Bulletin of the British Museum (Natural History). British Museum (Natural History) Converted as part of the ABLE project by Dauvit King London : BM(NH) Continued as: Bulletin of the Natural History Museum. Entomology series Vol.1 (1950) - vol.61 (1992) 2 4 This document has been converted to TEI XML as part of the ABLE project to make it more widely available to biodiversity researchers in a useful format. eng text No corrections have been made in the text. The original source has not been regularized or normalized. Quotation marks have not been processed. They are as in the original DjVu XML document. Hyphens, including end-of-line hyphens, have not been processed. They are as in the original DjVu XML document. The text has been segmented based purely on layout based on page breaks. No language level segmetation, such as sentences, tone-units or graphemic, has been applied. Additional mark up using taXMLit has been applied to the TEI XML based on analysis of the original source through the uBio and OpenCalais web services. (Add comment for fuzzy matching once this has been brought into the final workflow too.) 101 t' Bulletin of the L British Museum (Natural History) Entomology series Vol 52 1986 British Museum (Natural History)London 1986 Dates of publication of the partsNol 30 January 1986 No 2 24 April 1986 No 3 24 April 1986 No 4 26 June 1986 ISSN 0524-6431 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset ContentsEntomology Volume 52 No 1 The sandflies of Egypt (Diptera: Phlebotominae) R. P. Lane 1 No 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae) Gaden S. Robinson 37 No 3 A revision of the European Agathidinae (Hymenoptera: Braconidae) G. E.J.Nixon 183 No 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revisionof certain generaJ. Quinlan 243 Bulletin of the British Museum (Natural History) BRITISH .Wt'- ' The sandflies of Egypt (Diptera:Phlebotominae) R. P. Lane Entomology series Vol 52 No 1 30 January 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History) ,Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1986 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06015 5 ISSN 0524-6431 Entomology series Vol 52 No 1 pp 1-35British Museum (Natural History)Cromwell RoadLondon SW7 5BD Issued 30 January 1986 The sandflies of Egypt (Diptera: Phleboto R. P. Lane Medical Diptera Section,* Department of Entomology, British Museum (Natural History),'Cromwell Road, London SW7 5BD Contents Synopsis Introduction Medical importance of sandflies in Egypt Previous studies of Egyptian sandflies 3 Materials and methods 3 Abbreviations 3 Composition and distribution of the sandfly fauna in Egypt 4 Key to the species of Phlebotominae in Egypt 5 Review of species 7 Faunal associations 27 Acknowledgements 30 References 31 Index 35 Synopsis The medical importance of, and previous studies on, the sandflies of Egypt are reviewed. Phlebotominesandflies transmit phlebotomus fever (= sandfly fever) and two forms of leishmaniasis to man in Egypt.The number of species known from Egypt is increased from seven to 21 (8 Phlebotomus, 13 Sergentomyia).Following an identification key, a diagnosis is given for each species together with a discussion oftaxonomic and biological data, including vector status. The Egyptian sandfly fauna is composed of threefaunal elements, one clearly Palaearctic, the others Afrotropical but having particular affinity to thesandflies of the Arabian peninsula. Introduction The study of the phlebotomine sandflies of Egypt, including the Sinai peninsula, is important fortwo reasons: these flies act as vectors of pathogenic organisms there (phlebotomus fever virusand two species of Leishmania) and secondly, the area in which they occur is an importantinterface between the Palaearctic and Afrotropical zoogeographic regions. In a review ofleishmaniasis in the eastern Mediterranean, Zahar (1980: 11) commented on the dearth ofknowledge of Egyptian sandflies and recommended that a 'thorough revision of the sandflyfauna' should be made. The principal objective of this paper is to provide descriptions and keys to the sandflies ofEgypt and indicate established or potential vectors of disease, thus facilitating future studies ontheir biology and assessment of their medical importance in the area. The biogeographicalrelationships of the fauna are discussed in relation to other parts of the Middle East and Africa.Before dealing with the taxonomy of the phlebotomines, a review of the pertinent literature ismade. This is divided into the medical importance of phlebotomines and previous taxonomicstudies. Medical importance of sandflies in Egypt In Egypt phlebotomine sandflies are important as vectors of phlebotomus fever virus and twospecies of Leishmania: Le. major and Le. donovani, the causative agents of cutaneous andvisceral leishmaniasis respectively. Phlebotomus fever (sandfly fever) is a systemic and well-studied febrile disease of man and is * WHO Collaborating Centre for the study of Phlebotomine sandflies in relation to leishmaniasis. Bull. Br. Mus. not. Hist. (Ent.) 52 (1): 1-35 Issued 30 January 1986 2 R. P. LANE endemic in many parts of Egypt (Schmidt et al., 1971; Darwish & Hoogstraal, 1981; Taylor,1958). Phlebotomus papatasi was incriminated as the vector in Egypt by Schmidt et al. (I960)who isolated the virus from flies collected in human dwellings in suburban Cairo. Phlebotominesmay also be vectors of another viral fever, Rift Valley Fever. RVF is an arthropod-borne diseasewhich usually attacks domestic ungulates but, more importantly, has caused considerablehuman disease and mortality in two recent outbreaks in Egypt (Meegan, 1979; Hoogstraal etal. ,1979). The vector of RVF virus has not been discovered yet, although mosquitoes are thought tobe the most likely, particularly Culex pipiens (Meegan etal. , 1980). However, authors regularlystate that Phlebotomus species should still be examined as potential vectors in view of theirabundance and distribution throughout the affected areas. This cautious view is supported byHoch & Bailey's (1983) findings that five out of 34 attempts to transmit mechanically RVF virusto hampsters by using sandflies (Lutzomyia longipalpis) were successful. Until recently, neither cutaneous nor visceral leishmaniasis was considered common in Egyptas they were only known from occasional cases diagnosed clinically. Cutaneous leishmaniasishad been recorded by Khalil (1934), Halawani (1940) and Cahill etal. (1966) who found an areawhere Le. tropica is endemic in the Hihya district of Sharqiya Governate. Cahill (1965) alsomade a leishmanin skin test survey in various villages of the Nile Delta and found that mostpositive cases were either in Sharqiya Governate or originated there. In a sample of 612 patientsin the Faquus area of the governate, 17% of those tested were positive (i.e. had been challengedby Leishmania) with an equal sex distribution. However, recent studies by Soliman & Abo-Shady (1981), Morsy et al. (1982) and Rifaat et al. (1983a, 19836) have shown cutaneousleishmaniasis to be more widespread and common than previously thought. The sandfly P.papatasi is a common peridomestic species throughout Lower Egypt and is probably the vectorof Le. major there as it is in neighbouring Israel (Schlein et al. , 1982, 1984; Adler & Theodor,1925). Furthermore, P. papatasi was the only Phlebotomus species caught in a survey in Sinai(30 50'N, 34 20'E) where cutaneous leishmaniasis is common amongst members of theMultinational Peacekeeping Forces. P. papatasi is probably the vector there also as this area ofSinai is a very similar habitat to that described by Schlein et al. (1984) who are working in theNegev and central Arava regions of Israel , which are contiguous with northern Sinai . Some casesof cutaneous leishmaniasis have recently been recorded formally from southern Sinai (Bassili etal., 1983), but it is undoubtedly more prevalent there as numerous cases are known from centralSinai (J. Zimmerman, pers. comm.) and along the coast of the Red Sea and Gulf of Aqaba ofSaudi Arabia (R. Cross, pers. comm.). In southern Sinai several vector species other than P.papatasi have been found (see below). To date, there have not been any isolations of Leishmaniafrom wild-caught sandflies in Egypt and therefore any suggestion that a species is a vector mustbe speculative, although the circumstantial evidence for P. papatasi is considerable. Until recently, the only autochthonous cases of visceral leishmaniasis recorded from Egyptwere those of Phillips (1904), who found that 32% of patients with splenomegaly in a Cairohospital had Leishmania bodies detected in smears following spleenic puncture. The casedescribed by Hassan (19686) was thought to be imported from Saudi Arabia. Cahill (1968)reports that no visceral infections were found in an extension of his earlier leishmanin survey(Cahill, 1965) to different ecological zones of Egypt. Rifaat et al. (1968) examined potentialreservoirs (443 dogs and 324 rodents) and 'vectors' (P. papatasi and P. sergenti} of visceralleishmaniasis in 13 'indicator areas' of Egypt but did not find any infected with Leishmania.However, Morsy et al. (1982) found that 21-3% of Rattus norvegicus and 12% of R. rattusreacted positively in serological tests for Leishmania in Ismailiya Governate. Subsequently,smears from only one R. norvegicus and four R. rattus were found to have Leishmania bodies,but this was sufficient evidence to consider the rats to be reservoirs of the disease. Recently,Tewfik et al. (1983) detected a case of infantile visceral leishmaniasis from Al Agamy, nearAlexandria. Subsequently, 20 patients have been detected (to June 1984) and the parasiteidentified as Le. donovani donovani (Mansour et al. , 1984) . Al Agamy is an area of new housingdevelopment on a narrow limestone strip between the Mediterranean and a large brackish lake.There is very little vegetation around the patients' houses other than small fig orchards.Intensive sandfly collecting in and around houses produced only two species of Phlebotomus: SANDFLIES OF EGYPT 3 P. papatasi and P. langeroni (J. Beier, H. Kassem and B. Sawaf, pers. comm.). Sawaf et al.(1984) have suggested that P. langeroni is the most likely vector species, for two reasons: it is aclose relative of P. orientalis Parrot , a well-established vector of visceral leishmaniasis in Sudan ,and P. papatasi is the only other Phlebotomus species present and is known to be a poor vector ofLe. donovani (Lewis & Ward, in press). The unlikely possibility remains that the vector isneither of these species but another, such as P. tobbi Adler & Theodor, living well away fromhuman dwellings. Previous studies of Egyptian sandflies Previous studies of the sandfly fauna of Egypt have been fragmentary, and the Sinai and upperEgypt have not been studied at all. Willcocks (1917) recorded the ubiquitous vector speciesP. papatasi from lower Egypt and in 1948 Theodor recorded another anthropophilic species,P. sergenti, from metropolitan Cairo. Khalil (1934) reported Sergentomyia squamipleuris fromSharkiya Governate during a leishmaniasis study. Zein el Dine (1972) made the first attempt at aphlebotomine survey in Egypt, concentrating on the Bahariya, Kharga and Dakhla oases, butonly found P. papatasi and S. palestinensis . She also reports the collection of S. minuta, acommon Mediterranean species, by Eflatoon in 1922. During a survey to find visceral leish-maniasis vectors, Rifaat et al. (1968) added 5. tiberiadis to the faunal list. Recently, P. langeronihas been discovered at Al Agamy, a focus of visceral leishmaniasis near Alexandria (Sawaf et al. ,1984), bringing the known sandfly fauna to seven species: three Phlebotomus and fourSergentomyia. Based on recent collecting and material in the British Museum (Natural History),the present study increases this to 21 species: eight Phlebotomus and 13 Sergentomyia. Nocomprehensive taxonomic work exists facilitating the identification of these species. Materials and methods Specimens for this study of Egyptian sandflies came from several sources and were collected by avariety of methods. Details of the methods used by early collectors such as Omer-Cooper, whocollected in Siwa Oasis in 1909, are not available. However, the methods of recent collectorswho submitted material for identification are available. Rifaat (Cairo) and Braverman (Israel)both used CDC light-traps (with incandescent, not fluorescent, bulbs) in epidemiologicalsurveys, the first in a visceral leishmaniasis survey, the second during a search for Rift ValleyFever vectors. The author's collections were made with sticky traps, with a small lightweightchemical light-trap, and by aspirating from walls and inside animal houses. With the exception of the P. sergenti material collected by Theodor and mounted in Balsam,all specimens were mounted in Berlese medium. The head was removed from the body andmounted ventral side uppermost in a thin film of medium, thus allowing examination at highmagnification. The body was mounted laterally under a separate coverglass. It was necessary toremount many female specimens to show the spermathecae which were usually obscured bydeveloping ova or fat-body. The specimens were immersed in a 1% detergent solution for up to24hrs to remove fat; ova, if present, were dissected out. Abbreviations DEPOSITORIES BMNH British Museum (Natural History) , London , U . K . FM Laboratoire de Parasitologie, Faculte de Medicine, Paris, France. IPA Institut Pasteur, Algiers, Algeria. IPH Institute of Public Health , Teheran , Iran . MI Institute of Tropical Medicine and Parasitology , Moscow , U.S.S.R.U,Pavia University of Pavia, Italy. R. P. LANE NAMES OF COLLECTORS MENTIONEDK.Z.D. K. ZeinelDine M.A.R. M. A. Rifaat R.P.L. R. P. Lane Z.A.H. Z. A. Hassan Y.B. Y. Braverman VARIOUSA3, A4etc.Le. Antennal segment 3, 4, etc.Leishmania , JORDAN SAUDIARABIA airoLOWER EGYPT FAIYUM* SUDANMap 1 The principal towns, oases and areas mentioned in the text. Composition and distribution of the sandfly fauna in Egypt LOWEREGYPT UPPEREGYPT SOUTHERN SINAI Phlebotomus alexandri SintonP. arabicus TheodorP. kazeruni Theodor & MesghaliP. langeroni Nitzulescu SANDFLIES OF EGYPT P. major Annandale 'Sinai form' P. orientalis Parrot P. papatasi (Scopoli) P. sergenti Parrot Sergentomyia adleri (Theodor) 5. antennata (Newstead) 5. christophersi (Sinton) 5. cincta (Parrot & Martin) 5. clydei (Sinton) 5. fallax (Parrot) S. minuta (Rondani) 5. palestinensis (Adler & Theodor) S. schwetzi (Adler, Theodor & Parrot) 5. squamipleuris (Newstead) 5. taizi Lewis S. theodori (Parrot) 5. tiberiadis (Adler, Theodor & Lourie) * denotes presence of a species. LOWEREGYPT UPPEREGYPT SOUTHERNSINAI Key to the species of Phlebotominae in Egypt 1 Female without cibarial armature or pigment patch; male with four or five spines on style; all abdominal sclerites with erect hairs (PHLEBOTOMUS) 2 - Female with cibarial armature and pigment patch present; male with four spines on style; abdominal sclerites with recumbent hairs only, except in subgenus Sintonius (SERGEN-TOMYIA) 16 2 Males 3 - Females 9 3 Style with five short tooth-like spines (Fig. 39), surstyle with spines apically, paramere with three lobes. Small basal tubercle on coxite with non-deciduous hairs papatasi (p. 15) - Style with five long spines, surstyle without spines apically, paramere simple 4 4 Coxite with basal lobe bearing long hairs 5 - Coxite without basal lobe 7 5 A3 short and thick (Fig. 8). Plunger of sperm pump not much wider than barrel (Fig. 11), barrel not much longer than wide. Basal coxite tuft short and broad (Fig. 10) alexandri (p. 9) - A3 long (Fig. 41). Plunger of sperm pump wider than barrel, barrel longer than wide (Fig. 43). Basal coxite tuft long and slender (Fig. 17) 6 6 Style less than half length of coxite, mesonotum pale sergenti (p. 17) Style about half length of coxite, mesonotum dark kazeruni (p. 9) 7 Aedeagus with keel just before tip (Fig. 6); coxite with hair group of more than 30 hairs; sperm ducts with transverse ridges, more than eight times pump length arabicus (p. 7) - Aedeagus smooth to tip; hair group less than 20 hairs; sperm ducts smooth, less than 5 times pump length 8 8 Two ascoids on antennal segments 8-12 langeroni (p. 11) - One ascoid on antennal segments 8-12 orientalis (p. 13) 9 Spermathecae with single segmented capsule. Pharynx narrow, with scale-like armature, posterior margins of hind scales minutelyserrated (Fig. 15) kazeruni (p. 9) - Spermathecae either indistinctly (Fig. 5) or distinctly segmented (Fig. 34) 10 10 Spermathecae indistinctly segmented (Fig. 5) arabicus (p. 7) - Spermathecae distinctly segmented 11 1 1 Pharyngeal armature composed of broad scale-like teeth (Fig. 7) - Pharyngeal armature composed of rows of minute teeth or ridges (Fig. 38) 13 12 Pharynx triangular, lateral and posterior margins straight (Fig. 7); ascoids on A3 and A4 short and stout (Fig. 8); combined length of A3 -I- A4 shorter than labrum alexandri (p. 9) O R. P. LANE Pharynx indented posteriorly (Fig. 40); ascoids on A3 and A4 long and slender, almostreaching end of segment (Figs 41, 42); combined length of A3+A4 longer than labrum sergenti (p. 17) 13 Pharyngeal armature extending to half length of pharynx (Fig. 32) major 'Sinai form' (p. 12) Pharyngeal armature not extending beyond posterior third of pharynx 14 14 Spermathecae with segments subequal, apical segment short (Fig. 37) papatasi (p. 15) Spermathecae with segments unequal in size, middle segments larger than others, apical segment long (Fig. 26) 15 15 Ascoids on A4 more than 0-5 length of segment (Fig. 20) langeroni (p. 11) Ascoids on A4 less than 0-4 length of segment (Figs 21, 22) orientalis (p. 13) 16 Males 17 Females 28 17 Aedeagus finger-shaped, with blunt rounded tip (Fig. 67) 18 Aedeagus with straight tapering sides, pointed (Figs 53, 64) 23 18 Aedeagus curved downwards towards surstyle (Fig. 67) 19 Aedeagus straight or curved upwards 21 19 Style with two apical and two subapical spines (Fig. 64). Cibarial teeth irregular, outer teeth only slightly larger than central teeth, pharyngeal armature a series of fine transverse ridges schwetzi (p. 23) Style with all spines apical 20 20 Outer cibarial teeth scale-like , very much larger than central teeth taizi (p . 25) - All cibarial teeth small, outer teeth same size as central teeth. Aedeagus tapering minutu (p. 21) 21 Cibarium with 18-22 teeth, the central teeth distinctly smaller than lateral teeth .... theodori (p. 25)Cibarium with 16-18 subequal teeth 22 22 Style slender, five to seven times as long as wide (Fig. 58) , accessory seta on style close to apical spines fallax (p. 21) - Style stout, four times as long as wide (Fig. 57), accessory seta at about 0-75 style length antennata (p. 19) and cincta (p. 20) 23 Aedeagus truncated (Fig. 64), style with two subterminal and two terminal spines. Cibarium with straight row of 12 teeth palestinensis (p. 23) - Aedeagus tapering to pointed tip, all spines on style terminal 24 24 Cibarium with small process in front of teeth, mesanepimeron with setal sockets; aedeagus short and gently tapering squamipleuris (p. 25) Cibarium without any processes , mesanepimeron without setal sockets , aedeagus otherwise .... 25 25 Cibarium with convex row of uneven-sized, curved horizontal teeth (Fig. 75) tiberiadis (p. 26) Cibarium with straight horizontal teeth 26 26 Cibarium with 3-4 horizontal teeth (Fig. 51) , large cornua Christophers! (p. 19) Cibarium with 16-26 fine horizontal teeth, cornua small (not wider than long) 27 27 Cibarium with a single row of vertical teeth, horizontal teeth usually small and in groups clydei (p. 20)Cibarium with two or three rows of vertical teeth, horizontal teeth usually well developed adleri (p. 17) 28 Spermatheca clearly segmented (Fig. 52), ducts long and narrow. Abdominal tergites 2-6 with large setal sockets (erect hair sockets) 29 Spermathecae not segmented but sometimes indistinctly striated, ducts not long and narrow.Abdominal tergites 2-6 without large setal sockets (recumbent hair sockets) 32 29 Cibarium with 4-5 long slender widely spaced horizontal teeth (Fig. 49) (numerous spicules may also be present) Christophers! (p. 19) Cibarium with 12-14 closely packed horizontal teeth 30 30 Cibarium with strong, curved horizontal teeth, those at sides longer than central teeth (Fig. 78) tiberiadis (p. 26)Cibarium with straight , closely packed horizontal teeth 31 31 Cibarium with 40-80 vertical teeth in 3-5 rows, more than 20 horizontal teeth (Fig. 48) adleri (p. 17)Cibarium with 12-30 vertical teeth in one or two rows, 12-13 horizontal teeth clydei (p. 20) 32 Spermatheca with capsule covered in numerous small spicules, cibarial teeth in convex row (Fig. 69) squamipleuris (p. 25) - Spermatheca smooth , without spicules , cibarial teeth in straight or concave row 33 SANDFLIES OF EGYPT 7 33 Spermatheca with capsule (Fig. 62). Well-developed cibarial teeth in straight row (Fig. 61) palestinensis (p. 23) - Spermatheca simple , tubular. Cibarial teeth not straight and palisade-like 34 34 Cibarium with 50-60 equal-sized, horizontal teeth in a comb-like row minute (p. 21) - Horizontal cibarial teeth either unequal or row concave 35 35 Cibarium with central horizontal teeth at least half size of lateral horizontal teeth 36 - Cibarium with all horizontal teeth subequal, or lateral teeth only slightly larger than medial teeth 37 36 Cibarium with lateral horizontal teeth scale-like , very much larger than medial horizontal teeth taizi (p. 25)Cibarium with lateral horizontal teeth same shape as medial horizontal teeth theodori (p. 25) 37 Pharynx narrowing posteriorly with transverse striations or vague scale-like patterns schwetzi (p. 23) - Pharynx widening posteriorly with distinct teeth 38 38 Pharynx without distinct shoulder (Fig. 54), sides almost straight, hind margin straight without notch. Cibarium with 16-18 horizontal teeth (Fig. 55) cincte (p. 20) - Pharynx with distinct shoulder to give cordiform or funnel shape, hind margin with medial notch or depression. Cibarium with 16-26 horizontal teeth 39 39 Pharynx cordiform, hind pharyngeal teeth punctate, much smaller than anterior teeth (Fig. 59). Cibarium with 16-22 horizontal teeth, pigment patch usually rounded anteriorly (Fig. 60) fallax (p. 21) - Pharynx not cordiform, without notch in anterior margin. Pharyngeal teeth large, little difference in size between posterior and anterior teeth (Fig. 56). Cibarium with more than 22horizontal teeth, pigment patch usually angular anteriorly antennata (p. 19) Review of species Specimens were examined from many countries but, for brevity, only those from Egypt arelisted. All type-specimens in the BMNH were examined. Phlebotomus arabicus Theodor(Figs 1-6) Phlebotomus (Adlerius) chinensis arabicus Theodor, 1953: 120 [cf $]. Lectotype cf , YEMEN (BMNH), designated by Lewis & Buttiker (1982: 362). Phlebotomus (Adlerius) arabicus Theodor; Artemiev, 1980: 1190. [Raised to species.]Phlebotomus (Adlerius) davidi Artemiev, 1980: 1191 [cf $]. Holotype cf , YEMEN (MI). [Synonymised by Lewis & Buttiker, 1982: 362.] This species is currently placed in the subgenus Adlerius Nitzulescu. FEMALE. Pharynx narrowing after posterior bulge, rounded posteriorly; armature a series of longbackward-pointing teeth. A3 slender, as long as labrum, longer than A3+4. Ascoids short, slender, on A4half length of segment. Palp segments slender, segments 2 and 3 approximately same length. Spermathecadelicate, elongated, ovoid and incompletely striated (Fig. 5) with small terminal knob and thick individualduct. MALE. Wing length 2-29 mm (2-18-2-49 mm; n = 5). Pharynx slender; armature numerous smallbackward-pointing triangular teeth. Genitalia slender (Fig. 4). Aedeagus with pointed keel (Fig. 6). Stylewith two terminal spines, two on protuberance at 0-55 length style and one at 0-45 length style. Coxites with58-4 (54-63, n = 4) hairs. Surstyles longer than coxites. Sperm ducts nines times length of pump. MATERIAL EXAMINED Egypt. Sinai: 3 cf , 3 $ , Mt Katherina, 15-16.vi.1979 (Y.B.); 3 cf , 2 $, Tharfet el Qidren, 13-14.vi.1979(Y.5.); 1 $, Upper WadiNasb, 14-15.vi.l979(Y.B.); 1 $,Feiran, 12-13.vi.1979 (Y.B.). DISTRIBUTION. Egypt, Ethiopia, Yemen, Saudi Arabia. This species has not previously been recorded fromEgypt. Lewis & Buttiker (1982) discussed the variation in this species and distinguished another taxon, P. 'Naqbensp.', from Naqben in northern Saudi Arabia, on four main features: darker thorax, larger size, highernumber of coxite hairs (significantly different at P<0-001) and relatively longer sperm ducts. They did not R. P. LANE Figs 1-6 Phlebotomus arabicus. 1, $, head; 2, $, antennal segments 3 and 4; 3, $, pharynx; 4, cf,genitalia; 5, 9, spermatheca; 6, cf, tip of aedeagus. (Figs 1-3 and 5, 6 from southern Sinai; Fig. 4Ethiopia, Langano.) formally name this species because of the variation in this group and the lack of female specimens fromNaqben. Naqben is in the Jebel Aja mountain system well to the north of the Asir mountains and Yemen,where most P. arabicus in the Arabian peninsula have been found, therefore the specimens of P. arabicusfrom Sinai might reasonably be expected to show closer affinity to 'Naqben sp.' than to Asir P. arabicus.Table 1 compares three of the four features which distinguish 'Naqben sp.' and P. arabicus. Pigmentationwas not included because the Sinai specimens were collected into, and stored in, alcohol which tends tomake them darker. In the features measured, the Sinai arabicus clearly show a greater similarity to typicalP. arabicus (as well as the lectotype) than to 'Naqben sp.', suggesting that the latter is a distinct taxonand not simply a geographic variant of P. arabicus. Table 1 also demonstrates that 'Naqben sq.' is distinctfrom P. arabicus only in the number of coxite hairs and not in overall size and relative length of the spermducts. SANDFLIES OF EGYPTTable 1 Comparison of P. arabicus and P. 'Naqben sp.' P. arabicus is a high-altitude species which Buttiker & Lewis (1984) found in association with P.orientalis and 5. taizi in areas with cool or even cold winters. The vectorial status of P. arabicus is unknown but, as several species of the subgenus Adlerius are vectorsof visceral leishmaniasis in Asia, including the southern U.S.S.R. (P. halepensis; Sergiev, 1979: 208),Afghanistan (P. longiductus; Artemiev, 1978: 20) and the eastern Mediterranean (P. simici; Theodor,1964: 480), P. arabicus should be investigated in any visceral leishmaniasis studies. Phlebotomus alexandri Sinton (Figs 7-14) Phlebotomus sergenti var.; Newstead, 1920: 309 [of]. Phlebotomus sergenti var. alexandri Sinton 1928a: 308 [cf]; Adler, Theodor & Lourie, 1930: 533 [$]. Lectotype cf , IRAQ (BMNH), designated by Lewis (1982: 143).Phlebotomus alexandri Sinton; PernTev, 1966: 72-74. [Raised to species.] This species is currently placed in the subgenus Paraphlebotomus Theodor. FEMALE. Head round (Fig. 9). Pharynx triangular with sides and posterior margin almost straight (Fig. 7);armature with large scale-like plates, each fringed anteriorly with long fine teeth. A3 much shorter thanlabrum (Fig. 9). Ascoids on A3 short and stout (Fig. 8). Spermatheca with 6-7 segments.MALE. Pharynx narrow, not tapering after posterior bulge, posterior margin straight; armature a series ofirregular transverse striations. A3 stout, very much shorter than labrum, A3 +4 only slightly longer thanlabrum. Genitalia overall very short and thick; coxite lobe short and broad (Figs 11, 12). Sperm pump withbarrel as long as or only slightly longer than wide; plunger narrower than barrel (Figs 13, 14). Surstyles aslong as coxite. MATERIAL EXAMINED Egypt. Sinai: Wadi Sa'al, 4 cf , 2 $, 15-16.X.1979; 1 cf, 9-10.vi.1979; 1 $, 14-15.vi.1979 (Y.B.); 1 $,Feiran, 12-13.vi.1979 (Y.B.). DISTRIBUTION. North Africa (including Egypt), Turkey, southern U.S.S.R., Israel, Ethiopia, Yemen,Saudi Arabia, Iraq, Iran, Kuwait, United Arab Emirates, Afghanistan, Pakistan. This species has notpreviously been recorded from Egypt. P. alexandri is usually considered a highland species but it also occurs in suitable lowland areas (e.g.Kuwait: Lane & Al Taqi, 1983). It has been found naturally-infected with unidentified promastigotes inTurkmenia, where it is thought to be an important vector of cutaneous leishmaniasis (Dedet, 1979: 72;Petrishcheva, 1971: 573), and in Iran (Javadian et al., 1977). It is a suspected vector of cutaneousleishmaniasis in several countries and Xiong et al. (1963) suggested that it may be involved in thetransmission of visceral leishmaniasis in China. Phlebotomus kazeruni Theodor & Mesghali(Figs 15-19) Phlebotomus (Paraphlebotomus) kazeruni Theodor & Mesghali, 1964: 289, partim [cf ($ = sergenti)].Holotype cf , IRAN (IPH). This species is currently placed in the subgenus Paraphlebotomus. FEMALE. Pharynx slender with sides straight or slightly concave after posterior bulge (Fig. 15); hind margin 10 R. P. LANE Figs 7-19 7-14, Phlebotomus alexandri. (7) $ , pharynx; (8) $ , antennal segments 3 and 4; (9) $ , head;(10) cf , basal process of coxite; (11, 12) cf, sperm pump; (13, 14) cf , aedeagus. (Figs 7-14 from Sinai,WadiSa'al.)15-19,P. kazeruni. (15) $, pharynx; (16) cf , genitalia; (17) cf, basal process of coxite; (18)Cf , sperm pump; (19) cf , aedeagus. (Fig. 15 from Saudi Arabia, Wadi Khumbra; Figs 16-19 Sinai, MtKatherine.) SANDFLIES OF EGYPT 11 convex; armature extending to 0-25 pharynx length; a series of scales with those at anterior diagonal,becoming progressively transverse posteriorly; rows of minute teeth present on posterior scales; anteriormargin of armature straight or slightly convex. A3 only slightly shorter than labrum, ascoid slender.Spermatheca a single capsule. MALE. Pharynx narrowing after posterior bulge, convex posteriorly; armature a series of fringed scales. A3longer than labrum. Parameres long and slender in lateral view, broad dorsally. Coxite lobe long and broad(Fig. 17). Style three and half times as long as wide. Barrel of sperm pump longer than wide, plunger atleast as wide as barrel (Fig. 18). Surstyles longer than coxites. MATERIAL EXAMINED Egypt. Sinai: 2 cT, St Katherina, 15-16.vi.1979 (Y.B.); 1 cf , Tharfet el Qidren, 13-14.vi.1979 (Y.B.).Several females from various parts of southern Sinai may be this species or P. sergenti, but theirspermathecae, the diagnostic character, are not visible. DISTRIBUTION. Egypt, central Saudi Arabia, southern Iran, Afghanistan. This species has not previouslybeen recorded from Egypt. In Afghanistan, kazeruni occurs in low rocky deserts (Artemiev, 1978). In Saudi Arabia it is notperidomestic but is only found in remote wadis and hills, some very dry with sparse vegetation (Buttiker &Lewis, 1984). Lewis (1982) suggests that this species is sufficiently common in Saudi Arabia to transmitLeishmania among rodents. Phlebotomus langeroni Nitzulescu(Figs 20, 23-28) Phlebotomus perniciosus var.; Nitzulescu, 1930a: 382 [cf]. Phlebotomus langeroni Nitzulescu, 19306: 548 [cf]. Holotype cf , TUNISIA (?FM). This species is currently placed in the subgenus Larroussius Nitzulescu. FEMALE. Pharynx not tapering after posterior bulge, posterior margin straight or slightly convex, armatureextending 0-27 (0-24-0-29) length of pharynx and consisting of series of tightly packed irregular rows ofminute denticles and lateral scales without teeth. A3 slightly shorter than labrum (0-90). Ascoids on A4more than 0-50 length of segment (Fig. 20). Spermatheca with 10-11 segments, neck long, over half lengthof spermathecal body, individual ducts with transverse striations (Fig. 26). MALE. Pharynx slender, parallel-sided after posterior bulge, armature a series of transverse ridges ofminute teeth. Antennal segments 3-12 with two ascoids. Aedeagus slender with ventral subapical opening(Fig. 25). Parameres finger-like. Surstyles longer than coxites. Coxites slender, 5-42 (5-25-5-58) timeslonger than wide (Fig. 24). The female of this species has recently been described (Sawaf, Kassem & Said, in press) (see under P.orientalis for comments on identification). Some aberrant male specimens with six spines on the styleinstead of five were reared in culture (Fig. 28). MATERIAL EXAMINED Egypt. 2 $ , Al Agamy, 15.viii.1983; 1 $ , in poultry house, ll.x.1983 (R.P.L.) [used to establish colony];1 Cf , rodent burrow, 1 cf , sticky trap near house, 5.x. 1982. DISTRIBUTION. Morocco, Tunisia, Libya, Egypt. Sawaf et al. (1984) record this species from Al Agamy,near Alexandria (Egypt), based on the identification of two males listed below (det. R.P.L.). Until recently, P. langeroni was considered a very rare species known only from Morocco (Ristorcelli,1945), Tunisia (Croset et al. , 1978) and Libya (Nitzulescu & Nitzulescu, 1933). In their survey of Tunisia,Croset et al. (1978) found this species in coastal areas only, where it was rare - in one sample of over 9600sandflies from Fondouk-Choucha (near Tunis) there was only a single specimen of P. langeroni. P.langeroni comprised onlyO-l%of 5000 sandflies collected in various parts of Tunisia over a two-year period(Chadli et al. , 19706). Chadli et al. (1970a) collected P. langeroni in the town of Tunis. In Egypt this specieshas only been found during an epidemiological survey at a focus of visceral leishmaniasis at Al Agamy, nearAlexandria. Here, P. langeroni constituted only 1-2% of aspirator catches made inside houses, but up to50% of catches made with sticky traps placed in animal houses and animal burrows near houses (J. Beier,pers. comm.). The only other Phlebotomus species found at this focus was P. papatasi. Sawaf et al. (1984)have suggested that P. langeroni is the vector of visceral leishmaniasis in Al Agamy, because P. papatasi is avery poor vector of Le. donovani and P. langeroni is closely related to other vectors of visceralleishmaniasis (P. orientalis and P. perniciosus Newstead). 12 R. P. LANE Figs 20-28 20-22, cf , antennal segment 4 of (20) Phlebotomus langeroni, Egypt, Al Agamy; (21) P.orientalis, Sudan, Paloich District; (22) P. orientalis, Yemen, Taiz. 23-28, P. langeroni. (23) spermpump; (24) cf , genitalia; (25) cf , tip of aedeagus; (26) $ , spermatheca; (27) $ , spermathecal ducts; (28)Cf , complementary styles, one showing additional spine. (Figs 23-28 all ex laboratory culture originatingin Al Agamy, near Alexandria.) Phlebotomus major Annandale 'Sinai form'(Figs 32-34) Phlebotomus major Annandale, 1910: 46. Phlebotomus (Larroussius) major Annandale; PernTev, 1966: 254. P. major is the type-species of the subgenus Larroussius. Four female specimens from Tharfet el Qidren in the southern Sinai represent a distinct form of P. major.The pharynx, with ridges of fine teeth, is typical of the subgenus Larroussius. However, the pharyngealarmature extends to the mid point of the pharynx (Fig. 32) and therefore closely resembles that of P. majorneglectus from Italy, Malta, Albania, Yugoslavia. It differs from neglectus in having A3 much shorter thanthe labrum. Another subspecies, P. major syriacus, is known from the eastern Mediterranean, andrecently, from northern Saudi Arabia (Lewis & Buttiker, 1982). Table 2 summarises some biometriccharacters of the Sinai form as well as the syntypic series of syriacus in the BMNH from Jerusalem andSyria. The two forms differ in the ratio length of the pharyngeal armature/length of pharynx:- 0-25 insyriacus and 0-5 in 'Sinai form', and the A3/labrum ratio: 0-96 in syriacus and 0-87 in 'Sinai form'. The only other Middle East species with an extensive pharyngeal armature in the subgenus Larroussius SANDFLIES OF EGYPT 13 Table 2 Comparison of some biometric characters of P. major from Sinai and syntypes of P. majorsyriacus from Jerusalem and Syria. is P. wenyoni, which is restricted to certain areas of Iran. P. major 'Sinai form' can be differentiated by theshape of the pharynx and the detailed morphology of the armature. In P. wenyoni the teeth are all on smallelliptical scales with serrated edges whereas in 'Sinai form' the teeth are more isolated and in smallergroups. This difference is especially clear in the anterior part of the pharyngeal armature. Unfortunately, no males of P. major were collected in Egypt. The record of P. major syriacus from SaudiArabia was based on two males and therefore the material is not directly comparable with the Sinaispecimens. P. major was divided into three subspecies by Theodor (1958), and Perfil'ev (1966) added another (P.major krimensis). However, the distribution of some subspecies (given in Lewis, 1982) overlaps to aconsiderable degree and these populations cannot therefore be considered true subspecies (see Lane &Marshall, 1981, for discussion of subspecies). Until they are taxonomically revised, probably as aspecies-group composed of several species, it is not prudent to name formally the distinctive southern Sinaiform, particularly as only females are available. The closely related P. major syriacus has been infected with Le. donovani (see Adler & Theodor, 1957)and is considered to be a vector of visceral leishmaniasis in the eastern Mediterranean area (Hoogstraal &Heyneman, 1969: 1185; Theodor, 1964: 480; Leger et al., 1979: 20). The occurrence of P. major 'Sinai form' in the mountains of the southern Sinai is consistent with thehabitat-characteristic of this species-group. For example, P. major syriacus is only abundant at altitudesabove 300 m in Greece (Leger et al. , 1979). P . major has not previously been recorded from Egypt. MATERIAL EXAMINEDEgypt. Sinai: 4 $, Tharfet el Qidren, 13-14.vi.1979 (Y.B.). Phlebotomus orientalis Parrot(Figs 21, 22, 29-31) Phlebotomus (Phlebotomus) langeroni var. orientalis Parrot, 1936: 30 [C?