Znvertebr. Taxon., 1988, 2, 841-83

The Psychopsidae (Insecta :) of and the Oriental Region

T. R. New Department of Zoology, La Trobe University, Bundoora, Vic. 3083, Australia.

Abstract An illustrated synopsis is given of the described Psychopsidae of Australia (12 spp.) and the Oriental Region (2 spp.), together with the description of one new Australian species of Psychopsis. Generic limitations in the are reassessed and several genera raised on venational features (Balmes Navas, Magallanes Navas, Wernzia Navas) are shown to closely resemble typical Psychopsis Newman on genitalic characters; they are synonymised with Psychopsis, and keys are given to world genera of the family and the species of Psychopsis (14 spp.). Megapsychops Tillyard (1 sp.) is retained as a distinct .

Introduction The silky lacewings, Psychopsidae, are one of the smaller families of Neuroptera, containing only 20 described species. They occur in three major geographical areas (Tjeder 1960), namely Australia, the Oriental region and southern . Extant distri- bution of the family therefore suggests a Gondwanan origin, and there seems little doubt that Psychopsidae are an archaic group (Tillyard 1919, Kriiger 1922). Psychopsidae are generically confusing, although the family is unmistakable in appearance and integrity (Tillyard 1919, Kimmins 1939, Tjeder 1960), and the eight genera recognised in the more recent published literature have traditionally been separated on venation, wing shape and wing markings, with little appraisal of other features. A detailed appraisal of the African species by Tjeder (1960) has clarified intergeneric relationships for that fauna, and this paper is an attempt to elucidate relationships of the species from the Orient and Australia. None has yet been recorded from , although several Australian species are predominantly northern. With one exception, putative genera seem to be endemic to one of the three above-mentioned areas, but no detailed structural information on genitalia, particularly, has been available for any non-African psychopsid. The distributional exception is Balmes Navas, to which both Oriental and Australian taxa were attributed by Kimmins (1939). Clarification of the integrity of this genus is needed to fully assess relationships and possible origins of Australian Psychopsidae. All described species, following the arrangement of Kimmins (1939), have been examined, mostly from type material, and figures of wings and genitalia provided to form the basis for a fuller discussion of their relationships. The major controversy on the family has been the opposing views of generic limits expressed by Tillyard (1919) and Kimmins (1939). Tillyard noted that intraspecific variation in wing features of Australian species trans- cended generic limitations expressed by Navas (1P12, 1916), and regarded Magallanes Navas and Wernzia Navas as synonyms of Psychopsis Newman. He believed that: I, tribal divisions 08 18-0164/88/070841$09.00 842 T. R. New in the family (as raised by Navas 1916) were both unnecessary and misleading; and 2, several genera, listed above, raised on venational features (including the numbers of gradate series and the form and presence of the M-Cu anastomosis in the forewing) were not integral units, as different specimens-even in type series-of the same species could readily be allocated to different genera on such characters! Kimmins (1939), in an appraisal which has been adopted by more recent students, believed that venational features were useful for expressing generic limits and based his generic key largely on the form of the M-Cu anastomosis, if present, and on wing markings. Clearly, a broader range of characters is needed to appraise the relative merits of these two schemes, and Tjeder's (1960) treatment of African taxa showed clearly that Cabralis Navas, Notopsychops Tillyard and Silveira Navas are valid genera on genitalic features. In separating Cabralis and Notopsychops, for example, Kimmins (1939) relied on wing coloration, whereas Tjeder demonstrated differences in male genitalia (Notopsychops has entoprocessus, Cabralis does not) and spermatheca (Cabralis has one pair of accessory glandulae, Notopsychops has two pairs). The aim of this account is to determine whether equivalent non-wing features support maintenance of the remaining five genera, in particular the three which Tillyard amalgamated as Psychopsis, and Balmes. The detailed evidence is presented below, in a synoptic account of the species. Further background information on the relevant genera precedes this, together with the major conclusions of this study. In justifying his generic synonymy, Tillyard (1919, p. 768) commented (in part) as follows: 'The genus Wernzia, as defined by Navas, does not even include all the specimens of its type species. If, however, the definition of the genus be widened, then it at once forms a part of the Psychopsis-series of species, and should be included in that genus . . . The genus Magallanes is founded upon a type-species which grades into Wernzia in the characters of some of its individual members. This should also, therefore, be removed back to Psychopsis'. Tillyard (1919) recognised the unusual nature of his genus Psychopsella Tillyard, and this (as well as Orientichopsis Kuwayama) was synonymised with Balmes by Kimmins (1939), so that Balmes nominally occurs in Australia and includes the two described Oriental species. Many species of Psychopsidae are rare, and it has not been possible to trace represent- atives of both sexes of all species, a circumstance currently hampering complete reappraisal. Available genitalic structures are described and illustrated, and it is notable that several type specimens have proved to belong to a sex other than that cited in original diagnoses. The taxa treated here are much more similar genitalically than are the African species, and it is implied that structural diversification of the family may have been more conservative outside Africa. They reflect a featcre all too common in macy other groups of , and one which is challenging to interpret, namely, that groupings of species appearing to be both reliable and natural on relatively superficial characters (such as wing pattern) are not borne out by features which specialists intuitively feel should be of greater phylogenetic importance. In Neuroptera, genitalic structure is usually considered of greater significance than wing markings and, often, than small differences in venation, especially when the latter is known to be individually labile. Wing pattern has historically been a prime feature in division of the 'Psychopsis-group' of genera in Australia: the six species conventionally attributed to Psychopsis all have fasciated forewings (Figs 42, 59-62, 135) and a hind wing spot, the single species of Balmes lacks the latter, Wernzia has a heavily mottled forewing (Figs 153, 154) and that of Magallanes (Fig. 177) is more subtley mottled. Tillyard (1919) showed that forewing shape, earlier considered to be of generic importance, may merely reflect individual size. Larger individuals of P. elegans (Gukrin) have the tornus somewhat produced, and smaller specimens have this rounded as in Magallanes and Wernzia. The data presented later in this paper clearly indicate the overall similarity between species of these four 'genera'. Indeed, perhaps the most anomalous species on genitalia is Psychopsis barnardi Tillyard, in which the modified arcessus form is apparently unique. In wing features, barnardi is clearly congeneric with other 'banded' Psychopsis. The information now available on Psychopsidae, including that presented below, leads to the following conclusions. Australian and Oriental Psychopsidae 843

1. The African genera, as delimited by Tjeder (1960), appear to be well founded, on characters of male genitalia, spermatheca and praegenitale. Notopsychops is apparently unique in the family in having well-defined entoprocessus, and Silveira is the only known genus lacking superprocessus and praegenitale. 2. Of the remainder, Megapsychops is very distinctive on wing features and male genitalia, and less so on female genitalia, which approach the form of many other Australian species. This is not inconsistent with Tillyard's (1919) view that this is an archaic taxon. 3. The African taxa apparently have the arcessus simple. In all Australian species for which males are known, and birrnana, the arcessus is apically bifid. This feature may be an important difference delimiting the African from other psychopsid species. 4. The species comprising Psychopsis, Balmes, Wernzia and Magallanes are overall very similar in genitalic complement and form, and appear not to differ in characters of any major significance. Variation between species of Psychopsis sensu Kimmins is at least as great as that between the generotype species of the four genera. 5. If these four genera are to be retained as distinct, this can be done predominantly by accepting venational criteria on which they were originally diagnosed, in conjunction with wing pattern, and species allocation should remain as implied by Kimmins (1939). However, Tillyard's (1919) comments on individual variation in wing venation, which I can now endorse, cast doubt on the wisdom of this. Other than by using wing markings, I have not been able to produce a convincing key to separate the four, and more extensive 'definitions' of the genera overlap considerably. Balmes, Magallanes and Wernzia are regarded as synonyms of Psychopsis, and the contained species are individually transferred below. Whether or not 'species-groups' should be retained, reflecting former generic divisions, is a matter for personal preference, but I do not believe that they can be accorded formal status as subgenera. This account, although essentially a revision of extra-African Psychopsidae, is based on the contents of the following major collections: Australian National Collection, (ANIC) British Museum (Natural History), London (BMNH) South Australian Museum, Adelaide (SAM) Museum of , (NMV) I have not exhaustively appraised other collections in Australia. However, it seems that Tillyard's (1925) suspicion that 'a considerable number of new species might yet be found in out-of-the-way parts of Australia' may not be so, as virtually all specimens seen (about 180) are easily referable to described species. One exception is described below as a new species: in wing pattern it is intermediate between 'traditional' Psychopsis and Magallanes. Measurements are in rnm for the fo!!owing: forewing length (FW), hindwing !ecgth (HW), antenna length (A) and body length (B). Terminology follows that of Tjeder (1960) and is summarised in Figs 218-222. Wing venation drawings omit trichosors, which are present in all species. Drawings of abdominal apices are simplified by omission of details of the trichobothrial field and of setation. There is usually a large field of at least 30 small weakly defined trichobothria. The apex of the ectoproct, of both sexes, and the ventral and posterior borders of tergite IX are densely fringed with long hairs, especially in the female. Figures of genitalia are referred to a scaled figure of the relevant abdominal apex. Tjeder (1960) noted that the genital chamber of African female psychopsids he dissected was often filled with dark grains, which he considered to be bark powder, and others contained sand grains. Virtually all females dissected in the present work were similarly complemented, most commonly with bark scrapings, rarely also with algal spores and fungal hyphae, but occasionally with soil or sand.

