monomachus. The Chiromyzinae are a primitive element within the Australian Stratiomyidae. They are represented elsewhere by several genera AUSTRALIAN MONOMACHIDAE 269

M. antipodalis A M. australicus M. hesperius #: Boreoides spp.

FIG.39-Distribution of M0nomuchrr.s spp., Boreoidrs spp. and Chiromyzinae in Australia. in New Zealand and South America and by the more widespread Inopus (Colless pers. comm.). Their largely Gondwanan distribution resembles that of Monomachus, except that the latter is absent from New Zealand. Monomachus Klug MonomachusKlug, 1841: 378: Westwood, 1841: 535; 1843: 252; Brulle. 1846: 533; Schletrerer, 1889: 209-210; Ashrnead, 1902: 243, 258-259; Schulz, 191 I: 407-411; Riek. 1955: 258-260; 1970: 905-906; Muesebeck and Walkley. 1956: 372: Townes. 1977: 1: Johnson. 1982: 661, Type species: P~./rcirru.\,fu.c.cu/orPerly (by rnonotypy). Diugnosis-See key to genera above.

Key to Australian species of Monomachus 1. Occipital carina (oc, Fig. 9) widely separated from oral margin; external surface of mandible with distinct, transverse, subbasal fold (mf, Figs I, 6) ...... antipodalis Westwood Occipital carina reaching oral margin (Fig. 16) or separated from margin by 2 or 3 setigerous punctures; external surface of mandible smoothly curved, without distinct, transverse, sub- basal fold (Figs 2, 15) . . .. 2 2. In hind wing all (Fig. 38) or basal 0:5 (Fig: 37) of M be;ween.Cu, and lrm/Rs absent; never with basal stump of M beyond Cu ,; ventral mandibular tooth distinctly longer than dorsal tooth (Fig. 2); clypeal margin of female medially broadly triangular (Fig. 15) 270 1. D. NAUMANN or rounded (Fig. 2); apodeme of S8 of male anteriorly strongly expanded (Fig. 29) . . . . australicus Girault In hind wing M continuous between CU, and'lrm/Rs (Fig. 35) or narrowly interrupted near Cu, (Fig. 36); always with at least basal stump of M beyond Cu,; mandibular teeth subequal (Figs 24, 26); clypeal margin of female medially distinctly bidentate (Figs 24, 26) or indistinctly tridentate; apodeme of S8 of male anteriorly weakly expanded (Fig. 30) . . . . hesperius sp. n.

Table 1. Ranges of body and wing lengths, ratios of measurements: Australian Monomachus spp.

M. anripodalis M. ausrralicus M. hesperius Female Male Female Male Female Male

BL (mm) 11.0-21.0 7.1-10.3 11.0-14.6 6.4-7.8 10.8-13.7 6.1-6.7 FWL (mm) 4.5-8.1 5.2-7.0 4.7-5.9 4.2-5.0 4.8-5.7 4.2-4.9 HWjHL I .32-1.54 1.29-1.44 I .24- I .35 I .26-1.39 I .32-1.38 I .33-1.38 POLjOOL 0.82-1.14 1.15-1.42 I .OO- 1.42 1.10- I .so 1.06-1.17 1.40-1.44 ODjPOL 0.40-1 .OO 0.57-0.75 0.53-0.82 0.57-0.91 0.55-0.79 0.57-0.65 FW/MAE 1.07-1.35 1.03-1. I7 0.97-1.08 0.98-1.09 1.04- 1.30 1.07-1.14 ML/MAE 0.38-0.58 0.28-0.37 0.26-0.36 0.17-0.27 0.32-0.44 0.24-0.26 FlL/FlW 5.29-6.60 5.67-7.20 5.50-6.50 6.00-7.67 5.80-6.50 6.38-7.22 FI 3LjFI 3W 2.91-3.90 6.89-8.78 2.44-3.25 6.67-8.40 2.26-2.89 7.50-9.20 NLjCL 0.73-1 .OO 0.83-0.94 0.80-0.88 0.67-0.77 0.84 0.85 HWjSCW I .28- I .47 I .02- I. 34 1.16-1.21 1.07-1.19 I. 17-1.30 I. 14- I. 19 MI 0.35-0.43 0.38-0.45 0.39-0.46 0.43-0.57 0.32-0.43 0.23-0.39 R1 3.26-4.50 3.27-4.09 3.58-4.53 2.78-3.86 4.13-5.40 5.40-6.00 SI 3.00-4.50 3.67-4.67 4.17-4.50 3.83-5.75 3.75-4.50 3.29-5. I I PLjPW 3.00-4.64 7.50-11.30 4.91-5.84 7.88- 10.00 5.10-6.36 8.3I- 12.36 PLjPH 4.00-5.13 7.50-1 1.30 5.25-7.91 5.80-9.43 5.42-7.00 7.94-1 1.33 PL/T2L 0.78-0.99 1.50-2.06 0.93-1.03 I .52-2.06 0.96-1.07 I .94

