i ate tert ae Aeatetotat sae ft teste ma, ss. m ee av Aa ac AP ees ‘, anes, eR Agen. Ah = * tare eles gaeees ees yi ahh St AMS “4 Rr atared ROA ne sh a le A pair of Euprosopia subula (female above) on bark of Eucalyptus sp., McCarr’s Creek, near Sydney. [Photo: D. Clyne, black and white print: A. Healy. | 77811—b THE AUSTRALIAN MUSEUM, SYDNEY MEMOIR 15 The Australian Platystomatidae (Diptera, Schizophora) with a Revision of Five Genera’ By DAVID K. McALPINE The Australian Museum, Sydney Published by Order of the Trustees of the Australian Museum Sydney, New South Wales, Australia 1972 soe a er RT eee la eng wie aR Rm *This paper contains part of the results of research carried out at the Imperial College of Science and Technology in fulfilment of the requirements for the degree of Doctor of Philosophy at the University of London. oo EE ee ee ar G 77811—1 Registered in Australia for transmission by post as a book SYDNEY: V. C. N. BLIGHT, GOVERNMENT PRINTER, 1973 Contents T Introduction (A) General we (B) Materials and Methods (CQ) Acknowledgements II Biology and Habits III Geographic Distribution IV Morphology (A) Introductory (B) The Cuticular Surface .. (C) The Head (D) The Thorax (E) The Abdomen V Relationships and Classification (A) Status and Relationships B) Subfamily Classification. . ( ; (C) List of Australian Species, with Notes ( D) Key to Australian Genera VI Genus Mesoctenia VII Genus Plagiostenopterina .. VIII Genus Lamprogaster IX Genus Duomyia X Genus Euprosopia XI List of New Names, Synonymy, and Type Designations XII References XIII Index 00° ON OV ON 132 192 ABSTRACT A general survey is made of the Australian flies of the family Platystomatidae on the basis of large collections of preserved material and type specimens in European museums. Some information is provided on the biology and habits of the group. The distribution and abundance of the family in the zoogeographic regions of the world is described and particular attention paid to the distribution of genera occurring in Australia, both within and outside the Australian continent. The external morphology of the Platystomatidae is described in some detail and the principal trends in morphological diversity are outlined. The apparent relationships of the Platystomatidae to other families of Diptera are given together with their morphological basis. The family is defined and a scheme is presented for the division of the Platystomatidae into five subfamilies. A key to the Australian genera of Platystomatidae is given and a list of the known Australian species is presented. The genera Mesoctenia, Plagiostenopterina, Lamprogaster, Duomyia, and Euprosopia are defined. Keys to the 139 recognized Australian species of these five genera are provided together with descriptions of the species. Distributional data and, when available, habitat data are recorded for each species. I. INTRODUCTION (A) General Observations The Platystomatidae are a family of two-winged flies (Diptera) found in most vegetated parts of the world with the notable exception of the temperate zone of South America. ‘The number of known world species is slightly less than 1,000. Until recently it was usual to regard the group as a subfamily of the family Otitidae (Ortalidae) but specialist opinion has now swung towards giving the Platystomatidae family rank. ‘Though clearly related to other “‘acalyptrate’’ families of Diptera, the very large squamae (calyptrae) of certain forms have led some authors to mistake them for calyptrates (e.g. Walker, 1849; Ouchi, 1939). Use of the family-group name Platystominae in Hendel’s monographic treatment (1914a, 1914b) has stabilized the name of this group (now corrected to Platystomatidae). Universality of usage since that time clearly makes substitution of any alternative name undesirable. However, Wiedemann referred to the family “‘Achiidarum’”, using the name in the genitive case, in 1830. Bigot (1852) used a family name Achiadae, and Walker (1857) also used a subfamily name Achiides before Schiner (1864) introduced the name Platystominae. Unlike the related family Tephritidae (Trypetidae) the Platystomatidae are of very slight economic importance so far as is known. However a species of Rivellia has been recorded. as causing significant damage to a pasture legume in Queensland (Diatloff, 1965). Perhaps species of Duomyia play a role in pollination of some Eucalyptus species. For the purposes of this work the limits of Australia have been interpreted in the political sense. ‘Thus I include the six Australian states and the Northern Territory, but external territories of Australia are excluded. The Torres Strait Islands are included as part of the state of Queensland and Lord Howe Island is part of the state of New South Wales. Norfolk Island is excluded as an external territory. For practical reasons I do not use political boundaries for defining New Guinea, but apply the term to the main island of New Guinea together with only the most closely adjacent small islands. Thus defined New Guinea includes three zones: West New Guinea (West Irian); North East New Guinea (the Sepik, Highland, Madang, and Morobe Districts of the ‘Trust Territory of New Guinea); and the Australian Territory of Papua. New Guinea material is only treated here when it is desirable to do so for comparative purposes. Up to the present approximately 65 valid species of Platystomatidae have been recorded as occurring in Australia. Possible synonymy among the recorded nominal species of Rivellia might reduce this number slightly. (B) Materials and Methods The total number of specimens of Australian Platystomatidae available for this work is well over 5,000. Most types of previously described species have been examined and reassessed. In the lists of material examined I have abbreviated the names of museums and collections as follows: AM Australian Museum, Sydney. BM British Museum (Natural History), London. BPB Bernice P. Bishop Museum, Honolulu, Hawaii. CNC Canadian National Collection, Entomology Research Institute, Ottawa. df CSIRO Australian National Insect Collection, Commonwealth Scientific and Industrial Research Organisation, Canberra. DEI Deutsches Entomologisches Institut, Eberswalde, East Germany. GLB Collection of Dr G. L. Bush, University of Texas, Austin. MNM _ Természettudomanyi Mizeum, Hungarian National Museum, Budapest. NMV_ National Museum of Victoria, Melbourne. NSWDA New South Wales Department of Agriculture, Rydalmere. OXN Hope Entomology Collection, University Museum, Oxford. PM Muséum National d’Histoire Naturelle, Paris. QM Queensland Museum, Brisbane. SAM South Australian Museum, Adelaide. SPHTM School of Public Health and Tropical Medicine, University of Sydney. U@) Entomology Department, University of Queensland, Saint Lucia, Brisbane. USNM_ United States National Museum, Washington, D.C. WM Naturhistorisches Museum, Vienna. ZMB Zoologisches Museum der Humboldt—Universitat, Berlin. The names of the following collectors are abbreviated to the initials: J. H. Ardley, T. L. Bancroft, E. B. Britton, A. N. Burns, G. L. Bush, T. G. Campbell, P. B. Carne, C. E. Chadwick, D. H. Colless, I. F. Common, M. F. Day, F. P. Dodd, A. L. Dyce, K. English, F. Evans, W. W. Froggatt, M. Fuller, N. Geary, M. and V. Gregg, H. Hacker, G. H. Hardy, E. J. Harris, D. E. Havenstein, R. Helms, G. A. Holloway, C. M. Kelsall, K. H. Key, A. M. Lea, Z. R. Liepa, R. Lossin, D. K. McAlpine, I. M. Mackerras, G. Monteith, A. Musgrave, A. Neboiss, K. R. Norris, S. J. Paramonov, F. A. Perkins, E. F. Riek, C. N. Smithers, A. Snell, R. Straatman, K. L. Taylor, N. B. Tindale, A. L. Tonnoir, R. E. Turner, M. S. Upton, J. P. Verreaux, A. R. Wallace, M. Wallace, F. E. Wilson, W. W. Wirth, I. C. Yeo. For study of external features, dry pinned specimens were used. The aedaegus can be examined in such specimens by relaxing over damp sand in a sealed plastic container for 2 or 3 hours, then placing a drop of 5 per cent trisodium phosphate (NagPO,) at the apex of the abdomen under the microscope and carefully extending the aedeagus. This often involves splitting tergite 5 longitudinally a little to the right of the centre. Specimens should not be relaxed for longer periods or discoloration is likely to result, particularly with species of Euprosopia. For more detailed examination of cuticular genitalic structures the abdomen is removed and immersed in cold lactic acid for several days till the muscle is cleared and the cuticle extended. Lactic acid is also an ideal medium for microscopic examination of the genitalia. Specimens are not left indefinitely in lactic acid as gradual bleaching and deterioration takes place after several months. Genitalia are stored in phials of glycerine with plastic stoppers together with label data and number corresponding to that on the pinned specimen. Measurements were made with a squared eyepiece (graticule), calibrated by comparison with a millimetre scale on the stage of the microscope. It was not found possible to make direct measurements of the long and often coiled terminal filaments of the aedeagus in this way as they could not be straightened. ‘The method used was as follows. The aedeagus was placed in lactic acid in a Petri dish and gently flattened under the weight of a small square of glass cut from a microscope slide. ‘The specimen was placed under the microscope and viewed through a squared eyepiece. An outline drawing of the distal part of the aedeagus with filaments was prepared on squared paper so that the scale was known. Measurements of the length of the filaments in the drawing were made with an opisometer and expressed as a fraction or multiple of the length of the sclerotized glans. 8 The scanning electron microscope was used for study of some cuticular structures. The technique is outlined by Hockley (1968). ‘The image obtained is remarkable for its clarity and depth of focus, the latter being far superior to that obtained with a light microscope. (C) Acknowledgements Firstly I would like to thank Mr R. G. Davies for his help and advice during the preparation of this work. Also Mr B. Cogan, Dr P. Freeman, Mr A. Pont, and Mr K. G. V. Smith of the British Museum (Natural History) for access to the collections and facilities and help offered while working at the Museum. I am indebted to Dr C. N. Smithers of the Australian Museum, Dr D. H. Colless of CSIRO Division of Entomology, Mr G. Monteith of the Entomology Department, University of Queensland, Mr A. Neboiss of the National Museum of Victoria, Mr G. F. Gross and Mr N. McFarland of the South Australian Museum for loan of the Australian material of Platystomatidae in the collections under their care. Dr G. L. Bush also made available material in his collection. I am grateful to Dr L. Tsacas and Dr L. Matile of the Muséum National d’Histoire Naturelle, Paris, Dr A. Kaltenbach of the Naturhistorisches Museum, Vienna, Dr F. Mihalyi and Dr A. Sods of the Természettudomanyi Muzéum, Budapest, Mr G. C. Steyskal and Dr C. W. Sabrosky of the U.S. National Museum, Washington, D.C., for study facilities in these museums. Dr H. Schumann made available types from the Zoologisches Museum der Humboldt-Universitat zu Berlin, and Dr G. Morge those from the Deutsches Entomologisches Institut. Many friends have discussed parts of this work or provided various forms of assistance, including Dr P. M. Barrer, Dr M. A. Bateman, Dr G. L. Bush, Mr B. Cogan, Dr D. H. Colless, Dr R. W. Crosskey, Dr W. D. Hamilton, Mr E. Osuna, Dr M. Speight, Mr G. C. Steyskal. Assistance with field work in Australia was given by Mr R. Lossin and Mr G. A. Holloway of the Australian Museum, while Mrs K. Kota carried out much of the laborious processing of material. Mr Holloway also provided invaluable notes on his field observations of Platystomatidae. Mr H. E. Goto, Dr B. Martin, and Mrs M. Culpan gave substantial assistance with work involving use of the scanning electron microscope. ‘Thanks are due to the Trustees and Director, Dr F. H. Talbot, of the Australian Museum for financing the return trip from Sydney to London. Study at museums in Continental Europe was facilitated by grants from the University of London Grants Council and the Department of Zoology, Imperial College, through Professor T. R. E. Southwood. II. BIOLOGY AND HABITS Adults Many species of Platystomatidae have characteristic resting places which may serve as rendezvous points for courtship. Tree trunks are a usual resting place for many species of Euprosopia, Loxoneuroides varipennis, Achias kurandana and Achias australis, and probably for a number of species of Lamprogaster. On the Port Hacking River to the south of Sydney Euprosopia subula may often be seen in numbers on rock surfaces either on the bank or on projecting rocks in midstream. Much further north at Finch Hatton Gorge the same species was taken on trunks of palms and trees. Perhaps local conditions determine this difference in habits, as the tree-trunks along the Port Hacking River are rather densely shaded. On the other hand rock surfaces at Finch Hatton Gorge may become very hot in the tropical sun. At