$]. Syntypes 32 cf, 24 $, ETHIOPIA (IP A).Phlebotomus orientalis Parrot; Parrot & Clastrier, 1946: 64. [Raised to species.] This species is currently placed in the subgenus Larroussius. FEMALE. Pharyngeal armature consisting of irregular rows of nine pin-like teeth (often only the tooth basescan be seen as dark spots), pharynx with indistinct posterior margin (Fig. 30). Ascoids on A3 and A4 short(Fig. 29). A3 almost as long as labrum (Fig. 31). MALE. Pharynx narrowing slightly after posterior bulge, armature a series of short transverse ridges ofminute denticles extending 0-20-0-26 length of pharynx. Antennal segments 8-12 with a single ascoid.Aedeagus long, slender, with lateral subapical opening. Coxite slender, 5-1 times as long as wide. Parrot & Clastrier (1946) elevated P. langeroni orientalis to species rank, although it continued to betreated as a subspecies of P. langeroni by some authors until quite recently, e.g. Lewis et al. (1974) andTheodor (1958). The males of P. orientalis and P. langeroni may be distinguished by the distribution of theascoids on antennal segments 8-12 (1 per segment in P. orientalis, 2 in P. langeroni) and by subtledifferences in the position of the subapical opening of the aedeagus, which is ventral in P. langeroni andlateral in P. orientalis. This aedeagal character can only be observed in perfectly mounted specimens. Thefemales can be distinguished by the relative lengths of the ascoids on A4 (Figs 21, 22). The male of P. orientalis can be distinguished from that of P. major by the parameres which are broad in 14 R. P. LANE Figs 29-34 29-31, Phlebotomus orientalis. (29) 9, antennal segments 3 and 4; (30) $, pharynx; (31) $,head. (Figs 29-31 Sinai, Tharfet el Qidren.) 32-34, P. major. (32) $, pharynx; (33) $, head; (34) $,spermatheca. (Figs 32-34 S. Sinai.) SANDFLIES OF EGYPT 15 P. orientalis but slender in P. major, and by the aedeagal tip which is pointed in P. orientalis, but blunt androunded in P. major. MATERIAL EXAMINED Egypt. Sinai: 32 cf, 2 $, Upper Wadi Nasb, 14-15.vi.1979 (Y.B.); 10 cT, 9 $, Tharfet el Qidren,13-14.vi.l979(y.fl.). DISTRIBUTION. Chad, Niger, Ethiopia, Kenya, Sudan, Egypt, Yemen, Saudi Arabia (SW.). This is the firstrecord of P. orientalis from Egypt. The record from the southern Sinai is the furthest north P. orientalis has been found. This species shows aclassical East African-Arabian highland distribution, with the important exception of the lowland recordsfrom Sudan (Kirk & Lewis, 1940; Quate, 1964). In western Saudi Arabia, Buttiker & Lewis (1984) foundthat P. orientalis was predominant at altitudes above 1750 m and was absent in the lowlands. P. orientalis is a pernicious man-biter in many parts of its range with biting rates of 108-208 per hour inEthiopia (Ashford, 1974). It is a vector of Le. donovani donovani in the Acacia- Balanites forests ofsouthern Sudan (Hoogstraal & Heyneman, 1969). Ashford (1974) concluded that P. orientalis did nottransmit visceral leishmaniasis regularly in the highlands of Ethiopia (around Arbaya), although importedinfections might be transmitted occasionally. Phlebotomus papatasi (Scopoli)(Figs 35-39) Bibio papatasi Scopoli, 1786: 55 [<j>] Type(s), ITALY (U, Pavia?). Phlebotomus (Phlebotomus) papatasi (Scopoli) [see Lewis (1982) for complex nomenclatural history]. This species is the type-species of Phlebotomus . FEMALE. Pharynx stout, narrowing after posterior bulge; armature consisting of numerous small scales with fringe of fine backward-pointing teeth (Fig. 38). A3 shorter than labrum. Spermatheca with segments (Fig. 37). MALE. Pharynx slender, with numerous small teeth. Paramere with three lobes, dorsal lobe very much longer than broad median lobe. Coxite with small tuft of hairs on plate basally, and another clump of long hairs distally. Style long, slender, with short pointed spines: three terminal, distance between median and subapical spine less than between subapical and terminal spine (Fig. 39). MATERIAL EXAMINED Egypt. As numerous specimens of this common species were examined only localities and collectors aregiven. Alexandria (M.A.R.); Alexandria, El Amriya (Z.A.H.); Bahteem (M.A.R.); Beni Suef (M.A.R.);Birqet Qarun (R. L. Coe); Cairo, El Amirga, caves (M. L. Schmidt); Cairo, El Amiriya, Apartmentbuildings (M. L. Schmidt); Cairo, Mena House (J. Wakeling); Dakhaliya (M.A.R.); Fayoum (R. L. Coe);Imbaba (M.A.R.); Khasm (H. King); Luxor (S. Hirst); Matruh (M.A.R.); Quahsbieh (Z.A.H.); Sinai,Abu Aweigila (Y. Schlein); 30 50'N, 34 20'E, military bunkers (/. Zimmerman); Siwa (Omer-Cooper);Tanta(M.Afl.). DISTRIBUTION. This is the most widespread of any sandfly species, and is distributed from Portugal andMorocco in the west to Bangladesh in the east and from southern U.S.S.R. in the north to Sudan in thesouth. This species has been reported from Egypt by the following: Willcocks (1917); Whittingham & Rook(1923: suggested vector of phlebotomus (= papataci) fever); Khalil (1934: suggested vector of cutaneousleishmaniasisi); Sabin, Philip & Paul (1944: vector of phlebotomus fever virus); Schmidt etal. (1960, 1971:isolation of phlebotomus fever virus from Cairo P. papatasi); Zein el Dine (1972: found in Dakhla Oasisbut not Baharia or Kharga oases); Hafez (1977); Rifaat et al. (1968: no localities given); Hassan (1968a:extensive survey, found throughout country except non-peridomestic areas in Sinai and around Aswan);Kammah (1972: autogeny); Sawaf etal. (1984: Al Agamy, nr Alexandria). During rapid field examination of samples, female P. papatasi may be confused with Sergentomyiachristophersi because of the similarity in the shape and segmentation of the spermathecal capsule, and thepresence of a large number of erect hair sockets on the abdominal tergites. S. christophersi can easily bedistinguished, however, by the presence of cibarial teeth. Schmidt & Schmidt (1963) give a detailed account of the morphological variation in a single populationfrom a suburb of Cairo and conclude that the Egyptian P. papatasi bear a 'marked resemblance to 16 R. P. LANE 41 42 Figs 35-47 35-39, Phlebotomus papatasi. (35) $, head; (36) $, antennal segments 3 and 4; (37) $,spermatheca; (38) $, pharynx; (39) cf , style. (Figs 35-39 Sinai 30 50'N 34 20'E.) 40-47, P. sergenti.(40) $, pharynx; (41, 42) , antennal segments 3 and 4; (43) cf , sperm pump; (44) cf, style; (45) cf,aedeagus; (46) cf , basal process of coxite; (47) $, spermatheca. (Fig. 40 Libya, Jarian Prov.; 41 Libya,Yefran ; 42 Cairo , Maadi , 43-46 Jordan , Mowaggar ; 47 Egypt , Sharkeyia . ) SANDFLIES OF EGYPT 17 specimens from other areas [but] several differences are apparent in the Egyptian material [which]constitute preliminary evidence for infra specific variation'. This is probably the most studied of all sandflies because of its abundance, widespread distribution,anthropophily and peridomestic habits. It is the main vector of cutaneous leishmaniasis to man (Abon-nenc, 1972: 100; Lewis, 1974a; Theodor, 1964; Williams & Coelho, 1978) in many parts of its range. In theMediterranean basin and the Middle East P. papatasi has been proved a vector in several countries, e.g.Israel (Schlein et al. , 1982; Adler & Ber, 1941), and strong circumstantial evidence exists for its vectorialstatus in many other countries, e.g. Saudi Arabia (Nadim, Rashti & Ashi, 1979); Afghanistan (Nadim etal, 1979); Iran (Adler, 1964) and North Africa (review: Dedet, 1979). It is undoubtedly the vector ofcutaneous leishmaniasis in Egypt, but other species may also be involved (e.g. P. sergenti). P. papatasi hasbeen suspected of transmitting visceral leishmaniasis in areas where no other likely vector has been found:Saudi Arabia (Lewis & Buttiker, 1980); Iraq (Abu-Hab & Azawia, 1978; Adler & Theodor, 1957; Sukkar,1972). However, it is not an efficient vector of Le. donovani experimentally but might transmit it rarely(Hoogstraal & Heyneman, 1969; Zahar, 1980: 45, 53). The feeding habits of P. papatasi in Lower Egyptare given by Schmidt & Schmidt (1965). Phlebotomus fever virus was isolated from wild caught P. papatasi collected from human dwellings insuburban Cairo (Schmidt et al., 1960). This, together with the earlier work of Sabin et al. (1944), whichclearly demonstrated that P. papatasi was capable of transmission after the necessary incubation,incriminated P. papatasi as the vector of phlebotomus fever virus in Lower Egypt. Phlebotomus sergenti Parrot (Figs 40-47) Phlebotomus sergenti Parrot, 1917: 564 [cf]; Franca, 1918: 731 [$]. Syntypes cf , ALGERIA (IPA).This species is currently placed in the subgenus Paraphlebotomus Theodor. FEMALE. Pharynx with large scales anteriorly, some produced into long broad spines, scales becomingbroader and flatter posteriorly with hind margin serrated with fine teeth (Fig. 40). Third and fourthantennal segments with slender ascoids almost reaching tip of segment (Figs 41, 42). Spermatheca withequal segments (Fig. 47). MALE. Pharynx tapering after posterior bulge, posterior margin convex; armature series of scales withminute teeth along anterior margins. A3 slender, longer than labrum. Coxite short and stout, less thantwice as long as wide. Coxite lobe slender and relatively long (Fig. 46). Surstyles longer than coxite. MATERIAL EXAMINED Egypt. 1 $, Cairo, Maadi, 7.vii.l943 (Theodor); 1 $, Sharkeyia, 1967 (M.A.R.); Sinai: 1 cf, UpperWadi Nasb, 14-15.vi.1979 (Y.B.); 1 cf , Wadi Baaba, 8-9.vi.1979 (Y.B.). DISTRIBUTION. Mediterranean Basin eastwards through Israel, Iraq, Iran, Afghanistan, Pakistan, India andsouthwards to Saudi Arabia, Yemen and Ethiopian highlands.Theodor (1948) found P. sergenti around Cairo and Rifaat et al. (1968) record it but give no localities. P. sergenti is a proven vector of cutaneous leishmaniasis in the U.S.S.R. (Sergiev, 1979: 206), Crete(Molyneux, 1977), Iraq and India (Abonnenc, 1972). It is a suspected vector of Le. tropica in manycountries including Iran (Nadim & Rashti, 1971) and Yugoslavia (Lupascu et al., 1977). Schlein et al.(1984) found promastigotes in P. sergenti from the Arava in Israel. Sergentomyia adleri (Theodor) (Fig. 48) Phlebotomus adleri Theodor, 1933: 543 [cf $]. Syntypes, GHANA (BMNH).This species is currently placed in the subgenus Sintonius Nitzulescu. FEMALE. Cibarium with 54 vertical teeth in three rows and 17 subequal irregularly spaced horizontal teeth;pigment patch almost as wide as cibarial armature with pronounced anterior projection. Pharynx slender,tapering slightly after posterior bulge, armature only a series of vague transverse striations.MALE. Very similar to S. clydei except that cibarium has two or three rows of vertical teeth, usually 6 in eachrow, and horizontal teeth evenly spaced (not in groups like S. clydei). MATERIAL STUDIEDEgypt. Sinai: 1 $, Feiran 12-13.vi.1979 (Y.B.). 18 R. P. LANE Figs 48-56 48, Sergentomyia adleri $ , cibarium, Sinai, Feiran. 49-53, 5. christophersi. (49) 9 > cibarium;(50) cf , pharynx; (51) d", cibarium; (52) $ , spermatheca; (53) cf , genitalia. (Figs 49-53 Aswan.) 54, 55,S. cincta. (54) $ , pharynx; (55) $ , cibarium. (Figs 54, 55 Siwa Oasis.) 56, S. antennata 9 , pharynx, Sinai30 50'N 34 20'E. SANDFLIES OF EGYPT 19 DISTRIBUTION. Senegal, Ivory Coast, Ghana, Cameroun, Nigeria, Togo, Upper Volta, Chad, Republic ofCentral Africa, Sudan, Egypt, Kenya, Saudi Arabia. This is the first record from Egypt. This species is closely related to 5. clydei and has been considered synonymous by many authors (seediscussion under 5. clydei). Sergentomyia antennata (Newstead) (Figs 56-57) Phlebotomus antennatus Newstead, 1912: 365 [$]. Holotype <j>, GHANA (BMNH).This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with concave row of delicate unequal-sized teeth; pigment patch almost triangular, withanterior projection acutely angled, posterior margin broad, indented. Pharynx stout with armature ofdistinct teeth less than 0-20 pharynx length; posterior margin with shallow notch (Fig. 56).MALE. Cibarium with row of about 16 irregular teeth and small spicules; some vertical teeth also present.Pharynx slender with small triangular teeth. Style stout, less than three times as long as wide; two terminal,two subterminal spines (Fig. 57). MATERIAL EXAMINEDEgypt. Sinai: 1 cf , 2 $, 30 50'N 34 20'E, 19.vii.1983, light-trap in bunkers (J. Zimmerman). DISTRIBUTION. Occurs in a broad band south of the Sahara desert from Ghana through Central AfricanRepublic, East Africa, Yemen, Saudi Arabia to Kuwait. It has also been recorded from Algeria andTunisia. This is the first record of this species from Egypt. This is a very variable taxon and opinions differ on whether it is one or many species. The shape of thepharynx on which most differentiation is made is very variable, e.g. see Lewis & Buttiker (1982: 368). 5. antennata is commonly found outside houses associated with P. papatasi (Buttiker & Lewis, 1983;Lane & Al Taqi, 1983). Schlein etal. (1984) found 'promastigotes, possibly of reptilian leishmanial species'in 7 out of 33 antennata caught together at Arava (Israel). Sergentomyia Christophers! (Sinton)(Figs 49-53) Phlebotomus christophersi Sinton, 1927: 33 [$]. Lectotype $ , PAKISTAN (BMNH), designated by Lewis &Buttiker (1982: 365). This species is currently placed in the subgenus Sintonius. FEMALE. Cibarium with four or five long teeth, many small denticles visible in some specimens; smallpigment patch present (Fig. 49). Pharyngeal armature with a few scale-like folds anteriorly and a series ofvague ridges posteriorly. Spermathecae smooth- walled with approximately eight segments and longindividual ducts (Fig. 52). MALE. Cibarium with several long teeth and some small denticles (Fig. 51). The distinction between thelarge teeth and denticles is less marked in males than females. Pharynx constricted after bulge; armature aseries of indistinct ridges (Fig. 50). Two apical and two subapical spines on style. Paramere simple. Surstylelonger than parameres. Aedeagus slender, tapering from base (Fig. 53). MATERIAL EXAMINED Egypt. Aswan: 55 cf, 60 9, Elephantine Island, 3-5. x. 1983, sticky traps amongst rocks harbouringnumerous geckoes (R.P.L.). DISTRIBUTION. Guinea, Chad, Ethiopia, Sudan, Egypt, North Yemen, Saudi Arabia, Oman, Pakistan.This is the first record of S. christophersi from Egypt. This species is closely related to S. clydei, from which it can be distinguished by the number of cibarialteeth. male female christophersi 3-5 4 clydei 16-26 12 20 R. P. LANE Sergentomyia cincta (Parrot & Martin)(Figs 54, 55) Phlebotomus antennatus var. cinctus Parrot & Martin 1944: 55 [cf $]. Syntypes, DJIBOUTI, SUDAN (IP A), 1 $ syntype (BMNH).Sergentomyia cincta (Parrot & Martin) Theodor, 1958: 38. This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with concave row of 16-18 teeth, lateral teeth slightly smaller than central teeth (Fig.55); pigment patch a rounded triangle, hind margin convex. Pharynx without distinct shoulder, hindmargin straight or slightly convex, posterior teeth only slightly smaller than anterior teeth.MALE. Indistinguishable from S. antennata. MATERIAL EXAMINEDEgypt. 13 $, Siwa Oasis, 6.viii.l935 (Omer-Cooper), det. O. Theodor. DISTRIBUTION. Ghana, Central African Republic, Kenya, Uganda, Sudan, Ethiopia, Djibouti, Egypt. Thisis the first record of cincta from Egypt. This species is considered to be conscpecific with S. antennata by some authors (e.g. Duckhouse & Lewis,1980), but as a distinct but closely related taxon in the S. fallax-group by others (Abonnenc, 1972; Theodor,1958; Davidson, pers. comm.). Sergentomyia clydei (Sinton) Phlebotomus clydei Sinton, 1928ft: 179 [cf $]. Lectotype cf, PAKISTAN (BMNH), designated by Lewis(1967: 42). This species is currently placed in the subgenus Sintonius. FEMALE. Cibarium with row of 13-16 triangular, slender teeth; 25-37 vertical teeth in one or two rows,some teeth may be present centrally so as to produce a short third row; pigment patch triangular, broad, aswide as cibarium. Pharynx tapering markedly after bulge, no armature apparent although a few finetransverse ridges present in some specimens. Spermatheca distinctly segmented (collapsed in all speci-mens, therefore segment number unavailable); individual ducts rugose. MALE. Cibarium with tufts of small denticles posteriorly; pigment patch triangular, not as broad ascibarium. Pharynx more slender than in female and constriction after bulge less obvious. Abdominaltergites 5 and 6 subequal in length. Paramere with beak-like apex, style with 2 apical and 2 subapical spines.Aedeagus slender, pointed. MATERIAL EXAMINEDEgypt. Sinai: 1 cf , 5 $, Tharfet el Qidren, 13-14. vi.1979 (Y.B.). DISTRIBUTION. Algeria, Egypt, Mali, Senegal, Ghana, Niger, Chad, Cameroun, Sudan, Ethiopia (includ-ing Eritrea), Somalia, Kenya, Iraq, U.S.S.R. (Kazakstan, Tajikistan, Turkestan), Saudi Arabia, Kuwait,northern India. S. clydei has not previously been found in Egypt. This species is recorded from southern Sinai and is likely to occur in Upper Egypt because it also isknown from Wadi Haifa in Sudan, near the Egyptian-Sudanese border (Lewis & Kirk, 1954). 5. clydei is very closely related to S. adleri and there has been much discussion on whether they areconspecific. Quate (1964) treats them as separate species but concludes 'I suspect they eventually will beshown to be two forms of the same species'. Lewis & Buttiker (1980) also considered them as separatespecies and stressed the importance of females as a means of identification. Later (Lewis & Buttiker, 1982)they confirmed their earlier proposition by finding distinct specimens representing each species at a singlelocality (Artawiyah, Saudi Arabia). The two species are distinguished by the number of horizontal andvertical teeth on the cibarium, 5. adleri having a higher number than 5. clydei (summarised in Table 3). Theform of S. clydei originally described as latiterga Theodor has the sixth abdominal segment much largerthan the fifth (up to twice the length and width). This form was originally considered a distinct species butwas subsequently synonymised by Theodor & Mesghali (1964: 297). Form latiterga has not been found inEgypt. S. clydei is unusual in that it is one of several species of Sergentomyia which bite man (Quate, 1964), theremaining species of the genus feeding on lizards. 5. clydei has been used to transmit experimentally Le. SANDFLIES OF EGYPT 21Table 3 Differences in the number of cibarial teeth in female S. adleri and S. clydei. Sergentomyia clydei Sergentomyia adleri number of number of number of number of horizontal teeth vertical teeth horizontal teeth vertical teeth n.g. = not given. adleri from lizards to man, causing a transient infection and giving immunity against one form of Le.donovani (Garnham, 1971). Sergentomyia fallax (Parrot)(Figs 58-60) Phlebotomus minutus var. fallax Parrot, 1921a: 37 [cf]. Syntypes, ALGERIA, TUNISIA (IPA).Phlebotomus fallax (Parrot); Parrot, 19216: 99. [Raised to species.] This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with deeply concave row of 16-23 unequal, closely packed, delicate teeth, lateral teethlarger than medial teeth, pigment patch usually ovoid, posterior margin not indented (Fig. 60). Pharynxvariable in shape but typically cordiform with distinct posterior notch, posterior teeth punctiform, verymuch smaller than slender anterior teeth (Fig. 59). Length of A3 0-11 mm (s.d. = 0-009, range 0-083-0-113,n = 10). Spermathecae simple. MALE. Cibarium with concave row of unequal pointed teeth; pigment patch circular or oval. Pharynxslender, narrowing posteriorly, with distinct triangular teeth. Aedeagus broad, finger-shaped. Coxite andstyle long and slender, style more than five times as long as wide (Fig. 58), accessory seta subterminal,distance from setal base to closest terminal spine not greater than length of accessory seta. MATERIAL EXAMINED Egypt. Sinai: 1 cf , 1 $, Feiran, 12-13.vi.1979 (Y.B.); 2 $, Ain Higia, 9-10.vi.1979 (Y.B.); 2 cf , 6 $,TharfetelQidren, 13-14.vi.l979(y..); 3 cf,9 $, Upper WadiNasb, 14-15.vi. 1979 (y.fl.). DISTRIBUTION. From West Africa to eastern Afghanistan, and North Africa from Algeria to Egypt. This isthe first record of this species from Egypt. 5. fallax has been divided into three subspecies: 5. fallax cypriotica Theodor, 5. fallax afghanica Artemievand 5. fallax fallax (Parrot), principally based on absolute size. Thus Lewis (19746) discriminated betweenfemales of 5. fallax cypriotica and 5. fallax fallax on the size of the third antennal segment, 0-07-0-10 mmand 0-12-0-15 mm respectively (males cannot be reliably separated). The specimens from Sinai have amean A3 length of 0.11 and therefore are intermediate between the two 'subspecies'. Furthermore, in thischaracter the Sinai specimens are similar to those from Yemen which also have a A3 mean length of 0-11mm (range 0-09-0-12 mm) (in Lewis, 19746: 192). Until a study is made of this species throughout thewhole of its geographical range the validity of these subspecies must remain in doubt. Sergentomyia minuta (Rondani) Hebotomus minuta Rondani, 1843: 265 [cf]. Type(s) ITALY (depository unknown).This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with a row of long, fine parallel teeth, row slightly convex posteriorly; pigment patchdark, transverse-ovoid, almost as wide as tooth row. Pharynx narrowing markedly after posterior bulge;armature of numerous individual teeth. Spermatheca simple, tubular.MALE. Cibarium with vague row of irregular triangular teeth. Pharynx slender with ridged armature. 22 R. P. LANE Figs 57-64 57, Sergentomyia antennata cf , style, Sinai, 30 50'N 34 20'E. 58-60, S. fallax. (58) cf , style,Sinai, Tharfet el Qidren; (59) $, pharynx; (60) $, cibarium. (Figs 59, 60 Sinai, Upper Wadi Nasb.)61-64, S. palestinensis . (61) $ , cibarium, Aswan; (62) $ , spermatheca, Iraq, Jadizyah; (63) $ , pharynx,Wadi Gaamah; (64) cf , genitalia, Baharia Oasis. SANDFLIES OF EGYPT 23 Aedeagus finger-shaped, tapering. Style with four terminal spines, distance from accessory seta to terminalspine greater than length of seta. MATERIAL EXAMINED Examples of this species from Egypt have not been examined. Some specimens of 'minuta' collected byEflatoon in 1922 are reported by Zein el Dine (1972) to be in the Egyptian Entomological Society, Cairo,but no further details are given. It is possible that these specimens were misidentified because 5. minuta hasnot been collected in any sandfly surveys since. However, as 5. minuta occurs around most of theMediterranean Basin it is equally possible that this species will be found in the more humid areas of coastalLower Egypt. DISTRIBUTION. Portugal, Spain, France, Italy, Malta, Yugoslavia, Greece, Cyprus, Tunisia, Algeria,Morocco, Egypt (?). Two subspecies have been described: S. minuta minuta from continental Europe, with a mean cibarialtooth number of 40; and S. minuta parroti (Adler & Theodor, 1926) from North Africa, with 70 teeth(Theodor, 1958; Rioux & Golvan, 1968). Recent work by Belazzoug et al. (1982) in different ecologicalzones of Algeria has shown that the number of cibarial teeth varies according to certain climatic factors(mainly humidity). Sergentomyia palestinensis (Adler & Theodor)(Figs 61-64) Phlebotomus palestinensis Adler & Theodor, 1926: 64. Holotype $ , PALESTINE (BMNH).Phlebotomus lewisi Parrot 1948: 125. [Synonymised by Lewis & Buttiker, 1982: 370.] This species is currently placed in the subgenus Parrotomyia Theodor. FEMALE. Cibarium with straight row of 18-19 strong, straight, subequal horizontal teeth and a single row ofvertical teeth; pigment patch broad, occupying most of cibarium width and with flattish posterior margin;cornua large (Fig. 61). Pharynx broad with distinct radiating teeth posteriorly (Fig. 63). Spermathecatubular, capsule with thickened walls; collar surrounding spermathecal head well developed and oftenpigmented (Fig. 62). MALE. Cibarium with straight row of 16 delicate horizontal teeth and a single row of vertical teeth. Pharynxwith only slight constriction after post-medial bulge; armature with series of weakly developed scalesposteriorly. Parameres simple, almost as long as surstyles, with hooked tips (Fig. 64). MATERIAL EXAMINED Egypt. 1 9> Aswan, Elephantine Island, sticky trap amongst rock by riverine vegetation, 3-5. x. 1983(R.P.L.); 1 $, 5 cT, Baharia Oasis, 15.v.-7.viii.l971 (K.Z.D.); 2 $, Dakhla Oasis, Rashda, 17.viii.1971(K.Z.D.); 2 $, Sinai, Tharfetel Qidren, 13-14.vi.1979 (Y.B.). DISTRIBUTION. Egypt, Israel, Iraq, Iran, Saudi Arabia, Pakistan, Sudan, Ethiopia. Zein el Dine (1972)recorded this species from Dakhla and Baharia oases. This species may be separated from other species of the subgenus Parrotomyia by the shape and number ofpharyngeal teeth in both sexes, and by the shape of the spermathecal capsule and collar in the female. Sergentomyia schwetzi (Adler, Theodor & Parrot) (Fig. 65-67) Phlebotomus schwetzi Adler, Theodor & Parrot, 1929: 75 [cf $]. Syntypes, CONGO (BMNH).This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with 13-22 horizontal teeth, lateral teeth larger and broader than central (Fig. 65).Pharynx narrowing posteriorly with an armature of broad scales. Spermatheca simple, tubular andsmooth- walled. MALE. Cibarium with concave row of large irregular teeth. Aedeagus finger-like with gentle ventral curve(Fig. 67); style with two apical and two subapical spines at about 0-75 length of style (Fig. 66). MATERIAL EXAMINEDEgypt. 1 $, Aswan, Elephantine Island, 3-5.X.1983, sticky trap (RPL). Figs 65-74 65-67, Sergentomyiaschwetzi. (65) $, cibarium, Aswan; (66)d",genitalia;(67) C?,aedeagus.(Figs 66, 67 S. Sinai.) 68, 69, S. squamipleuris . (68) d", genitalia, Cairo; (69) $, cibarium, Kuwait.70-72, S-. taizi. (70) $, head; (71) $, spermatheca; (72) $, cibarium. (Figs 70-72 Sinai, Mt Katherine.)73, 5. theodori $, cibarium, Gaza. 74, .S. tiberiadis. cf , cibarium. (Fig 74, Aswan.) SANDFLIES OF EGYPT 25 DISTRIBUTION. Widespread in Africa, Yemen, Saudi Arabia. This is the first record of this species fromEgypt. Sergentomyia squamipleuris (Newstead) (Figs 68-69) Phlebotomus squamipleuris Newstead, 1912: 366 [$]. Syntypes, SUDAN (BMNH).This species is currently placed in the subgenus Grassomyia Theodor. FEMALE. Cibarium with convex row of fine, parallel horizontal teeth; undulating row of vertical teeth;pigment patch small, tapering anteriorly; broad, chitinised, transverse band present (Fig. 69). Pharynxtapering abruptly posteriorly, with rows of angular teeth. Spermatheca single capsule with dense coveringof fine ductules. MALE. Cibarium with slightly convex row of minute triangular teeth; faint pigment patch present, almostcircular. Pharynx narrow, tapering posteriorly. Coxite broad, style with two spines terminal, twosubterminal. Paramere thick, bluntly rounded. Aedeagus tapering gently from base to three-quarters oflength, then tapering abruptly (Fig. 68), dorsal margin concave. Surstyles short and thick, as long as coxite. MATERIAL EXAMINEDEgypt. 1 cf , Cairo, l.xi.1910 (Wakeling); 1 $, Aswan, xi.1969 (M.A.R.). DISTRIBUTION. Most of Africa (including Egypt), Israel, Iraq, Iran, Saudi Arabia, Kuwait. Khalil (1934)found this species in Sharqia Governate and Rifaat et at. (1968) recorded it but gave no localities. Sergentomyia taizi (Lewis) (Figs 70-73) Sergentomyia (Sergentomyia) taizi Lewis, 19746: 193 [cf $]. Holotype $, YEMEN (BMNH).This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with nine teeth, two outer teeth broad, scale-like. Pigment patch dark, less than halfwidth of cibarium, with rectangular anterior projection (Fig. 72). Pharynx with numerous fine transverseridges posteriorly, bearing small distinct teeth. In holotype, pharynx tapers posteriorly but not in Sinaispecimens (possibly artefact of preparation). Spermatheca broad, tubular, with apical knob in depression(Fig. 71).MALE. Cibarium with concave row of teeth, outer teeth large, scale-like. Style with all spines terminal. MATERIAL EXAMINEDEgypt. 1 9, Sinai, St Katherina, 15-16.vi.1979 (Y.B.). DISTRIBUTION. Yemen, SW. Saudi Arabia, Egypt. This is the first record of this species from Egypt. This species was described from the mountainous areas of Yemen and only subsequently found in thecontinguous mountain chain in Asir (SW. Saudi Arabia). Sergentomyia theodori (Parrot)(Fig. 73) Phlebotomus minutus (Rondani) sensu Adler & Theodor, 1926: 403 [C?$]. Misidentification.Phlebotomus (Prophlebotomus) theodori Parrot, 1942: 322. Syntypes, PALESTINE: Jericho (BMNH). This species is currently placed in the subgenus Sergentomyia. FEMALE. Cibarium with 20-22 sharply pointed horizontal teeth in concave row, lateral teeth at least twice aslong as central teeth (Fig. 73); pigment patch broad, almost as wide as cibarium and approximatelyrectangular. Pharynx triangular, posterior margin concave or notched; anterior teeth long and slender,posterior teeth appear shorter as they are usually viewed at an acute angle. MALE. Cibarium with concave row of pointed teeth, lateral teeth much longer than tightly packed medialteeth. Pharynx narrowing after posterior bulge (lamp-glass shaped), with fine teeth. Style 4-5 times as longas wide; accessory seta further from apex than length of seta. 26 R. P. LANE MATERIAL EXAMINEDEgypt. 2 cf , 4 $, Gaza, vii.1940 (0. Theodor}. DISTRIBUTION. Yugoslavia, Turkey, Lebanon, Syria, Israel, Cyprus, Egypt, Iraq, Iran, Pakistan, N. India,Afghanistan. This is the first record of this species from Egypt. Sergentomyia tiberiadis (Adler , Theodor & Lourie)(Figs 74-77) Phlebotomus tiberiadis Adler, Theodor & Lourie, 1930: 537 [cf ?]. Lectotype <j>, PALESTINE (BMNH),designated by Lewis & Biittiker (1982: 367). This species is currently placed in the subgenus Sintonius. FEMALE. Head narrow. Cibarium with straight row of about 16 strong, curved, closely packed horizontalteeth; these characteristic teeth may appear straight in some poorly mounted specimens; two or three rowsof distinct vertical teeth; pigment patch broad, almost as wide as tooth row (Fig. 77). Pharynx slender withbarely discernible armature in form of transverse striations. Spermatheca with 6-8 segments.MALE. Cibarium with a row of 12-14 curved horizontal teeth and one or two rows of vertical teeth (Fig. 74);pigment patch vague, almost as wide as tooth row. Pharynx slender, gently narrowing after bulge;armature a series of indistinct ridges (Fig. 75). Paramere with distinct beak apically (Fig. 76). Style with twoapical, two subapical spines. Aedeagus slender, tapering from base. MATERIAL EXAMINED Egypt: 1 cf, Aswan Governate, Aswan, Elephantine Island, 3-5. x. 1983, sticky traps amongst rocks(R.P.L.); 2 cf , Aswan, xi.1967 (M.A.R.); 2 cf , Luxor, 8.xi.l966 (Z.A.H.). DISTRIBUTION. Egypt, Israel, Djibouti, Ethiopia, South & North Yemen, Sudan. Rifaat et al. (1968) recordthis species from Egypt but give no specific locality. This species is easily distinguished from 5. christophersi and S. clydei by the shape and number of cibarialteeth in the male and female. The Oriental subspecies S. tiberiadis pakistanica Artemiev & Saf yanovadiffers from the nominate subspecies in the spermathecal capsule which narrows towards the tip and hasmore segments (9-12), and the lower mean number of cibarial teeth (13 compared to 17-18). Thesubspecies pakistanica is restricted to Pakistan and Afghanistan. 76 Figs 75-77 Sergentomyia tiberiadis. 75, cf, pharynx; 76, paramere; 77, $ , cibarium. (Aswan.) SANDFLIES OF EGYPT 27 Faunal associations Based on the distributions of sandflies collected in Egypt and described above, the fauna can bedivided into three distinct elements. Each element has affinities to faunas in areas outside Egyptand these are discussed below and compared to the distribution of other insects. Elements of the Egyptian sandfly fauna Each of the three faunal elements is composed of a characteristic, although not mutuallyexclusive, group of species. The areas occupied by these three faunal elements are: Mediterranean/Lower Egypt element: coastal Egypt, Siwa Oasis, the Delta and Nile valley as far south as Asyut, and northern Sinai.Upper Egypt: Nile valley to Lake Nasser and the oases of Bahariya, Farafra, Dakhla and Kharga. S. Sinai: the mountain system from Gebel el Igma southwards to Sharm el Sheikh.The fauna of the mountainous southern Sinai is clearly distinct from that of northern Sinai, which has a typical lowland Middle Eastern fauna composed of P. papatasi and 5. antennata. The species representing each of the three faunal elements are listed on pages 4-5. All arerestricted to their component areas except P. papatasi, P. sergenti and 5. palestinensis , which arewidespread in distribution. P. papatasi is the only species found predominantly in peridomestichabitats. Clearly, this division of the fauna into three elements is greatly influenced by the distributionof collecting sites, and at present it is not possible to delimit precisely each area. For example,the Upper Egypt component is based on collecting around Aswan and in the Kharga and Dakhlaoases, but the extent to which the species of this area penetrate northwards along the narrowriverine vegetation to Beni Suef and Faiyum in Lower Egypt is unknown. The faunas of Upperand Lower Egypt (including northern Sinai) are more similar (i.e. have more species incommon) to each other than either is to the southern Sinai fauna, presumably because of theNilotic connection suggested above, and the presence of high mountains in southern Sinai andassociated habitats not available in the rest of Egypt. Several geographically important areasneed to be surveyed to complete the faunistic picture of Egyptian sandflies, particularly theremote wadi systems of the southern Sinai and Upper Egypt, the oases of Siwa and Farafra, andthe Red Sea Hills. The largest and most diverse faunal component in Egypt is that of the southern Sinai, with 12species (6 Phlebotomus , 6 Sergentomyia) compared to the six (1 Phlebotomus, 5 Sergentomyia)found in Upper Egypt and eight (3 Phlebotomus, 5 Sergentomyia) in the Mediterranean/LowerEgypt component. It is not clear which environmental factors have the greatest effect on the distribution ofdifferent sandfly species, although it is clear that some species have a marked association with aparticular habitat. Vegetation affects sandfly distribution indirectly through its effect on thehosts of sandflies, which can be either carnivores (dogs, foxes etc.), herbivores (rodents) or manand domestic animals (goats, poultry). However, the presence of some plants may be importantas sources of sugar meals for the adults. The shade provided by vegetation probably has aminimal direct effect on sandfly distribution as sandflies rarely use plants as resting sites (e.g.tree holes) in arid regions, although vegetation structure clearly has an effect on some sandflyhosts. Temperature has some role in governing distribution because some species found in thehigh mountains of the southern Sinai (e.g. Gebel Katherina which rises to 2637 m) are exposed,during the winter snowfall, to low temperatures which would be fatal to other species. Association of Egyptian fauna with neighbouring areas Throughout this discussion, the affinity of the different elements of the Egyptian fauna with thefaunas of neighbouring areas is measured by the number of species they share. Figs 78-80 28 R. P. LANE summarise the proportion of each faunal element found in seven neighbouring areas. Severalfactors affect these estimates of affinity, including the number of species in a faunal element,differences in the taxonomic status of a species (e.g. S. cintus is treated as a distinct species herebut as a synonym by some authors) and differences in sampling (some areas are better knownthan others). Hence, such estimates, like so many others which take no account of relativeabundance of species or the range of habitats available in an area, can only be used to give ageneral impression of the affinity of faunas. The Southern Sinai is known as an area of endemicity in some groups, e.g. Lepidoptera andOrthoptera, but it does not have any endemic sandflies. However, the Sinai populations of somespecies show subtle morphological differences from populations in other parts of their range,e.g. P. orientalis and P. major. More refined taxonomic techniques may demonstrate substantialdifferences, the lack of endemicity in sandflies may be because the group is well adapted, evencommon, in arid areas and therefore the deserts of the region do not pose a very significantbarrier to dispersal and introgression of populations as they do in most other groups of insects. The sandfly fauna of the southern Sinai shows a marked affinity with the fauna of the ArabianPeninsula (see Lewis & Buttiker, 1982), particularly Asir, Yemen (in Lewis, 1974ft) and, to alesser but still significant extent, to the highlands of eastern Africa (Table 4; Fig. 80). Themountains of the southern Sinai represent an isolated part of a tongue of the AfrotropicalRegion extending northwards to the end of the Rift Valley System in northern Israel (UpperGalilee). The principal species indicating this association are P. arabicus, P. orientalis, P.kazeruni, S. taizi and S. adleri, which dominate the fauna of south-west of the Arabian Peninsulaand are closely related to species in the Afrotropical Region. The absence of the commonPalaearctic P. papatasi further supports this association and the conclusion that sandflies of thesehighland areas are a distinct part of the Afrotropical fauna and not a relict intrusion of thePalaearctic fauna. S. taizi has only been found in the mountains of Yemen (near Ta'izz) and atthe top of Gebel Katherina. Unlike parts of the sandfly fauna of the Arabian Peninsula, the Sinaifauna does not contain any Oriental species, although some species are east European/westAsian (P. alexandri, P. kazeruni). Therefore the southern Sinai does not constitute part of theTriad Zone as defined by Lewis & Buttiker (1980). The sandfly fauna of Upper Egypt has affinities with both the Arabian peninsula and the Sahelof Africa, through species such as S. christophersi, S. clydei, S. adleri, S. schwetzi and 5.tiberiadis (Table 4; Fig. 79). P. papatasi is not typically African and its presence in Upper Egyptis part of a southernly extension of its range along the Nile Valley as far as northern Sudan. Sandflies of Lower Egypt and the northern Sinai are typical of the North African and easternMediterranean faunas (Table 4; Fig. 78). The rarity of P. langeroni has already been noted(p. 11). Its presence in Egypt indicates that it may be widely distributed along the North Africancoast, perhaps in association with outcrops of limestone which are such a feature of the area inEgypt where it is comparatively common. 