Monomachus antipodalis Westwood (Figs 1, 4- 13, 3 1-36, 39; Table 1) Monomuchusunlipodalis Westwood, 1874: 126, plate 24; Schletterer, 1889: 225-226; Dalla Torre, 1902: 1088; Schulz, 1907: 309-310; 191 I: 414 (in part). Monomachus anripodalis anripodalis Westwood. Riek, 1955: 260-261. Monomachus anripodalis bendora Riek, 1955: 260-262. Syn. n. Monomachus osmondi Riek, 1955: 260,263. Syn. n. Tvpes-M. anripodalis: VICTORIA:holorype 9 (in good condition; lacking apical 1 I segments from left antenna), Melbourne, R. Bakewell, in OUM. M. antipoclalis bendoru: AUSTRALIANCAPITAL TERRITORY: holor.vpe (in good condition), Bendora, 21.iii.1951, E. F. Riek, in ANIC (type number 7593). M. osmondi: SOUTHAUSTRALIA: holorype 0 (in good condition), Glen Osmond. 13.v.1937, D. Swan, in ANIC (type number 7594). Orher mureriulc,samine~I-NEw SOUTHWALES: I 5, Bathurst, 27.iv.1970. J. Turner. in BCRI; I i,MI Wilson, 15.iv.1971, D. K. McAlpine, in AM; 1 0, Katoomba, 17.iv.1952, in AM; 6 09, Fish R., Tarana, 1940. in ANIC, AM; I 9,Vulcan State Forest, 24 km W ofOberon, l.iv. 1984, N. Drydale, in AM; I $2, Clyde Mtn, 5.v.1965, D. H. Colless, in ANIC; 15 99,107 5$,Kiandra, 6 and IO.iii.1960, E. F. Riek, ex Boreoides sp., in ANIC. BMNH, QM. AUSTRALIANCAPITALTERRITORY: 49 YY, 2 Sd, Canberra, variousdates between 19.iv and 2.viii. 1956-1983, T. G. Campbell, J. C. Cardale, S. Kohlhagen, D. Morgan, 1. D. Naumann, R. Powning, E. F. Riek, in ANIC, BMNH, QDPI, QM; I 3, Bendora, 2.iv.1958, E. F. Riek, in ANIC. VICTORIA: I Y. junction of Soldier Ck Spur track and Takes Ck track, 3.iv.1974, in MV; I 0, Melbourne, iii.1929, Blackbourne, in QDPI; 1 Y, same locality, 15.v.1982, in MV; 1 y, Port Phillip, Coulon, in ZMHU; 1 $2, Caulfield, 1910, C. French, in ANIC; 2 99,Berwick, Woostercoll., in ANIC; 1 (2, Nilma, 9.xi. 1955, in ANIC; I Y, Ocean Grove, 5.v. 1982, T. Pescott. in MV. UNLOCALISED:2 YV, no data, in ANIC, UQIC.