Kurnell, near Sydney, E. tenuicornis and E. anostigma occur together on the main trunks of various native and introduced trees. E. filicornis shows a distinct preference for settling on the lower branches of Cupaniopsis anacardioides in the same area. At Otford, south of Sydney, Euprosopia megastigma appeared to rest mainly on foliage of trees, and despite intensive searching was never swept from herbage or observed on tree-trunks. It was also taken in numbers on wet stones at the edge of a small stream which it apparently visits briefly for drinking. Achias australis and. Achias kurandanus often rest on tree-trunks, the latter showing a preference for Ficus trees. When large numbers occur together most of the individuals may be males. On one such occasion females of A. kurandanus were eventually found by sweeping {foliage of nearby trees. In this genus, as in a number of other platystomatid genera I have observed, there is a strong preference for resting on the lower surface of the leaf. Possibly males of Achias congregate on tree-trunks where they wait for receptive females. The females only visit the tree-trunks when ready to copulate. As males can presumably copulate effectively at frequent intervals their numbers on the trunks will be much greater than those of females which need only copulate occasionally (perhaps, as in some other Diptera, only once in a lifetime) for maximum reproductive activity. Near Cardstone Lamprogaster viola was seen on tree-trunks and was also swept from low foliage. L. violacea (in New Guinea and Queensland), L. tricauda (Mary’s Creek), L. indistincta (Mulgrave River), and L. stenoparia (Kuranda and Mulgrave River) rest on lower surfaces of leaves of trees. Lasioxiria sp. at The Crater, Atherton Tableland, rests on the large leaves of Alocasia macrorrhizos (Araceae). At least four species of Lenophila (but not L. dentipes) rest on leaves of Xanthorrhoea spp., always selecting the larger or more prominent plants. While walking on a surface some platystomatids exhibit a characteristic manner of movement. The wings are extended horizontally and moved backwards and forwards with a rowing motion. At the same time the proboscis is alternately raised and extended. ‘This behaviour has been especially noted in Pogonortalis and Rivellia but is, I think, common to a number of other genera. Similar wing movements occur in several other acalyptrate families. The mode of carrying the wings at rest differs among different genera and species of Platystomatidae. In Duomyia, Microepicausta, and Rhytidortalis rugifrons the wings are folded roof-wise along the abdomen with their posterior margins uppermost and almost touching above the median line of the abdomen. In the miliaria and ventralis groups of Euprosopia the manner of holding the wings is somewhat similar, but in the tenuicornis and separata groups of the genus the wings are held horizontally, slightly spread so that their long axes form a V. In the scatophaga group the position of the wings is usually intermediate, their costal margins being held a little lower than the posterior margins which almost meet over the abdomen. In Angitula longicollis the wings are extended horizontally almost at right angles to the body much of the time. Platystomatid species which mimic Hymenoptera may have unusual habits. Xenaspis sp. in New Guinea folds its wings longitudinally, thus increasing its resemblance to a 10 vespid wasp. Achiosoma nigrifacies in New Guinea and a related species in Queensland greatly increase their resemblance to Vespidae by extending the forelegs in front of the head and waving them rapidly like the antennae of wasps. This habit is shared by the tephritid fly Phytalmia wallacet which also mimics wasps in New Guinea. Platystomatid flies are attracted to a number of substances for feeding purposes. I have seen the following species on flowers: Duomyia decora (on Leplospermum), Rhytidortalis rugifrons (on Kunzea and other plants), Rivellia sp. (on unidentified cultivated plant). Specimens of Lenophila sp. (near coerulea) have been taken on flowers of Xanthorrhoea by R. Mulder. Hennig (1945) records Platystoma on decaying fruit and I have observed Plagiostenoplerina on broken cucurbitaceous fruit. Hennig also mentions Platystoma as attracted to vinegar. KR. Lossin and I collected specimens of Lamprogaster flavipennis apparently eating sweat. Species of Huprosopia, Rivellia, Genus C, Plagiostenoplerina, and occasionally Duomyia and Lenophila have been taken on fresh mammalian faeces (cow, horse, dog, possum, man) to which they are apparently attracted for feeding. Some other genera, notably Lamprogaster and Achias, seem not to be attracted to faeces when they are plentiful in its vicinity. Rivellia and Euprosopia spp. are attracted to decaying snails and to ammonium sulphide solution. Rivellia viridulans is known to obtain honey-dew from aphides by stroking them with the forelegs in the manner of an ant (Hennig, 1945). Detailed observations of courtship or mating have been made for very few species. Piersol (1907) has made observations on Rivellia boscti and Michelmore (1928) described copulation in Plalystoma seminationis. In the former there appears to be no complex preliminary courtship and the wings of the male are vibrated during copulation (as they probably would be at other times). In both species mentioned it seems that the male feeds the female by regurgitation while mounted. Mr G. A. Holloway informs me that he has observed courtship and mating in Lenophila dentipes on the trunk of a smooth-barked Eucalyptus species. He states: “T noticed several pairs before copulation facing each other for up to five minutes, but I was not able to get too close. ‘They do not wave their wings before copulation.” ‘The author is at present making further studies of the sexual behaviour of platystomatids. Seasonal Occurrence of Adults ‘The data on seasonal occurrence is very incomplete for Australian species. Most collectors’ labels bear dates indicating summer occurrence of adults (December to March) but some genera and species appear exceptional in this respect. Most species of Huprosopia, Achias, and Lamprogaster are to be found only in the summer months and are not to be found in the same habitat at other times. In the case of Huprosopia scatophaga at Upper Allyn and Duomyia spp. at Iluka, only females were found in March. — It is probable that these either emerge later or live longer than the males. Lamprogaster hilaris has been found only from late February to May and appears to be an autumneflying species. ‘Phese, and many other forms, both in tropical and temperate latitudes are probably univoltine. Adults of Huprosopia macrotegularia have been collected in October, November, December, January, and May in the Cairns district (latitude 17° S). This suggests that the species passes through at least two generations annually. ‘The closely related /, subula is to be found only in summer at Royal National Park, near Sydney (latitude 34° 5.), where it is apparently univoltine. Adults of Pogonortalis doclea may be found in every month in coastal areas near Sydney, the species being evidently multivoltine. Species of Rivellia occur as adults in winter as well as summer in various parts of New South Wales including Mount Wilson (altitude 3,000 feet), and are presumably multivoltine. No cases are known to me of species which fly principally or exclusively in the winter months. 11 Larvae Hennig (1945, 1952) has listed the recorded habitats of platystomatid larvae. These are very diverse and include fresh, damaged, and rotting vegetable material (including roots and bulbs), human corpses, and humus soil. Steyskal (1965a) has recorded larvae of Poecilotraphera spp. from guava, sugar cane, rice and maize. Diatloff (1965) has recorded larvae of Rivellia sp. in fresh and rotting root nodules of the pasture legume Glycine javanica in Queensland, causing considerable damage. Label data from a few reared specimens provide some information on larval habitat. A specimen of Duomyia (?tomentosa) in poor condition was reared from a mandarin (Citrus) in Queensland. A reared specimen of the closely related D. pallipes from Uriarra, near Canberra, is labelled “pupa in sand.” A specimen of Euprosopia megastigma collected at Roseville near Sydney, 14th October, 1957, by Miss K. English bears the following information: “Larva found eating curl grub pupa. Oct. 24 had pupated. Nov. 29 emerged deformed.”” Two specimens of Lenophila sp. (near coerulea) from Ballandean, near Stanthorpe, were bred from a rotting grass-tree (Xanthorrhoea) by H. Jarvis. A series of Scholastes bimaculatus in the Australian Museum is labelled ‘Ex imported coconuts.” I have observed a specimen of Pogonortalis doclea newly emerged on an indoor pot plant, the soil of which had been taken a short time previously from Dee Why near Sydney. The larvae of Elassogaster sepsoides attack the egg capsules of Locusta migratoria in the Philippines (Lopez, 1934), and presumably have similar habits elsewhere in the range of this species. Mr G. A. Holloway informs me that he has observed apparent oviposition by Euprosopia tenuicornis and Lenophila dentipes at Blue Lagoon Reserve, near The Entrance, New South Wales, in March, 1969. This is mentioned here as probably indicative of larval habitat. On the shaded side of a Eucalyptus tree he found more than forty females of both these species which appeared to be ovipositing in the sap that was exuding from bectle damage under a piece of dead bark. On investigation he found dipterous larvae in the sap, but those preserved do not resemble the known larvae of Platystomatidae. Attempts at rearing were unsuccessful. Natural Enemies No predators or parasites of this family appear to be recorded previously. In New Guinea I have collected several adult platystomatids parasitized by Strepsiptera. In some cases the stylopized individuals had triungulin-stage larvae attached to the legs. The platystomatid genera affected include Brea, Mesoctenia, Naupoda, and an unnamed genus (near Scholastes). According to Riek (1970), who has examined this material, the parasites should probably be referred to the subfamily Elenchinae of the Halictophagidae. A specimen of Lamprogaster corusca taken indoors at Bayview, N.S.W., by Mr L. C. Haines is affected by the parasitic fungus Empusa. I have observed the remains of a specimen of Pogonortalis doclea in a spider’s web. Platystomatids are no doubt frequently eaten by such general predators of flying insects. Ill. GEOGRAPHIC DISTRIBUTION The family Platystomatidae is best represented in the tropical, subtropical and warm temperate areas of the Old World. In the New World the family is less well represented; in fact it is very poorly represented if the almost cosmopolitan genus Rivellia is excluded from consideration. Steyskal (1965) listed 41 species from America north of Mexico of which 34 are species of Rivellia. In 1968 he recorded 25 species from America south of the United States, of which 11 belong to Rivellia. No species are known from America south of the tropics. Africa is the continent with the greatest number of known platystomatid species, the approximate number of described species being 240 (calculations based on figure for Otitidae s.l. given by Steyskal, 1960). ‘The Oriental Region (excluding the Papuan Subregion) appears also to have a large representation of Platystomatidae but this is less well known than the Ethiopian fauna. The Palearctic Region on the other hand has a very restricted platystomatid fauna. Despite the richness of the adjacent Ethiopian and Oriental faunas, these appear to have produced only a few temperate-adapted offshoots to the north. Platystoma is the only endemic Palearctic genus and Rivellia is the only other generally distributed genus in the region. ‘The Oriental-Australian genus Euprosopia has several species in the eastern Palearctic. The approximate number of valid described species recorded from New Guinea (excluding the Moluccas and Bismarck Archipelago but including Waigeo, Key, and Aru Islands) is 126, most of which are mentioned by Malloch (1939a). ‘This number is nearly twice as great as the number of valid described species previously recorded from Australia and. numbers of undescribed species have been seen from New Guinea. It is therefore evident that New Guinea has a greater concentration of platystomatid species than any other part of the world of similar area. The Papuan Subregion is also interesting for the number of characteristic Papuan genera which at most extend beyond New Guinea only to a few nearby island groups or to North Queensland. Some of these genera are of very bizarre form, e.g. Angitula, Laglaizia, Achiosoma, Asyntona. There is a relatively small Oceanic element in the Platystomatidae. ‘The closely related genera Giraffomyia and Angituloides form a Melanesian component, related to the Papuan Angitula. 