5. squamipleuris is a member of a species group widelydistributed along a band south of the Sahara, through the Arabian Peninsula and into northernIndia. Its presence in Lower Egypt represents a northerly extension of its range, in contrast tothat postulated above for other species (e.g. P. papatasi). S. squamipleuris is more rigidlyconfined to the riverine vegetation along the Nile than are many other sandflies, where it feedson amphibians. Distribution of insects other than sandflies The comparison of distribution patterns of sandflies with those of other biting flies (Simuliidae,Ceratopogonidae and Culicidae) is germane to understanding sandfly faunal associations,because these other flies also are influenced by the search for hosts. However, other biting fliesare restricted environmentally by the need for free water (running water in Simuliidae) for thedevelopment of the immature stages. The mosquito Culex sinaiticus Kirkpatrick was thought to be confined to southern Sinai, butwas later found in Eritrea and northern Sudan (Red Sea coast around Port Sudan and in centralSudan) (Lewis, 1956), although it has not been found elsewhere in Egypt. A similar situation isfound in Culex arbieeni de Meillon which is known from southern Sinai and the upper slopes of Lower Egypt SANDFLIES OF EGYPT Upper Egypt 29 S. Sinai 100 80 60 40 20 A B C D E F G A B C D E F G A B C D E F G 78 79 80 Figs 78-80 Histograms showing the percentage of the species from Lower Egypt (78), Upper Egypt (79)and southern Sinai (80) which also occur in neighbouring areas. A = east Saudi Arabia + Iran; B = south-west Saudi Arabia + Yemen + Ethiopia; C = Sudan; D =Sahel; E = north Africa; F = Israel; G = east Mediterranean. X = one species as percentage of totalnumber in faunal element. the Marra mountains in west Sudan (Lewis, 1954). In the Simuliidae, only two species are knownfrom Egypt. Simulium griseicolle Becker which breeds in large rivers in Africa, has only beenfound once in Egypt, at Aswan, its type-locality. The second species, Simulium ruficorneMacquart, has a very wide distribution throughout the savannah areas of Africa (south of theSahara), North Africa, Spain and African islands. It is not clear whether it is an Afrotropical orPalaearctic species but it probably represents a species complex. Specimens of S. ruficorne fromFeiran Oasis in southern Sinai are similar in their pupal gill structure (Crosskey , pers. comm.) tospecimens from Israel and the Hejaz. Unfortunately very little is known of the EgyptianCeratopogonidae, particularly Culicoides, but the known fauna, based on collecting in theDelta, is Palaearctic. Thus the faunal associations of other biting flies are broadly concordantwith those of the Egyptian sandflies, showing connections between Sinai and the AfrotropicalRegion, and Lower Egypt and the Palaearctic Region. As noted above, southern Sinai is a known area of endemicity in other insect orders. Forexample, Uvarov (1929) studied a small collection of Orthoptera from southern Sinai and drewthe following conclusions: 'in general composition the known fauna of the Sinai peninsula isclearly a local division of the Palaearctic eremian fauna ... it possesses certain features of itsown [endemics]. The Sinai peninsula must be regarded as one of the dry mountainous centreswhere an ancient Mediterranean fauna survives, where the present eremian fauna was born anddeveloped and from where it spread over the whole great desert belts . . . the Sinai fauna has aclose similarity, perhaps an intimate relation to that of Arabia' [although at the time the latterfauna was very poorly known]. In a comprehensive account of the biogeography of Arabian butterflies, Larsen (1984) makesrelatively little comment on the Egyptian fauna other than that it is typical of the 'Palaearcticeremic zone'. However, there are several endemic species of butterflies in the southern Sinaimassif. In conclusion, the sandfly fauna of Egypt is exceptional, compared to the sandfly fauna ofother countries, in being so clearly divided into Afrotropical and Palaearctic elements. WhetherEgypt and surrounding areas have always been a point of faunal exchange of sandflies betweenthe two zoogeographic regions, with the Nile valley and the mountains of the Red Sea coast andSinai acting as major dispersal routes, is not clear. Lewis (1982) has suggested that the genusPhlebotomus is Palaearctic in origin and subsequently extended southwards into Africa duringthe pluvial periods, presumably through the Nile and Rift valleys. However, the Sahara desert 30 R. P. LANE Table 4 Comparison of the species shared between each of the three components of the Egyptian sandflyfauna and the faunas of neighbouring areas. has not always been such a barrier to dispersal and possibly, in an arid-adapted group such as thesandflies, substantial dispersal may have taken place either through or around the Sahara. Amore comprehensive study of sandflies throughout the whole of the Eremic Zone (Morocco toAfghanistan) is required to answer these questions fully. Acknowledgements I am grateful to: staff of the Research and Training Centre on Vectors of Disease at Ain Shams University,Cairo, particularly Dr S. El Said, Professor B. El Sawaf and Dr J. Beier, for assistance during visits toEgypt; Dr Y. Braverman, Israel, for valuable material from Sinai; the UNDP/World Bank/WHO SpecialProgramme for Research and Training in Tropical Diseases, and the National Institutes of Health(National Institute for Allergy and Infectious Diseases) for financial assistance; and my wife for theillustrations. SANDFLIES OF EGYPT 31 References Abonnenc, E. 1972. Les Phlebotomes de la region ethiopienne (Diptera, Psychodidae). Memoires de I' Office de la Recherche Scientifique et Technique Outre-Mer no. 55: 1-289.Abu-hab, J. K. & Azawi, B. M. al 1978. Trials to artificially infect Phlebotomus papatasi (Diptera: Phlebotomidae) with the causitive agent of kala-azar. Proceedings of the Third Pest Control Conference, Ain Shams University, Cairo: 403-407. Adler, S. 1964. Leishmania. Advances in Parasitology 2: 35-96.Adler, S. & Ber, M. 1941 . The transmission of Leishmania tropica by the bite of P. papatasi. Indian Journal of Medical Research 29: 803-809.Adler, S. & Theodor, O. 1925. The experimental transmission of cutaneous leishmaniasis to man from Phlebotomus papatasi. Annals of Tropical Medicine and Parasitology 19: 365-371.1926. On a collection of Phlebotomus species of the minutus group. Annals of Tropical Medicine and Parasitology 21: 61-68.1929. The distribution of sandflies and leishmaniasis in Palestine, Syria and Mesopotamia. Annals of Tropical Medicine and Parasitology 23: 269-306. 1957. Transmission of disease agents by Phlebotomine sandflies. Annual Review of Entomology 2: 203-226.Adler, S., Theodor, O. & Lourie, E. M. 1930. On sandflies from Persia and Palestine. Bulletin of Entomological Research 21: 529-539.Adler, S., Theodor, O. & Parrot, L. 1929. Phlebotomes du Congo Beige. Revue de Zoologie et de Botanique Africaines 18: 72-89.Annandale, N. 1910. The Indian species of Papataci fly (Phlebotomus). Records of the Indian Museum 4: 35-52.Artemiev, M. M. 1978. Sandflies (Diptera, Psychodidae, Phlebotominae) of Afghanistan, iv + 87 pp. Kabul.1980. A revision of sandflies of the subgenus Adlerius (Diptera, Phlebotominae. Phlebotomus). [In Russian.] Zoologicheskii Zhurnal 59: 1177-1192.Ashford, R. W. 1974. Sandflies (Diptera: Phlebotomidae) from Ethiopia. Taxonomic and biological notes. Journal of Medical Entomology 11: 605-616.Bassili, W. R., Morsy, T. A. & Michael, S. A. 1983. Specificity and sensitivity of indirect haemagglutination in patients with cutaneous leishmaniasis. Journal of the Egyptian Society of Parasitology 13: 291- 295.Belazzoug, S., Mahzoul, D., Addadi, K. & Dedet, J.-P. 1982. Sergentomyia minuta parroti (Adler & Theodor, 1927) en Algerie (Diptera: Psychodidae). Annales de Parasitologie, Humaine etcomparee 57: 621-630.Biittiker, W. & Lewis, D. J. 1984. Insects of Saudi Arabia. Some ecological aspects of Saudi Arabian Phlebotomine sandflies (Diptera: Psychodidae). Fauna of Saudi Arabia 5: 479-530.Cahill, K. M. 1965. Leishmanin skin testing in Africa and the Middle East. 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Index Invalid names are in italics; principal references are in bold. adleri (Leishmania) 21 adleri (Sergentomyia) 5, 6, 17, 20, 21,28,30Adlerius 7, 9afghanica 21alexandri 4, 5, 9, 28, 30antennata 5, 6, 7, 19, 20, 26, 30arabicus4, 5,7,28, 30 Christophers! 5, 6, 15, 19, 26, 28, 30cincta5,6, 7,20,28clydei 5, 6, 17, 19, 20, 26, 28, 30cutaneous leishmaniasis 1, 2, 9, 15, 17cypriotica 21 davidi 7 donovani (Leishmania) 1, 2, 13, 15,17,21 fallax5,6,7,20,21,30Grassomyia 25halepensis 9kazeruni4,5,9,28, 30 krimensis 13 langeroni 3, 4, 5, 6, 11, 13, 28, 30 Larroussius 11, 12, 13 latiterga 20 leishmaniasis 1, 2, 3, 9, 11, 13, 15, 17 lewisi 23 longiductus 9 major (Leishmania), 1, 2 major (Phlebotomus) 6, 12, 15, 28, 30minuta3,5,6,7,21,25,30 Naqben sp. 7neglectus 12 orientalis5,6, 13, 28,30 pakistanica 26 palestinensis 3, 5, 6, 7, 23, 27, 30 papatasi 2, 3, 5, 6, 11, 15, 19, 27, 28, 30 Paraphlebotomus 9, 17parroti 23Parrotomyia 23perniciosus 11 Phlebotomus 2, 5, 27, 29phlebotomus fever 1, 15, 17 rift valley fever 2 sandfly fever 1schwetzi5,6,7,23,28,30sergenti2, 3,5,6,9, 17,27,30Sergentomyia 5, 27Sergentomyia (s.str.) 19, 20, 21, 23, 25 Sinai form 5, 6, 11simici 9 Sintonius 17, 19, 20, 26squamipleuris 3, 5, 6, 25, 28, 30syriacus 12 taizi5,6,7,25,28,30theodori5,6,7,25,30tiberiadis3,5,6,26,28,30tropica (Leishmania) 2, 17 visceral leishmaniasis 2, 3, 9, 11, 13,15,17 wenyoni 13 Some other works on Diptera The following papers were published in the Bulletin of the British Museum (Natural History)Entomology series. Copies are still available: each is virtually a monograph and complete initself with paper covers. Vol. 4 No. 3 The mosquitoes of Arabia. I. P. F. Mattingly and K. L. Knight. 1956. pp. 89-141. 4 textfigures 3. 70 Vol. 7 No. 9 A study of the New Zealand Chironomidae (Diptera, Nematocera). P. Freeman. 1959.pp. 393-437. 1 plate, 6 text figures 3.30 Vol. 21 No. 6 A revision of the Ethiopian species of the tribe Notiphilini (Diptera: Ephydridae). B. H.Cogan. 1968. pp. 279-365. 1 plate, 96 text figures 6.25 Vol. 24 No. 7 Studies of African Asilidae (Diptera) I. Asilidae of the Congo Basin. H. Oldroyd. 1970.pp. 207-334. 96 text figures 8.95 Vol. 26 No. 8 A revision of Francis Walker's types of North American Empididae (Diptera). K. G. V.Smith. 1971. pp. 345-370. 3 plates, 16 text figures 2.25 Vol. 30 No. 7 A revision of the genus Passeromyia Rodhain and Villeneuve (Diptera: Muscidae). A. C.Pont. 1974. pp. 339-372. 9 text figures 2.40 Vol. 36 No. 1 A review of the Rhinophoridae (Diptera), and a revision of the Afrotropical species.R. W. Crosskey. 1979. pp. 1-66. 46 figs 8.35 Vol. 37 No. 6 The phlebotomine sandflies (Diptera: Psychodidae) of the Oriental Region. D. J. Lewis1978, 128 pp including 345 references, 273 text figures and 13 maps 17.85 Supplement No. 14 A re-classification of the Simuliidae (Diptera) of Africa and its islands. R. W.Crosskey. 1969. pp. 195. 1 plate. 331 text figs. 11 maps 9.50 Supplement No. 19 A revisionary classification of the Rutiliini (Diptera: Tachinidae), with keys to thedescribed species. R. W. Crosskey. 1973. pp. 167. 109 text figs, 1 map 9.50 Supplement No. 21 A conspectus of the Tachinidae (Diptera) of Australia, including keys to thesupraspecific taxa and taxonomic and host catalogues. R. W. Crosskey. 1973. pp. 221. 95 text figs 11.50 Supplement No. 26 A taxonomic conspectus of the Tachinidae (Diptera) of the Oriental Region.R. W. Crosskey. 1976. pp. 1-357. 150 text figs 35.00 Complete catalogue of BM (NH) Bulletins free on request. Titles to be published in Volume 52 The Sandflies of Egypt (Diptera: Phlebotominae) by R. P. Lane Fungus moths: a review of the Scardiineas (Lepidoptera: Tineidae) by G. S. Robinson A revision of the European Agathidinae (Hymenoptera: Braconidae) by G. E. J. Nixon A key to the Afrotropical genera of Eucoilidae (Hymenoptena) with a revision of certain genera byJ. Quinland Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester BRr .(N PRESENTED Bulletin of the British Museum (Natural History) Fungus moths: a review of theScardiinae (Lepidoptera: Tineidae) G. S. Robinson Entomology series Vol52 No 2 24 April 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History) ,Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1986 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06016 3 ISSN 0524-6431 Entomology series Vol52No2pp37-181British Museum (Natural History)Cromwell RoadLondon SW7 5BD Issued 24 April 1 986 Fungus moths: a review of the Scardiinae(Lepidoptera: Tineidae) Gaden S. Robinson Department of Entomology, British Museum (Natural History), Cromwell Road, LondonSW7 5BD StiM ! hU ARY Contents Synopsis 37 Introduction 37 Techniques 40 Check-list of Scardiinae 41 Taxa excluded from the Scardiinae 43 Classification and characters - but phylogeny? 43 Objectives 43 Methods 43 Supraspecific groups (OTUs) classified 46 Characters used in classification 46 Results 50 Geographical distribution 58 Abbreviations 60 Acknowledgements 60 Scardiinae Eyer 61 References 135 Index to hostplants and fungi 180 Index to Lepidoptera 180 Synopsis This paper gives an account of the subfamily Scardiinae, the fungus moths. The group is redefined and 111species in 23 genera are ascribed to it. Its previous taxonomic history and its biological and morphologicalcharacteristics are reviewed. Le Quesne tests show homoplasy to be rampant in the Scardiinae. Numericalphylogenetic methods are used with the aim of deriving a phylogenetic classification; their results arecontrasted with those from various phenetic methods. Character-compatibility analysis is used to pinpoint'robust' characters with minimal homoplasy. Groups defined by such characters are found to be also'robust', being revealed by most of the analytical methods employed. The shortest optimized Wagner treeis adopted as the classification but the limited phylogenetic implications of this classification are stressed.The biogeography of the group is reviewed. Keys are provided to genera and species. Eight genera aredescribed as new and one genus is recalled from synonymy. Twenty-six new species are described and 18species are placed in new generic combinations; four new synonymies are established. Introduction The subfamily Scardiinae includes more than one hundred species of generally large and robusttineid moths. While the smallest species have a wingspan of only about 12 mm, the largestScardiinae are among the biggest ditrysian Microlepidoptera, with wingspans of up to 60 mm.Despite their size, however, Scardiinae are not conspicuous. It is the biology of the Scardiinae that sets them apart, with one exception, from all othersubfamilies of Lepidoptera. The larvae feed in either the fruiting body (sporophore) ofpersistent (hard) bracket-fungi, or in the wood of dead or moribund trees that has beenpermeated by the hyphae of such fungi. The traces left by feeding scardiinae larvae arecharacteristic - ramifying accretions of loosely webbed coarse frass forming a cover for surfaceand sub-surface feeding, usually combined with similar volcano-shaped accretions of frass or Bull. Br. Mus. nat. Hist. (Ent.) 52 (2): 37-181 Issued 24 April 1986 38 GADEN S. ROBINSON loose mounds above deeper tunnels. Larvae pupate just below the surface of the food-substrate.The mobile pupa is protruded half-way out of the substrate (using two rows of backward-pointedspines on each abdominal segment to force its exit) before it ruptures and the adult emerges. Fungus-feeding Lepidoptera Examples of fungivory are found throughout the Lepidoptera and the reader is referred to therecent review by Rawlins (1984) for further details. Within the Tineoidea fungivores are foundpossibly among the species of Compsoctena (Eriocottidae). Lichenivory, a form of fungivory, iswidespread among Psychidae. The biology of one of the eight species of Arrhenophanidae isknown and it is a fungivore, feeding on 'Polyporus sp.' (Costa Lima, 1945; Becker, pers.comm.). Within the Tineidae fungivory is found in the subfamilies Hieroxestinae, Tineinae andMeessiinae (Lawrence & Powell, 1969); many other species of Meessiinae are lichenivores.However, apart from the Scardiinae, the tineid subfamily Nemapogoninae is the only othersubfamily of Lepidoptera that feeds exclusively on fungi. Nemapogoninae are virtually restrictedto the Holarctic region whereas Scardiinae are found also in both the Old and New Worldtropics. There is no obvious division of niche between the two groups but there is a considerablesize difference, most adult Nemapogoninae having a wingspan of less than 18 mm, with thelarvae being correspondingly smaller than those of Scardiinae. General morphology The wing patterns of most genera of Scardiinae are cryptic, taking the form of a mottled creamand brown pattern resembling moss or tree-bark. Other genera have disruptive patterns: thebrown fore wing with a pale apex and dorsum. A few genera have a bronze or golden forewingground-colour blotched with dark brown. The Morophaga bucephala-group is quite atypical inhaving a conspicuously white forewing ground-colour with bold purple-brown markings. InAmorophaga the cryptic wing-pattern is corrupted by being drawn into longitudinal streaks. The scardiine head carries the erect scales on frons and vertex typical of most tineid groups.The maxillary palpus is 5-segmented, elongate and folded in most species. The labial palpus iscylindrical, the second segment with only lateral bristles rather than lateral and terminal bristlesas in, for example, the Tineinae. The antennal pecten may be very large with up to about 40strong, bristle-like scales. Most species are broad-winged and have all veins present. However, some have narrowfore wings and in these one or two pairs of veins are stalked. The narrowing of the forewing ismost extreme in Diataga, in which two pairs of veins are stalked. Externally, however, themorphology of adult Scardiinae is conservative and, with the exception of the enlarged antennalpecten (not found in all scardiinae genera but not known from any other tineid), no adultcharacter or combination of characters can be used to define the group. It is the larva thatprovides the single defining character of the Scardiinae (see below). History of classification of the Scardiinae Although Eyer (1924) is the first recorded user of the family-group name Scardiinae, theassociation of typical scardiine taxa was well-established by earlier authors (e.g., for the NewWorld, by Dietz, 1905, and Walsingham, 1914). No significant usage of the family-group nameoccurred again, however, until Hinton (1955; 1956) emphasized the characteristic for definingScardia (i.e., Scardia s.l: Morophaga + Scardia + Morophagoides) and its allies as a discretegroup. The larvae of Scardiinae differ from those of all other Tineoidea in that the prothorax bearsonly two L-group setae. All other known tineid larvae have three L-group setae. The generalityof this important character within the Scardiinae is, however, uncertain as larvae of onlyMorophaga, Diataga, Scardia, Daviscardia and Morophagoides are known. The characteristiclarval setation was first noted by Gerasimov (1937) in Scardia. Fracker (1915) had earlier figuredthe larva of Scardia anatomella but with three prothoracic L-group setae. However, this was FUNGUS MOTHS 39 probably an error as later workers (Kuznetzov, 1941; Hinton, 1956) found only two. Kuznetzov(1941), finding a fossil larva with only two L-group setae, searched for this character amongother Tineidae and confirmed it in Morophaga choragella (as boleti), Scardia boletella, S.anatomella and Daviscardia coloradella. Strangely, his mention en passant of coloradella is theonly reference to the setal pattern of this species and confirms the bisetose prothoracic L-groupin Daviscardia. Amsel (1952) figured the genitalia of the few European species of Scardiinae. Petersen (1957)placed together all those Palaearctic species of Scardiinae then recognized and described thegenitalia. He later (Petersen, 1959; 1960) described two new species and revised those known tohim. Zagulajev (1965; 1966; 1968) described a number of new species from the Caucasus and fromthe easternmost provinces of the U.S.S.R. and later (Zagulajev, 1973) combined and expandedhis earlier work into a full-scale monograph of the Palaearctic Scardiinae . This work remains theprimary reference to the group in this area and no attempt is made here to duplicate its content.Gozmany & Vari (1973), in their monograph of the Afrotropical Tineidae, also recognized thesubfamily as discrete and revised the Afrotropical species. Their concept of the Scardiinae was,however, broader than that of other authors. They included Afrocelestis , Ateliotum andPelecystola, genera placed by other authors as Meessiinae (Afrocelestis) and Myrmecozelinae(Ateliotum and Pelecystola), placements that I endorse. They also erected a further subfamily,the Tinissinae. Three years later I revised this diverse group of predominantly south-east Asianspecies (Robinson, I916a) and later proposed a phytogeny for the group, drawing attention to itsmorphological similarities to the Scardiinae and suggesting that the Scardiinae might beparaphyletic with respect to the Tinissinae (Robinson, 1981). This suggestion is supported in thepresent work and the Tinissinae are synonymized with the Scardiinae below. Davis (1983) has provided an up-to-date list of the North American Scardiinae, including newsynonymy: work in progress includes redescription of the North American Scardiinae (Davis,pers. comm.). The action of Capue (1971) in erecting the tribe Semeolonchini for the single genusSemeoloncha is based on no more than Gozmany's original description of the genus (Gozmany,1968); no characteristics are offered to define the tribe and it is not placed in any highertaxonomic category. Fossil 'Scardiinae' Kuznetzov (1941), in his revision of the Lepidoptera of Baltic amber (~40 million years BP),described four genera (Scardiites, Proscardiites, Palaeoscardiites and Glessoscardia) and alliedthem with Scardia (and thus with the Scardiinae). Zagulajev (1966; 1973) agreed with Kuznet-zov's placement. The first three genera were described from single adults, the last from a larva.The possibility that Scardiites is a monotrysian is suggested by a row of postfrenular bristles inKuznetzov's illustration, the shape of the apex of the abdomen of Proscardiites suggests thepossibility of its being a yponomeutoid (although the length of the maxillary palpi suggestsotherwise), and the outline of the valva of Palaeoscardiites as figured by Kuznetzov isreminiscent of Compsoctena (Eriocottidae). There are no characters figured or cited byKuznetzov to suggest affinity of Scardiites, Proscardiites or Palaeoscardiites to the Scardiinae.However, the larva, Glessoscardia, could well be a Scardiinae although from its size (only 9 mmlong) it is unlikely to be mature. Kuznetzov noted that it had only two prothoracic L-group('prestigmatal') setae. General biology A summary of the biologies of the Palaearctic Scardiinae (including novel information on somespecies, and figures of the larva and pupa of Scardia boletella) has been given by Zagulajev(1973). Powell ([1968]) and Lawrence & Powell (1969) bred Lepidoptera from a variety of deadwood and fungus samples in California. This rearing programme yielded a large number of newhost records for North American Scardiinae. Much earlier, Walsingham (1882) described his 40 GADEN S. ROBINSON experiences rearing Scardia anatomella in Oregon. Hinton (1956) has noted the few detaileddescriptions of the biology of European species. Petersen (1969) has summarized records of thebiology of Morophaga choragella (as boleti}. More recently, Moriuti (1976) has described thelife history of Morophagoides moriutii (as ussuriensis) and figured the larva and pupa. Scardiinae are only rarely encountered in collections made at light (Robinson, 1981 - Tinissd).Sixteen nights' collecting at light in Brunei in 1982, for example, yielded 260 Tineidae of about60 species: only six species and eight individuals were Scardiinae and all but one of those wereTinissa (Robinson, 1984). Species of Scardiinae of which large numbers are available havealmost invariably been bred. Zagulajev (1973) has collected numbers of Morophagoidesiranensis swarming at dusk near fungus-riddled trees and Fletcher (1933) collected Morophagacremnarcha (q.v.) under similar circumstances. Other specimens have been collected at rest byday close to a putative food-substrate. Scope and aims of the present study The present study is intended to complement existing studies of the Palaearctic Scardiinae,notably that of Zagulajev (1973), by integrating the comparatively unstudied species and generaof the Old World tropics and of the New World with the Palaearctic taxa. In so doing, it has beennecessary to revise substantially the existing classification of the Scardiinae and, to some extent,to adapt the restricted definition of the sub-family to accommodate the morphological differ-ences of the extra-Palaearctic taxa. The bibliography provided is limited in that references providing only minimal information(such as mention in checklists, or parochial distribution records of common European species)have been omitted. Zagulajev (1973) has collected together many references to Europeanspecies and others are given by Petersen (see bibliography): they are not reiterated here. This review is based primarily upon the collections of the British Museum (Natural History)but most specimens in the National Museum of Natural History, Washington, the University ofCalifornia, Berkeley, and the Zoological Institute of the Academy of Sciences, Leningrad, havealso been studied. Material in other institutions was requested selectively. Specimens inLeningrad and California and some of those in Washington were examined early on in theproject and, at that time, not all the characters finally adopted for use in classification had beenrecognized. Where observations are deficient, this is stated. Attempts to resolve a phylogeny for the Scardiinae have highlighted the problems caused bythe high degree of homoplasy encountered. Gauld & Mound (1982) have drawn attention to thisas a feature of several other insect groups and emphasized the difficulty in providing anacceptable generic classification in these circumstances. The data set obtained in the course of this study has been used in the exploration of techniquesfor numerical classification by phenetic and cladistic methods using a microcomputer. Terminology The external and genital morphology of the Scardiinae is unexceptional and the terminologyused here follows that of Common (1970) and Klots (1956). Further discussion of certain of thefeatures examined is given under 'Characters used in classification'. In the male genitalia theuncus is composed of a pair of soft lobes: these may be more or less strongly sclerotized, may befused with each other and/or with the tegumen, or may be widely separated. In my previouspaper (Robinson, 1981) I adopted Kuznetzov & Stekolnikov's (1976; 1977) term 'tegumenlobes' for these, but their clear differentiation from the tegumen in all taxa examined, and theirevident homology with the uncus of, for example, the Tineinae, make this term inappropriateand I here revert to my earlier (19760) usage of 'uncus lobes'. Techniques Pinned adult specimens were examined at magnifications of x 6 to x 50 in artificial light using a FUNGUS MOTHS 41 Wild M5 microscope. Venation, if not visible in oblique light, was examined by transmitted lightafter the wing had been wetted with toluene. Size measurements given were made with a ruler and are those of a specimen set inconventional fashion; other comparative measurements were made using an eyepiece graticuleon a Wild M5 microscope. The variety of male genital types in the Scardiinae required the utilization of a variety oftechniques to display the characters adequately. General methods for the preparation ofgenitalia slides have been described by Clarke (1941) and Robinson (1976b). For certainScardiinae it was necessary to employ the technique of cutting the vinculum laterally andunrolling the genital armature as described previously for Tinissa (Robinson, 1976a). In othergroups the internal surface of the uncus lobes and valvae could be displayed simply by folding thevalvae ventrad and forward and splaying them slightly. In other groups it was necessary todissect away one valva and mount it separately. Chlorazol Black E (Azo Black) was employed asa stain for preparations made early on in this study but this was superseded by Mercurochrome. Inow prefer the latter stain as it is more controllable and consistent. Although not applicable tothis study, it also offers better photomicrographic results. Preparations were examined and drawn using a Wild M5 stereoscopic microscope withcamera lucida drawing attachment. More detailed examination was carried out with a LeitzOrtholux microscope using phase-contrast and magnifications to x600. Phase-contrast wasparticularly useful in identifying microtrichia lining the vesica or the ductus bursae, in identify-ing the sensilla on the eighth tergite of females, and in locating and counting the setae on theabdominal sternites. Numerical analysis was carried out using a Commodore 64 microcomputer. Programs writtenby the author are in CBM BASIC; copies are available on request. When a draft of this paper was completed, David Swofford's computer program 'PAUP'(Phylogenetic Analysis Using Parsimony - Illinois Natural History Survey) became available,running on the Rutherford Laboratory's IBM Multiprocessor. This program was used withmultiple parsimony and global branch-swapping options. Check-list of Scardiinae MOROPHAGOIDESPetersen, 1957 iranens/sPetersen, 1960 ussuriens/s(Caradja, 1920) moriutiisp. n. berkeleyella (Powell, [1968]) burkerella (Busck, 1904) gracilis (Walsingham, 1907)caryophylella (Busck, 1908)errandella (Busck, 1908) montium (Walsingham, 1914) pythium sp. n. nimbiferum sp. n. inlinu (Walsingham, 1914) MONTESCARDIA Amsel, 1952tessulatellus (Zeller, 1846) /fiirenzov/(Zagulajev, 1966) comb. n.fuscofasciella (Chambers, 1875) comb. n.pravatella (Busck, 1908) BYTHOCRA TES Meyrick ,1919drosocjc/aMeyrick, 1919 DAVISCARDIAgen. n. coloradella (Dietz, 1905) comb. n. radulella sp. n. bimendella (Zeller, 1863) comb. n. beckerisp. n. luctuosa (Walsingham, 1914) comb. n. tnackieisp. n. bicolorella sp. n. sp. A lupulella sp. n. hypocritella sp. n. SCARDIA Treitschke, 1830 Agarica Sodoffsky, 1837Fernaldia Grote, 1881Duomitella Koshantschikov, 1923anatomella (Grote, 1881) comb. rev. fiskeella Busck, 1908assamensissp. n.amurensisZagulajev, 1965allenisp. n.boletella (F., 1794) nom. rev. $boleti (F., 1798) (unjustified emenda-tion) polypori (Esper, [1804])relicta (Koshantschikov, 1923)caucasj'caZagulajev, 1965 42 PERILICMETIS Meyrick, 1932ftdip/aca Meyrick, 1932ft MOSCARDIA gen. n. re/iite/is( Meyrick, 1922ft) comb. n.var/iasp. n. GENTINGIA gen. n.hollowayisp. n. SEMEOLONCHA Gozmany, 1968penicillata Gozmany , 1968 CR AN AODES Meyrick, 1919stereopa Meyrick, 1919oroya sp. n.seguesfrafa Meyrick, 1926. PECTINISCARDIA gen. n. prosfy//as (Meyrick, 1927) comb. n. HORMANTRIS Meyrick, 1927asfraga/opa Meyrick, 1927 CNISMORECTIS Meyrick, 1936c/ior/f/ca Meyrick, 1936 MINISCARDIA gen. n. mininwllu (Busck, 1914) comb. n.sp. A NECROSCARDIA gen. n. funeratella (Zeller, 1863) comb. n.morticina sp. n. TINISSA Walker, 1864 Polymnestra Meyrick, 1927polystacta (Meyrick, 1918) perilithios (Meyrick, 1927)cultellata (Gozmany & Vari, 1973)yaloma Robinson, 1981phrictodes Meyrick, 1910philippinensis Robinson, 1976apo/yse/na Zagulajev, 1972albipuncta Robinson, 1976ains/gfl/s Zagulajev, 1972eumetrota Meyrick, 1926pa/modes Meyrick, 1917con voluta Robinson, 1976achalcites Robinson, I976aaraucar/ae Robinson, 1976aamboinensis Robinson, 1976acinerascens Meyrick, 1910krakatoa Robinson, 1976adistracta Meyrick, 1916errantia Robinson, 1976atorvella torvella Walker, 1864torvella mysorensis Robinson, 19760classeyi Robinson, 1981 GADEN S. ROBINSON ruwenzorica Gozmany, 1966 spaniastra Meyrick, 1932 poliophasma Bradley, 1965 dohertyi Robinson, 1976a transversella (Walker, 1864) indica Robinson, 1976a bakeri Robinson, 1976a baliomicta Meyrick, 1928 rigida Meyrick, 1910 heterograpta Meyrick, 1928chloroplocama Meyrick, 1938 parallels Robinson, 1976a goliath Robinson, 1976a kidukaroka Robinson, 19760 insularia Robinson, 1976o chaotica Robinson, 1976a SCARDIELLA gen. n. approximated (Dietz, 1905) comb. n. AFROSCARDIA gen. n. capnochalca (Meyrick, 1932ft) comb. n. AMOROPHAGA Zagulajev, 1968rosemariae sp. n. cryptophori (Clarke, 1940) comb. n.hyrcanica Zagulajev, 1968japonicasp. n. DIATAGAWahingham, 1914/eptosce/esWalsingham, 1914frustraminis sp. n. brasiliensis (Zagulajev, 1966) comb. n.compsacma Meyrick, 1919levidensis sp. n.mercennaria sp. n.direpta sp. n. MOROPHAGA Herrich-Schaffer, 1853Atabyria Snellen, 1884Osphretica Meyrick, 1910Microscardia Amsel, 1952 feucepAa/a-group cremnarcha (Meyrick, 1932ft) comb. n.nigrocapitella Petersen, 1959 syn. n. bucephala (Snellen, 1884) chomatias (Meyrick, 1910)rotundata (Matsumura, 1931) soror Gozmany, 1965 vadonella (Viette, 1954) more//i/s-group more/7us(Duponchel, 1838) fungicolella Dumont, 1930 syn. n. s/sfrafa-groupborneensissp. n.sistrata (Meyrick, 1916) comb. n. FUNGUS MOTHS 43 formosana sp. n. fasciculata sp. n. iriomotensis sp. n. kobella sp. n. clonodes- group Taxa incertae sedis donodes (Meyrick, 1893) comb. n. LEPTOZANCLA Meyrick, 1920 porphyrea (Lower, 903) syn. n. ^^ / 192Q maculosa (Diakonoff, 1949) syn. n. PH1LAGR1AS Meyrick, 1932 gen. rev. cflorageiia-group ze/otica (Meyrick, 1932) comb. rev. choragella ([Denis & Schiffermuller], 1775) boleti (F. , 1777) SCARDIA s 1 fungella (Thunberg 1794) ' ttataw|esM rick> 1894 mediella (Hubner, 1796) ,/ ., ino/< , n rz i m/:*: pnaretrodes Meyrick, 1934 hyrcanella Zagulajev, 1966 . ., . , J ini/l ' . , ? r, } , . 