Female Body and wing lengths, indices as in Table 1. Head-Vertex: with dense setigerous punctures (Figs 5,7); ocellar area irregularly foveolate or rugose; postocellar area at least in part smooth. Occipitalcarina (Fig. 9) widely separated from oral margin. Gena: in dorsal view strongly swollen, without tubercle; with dense setigerous punctures; with or without smooth area AUSTRALIAN MONOMACHIDAE 27 I adjacent to compound eye. Clypeal margin medially abruptly produced. usually tridentate (Figs 6, 8). Mandible: apically broad, bidentate; teeth subequal; external surface with transverse, subbasal fold (Figs I, 6) and usually with weak excavation immediately distal to fold. Mrsosoma-Pronotum: middorsally with sparse setigerous punctures; dorsolaterally setigerous punctures dense. Notaulix punctate. Tegula larger than pedicel. Metanotum medially shorter than mesoscutellum. Propodeum: punctate to rugose (Figs 12-13), interspaces smooth or microreticulate; usually with poorly defined dorsolateral and posterodorsal unsculptured areas; median carina variable, sometimes absent . Macropterous. Fore wig-Radial cell closed. Junction of m-cu and Cu, proximal to bifurcation of Cu,. Hind wing-M continuous between Cu, and lrm/Rs (Fig. 35) or narrowly interrupted near Cu, (Fig. 36); always with at least basal stump of M beyond Cu,. Merasoma-Petiole: anterodorsally irregularly rugose or smooth; in lateral view more or less sinuate. Colour-Fore wing hyaline. Variation continuous between following forms: (a) melunisticform: body predominantly black; antenna, mandible (in part), small median patch on mesoscutum posteriorly, femora (apically). tibiae, tarsi reddish mange to reddish brown; and (b)rt@s/irform: body predominantly reddish brown to reddish orange; head (excluding most of mandible, clypeus), lateral pronotum. propleuron, mesopleuron, metanotum, propodeum brown to black; mandibular teeth black.

Mule Differing from female as follows: Lengths, indices as in Table 1. Head-Ocellar area sometimes smooth with only a few wrinkles close to ocelli. Postocellar area usually broadly smooth. Gena not as strongly swollen. Clypeal margin produced, usually rounded, rarely distinctly tridentate. Mrsosoma-Notaular punctures sometimes indistinct. In occasional specimens propodeum dorsally almost entirely smooth. Fort wing-Distal remnant of Rs + M proximal to m-cu sometimes present. Merasoma-Petiole straight. Apodeme of S8 anteriorly weakly expanded (Fig. 31 ). Colour--Body predominantly black; antenna brown: mandible (excluding teeth). mesoscutum and mesoscutellum (some specimens only). trochanters. femora. tibiae. tarsi. T2-T4 contrasting reddish orange.

Discussion Riek (1955) distinguished M. osmondi, M. untipodalis and M.antipodulis bendora on differences in the microsculpture of the head, pronotum and petiole, and some slight differences in colour. Long series now available for examination reveal in these characters continuous variation which encompasses the forms studied by Riek. The median carina and punctation of the propodeum is also variable (cf. Figs 12-13). The margin of the clypeus is always distinctly produced medially and in some females is almost tridentate.

Monomachus australicus Girault (Figs 2, 14-22, 29, 37-39; Table 1) Monomachus ausrralicus Girault, 1925: [2]; Riek. 1955: 260. 262; De Santis. 1961: 166. Type-QUEENSLAND: holory e (in fair condition; head + prothorax and rest of body mounted separately on triangular card; lading left antenna, right antenna beyond scape, right fore leg beyond coxa), Cleveland. 14.v.1925. in QM. Orher marerial elUmined-QUEENSLAND: 1 j,Crystal Cascades, near Cairns. IO.vi.1971, E. F. Riek. in ANIC; 1 ',, I 2. Bellenden Ker Range. Cable Tower No. 3. 1054 m. 17.x-5.xi.1981. in QDPI; 1 $, 0.5 km WSW of Mt Bellenden Ker Peak. 1560m. 3.xi.1981. E. D. Edwards. in ANIC: 12 ,j;. 10 km W of Paluma. 12.v.1980. I. D. Naumann. J. C. Cardale. in ANIC: 1 I+. Highfields. IO.viii.1965, G. L. Jocumsen, in UQIC; l6'Vi. Mt Tamborine. I-25.v.1935. R. E. Turner. in BMNH. NEWSOUTH WALES: 1 '+. Kiora. near Moruya. 6.iii.1966. 2. Liepa. in ANIC.