'Vhey are accompanied by the more widespread Scholastes in this part of the Pacific. The genus Pseudorichardia occurs only on oceanic islands and is widely distributed in the Pacific. It appears to have no closely related forms on any major land mass. New Zealand has only two known species of Platystomatidae both belonging to the endemic genus ealandortalis, which is perhaps related to the widely distributed Rivellia (Harrison, 1959). In Australia there are 23 known genera and 212 known species (including described and undescribed forms). It is probable that further genera will be found within Australian limits in the more northerly of the ‘Torres Strait Islands, some of which are very close to the New Guinea coast. ‘The genera occurring in Australia may be divided into 6 groups according to their world distribution patterns. (1) Almost cosmopolitan genus: Rivellia. (2) Widely distributed Old World tropical genera: Pterogenia, Naupoda, Elassogaster, Plagiostenopterina. (3) Oriental—Australian genera: Rhytidortalis, Pogonortalis, Euprosopia. (4) Papuan—Australian genera: Microepicausta, Lamprogaster. 13 (5) Papuan genera of restricted Australian occurrence: Lasioxiria, Asyntona, Mesoctenia, Brea, Achias, Achiosoma, genus A. (6) Endemic Australian genera: Lenophila, (?) Scotinosoma, Loxoneuroides, Duomyia, genus B, genus C. Several of the genera occurring in Australia are restricted to the tropical north-east coast of Queensland. Most of these genera are to be classed as primarily Papuan, because of their better representation in that subregion. They are all inhabitants of rain forest and riverine forest (see list of Papuan genera above). The genera Plagiostenopterina, Elassogaster, and Pterogenia, though mainly occurring in the wet tropical zone of Queensland have marked extensions outside this zone in Australia, and also have extensive distributions outside Australia. A possibly endemic Australian genus, which I have designated genus B in the generic key, has only been collected in tropical rain forest in Queensland. The endemic genus Loxoneuroides is a tropical and subtropical rain forest form of coastal eastern Australia, and Scotinosoma is perhaps confined to the same areas. An endemic unnamed genus (‘“‘genus C”) is known only from the temperate rain forests of central eastern New South Wales. The genus Rivellia is very widely distributed over the Australian continent but occurs mainly in forested areas. It is probably absent from arid parts. Lamprogaster has a similar distribution to Rivellia, occurring mainly in forests and woodlands, but is more restricted within this general habitat. One species occurs in low rainfall areas of western New South Wales, perhaps only in the vicinity of rivers. _Duomyia also occupies much the same areas as Rivellia but is known to occur in arid areas as well. It is the only genus of the family known from Central Australia and occurs in all parts of Australia where platystomatids have been collected. Euprosopia is represented mainly in Eastern Australia but one species extends west to Darwin in the north and another to Adelaide in the south. Together with Lamprogaster, Duomyia, and Rivellia, it is represented at higher altitudes in the Australian Alps. A specimen of Euprosopia alpina n.sp. has been collected at an altitude of 7,300 feet near the summit of Australia’s highest mountain. Pogonortalis occurs throughout coastal eastern Australia and also in the southwest. It is the only genus which is common in urban gardens in the eastern states. The Australian species P. doclea (Walker) has been introduced into California (Steyskal, 1961). A group of species is restricted to the vicinity of coastal sand dunes. Here it would appear to be edaphic conditions rather than climate which determine distribution. Some species of this group are threatened by habitat destruction through rutile mining in eastern Australia. The components are Microepicausta, the cribrata group of Rhytidortalis, and certain species of Duomyia and Rivellia. The pale coloration of some of these forms is perhaps an adaptation to the sandy environment. The temperate heathlands and xeromorphic scrubs of southern Australia have some characteristic forms. The genus Lenophila is more or less restricted to this habitat, as is possibly the rugifrons group of Rhytidortalis. The fauna of Lord Howe Island, 350 miles east of the coast of northern New South Wales in the warm temperate zone, is especially interesting as the island is considered to have been isolated from other land masses since the beginning of the Tertiary. The five genera of Platystomatidae, each represented by a single known species on the island, have almost certainly reached there by dispersal across the ocean. Pogonortalis howei Paramonov and Duomyia howensis n.sp. though endemic to Lord Howe Island are each very closely related to mainland Australian species. Naupoda insularis Paramonov is less closely related to the only other Australian species of its genus. Naupoda is also represented in Fiji and elsewhere in the Pacific. It evidently has considerable powers of dispersal. The genus which I have referred to as genus A in the generic key is known from a single specimen from Lord Howe Island, and a quite distinct undescribed species in New Guinea. A possible explanation for these isolated occurrences is the extinction of the group elsewhere through increased competition or predation. Ifa formerly 14 widespread form crossed the sea to become established on a remote island, its chances of escaping these pressures would be increased. The status of the Lord Howe Island form of Rivellia has not yet been investigated. As stated above the genus is well represented on the Australian mainland. With the exception of ten species mentioned below all the Australian species appear to be endemic. Asyntona tetyroides (Walker), Naupoda regina Hendel, Lamprogaster violacea (Macquart), L. macrocephala Hendel, Euprosopia ventralis (Walker) and E. xanthops n.sp. extend to New Guinea and a few of the adjacent islands. Llassogaster sepsoides (Walker), Plagiostenopterina aenea (Wiedemann), and P. enderleini Hendel have rather wide distributions, all extending to South East Asia. Pogonortalis doclea (Walker) is perhaps naturally endemic to Australia but has been recently introduced into California as mentioned above, and there is a record from Java (Hendel, 1914b). 15 IV. MORPHOLOGY (A) Introductory The structure of the higher Diptera (Series Schizophora) is in many respects very uniform. For this reason it is not the aim of this chapter to provide a comprehensive description of platystomatid structure. Instead an attempt is made to give a general review of outstanding characters of the Platystomatidae and to provide a basis for the descriptions of morphological diversity presented in the evolutionary and systematic studies. This applies as much to the studies of the cuticle and its processes made with the scanning electron microscope as to the classical methods of study. (B) The Cuticular Surface The visual impression of the surface texture of the cuticle has been much used as a taxonomic character in higher Diptera. Unfortunately there has been little attempt to define accurately the surfaces described as “pruinose”, “dusted”, “pubescent”, etc. The present study has convinced the author that it is important to discriminate between kinds of surface texture for taxonomic purposes. The use of the scanning electron microscope (SEM) shows great promise in enabling the often rather vague visual impressions of surface texture to be more accurately defined. This approach has been only touched on here. The cuticle, which covers the entire surface of the insect, is divisible into two main types—sclerotized and membranous, sometimes erroneously referred to as chitinized and non- chitinized. The former type, which is hardened by deposition of scleroproteins, is again divisible into catogories, according to the nature of its vestiture. There is also some diversification of the vestiture of membranous cuticle. The simplest type of surface for sclerotized cuticle is smooth, without processes of any kind. When present in sufficiently large areas smooth cuticle presents a strongly shining appearance (regular reflection). Even where there is a dense vestiture on the cuticle, the spaces between the processes often appear almost smooth, with only slight irregularities under high magnification (fig. 97). That this is a fairly general condition in the higher Diptera is seen from the fact that initially dull cuticle frequently becomes shining when the vestiture is removed by rubbing or abrasion. This may happen under natural conditions when movement against the abdomen causes the posterior surfaces of the hind femora to become shiny, or when parts of the female abdomen are caressed by the male during courtship. It also happens when captured specimens are carelessly handled before mounting. Cuticle which is devoid of vestiture may have its shining quality reduced or interrupted by fine sculpturing. This sculpturing generally takes the form of irregular pits, rugosity, or scratching, but may form a very regular grooving reminiscent of the convolutions of the skin on human fingertips. The latter type of grooving often occurs on the check region of the head in Platystomatidae. The dull, lustreless appearance of pruinescent cuticle is sometimes strengthened by sculpturing. Two broad categories of hair-like cuticular processes are generally recognized, viz. microtrichia and macrotrichia. Microtrichia are superficial extensions of the cuticle, not individually connected with the subcuticular cells of the epidermis. ‘They are immobile, being without a basal socket. Each macrotrichium arises from a specialized hypodermal trichogen cell and is often associated with a nerve cell. It is surrounded at its base by a membranous ring or socket enabling movement. A further difference between microtrichia and macrotrichia is in size. In the Platystomatidae microtrichia are generally less than gop. 16 in length and macrotrichia are of greater length, but there may be slight overlap in size and this limit varies with the size of the insect. Many of the specialized sensory setae, though structurally allied to macrotrichia, are within the size range for microtrichia. Microtrichia are of general occurrence on the wing surface where they are widely spaced and may be readily appreciated under the light microscope. Pruinescence is a general term for vestiture which appears as a close, dust-like or powder- like covering on the cuticle more or less obscuring the surface shine under the light microscope. As the true structure of pruinescence is not readily discernible under the light microscope it is possible that more than one kind of vestiture is included under the term. Under the SEM pruinescence on the thoracic pleura and abdominal tergites of Euprosopia spp. is seen to consist of very short, curved, hair-like microtrichia (figs 98, 108). It is possible in other cases that the microtrichia constituting the pruinescence are scale-like, but this is yet to be demonstrated. Microtrichia which are sufficiently long to be distinguishable at a magnification of x 100 to x 150 under the light microscope are termed pubescence. Pubescence is often seen to intergrade gradually into pruinescence in certain areas. In the genus Huprosopia a specialized kind of vestiture occurs on the fifth abdominal tergite of the female in a number of species. This consists of exceedingly fine microtrichia which, at a magnification of X 50 under the light microscope appear as a translucent covering above the surface of the cuticle. I have termed this velvety pubescence. Under the SEM this appears very similar to pruinescence but the microtrichia are much finer and more erect (figs 95, 96). Macrotrichia may occur as a sparser over-storey on cuticle clothed with microtrichia, or they may occur on smooth or sculptured sclerotized cuticle. More rarely they are seen on membranous cuticle. ‘They may occur on certain wing veins or, in a very few instances, on the wing membrane. The majority of macrotrichia of the body surface, including the legs, may be referred to as either hairs or setulae according to thickness and pigmentation, the setulae being thicker and often darker than the hairs. ‘The division is arbitrary and not much importance is attached to it for taxonomic purposes, though sometimes the colour of the macrotrichia is a useful taxonomic character. Macrotrichia on some areas of the body may be broadened into flat scales of various shapes. On the surface of the cuticle there is a much smaller number of greatly enlarged macrotrichia called bristles or setae. ‘These are arranged according to a definite pattern so that each bristle or linear series of bristles can be designated by a name. ‘The bristle arrangement or chaetotaxy of the Platystomatidae is dealt with below in the sections on the head, thorax and legs. Modified bristles may occur in certain parts or in a particular sex. Spatulate bristles may occur in the males of a few species, but not in the females. Bristles on the ventral surface of the femur are sometimes thickened into spines. Certain apical bristles of the tibiae are particularly strongly