1ft/: , isf/mMe//aBusck, 1914talyshensis Zagulajev, 1966 Taxa excluded from the Scardiinae Morophaga angulatella Walsingham, 1897: 168. Transferred to Acrolophus Poey (Tineidae: Acrolophinae) comb. n.Morophaga hirsutevestita Walsingham, 1897: 167. Transferred to Acrolophus Poey (Tineidae: Acrolophinae) comb. n.Scardia conglomerata Meyrick, 19220: 12. Transferred to Paradystis Meyrick (Tineidae: Tineinae) comb. n.Scardia lochaea Meyrick, 1911: 307. Transferred tentatively to Afrocelestis Gozmany (Tineidae: Meessiinae) comb. n.Scardia saccharata Meyrick, 1914: 205. Transferred to Narycia Stephens s.l. (Psychidae) comb. n. Classification and characters - but phylogeny? Objectives The object of this section is to derive a phylogenetic classification of the Scardiinae. Alternativesto the now widely-used 'Hennigian' phylogenetic classifications are those derived by pheneticmethods. Arguments for and against different procedures of classification are rife and the readeris referred to the plethora of recent textbooks and papers on the subject for further enlighten-ment. It was obvious by inspection that considerable homoplasy occurred within the Scardiinae andwithin the data matrix compiled for this exercise (Table 1). No 'hand-made' phylogeny could bederived. Techniques for measuring the degree of homoplasy and assessing character 'robust-ness' were therefore used to pinpoint 'reliable' characters and to clean up the data matrix byremoving characters that exhibited high levels of homoplasy. Both phenetic and cladisticmethods were employed, to gain familiarity with and to evaluate the techniques, and to contrastthe results and assess the degree of consensus between them. Methods Character compatibility and homoplasy The more characters in a matrix of character-scores that are mutually compatible, the moreeasily may a minimum-length rooted or unrooted tree be resolved. By current convention theminimum-length rooted tree is adopted as the hypothesis of phylogeny of a group. The mostparsimonious tree (that with fewest steps) will be shorter and have a greater probability of beingunique the greater the degree of character compatibility within the matrix. Le Quesne (1969; 1972; 1979) has shown the incompatibility of a pair of binary-statecharacters to involve the presence of all four possible combinations of character-states. This testfor incompatibility has become known colloquially as Le Quesne's test. Character incompatibil- 44 GADEN S. ROBINSON Table 1 Character matrix for Scardiinae: 26 OTUs and 54 characters. DATA MATRIXTAXON 1TAXON 2TAXON 3TAXONTAXONTAXONTAXONTAXONTAXONTAXON 10TAXON 11TAXON 12TAXON 13TAXON 14TAXON 15TAXON UTAXON 17TAXON ISTAXON 19TAXON 20TAXON 21TAXON 22TAXON 23TAXON 24TAXON 25TAXON 26 1 1 1 10000010000000001 10000010000000000001 101 1 1 100000000 APOMORPHIES: 14 111100000100000000111010001101100001010100000100000000 APOMORPHIES: 17 1111 00000 1 000000000 1101 000 1 1 000000 110101 00 11001 1000000 APQMORPH I ES : 1 8 111100000100001000111010000000000000000000000000000000 APOMORPHIES: 10 1 1 110000 1 1 000 1 0000 1110 10000 1 1 000000000 10011011 1001000 APOMORPHIES: 20 111100000100010000011010100000000000000000000111001000 APOMORPHIES: 14 1111 00000 1 000000 1011101111 100000000 101011 1101 100 1 0000 APOMORPHIES: 23 1111 0000 1 1 0000 1 0000 1 1000 1 1 10000000 1111110100111001 000 APOMORPH I ES : 23 101011000010000001000000000000100010000001000000000000 APOMORPHIES: 9 1 0001000000 1 000 111101 00000 1 0000 11001110100010111001010 APOMORPHIES: 2 1 100011000001000111001000001000011001110110110111001010 APOMORPHIES: 23 10101 1000001001010000001 1 10000000000000001000000000010 APOMORPHIES: 12 1101 0000000 1 0000 110011011001 0000000 1111100110111 00 1 1 APOMORPHIES: 23 110111 00000 1 1 00 110011011 00 10000000 1011110110111 0000 1 APOMORPHIES: 25 1 1 000000000 1 00 1 1 1 1 0000000 1 1 0000000 1 1 1 1 1 1 1 1 1 1 1 1 1 APOMORPH I ES : 22 000 1 000000000000 101001011 1 00000000 1011 0000 1 00 10000000 APOMORPH I ES : 12 000 1 000000000 1 00 1 000 1 1 00 1 00 1 1 000000 1 000 1 000000 1 1 000000 APOMORPHIES: 1 2 00 1 1 000000000 1 00 1 000 1 1 00 1101 1000000 1 000 1 1 00000 1 1 000000 APOMORPH I ES : 15 0010000000000 1001 1 1 00000000001 00000000000000000000000 APOMORPHIES: 6 00 1101 0000000 1 000 1111 00000 1 000 1 000000000010000000 1 0000 APOMORPH I ES : 12 001000000000010001001000000000100000010110110011000111 APOMORPHIES: 15 00000 1 0000 1 00 1 00001 00 1 1000 1 000000 1 1 000 1 000 1011100001 APOMORPH I ES : 15 1 0110 1 1 1 1 110010 1 010111000010 A P M R P H I E S : ib 11111111 00000 1 0000000000 1 00 1 000 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 APOMORPHIES: 30 1 1 000 1 1 00000 1 00000000000000000 10101111100111011110111 APOMORPHIES : 22 10101 10001 00 1 000 1 1 000 1 00 1 00000000 1011101 00 1 1 100000 1 1 APOMORPHIES: 20 ity (= Le Quesne test failure) is due to homoplasy, itself the result either of character reversal orconvergence. Gauld & Mound (1982) have. discussed the high frequency of Le Quesne testfailure found in data from some insect groups and found it to be concordant with difficultiesencountered in classification at generic level in those groups. Le Quesne's test as conceived originally (1969; 1972) is independent of any definition ofcharacter-state polarity; i.e., it is not necessary to define or know which state is the primitive(plesiomorphic) and which the derived (apomorphic) before performing the test. However, acorollary to the test must be that, in cases in which polarity is defined, the occurrence of onlythree of the four possible combinations when the missing fourth combination is that of the twoplesiomorphic states (00) must also be grounds for failure (Le Quesne, 1979; Gauld, pers.comm.). Le Quesne (1972) has described a series of methods for the selection of mutually compatiblecharacters. He has embodied his methods I-IV in a program (for the Commodore 64 micro-computer) and I am most grateful to him for providing this and other programs for use ininvestigating the properties of the Scardiinae data set. The methods are designed to yield a suiteof mutually compatible characters (cliques) by progressive elimination of those characters withthe highest frequency of Le Quesne test failure (I), by selection of those characters compatiblewith that having the lowest rate of failure (II), by selection of those characters compatible withthat having the lowest coefficient of character-state randomness (CCSR - see below) (III), andby selection of those characters compatible with that having the lowest value of normal deviate(see Le Quesne, 1972) (IV). Lack of certainty as to the polarity of certain character-states within the Scardiinae makes itappropriate to adopt in the first instance the uncritical version of Le Quesne's test that assumesunspecified character-polarity. It is this version that is used in Le Quesne's own program.However, combining this test with the more critical test in which failure requires the pairwisecomparison to give only the 01, 10 and 11 combinations may point up those characters that are,by comparison with all others, most probably allotted the wrong polarity. Thus in a table of LeQuesne test comparisons (e.g. , Gauld & Mound, 1982) failure may be denoted by an 'X', a passby ' ' and failure that is consequent upon a '00' pairing in a hypothetical taxon by a '*'. Acharacter that exhibits a large number of '*'s in the table has probably had the polarity of its FUNGUS MOTHS 45 constituent states assessed wrongly. Once this check has been made, the version of Le Quesne'stest that assumes defined character-polarity is the appropriate one to use. In addition, therefore, to Le Quesne's program utilizing his clique-indicating methods I-IV, afurther program was developed which makes this comparison and draws the table. The programperforms two further kinds of tests. The value of P (the probability of Le Quesne test failure by apair of characters assuming random distribution of the character-states among the species) iscalculated using the modified formula of Le Quesne (1979) and used to derive the coefficient ofcharacter-state randomness (CCSR) (Le Quesne, 1972). This is the ratio (expressed as apercentage) of the observed frequency of Le Quesne test failure to the expected frequency (fromsummation of P) - i.e., the proportion of test failures that would occur if the character-stateswere distributed at random among the species. It is a measure of the 'robustness' of a character.Characters exhibiting little homoplasy have a low CCSR; those with a high level of homoplasyhave a high CCSR. This measure may be used to provide a weight for each character: thereciprocal of the observed/expected ratio has been utilized for this purpose in weighted centroidcluster analysis. The program provides also, for each character, a list of the other characters thatare compatible with it and with each other: this list of cliques of wholly-compatible charactersmay be used to find, by inspection, the largest clique or cliques. The method is complementary toLe Quesne's clique-indicating methods I-IV. The individual test failures of pairs of characters may be examined individually to ascertainthose character-scores for particular taxa that contribute most to the overall homoplasy of thedata set. Pairs of characters that fail the test are examined in turn: if a single combination of '01','10' or '11' occurs, the taxon responsible for the single failure-causing score is allocated one 'hit'for each character on a taxa x characters matrix. This procedure is repeated for eachcharacter-combination that fails the Le Quesne test. The resulting table resembles the originaldata matrix but the individual character-scores are replaced by the total 'hits' for which eachcharacter-score was responsible. This technique pinpoints, by high numbers of 'hits', wronglycoded data and unique instances of homoplasy. Some falsehoods occur, however; the scores ofcharacter-states that occur in only two taxa tend to attract 'hits'. This is because Le Quesne testfailure of characters with such a distribution of states must involve each member of the pair ofstates (say, T) forming a different combination (thus, '1-0' and '1-1') with the states of theopposed character in order for the test to fail. Both combinations, being unique within the test,attract a 'hit'. Comparison of the sums of the 'hit' scores for the taxa will indicate those taxa inwhich homoplasy is concentrated. Clustering and tree-building Camin-Sokal trees are rooted cladograms constructed on the assumption that a descendentcharacter-state cannot revert to an ancestral character-state - i.e. , that homoplasy is always theresult of convergence. By contrast, in the construction of a rooted or unrooted Wagner treeeither convergence or reversal may be invoked (whichever is the more parsimonious - i.e.,requiring fewer steps) to account for the distribution of character-states among taxa (Sneath &Sokal, 1973). Camin-Sokal trees were derived using I. M. White's 'Hennig' program adapted forthe Commodore 64. Wagner trees, optimized by branch swapping and the testing of all possibletrees for up to nine OTUs, were constructed using a suite of programs written by W. J. LeQuesne. At the very end of the project, Swofford's sophisticated mainframe program 'PAUP'was utilized, employing options for multiple parsimony and optimization by global branch-swapping. Five methods of cluster analysis have been used to generate dendrograms as alternatives tocladistic analysis using Wagner and Camin-Sokal techniques. Three of these methods utilized acommon matrix of Gower similarity coefficients to yield dendrograms by, respectively, single-link, complete link and average-link cluster-analysis. Analysis was performed using programsprovided by W. J. Le Quesne. The fourth and fifth methods were, respectively, weighted andunweighted centroid cluster analysis using the technique described by Robinson (1975) butmodified for binary-state data. Characters were each weighted by the reciprocal of their ratio ofobserved to expected Le Quesne test failure as described above. 46 GADEN S. ROBINSON Supraspecific groups (OTUs) classified Twenty-six groups (OTUs) were subjected to numerical analysis. Twenty-one of these aregroups represented in the subsequent taxonomic treatment as either genera or species-groups:the remaining five OTUs are the two species of Moscardia, Morophaga borneensis, and the OldWorld and New World species of Morophagoides. Each taxon was allocated an identifyingnumber, as follows. 1 : Morophaga bucephala-group 2: Morophaga borneensis 3: Morophaga morellus-group (monobasic) 4: Morophaga choragella-group 5: Morophaga sistrata-group 6: Morophaga clonodes-group (monobasic) 7: Amorophaga 8: Diataga 9: Miniscardia10: Moscardia renitens11: Moscardia varna12: Necroscardia13: Scardia 14: Daviscardia 15: Cranaodes 16: Montescardia 17: Morophagoides (old World species) 18: Morophagoides (new World species) 19: Scardiella (monobasic) 20: Afroscardia (monobasic) 21: Pectiniscardia (monobasic) 22: Bythocrates (monobasic) 23: Perilicmetis (monobasic) 24: Cnismorectis (monobasic) 25: Hormantris (monobasic) 26: Gentingia (monobasic) The identifying number for each taxon is the same throughout all analyses and is that used inTable 1. Semeoloncha has been omitted from phenetic and cladistic assessment. The twospecimens are in poor condition and, in both, most of the abdomen has been discarded duringthe preparation of the genitalia slide. As a result, the set of character scores for Semeoloncha isso incomplete as to render analysis impossible. Tinissa has also been omitted: a cladisticclassification of this undoubtedly monophyletic group has already been proposed (Robinson,1981) and the marked homoplasy within the genus makes its scoring particularly difficult. It isplaced here as the sister-group of Necroscardia, sharing with that genus an extraordinarilymodified juxta (Robinson, 1976a; 1981). Characters used in classification Extraordinary conservatism in the external structure of Scardiinae contrasts sharply with themorphological diversity of the genitalia which is just as remarkable as that described previouslyfor Tinissa (Robinson, 1976a). Characters were selected for inclusion that were stable withingenera or species-groups defined on the basis of close similarity in genital structure. Two distinctgroups of characters were used. The first group (characters 1-30; 50-54) includes 'conventional'characters of the head, legs, wings and genitalia (including modification of the terminalabdominal segment). Genital characters were restricted to those of males (with the exception of29, a synapomorphy of all Morophagoides species) as the inclusion of characters of the femalegenitalia would have reduced substantially the number of taxa upon which analyses could beperformed. Thirty-five binary-state 'conventional' characters were selected for inclusion in theanalyses. In addition to the thirty-five 'conventional' characters, it was found that the number anddisposition of sensory setae on the ventral surface of the abdomen, visible under phase contrastat a magnification of x 150 or x 250, provided a second group of 'unconventional' characters thatwere of use in classification at generic and subgeneric level. Accordingly, a further 19binary-state characters (31-49) were added to the matrix of character-scores (Table 1). The arrangement of setae on the abdominal sternites follows a basic plan. A cluster of setae(more than 30 in most Scardiinae) occupies a medial position close to the posterior margin ofsternite 2. Sternites 3 to 7 each carry a pair of widely separated setae close to the anterior margin.The region between these 'fixed' setae may, in some taxa, contain an arc of irregularly-placedaccessory setae: their presence or absence and their concentration (although not their precisenumber) on a particular sternite appears to be reasonably consistent at species-group level. FUNGUS MOTHS 47 Accessory setae are present on sternite 7 in all Scardiinae. Sternite 8 does not possess the 'fixed'pair in all taxa but, in most taxa, accessory setae are present, scattered close to the anteriormargin. Amorophaga does not possess the 'fixed pair' and only a single pair of accessory setae ispresent. These, from their position, do not represent the 'fixed pair'. Setae are frequently lost during the process of cleaning the abdomen. However, the sockets ofsensory setae are easily distinguished from the surrounding scale-bases when viewed underphase-contrast and can be counted. Character-polarities were estimated initially by outgroup comparison with other subfamiliesof Tineidae, notably the Nemapogoninae, Myrmecozelinae (including Hapsifera), Tineinae andHieroxestinae. Inappropriate comparisons or those yielding ambiguous results were reinforcedby in-group comparisons in the light of preliminary classifications made using phenetic andCamin-Sokal methods. Further refinements to polarity estimation were made in the light ofcharacter-compatibility analysis using Le Quesne's test and its derivatives. In the absence of detailed information from other groups, the polarities of setal characters areinevitably tentative. The characters used in classification are as follows. The presumed apomorphic state isdescribed first, and the plesiomorphic state second. Some comments on the characters, theirvariation, occurrence, and significance are included. A measure of the homoplasy of eachcharacter (the coefficient of character-state randomness - CCSR) is given. The 14 charactersthat survive one or more of Le Quesne's elimination procedures (Le Quesne, 1972) areindicated. 1: Antenna of male lacking cilia from the dorsal surface/cilia developed on the wholecircumference of each flagellar segment. The states of this character are clear-cut and thearrangement of cilia is easily visible under low magnification if the specimen is rotated along theaxis of the flagellum. CCSR = 68%. 2: Antenna of male lacking scales on the ventral surface of the flagellum/complete circumfer-ence of each flagellar segment scaled. The states of this character are not as clear-cut as those ofthe preceding character, nor as easy to observe. Some loss of scales seems to occur naturally inworn specimens; however, the apomorphic state of this character is usually recognizable as atleast a narrow bare line on the undersurface of the flagellum. In some groups it is very obvious,and in Scardia the bare ventral area accommodates a distinct swelling of each segment (seeZagulajev, 1973: figs 5, 6). CCSR = 81%. 3: Antennal cilia of male longer than l-5x the flagellar diameter/cilia shorter than l-5x theflagellar diameter. In practice, this division is realistic although occasional 'borderline cases'have been noted at individual level. Both states have, however, been noted in different speciesof Daviscardia: the plesiomorphic state has been entered in the data matrix for that genus.CCSR = 68%. Survives Le Quesne elimination procedure 4. 4: Antennal pecten with more than 15 bristles/pecten with fewer than 15 bristles. Thischaracter has been discussed previously with respect to Tinissa (Robinson, 1981). The differ-ence between the typical tineid pecten with sparse and slender bristles and the apomorphic'brush' of sometimes more than 40 flattened bristles present in some Scardiinae is obvious infresh specimens. However, the pecten is subject to rapid attrition during life and the confirma-tion of the plesiomorphic state usually requires preparation of the antenna for microscopicexamination in transmitted light. Pectens with large numbers of bristles do occur in a few otherscattered genera in the Tineidae. For example, Dasyses (Hapsiferinae) has a pecten of about 30filamentous bristles but these are so thin that the pecten never appears brush-like as in theScardiinae. CCSR = 85%. 5: Interocular index greater than 1-0/interocular index equal to or less than 1-0. This index isthe vertical diameter of the eye divided by the interocular distance as measured at about the levelof the tentorial pits. Although the measurement is best made on a microscopic preparation of thehead, the paucity of specimens of most groups necessitated its being taken from pinnedspecimens in this study. The range observed in the Scardiinae is narrow -from about 0-7 to 1-3 - 48 GADEN S. ROBINSON and continuous; it is unlikely that this character is of value for classification in this group. CCSR= 92%. 6: Maxillary palpus with fewer than five segments (usually three, occasionally four segments:palpus usually short, only reaching the base of the second segment of the labial palpus/maxillarypalpus 5-segmented (usually elongate, reaching the apex of the second segment of the labialpalpus). Resolution of the segmentation of the maxillary palpus is difficult without preparationof the head for examination in transmitted light. This has not been possible for many groups andthe character-scores given here should be treated with some caution. Only a single example of anelongate three-segmented palpus has been noted - in one species of Daviscardia\ however, inMontescardia the maxillary palpus, although five-segmented, is as short as in most groups with athree-segmented palpus (Zagulajev, 1973: fig. 4). CCSR = 93%. 7: Pilifers absent/pilifers present. CCSR = 23%. Survives Le Quesne elimination procedures1-4. 8: Second segment of labial palpus exceptionally slender and elongate, longer than width ofhead/second segment shorter than width of head (see 54). CCSR = 23%. Survives Le Quesneelimination procedures 1-4. 9: Outer mid-tibial spur and outer proximal hind tibial spur short, less than 0-4 the length ofthe corresponding inner spur/outer spurs of normal length, about 0-5-0-7 the length of the innerspurs. While consistent and conspicuous, the significance of this character is uncertain. Shortspurs are found in Diataga, the Morophaga sistrata-group and in Morophagafasciculata. CCSR= 37%. Survives Le Quesne elimination procedure 2. 10: Forewing veins R 3 and R 4 fused, stalked or approximated at base (connate)//? 3 and R 4separate. This character is subject to some intraspecific variation and both stalking and basalapproximation of the two veins may occur in the same species. However, fusion is a rareindividual variation. CCSR = 73%. 11: Forewing veins M 3 and CuA\ stalked or approximated at base (connate)/M 3 and CuA\separate. Some individual variation occurs in this character: fusion has not been noted. Thestalking of M 2 and M 3 is an autapomorphy of Diataga. CCSR = 58%. 12: Forewing pattern composed of dark purple-brown ground-colour with contrasting whitishterminal and posterior fasciae (continuous in many taxa)/forewing pattern irrorate and cryptic,resembling moss or speckled tree-bark, or otherwise modified. The state of this character hasbeen scored as apomorphic for Moscardia renitens but is uncertain in this species as the onlyknown specimen is very worn. Several of the New World species of Morophagoides andCranaodes oroya have wing-patterns that approach the apomorphic state of this character butthese are considered to be cases of convergence. CCSR = 71%. 13: Forewing pattern composed of light greenish bronze ground-colour with bold purple-brown marginal spots/fore wing pattern irrorate and cryptic, resembling moss or speckledtree-bark, or otherwise modified (see 12). The apomorphic state of this character is shared bySemeoloncha as well as Cranaodes (highly modified in oroya} and Gentingia. The somewhatsimilar, strongly-marked pattern of certain species of Morophagoides (notably iulina) isconsidered to be a convergent development: it is not strictly comparable as the dark blotches arenot at or arising from the wing margin. CCSR = 45%. Survives Le Quesne eliminationprocedures \-$. 14: Coremata absent from eighth abdominal segment/pair of lateral coremata present ineighth abdominal segment. This character was found to be highly unstable in Tinissa (Robinson,1981) and a similar high level of homoplasy seems to occur in the remainder of the Scardiinae.CCSR = 87%. 15: Pair of elongate and tubular apodemes arising from sides of eighth sternite and associatedwith coremata/eighth sternite without apodemes or with flap-like apodemes arising fromposterior corners. CCSR = 73%. 16: Pair of short, flap-like apodemes arising from posterior corners of eighth sternite andassociated with coremata/eighth sternite without apodemes or with elongate rod-like apodemesarising from sides. The apomorphic state of this character is expressed in Semeoloncha (notincluded in analyses) as well as in both species of Moscardia and in Cranaodes. CCSR = 38%. FUNGUS MOTHS 49 17: Uncus complex , modified usually by strong sclerotization and the development of spines ormore elaborate processes/uncus simple, a pair of soft and setose lobes. CCSR = 79%.18: Uncus lobes fused with tegumen/uncus lobes separated from tegumen by at least a narrowand recognizable line of flexion. CCSR = 80%. 19: Tegumen broken mid-dorsally by at least a narrow membranous suture-line/tegumenfused dorsally and thus complete. CCSR = 90%. 20: Valva with setose lobe arising from internal membrane close to base of costa andfrequently associated (most markedly in Diataga) with a strongly developed transtilla/valvalacking basal setose lobe. The occurrence of an internal lobe in Scardiella and Afroscardia isconsidered to be a convergent development unrelated to the occurrence of a similar lobe inMorophaga, Diataga and Amorophaga. CCSR = 64%. Survives Le Quesne eliminationprocedures 1, 2 and 4. 21 : Valva with ventroapical processes (or with apex modified) forming a functional hook/apexof valva smoothly rounded and/or incapable of use as a hook. CCSR = 84%.22: Valvae fused ventrally to form a single movable complex/valvae separate, each capable ofat least some independent movement. In the case of, for example, Scardia, fusion of the valvaeinvolves also the juxta which is interposed between them: cases such as this have beeninterpreted as being the apomorphic state of the character. CCSR = 72%.23: Valva with deep longitudinal cleft effectively dividing valva into dorsal and ventralarms/valva entire. The apomorphic state of this character is distinctive. It occurs in all species ofAmorophaga, and in all species of Morophaga except the bucephala-group; however, the cleft isill-defined and shallow in Morophaga morella. The emarginate dorsoapical region of the valva inthe bucephala-group may in fact be a very shallow cleft but has not been scored as such. CCSR =49%. Survives Le Quesne elimination procedures 1, 3 and 4. 24: Saccus broad, wider than long/saccus elongate, longer than wide. CCSR = 80%.25: Juxta complex, bearing or forming processes, in some groups partially fused with andforming a functional part of the valvae, or apparently entirely fused with the valvae (e.g. , somespecies of Daviscardia)/ 'juxta simple, an ovate and usually ill-defined sclerite on the ventralsurface of the anellus. CCSR = 89%. 26: Juxta divided medially into two arms or lobes/juxta entire. CCSR = 82%.27: Vesica lacking spicular cornuti/vesica with spicular cornuti. Spicular cornuti are conspi-cuously developed in Daviscardia. In some other groups they are very small and sparse andshould not be confused with the microtrichia lining the ductus ejaculatorius of all Tineidae. Insome taxa the cornuti are minute, resembling microtrichia and probably of similar structure.There is, however, a clear demarcation between the vesica and the ductus ejaculatorius in allspecimens examined. CCSR = 86%. 28 : Aedeagus smooth-surf aced/aedeagus with spinose carinae . The homology of, for example ,the minute spicular carinae of Daviscardia luctuosa with the comparatively enormous carinae ofMoscardia is uncertain. CCSR = 91%. 29: Female with pair of strong pocket-shaped signa in bursa copulatrix/bursa copulatrix eitherwithout signa or signa not of this form. The apomorphic state of this character is a synapomorphyof all Morophagoides species. CCSR = 8%. Survives Le Quesne elimination procedures1-4. 30: Apices of otherwise simple pair of uncus lobes fused together and tapered to form a minutehook/apices of uncus lobes not so modified. The apomorphic state of this character is asynapomorphy for the Morophaga sistrata-group. CCSR = 31%. Survives Le Quesne elimina-tion procedures 1, 3 and 4. 31: Medial group of setae on sternite 2 with fewer than 15 pairs/medial group with more than15 pairs of setae. CCSR = 71%. 32: Longitudinal line of microtrichia present, running through medial group of setae onsternite 2/microtrichia absent. CCSR = 66%. 33: Microtrichia minute, requiring magnification of x600 and phase-contrast for definition/microtrichia comparatively large, visible at x!50. CCSR = 28%. Survives Le Quesne elimina-tion procedures 1-4. 50 GADEN S. ROBINSON 34: Medial group of setae broad, scattered across more than 0-4 width of sternite/medial groupcompact, less than 04 width of sternite. CCSR = 58%. 35: Medial group with isolated outlying pair of setae/medial group uniformly spaced. CCSR =98%. 36: Sternite 3 with accessory setae present/accessory setae absent. CCSR = 79%.37: Sternite 3 with more than 4 pairs of accessory setae/fewer than 4 pairs of accessory setae.CCSR = 82%. 38: Sternite 4 with accessory setae present/accessory setae absent. CCSR = 77% .39: Sternite 4 with 3 or more pairs of accessory setae/fewer than 3 pairs of accessory setae.CCSR = 82%. 40: Sternite 5 with accessory setae present/accessory setae absent. CCSR = 70%.41: Sternite 5 with 3 or more pairs of accessory setae/fewer than 3 pairs of accessory setae.CCSR = 97%. 42: Sternite 6 with accessory setae absent/with accessory setae present. CCSR = 51%.Survives Le Quesne elimination procedure 3. 43: Sternite 6 with 3 or more pairs of accessory setae/fewer than 3 pairs of accessory setae.CCSR = 77%. 44: Sternite 7 with 3 or more pairs of accessory setae/fewer than 3 pairs of accessory setae.CCSR = 76%. 45: Sternite 7 with 10 or more pairs of accessory setae/with fewer than 10 accessory setae.CCSR = 68% 46: Sternite 8 lacking 'fixed pair' of setae/'fixed pair' present. CCSR = 81% .47: Sternite 8 with accessory setae present/accessory setae absent. CCSR = 80%.48: Sternite 8 with more than one pair of accessory setae/one pair or no accessory setaepresent. CCSR = 80%. 49: Sternite 8 with 10 or more pairs of accessory setae/with fewer than 10 pairs of accessorysetae. CCSR = 61%. 50: Sockets of ventral abdominal scales scattered randomly/sockets arranged in irregulartransverse pattern. CCSR = 91%. 51: Sockets of ventral abdominal scales arranged in regular and strongly linear transversepattern/sockets arranged in irregular transverse pattern. CCSR = 74%.52: Ground-colour of fore wing gold or pale bronze/ground-colour not thus, usually cream oroff-white (but dark brown in Afroscardia) . CCSR = 64%. Survives Le Quesne eliminationprocedures 1-4. 53: Scales of second segment of labial palpus short, giving the segment a cylindrical appear-ance/scales elongate, particularly distally, giving the segment a triangular appearance in lateralview. CCSR = 71%. Survives Le Quesne elimination procedures 1 and 2.54: Labial palpus slender, longer than 0-9 the width of the head/labial palpus shorter andstouter, of typical tineid length. The apomorphic state of this character is expressed byCnismorectis , Hormantris and Pectiniscardia but is not as pronounced in the latter species as inthe other two. The extreme length of the palpus shared by these two is scored as character 8.CCSR = 51%. Survives Le Quesne elimination procedures 1-4. Results Investigation ofhomoplasy Of the 2862 possible combinations of pairs of characters in the data set for the Scardiinae (Table1), 1772 (61-9%) fail the critical version of Le Quesne's test. The number of test failuresexpected if scores were randomly distributed is 2495 and the overall coefficient of character-staterandomness is thus 71%, indicating overall a high degree of homoplasy. Homoplasy appears tobe spread throughout the data: only 14 of the 54 characters survived one or more of Le Quesne'selimination procedures (see above). Cliques of entirely compatible characters are smaller: twocliques of 10 characters, five cliques of nine, and larger numbers of smaller cliques were found.The seven largest cliques include 25 different characters. Only one of the characters that FUNGUS MOTHS 51 survived Le Quesne's elimination procedures (23) is not included among them. These cliques ofcharacters are too small to provide an adequate classification. Only three characters exhibited Le Quesne test failures consequent upon a '00' pairing in ahypothetical taxon. These were characters 3 (two occurrences), 36 and 47 (one occurrenceeach). 'Hits' of particular scores for particular OTUs were not markedly concentrated in particularcharacters: totalled hits per character ranged from six (characters 3 and 29) to 37 (character 35),43 characters attracting between 10 and 24 hits. The distribution of total hits among OTUs wasmuch more erratic, however, ranging from two (OTU 14) to 110 (OTU 24). Five OTUs attracted72 or more hits: these were Diataga, Necroscardia, Cranaodes, Cnismorectis and Gentingia. Theremaining OTUs attracted 58 or fewer hits, seven attracting 10 or less. This observation suggeststhat certain OTUs may be 'reservoirs of homoplasy' and that their position within a classificationshould be treated with appropriate caution. Nineteen individual character-scores (all apomorphic occurrences) attracted 10 or more hits.These were : OTU 1 - character 45 ; OTU 2 - character 30 ; OTU 5 - characters 9 and 30 ; OTU 8 -characters 9 and 15; OTU 9 - character 11; OTU 12 - characters 12, 15 and 26; OTU 15 -characters 13, 16 and 49; OTU 20 - character 50; OTU 21 - character 54; OTU 24 - character 35;OTU 26 - characters 10, 13 and 34. Characters 9, 13 and 30 have only a pair of apomorphicoccurrences among the OTUs and their attraction of large numbers of hits may be artefactual(see above) or might suggest that the occurrences are the result of convergence. The latterexplanation is unconvincing in the case of character 30 (the fusion and hooking of the apex of theuncus in the Morophaga sistrata-group and M. borneensis). However, the majority of theremaining scores attracting high numbers of hits would, by inspection and in the light ofsubsequent analyses, appear to be genuine instances of homoplasy. The degree of homoplasy of the data set of the Scardiinae is such that attempts to definecliques of compatible characters, 'refine' the data set by removal of characters with high CCSRsor by deletion of obvious convergent occurrences of apomorphic scores resulted either in almostcomplete deletion of the matrix or in little improvement in the overall level of homoplasy.Accordingly, trees were prepared from the data set in Table 1 without modification, using thetechniques described above. Cladistic classification The shortest cladogram found using a Camin-Sokal technique (no reversals permitted) involved217 character-steps (Fig. 1); the shortest trees found originally using a Wagner method(convergence and reversal permitted) on a microcomputer involved 199 steps (Figs 2,3). Moresophisticated subsequent analysis using Swofford's PAUP program gave eight trees (rooted toan all-zero 'ancestor') of closely similar topology, each of 198 steps. The first tree is illustrated inFig. 3a: the consensus tree of all eight variants is indicated by the dashed lines. Characterchanges on all branches are indicated; reversals (character-state changes from T to '0') areindicated by an asterisk. Topological variation within the eight 198-step trees involves inter-change of Morophaga borneensis and M. morellus, interchange of Morophagoides and Monies-car dia, and interchange of four groups: Scardia + Daviscardia, Gentingia, Moscardia +Perilicmetis , and Cnismorectis + Hormantris + Pectiniscardia + Cranaodes. The topology of the right-hand halves of the 199- and 198-step trees is the same. However, thesequencing of the left-hand halves differs considerably. The Camin-Sokal tree (Fig. 1) differsfurther from the 199- and 198-step optimized Wagner trees (Figs 2, 3, 3a). However, thespecies-groups of Morophaga, plus Diataga and Amorophaga together form a monophyleticgroup as they do in the Wagner trees (although interrelationships within the group differ). Thetwo Moscardia species, Scardia + Daviscardia, Cnismorectis + Hormantris + Pectiniscardia,Miniscardia + Necroscardia and the two groups of Morophagoides species also formmonophyletic groups as in the optimized Wagner trees. However, the sequence of branching tothese groups is quite different in the Camin-Sokal tree. The 198-step trees found by PAUP involve only seven uniquely derived characters (out of a 52 GADEN S. ROBINSON AMOROPHAGADIATAGA M. bucephal a-group M. morel 1 us M. sistrata-group M. borneensi s M.choragel la-group M. cl onodes MINISCARDIA NECROSCARDIA MOSCARDIA renitens MOSCARDIA varna CRANAODES PERILICMETIS SCARDIA DAVISCARDIA GENTINGIA MONTESCARDIA MOROPHAGOIDES (OW) MOROPHA6QIDES (NW) BYTHOCRATES CNISMORECTIS HORMANTRIS PECTINISCARDIA SCARDIELLA AFROSCARDIA 1 Fig. 1 Classification of Scardiinae by Camin-Sokal technique. This was the shortest tree found with 21steps. AMOROPHAGA DIATAGA M. morel 1 us M. sistrata-group M. bucephal a-group M. borneensi s M. cl onodes M. choragel la-group SCARDIELLA AFRQSCARDIA CNISMORECTIS HORMANTRIS PECTINISCARDIA CRANAODES GENTINGIA MOSCARDIA renitens MOSCARDIA varna PERILICMETIS SCARDIA DAVISCARDIA BYTHOCRATES MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) MONTESCARDIA MINISCARDIA NECROSCARDIA AMOROPHAGA DIATAGA M. morel 1 us M. sistrata-group M. bucephal a-group M. borneensi s M. clonodes M. choragel 1 a-group MINISCARDIA NECROSCARDIA SCARDIELLA AFROSCARDIA CNISMORECTIS HORMANTRIS PECTINISCARDIA CRANAODES GENTINGIA MOSCARDIA renitens MOSCARDIA varna PERILICMETIS SCARDIA DAVISCARDIA BYTHOCRATES MONTESCARDIA MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) Figs 2, 3 Classification of Scardiinae by Wagner technique. Three alternative 199-step trees were foundthe third a variant of that in Fig. 3 with the Miniscardia + Necroscardla and Scardiella + Afroscardubranches transposed. FUNGUS MOTHS 53 Fig. 3a Classification of Scardiinae by optimized Wagner tree generated by PAUP. The consensus treefor the eight 198-step trees obtained is indicated by dashed lines. Character changes on each branch areindicated; reversals are signified by an asterisk. There are no autapomorphic changes for Scardiella orMorophagoides (Old World spp.). potential maximum clique of 10). These are 7, 8, 20, 29, 33, 52 and 54. All these were correctlypredicted by Le Quesne elimination procedures. Characters 7 and 8 are synapomorphies ofCnismorectis and Hormantris; 20 (presence of valval lobe) is a synapomorphy of Morophaga,Diataga,Amorophaga, Afroscardia and Scardiella )but the lobe in the latter two genera may notbe homologous with that in the others - see above); 29 is a synapomorphy of Morophagoides 54 GADEN S. ROBINSON species, 33 a synapomorphy of Moscardia + Perilicmetis; 52 is a synapomorphy of the branchCnismorectis - Gentingia and 54 a synapomorphy of the group Cnismorectis + Hormantris +Pectiniscardia. Thirteen of the 19 character-scores that attracted 10 or more 'hits' (see above)are explained as single convergent occurrences in individual taxa. The remaining six involvechanges to characters 9, 13 and 54. Character 54 is uniquely derived in the PAUP tree. Character9 occurs in its apomorphic state in Diataga and the Morophaga sistrata-group; 13 occurs in itsapomorphic state in Cranaodes and Gentingia and is accounted for in the PAUP analysis by a'switch on - switch off sequence as is character 9. Characters undergo up to nine changes (characters 19 and 25) in the PAUP tree. The numberof characters undergoing two to nine changes is, respectively, 11,9, 8, 12, 3, 2 and 2. Characterssurviving Le Quesne elimination procedures appear to be 'robust' on the PAUP tree. Seven ofthe 14 are uniquely derived (see above; characters 9, 13 and 23 are derived only once butundergo reversal once (9, 13) or twice (23). Characters 30, 42 and 53 are derived twice whilecharacter 3 is derived three times. Phenetic classification Trees were derived using single-link, complete-link and average-link cluster analysis (Figs 4,5,6) and by unweighted and weighted centroid cluster analysis (Figs 7, 8). Single-link clustering(Fig. 4) recovers the left half of the optimized Wagner tree produced by PAUP (Fig. 3a) as adiscrete cluster but the sequence of linkage of its component OTUs is quite different. The groupsMoscardia + Perilicmetis, Cnismorectis + Hormantris, Morophagoides, and Scardia + Davis-cardia are also recovered. Complete-link clustering (Fig. 5) recovers Morophaga + Diataga +Amorophaga as a single cluster along with Miniscardia + Necroscardia, Moscardia + Perilicme-tis, Scardia + Daviscardia, Morophagoides, and Cnismorectis + Hormantris. Average-linkclustering (Fig. 6) additionally clusters Pectiniscardia with Cnismorectis + Hormantris. Centroidcluster analysis (Fig. 7) recovers only Moscardia, Scardia + Daviscardia, Morophagoides, andCnismorectis + Hormantris: the technique is highly discriminatory and the result is somewhat at M.choraqella-group M. cl anodes M. sistrata-group M. borneensi s M. morel 1 us M. bucephal a-group SCARDIELLA AFROSCARDIA AMOROPHAGA MINISCARDIA NECROSCARDIA DIATAGA MOSCARDIA renitens MOSCARDIA varna PERILICMETIS CRANAODES BYTHOCRATES MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) MONTESCARDIA SCARDIA DAVISCARDIA PECTINISCARDIA CNISMORECTIS HORMANTRIS GENTINGIA oooooooooooooooooooooooooo Fig. 4 Classification of Scardiinae by single-link cluster analysis. FUNGUS MOTHS 55 M. borneensi s M. morel lus M. bucephal a-group AMOROPHA6A M. choragel 1 a-group M. cl onodes M. si strata-group DIATAGA MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) MONTESCARDIA BYTHOCRATES SCARDIELLA AFROSCARDIA MINISCARDIA NECROSCARDIA MOSCARDIA renitens MOSCARDIA varna PERILICMETIS SCARDIA DAVISCARDIA GENTINGIA CRANAODES PECTINISCARDIA CNISMORECTIS HORMANTRIS b ooooooooooooooooooooooooooooo Fig. 5 Classification of Scardiinae by complete-link cluster analysis. M. borneensis M. morel lus M. bucephal a-group M. choragel 1 a-group M . clonodes M. si strata-group AMOROPHAGA DIATAGA MOSCARDIA renitens MOSCARDIA varna PERILICMETIS CRANAODES GENTINGIA MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) MONTESCARDIA BYTHOCRATES SCARDIA DAVISCARDIA CNISMORECTIS HORMANTRIS PECTINISCARDIA SCARDIELLA AFROSCARDIA MINISCARDIA NECROSCARDIA ooooooooooooooooooooo 6 Fig. 6 Classification of Scardiinae by average-link cluster analysis. 