Key to Genera of Recent Psychopsidae 1. Forewing with, at least, 4 branches to vein M; hindwing c. as wide as forewing; forewing with strongly embossed central areas (very large, FW 25 or more) (Australia) ...... Megapsych ops T. R. New

Forewing with, at most, 3 branches to vein M, more commonly 2; hindwing distinctly narrower than forewing; forewing without embossed central areas (generally smaller than above, FW rarely >20) ...... 2 Male: gonarcus with entoprocessus; female: spermatheca with 2 pairs of glandulae accessoriae (Africa) ...... Notopsychops Male: gonarcus without entoprocessus; female: spermatheca with, at most, 1 pair of glandulae accessoriae ...... 3 Male: superprocessus absent from parameres; female: praegenitale absent (Africa) ... Silveira Male: superprocessus present on parameres; female: praegenitale present ...... 4 Male: apex of arcessus simple; female: spermatheca with 1 pair of glandulae accessoriae (Africa) ...... Cabralis Male: apex of arcessus bifid; female: spermatheca without glandulae accessoriae (Orient*, Australia) ...... Psychopsis

*Male of P. formosa unknown; see later notes for recognition of this species.

Genus Psychopsis Newman Psychopsis Newman, 1842, p. 415. Arteriopteryx GuCrin, 1845, p. 389; partim Navas, 1916, p. 189. Zygophlebius NavBs, 1910, p. 82 (partim) [not sensu Navas, 1916, p. 1811. Wernzia Navas, 1912, p. 63; Tillyard, 1919, p. 773. Magallanes Navas, 1912, p. 69; Tillyard, 1919, p. 773. Balmes Navas, 1910, p. 85, syn. nov. Psychopsella Tillyard, 1919, p. 780; syn. of Balmes: Kimmins, 1939, p. 153. Orientichopsis Kuwayama, 1927, p. 123; syn. of Balmes: Kimmins, 1939, p. 153.

Type species: Psychopsis mimica Newman. Hindwing narrower than forewing; usually c. $ forewing width, occasionally c. fore- wing width; tornus of both wings usually rounded or slightly produced, sometimes angled; wings with 2-3 series of transverse gradate veins, occasionally more, or more numerous crossveins in of wing; costal series present or absent; forewing with or without M-Cu anastomosis; if present, the veins usually conjoint for distance, more rarely meeting at a point. Forewing markings various but always present: usually some indication of transverse banding or extensive suffusion, rarely a few small discrete spots; hindwing with or without dark spot on or near apex of vena triplica. Female. Tergite IX deep; praegenitale and subgenitale both present, often with sclerotised lateral areas and/or 2 setose tubercles between them; spermatheca slender, simple, without glandulae accessoriae; praegenitale present, usually extending well beyond rounded dorsal lobe, usually with central setae, often also with basal setae, dorsal lobe usually with 2 kinds of setae, stylus with 1 kind of thickened spine. Male. Ectoproct variously tapered; sternite IX often ornamented, usually with at least traces of lateral apodeme; arcessus bifid, usually with short dorsal setae near base; entoprocessus absent; superprocessus present, often setose and relatively small; parameres often medially convergent; hypandrium internum small.

Key to Species of Psychopsis The following simple key is based on features of wing pattern: anomalous specimens should be checked against the illustrations of wings and genitalia provided later in this account. 1. Hindwing without any large dark spot ...... 2 Hindwing with conspicuous large dark spot on or near apex of vena triplica (Australia) ...... 4 2(1). Forewing with markings predominantly limited to central f (Taiwan) ...... formosa Forewing with markings more diffuse or scattered (not Taiwan) ...... 3 3(2). Forewing markings diffuse (Fig. 11) (Orient) ...... birmana Forewing with darkest spots along vena triplica (Fig. 36) (Australia) ...... gallardi Australian and Oriental Psychopsidae 845

Forewing mottled, but without distinct transverse lines or banding ...... 5 Forewing with transverse lines or distinct banding, usually on pale ground ...... 6 Forewing with extensive brown or steely-blue suffusion on white ground; usually many irregular crossveins in basal of wing ...... coelivaga Forewing with few spots along vena triplica and on posterior margin; few irregular crossveins near internal gradate series ...... maculipennis Forewing predominantly brown or fawn, with extensive transverse brown suffusion and/or banding ...... 7 Forewing predominantly white or off-white, with few well-defined striae and suffusion associated with these ...... 9 Forewing fasciae very dark brown, with pronounced 'Y' across broadest part of wing (Fig. 135) ...... dumigani Forewing markings more diffuse (Fig. 177); no pronounced 'Y' across broadest part of wing ...... 8 Hindwing spot covering apex of vena triplica ...... meyricki Hindwing spot between apex of vena tripiica and wing apex ...... insolens Most transverse fasciae continuing unbroken to costal edge of forewing ...... mimica Most transverse fasciae not continuing unbroken to costal edge of forewing ...... 10 Broadest part of forewing without 2 or 3 major fasciae converging posteriorly to form 'Y' or trident marking; few, if any, markings along costal edge ...... 11 Broadest part of forewing with 2 or 3 fasciae converging posteriorly to form 'Y' or trident marking; sometimes traces of striae along costal edge, but these not continuous with major markings ...... 12 Main forewing markings 4 pairs of narrow brown striae, 2 pairs of which are near centre of wings ...... fillyardi Main forewing markings basal and preapical fasciae, central f of wing with only single stria ...... margarita Basal :of wing with extensive pinkish suffusion, without distinct outlining with dark brown or black (Fig. 60) ...... elegans Basal f of wing with less extensive pinkish suffusion, dark striae usually distinct (Figs 59, 62) ...... 13 Tornus of forewing strongly produced; a pair of short central striae between basal fascia and basal fork of 'Y' mark ...... gracilis Tornus of forewing not strongly produced; usually no central striae between fascia and basal form of 'Y' mark, rarely traces of one central mark ...... barnardi The following systematic notes on species do not repeat details of body coloration and markings given in earlier descriptions and are deliberately brief, as illustrations of available wing venation and pattern and genitalic features are provided.

Oriental Species Psychopsis birmana McLachlan, comb, rev. (Figs 6-23) Psychopsis birmana McLachlan, 1891, p. 321. (Burma). Balmes birmana (McLachlan), Kimmins, 1939, p. 153. Balmes terissinus Navis, 1910, p. 85 (Tibet); Kimmins, 1939, p. 153. Balmes notabilis Navas, 1912, p. 65 (China); Kimmins, 1939, p. 153. Psychopsis (Orientichopsis) birmana McLachlan; Kuwayama, 1927, p. 126.