Female Body and wing lengths, indices as in Table 1. Heacl-Vertex: with sparse setigerous punctures (Fig. 14) except near occipital carina; ocellar area with at most a few short rugae; postocellar area predominantly smooth. Occipital carina reaching oral margin (Fig. 16) or separated from it by at most 2.0-3.0 puncture diameters. Gena: in dorsal view not strongly swollen (Fig. 14). without tubercle; with dense setigerous punctures: with or without narrow smooth area adjacent to compound eye. Clypeal margin distinctly produced, broadly triangular (Figs 15, 17) or rounded (Fig. 2). Mandible apically slender, bidentate; ventral tooth distinctly longer than dorsal tooth (Fig. 2); external surface smoothly curved, without transverse fold or excavation. 272 I. D. NAUMANN Mesosoma-Pronotum (Figs 18-19) usually with sparse to very sparse setigerous punctures; sometimes dorsolaterally more densely punctate. Notaulix punctate, punctures sometimes weak. Tegula larger than pedicel. Metanotum medially shorter than mesoscutellum. Propodeum: punctate to rugose (Fig. 20); median carina variable, sometimes absent. Macropterous. Fore wing-Radial cell closed. Junction of m-cu and Cu, proximal to bifurcation of Cu ,. Hindwing-All (Fig. 38) or basal 0.5 (Fig. 37) of M between Cu, and Irm/Rs absent; never with basal stump of M beyond Cu,. Merasuma-Petiole: anterodorsal 0.5 with predominantly longitudinal rugae, sometimes also microreticulate; in lateral view more or less sinuate. Colour-Fore wing hyaline. Variation more or less continuous between following forms: (a) melanisric ,/arm: body predominantly black to dark brown; mandible, antenna, pronotal neck, axilla, mesoscutellum anterolaterally and posteriorly, legs, petiole posteriorly, T2-T7 brown to reddish orange; and (b) yellow /om: body predominantly bright yellow; ocellar area, upper frons, lower gena, mandibular teeth, antenna, pronotum laterally, propleuron anteriorly, lateral lobe of mesoscutum, scutellar fovea, mesopleuron laterally, propodeum laterally, small patch externally on each of mid and hind coxae. petiole, posterior metasomal segments contrastingly brown to black.