56 GADEN S. ROBINSON M . borneensi s M. morel lus M. si strata-group AMOROPHAGA DIATAGA MOSCARDIA renitens MOSCARDIA varna CRANAODES SCARDIA OAVISCARDIA M. bucephal a-group M.clonodes CNISMORECTIS HORMANTRIS GENTINGIA BYTHOCRATES PERILICMETIS MOROPHAGOIDES <OW) MOROPHAGOIDES (NW) MONTESCARDIA PECTINISCARDIA MINISCARDIA AFROSCARDIA NECROSCARDIA M.choragel la-group SCARDIELLA Eb- ID- ooooooooooooooooooo Fig. 7 Classification of Scardiinae by unweighted centroid cluster analysis: the linkage sequence isclarified on the right. M. borneensi s M. morel 1 us M. sistrata-group AMOROPHAGA DIATAGA M.choragell a-group M. clonodes M. bucephal a-group MOSCARDIA renitens MOSCARDIA varna CRANAODES SCARDIA DAVISCARDIA PERILICMETIS GENTINGIA CNISMORECTIS HORMANTRIS MONTESCARDIA BYTHOCRATES MINISCARDIA AFROSCARDIA NECROSCARDIA PECTINISCARDIA MOROPHAGOIDES (OW) MOROPHAGOIDES (NW) SCARDIELLA 8 ooooooooooooooooooooooo Fig. 8 Classification of Scardiinae by weighted centroid cluster analysis: the linkage sequence is clarifiedon the right. FUNGUS MOTHS 57 variance with those obtained using other methods. Weighting the characters in centroid analysis(Fig. 8) permits recovery of a further group found by other methods - Morophaga + Amorphaga+ Diataga. Within Morophaga, Diataga and Amorophaga, certain OTUs are grouped consistently by thevarious phenetic methods. The Morophaga choragella-group and M. clonodes are paired by fourof five methods; M. borneensis is paired with M. morellus by four of five methods. The M.sistrata-group is paired with the M. choragella-group + M. clonodes by three of five methods.None of these groups is supported by the PAUP tree but the Camin-Sokal tree also pairs the M.choragella-group and M. clonodes. Classification adopted The classification of the Scardiinae adopted here is a modified version of the 198-step tree shownin Fig. 3a. Tinissa is placed tentatively as the sister-group ofNecroscardia, and Semeoloncha asthe sister-group of Cranaodes. For the purposes of the present paper, Amorophaga and Diatagahave been retained as separate genera, but it is probable that Morophaga is paraphyletic withrespect to both. Morophaga borneensis is considered to be a member of the M. s is trata- group,sharing character 30 which is considered to be uniquely derived. Relationships of Amorophaga,Diataga and the species-groups within Morophaga are debatable; drastic rearrangement of thetree is possible with very little increase in the number of character-steps involved. The sequenceadopted here for Amorophaga, Diataga and the species-groups of Morophaga is thereforearbitrary. The integrity of Morophaga + Diataga + Amorophaga is supported by all methods exceptsingle-link and unweighted centroid cluster analysis. This group appears to be genuinelymonophyletic, defined by the derivation of characters 1, 2, 10 and 23 and the reversal of 31 in thePAUP tree: character 20 (basal lobe of valva) also supports the monophyly of the group if theoccurrence of valval lobes of rather different type in Afroscardia and Scardiella is ignored (seeabove). Characters 20 and 23 (modified interpretation) are uniquely derived but 23 undergoesreversal twice. Characters 1 and 2 are derived three times, character 10 twice (it is otherwise anautapomorphy of Gentingia) . All analyses except single-link and centroid clustering support the sister-group relationship ofMiniscardia and Necroscardia but the two groups are united only by the convergent derivation ofcharacters 1 and 5 in the PAUP tree. Both characters occur twice elsewhere. The sister-group relationship of the two Moscardia species is supported by all analyses: fivecharacter changes are involved but these all occur also elsewhere. The sister-group relationshipof Perilicmetis to Moscardia is supported by cladistic and most phenetic analyses. The fourcharacter changes involved include one uniquely derived character (33). The grouping is furthersupported by character 32 which, however, occurs also in Cnismorectis and Hormantris. Scardia and Daviscardia are paired by all analyses. On the PAUP tree their relationship issupported by changes to characters 2, 4, 12, 21, 25 and 39. Each of these is very homoplasiouswith a minimum of three independent apomorphic occurrences. The sister-group relationship of Cnismorectis and Hormantris, supported by all analyses,involves derivation of eight characters and reversal of one. Two of the characters (7, 8) areuniquely derived and three more (32, 34, 45) are derived elsewhere only once. The sister-grouping of Pectiniscardia with Cnismorectis + Hormantris, found by all cladistic methods andaverage-link cluster analysis, is supported by two derived characters and three reversals. Onecharacter (54) is uniquely derived; the other is multiply derived. One of the reversals (13) is of acharacter that is otherwise uniquely derived; the other two characters are multiply derived andreversed. The Old World and New World species of Morophagoides are grouped in all analyses; on thePAUP tree their relationship is supported by two derived characters, one unique (29) and theother multiply derived, and two reversals, both of characters multiply derived and reversed. Two other groups in the PAUP tree are defined by uniquely derived characters that exhibitalso a single reversal. Characters 9 and 13 appear in the data matrix as synapomorphies of 58 GADEN S. ROBINSON Diataga and the Morophaga strata-group and of Cranaodes and Gentingia respectively butattract a high number of 'hits' (see above) and are accounted for in the PAUP tree ashomoplasious. No method of analysis recovers these pairs of OTUs together. Other groups are not consistently recovered by either phenetic or cladistic methods althoughthere is strong suggestion of a close relationship between the Morophaga choragella-group andM. clonodes from phenetic analysis. Groups other than the six discussed above may be defined only in terms of characters that arehighly homoplasious; none of the groups is convincing. The limited phylogenetic hypothesisproposed here is therefore that the six groups (with the possible exception of Miniscardia +Necroscardia) constitute monophyletic entities. The remainder of the PAUP tree is adopted as acuratorial convenience. Discussion The impossibility of obtaining a consistent classification of OTUs within the Scardiinae reflectstwo factors, the high degree of homoplasy encountered throughout the group, and thelimitations placed upon the range of characters with which it was possible to work throughpaucity of material. Studies such as this underline the dangers of subjective character-assessment. Spectacularspecialized structures such as elongate corematal apodemes (character 15) or the valvalpectinifer (present in two Morophaga species and Pectiniscardia) prove to be homoplasious,while apparently insignificant characters such as abdominal microtrichia (32, 33) would appearto have much greater 'information value' in classification. Certain characters are alarminglyhomoplasious, for example dorsal fusion (or otherwise) of the tegumen, modification of thejuxta, presence or absence of coremata, reduction of the maxillary palpus, and surface structureof the aedeagus. The homoplasy of the coremata has already been alluded to (Robinson, 1981)but the other characters are 'traditional' taxonomic characters and, if their behaviour in othergroups parallels that in the Scardiinae, our trust in them is perhaps misplaced. It is not suggested here, of course, that phenetic methods should produce results congruentwith those of cladistic techniques. However, given that the criterion for clustering or agglomera-tion is the pattern of distribution of Ts in the data matrix (apomorphies among OTUs) thenmimicry of pattern will tend to emerge in the form of the common occurrence of (usuallyterminal) groups or clusters. However, it is axiomatic that phenetic techniques cannot be reliedupon to identify monophyletic groups. Cladistic testing of phenetic clusters is necessary toestablish that they are natural groups in the phylogenetic sense. The weaknesses of bothmethods in situations involving homoplasy are obvious. Phenetic classifications are even moreunlikely to be natural classifications. Use of the methods described above permits the identification of groups that are recoveredconsistently and which might be described as 'robust groups'. Cladistic methods identify thosecharacters by which such robust groups may be defined. Character-compatibility methodsmeasure objectively (within the context of the data) the robustness of characters. Robust groupsof OTUs are usually defined by robust characters. Character-compatibility methods furtheridentify and predict (by the 'hits' procedure) the occurrence of single instances of convergenceor reversal. A consensus obtained using these methods provides a strong (but, admittedly,subjective) indication that robust groups are natural groups, an indication that cannot beobtained by any one method in isolation. The classification offered here is, therefore, a very conservative one. Nevertheless, it isentirely at variance with that proposed by Zagulajev (1973). He suggested Amorophaga andMorophagoides as sister-groups, Scardia as the sister-group of these, and Morophaga as thesister-group of all three (Zagulajev's Scardia includes Montescardia, separated in the presentwork). Geographical distribution It has been impossible to resolve fully a phylogeny for the Scardiinae so comment on FUNGUS MOTHS 59 biogeography in relation to phylogeny must needs be restricted to the few unequivocalsister-groupings recognized. The overall pattern of diversity of the Scardiinae is dominated by Tinissa, with 34 species.These range from the Afrotropical region (5 species) through the Oriental region (19 species) tothe Australian region (New Guinea, Solomon Is, Australia - 13 species) but are poorlyrepresented in Australia itself where there are only two species, both of these found also in NewGuinea. Further comments on the distribution of Tinissa have been given elsewhere (Robinson,19760; 1981). Apart from Tinissa, Australia has only one other species of Scardiinae, Moropha-ga clonodes. Its possible sister-group (suggested by only phenetic analyses) is the M. choragella-group which consists of four Palaearctic species. Penetration of the Australian region byScardiinae is restricted otherwise to Morophaga bucephala which ranges from Japan to NewGuinea. Excluding Tinissa, Scardiinae are represented in the Afrotropical region almost as poorly asthey are in Australia. Two of the four species of the Morophaga bucephala-group occur there,one widespread in forested areas of the African mainland and the other restricted to Madagas-car. In addition to these, four monobasic Scardiinae genera have been found in Africa. Each isknown only from a few specimens and the phylogenetic position of each is uncertain. Three ofthe genera, Afroscardia, Leptozancia and Philagrias, occur on the mountains of east Africa ataltitudes over 2400 m (8000'). The fourth, Semeoloncha, occurs in the rain forests of west Africa.The current picture of Scardiinae diversity in the Afrotropical region may, however, bemisleading. Few specimens have been collected yet the generic diversity among these is great. Itis probable that the material available in collections falls far short of being a true representationof the group. It is likely that further collecting and, particularly, the use of specialized collectingtechniques, may increase substantially the range of Scardiinae genera known from the Afro-tropical region. No Scardiinae are known from southern South America (Patagonia) despite extensive recentcollecting of Microlepidoptera by workers from the U.S.A. and Denmark. The virtual absenceof Scardiinae from the southern continents suggests strongly that the group evolved after thebreak-up of Gondwanaland. In contrast, the Neotropical region has a greater concentration of Scardiinae genera (13) and(if Tinissa in the Oriental region is excluded) of species (32) than any other region. Again, thenumber of specimens known of each species is small and the diversity suggested here is probablya considerable underestimate. Three genera, Morophagoides , Daviscardia and Diataga, areparticularly diverse, accounting for 19 of 32 species. All are found in forest habitats and have aconsiderable altitude range, Morophagoides especially so, M. iulina having been collected at atleast 2600 m (8500'). The remaining Neotropical genera occur in forested habitats generallybelow about 2000 m (6550') from the eastern slopes of the Andes and the mountains of CentralAmerica (Cnismorectis , Necroscardia, Moscardia, Cranaodes} to the delta forest of the Amazonbasin (Perilicmetis, Miniscardia); Bythocrates is known only from Trinidad, and Diataga iswidespread. Pectiniscardia and Hormantris are exceptional in that they are found at altitudescomparable to those at which live the Scardiinae of the east African mountains. Both genera aremonobasic and represented by single specimens collected in the central Cordillera of Colombiaat over 3500 m (11,500'). Hormantris is the sister-group of Cnismorectis. A single example ofScardia anatomella is known from the Neotropical region. The Nearctic Scardiinae fauna is poor with only eight genera and 10 species. One species ofDiataga, one of Miniscardia, two of Morophagoides and two of Daviscardia represent thediverse Neotropical element at its northern limit. Montescardia, Scardia and Amorophaga areHolarctic genera each represented by single species in North America. The monobasicScardiella is the only Scardiinae genus restricted to the Nearctic region. The Palaearctic region has a similarly sparse Scardiinae fauna, with five genera, theserepresented in both the eat and the west of the region. The 16 Palaearctic species are dividedevenly between west and east. Morophagoides (also in the Neotropical and Nearctic regions) isrepresented by three species, the Holarctic genera Montescardia, Scardia and Amorophaga bytwo, three and three species respectively. The remaining six species are those of Morophaga. 60 GADEN S. ROBINSON This genus is also well-represented in the Oriental region with a further six species. The paucityof Scardiinae in the Holarctic region may be connected with the presence of Nemapogoninae aspart of the guild of fungivorous larvae. Nemapogoninae are absent from both the Old and NewWorld tropics. Four genera and 10 species of Scardiinae, excluding Tinissa, are now recorded from theOriental region. In addition to representatives of Morophaga (including the strata-group,endemic to the Oriental region), there are two species of Scardia, one of Cranaodes (knownotherwise from the Neotropical region), and the monobasic genus Gentingia, of uncertainaffinities. The sister-group relationship of Daviscardia and Scardia provides a scenario of dispersal andspeciation that may be mirrored also in other groups. Daviscardia is a Neotropical genus with afurther two species in the U.S.A. Scardia anatomella (also from the U.S.A.) is suggested(below) to be the sister-group of all other Scardia. The pattern of present-day distribution ofScardia species suggests dispersal of the Old World Scardia ancestor from North America toeastern Asia with progressive southward and westward dispersal and isolation to provide thehypothesized phyletic sequence: assamensis (Himalayas/N. India), amurensis (E. Palaearctic/Japan), alleni (Borneo), and boletella and caucasica (W. Palaearctic). In this case speciation hasinvolved adaptation within one genus to a wide variety of climates. It is tempting, particularly in view of the presence in the Oriental region of what may be agenuine Cranaodes species, to view the phylogeny of the Scardiinae in terms of a series ofdispersal events from a Neotropical centre. This model is fitted, for example, by Morophagoidesand by the hypothesis of a sister-group relationship for Necroscardia and Tinissa, the Afro-tropical origin for Tinissa (Robinson, 1976a) now being unlikely in the light of the discovery of a'primitive' species in the New Guinea subregion (Robinson, 1981). However, in the absence of amore sound phylogenetic hypothesis this can only be speculative. Abbreviations BMNH British Museum (Natural History), London, England CAS California Academy of Sciences, Los Angeles, California, U.S.A. EIHU Entomological Institute, Hokkaido University, Sapporo, Japan HNHM Hungarian Natural History Museum (Termeszettudomanyi Muzeum), Budapest, Hungary LN Landessamlungen fur Naturkunde, Karlsruhe, West Germany MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. MINGA Muzeul de Istoria Naturala 'Grigore Antipa', Bucharest, Rumania MNHN Museum National d'Histoire Naturelle, Paris, France MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, East Germany NM Naturhistorisches Museum, Vienna, Austria NMNH National Museum of Natural History, Smithsonian Institution, Washington D.C. , U.S.A. RNH Rijksmuseum van Natuurlijke Historic, Leiden, Netherlands SAM South Australian Museum, Adelaide, Australia SM Sarawak Museum, Kuching, Sarawak UC University of California, Berkeley, California, U.S.A. ZI Zoological Institute of the Academy of Sciences, Leningrad, U.S.S.R. ZM Zoologisk Museum, Copenhagen, Denmark Acknowledgements For the loan, exchange or presentation of specimens I am most grateful to: Dr V. O. Becker, Brasilia; DrD. R. Davis, NMNH, Washington; Dr A. Diakonoff, RNH, Leiden; Dr F. Kasy, NM, Vienna; Dr N. P.Kristensen and Dr E. S. Nielsen, ZM, Copenhagen; Dr E. G. Matthews, SAM, Adelaide; Dr S. Moriuti,Osaka; Dr A. Popescu-Gorj , MINGA, Bucharest; Dr J. A. Powell, UC, Berkeley; Dr P. Viette, MNHN,Paris, and Dr A. K. Zagulajev, ZI, Leningrad. I am most grateful to my colleagues, Mrs L. M. Pitkin, Dr K. Sattler and Mr K. Tuck, for their help andadvice during the preparation of this paper and for their constructive criticism of the manuscript. I amparticularly grateful to Dr I. D. Gauld and Dr I. J. Kitching for invaluable and stimulating discussions ofnumerical methods in the investigation of character compatibility and the construction of phy logenies . I am FUNGUS MOTHS 61 indebted to Dr W. J. Le Quesne and Dr I. M. White for the provision of computer programs and for theiradvice on computing problems. Recently collected material from West Malaysia, Brunei and Nepal was obtained either by the collectingefforts of myself and my colleague Kevin Tuck or through the generosity of Lt-Col. M. G. Allen who hasdonated large numbers of specimens from Brunei and Nepal to the BMNH collection. Tuck's collecting inWest Malaysia was supported through the generosity of Mr H. Barlow. My collections in Brunei weremade during participation in Exercise Ulu Temburong under the aegis of the Ministry of Defence:comprehensive acknowledgements for this expedition are published elsewhere (Robinson, 1984). Ourjoint collecting in Nepal was made possible through the unstinting help and hospitality of Lt-Col. and MrsM. G. Allen. Examination of material in the Zoological Institute, Leningrad, was carried out under the RoyalSociety/Soviet Academy of Sciences Scientific Exchange Programme. SCARDIINAE Eyer Scardiinae Eyer, 1924: 320; Zagulajev, 1966: 634; Capue, 1968: 122; Zagulajev, 1973; Gozmany & Vari, 1973: 147. Type-genus: Scardia Treitschke. Semeolonchini Capue, 1971: 232. Type-genus: Semeoloncha Gozmany. Syn. n.Tinissinae Gozmany & Vari, 1973: 84; Robinson, 1976; Robinson, 1981. Type-genus: Tinissa Walker. Syn. n. ADULT. Vertex with tufts of erect scales convergent towards mid-line of head and tending to form whorls atback of head; scale bases often divided medially and transversely into four groups by narrow bare areas.Frons with tufts of erect and slightly upward-directed scales convergent with scale tufts of frontovertex;scale-bases either scattered evenly or forming upper medial group (sometimes divided vertically) and pairof lateral groups close to tentorial pits. Epipharynx triangular or hardly developed. Pilifers present exceptin Hormantris and Cnismorectis (Figs 180-182). Mandible present, elongated transversely. Galeae presentin all species examined except Tinissa poly stacta, only rarely associated, if at all, rarely much longer thansecond segment of labial palpus. Cibarium without chemoreceptor pits in Morophaga, Scardia, Tinissa andMontescardia but pits present in Bythocrates, Perilicmetis and Cnismorectis (not checked in other genera).Maxillary palpus elongate, 5-segmented and folded in many species but in some substantially reduced inlength and with five, four or three segments; second segment frequently with thick patch of dark scalesabove; first segment lacking sensilla trichodea; terminal segment with whorl of pale scales in most speciesof Tinissa. Labial palpus three-segmented, elongate, upturned at about 60; second segment enormouslyelongated in Hormantris and Cnismorectis; lateral bristles of second segment sparse, scattered along lengthand never concentrated apically to form a terminal whorl; third segment with vom Rath's organ eitherpresent but small or apparently absent. Antenna one-half to four-fifths length of forewing. Scape elongatebarrel-shaped; pecten present with 4-10 sparse bristles or large and thick with 15 to more than 40 bristles.Pedicel short, barrel-shaped, less than twice as long or broad as adjacent flagellar segments. Flagellarsegments ciliate, cilia short in females, rarely longer than flagellar diameter, more elongate in males, up tofour or five times as long as flagellar diameter, in some taxa restricted to ventral half of segment; eachsegment with whorls of overlapping scales arranged in longitudinal rows, in some genera the whorlincomplete ventrally; most segments with well-developed palisaded sensilla coeloconica. Forewing with allveins present; cell with at least traces of forked M and with at least traces of chorda; stalking of /? 3 and R 4 ,R 4 and R 5 , M 2 and M 3 , or M 3 and CuA\ occurs in some genera (Figs 183, 184). Pattern usually cryptic,resembling mottled tree-bark, or with dark ground colour and pale apex and dorsum, or consisting of boldblotches on a paler background, rarely almost unicolorous (Bythocrates, Afroscardia, some Tinissa).Hindwing with all veins present, trace of forked M in cell; female with three (or, occasionally, four)frenular bristles (but with about 15 in Scardia). Legs without significant modification; spines absent;epiphysis present, strong, elongate peg-shaped; mid-tibia of many species with oblique pale band on outersurface; hind tibia with conspicuous distal whorl of erect scales in Tinissa. GENITALIA d*. Eighth segment with elongate tubular sternal apodemes associated with coremata inMorophaga choragella-group, Diataga and Necroscardia; flap-like apodemes from corners of sternitepresent in Semeoloncha, Moscardia and Cranaodes; sternite with postero-lateral processes in Tinissatorvella and T. ruwenzorica; eighth segment otherwise unmodified; coremata present or absent in pleuralmembrane. Saccus well developed, longer than broad in most genera. Tegumen broad, angled dorso-ventrally in most species and, in most, fused dorsally. Uncus bilobate, lobes sometimes fused with tegumenand with each other, complexly modified in some taxa but in others a pair of simple setose lobes. Gnathosabsent. Subscaphium, if present, ribbon-like, ill-defined, but modified with pair of setose processes in one 62 GADEN S. ROBINSON species of Tinissa. Juxta, if present, simple or strongly modified, fused with valvae or free, in Tinissa andNecroscardia enormously enlarged and forming an integral part of a valve-juxta complex. Transtilla, ifdeveloped, forming a dorsal collar (Morophaga, Diataga) or, rarely, modified with labides (Tinissa,Leptozanda). Valva present (except, possibly, in Leptozanda), variously modified; valvae free or, in sixgenera, fused ventrally to form a single movable complex; pectinifer present in two species of Morophagaand in Pectiniscardia; articulated ventrodistal process absent. Aedeagus simple, free, with or withoutcarinae, straight or almost so (but highly modified and partially fused with the valvae in Montescardia);inception of bulbus ejaculatorius laterobasal or subbasal; vesica smooth or with fine spicular cornuti(microtrichia) or, in a few species, with stout cornuti. GENITALIA $. Seventh segment with strongly spined sternite in Necroscardia funeratella; reduced to anarrow band and with corethrogyne in most Tinissa species; corethrogyne also present but segment ofnormal size in Diataga and Cnismorectis; tergite with spined posterior lobe in a few Tinissa species;otherwise unmodified. Eighth tergite of most species with 2-5 pairs of strong terminal or subterminal setaeplus sparse smaller setae and, in some species, with a few scattered sensilla coeloconica and/or sensillabasiconica. Eighth sternite usually strongly sclerotized and modified to accommodate broad ostium,frequently with paired processes ventral, lateral or immediately dorsal to ostium each bearing a pair ofstrong setae. Antrum invariably developed, variously modified, incorporating inception of ductus semin-alis dorsally or paradorsally or terminating just posterior to inception of ductus seminalis. Inception ofductus seminalis very close to ostium in some taxa, these frequently with bulbous outgrowth from ventralwall of ovipositor overlying ostium and possibly functioning as a closure. Ductus bursae looped to the rightin some Tinissa and Morophaga, its internal surface with microtrichia or overlapping plaques in manyspecies; some genera (notably Tinissa) with regular transverse constrictions such that the ductus resemblesa shower-hose. Corpus bursae thin- walled, with signa in Gentingia, Bythocrates and Morophagoides only.Ovipositor elongate, slender, with single infold at rest, in most taxa about as long as the last threeabdominal segments; posterior half with ventral rods arising from bases of papillae anales; apophysesposteriores extending anteriorly beyond margin of eighth sternite by about once or twice length of sternite. PUPA (Figs 185-192). Antennae almost reaching wing-tips; wing-tips reaching fourth abdominal segment.Anterior and posterior transverse bands of dorsal spines present on abdominal segments 3-7 in males and3-6 in females (anterior band on segment 3 represented only by a smooth ridge in Diataga); single(anterior) band present on segments 8 and 9 in males and 7-9 in females (but no spines on segments 8 and 9and only anterior band present on segment 7 in Diataga; band on segment 9 reduced to only four spines inone example of Morophaga sistrata); trace of anterior band present also on second abdominal segment inScardia. Band of spines on segment 8 incomplete medially in Morophagoides, complete in all other taxaexamined (see 'Remarks'); band of spines on segment 9 incomplete medially in all taxa except Morophagacremnarcha and in two of three males of Morophaga morellus. Cremaster with two pairs of ventral hooks inScardia and some Morophaga but smaller inner pair absent in Diataga and Morophagoides and stronglyreduced or absent in Morophaga. In rotting wood or in fungal sporophores, protruded prior to emergenceof adult. LARVA (Figs 193-197). Al of head only one-third length of A2; six poorly defined ocellar lenses present.L-group of prothorax bisetose; Dl setae of abdominal segments more widely separated than D2 setae;L-group of ninth abdominal segment bisetose. Dorsal cuticle with strong microtrichia. In rotting woodpermeated by fungal hyphae or in hard (persistent) fungal sporophore. REMARKS. Description of the scardiine pupa is based on the study of Morophaga choragella (severalexamples of both sexes), M. morellus (3 cf), M. cremnarcha (1 $), M. hyrcanella (1 $), M. sistrata (2 $),Diataga leptosceles (1 cf , 1 $), Scardia anatomella (several examples of both sexes) and Morophagoidesburkerella (1 $). Additionally, figures of female pupae of Scardia boletella and Morophagoides moriutii aregiven by Zagulajev (1973) and Moriuti (1976) respectively. The larval diagnosis is based on study of threelarvae of Diataga leptosceles from Turrialba, Costa Rica, on the detailed description of Morophagachoragella by Hinton (1956), and on figures of Scardia boletella by Zagulajev (1973) and of Morophagoidesmoriutii by Moriuti (1976). The terminology of larval setae adopted here for Diataga leptosceles follows that used by Hinton (1956)for Morophaga choragella. However, Moriuti (1976) uses the earlier terminology of Hinton (1946) and thereader is referred to MacKay (1963) for a discussion of the still unresolved differences between the twosystems. Hinton (1946) terms the four anterior macrosetae of the prothoracic shield XD1, XD2, SD1 andSD2 (Dl, XD1, XD2 and SD2 of Hinton, 1956) and the two posterior macrosetae Dl and D2 (D2 and SD2of Hinton, 1956); the L-group setae of the ninth abdominal segment are LI and L3 (LI and L2 of Hinton,1956). FUNGUS MOTHS 63 The larva of Diataga leptosceles is similar to that of Morophaga choragella as described by Hinton but thepositions of Va and Pb on the head are variable ; the labrum has only three pairs of marginal setae , not fouras in choragella or Morophagoides moriutii (Moriuti, 1976). On the abdominal segments Dl is stronglydisplaced ventrad in Diataga in comparison with Morophaga and Morophagoides. In Morophagoides theD-group setae are almost level; in Morophaga Dl is displaced slightly ventrad. In Morophagoides L3 onthe abdominal segments is placed slightly further posteriorly than in Diataga or Morophaga. Key to genera and species-groups of Scardiinae (males) 1 Hind tibia with strongly developed apical and subapical tufts of elongate, dark-tipped scales TINISSA(p. 108)Hind tibia rough-scaled above but scales never concentrated into conspicuous distal tufts 2 2 Antenna lacking dorsal cilia -cilia roughly restricted to ventral 180 degrees of each segment 3 - Antenna with dorsal cilia - cilia arranged through 360 degrees of each segment 10 3 Antennal segments completely scaled 4 - Antennal segments with ventral surface lacking scales 17 4 Antennal cilia shorter than 1 -5 x flagellar diameter 5 Antennal cilia longer than 1 -5 x flagellar diameter 6 5 Forewing with M 3 and CuA\ stalked; forewing speckled greyish brown with dark brown subterminal fascia (Fig. 40) PERILICMETIS (p. 92) Forewing with A/ 3 and CuA\ separate; forewing dark purple-brown with pale termen and dorsum(Figs41,42) MOSCARDIA (p. 93) Pilifers absent; second segment of labial palpus longer than width of head (cf. Fig. 182) HORMANTRIS(p. 102)Pilifers present; second segment of labial palpus shorter than width of head (Figs 180, 181) 7 7 Forewing with M 3 and CuAi stalked MINISCARDIA(p. 104) Forewing with M 3 and CuA i separate 8 8 Forewing pattern consisting of dark purple-brown ground-colour with pale terminal fascia, and small pale spots on posterior margin (Figs 53, 54); juxta complex, divided, forming afunctional part of the valvae (Figs 108, 109) NECROSCARDIA (p. 106) - Forewing pattern consisting of dark blotches on a pale bronze or cream ground-colour (but see Cranaodes oroyd) ; juxta simple , undivided , or fused with valvae and not recognizable 9 9 Forewing with RT, and R 4 stalked (Fig. 184); tegumen unbroken, completely sclerotized dorsally; valvae fused ventrally; aedeagus with fine spicular carinae (Fig. 103) GENTINGIA (p. 95) - Forewing with RT, and R 4 separate; tegumen broken dorsally by at least a membranous suture-line; valvae separate; aedeagus smooth-surfaced (Fig. 106) CRANAODES (p. 98) 10 Antennal cilia longer than 1-5 x flagellar diameter 11 Antennal cilia shorter than 1 -5 x flagellar diameter 14 1 1 Antennal scape with more than 15 pecten bristles 12 Antennal scape with fewer than 15 pecten bristles 13 12 Maxillary palpus with fewer than 5 segments; forewing uniformly brown (Fig. 51); juxta simple, entire. (Afrotropical region) AFROSCARDIA(p. 110) Maxillary palpus with 5 segments; forewing with mottled pattern or with pale markingsconcentrated at termen, costa and dorsum; juxta complex, divided, fused with valvae. (NewWorld) MOROPHAGOIDES (part) (p. 65) 13 Small (13-16 mm) Nearctic species; uncus complex, forming pair of widely separated digitate processes, not fused with tegumen; valva with simple ventral margin (Fig. 107) SCARDIELLA(p. 108) Large (27 mm) Neotropical species; uncus simple - a pair of setose lobes fused with tegumen;valva with ventral pectinifer of spines PECTINISCARDIA (p. 101) 14 Antennal scape with more than 15 pecten bristles 15 Antennal scape with fewer than 15 pecten bristles 16 15 Coremata present in eighth abdominal segment; juxta complex, enveloping aedeagus MONTESCARDIA (p. 74)Coremata absent from eighth abdominal segment; juxta, if present, fused with valvae and not recognisable MOROPHAGOIDES (part) (p. 65) 16 Large (20 mm) Afrotropical species; maxillary palpus 5-segmented; coremata present in eighth abdominal segment SEMEOLONCHA (p. 97) 64 GADEN S. ROBINSON - Small (11-14 mm) Neotropical species ; maxillary palpus with fewer than 5 segments ; coremata absent from eighth abdominal segment BYTHOCRATES (p. 77) 17 Forewing with /? 3 and R 4 free, widely separated at base (if /? 3 and R 4 close together at base, then forewing pattern mottled purple-brown with pale termen and dorsum) 18 - Forewing with R$ and R 4 fused, stalked or approximated at base; forewing lacking paler apex and dorsum 20 18 Pilifers absent; second segment of labial palpus longer than width of head (Fig. 182) CNISMORECTIS(p. 102)Pilifers present ; second segment of labial palpus shorter than width of head 19 19 Coremata present in eighth abdominal segment; juxta complex, fused with valvae SCARDIA (p. 86) Coremata absent from eighth abdominal segment; juxta, if present, fused with valvae and notrecognizable DA VISCARDIA (p. 78) 20 Forewing with white or off-white ground-colour, marked with bold purple-brown basicostal and postero-medial blotches, with a group of smaller spots along costa becoming progres-sively larger towards apex (Figs 67-70) MOROPHAGA bucephala-group (p. 121) - Forewing pattern cryptic, resembling moss or tree-bark, or longitudinally streaked 21 21 Forewing pattern consisting of either brownish grey or khaki longitudinal streaks (Figs 57-59) ; uncuscomplex AMOROPHAGA(p. Ill) - Forewing pattern resembling moss or tree-bark; uncus simple 22 22 Forewing slender; veins R 3 + R 4 and M 2 + M 3 fused, stalked or closely approximated at base (Fig. 183) ; outer mid and proximal hind tibial spurs very short - less than 0-4 length of innerspurs (Neotropical species) DIATAGA(p. 114) - Forewing broader; only /? 3 and R 4 fused, stalked or closely approximated at base; outer and proximal hind tibial spurs usually more than 0-4 length of inner spurs (but slightly less than 0-4 in the Oriental Morophagasistrata-group) 23 23 Coremata absent from eighth abdominal segment 24 - Coremata present in eighth abdominal segment 25 24 Outer mid and proximal hind tibial spurs less than 0-4 length of inner spurs; apices of uncus lobes fused together and tapered to form a small hook; vesica with line of strong thorn-likecornuti (Figs 131-137) MOROPHAGA s/strafa-group (part) (p. 126) - Outer mid and proximal hind tibial spurs more than 0-4 length of inner spurs; apices of uncus lobes separate; cornuti small and spicular (Fig. 138) MOROPHAGA c/onodes-group (p. 129) 25 Apices of uncus lobes fused and tapered to form a small hook; vesica lacking any ornamenta- tion (Fig. 135) MOROPHAGA sistrata-group (part - borneensis) (p. 126) - Apices of uncus lobes separate ; vesica with minute spicular cornuti 26 26 Uncus lobes short, hardly extending beyond apices of valvae, square-ended MOROPHAGA morellus-group(p. 125) - Uncus lobes elongate , extending well beyond apices of valvae , apices tapered and rounded MOROPHAGA choragella-group(p. 130) Key to genera and species-groups of Scardiinae (females) (Females of Moscardia, Semeoloncha, Afroscardia, Pectiniscardia, Perilicmetis and Hormantris areunknown.) 1 Hind tibia with strongly developed apical and subapical tufts of elongate, dark-tipped scales; most species with seventh segment reduced, tergite and sternite more than twice as wide aslong, with corethrogyne of dense, fine, elongate hair-scales TINISSA (p. 108) - Hind-tibia rough-scaled above, but scales never concentrated into conspicuous distal tufts; seventh segment not reduced , corethrogyne present in Cnismorectis and Diataga only 2 2 Forewing pattern consisting of longitudinal streaks or a 'moss' pattern disrupted by longitudin- al streaking (Figs 57-59) AMOROPHAGA(p. Ill) - Forewing pattern not consisting of longitudinal streaks 3 3 Forewing uniformly dark purple-brown BYTHOCRATES (p. 77) - Forewing variegated 4 4 Forewing pattern consisting of large purple-brown spots on a paler ground-colour 5 - Forewing coloration forming either a cryptic 'moss' or 'tree-bark' pattern or consisting of a pale termen and dorsum on a darker ground-colour 7 5 Forewing ground-colour white or off-white (Figs 67-70) MOROPHAGA bucephala-group (p. 121) FUNGUS MOTHS 65 - Forewing ground-colour pale bronze 6 6 Maxillary palpus short, with fewer than 5 segments; forewing with R 3 and R 4 stalked (Fig. 184) GENTINGIA (p. 95) - Maxillary palpus elongate, 5-segmented; forewing with 7? 