Types. Holotype, 9, of P. birmana 'Birmah' (BMNH); Holotype, 0, of Balmes notabilis Navas (BMNH) (both seen); Holotype of Balmes terissinus Navas, specimen interpreted by Navas as u, Tibet, Ta-Tsien-Lou, J. Terisse, 1903 (? Paris, not traced). Description based on specimens compared with type in general appearance (BMNH) from Burma (Maymyo) (4), S. Shan States, 4000 ft (c. 300 m), Apr. 1916 (I), Laos, Xieng Khouang, 22.iv. 1919 (1). Wing venation as in Figs 6-10; only 2 gradate veins (FW) or 1 series (HW) in specimens seen; forewing extensively mottled (Fig. 11); no hindwing spot; forewing broad, hindwing broad, hindwing c. 4 forewing width. 846 T. R. New

Female. Apex of abdomen as in Fig. 12: ectoproct relatively small; tergite IX deep and ventrally rounded; tergite VIII with anterior border strongly arched; sternite VII deep; subgenitale (Figs 13-15) broad, with small lateral lobes; praegenitale (Figs 13-15) arched, rather narrow; spermatheca (Figs 16, 17) sinuous, slender; postgenitale (Fig. 18) small; gonapophyses laterales (Fig. 18) with large group of central ventral simple setae, a dorsal group of c. 40 long spatulate setae, most dorsal setae the longest; stylus with c. 18-20 thickened spines. Male. Apex of abdomen as in Figs 19,20: ectoproct narrow dorsally, elongate ventrally; sternite IX (Fig. 21) broad, rounded, with strong lateral apodeme, otherwise unornamented; gonarcus (Figs 22, 23) broad, with median dorsal expansion; arcessus arched, apically bifid, tapered only near apex; parameres with small dorsal superprocessus; arcessus, parameres, superprocessus all naked; hypandrium internum small and triangular. Dimensions. FW 14-15. Comments. This species is the generotype of Balmes Navas. It differs from the two other species formerly attributed to that genus as follows: formosa (female only known) has the gonapophyseal stylus shorter and about equal in length to the dorsal lobe, and the apex of the ectoproct longer; gallardi (male only known) has the superprocessus setose. All three species lack a hindwing spot (the only non-African Psychopsidae to do so) and the paratype of formosa has three forewing gradate vein series. P. birmana appears to be widely distributed in the Oriental region.

P. formosa Kuwayama, comb. rev. (Figs 1-5, 24-30) Psychopsis (Orientichopsis)formosa Kuwayama, 1927, p. 123. Balmes formosana (sic) (Kuwayama): Kimmins, 1939, p. 153.

Types. Described from 2 individuals stated to be males. Holotype: Formosa, Rengechi, near Horisha, Taichiu-district, l.viii.1925, T. Uchida; Paratype: Formosa, Kagi, Tainan- district Aug. 1921, S. Hirayama. (both Hokkaido University, Japan). The holotype (not seen) has the abdomen detached and reglued, and is in a fragile condition: it could not reasonably be shipped (S. Takagi in litt. 1986). The following notes are based on the paratype, a female, not a male as stated in original description. No further specimens of this species have been traced. Wing venation 2s in Figs 1-5; forewing markings as in Fig. 24; hindwing without spot; forewing with 3 transverse series of gradate veins, costal series incomplete; hindwing with 3 series of gradate veins; hindwing >) as broad as forewing. Female. Apex of abdomen as in Fig. 25; ectoproct with long tapered setose apex; tergite IX very deep and ventrally tapered; tergite VIII c. ) depth of tergite IX; sternite VII bilobed at apex; subgenitale (Figs 26, 27) broad; praegenitale (Figs 26, 27) arcuate; sperma- theca (Figs 28, 29) simple, slender; postgenitale small; gonapophyses laterales (Fig. 30) with stylus c. the same length as dorsal lobe; 2 rows of 10 and 21 short central setae, 3 longer basal setae; dorsal lobe with c. 35 long apically expanded setae, stylus with c. 18 thickened spines. Dimensions. FW 2 1. Comments. See under P. birmana. The forewing pattern is more restricted than in either of the other species hitherto attributed to Balmes, and the ectoproct form of the female is unusual. Kuwayama (1972) followed Kriiger's (1922) appraisal of Psychopsis in dividing it into species groups based on wing markings: he raised Orientichopsis Kuwayama to contain species without a hindwing spot, including P. (0.) birmana. Kuwayama also commented on the resemblance of formosa to gallardi, but did not formally include the latter in Orientichopsis. Orientichopsis was treated by Kimmins as a synonym of Balmes, and he also synonymised Psychopsella Tillyard (erected for gallardi) with that genus. Australian and Oriental Psychopsidae 847

Kimmins' synonymy of Orientichopsis was apparently accepted by Kuwayama, as the paratype bears an additional label: Balmes formosa (Kuwayama) det. Satoru Kuwayama 1960.

Australian Species Psychopsis gallardi (Tillyard), comb, nov. (Figs 31-41) Psychopsella gallardi Tillyard, 1919, p. 780. Balmes gallardi (Tillyard): Kimmins, 1939, p. 153.

Type. Holotype, 0,unlabelled (Tillyard: 'Mr. Gallard . . . informs me that he bred it from a larva found near Gosford, N.S.W.') (BMNH) (seen). Description based on type and one other male: Kenthurst, , in rocky cave, 14.i.1923 (ANIC). The female has not been seen. Wing venation as in Figs 31-35: both wings with 2 incomplete series of gradate veins; forewing also with few costal gradates; hindwing slightly more than width of forewing; forewing with irregular, predominantly transverse, markings (Fig. 36); hindwing white. Male. Apex of abdomen (Fig. 37): ectoproct ventrally broad, rounded, not conspicuously elongate; sternite IX (Fig. 38) rounded, with slender lateral apodeme; genitalia (Figs 39-41): gonarcus broad, dorsally angled; arcessus with broad apical bifurcation, basally with group of short setae; parameres slender, incurved, with membranous setose superprocessus; hypandrium internum small. Dimensions. FW 12;. Comments. See under P. birmana and P. formosa. Tillyard separated Psychopsis and Psychopsella on hindwing width relative to the forewing, and presence or absence of a hindwing spot.

Psychopsis mimica Newman (Figs 42-58) Psychopsis mimica Newman, 1842, p. 415, not Psychopsis mimica Froggatt, 1902, p. 367.

Type. Holotype, u, (BMNH) (seen). This is the most widely distributed and common species of Psychopsis in Australia: I have seen about 100 individuals, from (Charleville, Milmerran, Noondoo, Toowoomba), New South Wales (Broken Hill, Cobram, Deniliquin, Hay, Leeton, Merriwa, Mullaky, Oxley, , Wakool, Young), Victoria (Echuca, Lurgs, Mooroopna, Murrabit, Pyramid Hill, Wangaratta), South Australia (Adelaide, Bucheringa Forge, Berri, Balacoona, College Town, Melrose, Mt Painter, Murray Bridge) and (1, label illegible: ?Pingolly, Beverley). Apparently most common in the south east, but not recorded from . Wing venation as in Figs 43-47: forewing with 3 complete gradate series and a variable number of crossveins behind apex of vena triplica, these only rarely forming a complete 4th series; hindwing > f forewing width, with 2 or 3 series of gradate veins, the innermost one irregular and sometimes with only a few veins; costal crossveins of both wings usually complete or nearly so; forewing with irregular transverse striae (Fig. 42), the outer 3 pairs of 'parallel lines' sometimes with irregular 3rd line between them; 2 or 3 small black spots on tornus; tornus usually slightly produced. Female. Apex of abdomen (Fig. 48): ectoproct rather narrow and strongly narrowed ventrally; tergite IX large and deep, ventrally broadly rounded; tergite VIII rather broad; subgenitale (Figs 49, 50) broad, linked by sclerotised arms with domed ovoid praegenitale, 2 small setose tubercles between arms; spermatheca (Figs 51, 52) arched, broadened near apex; postgenitale small; gonapophyses laterales (Fig. 53) with stylus longer than dorsal lobe, 848 T. R. New a row of long basal setae, group of c. 20 central setae, dorsal lobe with dorsal group of c. 30 long simple setae (some with apex slightly expanded) and ventral group of c. 50 setae with apex more conspicuously expanded; stylus with c. 20-30 thickened spines. Male. Apex of abdomen as in Figs 54, 55: ectoproct ventrally broad and angled; sternite IX reflexed dorsally at apex (Fig. 56) where strongly bilobed (Fig. 58), strong lateral apodeme and much of apex strongly sclerotised; genitalia (Figs 56-58): gonarcus rather narrow and deep; arcessus with narrow incipiently-divided apex, bare: parameres small with long slender spiculate superprocessus; hypandrium internum small, rather broad. Dimensions. FW 13-22. Comments. This is the type species of Psychopsis. The following five species all have predominantly transverse striae on a very pale background (Figs 59-62, 125), and four of them have conventionally been allied with it on this feature. Genitalia of these species also imply similarity: in all those known, the female praegenitale and subgenitale are linked and the stylus of the gonapophyses are relatively long; male sternite IX is usually substantially ornamented and the arcessus (other than barnardi) is of grossly similar form.

Psychopsis elegans (GuCrin) (Figs 60, 63-74, 87-90) Arteriopteryx elegans GuCrin, 1845, p. 389. Psychopsis elegans (GuCrin): Hagen, 1866, p. 458; Tillyard, 1919, p. 777; Kimmins, 1939, p. 151. Psychopsis mimica Froggatt, 1902, p. 367; Tillyard, 1919, p. 777. Psychopsis newmani Froggatt, 1903, p. 454; Tillyard, 1919, pp. 766, 777. Zygophlebius verreauxinus Navas, 1910, p. 84; Tillyard, 1919, pp. 759, 777.