developed and mobile. These are called spurs. In addition to the microtrichia and macrotrichia there are various kinds of immovable processes or excrescences of the cuticle which have no basal socket, but have a sclerotized covering continuous with that of the surrounding surface. These are dealt with under appropriate sections of this chapter or, in the case of more restricted types, under the individual species. (C) The Head The head capsule is characterized by the loss of most of the boundaries defining the principal head sclerites of more primitive insects. For this reason a modified nomenclature for the regions of the head is necessary, though a few of the general terms are retained with a modified meaning. At the summit of the head there is a transverse ridge, often much rounded 17 off, connecting the eyes. This is termed the vertex. It usually bears two pairs of strong bristles, the inner and outer vertical bristles. Ummediately behind the centre of the vertex there is often a pair of much shorter closely placed divergent postvertical bristles. The area between the vertex and the bases of the antennae, bounded laterally by the eyes, is the postfrons (or simply frons of many authors). The anterior marginal region, divided off by the ptilinal fissure, is the frontal lunule. It often forms a more or less horizontal ledge above the bases of the antennae. ‘There are three closely grouped ocelli on the posterior median part of the postfrons (absent in the African genus Bromophila). From between the ocelli there often arises a pair of ocellar bristles. ‘The one or two pairs of fronto-orbital bristles when present are situated on the upper lateral parts of the postfrons, not far from the eyes. The anterior surface of the head below the postfrons is the face or prefrons. The lateral areas, between the downwardly produced lateral arms of the ptilinal fissure and the eye on each side, are called the parafacials. Lying immediately inside the parafacial is usually an antennal groove into which the antenna may be folded. | Often, especially in the subfamily Platystomatinae, the median area of the face between the antennal grooves forms an elevated ridge or platform, the facial carina. When well-developed this may be flat topped with sharp lateral margins, and is narrowed towards the upper exiremity where it passes between the bases of the antennae. At its lower extremity the sclerotization of the face ceases abruptly on the transverse epistomal margin. The lower lateral angles of the face may be produced laterally beneath the cheeks from which they are separated by the laterofacial sutures which continue downwards and outwards from the outer margins of the antennal grooves. ‘I'he shape of these lateral extensions of the face has proved a helpful taxonomic character for separating some species of Euprosopia. The cheek is the lateral area of the head capsule lying below the eye. ‘The height of the cheek, a measurement used in the descriptions, is the vertical distance between a horizontal line passing through the lowest point of the eye margin and a similar line passing through the lowest point of the cheek. The cheek has usually a fairly strong downwardly directed cheek bristle differentiated from the surrounding hairs. ‘The occiput is the term for the entire posterior surface of the head capsule. In the Platystomatidae, and particularly in many of the Platystomatinae, the lower part of the occiput is often swollen or convex whereas the upper part, immediately below the vertex, is flat or slightly concave. The ventral surface of the head is the membranous subcranial region supporting the mouthparts. It is sometimes referred to as the subcranial cavity, but when the mouthparts are extended it ceases to be a cavity. Anteriorly the subcranial membrane supports a horseshoe-shaped or lip-like sclerite, the prelabrum. It has been erroneously called labrum, but may be homologous with the clypeus. The mouthparts are very similar to those of such better known forms as Musca and Calliphora. The maxillae are reduced to the large, unsegmented palpi attached to the membranous basal part of the proboscis. The distal part of the proboscis has on its anterior surface an oral groove leading to the mouth opening near its base. The oral groove and mouth opening are concealed by the elongate labrum which is attached just in front of the mouth, The posterior surface of the distal part of the proboscis is covered by a sclerite, the theca, always broadly developed in the Platystomatidae. At the terminal end of the proboscis there is a pair of broad semicircular labella, armed with numerous pseudotracheae which radiate from the end of the oral groove. (D) The Thorax (fig. 1) The prothorax in most of the Platystomatidae is of the greatly reduced kind generally met with in the Schizophora. In a very few genera, notably Angitula and Giraffomyia, the prothorax is prolonged forming with the cervical region a prominent neck (Speight, 1969). The prosternum, or more correctly the basisternum of the prothorax varies considerably in shape. In 18 the Trapherinae it is very broad, with broadly rounded lateral lobes. In the Scholastinae and Platystomatinae it is generally more or less quadrate and the anterior angles may be connected to the propleura by means of variably sclerotized precoxal bridges. Speight (1969) records the shape of the prosternum in many platystomatid genera. The humeral calli, a pair of prominent convexities on the anterolateral angles of the dorsal surface of the thorax, are usually considered to belong to the pronotum. Frequently a humeral bristle is borne on each humeral callus. The enlarged mesothorax of the higher Diptera is so highly modified that it is almost impossible to apply terms to most of the parts which indicate exact homologies with the sclerites of other insects. The greater part of the dorsal surface of the thorax forms the mesoscutum, often referred to as the mesonotum, but more correctly the latter term includes also the scutellum. Laterally, immediately behind the humeral callus and in front of the wing base, is the notopleural area which is not sharply defined in the Platystomatidae. It usually bears an anterior and a posterior notopleural bristle, the latter surmounting a tubercle, the posterior notopleural callus. ‘The pair of notopleural areas is considered to represent the mesothoracic proscutum by Snodgrass (1935). From the posterior end of each notopleural area a transverse suture runs for a short distance in a mesal direction on the mesoscutum. ‘The bristles of the mesoscutum (see fig. 1) include the following: supra-alar, postalar, intra-alar, dorsocentral, and prescutellar acrostichal. All these bristles are paired. At its posterior extremity the mesoscutum is separated from the scutellum by a deep secondary scutellar suture. The scutellum forms a posteriorly projecting platform, its dorsal surface almost level with that of mesoscutum. At the free edges it bears a number of scutellar bristles. The scutellar bridges are a pair of strongly raised ridges, one from each upper basal angle of the scutellum running anteriorly to the mesoscutum and bridging the scutellar suture. A longer ridge on each side of the thorax runs from the lower lateral angle of the scutellum to the wing base. This is termed the suprasquamal ridge by Malloch (1929). The presence of hairs on this ridge is sometimes a useful taxonomic character. Below the posterior edge of the scutellum there is usually a transverse convexity, the postscutellum. Below the postscutellum the postnotwum drops away almost vertically to the base of the abdomen. The pleura of the mesothorax are made up of the following sclerites: mesopleuron, pteropleuron, pleurotergite, and sternopleuron. This application of the term mesopleuron to a sclerite which is only part of the true mesopleuron is unfortunate but well established and no very suitable isin use. ‘The mesopleuron, pteropleuron, and sternopleuron may each have a distinct bristle, known as the mesopleural, pteropleural, and sternopleural bristles respectively. In many platystomatids some or all of these bristles are absent. Posteriorly the mesopleuron is limited by a broad membranous cleft, which is a modified suture. Normally this terminates where it meets the upper margin of the sternopleuron, but in Giraffomyia and Angituloides it penetrates deeply into the region of the sternopleuron. The basalare is a small sclerite bordering the posterior side of the upper extremity of the membranous cleft. In many species of Euprosopia, more particularly in the females, it bears a basalar process which projects upwards or forwards. The metathorax, apart from its appendages, is represented only by the pleura and sternum. The pleura tend to become continuous with the meron of the middle coxa to form the hypopleuron. In Angitula, Angituloides, and Giraffomyia the posterior surface of the metathorax between the bases of the hind coxae and the first abdominal sternite is extensively and heavily sclerotized to form a broad postcoxal bridge which is quite continuous with the hypopleuron on each side. A much less marked development of the postcoxal bridge occurs in some species of Achiosoma. It is present also in some other elongate acalyptrates e.g. in the families ‘Tephritidae (Phytalmia, Diplochorda, Dacus, Soita, Adrama), Micropezidae (Crosa), Megamerinidae, Gobryidae, Diopsidae, etc. ‘The degs of Platystomatidae vary much in length and thickness, and in the armature of macrotrichia which may or may not include distinct series of bristles. Apart from these superficial differences the structure of the legs is uniform throughout the family and similar to that of other higher Diptera, except for certain specific modifications more or less confined to the male sex. Most of these are mentioned in the specific descriptions. One character worth 19 mentioning here is the presence of the mid-coxal prong in all Platystomatidae, because this prong is absent in many species of the related family Pyrgotidae. The terms anterior, posterior, dorsal, and ventral, as applied to the surfaces of the legs, indicate the position of those surfaces when the leg is completely extended horizontally at right angles to the longitudinal axis of the body. As a further guide, the knee-joint between femur and tibia is taken as moving in a transverse plane dividing anterior from posterior, and the tarsus is considered to be dorsoventrally compressed. The wing-base includes several structures of taxonomic importance. The tegula is a normally scale-like process, armed with macrotrichia, which overlaps the anterior surface of the basal extremity of the wing. It is much modified in the females of certain species of Euprosopia. The axillary lobe is a membranous elastic lobe which forms an extension of the wing membrane between the posterior basal part of the wing and the body of the thorax. In most Diptera this extends back along the surface of the thorax to the lower lateral angle of the scutellum as a narrow fringe. In certain groups, notably the Tabanidae and the “‘calyptrate”’ families, this fringe is broadened into a conspicuous lobe, the squama or calyptra which may conceal the haltere from above. (Calypira is the form of the word in both Latin and Greek and is preferable to the forms calypter and calyptron sometimes used.). In many of the subfamily Platystomatinae and perhaps all of the subfamily Scholastinae the squama is a well-developed and often conspicuous lobe. This is a most unusual character among the “‘acalyptrate” families. The wing venation of the Platystomatidae is of the type characteristic of the Schizophora asa whole. The nomenclature of the veins is that employed by Harrison (1959: fig. 437) and Munro (1947: t. fig. 13) except for the following points: the anal vein of Harrison is here termed vein 6; the upper crossvein of Munro is here termed anterior crossvein; the posterior crossvein of Harrison, lower crossvein of Munro, is here termed the discal crossvein; the vein separating the second basal and discal cells is here termed the basal crossvein; the vein closing the distal end of the anal cell is here termed the anal crossvein. ‘The nomenclature of the cells here used is also that of Harrison and Munro except that the cell between the subcosta and vein 1 is here termed the subcostal cell. The costal vein in Platystomatidae is weakened or incised about halfway between the humeral crossvein and its basal extremity and again more distinctly a short distance beyond the humeral crossvein. The costa is not incised or broken at the end of the subcosta, a point which distinguishes the Platystomatidae from the Tephritidae. In most families of Tephritoidea the anal cell has a characteristic shape, the anal crossvein being indented and the outer posterior angle of the cell being produced into an acute lobe. In comparatively few of the species of Tephritidae, Otitidae, and Pyrgotidae, but in all species of Platystomatidae, the anal crossvein is not indented and the anal cell has no acute lobe. In a few platystomatid genera (Asyntona, Clitodoca, Giraffomyia, and Loxonevra) the anal cell is acute, but not lobed. Macrotrichia or setulae occur on the upper surfaces of veins 1 and 3 in all platystomatids that have been studied. In some forms there are also setulae on vein 5, most frequently confined to the basal section of that vein bordering the anal cell. The setulae on vein 1 generally extend from just beyond the level of the humeral crossvein to the distal extremity of the vein. In the genera Euprosopia and Pterogenia there is a group of setulae on the basal part of vein 1 (radial vein) before the level of the humeral crossvein. Similar variation in the extent of the setulae on vein 1 occurs within the family Tephritidae. In the family Otitidae the setulae on vein 1, when present, are usually more restricted to the distal part of the vein. In order to facilitate description I have introduced a system of nomenclature for the bands of pigmentation on the wing, which is shown in fig. 50. ‘This system was originally devised for Euprosopia species but its use has been extended to the genus Lamprogaster, and it could also be employed for description of Rivellia species. The metathoracic haltere shows no unusual characters in the Platystomatidae, at least under moderate magnification. For descriptive purposes it is divided into three parts, a swollen basal scabellum, an elongate stalk-like pedicel, and an expanded apical capitellum. 