3 and /? 4 free CRANAODES (p. 98) 7 Forewing with RT, + R 4 and M 2 + M 3 stalked or very closely approximated at base (Fig. 183) DIATAGA(p. 114) - Forewing with all veins free or only one pair of veins stalked 8 8 Forewing with M 3 and CuA l stalked MINISCARDIA(p. 104) - Forewing with M 3 and CuA l free 9 9 Forewing with all veins free 10 Forewing with R 3 and R 4 fused or stalked 17 10 Forewing pattern composed of pale termen and dorsum on darker ground-colour 11 - Forewing coloration forming a cryptic 'moss' or 'tree-bark' pattern 14 11 Frenulum with more than 10 bristles SCARDIA (p. 86) - Frenulum with fewer than 5 bristles 12 12 Bursa copulatrix with pair of large, opposed, pouch-shaped signa (Figs 140-144) MOROPHAGOIDES (part) (p. 65) - Bursa copulatrix with many fine spicular signa, or without signa 13 13 Antennal pecten with more than 16 bristles; antrum less than one-half length of apophyses anteriores; ductus bursae smooth-walled (Figs 147-151) DA VISCARDIA (p. 78) - Antennal pecten with fewer than 16 bristles; antrum more than one-half length of apophyses anteriores; ductus bursae with fine, irregular transverse constrictions (Fig. 162) NECROSCARDIA(p. 106) 14 Pilifers absent; second segment of labial palpus longer than width of head (Fig. 182) CNISMORECTIS(p. 102) - Pilifers present; second segment of labial palpus shorter than width of head (Figs 180, 181) 15 15 Bursa copulatrix with pair of large, opposed, pouch-shaped signa (Figs 140-144) MOROPHAGOIDES (part) (p. 65) - Bursa copulatrix without signa 16 16 Large (22-28 mm) species; antennal pecten with more than 15 bristles. . . MONTESCARDIA (p. 74) - Small (13-16 mm) species; antennal pecten with fewer than 15 bristles SCARDIELLA(p. 108) 17 Intersegmental membrane between eighth and seventh abdominal segments with pair of deep pouches with scobinate inner surfaces (Fig. 172) MOROPHAGA cJonodes-group (p. 129) - Intersegmental membrane lacking such pouches or, if present, pouches shallow and with smooth inner surfaces 18 18 Corpus bursae not extending anteriorly beyond apophyses anteriores MOROPHAGA morellus-group(p. 125) - Corpus bursae extending anteriorly beyond apophyses anteriores 19 19 Ductus + corpus bursae more than twice length of apophyses anteriores; antrum less than half length of apophyses anteriores (Fig. 179) MOROPHAGA s/sfrafa-group (p. 126) - Ductus + corpus bursae less than twice length of apophyses anteriores or, if more, then antrum extending anteriorly beyond apices of apophyses anteriores MOROPHAGA choragella-group(p. 130) MOROPHAGOIDES Petersen Morophagoides Petersen, 1957: 593. Type-species: Scardia ussuriensis Caradja, 1920: 167, by originaldesignation and monotypy. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter (but not in species from the Old World). Scape with more than 15 pecten bristles. Interocularindex (male) 1-0 or less. Maxillary palpus 5-segmented; pilifers present; second segment of labial palpusshorter than width of head. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewingwith /? 3 and R 4 separate; Af 2 , M 3 and CuA^ separate; mottled coloration forming cryptic, coarse 'moss'pattern (but pale markings of some species concentrated at termen, costa and dorsum). Male lackingcoremata in eighth abdominal segment. Male genitalia with complex uncus (simple in burkerelld)separated from tegumen by narrow band of membrane; tegumen unbroken, completely sclerotizeddorsally ; valva lacking basal setose lobe on inner surface; apex of valva forming ventral hook or hooks, orwith spines; valvae + juxta fused ventrally into a single movable complex, valvae without longitudinalcleft; saccus longer than wide; juxta complex, divided medially; however, juxta, if present, fused with 66 GADEN S. ROBINSON valvae and not recognizable as such in species from the Old World; vesica with spicular cornuti; aedeagussmooth-surfaced, without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Female with pair of strong, pocket-shaped signa in bursa copulatrix. DISTRIBUTION . Western and eastern Palaearctic region ; Oriental region - Taiwan ; Nearctic region - BritishColumbia, Washington State, Utah, California; Neotropical region. BIOLOGY. See under entries for individual species. Key to species of Morophagoides Males (males ofpythium, nimbiferum and montium are unknown) 1 Antennal cilia as long as or longer than 1 -5 x flagellar diameter; juxta complex, bipartite. (New World) 2 Antennal cilia shorter than 1-5 x flagellar diameter; juxta, if present, fused with valvae and notrecognizable. (Old World) 4 2 Uncus with lateral margins strongly infolded , sharply serrate ; valva with tuft of strong bristles on inner surface; juxta small, kidney-shaped; vesica without cornuti (Fig. 85) iulina (p. 73) - Uncus with lateral margins smooth, not strongly infolded; valva with scattered fine setae on inner surface ; juxta greatly enlarged , V-shaped ; vesica with single strong cornutus 3 3 Uncus with lateral margins slightly infolded, corners of uncus with blunt protuberance (Fig. 83) berkeleyella (p. 69)Uncus with lateral margins not infolded, corners of uncus not protuberant (Fig. 84) burkerella (p. 70) 4 Lateral margins of uncus straight, apices slightly tapered and hooked iranensis (p. 66) Lateral margins of uncus produced to form a ventrally-directed process 5 5 Lateral processes of uncus short, subapical, double-ridged at extremity (Fig. 80) ussuriensis (p. 67)Lateral processes of uncus elongate, apical, extremity irregularly rounded (Figs 81, 82) moriutii (p. 68) Females (females of iulina are unknown) 1 New World species 2 Old World species 6 2 Antrum one-half or more length of apophyses anteriores 3 Antrum less than one-quarter length of apophyses anteriores 5 3 Posterior margin of eighth sternite with deep, narrow medial emargination (m-shaped), with pair of short, blunt, setose subapical lobes (Fig. 140) burkerella (p. 70) - Posterior margin of eighth sternite ill-defined, surmounted by pair of strong caudally-directed setose digitate processes 4 4 Ostium narrow, ventral (anterior) margin deeply U-shaped; antrum tapered posteriorly (Fig. 143) pythium (p. 72) Ostium broad, ventral (anterior) margin transverse; antrum flared posteriorly . . berkeleyella (p. 69) 5 Ventral margin of ostium extended posteriorly as a pair of setose flaps, m-shaped (Fig. 144) montium (p. 71) - Ostium set in membrane at anterior margin of eighth sternite, with pair of strongly sclerotized setose digitate processes posterior to ostium (Fig. 142) nimbiferum (p. 72) 6 Antrum running anteriorly from ventral lip of ostium without modification iranensis (p. 66) Antrum with strongly sclerotized triangular pocket in ventral wall, lip of pocket forming ventral lip of ostium 7 7 Ventral margin of ostium only slightly concave; antrum narrow, no broader than width of ostium ussuriensis (p. 67) Ventral margin of ostium strongly concave; antrum broadened at one-half to 1-5 x width of ostium (Fig. 141) moriutii (p. 68) Morophagoides iranensis Petersen (Fig. 9) ^Morophagoides iranensis Petersen, 1959: 573, 577. [Nomen nudum.]Morophagoides iranensis Petersen, 1960: 1. Holotype cf , IRAN (LN) [not examined]. ADULT (Zagulajev, 1973: fig. 93; Fig. 9). cf, 17-23 mm; $, 19-28 mm. Vertex and frons cream, some FUNGUS MOTHS 67 brown scales close to eyes. Labial palpus cream, dark brown on outer surface of first and second segment.Maxillary palpus mixed cream and dark brown, 5-segmented, reaching a little more than one-half length ofsecond segment of labial palpus. Antennal scape, pedicel and basal flagellar segments dark brown above;cilia 1-0 x (cf)orO-4 x ($) flagellar diameter. Thorax cream flecked with light brown ;tegula dark brown,cream at apex only. Forewing cream, scattered orange-brown scales along veins, strongly marked withdark brown to form a coarse, cryptic pattern resembling tree-bark. Hindwing light greyiwh cream withslightly darker greyish flecks. Legs cream; fore- and mid-legs strongly marked above with dark brown butpale at articulations; outer surface of mid-tibia with distinctive transverse pale streaks; outer mid-tibial andouter proximal hind tibial spurs 0-7 length of inner spurs. GENITALIA cf (Petersen 1960: fig. 1; Zagulajev, 1968: fig. 4; Zagulajev, 1973: figs 15D, 94). Saccus almostan equilateral triangle; uncus lobes large, divergent, elongately triangular, hooked at the apices.Subscaphium not developed. Juxta, if present, fused with valvae and not recognizable as such; transtillanot developed. Valva triangular, with large conical dorsocaudally-directed medial process; smaller bluntprocess from internal surface of valva close to base of ventral margin. Aedeagus about 10 x as long asbroad in middle, with blunt subapical carina; vesica without cornuti (but large microtrichia possiblypresent). GENITALIA $ (Zagulajev, 1968: fig. 5; Zagulajev, 1973: figs 95, 96). Eighth tergite as long as eighth sternite,with three or four pairs of strong apical setae; eighth sternite forming pair of stout, setose, caudally-directed lobes lateral and dorsal to ostium; ventral margin of ostium slightly concave, protruded deeplyventrad. Antrum thick-walled, longitudinally constricted dorsoventrally, lined with microtrichia, twicelength of eighth sternite. Ductus bursae thin- walled, finely microtrichiate, extending well beyond apices ofapophyses anteriores. Corpus bursae ovate, with pair of small signa, each a flattened cone invaginated intowall of corpus bursae; bursa copulatrix overall more than twice length of apophyses anteriores. DISTRIBUTION. Iran; U.S.S.R. (Caucasus). BIOLOGY Zagulajev (1973) has bred this species from a variety of polypores (Fames, Ganoderma,Polyporus, Coriolus) on Quercus, Zelcova and Parrotia in the Caucasus. MATERIAL EXAMINED. 108 ex., 10 pupae. U.S.S.R.: 1 cf, Adzharia, 15.vi.1969 (Zagulajev); 1 $, Georgia, 19.viii.1972 (Zagulajev) (BMNH).'Central Europe': 1 cf (BMNH). Also a further 105 ex., 10 pupae (ZI) (see Zagulajev, 1973). REMARKS. The specimen from 'Central Europe' bears the registration number [18J76-75: this refers to anidentified synoptic collection of 2500 European Microlepidoptera purchased from Staudinger. Thisspecimen, with the reference number 687, was bought as an example of 'Scardia boleti' (i.e., Morophagachoragelld) (BMNH Accession Register). It must be assumed that its locality label is as incorrect as itsidentification. The three Old World Morophagoides species are allopatric and iranensis is the only Morophagoides tooccur in the western Palearctic region. It differs from ussuriensis and moriutii in lacking any dark scaling onthe third segment of the labial palpus but apart from this difference the three species are externally quitesimilar. The male genitalia of iranensis are distinguished by the divergent uncus lobes with hooked apices,and by the process from the valva being broadly conical: in the other two species the uncus lobes each bear aventrally-directed nodular process, and the process from the valva is slender. The female genitalia ofiranensis lack the conspicuous ventral triangular pocket that forms the ventral margin of the ostium inussuriensis and moriutii; instead, the form of the antrum is conventional. A tentative phylogeny for Morophagoides cannot be resolved because of the lack of material of mostspecies. However, the Old World and New World species may form separate monophyletic groups (theNew World species have a conspicuously modified juxta; the juxta is either lost or entirely fused with thevalva in species from the Old World; species from the Old World have short antennal cilia; the cilia are longin species from the New World). Within these groups three sister-group pairings are apparent: moriutii +ussuriensis (synapomorphies: uncus lobes with nodular processes, ventral margin of ostium infolded toform a triangular pocket), burkerella + berkeleyella (see 'Remarks' for burkerella) and pythium +nimbiferum (see 'Remarks' for pythium). Morophagoides ussuriensis (Caradja)(Figs 10, 80) Scardia ussuriensis Caradja, 1920: 167. Lectotype cf , U.S.S.R. (MINGA), designated by Petersen (1957:593) [not examined]. 68 GADEN S. ROBINSON ADULT (Zagulajev, 1973: fig. 89; Fig. 10). cf , 15-19 mm; <J>, 17-23 mm. Coloration and external structuresimilar to iranensis but maxillary palpus light greyish cream, outer surface of third segment of labial palpuswith a few brown scales. GENITALIA cf (Petersen, 1957: fig. 246; Zagulajev, 1973: figs 14b, 15G, 90; Fig. 80). Saccus triangular;uncus lobes broad, appressed to tegumen for only medial third of their width; lateral margin producedventrad to form a strongly sclerotized knuckle-shaped process. Subscaphium indicated only by slightthickening of diaphragma close to base of valval apodemes. Juxta, if present, fused with valvae and notrecognizable as such; transtilla not developed. Valva triangular, apex consisting of two overlapping lobes,inner lobe extended mesad to form an elongately triangular process with a sinuate apex, outer lobe forminga blunt process close to ventral margin of valva. Aedeagus about 14 x as long as broad in middle, withdigitiform subapical carina; vesica with patch of large spicular microtrichia. GENITALIA $ (Petersen, 1957: fig. 247; Zagulajev, 1973: figs 17G, 18b, 91, 92). Eighth tergite slightlylonger than eighth sternite , with three or four pairs of strong subapical setae ; eighth sternite forming pair ofslender, setose, caudally-directed lobes dorsal to ostium; ventral margin of ostium transverse, protrudingstrongly ventrad. Antrum with conspicuous and strongly sclerotized triangular pocket in ventral wall,antrum itself behind the pocket, elongate, longitudinally constricted dorsoventrally, lined with microt-richia, about 2-5 x length of eighth sternite. Ductus bursae thin-walled, microtrichia at posterior endaround inception of ductus seminalis at junction with antrum. Corpus bursae ovate, with pair of smallpouch-shaped signa invaginated into wall of corpus bursae (Zagulajev figures (1973: fig. 92b) a pair ofadditional minute conical signa situated further posteriorly - 1 believe this to be an individual variation);bursa copulatrix overall more than twice length of apophyses anteriores. DISTRIBUTION. U.S.S.R. (Amur, Chabarovsk, Maritime Territories). BIOLOGY. Zagulajev (1973) associates this species with several genera of trees but no identifications of thehost fungus have been made. Moriuti's detailed description (1976) of the life history of 'ussuriensis' fromJapan refers to moriutli (see below). MATERIAL EXAMINED. 26 ex. , 2 pupae. U.S.S.R.: 1 cf, Vladivostok, 21. vii. 1876 or 19. vii. 1877 (C/iristop/i) (genitalia slide no. 12356; BMNH); 1ex. (head, thorax and fore wings only), same data; 1 cf , Amur, Raddefka, 3.viii.l876 (Chris toph) (genitaliaslide no. 12357; BMNH); 1 $, Ussuri Railway, Chabarovsk, 14.vii.1910 (Borsow) (BMNH). Also afurther 22 ex., 2 pupae (ZI) (see Zagulajev, 1973). REMARKS. Externally similar to iranensis and moriutii, ussuriensis has a restricted distribution on theeasternmost mainland of the U.S.S.R. It is characterized by the conspicuous knuckle-shaped processes ofthe uncus lobes in the male genitalia. With moriutii it shares a conspicuous and diagnostic feature in thefemale genitalia - a conical ventral pouch, triangular in dorso-ventral view, which runs anteriorly from theventral (posterior) margin of the ostium: the antrum proper has its posterior inception in the dorsal wall ofthis pouch although the pouch should be considered a part of the antrum too. Females of moriutii have amuch more deeply concave ventral ostial margin than those of ussuriensis, and the medial region of theantrum is broader. However, the latter difference is subject to considerable intraspecific variation. Morophagoides moriutii sp. n. (Figs 11, 81, 82, 141)[Morophagoides ussuriensis (Caradja); Moriuti, 1976: 86; Moriuti, 1982: 164. Misidentifications.] ADULT (Moriuti, 1976: figs 1, 2; Moriuti, 1982: pi. 2, fig. 17, pi. 236, fig. 3; Fig. 11). cf, 16-17 mm; $, 19-24mm (Taiwanese specimens - cf, 24 mm; $, 24, 29 mm). Coloration and external structure similar toiranensis but, like ussuriensis, maxillary palpus light greyish cream, outer surface of third segment of labialpalpus strongly marked with brown in basal half. GENITALIA cf (Moriuti, 1976: fig. 3; Moriuti, 1982: pi. 248, fig. 9; Figs 81, 82). Saccus triangular, slightlytapered; uncus lobes divergent, somewhat square, outer corners with ventrally-directed blunt process.Subscaphium poorly developed, represented by slight thickening of diaphragma. Juxta, if present, fusedwith valva and not recognizable as such; transtilla not developed. Valva with lobed deep apex and withelongate mesally-directed triangular process on internal surface. Aedeagus 15 x as long as broad in middle(stouter, only 10 x in Taiwanese examples), with digitiform subapical carina; vesica with patch of largespicular microtrichia. FUNGUS MOTHS 69 GENITALIA $ (Moriuti, 1976: figs 4, 5; Fig. 141). Eighth tergite as long as eighth sternite, with three pairs ofstout apical setae and about 8 pairs of scattered smaller setae; eighth sternite forming pair of small setosecaudally-directed lobes dorsal to ostium; ventral margin of ostium concave, protruded strongly ventrad.Antrum with triangular and strongly sclerotized pocket in ventral wall, antrum itself behind the pocket,elongate, very thick-walled, longitudinally constricted dorso-ventrally, lined with strongly sclerotized andconspicuous nodular microtrichia, about 2-5 x length of eighth sternite. Ductus bursae lined with spicularmicrotrichia for almost entire length from inception of ductus seminalis anteriorly. Corpus bursaeelongately ovate , with pair of pouch-shaped signa invaginated into wall ; bursa copulatrix overall more thantwice length of apophyses posteriores. DISTRIBUTION. Japan; Taiwan. BIOLOGY. Moriuti (1976) has described the biology of this species (as ussuriensis) in some detail and figuredthe larva and pupa. Specimens were reared from the fungus Lentinus edodes and its host timber. Lentinus,the shiitake fungus, is a valuable commercial crop used in gourmet cookery. MATERIAL EXAMINED. 9 ex. Holotype cf , Japan: Honsyu, Okayama, Syootyo, ex cultured wood of Lentinus edodes, em. 15. vi. 1968(Inoue) (coll. S. Moriuti, Osaka). Paratypes. Japan: 2 cf, data as holotype (BMNH; coll. S. Moriuti); 1 cf, 1 $, Honsyu, Nara,Katuragisan, ex polyporaceous fungus, em. 31. v. 1961 (Saito) (BMNH; coll. S. Moriuti); 1 $, Honsyu,Nara, Koozindake, 1200 m, 2-3. viii. 1971 (Moriuti) (coll. S. Moriuti). Excluded from paratype series. Taiwan (Formosa): 1 Cf, 2 9, Arizan, 7000', 14, 17, 17. ix. 1906(Wileman) (genitalia slide nos. 12360, 12361; BMNH). REMARKS. Superficially similar to iranensis and ussuriensis, moriutii has a NW. Pacific distribution. Thegenitalia of both sexes are structurally close to those of ussuriensis but in the male the process from eachuncus lobe is apical and irregular, not subapical and knuckle-shaped as in ussuriensis. In the female theventral lip of the ostium is more strongly concave than in ussuriensis and the internal surface of the antrumis more markedly rugose. The marked difference between the illustration of the uncus lobes of the male by Moriuti (1976: fig. 3)and that given here for a Japanese specimen (Fig. 81) is entirely due to the degree of compression of thepreparation: in Fig. 81 the cover-slip is not in contact with the genital armature. Compression of thepreparation of a specimen from Taiwan (Fig. 82) splays out the ventral processes of the uncus in similarfashion to that shown in Moriuti's illustration. Specimens from Taiwan are substantially larger than those from Japan; the aedeagus is slightly thickerand the processes from the uncus are slightly narrower. However, at present I consider the two groups ofspecimens to be conspecific although I exclude Taiwanese material from paratype status. Morophagoides berkeleyella (Powell)(Figs 12, 83) 'Scardia' berkeleyella Powell, [1968]: 303. Holotype cf , U.S.A. (CAS) [not examined].Morophagoides berkeleyella (Powell) Davis, 1983: 5. ADULT (Fig. 12). cf, 14, 16mm. Vertex and frons whitish, brown close to eyes. Labial palpus purple-brownbut pale buff at apex and articulations and on inner surface. Maxillary palpus pale buff, 5-segmented,extending to about one-half length of second segment of labial palpus, basal three segments flecked abovewith purple-brown. Antennal scape, pedicel and flagellum dark brown, paler beneath; cilia 1-5 x (cf)flagellar diameter. Thorax and tegula dark brown flecked with paler scales. Forewing pale buff, stronglyflecked with dark brown to form a speckled cryptic pattern (similar to that of Morophaga morellus in oneexample; in the other much more variegated and similar to that of iulina but more strongly speckled);fringe with pale apical spot, otherwise yellow-grey with grey basal band. Hindwing greyish cream withslightly darker irregular apical spots. Legs greyish ochreous; fore- and mid-legs dark grey above and onsides, articulations conspicuously white; outer mid-tibial spur and outer proximal hind tibial spur 0-7 lengthof inner spurs. GENITALIA cf (Powell, [1968]: fig. 6; Fig. 83). Saccus narrowly triangular, concave laterally, as long as valva+ juxta; uncus lobes moderately sclerotized, sparsely setose, fused dorsally and curved ventrad at sides,thus semicylindrical; corners produced slightly to form pair of shallow triangular processes. Subscaphiumnarrow, hardly sclerotized, spatulate anteriorly. Juxta partially fused with valvae, bipartite, forming pair of 70 GADEN S. ROBINSON large, blade-shaped caudally-directed processes extending well beyond apices of valvae. Valva short,globose, with digitate apical process. Aedeagus slender, 20 x as long as broad, with strong subapicalcorniform carina one-seventh length of aedeagus; vesica with fine spicular cornuti (microtrichia). GENITALIA 9 (Powell, [1968]: fig. 7). (From Powell's description and illustration.) Eighth tergite slightlylonger than eighth sternite; eighth sternite forming pair of stout setose lobes caudal to ostium; ventralmargin of ostium irregular. Antrum short, elongately funnel-shaped, one-half length of apophysesanteriores. Ductus and corpus bursae pyriform, thin-walled, corpus bursae with pair of small pouch-shaped signa. DISTRIBUTION. U.S.A. (California). BIOLOGY. Powell ([1968]) reared the type-series of this species from Polyporus gilvus on fallen Quercusagrifolia, from dead stems of Lupinus, and from a log of Lithocarpus: these data are reiterated byLawrence & Powell (1969). MATERIAL EXAMINED. 2 ex. U.S.A.: 1 cf (paratype), California, 2 m. SE. of Canyon, Contra Costa County, ex Polyporus gilvus onfallen Quercus agrifolia, coll. 5.H., emerged 9.iv.l967 (genitalia slide no. 2240 [Powell]; BMNH); 1 cf ,California, Burney Mt, 14.viii.1930 (genitalia slide no. 13107; BMNH). REMARKS. Most closely related to M. burkerella, berkeleyella is, however, a much smaller species. Powell([1968]) suggests that it is less distinctly marked than burkerella but the male from Burney Mt is stronglyvariegated. The male genitalia of the two species differ substantially only in the form of the uncus lobes(compare illustrations) which are broader and flatter in burkerella than in berkeleyella. The femalegenitalia differ in that the signa are substantially larger in burkerella than in berkeleyella, the antrum is notas tapered and the ventral margin of the ostium is evenly convex. Whereas the caudal lobes of the eighthsternite are elongate and terminal in berkeleyella, they are very short and subterminal in burkerella. Morophagoides burkerella (Busck)(Figs 13, 84, 140) Scardia burkerella Busck, 1904: 777. Holotype $, U.S.A. (NMNH) [examined]. Scardia gracilis Walsingham, 1907: 225. Holotype cf , U.S.A. (NMNH) [examined]. [Synonymized by Davis (1983: 5).]Scardia caryophylella Busck, 1908: 92. Holotype $ , U.S.A. (NMNH) [examined]. [Synonymized by Davis (1983: 5).]Scardia errandella Busck, 1908: 94. Holotype cf , U.S.A. (NMNH) [examined]. [Synonymized by Davis (1983: 5).] ADULT (Fig. 13). cf$, 19-28 mm. Vertex and frons cream, brown close to eyes. Labial palpus brown,cream at apex and articulations and on inner surface. Maxillary palpus 5-segmented, extending to apex ofsecond segment of labial palpus, buff, strongly flecked above with brown. Antennal scape, pedicel andbasal segments of flagellum dark brown above, flagellum lighter distally; cilia 1-5 x (cf) or 0-3 x ($)flagellar diameter. Thorax and tegula cream, dark brown in anterior half but mixed with cream on thorax.Forewing light yellowish buff, strongly speckled and blotched with dark brown to form a dense, crypticpattern resembling tree-bark, usually with a strongly defined medial band; fringe barred, with well-definedgrey basal line. Hindwing cream tinted with grey, with irregular terminal grey line; with apical grey spotsextended into fringe; fringe with grey basal line. Legs pale yellowish buff; foreleg, mid-leg and hind tarsusstrongly marked above and on sides with dark brown but pale at articulations; outer mid-tibial spur 0-8length of inner; outer proximal hind tibial spur 0-7 length of inner spur. GENITALIA cf (Fig. 84). Similar to those of berkeleyella (q.v.) but uncus lobes broader, flatter, the sides notdeeply folded ventrad. GENITALIA $ (Fig. 140). Eighth tergite slightly shorter than eighth sternite, with three pairs of elongatesetae at caudal margin and a medial pair of sensilla basiconica; eighth sternite almost square but flaredanteriorly, with narrow medial emargination posteriorly and pair of shallow subterminal setose lobes;ostium protuberant, ventral margin slightly convex; lip of antrum with three pairs of strong setae oninternal surface. Antrum elongate, more than one-half length of apophyses anteriores, very thick-walled,with coarsely rugose internal surface. Ductus bursae short, thin-walled, lined with microtrichia, extendingto apices of apophyses anteriores. Corpus bursae ovoid, thin-walled, with pair of large and stronglysclerotized shark's fin-shaped signa. FUNGUS MOTHS 71 DISTRIBUTION. U.S.A.: California (Powell, [1968] - asgracilis) but possibly restricted to the Coast Range(Lawrence & Powell, 1969); Utah; Washington State; Pennsylvania (Forbes, 1923). Canada: BritishColumbia. BIOLOGY. Recorded by Lawrence & Powell (1969) from a wide range of host Polyporaceae in Californiaand recorded by Powell ([1968]) as possibly having two generations per year. Lawrence & "Powell foundthis to be the commonest scardiine that they encountered in breeding from fungi and found it to be 'limitedprimarily to Polyporaceae with relatively large and persistent sporophores, or large colonies with extensivesterile tissue in species with small sporophores'. MATERIAL EXAMINED. 37 ex. Holotype $ (of burkerella), U.S.A.: Washington, Hoquiam, bred from Tsuga heterophylla (Burke)(genitalia slide no. 18654; NMNH). Holotype cf (of gracilis), U.S.A.: California (Beutenmueller)(genitalia missing) (NMNH). Holotype $ (of caryophylella), U.S.A.: California, Fieldbrook, 28.V.1903(Barber) (abdomen missing) (NMNH). Holotype cf (of errandella), U.S.A.: [Washington State], (Beuten-mueller) (abdomen missing) (NMNH). U.S.A.: 30 ex., California, various hosts, dates and localities (genitalia slide no. 12400; BMNH; UC,Berkeley); 1 cf , 1 $, Utah, Dividend, vii.1926 (BMNH). Canada: 1 cf , British Columbia, Fraser Mills,12.vii.1921 (E.H.B.) (genitalia slide no. 12399; BMNH). REMARKS. This is the larger and commoner of the two North American Morophagoides species: fordifferentiation from berkeleyella see 'Remarks' for that species. M. burkerella and berkeleyella areprobably sister-groups, exhibiting close similarity in the male genitalia, notably in the modification of thejuxta of both species to form a pair of elongate, almost rectangular processes, and in the reduction of thevalva to a globose shape with a terminal digitate process. Morophagoides montium (Walsingham) comb. n. (Figs 14, 144)Phycis montium Walsingham, 1914: 359. Holotype $, PANAMA (BMNH) [examined]. ADULT (Fig. 14). $, 17 mm. Vertex and frons cream, mixed strongly with brown close to eyes. Labialpalpus brown but cream at apex and articulations and on inner surface except for brown-dusted medial areaon second segment. Maxillary palpus buff, brown above on basal segments, 5-segmented, extendingprobably no more than two-thirds length of second segment of labial palpus. Antennal scape, pedicel andbasal flagellar segments dark brown, distal segments paler; cilia 0-3 x flagellar diameter. Thorax and tegulabrown, slightly paler posteriorly. Forewing cream speckled with orange-brown and marked more stronglywith dark brown to form a complicated cryptic pattern; dorsum generally lacking dark markings and thusconspicuously pale; fringe strongly chequered. Hindwing light brownish grey. Legs damaged but forelegand mid-leg dull buff strongly marked above with grey-brown, pale at articulations. GENITALIA cf . Unknown. GENITALIA $ (Fig. 144). Eighth tergite almost circular, as long as eighth sternite, with three pairs- ofelongate setae and one or two shorter setae at caudal margin, with numerous pits (?sensilla coeloconica)scattered close to caudal margin; eighth sternite forming a strongly sclerotized plate ventral to ostium,terminating caudally in a pair of lobes; each lobe with a single elongate seta and numerous smallerspine-like setae at the tip; eighth sternite hardly sclerotized anteriorly; antrum triangular, short, stronglysclerotized; inception of ductus seminalis at apex of antrum. Ductus bursae thin-walled, lined with spicularmicrotrichia (broken and a section missing so length uncertain). Corpus bursae thin-walled, pyriform, witha pair of shallow pocket-shaped signa that are rectangular in lateral view. DISTRIBUTION. Panama.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype $, Panama: Volcan de Chiriqui, 2000-3000', 1881 (Champion} (genitalia slide no. 15279;BMNH). REMARKS. This species is small, of comparable size to M. nimbiferum, but considerably smaller thanpythium or iulina. Its wing pattern is not as variegated as that of iulina but not as uniform in the anterior halfof the wing as in pythium or nimbiferum. The hindwing is darker than that of iulina or pythium. The femalegenitalia are distinctive, characterized by the short antrum and by the lobes ventral to the ostium being 72 GADEN S. ROBINSON broad, situated far posteriorly, and with only a shallow emargination between them. In pythium the antrumis elongate, the ostium set in a deep emargination between the narrow processes of the eighth sternite; innimbiferum the ostium is anterior, the processes arising caudally, and the eighth sternite strongly andcomplexly modified dorsal to the processes. Walsingham's original description oi montium was based on five specimens. One of these was expresslycited as 'Type' and two others (in NMNH) cited as paratypes. The remaining two specimens are implicitlyexcluded from type-status. They are, in fact, not conspecific with the holotype and are described below asnew species -pythium and nimbiferum. Morophagoides pythiumsp. n. (Figs 15, 143)[Phycis montium Walsingham; Walsingham, 1914: 360. Partim - 1 $ only. Misidentification.] ADULT (Fig. 15). 9 , 24 mm. Vertex and frons cream, brown close to eyes. Labial palpus brown, cream atarticulations; cream flecked with light brown on inner surface. Maxillary palpus pale buff flecked withbrown; basal two segments strongly marked above with dark brown; 5-segmented, extending to aboutthree-quarters length of second segment of labial palpus. Antennal scape, pedicel and basal flagellarsegments dark brown above, distal segments lighter; cilia 0-5 x flagellar diameter. Thorax and tegulabrown anteriorly, cream posteriorly. Forewing cream, lightly flecked with orange-brown, strongly markedwith dark brown to form a cryptic 'moss-pattern' but termen, costa and dorsum less strongly marked andthus conspicuously paler; conspicuous small white subtornal spot. Hindwing pale greyish cream. Legs allwanting except foreleg - buff beneath, grey-brown above and on sides but pale at articulations. GENITALIA cf . Unknown. GENITALIA 9 (Fig. 143). Eighth tergite markedly longer than eighth sternite, with three pairs of strongsubapical setae; eighth sternite short, folded laterally to accommodate bases of apophyses, with pair ofnarrow setose processes either side of ostium, folded anteriorly to ostium to form a shallow pouch. Antrumelongate, thick-walled, about one-half length of apophyses anteriores. Ductus bursae lined with micro-trichia close to antrum, as long as antrum, thin-walled. Corpus bursae thin-walled, with pair of thin shark'sfin-shaped signa set in sclerotized base-plates, each base-plate with and surrounded by scattered micro-trichia. DISTRIBUTION. Costa Rica.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype 9, Costa Rica: Volcan de Irazu, 6000-7000', 18 (Rogers) (genitalia slide no. 12383;BMNH). Paratype. Costa Rica: 1 9 , Puntarenas Prov. , 6 km S. of San Vito, 2 or 5. v. 1967 (Veiro) (UC). REMARKS. With nimbiferum and montium, pythium forms a group of dark-patterened Neotropical speciesfrom which iulina is excluded by dint of its strongly variegated pattern. M. pythium is larger and has palerhindwings than montium and nimbiferum. The genitalia are distinctive in that the paired processes of theeighth sternite do not overly the ostium as in montium, are not entirely caudal to it as in nimbiferum, butform the lateral margins of the ostium and extend caudally. Like nimbiferum, the eighth tergite isconsiderably longer than the eighth sternite but the antrum is distinctly thicker and more elongate. Apartfrom nimbiferum, this is the only Morophagoides species to have microtrichia associated with the signa. M.pythium and nimbiferum may be sister-groups (synapomorphies: microtrichia in corpus bursae; eighthsternite 'shouldered' to accommodate bases of apophyses anteriores; ostium close to anterior margin ofeighth sternite). Morophagoides nimbiferum sp. n. (Figs 16, 142)[Phycis montium Walsingham; Walsingham, 1914: 360. Partim -19 only. Misidentification.] ADULT (Fig. 16). 9> 16 mm. Vertex and frons brownish cream, brown close to eyes. Labial palpus brownon outer surface but pale at apex and articulations; inner surface cream, small patch of brownish scales inmiddle of second and middle of third segment. Maxillary palpus light buff, strongly flecked above with FUNGUS MOTHS 73 brown, 5-segmented, reaching three-quarters length of second segment of labial palpus. Antennal scapeand pedicel dark brown above, flagellum damaged and mostly lost. Thorax and tegula brown anteriorly,light buff posteriorly. Forewing cream, sparsely flecked with orange-brown, strongly marked with darkbrown; however, termen, dorsum and costal spots conspicuously pale. Hindwing light grey. Legs dull buff,fore- and mid-legs dark grey-brown above and on sides but pale at articulations and across mid-tibia; outermid-tibial spur 0-7 length of inner; outer proximal hind tibial spur 0-6 length of inner spur. GENITALIA cf Unknown. GENITALIA 9 (Fig. 142). Eighth tergite considerably longer than eighth sternite, with three pairs of largeand two pairs of small marginal setae and about 10 scattered sensilla (basiconica and (?) coeloconica);eighth sternite short, strongly folded transversely, with strongly sclerotized 'shoulders' at bases ofapophyses; medially with three convergent sclerites forming a flexible blunt process overlain by a lobatemembranous outgrowth of the ovipositor; paired setose processes arising caudally to ostium, stronglydivergent, each bearing a pair of large setae and numerous smaller setae at apex; ostium close to anteriormargin of sternite. Antrum short, thick-walled, but hardly sclerotized, less than one-quarter length ofapophyses anteriores. Ductus bursae half length of apophyses, lined with microtrichia in posterior half.Corpus bursae ovoid, with pair of thorn-shaped signa set in sclerotized base-plates, each base-plate withand surrounded by scattered microtrichia. DISTRIBUTION. Guatemala.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype $, Guatemala: Volcan de Atitlan, 2500-3500', [xii.1880], (Champion) (genitalia slide no.12385; BMNH). REMARKS. This species is similar in size and external appearance to montium but the forewing pattern is lessvariegated. The genitalia are distinctive, however; the eighth sternite is strongly folded with elongateprocesses that arise posteriorly, and the ostium is close to the anterior margin, a configuration quite unlikethat of any other species. Additionally, the signa differ from those of all other Morophagoides in beingslender and thorn-like rather than laterally compressed and pocket-shaped. Withpythium, nimbiferum isthe only Morophagoides species to have microtrichia associated with the signa. Morophagoides iulina (Walsingham) comb. n. (Figs 17, 85)Phycis iulina Walsingham, 1914: 359. Holotype cf , GUATEMALA (BMNH) [examined]. ADULT (Walsingham, 1914: pi. 10, fig. 17 (colour); Fig. 17). cf , 26 mm. Vertex and frons whitish, lightbrown close to eyes. Labial palpus pale buff speckled strongly with dark brown on first segment and onbasal two-thirds of second segment. Maxillary palpus whitish, 5-segmented, elongate, extending beyondapex of second segment of labial palpus, basal three segments flecked above with dark brown. Antennalscape and pedicel buff flecked with dark brown; flagellum pale buff; cilia 2-5 x flagellar diameter. Thoraxand tegula dark purple-brown with only a few whitish scales posteriorly. Forewing cream mottled with paleorange-brown; strong W-shaped dark brown mark occupying most of wing; strong dark brown costal spots;terminal spots small but extending through fringe and giving a conspicuously chequered appearance to thefringe. Hindwing slightly greyish silk-white. Mid-legs missing; fore-leg strongly marked above with darkpurple-brown; hind-leg pale buff, tibial spurs damaged. GENITALIA cf (Fig. 85). Saccus broadly triangular, apex rounded; uncus lobes almost semicircular, stronglyfolded, with conspicuously serrate inner edge. Subscaphium a broad but almost imperceptible thickeningof the diaphragma. Juxta, if present, represented by a pair of shallow and inconspicuous ventral lobes atbases of valvae; transtilla represented by a pair of shallow membranous lobes dorsal to anellus. Valvasimple, with subapical tuft of inwardly-directed strong spicular setae. Aedeagus about 8 x as long as broadat base, lacking carinae; vesica with minute spicular cornuti (microtrichia). GENITALIA $. Unknown.DISTRIBUTION. Guatemala.BIOLOGY. Unknown. 