Type. Holotype (? Paris) not traced; determination based on specimens det. Tillyard, Kimmins et al. About 40 specimens seen, from Queensland: , Caloundra, Eidsvold, Herberton, Milmerran, Mt Tamborine, Paluma, Stanthorpe, Toowoomba; New South Wales: Sydney, Upper Colo, Wauchope; apparently widely distributed in both States, especially towards the east. Wing venation as in Figs 89-90: forewing with 3 or 4 gradate series, 3 series are usually complete, or nearly so (Fig. 87) and the 4th highly irregular in incidence; hindwing with outer 2 series of gradates almost always complete and innermost one variable, sometimes with only 2 or 3 irregular crossveins. Forewing pattern (Fig. 60) variable in detail, but large pinkish basal patch usually distinctive, and usually 2 dark dots in centre of distal f of wing, and 2 at tornus; tornus more produced in large individuals than in smaller ones. Female. Apex of abdomen as in Fig. 63: ectoproct broad, with apex rather broad; tergite IX very deep and ventrally rounded; tergite VIII and IX closely associated; sub- genitale (Figs 64, 65) with small central lobe and more prominent lateral lobes, relatively narrow; praegenitale simple, domed, with sclerotised lateral arms towards side of subgenitale; central setose membranous lobes; spermatheca and bursa (Figs 66, 67) broad, with strong lateral lobes and relatively membranous dorsal area; postgenitale slender; gonapophyses laterales (Fig. 68) with stylus long and slender, no basal setae; a group of c. 12 central setae near separation of dorsal lobe from stylus; dorsal lobe with c. 20 long curved setae and ventral group of c. 30 shorter apically expanded setae; stylus with c. 30 thickened spines. Male. Apex of abdomen as in Fig. 69: ectoproct expanded at apex; tergite IX ventrally tapered; sternite IX (Figs 70, 71) elaborately lobed and strongly divided at apex; genitalia (Figs 72-74): gonarcus slender, dorsally angled, lateral arms rather short; arcessus short, apically divided, dorsal group of short setae near base; parameres slender, with strong bare superprocessus; hypandrium internum small. Dimensions. FW 12-21 , Australian and Oriental Psychopsidae 849

Comments. As Tillyard (1919) noted, venational variation in this species would lead to different individuals being allocated to different genera should Navas' generic divisions be strictly followed. He also (p. 778) pointed out the close resemblance between larvae of elegans and coelivagus. The male sternite IX is more elaborated than in most other species of Psychopsis, and it is the only (known) species of Psychopsis, (conventional sense) in which the super- processus is entirely bare. The lack of basal setae on the gonapophyses laterales is also unusual in the 'striped wing' species.

Psychopsis gracilis Tillyard (Figs 59, 75-86, 92-95) Psychopsis gracilis Tillyard, 1919, p. 777.

Type. Holotype, Q (not w, as stated by Tillyard), New South Wales, Booyong, Nov. 1904 (BMNH) (seen). Apparently rather rare: specimens seen from Queensland (Bunya Mts, Mt Tamborine) and New South Wales (Type) (BMNH, NMV, ANIC). Description based on type and male from Bunya Mts. Wing venation as in Figs 92-95: forewing with 3 complete gradate series and irregular number of gradates between internal and discal series; hindwing with discal and terminal gradate series usually almost complete, internal series very irregular; forewing with tornus moderately to strongly produced; forewing pattern as in Fig. 59, with 4 distinct transverse bands, much less defined near costa than those of P. mimica: 1 or 2 black spots near junction of 3rd and 4th fasciae, 1 or 2 similar spots at tornus; distal shading pinkish; hindwing with tornus strongly angled, almost a right angle, c. width of forewing. Female. Apex of abdomen as in Fig. 75: ectoproct arched, apex narrow; tergite IX very deep and ventrally rounded; tergite VIII shallow; subgenitale (Figs 76, 77) small, with small lateral lobes; praegenitale ovoid, linked to subgenitale by broad lightly sclerotised arms; 2 small setose intermediate lobes; spermatheca (Figs 77, 78) broad and arched, no lateral ornamentation; postgenitale (Fig. 79) small; gonapophyses laterales (Fig. 79): stylus slender, well separated from dorsal lobe, a few basal and central ventral setae, a larger group of c. 7 thickened median setae near separation of stylus and dorsal lobe; dorsal lobe with dorsal group of c. 25 slender curved setae and c. 35 shorter apically thickened ventral setae; stylus with c. 20 short thickened spines. Male. Apex of abdomen as in Figs 80, 81: ectoprocts posteriorly convergent, dorsally narrow, ventrally broader and angled; sternite IX (Figs 82, 83) dorsally angled, with apex transverse and slightly bilobed; genitalia (Figs 84-86): gonarcus narrow and very deep, with lateral groups of setae; arcessus basally broad, narrowed and divided at apex with minute dorsal setae; parameres fused medially, with tapered setose superprocessus also bearing basal spicules. Dimensions. FW 19-21. Comments. Most similar in size and superficial appearance to P. mimica and P. barnardi but clearly distinct from both on forewing pattern and genitalia of both sexes.

Psychopsis barnardi Tillyard (Figs 62, 96-1 10) Psychopsis barnardi Tillyard, 1925, p. 388; Kimmins, 1939, p. 151

Type. Holotype, u, Queensland, Blackwater, 9.xi.1922, W. B. Barnard (BMNH) (seen). The following 2 individuals (Qld) were noted by Tillyard in his original citation in the paragraph 'Types' but not there specifically designated as paratypes. Both are labelled 'Paratype o" in his hand: Q, Huntley Stn, 10.xi.21; w,Clermont, 15.xi.24 E.J.D. (both ANIC). 850 T. R. New

Other material examined. Six ex. Queensland: Biloela, Blackwater, Clermont, Springwise, - hampton (BMNH, ANIC, NMV). Description based on type and other Queensland material.

Wing venation as in Figs 96, 108-110: both wings with 3 complete sets of gradate veins; forewing with few additional gradates forming short row between internal and discal series; costal series complete or nearly so in both wings; hindwing only slightly narrower than forewing, strongly angled at tornus; forewing with transverse fasciae (Fig. 62) and additional pinkish to reddish brown blotches; usually 3 black dots near convergence of major distal fasciae and 3 on margin at tornus; hindwing with black spot behind apex of vena triplica, tornus often suffused with brown and/or with 1 or 2 small dark spots. Female. Apex of abdomen as in Fig. 102: ectoproct rather slender; tergite IX deep and somewhat arcuate ventrally; tergite VIII strongly tapered ventrally; subgenitale (Fig. 103) with short lateral lobes; praegenitale (Figs 103, 104) elaborate, domed, annular, with strong sclerotised posterior arms to sides of subgenitale; spermatheca (Figs 105, 106) short, apically tapered and divided; postgenitale rather blunt and broad; gonapophyses laterales (Fig. 107) with slender stylus extending well beyond dorsal lobe, few basal setae, a central ventral group of c. 10 setae, dorsal lobe with c. 30 long curved dorsal setae and c. 40 shorter apically expanded ventral setae; stylus with c. 16-20 short thickened spines. Male. Apex of abdomen as in Fig. 97: ectoproct with tapered posterior lobe, ventrally excavated; tergite IX deep; sternite IX with strong lateral apodemes, apex (Fig. 98) reflexed dorsally and strongly lobed, Genitalia (Figs 99-101): gonarcus rather narrow, sides deep; arcessus with short bifid apex raised above broad rounded ventral process, short dorsal setae on apex, and ventral short setae below preapical constriction; parameres elongate, with apical ventral membranous lobe; long slender dorsal superprocessus with scattered short setae. Dimensions. FW 17-19. Comments. Although clearly allied with the three foregoing species on wing features and many genitalic characters, the arcessus form of P. barnardi is highly unusual: the subapical lobe is not present in any other species for which the male has been examined.

Psychopsis margarita Tillyard (Figs 61, 111-119) Psychopsis margarita Tillyard, 1922, p. 37; Kimmins, 1939, p. 151.