20 (E) The Abdomen The term preabdomen is applied to the relatively unmodified anterior segments of the abdomen. In the males of Tephritoidea the preabdomen consists of segments 1 to 5. Usually in females of Tephritoidea the preabdomen may be considered to consist of segments 1 to 6, but in the Platystomatidae segment 6 of the female is always reduced in size and in most genera its tergite is concealed or absent. Hence it is convenient to regard the first five segments as constituting the preabdomen as in the male. The first two tergites of the abdomen are invariably fused together, though a faint transverse line appears to indicate where fusion has taken place. In the males of some species of Euprosopia, Lamprogaster, Platystoma, and other genera of Platystomatinae tergite 5 is enlarged at the expense of the preceding tergites, apparently in connection with accommodating the genital pouch (see below). By contrast, in the subfamily Scholastinae tergite 5 of the female is always reduced in size or absent and tergite 4 may also be reduced or absent. Thus in the genera Mesoctenia, Naupoda, Neohemgaster, Scholastes, and Trigonosoma the female preabdomen is covered dorsally by two large sclerites only, viz. tergite 1 + 2 and tergite 3. The preabdominal sternites are much narrower than the tergites and are sometimes reduced in size and number. The pleural membrane between the tergites and sternites is well-developed and much of its surface faces ventrally because the narrowing of the sternites draws it on to the ventral surface. It may bear hairs or setulae, or, more exceptionally, scales. Normally there is a pair of spiracles for each of the five preabdominal segments, situated in the pleural membrane close to the lateral margins of its segmental tergite. In the females of many species of Scholastinae and Platystomatinae the spiracles of segment 5 and to a lesser extent those of segment 4 are displaced dorsally into the intersegmental membrane behind the tergite of their segment, and in extreme cases those of the fifth pair may be closely approximated on each side of the median dorsal line (figs 74, 82). More rarely the spiracles of segment 3 are similarly displaced. The abdominal spiracles vary also in the size and thickness of the sclerotized ring surrounding the opening and in the development of microtrichia on the ring. In many genera of Platystomatinae the microtrichia form a dense conical tuft (fig. 100). In Giraffomyia and Angituloides, unlike Angitula, there are no external microtrichia and the sclerotized ring is very thick in relation to its central aperture. In the Scholastinae the tuft of microtrichia is moderately developed. In Loxoneuroides there are no external microtrichia but on the inner surface of the ring there are numerous radially arranged microtrichia pointing towards the centre of the aperture. The male postabdomen (figs 2, 3), consisting of all the segments behind segment 5, may again be divided into the more anterior part, the protandrium (segments 6 to 8) and the posterior part or andrium including the genital and anal segments. The protandrium is without tergites or spiracles. Its sclerotized parts thus consist entirely of sternites which have become highly asymmetrical and spirally displaced in a manner that has only been adequately explained by Crampton (1942). The displacement results from the rotation of the genital segment on its longitudinal axis through 360°, and consequent torsion of the immediately preceding segments. On the right side and on the dorsal surface of the protandrium there is a membranous area which is more or less infolded beneath tergite 5 to form a genital pouch containing the distal part of the aedeagus. In those Platystomatinae in which the aedeagus has very long terminal filaments (see below) the genital pouch may be very deep and extend into segment 4 on the right side (fig. 2). The andrium is covered on most of its dorsal and lateral surface by tergite 9, often called the epandrium or, in Drosophilidae, the genital arch. ‘Tergite 9 bears two pairs of rigid processes on its lateral margins, the surstyli (also called paralobes). ‘The inner and outer surstyli of each side are closely united in their basal parts by membranous cuticle, their distal ends projecting freely. It is thus possible to distinguish a fused basal section and a free distal section in each surstylus. At its base the outer surstylus is sclerotically continuous with the surface Pel of tergite 9. The basal end of the inner surstylus either ends freely in the membranous part of segment 9 or is joined to a ventral sclerite of doubtful origin. ‘The distal end of the inner surstylus generally bears two short, strong teeth, the prensisetae of Munro (1947). Ventrally sternite 9 (the Aypandrium) extends anteriorly far beyond the anterior limit of tergite 9. It has no external appendages other than the aedeagus but there is usually a gibbosity on each side. Most of its surface is descloritized, but there is a sclerotized arm on each side, the two arms being joined posteriorly to form the genital ring or Gabelplatte of German authors. The intromittent organ or aedeagus emerges from near the posterior end of sternite 9. Its basal section, here termed the stipe (phallotheca of Munro, 1947), is a very long more or less looped tube which, in the Trapherinae (Steyskal, 1965a), constitutes virtually all of the aedeagus, as is also the case in the Otitidae. In the Platystomatidae, as in the Tephritidae and Pyrgotidae, and stipe is not spirally coiled as it is in the Otitidae, but from its origin passes forwards, downwards and upwards in a loop on the right side beneath tergite 5 to the dorsal surface of the protandrium when not in use (fig. 2). The distal end of the stipe may be differentiated into a usually short but highly variable segment, the preglans. In all Platystomatidae that have been examined, except the Trapherinae, the aedeagus has a variably sclerotized capsule, the glans, distal to the stipe. It is unfortunate that Munro (1947) restricts the term aedeagus to the glans. Because his usage is somewhat unusual, I have avoided many of Munro’s terms in this work. It is always very heavily sclerotized in the Platystomatinae except for a membranous window in some forms. In the Plastotephritinae and Scholastinae its structure may be very complex. In repose the glans usually lies on the dorsal surface of the protandrium or in the genital pouch. A membranous bulb is often present at the distal end of the glans and in the genus Duomyia usually has a pair of membranous caeca arising from its base. In the subfamily Platystomatinae, the aedeagus has a pair of hollow terminal filaments (three filaments in the violacea group of Lamprogaster). ‘The filaments may arise from the membranous bulb, or when the bulb is absent, they arise directly from the glans. The bases of the filaments may be enclosed in a sclerotized tunic. The gonopores are situated at the apices of the filaments. When the filaments are very long they may be coiled like a watch spring inside the genital pouch which is enlarged to receive them. ‘The basal end of the aedeagus connects with the posterior end of the internal rod-like aedeagal apodeme or phallapodeme. ‘The latter has a pair of lateral arms near the middle which fit into the inner surface of the lateral gibbosities of sternite 9, on which the apodeme pivots. The anterior end of the aedeagal apodeme is compressed in a sagittal plane and has muscles attached to it. An aedeagal apodeme of this kind is referred to as fultelliform, and is characteristic of the families Tephritidae, Pyrgotidae, and Platystomatidae, but not, according to Steyskal (1961), of the Otitidae and Richardiidae. The ejaculatory duct connects the base of the aedeagus to the sperm pump which occupies a median position in the posterior part of the abdominal cavity. The sperm pump has been best described by Drew (1969) for the tephritid Dacus (Strumeta) tryoni (Froggatt) in which it appears to be very similar to that of Platystomatidae. The ejaculatory duct connects to the gaculatory sac of the sperm pump. The posterior wall of the sac is often sclerotized to form an ejaculatory plate which, in the Platystomatinae has generally a pair of sclerotized convexities or tubercles. ‘The anterior part of the sperm pump is the large ejaculatory apodeme which is strongly compressed in a saggital plane and is often fan-like in shape. It is generously furnished with muscles. Drew has shown that in Dacus tryoni the ejaculatory apodeme varies greatly with the age of adult flies and recommends care in its use as a taxonomic character. ‘The only sclerotized parts behind the epandrium are the large paired unsegmented cerc? which are often partly fused or joined by a membrane. The proctiger, bearing the anus, is completely membranous and dorsal to the cerci. As previously stated segment 6 may be regarded as the first segment of the female postabdomen. In a few forms (Apiola, Giraffomyia, Angituloides, and the subfamily Trapherinae) segment 6 could perhaps be better associated with the preabdomen as its tergite is largely exposed and closely associated with tergite 5, though it is much shorter than the latter. In all other forms examined tergite 6 is very small and more or less hidden in a depression behind tergite 5 or it is completely absent. ‘The spiracles of segment 6 appear often to be present in Pup) these forms. Segment 7 forms a broadly flattened sclerotized ovipositor sheath. The last pair of abdominal spiracles is situated on this segment. ‘The terminal segments, forming the telescopic ovipositor are capable of being completely withdrawn into segment 7. Segment 8 has no tergite or sternite but has three pairs of longitudinal sclerotized strips. Its whole cuticular surface is highly flexible, as it is capable of being totally invaginated into segment 7. The aculeus is the terminal structure of the ovipositor complex, and appears to consist of the completely fused sclerites of segment 9, and perhaps segment 10, together with the cerci. In Xiria, Lule, and some other forms the ovipositor is very like that of many frugivorous Tephritidae, being apparently adapted for piercing or cutting. Segment 8 has decussate rows of minute file-like teeth and the aculeus is very broad and blade-like with acute apex. In the Scholastinae and Platystomatinae there are generally no file-like teeth on segment 8 and the aculeus is quite slender with obtuse apex. 23 V. RELATIONSHIPS AND CLASSIFICATION (A) Status and Relationships That the components of the family Platystomatidae are closely related to the otitid- tephritid group of families has long been accepted. Several important and consistent characters support this view particularly the following: preapical tibial bristles absent; subcosta always well separated from vein 1, never much reduced; vein 1 setulose dorsally ; anal and second basal cells complete; in male the absence of tergite 6 and the elongate, looped aedeagus; in the female the fusion of the terminal abdominal parts into an ovipositor which can be withdrawn into the enlarged, sheath-like segment 7. Hennig (1958) and Steyskal (1961) have done much to clarify the limits of this group of families. Hennig calls the group the superfamily Otitoidea; Steyskal calls it the Tephritoidea giving it the same limits. The name Tephritoidea is here preferred as it is based on an older family-group name. Hennig (1940a, 1945) is responsible for setting the currently accepted limits of the Platystomatidae. Steyskal (1961) has better defined the families of Tephritoidea than any previous worker though his views do not appear to differ widely from those of Hennig. The families