74 GADEN S. ROBINSON MATERIAL EXAMINED. 1 ex. Holotype cf, Guatemala: Totonicapam, 8500-10,500', [viii.1880], (Champion) (genitalia slide no.12398; BMNH). REMARKS. Among the American Morophagoides species, iulina is distinguished by its pale forewingground-colour with strong W-shaped mark; it is large, of similar size to burkerella, and the female couldwell be the largest of the genus. The male genitalia are distinguished by the serrate uncus lobes and by thecharacteristic modified setae on the valvae. It is the only species of Morophagoides that does not have ablunt subapical carina on the aedeagus. The affinities of iulina are obscure . The structure of its genitalia set it well apart from the other Americanspecies of which the males are known (berkeleyella and burkerella). However, the ventral lobes at the basesof the valvae may well be homologous with the enormously developed pair of lobes (assumed to be thejuxta) in these species. MONTESCARDIA Amsel Montescardia Amsel, 1952: 139. Type-species: Euplocamus tessulatellus Zeller, 1846: 178, by originaldesignation and monotypy. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia shorter than 1-5 x flagellardiameter. Scape with more than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpus5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outer mid andproximal hind tibial spurs >0-4 length of inner spurs. Forewing with 7? 3 and R 4 separate; M 2 , M 3 and CuA\separate; mottled coloration forming cryptic, coarse 'moss' pattern. Male with coremata in eighthabdominal segment; coremata not associated with apodemes. Male genitalia with complex uncus separatedfrom tegumen by narrow band of membrane; tegumen broken dorsally by at least a membranous sutureline; valva lacking basal setose lobe on inner surface; apex of valva not forming ventral hook or hooks,without spines; valvae fused ventrally into a single movable complex, valvae without longitudinal cleft;saccus wider than long; juxta complex, entire, not divided medially; vesica lacking spicular cornuti;aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Juxta complex, enveloping aedeagus to form an elaborate, double-layeredand partially fused intromittent complex; subscaphium broad, triangular; female with eighth tergiteexplanate, forming pair of broad lateral flanges (but not in fuscofasciella). DISTRIBUTION. Holarctic. BIOLOGY. See Montescardia tessulatellus (Zeller). Key to species of Montescardia Males (male genitalia of fuscofasciella are unknown) 1 Maxillary palpus elongate, reaching apex of second segment of labial palpus .... fuscofasciella (p. 76)Maxillary palpus short, only just reaching base of second segment of labial palpus 2 2 Caudal apex of juxta + aedeagus-complex fused, boat-shaped tessulatellus (p. 74) Caudal apex of juxta + aedeagus-complex bilobed, lobes separated by their own width kurenzovi (p. 75) Females 1 Maxillary palpus elongate , reaching apex of second segment of labial palpus ; posterior margin of ostium a protuberant ridge; ostium teardrop-shaped (Fig. 146) fuscofasciella (p. 76) Maxillary palpus short, only just reaching base of second segment of labial palpus; posterior margin of ostium not defined, eighth sternite merging into membrane of ovipositor 2 2 Eighth tergite 1 -3 x or more wider than long tessulatellus (p. 74) Eighth tergite roughly as wide as long kurenzovi (p. 75) Montescardia tessulatellus (Zeller)(Fig. 18) Euplocamus tessulatellus Zeller, 1846: 178. LECTOTYPE cf, AUSTRIA (BMNH), here designated[examined]. FUNGUS MOTHS 75 ADULT (Fig. 18). C?$, 22-29 mm. Vertex and frons straw-coloured, tufts above eyes darker. Labial palpusstraw-coloured, strongly flecked with brown on outer surface of second and third segments but pale atarticulations. Maxillary palpus whitish, very short, only just reaching second segment of labial palpus,5-segmented. Antennal scape and pedicel straw-coloured, dark brown above; flagellum grey-brownabove; cilia 1-0 x (cf) or 0-7 x ($) flagellar diameter. Thorax and tegula cream, strongly flecked withbrown. Forewing cream, marked with orange-brown along veins and strongly flecked with dark brown toform ill-defined jagged medial and postmedial bands; medial costal spot strongly defined; strong dark spotat end of cell. Hindwing pale grey, mottled at margin and fringe. Legs straw-coloured, fore-, mid- andhindlegs strongly marked above and on outer surfaces with dark brown, but pale at articulations; outermid-tibial spur 0-6 length of inner; outer proximal hind tibial spur 0-7 length of inner spur. GENITALIA cf (Petersen, 1957: fig. 242, pi. 12, figs 1, 2; Zagulajev, 1973: figs 16A, 80, 81). Saccus short,triangular; uncus lobes strongly sclerotized, separated from tegumen by relatively broad membranousarea, setose only apically and laterally, with irregularly serrate caudal margin. Subscaphium triangular.Juxta enveloping and partly fused with aedeagus to form an elaborate intromittent complex; transtilla notdeveloped. Valvae fused ventrally to form a united complex with a narrowly V-shaped medioventralemargination; apices of valvae simple, rounded. Aedeagus fused with juxta to form intromittent complex;vesica without cornuti. GENITALIA $ (Petersen, 1957: fig. 243; Zagulajev, 1973: figs 82, 83). Eighth tergite longer than eighthsternite, caudal one-quarter membranous, with four pairs of strong and elongate subapical setae posteriorto irregular margin of sclerotization and with about six scattered pairs of sensilla basiconica; tergiteexplanate anteriorly to form conspicuous pair of quadrate lateral flaps; eighth sternite U-shaped, bases ofarms of 'U' strongly folded, between arms a broad and deep depression leading to ostium which is overlainby a strong semicircular sclerite. Antrum short, longitudinally folded and with very fine transversestriations; inception of ductus seminalis at anterior end of antrum. Ductus bursae thick-walled, slightlylonger than antrum. Corpus bursae large, ovate, extending well beyond apices of apophyses anteriores;signa absent. DISTRIBUTION. Sweden; Norway; Finland (Krogerus et al., 1971); Italy (Petersen & Gaedike, 1979);Sardinia (Amsel, 1952); central Europe including East and West Germany; Yugoslavia; Austria; Ruma-nia; Hungary; Czechoslovakia; Poland; U.S.S.R. (European Region, Crimea, E. Siberia, Primorskregion); Mongolia. BIOLOGY. Petersen (1969) has recorded this species from pore-fungi and from dead wood of Picea andFagus. MATERIAL EXAMINED. 82 ex. Lectotype C", [Austria]: Semm[e]ring [approx. 80 km. SW. of Vienna], vi (Mann) (BMNH). Poland: 1 $ (paralectotype), [approx. 15 km W. of Klodzkol] 'Sfd' ['between Reinerz and theSeefelder'], 29.vii (Zeller} (BMNH). 80 ex., various localities (see 'Distribution') (BMNH; ZI). (Fordetails of ZI holdings see Zagulajev (1973).) REMARKS. This species is characteristically patterned and, with its geographical context, should beidentifiable without recourse to dissection. Unlike all other Palaearctic scardiines with this wing-pattern,all veins in the forewing are free and the forewing appears correspondingly broader than in, say,Morophaga (but beware of Morophagoides iranensis, worn specimens of which might be mistaken forMontescardia). Unlike most other scardiines, the hind tarsus in Montescardia is dark brown above, thepattern broken by pale scaling at the articulations. In the female, the eighth tergite is deeply recessed intothe intersegmental membrane, forming a well-defined dorsal pouch with a strongly microtrichiate dorsalinternal surface. This species and M. kurenzovi may be separated from the North American fuscofasciellaby their having very short maxillary palpi: in fuscofasciella the palpi would, if unfolded, reach the apex ofthe second segment of the labial palpus. Separation of tessulatellus and kurenzovi is more problematical:the latter taxon is probably no more than a slight geographical variant of tessulatellus from the edge of itsrange. However, further material of kurenzovi is required to clarify its status. It may be tentativelyseparated from tessulatellus using the characters described in the key and below. Montescardia kurenzovi (Zagulajev) comb. n. Scardia kurenzovi Zagulajev, 1966: 637. Holotype cf, U.S.S.R. (ZI) [examined].ADULT. C"$, 21-5-23-5 mm. Coloration and external structure similar to tessulatellus. 76 GADEN S. ROBINSON GENITALIA cf (Zagulajev, 1966: fig. 2A; Zagulajev, 1973: figs 15A, 15b, 16B, 84). Similar to those oftessulatellus but apex of juxta-aedeagus complex with pair of separated lobes. GENITALIA $ (Zagulajev, 1966: figs 2b, 2B; Zagulajev, 1973: figs 17A, 17b, 19A, 85, 86). Similar to those oftessulatellus but eighth tergite not as broadly explanate, outer corners of tergite acute. DISTRIBUTION. U.S.S.R. (Maritime Territory); Kurile Is (Kunashir) (Zagulajev, 1973).BIOLOGY. Unknown. MATERIAL EXAMINED. 5 ex. Holotype cf , U.S.S.R.: Maritime Territory, Khasan district, Verkhnyaya Sidimi, 8.vi.l950 (Zagulajev}(ZI). U.S.S.R.: 1 9 (paratype), data as holotype but nr Vladivostok, 8. i\.l956(Omelko); 1 $ (paratype),dataas holotype but Suchan, source of the Sitsa River, 28.viii.1928 (Kurentsov} (ZI); 1 $, data as first paratypebut 24.vii. 1950 (Zagulajev} (ZI); 2 $, Kurile Is., Kunashir, 5-7. vii. 1962 & 6.vii.l964 (Krlvolutskaya} (21-only one specimen present when examined by author). REMARKS. It is unlikely that this taxon represents a good species (see 'Remarks' for tessulatellus}; thedifferences between it and tessulatellus are subtle in the extreme. The BMNH collection contains a maleMontescardla from Khabarovsk ('Chabarovsk') on the Amur River. This locality is at the edge of thegeographical range of kurenzovi as recorded by Zagulajev (1973) and well beyond the eastern limit oftessulatellus recorded by the same author. However, this specimen is an entirely typical example oftessulatellus. Montescardia fuscofasciella (Chambers) comb. n.(Figs 19, 146) Euplocamus (?) fuscofasciella Chambers, 1875: 257. LECTOTYPE $, U.S.A. (MCZ), here designated [examined].Scardiapravatella Busck, 1908: 94. Holotype cf , U.S.A. (NMNH), here designated [examined]. [Synony- mizedbyDavis(1983:5).] ADULT (Fig. 19). , 27 mm. Vertex and frons pale buff. Labial palpus off-white, strongly flecked withbrown on outer surface. Maxillary palpus cream, flecked with brown above on basal segments, elongate,5-segmented, reaching apex of second segment of labial palpus. Antennal scape, pedicel and basal flagellarsegments dark brown above; flagellum medium brown; cilia 0-6 x () flagellar diameter. Thorax andtegula speckled brown and cream. Forewing brownish cream, ill-definedly orange-brown along veins,strongly speckled with dark brown to form strong solid blotches and a well-defined medial transversefascia. Hindwing light grey-brown, ill-defined paler mottling at apex. Legs greyish buff, fore- and mid-legsstrongly marked above with dark brown but pale at articulations; hind tarsus also dark brown above butarticulations pale; outer mid-tibial spur 0-4 length of inner; outer proximal hind tibial spur 0-5 length ofinner spur. GENITALIA cf . Unknown. GENITALIA $ (Fig. 146). Eighth tergite markedly longer than eighth sternite, with three or four pairs ofstrong subapical setae; eighth sternite moulded round large pyriform ostium, apex emarginate, laterocaud-al lobes each with pair of strong setae . Antrum very short , dorsal wall rugose ; inception of ductus seminalisalmost at level of ventral margin of ostium. Ductus bursae broad, with lepidote (?microtrichiate) innersurface, merging with elongately ovate corpus bursae. Corpus bursae thin- walled, extending slightlyanteriorly beyond apophyses anteriores; signa absent. DISTRIBUTION. U.S.A. (Kentucky (?), Pennsylvania, North Carolina, Texas (?)).BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Lectotype $ (of fuscofasciella}, U.S.A.: 'Kentucky' [but more probably Texas - see 'Remarks'](Chambers} (genitalia slide no. 2563 [Davis]; MCZ). Holotype cf (of pravatella} (abdomen missing),U.S.A.: Pennsylvania, New Brighton, 23.viii.1902 (Merrick) (NMNH). U.S.A.: 1 ex. (abdomen missing), North Carolina, 1884 (Morrison) (BMNH). REMARKS. Distinguished from its congeners by its elongate maxillary palpus, fuscofasciella is a distinctlydark, speckly species somewhat reminiscent of Morophaga morellus. Apart from Morophagoides bur- FUNGUS MOTHS 77 kerella it is the only North American scardiine of this size and wing-pattern. In burkerella the thorax andthe apices of the tegulae are cream , not with mixed brown and cream scales , and the hindwing is pale creamwith a slight greyish tint, considerably paler than the light grey-brown hindwing oi fuscofasciella. The female genitalia are very different from those oftessulatellus; the eighth tergite is not explanate, theinner surface of the ductus bursae is lepidote, and the shape of the eighth sternite is markedly different. Thestructure of the male genitalia, presently unknown, may prove fuscofasciella not to be a Montescardia. Itspresent placement should be considered provisional. Chambers (1875: 258) refers to 'the foregoing descriptions of "Teneina [sic] from Texas" . . . theconclusion of a series' . The block of descriptions of Texan material appears to begin on p . 250 with Gelechiasaphirinella and to include fuscofasciella. Although the lectotype is labelled 'Kentucky', the label is not byChambers and may be erroneous. The type-locality must remain in question as the distribution of thisspecies is hardly known. B YTHOCRA TES Meyrick Bythocrates Meyrick, 1919: 268. Type-species: Bythocrates drosocycla Meyrick, 1919: 268, by monotypy. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia shorter than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpuswith fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width of head.Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3 and R 4 separate ; M 3and CuAi stalked or very closely approximated at base; mottled coloration forming cryptic, coarse 'moss'pattern. Male lacking coremata in eighth abdominal segment. Male genitalia with simple uncus- a pair ofsetose lobes - separated from tegumen by narrow band of membrane; tegumen unbroken, completelysclerotized dorsally; valva lacking basal setose lobe on inner surface; apex of valva not forming ventralhook or hooks, without spines; valvae fused ventrally into a single movable complex, valvae withoutlongitudinal cleft; saccus wider than long; juxta simple, entire, not divided medially; vesica with spicularcornuti; aedeagus smooth-surfaced, without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Valvae entirely fused ventrally, with no trace of a suture line (line effusionalso difficult to discern in some species of Daviscardia); corpus bursae of female with four stronglysclerotized, pocket-shaped (or wedge-shaped) signa; ductus bursae with oblique regular constrictions;female eighth tergite entirely divided medially. DISTRIBUTION. Neotropical region - Guyana, Trinidad.BIOLOGY. Has been bred from Poly poms in Trinidad (label data). Bythocrates drosocycla Meyrick (Figs 20, 145)Bythocrates drosocycla Meyrick, 1919: 268. Holotype cf , GUYANA (BMNH) [examined]. ADULT (Clarke, 1970: pi. 16, figs 1, Ib; Fig. 20). cf , 11 mm; $, 14 mm. Vertex and frons dull buff. Labialpalpus dark brown but pale buff on inner surface of second segment. Maxillary palpus buff-cream, a fewdark brown scales above close to base, short, 4-segmented, extending only to basal one-fifth of secondsegment of labial palpus. Antennal scape and pedicel dull buff, flagellum dark brown; cilia 1-0 x (cf) or0-7 x ($) flagellar diameter. Thorax and tegula dark brown. Forewing purple-brown with very slightlydarker mottling forming an indistinct 'moss' pattern; costa with six or seven inconspicuous paler yellowishspots; termen with interrupted line of yellowish spots. Hindwing purple-brown. Legs dull buff; outermid-tibial spur 0-5 length of inner; outer proximal hind tibial spur 0-6 length of inner spur. GENITALIA cf (Clarke, 1970: pi. 16, figs Ic-le). Saccus shallow, triangular; uncus lobes slender, elongate,digitiform, widely separated. Subscaphium strongly sclerotized, shuttle-shaped. Juxta not identified;transtilla forming V-shaped arch between costae of valvae. Valvae fused; valval complex elongate,strongly sclerotized and setose at apex, with transverse ridge at three-quarters and pair of obliquesclerotized bars at one-half, these fused with transtilla. Aedeagus short, broad and stumpy, only 4 x as longas broad, smooth-surfaced; vesica with minute spicular cornuti (microtrichia). GENITALIA $ (Fig. 145). Eighth tergite divided medially, slightly longer than eighth sternite, with threepairs of stout terminal bristles; eighth sternite deeply folded transversely to form a triangular sterigma with 78 GADEN S. ROBINSON strongly sclerotized truncated apex overlying ostium and bearing pair of elongate setae plus single(?supernumerary) seta and a few scattered sensillae (?basiconica). Antrum not developed. Ductus bursaeas long as apophyses anteriores, thin-walled, lined anteriorly with granular microtrichia from inception ofductus seminalis (at twice length of eighth sternite from ostium), oblique constrictions forming acharacteristic pattern. Corpus bursae ovate, posterior half with regular transverse constrictions, thin-walled, with four stout, wedge-shaped and finely spinose signa. DISTRIBUTION. Guyana, Trinidad. BIOLOGY. Specimens have been reared from Polyporus in Trinidad. MATERIAL EXAMINED. 3 ex. Holotype d", Guyana: Bartica, ii.1913 (Parish) (genitalia slide no. JFGC 6639; BMNH).Trinidad: 1 cf , 1 $, reared from Polyporus, 1922 (Urich) (genitalia slide nos 1664, 13110; BMNH). REMARKS. This small, dark species may be recognized simply by its external appearance: it bears a slightresemblance to Diataga but is smaller, darker and broader-winged. Forewing veins RT, and R 4 are separate,unlike Diataga. Both male and female genitalia are distinctive, the male in that the valvae are stronglyfused and the elongate uncus lobes are widely separated, and the female in that it is the only scardiine withmore than two signa. DAVISCARDIA gen. n. Type-species: Scardia color adella Dietz, 1905: 25. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia longer or shorter than1-5 x flagellar diameter. Scape with more than 15 pecten bristles. Interocular index (male) greater than1-0. Maxillary palpus with fewer than 5 segments; pilifers present; second segment of labial palpus shorterthan width of head. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3and /? 4 separate; M 2 , A/ 3 and CuA\ separate; pattern consisting of pale termen and dorsum on darkerground-colour. Male lacking coremata in eighth abdominal segment. Male genitalia with complex uncusfused with tegumen ; tegumen unbroken , completely sclerotized dorsally ; valva lacking basal setose lobe oninner surface; apex of valva forming ventral hook or hooks, or with spines; valvae fused ventrally into asingle movable complex, valvae without longitudinal cleft; saccus wider than long; juxta, if present, fusedwith valvae and not recognizable as such; vesica with spicular cornuti; aedeagus smooth-surfaced, withoutcarinae, but minute spicular carinae present in two species. CONSPICUOUS AUTAPOMORPHIES. None; however, this is the only group of species with a pale apex anddorsum in the forewing which lacks coremata; the deep, almost box-like fused valvae are also characteris-tic. DISTRIBUTION. Neotropical region; Nearctic region -U.S.A. (southern states).BIOLOGY. See Daviscardia coloradella (Dietz). Key to species of Daviscardia Males (males of hypocritella and lupulella are unknown) 1 Tegumen with caudally-directed lateral digitate processes 2 - Tegumen not produced laterocaudally 6 2 Tegumen with two pairs of caudally-directed processes (Fig. 95) species A (p. 84) Tegumen with single pair of caudally-directed processes 3 3 Aedeagus slender, about 20 times as long as broad, with four or five small cornuti (Fig. 94) bicolorella (p. 84)Aedeagus stout, about 10 times as long as broad, with very many spicular cornuti 4 4 Processes of tegumen triangular; apex of saccus pointed (Fig. 93) mackiei (p. 83) Processes of tegumen elongately triangular with extended apex; apex of saccus rounded 5 5 Uncus lobes simple, flap-like; outer surface of valva with shallow ridge; aedeagus with spicular carinae on ventral surface (Fig. 89) luctuosa (p. 82) Uncus lobes with triangular anterior process; outer surface of valva smooth; aedeagus smooth-surfaced 6 Valvae extended ventrally and caudally to form a pair of hand-shaped processes (Fig. 87) coloradella (p. 79) FUNGUS MOTHS 79 Valvae extended ventrally and caudally in a single, fused, irregularly-shaped process (Fig. 91) radulella (p. 80) 7 Uncus lobe with minute triangular internal (anterior) process; setae of valvae spine-like;medioventral emargination between valvae extending to one-third their length (Fig. 92) beckeri (p. 81) Uncus lobe with large triangular internal (anterior) process; setae of valvae thin and hair-like;medioventral emargination between valvae extending to one-half their length (Fig. 90) hin wndcl In (p. 81) Females (females of bimendella, bicolorella, radulella and species A are unknown) 1 Bursa copulatrix with fine spicular signa 2 Bursa copulatrix without spicular signa (but spicular microtrichia may be present in the antrum) 3 2 Spicular signa arranged in an equatorial band (Fig. 148) beckeri (p. 81) Spicular signa arranged in a pair of opposed T-shaped clusters (Fig. 150) mackiei (p. 83) 3 Ductus bursae elongate and narrow, corpus bursae spherical; ventral margin of ostium trans- verse, with deep and narrow medial emargination (Fig. 151) hypocritella (p. 86) - Ductus bursae very short, corpus bursae elongately pyriform; ventral margin of ostium a pair of lobes 4 4 Emargination between lobes that form ventral margin of ostium broadly U-shaped; ductus bursae with sclerotized lateral patches anterior to inception of ductus seminalis (Fig. 147) coloradella (p. 79) - Emargination between lobes that form ventral margin of ostium narrowly V-shaped; ductus bursae without sclerotization anterior to inception of ductus seminalis (Fig. 149) lupulella (p. 85) Daviscardia coloradella (Dietz) comb. n.(Figs 21, 86-88, 147) Scardia coloradella Dietz, 1905: 25. LECTOTYPE cf , U.S.A. (MCZ), here designated [examined on author 's behalf by D. R. Davis].Fernaldia coloradella (Dietz) Davis, 1983: 5. ADULT (Fig. 21). cf $ , 26-33 mm. Vertex whitish, frons brownish yellow. Labial palpus grey-brown, paleron inner surface, whitish at base and apex of terminal segment. Maxillary palpus light grey-brown, short,3-segmented. Antennal scape, pedicel and flagellum dark brown; cilia 2-3 x (cf) or 0-5 x ($) flagellardiameter. Thorax whitish flecked with brown; tegula whitish, brown anteriorly. Forewing purple-brown,paler spots along costa, with continuous whitish posterior and terminal fascia strongly flecked withpurple-brown and tinged with orange-brown posteriorly ; pair of conspicuous dark spots at apices of R 3 andR 4 ; brown flecks coalescing at tornus and fringe ; posterior margin of dark fascia strongly sinuate. Hindwingvery light brownish grey. Legs greyish buff, hindlegs paler, foreleg and mid-leg grey brown above but paleat articulations; outer mid-tibial spur 0-5 length of inner; outer hind proximal tibial spur 0-6 length of innerspur. GENITALIA cf (Figs 86-88). Saccus shallow, almost square; uncus lobes very short, widely separated, butfused basally, each with setose ventral lobe bearing short triangular process with three or four stiff bristles;uncus lobes setose but strongly sclerotized, each with pair of spinose setae close to apex. Tegumenproduced laterocaudally to form pair of rounded 'shoulders'. Subscaphium not developed. Juxta, ifdeveloped, fused with valvae and not recognizable but possibly forming caudally-directed hand-shapedprocess. Valva sinuate, fused with medial pair of hand-shaped processes, apex with dorsal ridge andchisel-shaped ventral tip. Aedeagus stout, 8 x as long as broad at middle, without carinae; vesica withnumerous strong, spicular cornuti. GENITALIA 9 (Fig. 147). Eighth tergite longer than eighth sternite, with five pairs of stout subapical setaeand two or three very small marginal setae, with scattered pits anterior to large setae; eighth sternite deeplyfolded to accommodate broad ostium, ventral margin of ostium with pair of large lobes, each bearing a pairof stout subapical setae and numerous smaller apical and subapical setae. Antrum broadly funnel-shaped,inception of ductus seminalis at apex of 'funnel' just beyond anterior margin of eighth sternite. Ductusbursae and corpus bursae contiguous, posteriorly with sclerotized region on left and irregularly-marginedand more strongly sclerotized patch on right, wall between these patches with microtrichia; medial regionthick-walled, anterior region ovoid, signa absent. 80 GADEN S. ROBINSON DISTRIBUTION. U.S.A. (Colorado, Arizona, Utah, New Mexico, Maine; Texas- Dietz, 1905); Mexico(Durango - Powell, [1968]; Lawrence & Powell, 1969). BIOLOGY. Reared from fungus on Pseudotsuga, from Poly poms on Pinus and from Ganoderma on Populusin the U.S.A., and from Ganoderma on Quercus in Mexico (Lawrence & Powell, 1969). MATERIAL EXAMINED. 7 ex. Lectotype cf , U.S.A.: Colorado, Durango, 9.xii.l899 (Dietz) (MCZ). U.S.A.: 1 cf (paralectotype) , New Mexico, Beulah, end of viii. (genitalia slide no. 12387; BMNH); lex.,Maine, Lincoln Co., 3 m. SE. Demariscotta, 14.vii.1969 (Powers} (UC); 1 $, Arizona, Cochise Co., upperMiller Canyon, Huachuca Mts, 6-7000', 9.viii.l974 (D. & J. Powell) (BMNH); 1 $, Arizona, Redington(BMNH); 2 $ , Arizona, Huachuca Mts, 27.ix.1903 (Oslar) (genitalia slide no. 12388; BMNH); 1 $ , Utah,Wasatch Co., 7 m. E. of Springville, 6000', 28.vii. 1968 (Doyen) (BMNH). REMARKS. This is the only Daviscardia species known to occur north of Mexico. It is larger than all otherknown species, the pale forewing fascia is more strongly speckled with brown, and the antennal cilia of themale are the longest of the genus, approached only by those of an unnamed species from Mexico (seebelow). The male genitalia are distinctive in that the tegumen does not bear the laterocaudal processes thatare present in all other species except radulella; the valvae are produced ventrally to form a pair ofhand-shaped lobes. These lobes may be homologous with the irregular, quadrate mediocaudal processbetween the valvae of radulella and with the hemicylindrical structure in the same position in bicolorella.By analogy with other genera, it seems likely that the juxta is implicated in these structures but that it isclosely fused with the ventrobasal region of each valva and is not recognizable. However, in several species(for example, mackiei) there is no trace of any specialization at the ventral margin of the valva, and thejuxta is either entirely subsumed into the valval complex or else has been lost. For the purpose of scoringcharacter-states for numerical analysis, the juxta in Daviscardia is interpreted as primitively present andmodified into a complex structure. The female genitalia of coloradella are distinctive in their size, the presence of two sclerotized patches atthe posterior end of the ductus bursae, and the deep U-shaped medial emargination of the ventral lip of theostium. I am unable to formulate a hypothesis of phylogeny for Daviscardia. However, two groups of species arerecognizable - coloradella + radulella (with specialized ventral process(es) at the base(s) of the valvae,possibly derived from the juxta, and with the tegumen lacking laterocaudal processes) and all remainingspecies (with laterocaudal processes from the tegumen). Of the latter group, bicolorella has a ventralprocess between the valvae possibly homologous with that of radulella. The identity of Mexican specimens, left in some doubt by Powell ([1968]), has not been confirmed. Daviscardia radulella sp. n. (Figs 22, 91) ADULT (Fig. 22). cf , 16 mm. Coloration and external structure similar to bimendella (see below) but vertexand frons pale yellow-ochre; antennal flagellum light greyish brown above; cilia 1-0 x flagellar diameter;thorax and tegula brown, thorax flecked with brownish cream. Forewing with narrow posterior creamfascia continuous with terminal fascia but almost broken at tornus; distal margin of anterior brown fasciastrongly concave; costa with about eight cream flecks, two basal pairs of these coalesced and extendedposteriorly to form conspicuous spots; terminal fascia with large brown spots between R 3 and R 5 andbetween MI and M 2 . (All legs badly damaged or missing.) GENITALIA cf (Fig. 91). Saccus rounded, with slightly angled apex; uncus lobes very shallow, simple, widelyseparated, fused basally, ventral lobe little more than a ridge. Tegumen not extended laterocaudally.Subscaphium represented by thickening of diaphragma, with spatulate anterior end. Juxta, if developed,fused with valvae and not recognizable but possibly forming irregularly quadrate ventral process betweenarms of valvae; transtilla not developed. Valva simple, with small and thorn-like apical and subapicalprocesses. Aedeagus stout, about 10 x as long as broad in middle, lacking carinae; vesica with numerousstout spicular cornuti. GENITALIA $. Unknown.DISTRIBUTION. Costa Rica.BIOLOGY. Unknown. FUNGUS MOTHS 81 MATERIAL EXAMINED. 1 ex. Holotype cf , Costa Rica: Palo Verde, 5250', 1920 (genitalia slide no. 12386; BMNH). REMARKS. A small, speckly species, radulella may be recognized by the uniform head vestiture, shortantennal cilia and marked extension of the dark forewing fascia towards the tornus. The male genitalia aredistinctive in that there are no processes from the tegumen (as in color adella - q.v.) and there is anirregularly quadrate ventral process between the bases of the valvae.The holotype was originally identified by Meyrick in his collection as 'Cranaodes iulina Wals.'. Daviscardia bimendella (Zeller) comb. n. (Figs 23, 90)Tinea bimendella Zeller, 1863: 143. LECTOTYPE cf , VENEZUELA (BMNH), here designated [examined]. ADULT (Zeller, 1863: pi. 2, fig. 5; Fig. 23). cf , 22 mm. Vertex white, frons brown. Labial palpus brown,whitish at articulations, apex, and on inner surface of first and second segments. Maxillary palpusbrownish, very short, 3-segmented. Antennal scape and pedicel brown; flagellum ochreous; cilia 1-7 xflagellar diameter. Thorax and tegula whitish, anterior half of tegula brown. Forewing purple-brown withextensive continuous terminal and posterior whitish fascia flecked with brown; some orange-brown scalesclose to posterior margin at one-half; costa with large basal and smaller medial white spot. Hindwingoff-white, diffusely flecked with grey at apex. Legs pale buff; foreleg strongly marked with dark brownabove; mid-legs missing; outer proximal hind tibial spur 0-6 length of inner spur. GENITALIA cf (Fig. 90). Saccus short, rounded; uncus lobes very short, widely separated and apparently notfused basally, each with pendulous ventrally-directed lobe bearing subapical triangular process. Tegumenproduced caudally to form pair of lateral digitate processes. Subscaphium defined but only recognizable atspatulate anterior end. Juxta either not developed or entirely fused with valvae and unrecognizable. Valvasimple, with shallow spine on terminal margin and with spined apex. Aedeagus stout, 10 x as long as broadin middle, lacking carinae; vesica with numerous strong, spicular cornuti. GENITALIA 9- Unknown.DISTRIBUTION. Venezuela.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Lectotype cf , Venezuela (genitalia slide no. 13124; BMNH). REMARKS. D. bimendella, with beckeri, luctuosa, mackiei, species A and radulella, forms a group ofexternally very similar species. With the much larger coloradella, beckeri, and the unnamed species fromMexico, it is conspicuous in having the male antennal cilia longer than 1-5 x the diameter of the flagellarsegments; however, the male antenna of luctuosa is unknown. As in coloradella, beckeri, luctuosa andmackiei, the scales of the frons and vertex are of contrasting shades, the vertex whitish and the fronsbrownish. It may be separated from beckeri, luctuosa and mackiei by the broader pale terminal fascia in theforewing and by the large pale basicostal spot. This spot, although present in mackiei, is suffused withpurple-brown in that species. The male genitalia are very similar to those of luctuosa but each valva has a pair of terminal thorn-likeprojections and the uncus lobes each bear a triangular medioventral process; these are absent in luctuosa.The aedeagus is smooth-surfaced but in luctuosa it bears numerous minute spicular carinae. The malegenitalia resemble even more those of beckeri but differ in that the armature is larger, the tegumenprocesses are broader and not as strongly pointed, and the free arms of the valvae are more elongate. Thetriangular medioventral process on each uncus lobe is hardly developed in beckeri. Meyrick misidentified a male of mackiei and the holotype female of hypocritella as 'Cranaodesbimendella' in his collection. Daviscardia beckeri sp. n. (Figs 24, 25, 92, 148)[Phycis luctuosa Walsingham, 1914: 358. Partim-2 9 only. Misidentification.] ADULT (Figs 24, 25). cf 9, 14-21 mm. Coloration and external structure similar to bimendella but fronslight brown, inner surface of third segment of labial palpus greyish (white in one female from Mexico), 82 GADEN S. ROBINSON antennal flagellum grey-brown, cilia 2-0 x (cf) or 0-4 x (9) flagellar diameter; forewing costa with a fewill-defined paler flecks; hindwing light grey in females; outer mid-tibial spur 0-35 length of inner; outerproximal hind tibial spur 0-6 length of inner spur. GENITALIA cf (Fig. 92). Similar to those of bimendella but genital armature smaller, only about 1-05 mmlong (about 1-30 mm in bimendella); uncus lobes not as elongate, subapical triangular process on each lobevery small and insignificant. Subscaphium well defined only at slightly spatulate anterior end, ribbon-like.Ventral emargination between valvae infilled to two-thirds by membranous 'bridge' (only to one-half inbimendella); setae of valvae stout and spine-like. GENITALIA $ (Fig. 148) . Eighth tergite as long as eighth sternite , with three or four pairs of strong subapicalsetae and numerous terminal and subterminal pits; eighth sternite with lamellate surface and only slightlysclerotized dorsal to ostium , ventral lip of ostium m-shaped , lobes each bearing one pair of large setae andnumerous small setae. Antrum with nodular/microtrichiate internal surface, terminating in stronglysclerotized colliculum well beyond margin of eighth sternite. Ductus bursae short, thin- walled, inception ofductus seminalis at junction with corpus bursae. Corpus bursae pyriform (but exaggeratedly so), posteriorend sclerotized, sclerotization terminating anteriorly in equatorial band of spicular signa; corpus swollenanterior to signa, membrane thick-walled and finely reticulate, becoming thin-walled and smooth-surfacedin globular anterior region. DISTRIBUTION. Mexico; Costa Rica.BIOLOGY. Unknown. MATERIAL EXAMINED. 6 ex. Holotype cf , Costa Rica: Turrialba, 600 m, iii.1973 (Becker) (genitalia slide no. 9492; BMNH). Paratypes. 1 cf, 1 $, data as holotype, x.1971 and iii.1973 (coll. V. O. Becker, Brasilia); 1 $, data asholotype, x.1971 (genitalia slide no. 9493; BMNH). Excluded from paratype series. Mexico: 2 $ , Vera Cruz, Jalapa, 4500' , 1887 (Schaus; Trujillo) (genitaliaslide no. 13126; BMNH). REMARKS. This species, similar to bimendella, luctuosa and mackiei, is difficult to recognize by externalcharacteristics. It may be distinguished from mackiei by its considerably longer antennal cilia in the male,and from bimendella by the darker costal region of the forewing and the greater extension in the tornalregion of the dark forewing fascia. The outer mid-tibial spur is shorter in beckeri than in luctuosa or mackiei(0-35 as opposed to 0-50 the length of the inner spur). The male genitalia are distinguished from those ofbimendella as described above. The female genitalia are very similar to those of mackiei but the spicular signa are in an equatorial band,not in a pair of opposed T-shaped clusters, and mackiei has no posterior sclerotized patch on the corpusbursae. It is questionable whether the two females from Mexico are conspecific with the type-series. Thegenitalia of one specimen have been dissected (the other has the abdomen missing) and are illustrated (Fig.148). In comparison with the dissected female from Costa Rica, there are only five strong setae on theeighth tergite (i.e. two pairs plus one supernumerary) whereas there are three pairs plus one in the CostaRican specimen. The antrum is only very weakly sclerotized, the posterior region of the corpus bursae hasonly an irregular sclerotized patch, the shape of the corpus bursae is substantially different, and there is noreticulation of the membrane anterior to the signa. However, the Mexican specimen has mated whereasthe Costa Rican example is virgin; this may account for the differences in the structure and thickness of themembrane of the corpus bursae and for the difference in its shape. Daviscardia luctuosa (Walsingham) comb. n.