Type. Holotype, Q (not 0,as stated by Tillyard), New South Wales, Woodford, 29.xii.1920, M. Waterhouse (BMNH) (seen). Tillyard also designated a paratype (?) 0 taken with the holotype (not traced). Although the two individuals seen by Tillyard were stated to be part of 'a large number' seen together, the species appears to be rare, and no other material has been examined. Description based on type. Wing venation as in Figs 111-1 14: forewing with only irregular basal costal gradate veins, 2 main series of gradate veins: terminal series absent; hindwing with few basal costal gradates, few internal gradates, discal series complete, terminal series absent; hindwing slightly >$forewing width, tornus broadly rounded; forewing pattern (Fig. 61) with only basal mark and major fascia near apex well defined, trace of slender median fascia, markings predominantly brownish or reddish, with few small black dots; hindwing hyaline except large rounded black spot beyond and behind apex of vena triplica. Female. Apex of abdomen as in Fig. 115: ectoproct arched, dorsally strongly tapered; tergite IX deep and ventrally rounded; tergite VIII slender; subgenitale (Figs 116, 117) large, lateral lobes arched; praegenitale (Figs 116, 117) with transverse posterior apex, dorsally arched, no strong posterior arms, but 2 small setose lobes between praegenitale and sub- genitale; spermatheca (Figs 116, 118) slender, anteriorly curved dorsally; postgenitale small; gonapophyses laterales (Fig. 119) with stylus slender, much longer than dorsal lobe; a Australian and Oriental Psychopsidae 85 1 concentrated group of basal setae, a small preapical group of setae; dorsal lobe with group of c. 15 long simple setae and c. 40 shorter apically expanded ventral setae; stylus with c. 18 short thickened spines. Dimensions. FW 18. Comments. The gradate vein complement is considerably less than that of most other species with striated forewings but, because of variation known in other species, the significance of this is not wholly clear. In hindwing shape, margarita seems to be most closely related to elegans and mimica. The stylus is unusually narrow, and the lateral arms between praegenitale and subgenitale are unusually lightly represented.

Psychopsis tillyardi, sp, nov. (Figs 120- 13 1) Type. Holotype, 9, Northern , Narbalek, 12°19'S.,133019'E., ll.iv.1983, M. Wells (ANIC). Coloration. Creamy white. Eyes black. Palpi pale. Labrum pale; clypeus brown medially; frons anteriorly pale, a dark brown band immediately below pale antennal socket, interantennal space dark brown, a narrow transverse band above antennal socket, enclosing pale -shaped median spot. Vertex midline narrowly pale, an anterior brown spot each side, much of posterior pale greyish brown with large 'V' embracing much of dorsal orbit each side; long pale adpressed setae anteriorly. Antennae pale cream, brown from about segment 12 on. Pronotum with trace of narrow dark stripe along anterior half, dense erect pale setae. Pterothorax and abdomen apparently wholly pale (discoloured), most pterothoracic setae long and pale, some scutal setae black. Legs pale, TII, I11 with slight darkening along outer edge. Wings pale, off-white; forewing with dark brown striae (Fig. 125) not extending anterior to vena triplica, no defined posterior convergence of striae towards posterior of wings, a dark brown tornal spot and several striae broadened at posterior in basal of wing; much of rest of wing membrane pale greyish brown; most vein setae dark. Hindwing with large dark greyish brown spot immediately behind apex of vena triplica and enclosing anterior 7 or 8 discal gradate veins; much of apex of wing irregularly suffused with pale greyish brown; vein setae pale. Antennae sturdy, 38-segmented. No setose tubercles on vertex. Wing venation as in Figs 120-124: both wings with 3 complete series of transverse gradate veins, and costal gradates complete; forewing with few additional crossveins basal to anterior part of discal gradates; M and Cu anastomosing. Female. Apex of abdomen as in Fig. 126: tergite IX deep, with narrow rounded ventral edge; tergite VIII slender, fused with tergite IX ventrally, and heavily sclerotised at c. depth of distinct sulcus; praegenitale (Figs 127, 128) dorsally flexed, an outer rim enclosing small conical structure, strong lateral arms slightly bowed towards sides of subgenitale; subgenitale with curved lateral processes; pair of strong setose lobes c. 31 distance between praegenitale and subgenitale; spermatheca (Figs 129, 130) small, dorsally arched, simple; postgenitale rather broad; gonapophyses laterales (Fig. 131): stylus slender, extending well beyond rounded dorsal lobe, a group of c. 11 basal setae, 8 long blunt central setae, dorsal lobe with c. 12 long curved simple dorsal setae, c. 45 shorter dense apically-expanded ventral setae, stylus with c. 30 rather broadly rounded thickened spines. Male. Unknown. Dimensions. FW 22, HW 17f, A c. 4, B 11. Comments. The wing pattern of this species is in some features intermediate between the heavily striated pattern typical of Psychopsis sensu Kimmins and the more diffuse transverse shading of species formerly placed in Magallanes. It is most similar to that of P. margarita but is considerably paler, and the forewing venation differs considerably. 852 T. R. New

Female genitalia of the two species are also distinctive, and confirm that this individual cannot be allocated to any previously described species. This is the only psychopsid I have seen from the , and further collecting is needed in to determine its distribution.

Psychopsis dumigani Tillyard (Figs 132-141) Psychopsis dumigani Tillyard, 1922, p. 36; Kimmins, 1939, p. 151.

Type. Holotype (stated by Tillyard to be a 0,but abdomen now missing), Queensland, Clermont, ll.xi.1919, E. J. Dumigan (BMNH) (seen). Paratype (?)o, noted by Tillyard from same locality, not traced.

Other material examined. Three Q, Clermont (ANIC, NMV). The male remains unknown.

Wing venation as in Figs 132-134: forewing with 3 almost complete series of transverse gradate veins, costal series also nearly complete; hindwing with costal, discal and terminal gradate series almost complete; internal series irregular, usually with only c. 4 veins; hindwing c. $ forewing width, tornus broadly rounded; forewing tornus slightly produced; forewing extensively marked with dark brown to black, dark striae as in Fig. 135, with much of membrane mottled with paler brown; hindwing with irregular brownish suffusion and more conspicuous spots near end of vena triplica and near tornus. Female. Apex of abdomen as in Fig. 136: ectoproct arched; tergite IX very deep and ventrally tapered; tergites VIII and IX closely associated; subgenitale (Figs 137, 138) small, lateral lobes slender; praegenitale small, strong dorsolateral sclerotised arms towards sides of subgenitale, 2 small setose lobes c. distance between praegenitale and subgenitale; spermatheca (Figs 139, 140) short and broad; postgenitale rather narrow; gonapophyses laterales (Fig. 141) with stylus slender and extending well beyond dorsal lobe; c. 9 basal setae, radiating group of setae near base of stylus; dorsal lobe with dorsal group of c. 12 long curved simple setae and ventral patch of c. 40 shorter apically-expanded setae; stylus with c. 22 thickened spines. Dimensions. FW 13-16. Comments. The wing coloration of this species is very distinctive, but the transverse striae suggest that dumigani is related to the several foregoing species. Tillyard (1922) suggested that it forms a link between them and the insolens-group of smaller species with drabber markings and rounded tornus.

Psychopsis maculipennis Tillyard (Figs 142- 147) Psychopsis maculipennis Tillyard, 1925, p. 389. ? Wernzia maculipennis (Tillyard): Kimmins, 1939, p. 152.

Type. Holotype, Q (not 0, as stated by Tillyard), Western Australia, '1920-130/ Roebourne' (Western Australian Museum, Perth) (seen). This species remains known only from the holotype, which has not been dissected. Wing venation as in Figs 142-146: both wings with 3 complete sets of gradates and costal series almost complete; forewing internal series partially duplicated, and few additional crossveins between discal and internal series; hindwing only slightly narrower than forewing; tornus of both wings broadly rounded; forewing with small black spots along and beyond vena triplica and along posterior margin (Fig. 147); hindwing with ovoid black spot below end of vena triplica, otherwise pale. Dimensions. FW 19. Australian and Oriental Psychopsidae 853

Comment. Kimmins (1939) tentatively allied this species with coelivagus in the genus Wernzia, but did not comment on this alliance.

Psychopsis coelivagus (Walker), comb. rev. (Figs 148-166) Hemerobius coelivagus Walker, 1853, p. 279. Psychopsis coelivagus (Walker): Hagen, 1866, p. 458. Wernzia coelivaga (Walker): Navas, 1912, pp. 63, 195. Psychopsis coelivagus (Walker): Tillyard, 1919, p. 778. Wernzia coelivagus (Walker): Kirnmins, 1939, p. 152.