of Tephritoidea accepted by Steyskal are Richardiidae, Otitidae, Platystomatidae, Pyrgotidae, Tachiniscidae, and Tephritidae (= Trypetidae). Two other groups which have often been given family rank are the Pterocallidae and Ulidiidae, but Steyskal has convincingly demonstrated that these should be included in the Otitidae. It is now proposed to compare the Platystomatidae with each of the other tephritoid families in order to ascertain their relationships within the superfamily. The Richardiidae are a family of purely American flies which share with the Platystomatidae the absence of an acute lobe to the anal cell. This lobe is present in the greater number of forms belonging to the other tephritoid families. Despite this fact the Richardiidae and Platystomatidae seem to be further apart morphologically and perhaps phylogenetically than any of the other families in the superfamily. In Richardiidae the costa is broken near the end of the subcosta but not broken just beyond humeral crossvein, vein 1 is devoid of setulae, the aedeagal apodeme is reduced or absent, and the aedeagus is not of the usual platystomatid type. All these characters contrast strongly not only with the Platystomatidae but with other families that appear to have greater claim to relationship with the latter. The Otitidae (formerly called Ortalidae) are the family with which the Platystomatidae have been most frequently associated. Very often, as in all the writings of Malloch and the earlier writings of Steyskal, the Platystomatidae have been considered to be a subfamily of the Otitidae. Steyskal (1961), however, has shown that the Otitidae differ from the Platystomatidae (and also from the Tephritidae and Pyrgotidae) in some significant characters which suggest that they belong to a phylogenetic branch separate from that which gave rise to these three latter families. In the Otitidae the aedeagal apodeme is simple or Y-shaped, while in the Tephritidae, Pyrgotidae, and Platystomatidae it is, so far as known, always fultelliform. Again the apex of the aedeagus in the Otitidae lacks the sclerotized apical capsule or glans of the Platystomatidae and most Tephritidae. On this point the Pyrogotidae are too poorly known for comparison. Subsequently Steyskal showed that in the platystomatid subfamily Trapherinae the glans is absent. Despite this fact there appears to be a fairly consistent difference between the aedeagus of the Otitidae and the other families mentioned. In the Otitidae the aedeagus in repose is coiled in a rather tight spiral like a watch spring so that it can be accommodated in the genital pouch. In the other three families the aedeagus (or the stipe of the aedeagus) forms a loose complex loop generally passing forwards and then downwards from its base, backwards and upwards on the right side of the epandrium, then forwards on the dorsal surface 24 of the epandrium into the genital pouch. If, as in some Platystomatinae, the aedeagus has very long terminal filaments these may be spirally coiled within the genital pouch. Steyskal (1961) discarded the use of the setulae on vein 1, as a family character because of its variability within the family Otitidae. My further studies suggest its use as follows. In those otitids which have setulae on vein 1 these are confined to the distal part of the vein from about the level of maximum forward curvature of the subcosta (only a few exceptions, perhaps confined to the Neotropical Region). In all Tephritidae, Pyrgotidae, and Platystomatidae which I have examined the setulae on vein 1 extend basally almost to level of humeral crossvein and in some genera of ‘Tephritidae and Platystomatidae they extend considerably further basad. Examination of the genitalia figures of Otitidae provided by Steyskal suggests a further difference between the two groups. In Platystomatidae and ‘Tephritidae there is a pair of inner surstyli which generally have two (sometimes one) prominent distal teeth (termed prensisetae by Munro, 1947). In most Otitidae only the outer pair of surstyli appears to be present, but if there is an inner lobe or process (as appears to be the case in Diacrita) it is without the specialized teeth. In view of these contrasting characters I am in agreement with Steyskal that the Platystomatidae are well removed from the Otitidae and more closely aligned with the ‘Tephritidae and probably the lesser known Pyrgotidae. A further tephritoid family that should be mentioned here is the 'Tachiniscidae. It is possible that this rare and little known Neotropical and Ethiopian family is also a member of the narrow group which includes the ‘Tephritidae, Pyrgotidae and Platystomatidae. If this is the case the family should be readily distinguished from the Platystomatidae by the extraordinary development of the chaetotaxy and the well developed lobe of the anal cell. Mr B. Cogan of the British Museum (Natural History) is at present investigating the morphology and relationships of this family. The family Pyrgotidae has been distinguished from other tephritoid families (most recently by Steyskal, 1961) by the absence of ocelli. However ocelli are fully developed in the somewhat aberrant subfamily Teretrurinae. If the view of Malloch (1933) is accepted that this group is not really pyrgotid, then there are still many undoubted pyrgotid species in which the ocelli are present in various stages of development. In the platystomatid genus Bromophila ocelli ave absent, but they are present in all other platystomatids examined. In the Pyrgotidae the anal cell is generally produced into a short acute lobe at its outer posterior corner, though there are a few forms in which this lobe is indistinct. ‘This lobe is always absent in Platystomatidae. A more reliable method of distinguishing the Pyrgotidae from the Platystomatidae is provided by the structure of the female abdomen. In females of Pyrgotidae segment 6 is always well developed with tergite similar to and only slightly shorter than the preceding tergites. In platystomatid females tergite 6 is always much reduced in length, concealed, or altogether absent, except in the aberrant Apiola, the position of which is doubtful. Further the ovipositor sheath of Pyrgotidae (except in the ‘Teretrurinae) is very strongly developed and dorsally convex, more or less conical, cylindrical, or globose; segment 8 is extensively membranous, usually without longitudinal sclerotized bands, appearing amorphous. in dried specimens but probably inflatable; and the aculeus is much modified, usually thorn- like or reduced, though sometimes with accessory sclerites. By contrast, in the Platystomatidae the ovipositor sheath is dorsoventrally compressed with keeled lateral margins and is less strongly sclerotized; segment 8 forms a tubular inversion membrane with longitudinal sclerotized bands; the aculeus, though somewhat variable, is always simple, straight, with sensory hairs on each side near apex. ‘To summarize, the Pyrgotidae and Platystomatidae appear to be very closely related families; structurally there are some rather sharp distinctions so that their separate family status seems assured, but a much more thorough morphological study of the Pyrgotidae is to be desired. The morphology of the ‘Tephritidae is better known than that of most acalyptrate families and therefore provides a better basis for comparison with the Platystomatidae. Remarkably consistent differences exist between the two families as shown in the following table. 25 ‘TEPHRITIDAE PLATYSTOMATIDAE Lower incurved fronto-orbital bristles present. Lower fronto-orbital bristles absent. Costa broken near distal end of subcosta. Costa not broken near distal end of subcosta. Anal cell produced into an acute lobe (absent Anal cell not produced into an acute lobe. in a few forms). Segment 6 of 2 abdomen well developed, its Segment 6 of 2 abdomen much shortened, tergite only slightly shorter than tergite concealed, or absent. 5 Apart from these differences there is very close agreement between the families so that the Tephritidae appear to be at least as closely related to the Platystomatidae as are the Pyrgotidae. There is very close similarity in the male genitalia between many forms of both families. ‘The inner surstylus of most tephritids and platystomatids has a pair of large distal teeth and it is in the more primitive platystomatids such as Lasioxiria that this structure most resembles that of Tephritidae. The kind of aedeagus perhaps most common in the Tephritidae, with complex distal glans, is very similar to that of many platystomatids of the subfamilies Plastophritinae and Scholastinae. In the female postabdomen also the resemblance is remarkable. In many, perhaps primitive, genera (including Xiria, Pterogenomyia, Lasioxiria, and the Trapherinae) the aculeus is broad, flattened, often with acute apex, and apparently forms a piercing organ much as in Tephritidae. In some of these platystomatid genera segment 8 has rows of minute cuticular teeth as in many Tephritidae. Again in many tephritid genera (e.g., Dacus, Tephritis) the ovipositor sheath is dorsoventrally compressed in much the same way as in Platystomatidae. Conclusions. ‘The family Platystomatidae belongs in the superfamily Tephritoidea (or Otitoidea). Within the superfamily it is much less closely related to the Otitidae than has often been supposed, and is very closely related to the Pyrgotidae, Tephritidae, and perhaps Tachiniscidae. It is not closely related to the Richardiidae. There are adequate grounds for considering the Platystomatidae a distinct family from the other families mentioned above. Definition of Platystomatidae. ‘TYephritoid Diptera having the following combination of characters: ocelli usually present; no lower fronto-orbital bristles; vibrissa absent or rarely a rudimentary vibrissa present; costa broken just beyond humeral crossvein, but not broken near end of subcosta; vein 1 always with continuous dorsal series of setulae from immediately beyond humeral crossvein to its distal extremity, sometimes also some more basally situated setulae; anal cell not produced into an acute lobe at posterodistal corner, the anal crossvein either straight or convexly curved; g: inner surstylus well developed, generally with one or two distal teeth; stipe of aedeagus not forming a tight spiral coil in repose; 9: abdominal segment 6 always more or less reduced, with tergite 6 much shorter than preceding tergites, often concealed or absent altogether; segment 7 forming a flattened ovipositor sheath; segment 8 forming a tube which can be retracted by invagination, supported by several longitudinal sclerotized bands; aculeus always developed, straight, with sensory hairs on each side near apex. (B) Subfamily Classification The first serious attempt at major groupings of genera within the Platystomatidae is that of Hendel (1914a). Enderlein (1922; 1924) added several more groupings. These groupings were called tribes as the authors considered the Platystomatidae as a subfamily of either the Muscidae or Ortalidae (Otitidae). Frey (1932) and Brues, Melander, and Carpenter (1954) recognized a number of these as subfamilies, assigning family rank to the Platystomatidae. Hennig (1945) transferred the subfamily Myrmecomyiinae (Cephaliinae) to the family Otitidae, and more recent authors (Steyskal, 1962, 1965a, 1965c; Frey, 1964) appear only to recognize three subfamilies, viz., Trapherinae, Plastotephritinae, and Platystomatinae. Hennig (1940a) provided evidence that the subfamily Angitulinae of the obsolete family Phytalmiidae should be transferred to the Platystomatidae. 26 A thorough examination of most of the world genera would be desirable to produce a satisfactory reclassification, but a study of Australian genera, supplemented by a less complete study of certain other genera, shows that many of these are currently placed in the wrong subfamilies. The characters of the male genitalia and of the female abdomen appear to provide the most reliable characters but these are not always in accord with the characters of general form and chaetotaxy used by Hendel and Enderlein. Unfortunately the male genitalia remain unstudied for many genera. I have examined the aedeagus of a number of non-pletystomatine genera but in view of the available time and material I have examined the sperm pump in only a few of these. At present it appears that the five following subfamilies should be recognized: (1) Subfamily Trapherinae Hendel, 1914a Synonym: Xiriina Hendel, 1914a. I have not examined the male genitalia of the more typical members of this group. Steyskal (1965) has figured and briefly described the genitalia of Poecilotraphera and indicated that in the genera Traphera and Lule the structure of the aedeagus is similar. The stipe of the aedeagus is largely membranous and inflated with a single anterior sclerotized longitudinal strip which is expanded apically. ‘There is no well defined glans. In Lule abdominal tergite 5 is as long as tergite 4, and in the female tergite 6 is well-developed and exposed though much shorter than tergite 5; the aculeus of the ovipositor is very broad and acute. Traphera, Poecilotraphera, and Phasiamya (= Pachymyza) resemble Lule in these characters. Xiria is of rather different habitus to the above genera but agrees in the female abdomen, as well as in the absence of the glans of the aedeagus. I therefore include it in this subfamily. The group is represented in the Ethiopian and Oriental Regions. (2) Subfamily Plastotephritinae Enderlein, 1922 Diagnostic characters: squama reduced to a linear fringe; tergite 5 not much shorter than tergite 3; g: aedeagus with at least partly sclerotized glans which is always without terminal filaments; 9: abdomen with tergite 6 absent or vestigial and concealed below tergite 5; aculeus variable, often broad and acute. Of the genera I have examined I would include the following in this subfamily: Atopognathus (= Dasiortalis), Conopariella, Lasioxiria, Plastotephritis, Pterogenomyia. It is possible that an undescribed genus from Lord Howe Island and New Guinea belongs here. Chaetorwellia appears to be more in agreement with this subfamily than any other. The aedeagus has a distinct glans without filaments, the aculeus of the ovipositor is broadened much as in Lasioxiria, and the squama is little developed. However tergite 5 is much reduced in the female only. It may be considered to be a transitional genus between the Plastotephritinae and Scholastinae. Frey (1932) also includes the following genera: Acanthoneuropsis, Agrochira, Atopocnema, Federleyella, Rhegmatosaga. 