Male Differing from female as follows: Lengths, indices as in Table I. Head-Gena more sparsely punctate. Mesosoma-Pronotum more sparsely punctate. sometimes almost entirely smooth. Propodeum sometimes predominantly smooth with median, dorsal band of weak punctures and microreticulation. Merasoma-Petiole: anterodorsally more weakly sculptured; in lateral view straight. Apodeme of S8 anteriorly strongly expanded (Fig. 29). Colour-Body predominantly reddish orange or bright yellow; extreme lower gena, mandibular teeth, apical flagellar segments, lateral pronotum, propleuron, lateral lobe of mesoscutum, metanotum laterally, propodeum posterodorsally and laterally, petiole anteriorly, T2-T7 (in part) pale to dark brown. Discussion In females, at least, there is clinal variation in colour and sculpturing: specimens from northern Queensland (Crystal Cascades, Bellenden Ker) are of the yellow form and more polished, specimens from southern localities (Cleveland, Kiora) tend to be of the melanistic form and more densely punctate. Males from northern Queensland are also lightly coloured and smooth. No males of M.australicus have been collected from the southern half of the range of the species. Monomachus hesperius sp. n. (Figs 23-28, 30, 39; Table 1) Monomachus anripodalis 3en.w Schulz, 191 I: 414 (in part). TypeS-WESTERN AUSTRALIA:holoiype 0, 35.058 177.548, 7 km S by E of Albany, 18.iv.1983, E. S. Nielsen, E. D. Edwards, in ANIC (type number 7595). Paratypes: 2 99,Darlington, ii.1966, G. H. Lowe, in WAM; 1 d, 32.41s 115.398, YalgorupNational Park, 1.v.1983,E. S. Nielsen, E. D. Edwards, in ANIC: 1 ,T. 33.39s 115.42E. 14 km SW by S of Donnybrook, 27.iv.1983, E. S. Nielsen, E. D. Edwards, in ANIC; 1 i. 34.54s I17.02E, 7 km N of Kent R. Bridge, 19.iv.1983.E. S. Nielsen, E. D. Edwards, in ANIC. O/her marerid eSUmiflcd-wESTERN AUSTRALIA:1 ?Q, 1 .j, unlocalised, [?J.A.L.] Preiss, in ZMHU Female Body and wing lengths, indices as in Table 1. Head-Vertex: with dense setigerous punctures (Fig. 23); ocellar area with at most a few short rugae; postocellar area sometimes in part smooth. Occipital carina (Fig. 25) reaching oral margin. Gena: in dorsal view not strongly swollen, without tubercle; with dense setigerous punctures; with or without narrow smooth area adjacent to compound eye. Clypeal margin medially very weakly produced, bidentate (Figs 24, 26) or weakly tridentate. Mandible: apically broad, bidentate; teeth subequal; external surface smoothly curved, without transverse fold or excavation. Mesosomu-Pronotum: middorsally with sparse setigerous punctures and a few transverse striae (Fig. 27); dorsolaterally with dense setigerous punctures. Notaulix punctate, punctures sometimes weak. Tegula larger than pedicel. Metanotum medially shorter than mesoscutellum. Propodeum: punctate to rugose (Fig. 28); with or without poorly defined, posterodorsal unsculptured area; median carina variable, sometimes absent. Macropterous. Fore wing-Radial cell closed. Junction of m-cu and Cu,proximal to bifurcation of Cu ,. Hind wing-M continuous between Cu, and Irrn/Rs or narrowly interrupted near Cu,; always with at least basal stump of M beyond Cu,. AUSTRALIAN MONOMACHIDAE 273

Metasoma-Petiole: anterodorsally smooth or with a few weak wrinkles; in lateral view more or less sinua te. Colour-Fore wing hyaline. Body predominantly black to dark brown; clypeus, mandible (excluding teeth), flagellum, legs, ventral metasoma (in part) brown; small patch on vertex between lateral ocellus and compound eye, lateral lobe of mesoscutum, mesoscutellum reddish orange; or body more extensively reddish brown to reddish orange with black to dark brown restricted to lateral pronotum, mesoscutum anteriorly and laterally, mesopleuron, mesoscutellum laterally and posteriorly, metanotum (excluding dorsellum) and propodeum.

Male Differing from female as follows: Lengths, indices as in Table I. Head-Clypeal margin weakly produced, broadly convex. Mesosoma-Pronotum sometimes uniformly very sparsely punctate. Propodeum: punctation some- times uniformly very weak. Metasoma-Petiole: anterodorsally smooth; in lateral view straight. Apodeme of S8 anteriorly weakly expanded (Fig. 30). Colour-Body predominantly black to dark brown; lower gena, clypeus, mandible (excluding teeth), pronoturn dorsally (except for small black middorsal patch), mesoscutum, legs reddish brown to reddish orange; or body predr .ninantly reddish orange with pronotum laterally, mesopleuron, propodeum dorsolaterally, apex of metasoma brown.