Type. Sex unknown (abdomen missing) (BMNH) (seen). Apparently widely distributed in eastern Queensland; c. 30 ex. seen (BMNH, ANIC, NMV); SAM has a specimen labelled 'Vic: Horsham': this record needs confirmation. Wing venation as in Figs 148-152; forewing with costal, terminal and discal gradate series complete or almost so, internal series variable, often many irregular multiple gradates not forming distinct series (Fig. 148) or (occasionally) with very few veins present; hindwing with costal and terminal series complete, discal series variable, internal series absent; hindwing c. $ forewing width, tornus broadly rounded; forewing with extensive but variable metallic blue or brownish markings (Figs 153, 154) on pure white ground; hindwing with spot of variable size at apex of vena triplica, sometimes with more general brownish suffusion towards tornus. Female. Apex of abdomen as in Figs 155, 156: ectoproct with reflexed narrow apical lobe and long apical setae, tergite IX broad, ventrally tapered; tergite VIII very short; praegenitale (Figs 157, 158) transverse; subgenitale small; spermatheca (Figs 159, 160) slender, sinuous; postgenitale small; gonapophyses laterales (Fig. 161): stylus extending beyond dorsal lobe; no basal setae, row of c. 8 long central setae and marginal row of central spicules; dorsal lobe with c. 15 long dorsal setae, ventral group of c. 20-25 shorter apically expanded setae; stylus with c. 16 short thickened spines, concentrated at apex. Male. Apex of abdomen as in Fig. 162: ectoproct with short rounded apex; tergite IX deep; sternite IX (Fig. 163) with strong lateral apodeme, apex slightly reflexed and lobed; genitalia (Figs 164-166): gonarcus broad and deep; arcessus long, apically divided, a group of short dorsal setae; parameres medially convergent, with setose superprocessus. Dimensions. FW 10-13. Comments. This, the generotype of Wernzia Navas, does [as Tillyard (1919) indicated] have rather variable wing venation, and only some specimens within clearly conspecific series tally with Navas' generic diagnoses. The wing markings are unusual in pattern and imply that the species is relatively isolated, but genitalic features suggest a much closer relationship to other species here placed in Psychopsis. Male genitalia, especially the form of the superprocessus, indicate particularly close affinity with the next two species, both of which have conventionally been placed in Magallanes. Additionally, Tillyard (1919) showed that venational features of Wernzia and Magallanes, as defined by Navas, intergrade.

Psychopsis insolens McLachlan, comb. rev. (Figs 172-190) Psychopsis insolens McLachlan, 1866, p. 114. Magallanes insolens (McLachlan): Navas, 1912, p. 69. Psychopsis insolens (McLachlan: Tillyard, 1919, p. 779. Magallanes insolens (McLachlan): Kirnrnins, 1939, p. 152. Psychopsis coelivaga (Walker): Froggatt, 1902, p. 367; Tillyard, 1919, p. 779.

Type. Holotype, Q,Queensland (BMNH) (seen); 2 paratypes (BMNH) (seen). 854 T. R. New

Other material examined. About 40 ex., widely distributed in eastern Queensland: Brisbane, Glen Aplin, Herberton, Killarney, Mt Tamborine, Stanthorpe; New South Wales: Armidale, Asquith, Audley, Batemans Bay, Bawley Pt, Coffs Harbour, Emmaville, Nowra; with one specimen from Canberra (BMNH, ANIC, NMV). Description based on Queensland specimens compared with type.

Wing venation as in Figs 172-176: forewing with discal and internal gradate series complete or almost so, terminal series absent or represented by few crossveins in anterior 1 of wing, costal series usually incomplete distally; hindwing with discal series relatively complete, internal series usually represented by 1-4 crossveins, terminal and costal series absent; hindwing

Psychopsis meyricki McLachlan, comb. rev. (Figs 167-171, 191-195) Psychopsis meyricki McLachlan, 1887, p. 30. Magallanes meyricki (McLachlan): Navas, 1916, p. 197. Psychopsis meyricki McLachlan: Tillyard, 1919, p. 779. Magallanes meyricki (McLachlan): Kimmins, 1939, p. 152.

Type. Holotype, u and 6 paratypes (all u), New South Wales, labelled 'Kosciusko, 2800 feet' 20.i. 1885, Meyrick. Tillyard (1919) showed that the type locality is near Jindabyne, not actually on Kosciusko (all BMNH) (seen).

Other material examined. None. This species has apparently not been taken again since 1885, and the female remains unknown.

Wing venation as in Figs 167-171: very similar to that of P, insolens, but the forewing gradate series seems to be slightly more irregular; wing markings and shape very similar to those of insolens, except hindwing spot covering apex of vena triplica. Male. Apex of abdomen as in Figs 191, 192: apex of ectoproct broadly rounded; tergite IX deep; sternite IX broad, scarcely ornamented; genitalia (Figs 193-195): gonarcus anteriorly broad and deep; arcessus long, strongly bifid, with dorsal basal short setae; parameres long, with preapical rounded setose superprocessus, a membranous median connection. Australian and Oriental Psychopsidae

Dimensions. FW 11-1 4. Comments. Tillyard (1919) separated insolens and meyricki in his key on the position of the hindwing dark spot. The two species otherwise seem to be extraordinarily similar, and their male genitalia differ only in rather trivial features. They are here maintained as distinct species, pending discovery of fresh material of P. meyricki, but I believe it likely that meyricki may eventually be reduced to a synonym of P, insolens.

Megapsychops Tillyard Megapsychops Tillyard, 1919, p. 771; Kimmins, 1939, p. 149.

Type species: Psychopsis illidgei Froggatt. This genus has not been augmented, and contains the single most spectacular species of Australian Psychopsidae. Tillyard (1919) defined Megapsychops in terms of key characters, as follows: 'Ocelli vestigial. Fore- and hindwings of almost equal width, with the branches of Rs excessively numerous (27-31 in forewing, 18-24 in hind), exceedingly close together, so as to give the wing the appearance of the close texture of spun silk; dorsal margin strongly excavated before the tornus, the latter very prominent, especially in the forewing. M at least four-branched. Forewings with raised or embossed areas'. The 'ocelli' referred to appear to be merely raised setose tubercles above the antenna1 sockets, and these are present to a varying extent in many species here referred to Psychopsis. The contrasted key feature used by both Tillyard and Kimmins of 'ocelli vestigial' versus 'ocelli absent' is therefore invalid, but Megapsychops is very distinct on the highly complex venation of the sole included species. This venation (Figs 196-200) is more complex than that of any other known psychopsid and Tillyard (1919, p. 781) implied that M. illidgei may be closely related to found in Queensland. The venation bears some resemblance to that of . It seems likely that Megapsychops may be the most archaic living psychopsid, as all other known species have the wing venation considerably less complex. M, illidgei is also the largest known species. Despite considerable female genitalic similarity to some species of Psychopsis, the venation is clearly sufficient to retain Megapsychops as distinct. There is greater difference in venation between Mega- psychops and Psychopsis than between any other members of the Psychopsidae.

Megapsychops illidgei (Froggatt) (Figs 196-217) Psychopsis illidgei Froggatt, 1903, p. 455. Megasychops iNidgei (Froggatt): Tillyard, 1919, p. 771; Kirnmins, 1939, p. 149.

Type. Holotype, 9, Queensland, Mt Tamborine, W. W. Froggatt (E. F. Riek genitalic prep. no Neur. Gen. 89) (ANIC) (seen).

Other material examined. About 20, all Q,from: Queensland: mainly Mt Tamborine, also Bunya Mts; New South Wales: Sandy Flat, nr Tenterfield (BMNH, ANIC, NMV); lu, Mi Tamborine (Queensland Museum).