1 am unable to comment on the position of these genera, most of which are African. _Tylopterna, included here by Frey, is a chloropid (Sabrosky, 1951). (3) Subfamily Scholastinae Enderlein, 1924. This group of stout-bodied flies is somewhat intermediate between Plastotephritinae and Platystomatinae. Enderlein defined the group on the presence of the sternopleural bristle, a character much too unstable for use at the subfamily level. Other authors have not separated the group from the Platystomatinae. Diagnostic characters: squama always forming a definite lobe; tergite 5 much shorter than tergite 3, especially in 9 where it is often absent; 3: aedeagus with glans as in Plastotephritinae, but sometimes more complex, without terminal filaments; @: tergite 6 absent; aculeus very slender with rounded apex. Pj} I include the following genera in this subfamily: Asyntona, Lenophila, Mesoctenia, Naupoda, Neohemigaster, Pterogenia, Scholastes, Trigonosoma (= Tropidogastrella), Kygaenula. (4) Subfamily Platystomatinae Schiner, 1864, Synonyms: Achiadae Bigot, 1852; Achiides Walker, 1857; Cleitamiina Hendel, 1914a; Lamprogastrina Hendel, 1914a; Loxoneurinae Enderlein, 1912; Rivelliina Hendel, 1914a; Stenopterinina (correctly Senopterinini) Hendel, 1914a. Diagnostic characters: squama variable, often forming a well developed lobe; tergite 5 well developed, in 2 sometimes shorter than tergite 3; 3: aedeagus with simple oval or cylindrical sclerotized glans and normally 2 hollow terminal filaments, each with apical gonopore; vesicle of sperm pump with heavily sclerotized cap usually with a pair of swellings or tubercles; 9: tergite 6 vestigial and concealed below tergite 5, or altogether absent; aculeus slender with rounded apex. In the African species Bromophila caffra (Macquart) the aedeagus has a single terminal filament, but this appears to be the product of fusion of the usual two. In a few species of Lamprogaster there are three instead of two filaments. I have examined the following genera belonging to this subfamily: Achias, Achiosoma, Brea, Bromophila, Cleitamia, Cleitamoides, Duomyia, FElassogaster, Euprosopia, Euthyplatystoma, Euxestomoea, Laglaizia, Lamprogaster, Loxoneuroides, Loxonevra, Microepicausta, Plagiostenopterina, Platystoma, Pogonortalis, Pseudepicausta, Pseudorichardia, Rhytidortalis, Rivellia, Scotinosoma, Senopterina, Xenaspis. (5) Subfamily Angitulinae Enderlein, 1936 Diagnostic characters: occiput strongly convexly produced; prothorax forming a long neck; scutellum with pair of elongate horn-like tubercles, each with an apical bristle; membranous cleft of mesopleuron variably prolonged into sternopleuron; metathoracic postcoxal bridge broadly sclerotized; squama little developed; $: aedeagus with well developed glans, with or without pair of terminal filaments; sclerotized cap of vesicle of sperm pump with two gibbosities. Includes the following genera: Angitula, Angituloides, Giraffomyia. Giraffomyia and Angituloides together exhibit a number of features which contrast with those of Angitula. Some of the more noteworthy characters separating the two groups of genera are as follows: ANGITULA Palpus well developed, reaching to anterior edge of prelabrum. Head without vertical bristles. Submarginal cell much wider than marginal cell at and beyond level of anterior crossvein. Anal cell at distal end no wider than distal end of second basal cell. Abdominal spiracles 4 and 5 each with opening concealed by a tuft of pubescence. @ with tergite 6 of abdomen much reduced and concealed below tergite 5. 6: aedeagus with pair of terminal filaments. GIRAFFOMYIA and ANGITULOIDES Palpus small and narrow. A pair of strong vertical bristles. Submarginal cell not wider than marginal cell near level of anterior crossvein. Anal cell expanded distally and much wider than distal end of second basal cell. Abdominal spiracles each with opening surrounded by an almost glabrous sclerotized ring. Q with tergite 6 well developed and exposed, but much shorter than tergite 5. 6: aedeagus without terminal filaments. 28 In some of these characters, notably in the aedeagus, degree of development of tergite 6 in the female abdomen, and development of pubescence on the abdominal spiracles, Angitula appears to be aligned with the subfamily Platystomatinae, while the other two genera contrast sharply with that group. On the other hand the combination of apomorphic characters common to all three genera is sufficiently distinctive to establish them as a single monophyletic group. I therefore regard them as constituting a highly specialized subfamily, which probably originated from near the base of the platystomatine stem. (C) List of Australian Species SUBFAMILY PLASTOTEPHRITINAE Lasioxiria Hendel, 1914a (1 undescribed sp.) Genus A (unnamed) (1 undescribed sp.) SUBFAMILY SCHOLASTINAE Lenophila Guérin, 1843 dentipes (Macquart, 1843) coerulea (Macquart, 1846) (also 4. undescribed spp.) Asyntona Osten Sacken, 1881 tetyroides (Walker, 1859) Pterogenia Bigot, 1859 similis Malloch, 1939a nubecula Hendel, 1914b latericia Hendel, 1914b Naupoda Osten Sacken, 1881 regina Hendel, 1914b insularis Paramonov, 1957 Mesoctenia Enderlein, 1924. australis n. sp. SUBFAMILY PLATYSTOMATINAE Rhytidortalis Hendel, 1914a rugifrons (Thomson, 1869) solocufemur (Enderlein, 1924) n. comb. (also c. 3 undescribed spp.) Scotinosoma Loew, 1873 bistrigatum Hendel, 1914a completum (Malloch, 1931) attenuatum (Malloch, 1931) erasum Malloch, 1939a (also c. 3 undescribed spp.) Microepicausta Hendel, 1914a gracilis Hendel, 1914a terraereginae (Malloch, 1928a) n. comb. (also 1 undescribed sp.) Elassogaster Bigot, 1859 sepsoides (Walker, 1861) (also 1 undescribed sp.) Plagiostenopterina Hendel, 1914a (subgenus Plagiostenopterina) aenea (Wiedemann, 1819) enderleint Hendel, 1914b (subgenus Stenopterosoma Malloch, 1939a) claudiana n. sp. crinita n. sp. macies N. Sp. Pogonortalis Hendel, in de Meijere, 1911 doclea (Walker, 1849) commoni Paramonoy, 1957 howe: Paramonov, 1957 Rivellia Robineau-Desvoidy, 1830 connata (Thomson, 1869) viridis Hendel, 1914a mentissa (Walker, 1849) virgo Hendel, 1914a isolata Malloch, 1930b bipars (Walker, 1858) nigripes (Macquart, 1851) n. comb. (also c. 23 undescribed spp.) Genus C (unnamed) (1 undescribed sp.) Loxoneuroides Hendel, 1914a varipennis Hendel, 1914a Brea Walker, 1859 (1 undescribed sp.) Genus B (unnamed) (1 undescribed sp.) Lamprogaster Macquart, 1843 (violacea group) violacea (Macquart, 1843) tricauda n. sp. (impertalis group) indistincta Malloch, 1928a flavihirta n. sp. nigrihirta n. sp. rugifactes n. sp. imperialis n. sp. (stenoparia group) viola Malloch, 1929 stenoparia Hendel, 1914a corax Nn. sp. ( flavipennis group) bicolor Macquart, 1847 flavipennis Macquart, 1843 vella (Walker, 1849) Spat excelsa Nn. sp. nigripes (Macquart, 1851) corusca N. Sp. laeta (Macquart, 1835) relucens 1. sp. hilaris (Walker, 1849) Sp. 2 poecila Hendel, 1914b maculipennis Macquart, 1847 (macrocephala group) macrocephala Hendel, 1914a Achias Fabricius, 1805 kurandanus Hennig, 1940b australis Malloch, 1939a Achiosoma Hendel, 1914a apictipenne (Hennig, 1940b) n. comb (also 1 undescribed sp.) Duomyia Walker, 1849 scutellaris (Macquart, 1851) personata n. sp. commont n. sp. convallis n. sp. pallipes n. sp. tomentosa Hendel, 1914b monteitht n. sp. Joliata n. sp. triquetra n. sp. sericea Hendel, 1914b botulus n. sp. argentata n. sp. eremia Nn. sp. smaragdina n. sp. lacunosa n. sp. marginalis n. sp. chaetostigma n. sp. apicalis (Walker, 1849) lutea n. sp. spinifemorata Malloch, 1929 29 irregularis, Malloch, 1929 Serra, 1. sp. umbrosa n. sp. curta n. sp. octoseta 1. sp. Scipio n. sp. Sper aurantiaca n. sp. testacea (Macquart, 1855) dete n. sp. longicauda n. sp. capnodes n. sp. ustulata n. sp. brevicornis, n. sp. angustata n. sp. lonchaeina n. sp. capitalis n. sp. picta n. sp. rasa 0. sp. latipilus n. sp. parallela n. sp. adelaidae n. sp. iris 1. Sp. acrogenea Ni. sp. upton n. sp. viridaurea Nn. sp. azurea Hendel, 1914b amentna n. sp. montium n. n. scintilla n. sp. decora (Macquart, 1846) obscura Walker, 1849 hypene n. sp. gigas (Macquart, 1851) thalassina Walker, 1849 howensis n. sp. maculipenns Hendel, 1914b mithrax Hendel, 1914b glebosa n. sp. loxocerina n. sp. cancellata n. sp. nigricosta Malloch, 1929 hebes n. sp. brevifurca n. sp. rudis n. sp. Euprosopia Macquart, 1847 (miliaria group) rete n. sp. acula n. sp. (ventralis group) ventralis (Walker, 1859) xanthops n. sp. punctifacies Malloch, 1928a lenticula n. sp. Sericata n. sp. Spal crassa . sp. Jos D sp. 3 piperata n. sp. conferta n. sp. integra n. sp. kurandae n. sp. Ses ae mica Ni. Sp. (remota group) remota n. sp. (megastigma group) hollowayt n. sp. megastigma n. sp. crispa n. sp. (maculipennis group) albipila n. sp. maculipennis (Guérin, 1831) sp. 5 Jimbripes n. sp. 30 vitrea 1. sp. alpina n. sp. celsa Nn. sp. (scatophaga group) hypostigma n. sp. subacuta n. sp. filicornis n. sp. scatophaga Malloch, 1930a ramosa Ni. Sp. (macrotegularia group) macrotegularia Malloch, 1928a subula n. sp. (tenuicornis group) anostigma n. n. armipes Ni. sp. monodon n. sp. tenuicorns Macquart, 1847 (separata group) comes ni. sp. separata Hendel, 1914a imermis 1. Sp. conjuncta Hendel, 1914b biarmata Malloch, 1929 Notes on Genera and Species Lastoxiria. Hendel (19144; adjacent areas in North Queensland. New Guinea. 1914b) based this genus on a single species from New Guinea. An undescribed species occurs in the rain forests of the Atherton Tableland and I have collected a third species, also undescribed, in Genus A. This is a possibly new genus based on an undescribed species from Lord Howe Island, 350 miles east of the coast of New South Wales. The distinguishing characters are given in the key to genera, but as only a single damaged specimen is available it seems best to defer formal description for the present. A second species of the genus has been recently seen from New Guinea. It is possibly related to Guamomyia from Micronesia. Lenophila. Munro (1959) has pointed out that this is the correct name for the genus previously known as Celetor Loew. Rhytidortalis solocifemur (Enderlein) is a new combination for Pseudepicausta solocifemur Enderlein, 1924. ‘This species is related to R. rugifrons (Thomson) but differs in the smooth postfrons, horizontal grooving on the parafacial and narrower anal cell. I have seen the following material from coastal New South Wales: Sydney, no date (lectotype g, here designated, paralectotype 2, ZMB), Damel; Nelson’s Bay, Port Stephens, viii (or v) 1920 (1 9, AM), A.M. Microepicausta. and Elassogaster. Microepicausta terraereginae (Malloch) is a new combination for Elassogaster terraereginae Malloch (1928a). The species is very closely related to M. gracilis Hendel, the type-species of Microepicausta. I have examined the type material of both species. Elassogaster evitta Malloch (1939a) from New Britain also appears to belong in the genus Microepicausta. 