Discussion M. hesperius is most similar to M. australicus, especially with respect to the shape of the head and mandible, and the development of the occipital carina. The 2 species are distinguished as detailed in the key. Females and males of M. hesperius have been correlated by similarities in wing venation, sculpturing and colour, and by collecting records. The holotype and 3 of the paratypes were collected at light, by the same collectors, during a 14-day period, although at 4 different localities between 80 and 190 km apart. The specific name is from the Latin hesperius (western). A badly damaged ?female (mesosoma crushed; propodeum, metasoma, portions of antenna, legs lost) and a male (lacking portions of antennae; portions of legs and metasoma glued to rest of body) in ZMHU are each labelled: (1) “N. Holl. Preiss”; (2) “1 1557” and “1 1558” respectively; (3) “M. antipodalis”. Almost certainly these are the specimens regarded by Schulz (1911: 414) as M.anfipodalis and stated to be from Western Australia. These damaged specimens are excluded from the type series. Acknowledgments I thank CSIRO colleagues Ms K. A. Pickerd (for scanningelectron micrographs), Dr D. H. Colless (for host information and comments on the manuscript), Dr J. F. Lawrence (for comments on the manuscript) and Ms J. C. Cardale (for bibliographic assistance); and the following curators for loan of material: Mr G. Holloway (AM), Mr G. Brown (BCRI), Mr N. Fergusson (BMNH), Dr J. Papp (HNHM), Drs A. Newton and M. Thayer (MCZ), Dr A. Neboiss (MV), Mr C. O’Toole (OUM), Dr 1. D. Galloway (QDPI), Mr E. C. Dahms (QM), Dr T. Houston (WAM) and Dr F. Koch (ZMHU). References ASHMEAD,W. H. (1 902)-Classification of the pointed-tailed wasps, or the superfamily Proct0trypidae.-I. Jl N.Y. ent. Soc. 10 240-247. BR~THES,J. (1928)-Nouveaux hymenopttres parasites du Chili. Revra chil. Hisr. nut. 31: 194-200. BRULL~,A. (1846tHisroire naturelle des insectes. HymknopGres (Suites d Buffon) 4. Roret: Paris. DALLATORRE, C. G. (1902)-Trigonalidae, Megalyridae, Stephanidae, Ichneumonidae, Agriotypidae, Pelecinidae. Hymenoptm Car. 3: 1088-1089. DE SANTIS,L. (1961tLas publicaciones entornologitas privadas de Arsene A. Girault. Revta Mus. La Plata (Zool.) 7: 123-172. GIRAULT,A. A. (1925)-Somegem-like or marvellous inhabitants of the woodlands heretofore unknown and by most never seen nor dreamt of. Privately published: Brisbane. JOHNSON,N. F. (1982)-Monomachidae. In Parker, S. P. (Ed.). Synopsisandclassification of’living organisms p. 661. McGraw Hill Book Company: New York. KLUG,J. C. F. (1841)-Die Arten der Gattung Pelecinus. Germar. Z. Enr. 3: 377-388. MUESEBECK,C. F. W. and WALKLEY,L. M. (1956)-Type species of the genera and subgenera of parasitic wasps comprising the superfamily Proctotrupoidea (Order Hymenoptera). Proc. U.S. natn. MUS.105: 319-419. 2 74 1. D. NAUMANN

RIEK,E. F. (1955)-Australian Heloridae, including Monomachidae (Hymenoptera). Ausr. J. Zool. 3 258- 265. RIEK. E. F. (1970)-Hymenoptera. In CSlRO (Ed.). The of’ Australia pp. 867-959. Melbourne University Press: Carlton. SCHLETTERER,A. (1889)-Die Hymenopteren-Gattungen Stenophasmus Smith, Monomachus Westw., Pe1rcinu.r Latr. and Megulyra Westw. Berl. em. Z. 33: 197-250. SCHULZ,W. A. (1907)-Alte Hymenopteren. Berl. eni. Z. 33: 303-333. SCHULZ,W. A. (I91 I)-Systematische Uebersicht der Monomachiden. Proc. I int. Congr. Ent. (Brussels, 1910) 2 405-422. SZEPLIOETI.V. ( 1903)-Aus der Sammlung des ungarischen National-Museums. Annls hist.-nut. Mus. natn. hung. 1: 364-395. TOWNPS,H. (1977)-A revision of the Heloridae (Hymenoptera). Conir. Am. ent. hi.lS(2): 1-12. WESTWOOD,J. 0. (1841)-On the Evaniidae and some allied genera. Ann. Mag. nut. Hist. (I) 7: 535-538. WESTWOOD,J. 0. (1843)-0n Erania and some allied genera of hymenopterous insects. Trans. R. ent. SOC. Lond. 3 237-278. WESTWOOD,J. 0. (I 874)-Thesaurus entomologicus oxoniensis. Clarendon Press: Oxford. [Manuscript received 2 1 September 1984.1