Wing venation as in Figs 196-200: both wings with costal, terminal, discal and internal gradate series complete, or almost complete; forewing sometimes with considerable multi- plication of gradate veins in discal area, and main discal series sinusoidal; M with at least 4 main branches; both wings with tornus expanded and rounded; hindwing almost as wide as forewing; forewing with strongly embossed (raised) areas, especially at and behind apex of vena triplica; forewing with strong brown to black markings as in Fig. 201, rarelv also with strong basal costal patch anterior to main basal wing patch; hindwing with large dark spot over apex of vena triplica (Fig. 202) and patches of black vein hairs at tornus and near wing base. 856 T. R. New

Female. Apex of abdomen as in Fig. 203: ectoproct rather broad, tergite IX deep, ventrally angled; tergite VIII very deep, narrow; praegenitale (Figs 204, 205) arcuate, with anterior sclerotised lateral arms bordering margin of sternite; subgenitale (Figs 204, 205) with lateral processes prominent and slightly convergent; spermatheca (Figs 206, 207) basally broad and slightly bilobed posteriorly, dorsally slender; postgenitale moderately developed; gonapophyses laterales (Fig. 208): stylus extending beyond dorsal lobe; a group of long basal setae, larger group of ventral central setae, some slightly spatulate; dorsal lobe with all setae apically expanded, those of dorsal group longer than ventral group; stylus with c. 25 short thickened spines. Male. Apex of abdomen as in Figs 209, 210: ectoproct with long tapered slender ventral process, and additional similar process from near inner base; tergite IX slender; sternite IX broad, membranous, with slender dorsally-reflexed apical process (Figs 211, 212); genitalia (Figs 213-217): gonarcus broad, lightly sclerotised, with small setose dorsolateral flanges, few small lateral setae; parameres membranous, with broad setose apex, no distinct super- processus; a heavily sclerotised median structure (?gonapsis); hypandrium internum small. Dimensions. FW 25-30. Comments. The male of M. iNidgei appears to be substantially rarer than the female, and only one individual of this sex has been seen. Whereas the female genitalia closely resemble those of Psychopsis, the male genitalia are extremely distinctive, and differ consp:cuously from those of any other known psychopsid. Features of particular interest include: I, the twin elongate ectoproct processes; 2, the slender filamentous apex to sternite IX; and 3, the median structure between the parameres. This structure is tentatively referred to above as a 'gonapsis', but this name is used only as a matter of convenience and not to imply homology with similarly named structures in other families; no equivalent structure has been recorded in other psychopsids. The setose apex of the parameres possibly represent the superprocessus of many Psychopsis, but the latter do not appear to have a parallel median structure similar to that of Megapsychops. The apically divided arcessus resembles that typical of Psychopsis.

Acknowledgments I am very grateful to the following Curators, who permitted me to study collections in their care, loaned single types or critical specimens for examination, or responded to my queries: Dr P. C. Barnard and Mr S. J. Brooks (BMNH), Miss J. C. Cardale (ANIC), Dr A. Neboiss (NMV), Mr E. C. Dahms (Queensland Museum), Dr E. G. Matthews (SAM), Dr T. F. Houston (Western Australian Museum), Dr C. N. Smithers (Australian Museum) and Dr S. Takagi (Hokkaido University).

References Froggatt, W. W. (1902). Notes on Australian Neuroptera and their life-histories. Proc. Linn. Soc. N.S. W. 27, 358-69. Froggatt, W. W. (1903). Notes on the genus Psychopsis Newman, with descriptions of new species. Proc. Linn. Soc. N.S. W. 28, 453-6. GuBrin-MenCville, F. E. (1845). Iconographie du Regne . 111. Insectes. pp. 1-559. (Paris.) Hagen, H. A. (1866). Hemerobidarum synopsis synonymica. Stettin. Entomol. Ztg. 27, 369-462. Kimmins, D. E. (1939). A review of the genera of the Psychopsidae (Neuroptera) with a description of a new species. Ann. Mag. Nat. Hist. (11) 4, 144-53. Kruger, L. (1922). Psychopsidae. Beitrage zu einer Monographie der Neuropteren-Familie der Psychopsiden. Stettin. Entomol. Ztg. 83, 17-48. ,, Kuwayama, S. (1927). On a new species of ~s~cho~&aefrom Formosa. Ins. Matsum. 1, 123-6. McLachlan, R. (1866). On some new species of neuropterous insects from Australia and New Zealand, belonging to the family . J. Entomol. 2, 11 1-6. McLachlan, R. (1887). Psychopsis meyricki, nsp. Entomol. Mon. Mag. 24, 30-1. McLachlan, R. (1891). An Asiatic Psychopsis (Ps. birmana, n.sp.). Entomol. Mon. Mag. 27, 320-1. Navas, L. (1910). Hemerobidos nuevos. Con la clave de 10s tribus y generos de la familia. Broteria 9. 69-90. Australian and Oriental Psychopsidae 857

Navas, L. (1912). Crisopidos y Hemerobidos (Ins. Neur.) nuevos o criticos. Broteria 10, 98-113. Navas, L. (1916). Essayo monografico de la familia de 10s Sicopsidos (Ins. Neur.). Assoc. Espanola. para el Progr. de la Ciencias: Congreso de Valladolid, Oct. 1915, 181-210. Newman, E. (1842). Entomological notes. Entomol. 1842, 413-5. Tillyard, R. J. (1919). Studies in Australian Neuroptera. No. 6. The family Psychopsidae, with descriptions of new genera and species. Proc. Linn. Soc. N.S. W. 43, 750-86. Tillyard, R. J. (1922). Descriptions of two new Australian species of Psychopsis. Aust. J. Zool. 3, 35-8. Tillyard, R. J. (1925). Two new species of silky lacewings from Australia. Proc. Linn. Soc. N.S. W. 50, 387-90. Tjeder, B. (1960). Neuroptera-Plannipennia. The lace-wings of Southern Africa. 3. Family Psychopsidae. S. Afr. Anim. Life 7, 164-209. Walker, F. (1853). List of the specimens of Neuropterous Insects in the collection of the British Museum. 11. pp. 193-476. (London.)

Manuscript received 16 December 1986, accepted 23 February 1987 T. R. New

Figs 1-10. Psychopsis spp., Oriental species, venation: (1-5, P, formosa Kuwapama), I, forewing; 2, hindwing; 3, forewing, M-Cu area; 4, 5, apex of vena triplica of (4) forewing; (5) hindwing; (6-10, P. birmana McLachlan) 6, forewing; 7, hindwing; 8, forewing, M-Cu area; 9, 10, apex of vena triplica of (9) forewing, (10) hindwing. Australian and Oriental Psychopsidae

Figs 11-23. Psychopsis birmana McLachlan: 11, forewing pattern; (12-18, female) 12, apex of abdomen, lateral; 13, praegenitale and subgenitale, lateral; 14, same, ventral; 15, same, posterior; 16, spermatheca, lateral; 17, same, ventral; 18, postgenitale and gonapophysis, lateral, with inset of stylus spine; (19-23, male) 19, apex of abdomen, lateral; 20, ectoprocts and tergite IX, dorsal; 21, sternite IX, ventral; 22, genitalia, lateral; 23, same, dorsal. (Scales in mm to Figs 12, 19.) T. R. New

Figs 24-30. Psychopsis formosa Kuwayama, female: 24, forewing pattern; 25, apex of abdomen, lateral; 26, praegenitale and subgenitale, ventral; 27, same, lateral; 28, spermatheca, lateral; 29, same, ventral; 30, gonapophysis, with inset of dorsal lobe seta and stylus spine. (Scale in mm to Fig. 25.) Australian and Oriental Psychopsidae

Figs 31-41, Psychopsis gallardi (Tillyard), male: 31, forewing venation; 32, hindwing venation; 33, forewing, M-Cu area; 34, 35, apex of vena triplica of (34) forewing, (35) hindwing; 36, forewing pattern; 37, apex of abdomen, lateral; 38, sternite IX, ventral; 39, genitalia, lateral; 40, same, antero- dorsal; 41, posterior. (Scale in mm to Fig. 37.) T. R. New

Figs 42-46. Psychopsis mimica Newman: 42, forewing pattern; 43, forewing venation; 44, hindwing venation; 45, 46, apex of vena triplica of (45) forewing, (46) hindwing. Australian and Oriental Psychopsidae

Figs 47-58. Psychopsis mimica Newman: 47, forewing, M-Cu area; (48-53, female) 48, apex of abdomen, lateral; 49, praegenitale and subgenitale, lateral; 50, same, ventral; 51, spermatheca, lateral; 52, same, ventral; 53, postgenitale and gonapophysis, lateral; (54-58, male) 54, apex of abdomen, lateral; 55, ectoprocts and tergite IX, dorsal; 56, genitalia and sternite IX, lateral; 57, gonarcus and arcessus, dorsal; 58, genitalia and sternite IX, posterior. (Scales in mm to Figs 48, 54.) T. R. New

Figs 59-62. Forewing patterns of (59), Psychopsis gracilis Tillyard, (60), P. elegans (GuCrin), (61), P. margarita Tillyard, (62), P. barnardi Tillyard. Australian and Oriental Psychopsidae

Figs 63-74. Psychopsis elegnns (Guerin): (63-68, female) 63, apex of abdomen, lateral; 64, praegenitale and subgenitale, lateral; 65, same, ventral; 66, spermatheca, lateral; 67, same, ventral; 68, postgenitale and gonapophysis, lateral; (69-74, male) 69, apex of abdomen, lateral; 70, sternite IX, ventral; 71, same, lateral; 72, genitalia and sternite IX, posterior; 73, genitalia, lateral; 74, gonarcus and arcessus, dorsal. (Scale in mm to Figs 63, 69.) T. R. New