31 Pogonortalis. Wendel (1914a) gives P. barbifera Hendel (= P. doclea (Walker)) as the type species. I consider that P. uncinata de Meijere is the type species by monotypy, as given by Steyskal (1956b). A further study of P. uncinata (from Java) is now needed to decide whether it is congeneric with P. doclea and the other Australian species. As pointed out elsewhere Pogonortalis similis Hendel belongs in the genus Plagiostenopterina. Rivellia nigripes (Macquart) is a new combination for Urophora nigripes Macquart (1851). I have examined the type (PM). Genus C. This unnamed genus is related to Loxoneuroides and Rivellia. To date only a few females of the single species have been collected in rain forest areas of New South Wales. Loxoneuroides varipennis Hendel. This form has three distinctive geographic variants in northern New South Wales and Queensland. The status of these variants is not clear at present. Brea. This genus contains several species in New Guinea and adjacent islands. I have collected an undescribed species on the Cape York Peninsula, Queensland. Genus B. This apparently unnamed genus is represented by a single female specimen which I collected on the Atherton Tableland, Queensland. It may be distinguished by the characters given in the key to genera. Though its appearance is reminiscent of a small dark sepsid, it shares a remarkable number of structural characters with the Oriental genus Loxonevra. It is considered wise to defer description till more material is available. Achias and Achosoma. Mystia Walker (1861) is a new synonym of Achias Fabricius (1805). Achias attrahens (Walker) is a new combination for Mystia attrahens Walker (1861). Achias kurandanus Hennig is the species erroneously determined by Malloch (1939a) as A. brachyophthalmus Walker. Achias apictipennis Hennig belongs in the genus Achiosoma and. is quite closely related to Achiosoma dacoides (Walker). Scholastes. No species of this genus is known to be established in Australia. S. cinctus (Guérin, 1831-1838) was originally erroneously reported to be from Port Jackson, New South Wales. It is probable that this species occurs within Australian limits on the more northerly of the Torres Strait Islands, as it is common on the adjacent coast of Papua-New Guinea. S. bimaculatus Hendel has been reared in Sydney from imported cocoa-nuts. Apiola is a new name for Xenognathus Malloch (1930c, preoccupied by Xenognathus Gilbert, 1915 (Pisces), Case, 1928 (Pisces), Jordan, 1928 (Coleoptera)). ‘This genus, which is only known from Samoa, appears to be a platystomatid, but its subfamily position is doubtful. Pachymyza Frey, 1964, is a new synonym of Phastamya Walker, 1849. The type species of the two nominal genera, P. metallica Walker and P. coleoptrata Frey, appear to be synonyms. I have seen Walker’s type and an addition series of this Indonesian form in BM. (D) Key to Australian Genera of Platystomatidae I Ides Glonkelhy Inaiinyg ‘yea Semllese loos Gasoncoscdsanon node Lastoxiria Hendel BiveseWALCIOlga MOstsOne VEIN HmUsmally Wane sais eye ela neem de ce ieepe nes skies B 3. (i) eRe Or Wel An gelnil los BICONE 4 Sain cryin Sao ae cicka tis heen Bec nel eee Oe een B Wieitisiyathiout setae perore leveliof lumenal crossyeine) se. 2-2-6 sees. Sa. 4 3 (2) Arista bare, or haired near base only; face with a broad, flat-topped carina between BiMIEODOLEYS | oto i Soe Moe BIRO AC ORE ae aa ane Euprosopia Macquart Arista plumose throughout; facial carina obsolete .............. Pterogenia Bigot 52 4 (2) Middle femur swollen, much thicker than other femora, with 2 rows of stout ventra Ohi eames ohne oon AN Ane R ORSON oOo C0 Lod Ogu TOON CO oO Up Ae clos 5 ——— Middle femur not noticeably thicker than other femora, either without ventral spines, or with weakly spinescent bristles no more developed than on fore femur ...... 5 (4) Antennae separated by a distance at least equal to width of first segment; head appressed to thorax in repose; ¢: tergites 4 and 5 together much shorter than tergite 3; Q: tergites 4 and 5 absent, i.e., preabdomen with only 2 separate dorsal SCIERITES rere CE ae hee seo er mre cence itor Mesoctenia Enderlein Antennae exceedingly close together at their bases; head held away from main body of thorax by the neck-like prothorax; ¢ and Q: tergite 5 well-developed, not Shorter thiamin oitend, wap eects onic) ssc reel eee ease Newer ronment eter Brea Walker 6 (4) Second basal cell longer than discal cell; very stout or subglobose flies .......... Fi Second basal cell shorter than discal cell; form variable but never subglobose ....8 7 (6) Antennae widely separated at bases; anal cell acutely pointed at posterior distal INSMES Vera AL, Grave hirovey ie WMO! BIE coc ogaceoscasceoce Asyntona Osten Sacken Antennae separated basally by not more than width of first segment; posterior distal angle of anal cell not acute; vein 4 ending behind wing apex. Naupoda Osten Sacken 3) (0h Stemoplenralsbpnistlespresentiputioften rather tne mie ce lane trre rere erent ier 9 Stermopleuralibristle absembem. mre roiete ters oictoyetellelerbolstoletel a stokstet set ereet ere 10 g (8) Wing with a broad regular blackish band from end of vein 5, across discal and anterior crossveins to submarginal cell, thence bent back, completely traversing wing again through base of discal cell, also a large separate apical costal mark; Rowicllhnn Wasy Comes, Sistine BEE soc6nscsadensessoonrs< Lenophila Guérin — Wing without such a bent band, irregularly spotted or with broken bands; scutellum somewhat flattened, brown with whitish margin ............ Scholastes Loew 10 (8) Eyes protruding or stalked; arista long-plumose for most of its length; mesopleural bristle*absemt 43 sc0 say. nus eisai lls sce 2s aah are tie ec Ree, ena ene eee ere tee ai Eyes not protruding or stalked; arista bare or minutely haired, at least on distal thind!s smesoplemnallpristlenvatiallb| csr r rey ie eee 12 tr (10) Humeral, intra-alar, and prescutellar acrostichal bristles distinct; squama dilated posteriorly, its outer margin not describing a continuous curve; suprasquamal ridge with some fine erect hairs near middle, which are longer than the pile-like PUubescenceOnspOstenlOnspAlteny wrser a Sener ee cee eee Achias Fabricius Humeral, intra-alar, and prescutellar acrostichal bristles absent; squama not dilated posteriorly, its outer margin describing a continuous curve; suprasquamal ridge wWhlda Gxdneenelhy Savordt [UIKETGNCTS OMY Goocdaccnacacbancccccs Achiosoma Hendel 12 (10) Mesopleural bristle absent; suprasquamal ridge with a group of rather long, loose AaITSHOLES tule SCLULLAG es vce a cree. SU re on tenes, Semen Ce aaa Duomyia Walker Mesopleural bristle present; suprasquamal ridge with short pubescence only ....13 13 (12) Face with a broad, flat, usually sharply margined carina between antennae; lower squama very large, always greatly exceeding the upper squama ................ ROU GES UD CONDON ORM O ODOUR OSC ANCE Te OO oe GOs ca ees ads Lamprogaster Macquart Facial carina little developed or completely rounded off at lateral margins; lower SqUamamOtmuchilarcen tian Upper squalnaar oi cyiaie scissile eae eter 14. 33 14 (13) Lower part of parafacial with numerous short black setulae; lower margin of cheek with a series of longer setulae, the foremost corresponding in position to a vibrissa; subcosta angularly bent forward near distal end; wing blackish with variable clear MBAR Qj guda ood scaek obeuaoogeeeogeoo aU uoRUSUusnedanconmogager genus A Lower part of parafacial with at most a few inconspicuous hairs; lower margin of cheek without complete series of long setulae; subcosta not abruptly bent distally, usually curved forward; wing predominantly clear, usually with few blackish GaaOES OP YOOUS > soos ed coanonsouo Uo es osu OeduOOD OA aU Dd dcau GEO uUs 00 UR dO OOK 15 15 (14) Prelabrum much prolonged downwards and backwards; metathoracic spiracle with a few fine setulae on upper margin; middle tibia without strong apical spur .... oi" “SS PROLS Ws tn Gas eh Gin ere Ree SOs gi cs Nodal ne Maas run her rece ec genus B Prelabrum normal, transverse; metathoracic spiracle with soft hairs only; middle (inlorka, Wallen CIS Ge WO Garo GUS ocd oacncccccodrscodconssdsaucconnsa0ns 16 16) (15) Vems 9 and 4 converse towards apex’... --..-----+-- ++ e+e + sees sees 17 Veins 3 and 4 subparallel or slightly diverging towards apex .............++--- 20 17 (16) Face with fine hairs near middle; wing with a longitudinal blackish stripe covering Pima Seal ACN Ly Msn. caller clad dio ols iain a Ao woe e Omi Plagiostenopterina Hendel Face without hairs near middle; wing markings variable ..................-- 18 18 (17) Fronto-orbital bristles distinct; mesoscutum approximately as wide as long; wing with a longitudinal blackish stripe covering first basal cell .. Pogonortalis Hendel Fronto-orbital bristles vestigial; mesoscutum longer than wide; wing without stripe COVERING inet: oAgall Gell so ondloscooodnooeobugosshGndcdcdMuEmodkpEoodsbOoe 19 1g (18) Scutellum not haired or with few hairs at lateral margins in some non-Australian Species; veim Arencimeypetore winevapex 2-0. ies Elassogaster Bigot Scutellum with hairs on most of dorsal surface; vein 4 ending slightly behind wing BIDIENS” saio'g td d-dh Slain detroit Chern Sate Greet stan Socit ca aia cae Microepicausta Hendel 20 (16) Scutellum haired or setulose on central region of dorsal surface; vein 4 nearly always deeply curved down into discal cell before anterior crossvein ...........++--- QI Scutellum not haired in central region, but sometimes with very short pubescence or with basal hairs; vein 4 usually slightly curved or almost straight before anterior CROSS | fee 5 0. G8 0:4. tue Gioia 0 08 CORO OSS CREE OCG Cy ORAON ROSE Oe REET em eee ST 22 21 (20) Scutellum with three pairs of marginal bristles; prescutellar acrostichals surrounded by long hairs; second basal cell about three quarters as long as discal cell ...... Loxoneuroides Hendel Scutellum with two pairs of marginal bristles; usually no hairs between prescutellar acrostichals and scutellar suture; second basal cell not more than about half as long as chscallk calla” pith. eH oo se en Sa eR ee ae enn Rivellia Robineau-Desvoidy 22 (20) Third antennal segment short, rounded-ovate; discal crossvein longer than penultimate SECM OMMOPVEllIe ee ON rents Meet eren eA iret to fe NE erates os choterei ni ens sient lal nes 4 genus C Third antennal segment elongate; discal crossvein shorter than penultimate section OLGA cea 8 Sorewindts walang ORL dao earale calls Ban aca» Baneee eset ani ere rene errr 23 23 (22) Mesoscutum with one or more longitudinal greyish pruinescent stripes; arista with numerous short or minute hairs basally, which extend beyond the slightly swollen lenRaILiowNe: SoG bc ob adem be hbo s BO Ooo Up ot CRD ohn sir creme Scotinosoma Loew —— Mesoscutum without longitudinal pruinescent stripes; arista quite bare, or in one species with minute hairs confined to the strongly swollen basal part ............ Be eer cane che BON er fe 1c ome a ee rie a Rhytidortalis Hendel G 77811—2 34 VI. GENUS MESOCTENIA ENDERLEIN Mesoctenia Enderlein, 1924: 130; Malloch, 1939a: 123. Head broad, about as wide as thorax in 9, often wider in g, compressed from front to rear; postfrons broad, steeply sloping; facial carina only slightly elevated, flat-topped, at least as wide as antennal segment 1, often wider in 3, between bases of antennae; epistomal margin of face sometimes produced medially into a prominent, downwardly directed lobe in 3, which is only very slightly developed in Q; cheeks wide, somewhat gibbous laterally in 2, forming a vertical, anteriorly flattened ledge in this region in J; outer vertical bristle strong, inner vertical a little weaker; fronto-orbital vestigial or absent; ocellar absent; cheek bristle rather weak or absent. Antenna decumbent, with short oval segment 3, arista long, with pubescence on its entire length. Prelabrum moderately developed, narrowed medially; palpus of moderate width, usually tapered basally and apically. Thorax very short and broad, shining, devoid of pruinescence; mesoscutum about 1.3 times as wide as long; scutellum broad, rounded in outline, somewhat flattened but with rounded margin finely setulose on dorsal, lateral and apical surfaces; mesopleuron strongly convex; prosternum without sclerotized precoxal bridges; thorax with the following bristles: a rather weak humeral; 1 + 1 notopleurals; supra-alar; postalar; posterior intra-alar situated rather high up; no dorsocentral; prescutellar acrostichal; two or three pairs of scutellars; mesopleural; no sternopleural. Legs, especially the fore one, rather short; fore femur with rather weak pale posteroventral bristles, the dorsal bristles scarely differentiated from the long dorsal hairs; middle femur strongly swollen and much stouter than other femora, with anteroventral and posteroventral series of strongly thickened black spines. Wing. Subcosta curved forward at distal extremity; vein 3 ending in wing apex, not converging with vein 4 apically; second basal cell much dilated distally, fully as long as discal cell (both cells measured along posterior margins); anal crossvein almost straight forming on its proximal side an obtuse angle with vein 6. Abdomen. 3: tergites 4 and 5 distinct but much shorter than tergite 3; Q: tergite 3 very large; tergites 4, 5, and 6 not distinguishable; spiracles of segments 3, 4, and 5 situated dorsally in membranous area between tergites 3 and 7; aculeus of ovipositor slender. 3 postabdomen: segment 9 (genital segment) not concealed below tergite 5; aedeagus lying in a shallow dorsal, rather than lateral pouch; distal capsule (glans) divided by a transverse constriction, without terminal filaments. Type species: M. ralumensis Enderlein (=