Figs 75-86. Psychopsis gracilis Tillyard: (75-79, female) 75, apex of abdomen, lateral; 76, praegenitale and subgenitale, ventral; 77, same and spermatheca, lateral; 78, spermatheca, ventral; 79, postgenitale and gonapophysis, lateral; (80-86, male) 80, apex of abdomen, lateral; 81, same, dorsal; 82, sternite IX, lateral, with inset of apex, posterior; 83, same, ventral; 84, genitalia, lateral; 85, gonarcus and arcessus, dorsal; 86, genitalia, posterior. (Scales in mm to Figs 75, 80.) '%u!MPu!~(56) '8ur~aloj&6) JO eqd!ll euaA JO xadv '96 Iean n3-rn '8U!~al0J'&5 :uo!leuan lu!~aloj'26 :(pnLlp~s!1!3v~8 .d 'M-26) !8U!MpU!y (16) '~u!M~~oJ(06) JO ~3!1d!.11€?USA JO xade '16 '06 :~alt!n3-pJ '%u!~alOJ'68 :UO!~EU~ABu!~pu!y '88 :UO!lz?UaA 8U!MalOJ '~8:[(U!lanf)) suv8ap S!~dOy3&d '16-L8] '56-Lg S%!S

aop!sdoy3Lsd [euxayo pue umplsnv T. R. New

Figs 96-101. Psychopsis barnardi Tillyard: 96, forewing venation; (97-101, male) 97, apex of abdomen, lateral; 98, apex of sternite IX, ventral; 99, genitalia, lateral; 100, gonarcus and arcessus, dorsal; IOI, genitalia, posterior. (Scale in mm to Fig. 97.) Australian and Oriental Psychopsidae

Figs 102-110. Psychopsis barnardi Tillyard: (102-107, female) 102, apex of abdomen, lateral; 103, praegenitale and subgenitale, lateral, with inset of subgenitale, ventral; 104, praegenitale and associated structures, ventral; 105, spermatheca, lateral; 106, same, ventral; 107, postgenitale and gonapophysis, lateral; 108, forewing, M-Cu area; 109, 110, apex of vena triplica of (109) forewing, (110) hindwing. (Scale in mm to Fig. 102.) T. R. New

Figs 111-119, Psychopsis margarita Tillyard: Ill, forewing venation; 112, forewing, M-Cu area; 113, 114, apex of vena triplica of (113) forewing, (114) hindwing; (115-119, female) 115, apex of abdomen, lateral; 116, praegenitale, subgenitale and spermatheca, lateral; 117, praegenitale and subgenitale, ventral; 118, spermatheca, ventral; 119, postgenitale and gonapophysis, lateral. (Scale in mm to Fig. 115.) Australian and Oriental Psychopsidae

Figs 120-131. Psychopsis tillyardi, sp. nov.: 120, forewing venation; 121, hindwing venation; 122, forewing, M-Cu area; 123, 124, apex of vena triplica of (123) forewing, (124) hindwing; 125, forewing pattern; (126-131, female) 126, apex of abdomen, lateral; 127, praegenitale and subgenitale, lateral; 128, same, ventral; 129, spermatheca, lateral; 130, same, ventral; 131, postgenitale and gonapophysis, lateral. (Scale in mm to Fig. 126.) T. R. New

Figs 132-141. Psychopsis dumigani Tillyard: 132, forewing venation; 133, forewing, M-Cu area; 134, apex of vena triplica, forewing; 135, forewing, pattern; (136-141, female) 136, apex of abdomen, lateral; 137, praegenitale and subgenitale, lateral; 138, same, ventral; 139, spermatheca, lateral; 140, same, antero-dorsal; 141, postgenitale and gonapophysis, lateral. (Scale in mm to Fig. 136.) Australian and Oriental Psychopsidae

Figs 142-147. Psychopsis maculipennis Tillyard: 142, forewing venation; 143, hindwing venatior?; 144, 145, apex of vena triplica of (144) forewing, (145) hindwing; 146, forewing, M-Cu area; 147, forewing, pattern. T. R. New

Figs 148-154. Psychopsis coelivagus (Walker): 148, forewing venation; 149, hindwing venation; 150, forewing, M-Cu area; 151, 152, apex of vena triplica of (151) forewing, (152) hindwing; 153, 154, examples of forewing pattern. Australian and Oriental Psychopsidae

Figs 155-166. Psychopsis coelivagus (Walker): (155-161, female) 155, apex of abdomen, lateral, with inset of apex of ectoprocts, ventral; 156, same, dorsal; 157, praegenitale and subgenitale, ventral; 158, same, lateral; 159, spermatheca, lateral; 160, same, ventral; 161, postgenitale and gonapophysis, lateral; (162-166, male) 162, apex of abdomen, lateral; 163, sternite IX, ventral; 164, genitalia, lateral; 165, same, dorsal; 166, same, posterior. (Scales in mm to Figs 155, 162.) T. R. New

Figs 167-178. Psychopsis spp.: (167-171, P. meyricki McLachlan) 167, forewing venation; 168, hindwing venation; 169, forewing, M-Cu area; 170, 171, apex of vena triplica of (170) forewing, (I 71) hindwing; (1 72-1 78, P. insolens McLachlan) 172, forewing venation; 173, hindwing venation; 174, forewing, M-Cu area; 175, 176, apex of vena triplica of (175) forewing, (176) hindwing; 177, forewing pattern; 178, hindwing pattern. Australian and Oriental Psychopsidae

Figs 179-190. Psychopsis insolens McLachlan: (179-184, female) 179, apex of abdomen, lateral; 180, praegenitale and subgenitale, ventral; 181, same, lateral; 182, spermatheca, lateral; 183, same, ventral; 184, postgenitale and gonapophysis, lateral, with inset of lower dorsal lobe seta and stylus spine; (185-190, male) 185, apex of abdomen, lateral; 186, ectoprocts and tergite IX, dorsal; 187, sternite IX, ventral; 188, genitalia, lateral; 189, same, dorsal; 190, same, posterior. (Scales in mm to Figs 179, 185.) T. R. New

Figs 191-195. Psychopsis meyricki McLachlan, male: 191, apex of abdomen, lateral; 192, same, dorsal; 193, genitalia, lateral; 194, same, dorsal; 193, genitalia, lateral; 194, same, dorsal; 195, same, posterior. (Scale in mm to Fig. 191.) Australian and Oriental Psychopsidae

Figs 196-198. Megapsychops illidgei (Froggatt): 196, forewing venation; 197, hindwing venation; 198, forewing, M-Cu area. T. R. New

Figs 199-202. Megapsychops illidgei (Froggatt): 199, 200, apex of vena triplica of (199) forewing, (200) hindwing; 201, forewing pattern; 202, hindwing pattern. Australian and Oriental Psychopsidae

Figs 203-208. Megapsychops illidgei (Froggatt), female: 203, apex of abdomen, lateral; 204, prae- genitale and subgenitale, lateral; 205, same, ventral; 206, spermatheca, lateral; 207, same, ventral; 208, postgenitale and gonapophysis, lateral. (Scale in mm to Fig. 203.) T. R. New

Figs 209-217. Megapsychops illidgei (Froggatt), male: 209, apex of abdomen, lateral; 210, same, dorsal; 211, apex of sternite IX, lateral; 212, same, ventral; 213, gonarcus, arcessus and ?gonapsis, lateral; 214, parameres, lateral; 215, arcessus, posteroventral; 216, gonarcus and entoprocessus, dorsal; 217, parameres and ?gonapsis, ventral. (Scale in mm to Fig. 209.) Australian and Oriental Psychopsidae

Figs 218-222. Explanatory diagnosis: 218, forewing [venation in standard terminology; gv, gradate veins, divided into cs (costal series), is (internal series), ds (discal series), ts (terminal series); vt, (vena triplica)]; 219, female, apex of abdomen lateral; 220, gonapophysis, lateral; 221, male, apex of abdomen, lateral; 222, male genitalic complex, lateral (219-222; VIII-IX, segment Nos; up, apodeme; ar, arcessus; bs, basal setae; cs, central setae; dl, dorsal lobe; ds, dorsal setae; ect, ectoproct; gen, genitalia; gl, gonapophyses laterales; gs, gonarcus; pa, paramere; pg, praegenitale; pop, postgenitale; s, spermatheca; sg, subgenitale; sp, superprocessus; st, sternite; sts, stylus spines; sty, stylus; t, tergite; vs, ventral setae).