(Figs 26, 89) Phycis luctuosa Walsingham, 1914: 358. LECTOTYPE cf, COSTA RICA (BMNH), here designated[examined]. ADULT (Fig. 26). cf, 20 mm. Coloration and external structure similar to bimendella but frons light brown,inner surface of labial palpus pale greyish; (antennae broken). Forewing pattern similar to bimendella, buttornal region of pale fascia narrower as distal margin of dark fascia is expanded towards tornus; pale costalspots (particularly basal spot) smaller than in bimendella and suffused with purple-brown; outer mid-tibialspur 0-5 length of inner; outer proximal hind tibial spur 0-6 length of inner spur. FUNGUS MOTHS 83 GENITALIA cf (Fig. 89). Similar to those of bimendella but uncus lobes more widely separated, ventral lobeshorter and without triangular process ; lateral processes of tegumen broader basally ; valva with free caudalregion more elongate, outer surface swollen and slightly ridged, spine on terminal margin represented onlyby slight irregular swelling; aedeagus similar to that of bimendella but ventral surface with numerousminute spicular carinae. GENITALIA $. Unknown. DISTRIBUTION. Costa Rica.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Lectotype cf , Costa Rica: Volcan de Irazu, 6-7000', 18 (Rogers) (genitalia slide no. 13129; BMNH). REMARKS. This species is similar to bimendella (see 'Remarks', above) but the terminal pale forewing fasciais narrower and the pale costal spots much smaller. The male genitalia differ as described for bimendella.The external similarity of luctuosa and its allies is such that Walsingham's syntype series of this speciesincluded the male and (provisional) female ofmackiei as well as the (holotype) female of lupulella and twofemales of beckeri, and the holotype oiMoscardia varna. Daviscardia mackieisp. n. (Figs 27-29, 93, 150)[Phycis luctuosa Walsingham, 1914: 358. Partim - 1 cf , 1 $ only. Misidentification.] ADULT (Figs 27-29). cf , 18, 19 mm; $ , 16 mm. Coloration and external structure similar to bimendella andluctuosa but antennal cilia 1-3 x (cf) flagellar diameter (antennae of $ broken). Forewing pattern similarto luctuosa but pale costal spots larger, resembling more those of bimendella, but suffused with purple-brown as in luctuosa; female with purple-brown fascia extended towards tornus, reaching posterior marginand extending into fringe. GENITALIA cf (Fig. 93). Saccus shallowly triangular; uncus lobes very short, widely separated, but fusedbasally to form a kidney-shaped complex, each with pendulous setose ventral lobe bearing shallowsubapical triangular process; uncus lobes setose but strongly sclerotized. Tegumen produced caudally toform pair of lateral triangular blunt-tipped processes. Subscaphium not developed. Juxta, if present, fusedwith valvae and not recognizable; transtilla possibly represented by thickened band across diaphragmabetween bases of valvae. Valva simple, lobate, apex with strong internal ridge terminating in shallowpyramidal process. Aedeagus stout, 8 x as long as broad at middle, without carinae; vesica with numerousstrong spicular cornuti. GENITALIA $ (Fig. 150). Eighth tergite longer than eighth sternite, with three pairs of strong subapicalsetae and a few scattered pits in posterior half; eighth sternite with lamellate surface and only slightlysclerotized dorsal to ostium, ventral lip of ostium m-shaped, lobes each bearing one pair of large setae andnumerous small setae. Antrum with nodular/microtrichiate internal surface, terminating in colliculum wellbeyond margin of eighth sternite. Ductus bursae short, thin-walled, inception of ductus seminalis atjunction with corpus bursae. Corpus bursae elongately pyriform, sparsely microtrichiate just anterior toinception of ductus seminalis, with strongly sclerotized subequatorial band bearing numerous dense andstrong spicular signa arranged in two opposed inverted T-shaped clusters; membrane adjacent to theseclusters with regular honeycomb-like reticulation. DISTRIBUTION. (?)Guatemala; Colombia; Bolivia.BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Holotype cf , Bolivia: Yungas de la Paz, 1908 (Seebold) (genitalia slide no. 12393; BMNH) (paralecto-typeof luctuosa). Paratype. Colombia: 1 cf , La Crumbre, 6600', v.1914 (P.) (genitalia slide no. 12391; BMNH). Excluded from paratype series. Guatemala: 1 $, Alta Vera Paz, Sinanja, x.1879 (Champion) (genitaliaslide no. 13128; BMNH) (paralectotype of luctuosa). REMARKS. Externally very similar to bimendella, beckeri and luctuosa, mackiei has shorter antennal cilia inthe male than bimendella or beckeri although it has a large basicostal spot on the forewing similar to that in 84 GADEN S. ROBINSON bimendella. This spot is, however, suffused with brown whereas it is pure white in bimendella. The malegenitalia are distinctive in having a pointed saccus, stout valvae with pyramidal apical processes, and thetegumen with triangular laterocaudal processes with rounded apices. The female, provisionally placedhere , is distinguished by the elegant arrangement of spicular signa in a pair of opposed T-shaped clusters atthe mid-length of the corpus bursae.Meyrick originally identified the paratype of this species in his collection as 'Cranaodes bimendella Z.'. Daviscardia bicolorella sp. n. (Figs 30, 94) ADULT (Fig. 30). d", 17, 18 mm. Coloration and external structure similar to bimendella but frons whitishwith a few brown lateral scales; entire inner surface of labial palpus whitish; antennal flagellum pale buff;cilia only 1-0 x flagellar diameter. Forewing purple-brown with very broad posterior and terminal creamfascia, tinged with orange-brown on dorsum and with some scattered brown scales in terminal fascia.Hindwing slightly greyish cream, some darker grey flecks at apex; mid-leg light grey-brown above and onouter surface of tibia, pale at articulations, tibia with oblique pale band on outer surface at one-half; outermid-tibial spur 0-4 length of inner; outer proximal hind tibial spur 0-6 length of inner spur. GENITALIA cf (Fig. 94). Saccus triangular but rounded apically, with vinculum and tegumen forming adistinctly elongate genital armature; uncus lobes small, widely separated, strongly sclerotized, formingventrally-directed lobe with strongly setose inner surface. Tegumen extended laterocaudally to form pairof sharp, spine-like processes. Subscaphium present, narrow, only slightly sclerotized. Juxta, if developed,fused with valvae and not recognizable, but possibly forming hemicylindrical process between valvae;transtilla hardly sclerotized but represented by thickened diaphragma forming a rigid cowl-shapedstructure dorsal to valvae. Valva simple, apex ridged and sinuate. Aedeagus slender, about 20 x as long asbroad at middle, without carinae; vesica with four or five short sagittate cornuti. GENITALIA $. Unknown.DISTRIBUTION. Bolivia.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype cf , Bolivia: Cochabamba (Yunga del Espiritu Santo), 1888-9 (Germain) (genitalia slide no.13121; BMNH). Paratype. 1 $ , data as holotype (BMNH). REMARKS. This species, while superficially similar to bimendella, beckeri, luctuosa and mackiei, isdistinguished by its short antennal cilia, glossy and markedly contrasting forewing fasciae with very littlebrown speckling in the pale fascia, lack of pale costal spots, and its uniformly pale-coloured frons andvertex. The male genitalia are distinguished by the elongate genital armature, sharp-tipped and horn-likelaterocaudal tegumen processes (somewhat similar to those of beckeri), and by the hemicylindrical ventralstructure (?juxta) between the bases of the valvae. This is the only species of Daviscardia that has a longand narrow aedeagus with only a few cornuti. All other species have an aedeagus about 10 times as long asbroad and with numerous spicular cornuti. D. bicolorella has an aedeagus about 20 times as long as broadwith only four or five small cornuti. Daviscardia species A(Fig. 95) ADULT, d", 18 mm. Vertex and frons light ochre. Labial palpus dark brown, apex ochreous. Antennal cilia2-0 x flagellar diameter. Thorax and tegula dark brown anteriorly, whitish posteriorly. Forewing patternsimilar to that of bimendella but without pale costal spots; distal half and terminal area of continuous palefascia strongly flecked with purple-brown, strong spot at apex of R 4 . Hind wing light grey. Outer mid-tibialspur 0-4 length of inner; outer proximal hind tibial spur 0-75 length of inner spur. GENITALIA cf (Fig. 95). Saccus shallow, rounded; uncus lobes widely separated, but fused basally, eachforming shallow caudal and slightly longer ventral setose lobe. Tegumen extended laterocaudally to formtwo pairs of digitate processes. Subscaphium ribbon-like, ill-defined. Juxta, if developed, fused with valvaeand not recognizable; transtilla possibly developed as a broadly triangular region of thickened membrane. FUNGUS MOTHS 85 Valva ridged on internal surface, spatulate, with truncated apex. Aedeagus stout, about 10 x as long asbroad at middle, with subapical group of numerous spicular carinae; vesica with numerous strong spicularcornuti. GENITALIA $. Unknown.DISTRIBUTION. Mexico.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Mexico: 1 cf , Oaxaca, km 140 on highway 175, 4000', 22. v. 1969 (Howden) (UC). REMARKS. This species may be distinguished from all other Daviscardia by its having two processes fromeach corner of the tegumen rather than one. Superficially, it resembles bimendella and its allies but, likebicolorella, radulella and lupulella (and, possibly, hypocritella), the head vestiture is coloured uniformly.D. bicolorella has whitish head vestiture but the present species, radulella, lupulella and hypocritella havethe frons and vertex a light shade of ochre. This species may be separated from radulella (the only one ofthese species of which the male is known) by its longer antennal cilia (twice the flagellar diameter in thisspecies but only as long as the flagellar diameter in radulella). Daviscardia lupulella sp. n. (Figs 31, 149)[Phycis luctuosa Walsingham, 1914: 358. Partim- 1 $ only. Misidentification.] ADULT (Fig. 31). $, 23 mm. Vertex and frons light orange-brown (but very worn). Labial palpus brown,paler on inner surface. Maxillary palpus brownish grey, 3-segmented, but longer than in preceding species,almost reaching apex of second segment of labial palpus. Antennal scape, pedicel and flagellum darkbrown but scape and distal region of flagellum ochreous above; cilia 0-7 x flagellar diameter. Thorax andtegula dark brown in anterior half, light orange-brown posteriorly. Forewing purple-brown with ill-definedand slightly paler costal spots at one-third and two-thirds; with brownish cream terminal and posteriorfascia strongly tinted posteriorly and basally with orange-brown; strong dark spot at apex of R 4 ; furthersmaller spots on apices of M 1? M 2 and M 3 ; pale fascia sparsely flecked with purple-brown scales; posteriormargin of purple-brown fascia with shallowly V-shaped medial emargination. Hind wing grey. Foreleg andmid-leg grey-brown, darker above, pale at articulations (hind legs missing); outer mid-tibial spur 0-5 lengthof inner spur. GENITALIA cf Unknown. GENITALIA $ (Fig. 149). Eighth tergite longer than eighth sternite, with three or four pairs of strongsubapical setae and numerous terminal and subterminal pits; eighth sternite smooth dorsal to ostium,ventral lip of ostium m-shaped, lobes each bearing one pair of large setae and numerous small setae.Antrum with nodular/microtrichiate internal surface, strongly sclerotized but not forming a colliculumanteriorly, terminating well beyond anterior margin of eighth sternite. Inception of ductus seminalis atapex of antrum. Corpus bursae elongately pyriform, posterior region thin-walled, appearing finelyreticulate under phase-contrast; strongly wrinkled in a coarse reticular pattern and thick-walled medially;anteriorly very thin-walled; signa absent. DISTRIBUTION. Panama.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype $, Panama: Volcan de Chiriqui, 2000-3000', 1881 (Champion) (genitalia slide no. 12392;BMNH) (paralectotype of luctuosa). REMARKS. This and the following species, hypocritella, are distinctive in that they are large and plain-patterned (and probably glossy-looking when fresh). The coloration of the head vestiture is uniform. Themaxillary palpus is more elongate in this than in other Daviscardia species and the pale forewing fascia ismore strongly tinted with orange-brown. The genitalia are distinctive in that there is an elongate, stronglysclerotized antrum and, as in hypocritella, no sclerotization of the ductus bursae. The m-shaped ventral lipof the ostium is similar to that seen in luctuosa and mackie; but quite unlike that of hypocritella which isalmost transverse and with a deep and narrow medial emargination. 86 GADEN S. ROBINSON Daviscardia hypocritella sp. n. (Figs 32, 151) ADULT (Fig. 32). $ , 24 mm. Coloration and external structure similar to preceding species, but colorationof head , thorax and legs uncertain , owing to covering of fungal hyphae . Forewing with whitish fascia paler ;distal margin of purple-brown fascia slightly concave, turned at right-angles close to tornus, posteriormargin only slightly concave medially, otherwise nearly straight; posterior fascia only slightly tinted withorange-brown on dorsum; pale fascia sparsely flecked with purple-brown scales; purple-brown spot at apexof R 4 only. GENITALIA cf . Unknown. , GENITALIA $ (Fig. 151). Eighth tergite longer than eighth sternite, with four pairs of strong subapical setae,a few terminal pits and a pair of small setae; eighth sternite with terminal ostium, almost square-ended,ventral lip of ostium with deep and narrow medial emargination; posterior margin with two pairs ofelongate strong setae and numerous smaller setae. Antrum short, hardly sclerotized, internal surfacenodular/microtrichiate, apex forming a strongly sclerotized colliculum, its apex level with anterior marginof eighth sternite. Ductus bursae elongate, lined with sparse microtrichia, finely reticulate close to corpusbursae if observed under phase-contrast (x250); inception of ductus seminalis at one-fifth posteriorly.Corpus bursae spherical, posterior half thick- walled with strong transverse wrinkles; signa absent. DISTRIBUTION. Panama.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype $, Panama: Chiriqui, 1899 (/?.) (genitalia slide no. 13125; BMNH). REMARKS. The pale forewing fascia of this species is not as strongly spotted with dark brown nor as stronglysuffused posteriorly with orange-brown as in lupulella. The genitalia are distinctive and markedly differentfrom those of the four other species of Daviscardia of which the female is known. The elongate ductusbursae and spherical corpus bursae are peculiar to hypocritella, and it is the only Daviscardia species inwhich the ventral lip of the ostium is not even approximately m-shaped. The holotype of this species was originally identified by Meyrick in his collection as 'Cranaodesbimendella Z.' along with the present paratype of mackiei. SCARDIA Treitschke Scardia Treitschke, 1830: 291. Type-species: Phycis boleti Fabricius, 1798: 463, by subsequent designation byBusck,1914:65.Agarica Sodoffsky, 1837(6): 20 (93). Type-species: Phycis boleti Fabricius, 1798: 463. [Unnecessary objective replacement name for Scardia Treitschke.] Fernaldia Grote, 1881: 274. Type-species: Fernaldia anatomella Grote, 1881: 274, by monotypy. Syn. n.Duomitella Koshantschikov, 1923: 22. Type-species: Duomitella relicta Koshantschikov, 1923: 23, by monotypy. [Synonymized by Zagulajev, 1973: 83.] DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia shorter than 1-5 xflagellar diameter. Scape with more than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillarypalpus 5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outermid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3 and R 4 separate; M 2 , M 3and CuAi separate; pattern consisting of pale termen and dorsum on darker ground-colour. Male withcoremata in eighth abdominal segment; coremata not associated with apodemes. Male genitalia withcomplex uncus fused with tegumen; tegumen unbroken, completely sclerotized dorsally; valva lackingbasal setose lobe on inner surface; apex of valva forming ventral hook or hooks, or with spines; valvae +juxta fused ventrally into a single movable complex, valvae without longitudinal cleft; saccus wider thanlong; juxta complex, conspicuous, entire, not divided medially; vesica with spicular cornuti; aedeagussmooth-surfaced, without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Uncus fused with bases of valvae, with pair of elongate horn-like internalprocesses; female with frenulum of about 15 strong bristles. DISTRIBUTION. Western and eastern Palaearctic region; Oriental region - NE. India, Borneo; Nearcticregion; Neotropical region - Venezuela. FUNGUS MOTHS 87 BIOLOGY. See under entries for individual species. Key to species of Scardia Males (males of alleni are unknown) 1 Uncus lobes elongate , tapered, conspicuous 2 - Uncus lobes very short, inconspicuous 4 2 Tegumen extended caudally, dorsal to uncus lobes, forming a cowl-shaped process (Fig. 98). (eastern Palaearctic region) amurensis (p. 89) Tegumen with U-shaped medial emargination, membrane within emargination bearing strong setae . (western Palaearctic region) 3 3 Costa of valva with shallow flap at three-quarters; apex of valva with U-shaped emargination (Figs 99, 100). (Europe east to Siberia, south to Yugoslavia) boletella (p. 90) - Costa of valva lacking shallow flap; apex of valva only slightly concave. (Caucasus) caucasica (p. 92) 4 Uncus lobes each with pair of thorn-like processes; two groups of strong dorsal setae associated with uncus lobes - setae elongate, extending well beyond processes of uncus lobes (Fig. 96).(Nearctic and (?)Neotropical regions) anatomella (p. 87) - Uncus lobes reduced to a pair of small pads, lacking processes; single medial group of setae associated with uncus lobes - setae short, not extending beyond finger-like lateral projections of tegumen (Fig. 97). (NE. India) assamensis (p. 88) Females (females of assamensis are unknown) 1 Ventral lip of ostium with deep medial emargination at least as deep as wide 2 - Ventral lip of ostium hardly or only shallowly emarginate medially, emargination wider than deep 4 2 Ventral lip of ostium with deep, narrow medial emargination many times deeper than wide (Fig. 154). (eastern Palaearctic region) amurensis (p. 89) - Ventral lip of ostium with V-shaped emargination no more than twice as deep as wide 3 3 Ventral lip of ostium with emargination as deep as wide (Fig. 155). (Europe east to Siberia, south to Yugoslavia) boletella (p. 90) - Ventral lip of ostium with emargination deeper than wide (Fig. 156). (Caucasus). . . . caucasica (p. 92) 4 Ventral lip of ostium shallowly concave; pair of shallow, lateral, keel-like processes on eighth sternite bearing elongate, apical setae (Fig. 153). (Nearctic and (?)Neotropical regions) anatomella (p. 87) - Ventral lip of ostium with broad, U-shaped emargination; eighth sternite without keel-like processes, elongate setae scattered near ventral margin of ostium (Fig. 152). (Borneo) alleni (p. 90) Scardia anatomella (Grote) comb. rev.(Figs 33, 34, 96, 153) Fernddia anatomella Grote, 1881: 274; Davis, 1983: 5. LECTOTYPE $, U.S.A. (BMNH), here designated [examined].Scardia fiskeella Busck, 1908: 93. Holotype <j>, U.S.A. (NMNH) [examined]. [Synonymized by Davis, 1983: 5.]Scardia anatomella (Grote) Walsingham, 1882: 171; Dyar, [1903]: 568; Dietz, 1905: 24; McDunnough, 1939: 104. ADULT (Figs 33, 34). cf $, 20-34 mm. Coloration and external structure similar to boletella but antennalcilia onlyO-4 x ($)flagellar diameter and distal segments of antenna light brown. Forewing with little or noorange-brown scaling on veins; in specimen from Venezuela (Fig. 34) cream markings not as stronglyspeckled with brown as in boletella; cream markings at posterior margin reduced in several examples. GENITALIA cf (Fig. 96). Saccus broad, shallow; uncus lobes very short and reduced, highly modified, withelongate and spine-like dorsal processes and shallow, dentate dorsal ridge, setae few, restricted tomediodorsal region. Caudal margin of juxta-tegumen complex strongly emarginate, with four pairs ofstrong spines arising from fused base at dorsocaudal margin of each uncus lobe. Subscaphium not 88 GADEN S. ROBINSON developed. Juxta U-shaped, with further inverted U-shaped sclerotization within the 'U', fused laterallywith valvae, extended distally into pair of claw-shaped processes; transtilla not developed. Valva withventral margin strongly angled outward, apical margin strongly emarginate and with shallow subapical flaprunning obliquely across ventrocaudal region, with weak medial longitudinal ridge and with shallowtriangular process close to costa at one-half. Aedeagus tapered, simple, 4 x as long as broad at base, apexobliquely truncated; vesica with a few scattered minute spicular cornuti (microtrichia) close to inception ofductus ejaculatorius (i.e., at apex of vesica when everted). GENITALIA $ (Fig. 153). Eighth tergite slightly longer than eighth sternite, inverted shield-shaped anteriorregion strongly sclerotized, posterior quarter almost membranous but with five pairs of strong submarginalsetae and numerous smaller setae, these scattered almost to mid-length of tergite; eighth sternite folded toform strongly sclerotized posterior lobe overlying deep pocket (a 'false antrum'), then folded againventrally and slightly anteriorly to form pair of shallowly triangular lobes only slightly sclerotized mesallyand forming lateral margins of ostium; lobes each with three strong setae and four or five smaller setae.Anterior margin of ostium membranous and ill-defined; sternite deepened dorso-ventrally at anterior endto accommodate antrum. Antrum ill-defined, slightly tapered anteriorly, extending slightly beyondanterior margin of eighth sternite; inception of ductus seminalis close to anterior end of antrum. Ductusbursae slightly less than one-half length of apophyses anteriores, thin-walled. Corpus bursae elongatelyovoid, thin-walled except for posterior third which is apparently thicker and more heavily-staining,reaching 1-3 length of apophyses anteriores; signa absent. DISTRIBUTION. U.S.A. (New York - Grote, 1881; North Carolina - Busck, 1908; Louisiana; Arkansas;Oregon- Walsingham, 1882; California -Powell, [1968]; Florida -Kimball, 1965; Pennsylvania -Forbes,1923; Illinois, Texas, Utah - Dietz, 1905); Canada (Ontario - Forbes, 1923); Venezuela. BIOLOGY. Described by Walsingham (1882) who bred several specimens from larvae collected in Oregon inMarch. The larvae were boring round holes in a fallen dead pine tree. MATERIAL EXAMINED. 14 ex., 5 pupae. Lectotype $ (of anatomella) (abdomen missing), U.S.A.: New York (Grote) (BMNH). Holotype 9 (offtskeella), U.S.A.: North Carolina, Tryon, 8.vii.l904 (Fiske) (genitalia slide no. 18664; NMNH). U.S.A.: 9 ex., Oregon, Grant Co., Camp Watson, in dead wood (pine), coll. iii, em. vi.1872(Walsingham) (genitalia slide nos. 19195, 12389, 12390; BMNH); 1 cf, Louisiana, 1884 (Morrison)(BMNH); 1 $, Arkansas, Hope, vi.1926 (BMNH). Venezuela: 1 cf (ex Felder coll.) (BMNH). REMARKS. The only New World species of Scardia, anatomella is characterized by the genitalia of bothsexes: in the male the uncus lobes are short and modified into spine-like processes; the female genitaliadiffer from other Scardia in that the triangular membranous lobe dorsal to the ostium is lacking, as inamurensis, and the eighth sternite has a conspicuous and strongly sclerotized posterior margin that is dorsalto the ostium. This strongly developed posterior region also occurs in amurensis but in that species it isdeeply cleft and overlaid (in ventral view) by a pair of lateral processes directed caudally and extendingbeyond this posterior and dorsal region of the sternite. The specimen from Venezuela is from the Felder collection. It may be labelled incorrectly. It is lighterand with a more strongly marbled appearance to the forewings than specimens from the U.S.A. : this is dueto the lack of brown speckling within the cream forewing markings, and the comparatively larger palecostal spots. I am unable to resolve convincingly the phylogeny of the six Scardia species. 5. boletella and caucasicaare clearly sister-species, their sister-group being alleni (synapomorphy: membranous triangular lobeoverlying ostium dorsally). The sister-group of boletella + caucasica + alleni is probably amurensis(synapomorphy: elongate and tapered uncus lobes). The precise relationship of this grouping to ana-tomella and assamensis, which retain the short, squat uncus complex also observed in Daviscardia, theprobable sister-group of Scardia, is uncertain. However, the uncus lobes in anatomella are almostcompletely separated from the tegumen by membrane whereas in all other Scardia species they are almostcompletely fused; the degree of separation observed in anatomella is similar to that observed inDaviscardia and is considered to be the ground-plan state of the character. On the basis of this decidedlyweak feature, anatomella is the sister-group of all other Scardia. Scardia assamensis sp. n. (Figs 35, 97, 101)ADULT (Fig. 35). cf , 24 mm. Coloration and external structure similar to boletella but frons and vertex dark FUNGUS MOTHS 89 reddish brown, thorax and tegula deep purple-brown. Forewing (worn) lacking extensive pale posteriorfascia, with only scattered whitish scales close to posterior margin; pale markings of terminal fasciasilver-grey, not as extensive as in boletella. Hindwing fringe not chequered. GENITALIA cf (Figs 97, 101). Saccus broad, shallow; uncus lobes short, apices short and digitate and widelyseparated, strongly sclerotized and with setae restricted to laterodorsal region. Fused uncus-tegumencomplex strongly emarginate medially, membrane of emargination bearing conspicuous group of about 40strong spines. Subscaphium not developed. Juxta trapezoidal , fused laterally with valvae , extended distallyto form pair of hand-shaped processes; transtilla not developed. Valva simple, with small shallowhook-like process at apex. Aedeagus stout, about 5 x as long as broad at base, strongly sclerotized, apexlaterally emarginate; vesica with scattered minute spicular cornuti (microtrichia) for half its length closestto inception of ductus ejaculatorius (i.e., in apical half of vesica if it were everted). GENITALIA $. Unknown.DISTRIBUTION. India- Assam.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype cf , India: Assam, Khasi Hills, Cherrapunji, v.1895 (native collector] (genitalia slide no. 1514;BMNH). REMARKS. This is the only Old World Scardia species in which the lobes of the uncus are short. S.assamensis is small; it lacks the extensive pale fascia at the posterior margin of the forewing thatcharacterizes other Scardia with the exception ofalleni. It is the only species in which the head, thorax andtegulae are entirely dark brown and the only one in which the horn-like lateral processes from the juxta aremodified and hand-shaped, bearing small digitate subterminal processes. Scardia amurensis Zagulajev(Figs 36, 98, 154) Scardia amurensis Zagulajev, 1965: 411; 1973: 96; Moriuti, 1982: 163. Holotype cf, U.S.S.R. (ZI) [examined] [(?) Scardia boletella (F.); Caradja, 1939: 111. Misidentification.][Scardia boletella (F.); Issiki, 1957: 16. Misidentification.] ADULT (Issiki, 1957: pi. 2, fig. 46 (colour); Moriuti, 1982: pi. 2, fig. 12 ($), pi. 227, fig. 1 (cf) (colour); Fig.36). cf $ , 39-42 mm. Coloration and external structure similar to boletella. GENITALIA cf (Zagulajev, 1965: fig. 1; Zagulajev, 1973: figs 15B, 16b, 75; Moriuti, 1982: pi. 248, figs 10,lOa; Fig. 98). Saccus broad, shallow; uncus lobes flattened laterally, strongly sclerotized, setose, tapered toa point; tegumen extended dorsocaudally to form overlying triangular process, spined membranous dorsalrecess thus absent. Subscaphium not developed but membrane of diaphragma in this region formingshallow lateral pockets with transverse wrinkling. Juxta quadrate, fused laterally with valvae, extendeddistally into pair of claw-like processes, with doubly bulbed shallow ventral process; transtilla notdeveloped. Valva almost square, simple, costa extended caudally in a simple, rounded extension that isbulbous subapically; slight basal ridge on internal surface of valva close to patch of about 10 short, strongsetae. Aedeagus 5 x as long as broad at base, explanate apically, planarian-shaped; vesica with minutespicular cornuti (microtrichia) close to inception of ductus ejaculatorius (i.e., at apex if vesica wereeverted). GENITALIA $ (Zagulajev, 1973: figs 76, 77; Fig. 154). Eighth tergite slightly longer than eighth sternite,inverted shield-shaped anterior region strongly sclerotized, posterior quarter almost membranous but withfive pairs of strong submarginal setae and numerous smaller setae; eighth sternite strongly folded to formpair of lateral setose digitate processes, each overlying a cushion-shaped lobe; ostium a deep fold behindthese cushion-like lobes, ventral margin a narrow bridge between them. Course of antrum complex,oriented dorso-ventrally through base of cushion-like lobes then turned anteriorly and forming acolliculum-like apex just anterior to margin of eighth sternite; inception of ductus seminalis apparentlyclose to inner bases of cushion-like lobes (i.e., remarkably posterior). Ductus bursae very thin-walled,one-half length of apophyses anteriores, with fine and numerous irregular transverse constrictions. Corpusbursae ovoid, very thin-walled, reaching 1-3 length of apophyses anteriores; signa absent. 90 GADEN S. ROBINSON DISTRIBUTION. U.S.S.R. - Amur and Primorsk regions; (?) China (Caradja, 1939); Japan. BIOLOGY. Specimens have been collected on the trunk of a rotting tree and bred from larvae living in fungus(Zagulajev, 1973); the Japanese specimen examined (see below) was bred from Fames fomentarius . MATERIAL EXAMINED. 10 ex. Holotype cf , U.S.S.R.: Primorsk Terr., Spassk-Dalniy, at light, 5.viii.l950 (Zagulajev) (ZI). U.S.S.R.: 1 O" (paratype), Amur Prov., Svobodnyi Dist., Simonovo vill., at light, l.viii.1959 (Falko-vitsh) (ZI); 1 cf (paratype), Primorsk Terr., Suchan, on trunk of rotting tree, 16. viii. 1950 (Zagulajev) (ZI);1 $, Ussuri, Kaimanovka vill., 28-31. vii. 1964 (Tsvetayev) (ZI); 1 cf, 1 $, Suputinskiy Reserve, bred fromlarva in fungus, 14. v. 1967 (Mamaev) (BMNH; ZI); 2 $, Ussuri Railway, Chabarovsk, 14. vii. 1910 &23. vii. 1911 (Borsow) (BMNH); 1 ex., data not recorded (ZI). Japan: 1 cf , Ehime Pref., Mt Sara, bredfrom Fames fomentarius , em. 8. v. 1954 (Hisamatsu) (genitalia slide; coll. S. Moriuti, Osaka). REMARKS. The wing-pattern of this species and the morphology of the male genitalia are very similar tothose of boletella (see 'Remarks' for that species) and caucasica. The female genitalia, however, aremarkedly different, lacking the triangular membranous lobe dorsal to the ostium and having the eighthsternite conspicuously modified as described above. Caradja's record of 'boletella' from China, probably referable to this species, has not been confirmed byexamination of the original specimen. Scardia allenisp. n. (Figs 37, 152) ADULT (Fig. 37). $, 31 mm. Coloration and external structure similar to boletella but frons and vertexbrownish yellow, maxillary palpus dark purple-brown, thorax cream, tegula dark purple-brown. Forewingwith pale markings almost obliterated by dark purple-brown, a few silvery white scales close to posteriormargin at one-half; terminal fascia composed of discrete silvery white spots not speckled with brown as inboletella. Hindwing charcoal grey, a few slightly paler flecks towards apex, fringe not chequered. GENITALIA cf . Unknown. GENITALIA $ (Fig. 152). Eighth tergite slightly longer than eighth sternite, with four pairs of elongate,strong setae close to posterior margin, and with a few scattered minute thorn-like setae; eighth sterniteseparated by membranous sutures into roughly triangular mediocaudal and lateral sclerites; mediocaudalregion rugose, posterior margin strongly concave, with five pairs of strong setae and numerous smallersetae; ostium overlain dorsally by triangular membranous lobe, wall of ovipositor dorsal and slightlyposterior to apex of lobe ballooned ventrad to form pair of shallow lobes. Antrum very short, conical,sclerotized strongly at anterior end and forming a colliculum-like constriction. Ductus bursae thick-walled,with strong and regular transverse constrictions on inner surface (this apparent restriction to the innersurface may be an artefact caused by separation of the outer layer of the membrane of the ductus). Corpusbursae contiguous with ductus, elongately ovoid, reaching 1-5 length of apophyses anteriores; signaabsent. DISTRIBUTION. Borneo -Brunei.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype $, Brunei: Labi, lowland forest, 60 m, 12. i. 1980 (Allen) (genitalia slide no. 1517; BMNH). REMARKS. Of the six Scardia species, alleni is the darkest, the pale terminal and posterior fasciae of theforewing being markedly reduced in comparison with the other species. The hindwing is distinctivelydarker than in the other species. The genitalia resemble those of boletella and caucasica in that there is atriangular membranous lobe dorsal to the ostium, but the ventral margin of the ostium lacks the deep U- orV-shaped invagination of these species and the membranous lobe lacks a terminal sclerite. The locality in which the holotype was collected is on the Rampayoh River in an area of low hills forestedwith typical lowland dipterocarp forest (Allen, pers. comm.). Scardia boletella (Fabricius) nom. rev.(Figs 38, 99, 155) [Phalaena (Tinea) gigantella [Denis & Schiffermuller]; Hiibner, 1790: 38, pi. 2(2), fig. F. Misidentifica-tion.l FUNGUS MOTHS 91 Tinea boletella Fabricius, 1794: 287. LECTOTYPE (?) $, [AUSTRIA or SWEDEN], (ZM), here designated [examined]. Phycis \boleti Fabricius, 1798: 463. Unjustified emendation of boletella.Noctua polypori Esper, [1804]: pi. 196, fig. 1; [1805]: 64. Unnecessary objective replacement name for Phycis boleti F., 1798 (cited as Tinea Boleti') which is an unjustified emendation of Tinea boletella F., 1794.Duomitella relicta Koshantschikov, 1923: 23. Syntypes, 1 cf, 1 $, U.S.S.R. (ZI) [examined]. [Synony- mized by Zagulajev, 1973: 89.] ADULT (Hubner, 1796: pi. 3, fig. 18; Esper, [1804]: pi. 196, fig. 1; Zagulajev, 1973: figs 31, 32, 66, pi. 2, fig.1; Fig. 38). Cf $ , 37-60 mm. Vertex and frons brownish cream, brown close to eyes. Labial palpus cream,brown on outer surface of first and second segment and in middle of third segment. Maxillary palpus creamflecked with brown. Antennal scape, pedicel and flagellum dark brown; cilia 1-2 x (cf) or 0-7 x (9)flagellar diameter. Thorax cream flecked with brown; tegula brown, cream posteriorly. Forewing cream,strongly patterned with dark purple-brown, with orange-brown on veins; cream coloration almostobliterated by brown and restricted to termen and posterior margin, with a few pale flecks towards costa;fringe conspicuously chequered. Hind wing light grey-brown, pale flecks towards apex; fringe chequered.Legs cream, strongly flecked with brown but pale at articulations; outer mid-tibial spur 0-6 length of inner;outer proximal hind tibial spur 0-7 length of inner spur. GENITALIA cf (Petersen, 1957: fig. 238; Zagulajev, 1973: figs 14B, 67; Fig. 99). Saccus broad, shallow;uncus lobes flattened laterally, strongly sclerotized, setose and tapered to a point; tegumen with dorsalemargination bearing five or six pairs of strong spines; dorsal region of fusion of uncus lobes and tegumenbroad, with shallow lateral invaginations. Subscaphium not developed, but membrane in this region withlateral patches of lamellate microtrichia. Juxta broadly V-shaped, fused laterally with valvae, extendeddistally into pair of horn-shaped processes; transtilla not developed. Valva simple but with shallow andrugose costal flap and with mediobasal fold; apical margin with broadly U-shaped emargination. Aedeagus10 x as long as broad, strongly sclerotized but simple; vesica with minute spicular cornuti (microtrichia)close to inception of ductus ejaculatorius (i.e. at apex if vesica were everted). GENITALIA $ (Petersen, 1957: fig. 239; Zagulajev, 1973: figs 68, 69; Fig. 155). Eighth tergite slightly longerthan eighth sternite, with slightly concave caudal margin, with four pairs of strong marginal setae atposterior corners and seven or eight pairs of smaller submarginal setae; eighth sternite oval, stronglykeeled, ventral margin of ostium V-shaped, with small medial emargination; lobes either side of ostiumeach bearing six stout setae and a few smaller setae; ostium surmounted by membranous posteriorly-extended lobe bearing elongate heart-shaped sclerite with shallow lateral invaginations; posterior to lobeand sclerite a deep anteriorly-directed pocket forming a 'false antrum'. Antrum hardly sclerotized, taperedanteriorly, reaching anterior margin of eighth sternite; inception of ductus seminalis just anterior tocolliculum-like apex of antrum. Ductus bursae one-half length of apophyses anteriores, with irregulartransverse constrictions, very thin-walled. Corpus bursae ovoid, very thin-walled, reaching 1-3 length ofapophyses anteriores. DISTRIBUTION. Norway (Aarvik & Midtgaard, 1982); Sweden (Petersen, 1957); Finland (Jalava, 1977);U.S.S.R. - Baltic and European regions, Crimea, Siberia (Zagulajev, 1973); West Germany (Petersen,1968); Czechoslovakia (Petersen, 1965); Hungary (Petersen, 1957); Rumania; Austria (Petersen, 1957);Switzerland (Rebel, 1901); Italy (Petersen & Gaedike, 1979); Yugoslavia. BIOLOGY. See Zagulajev (1973: 93) - bred from Fomes, Ganoderma and Polyporus. Zagulajev (1973: figs20, 22, 26, 27) illustrates the larva. The biology is also described briefly by Mitterberger (1910: 171) andKoshantschikov (1923). The latter author found three live pupae enclosed in webbed frass on fungi on adead birch, and he also found three large larvae (which he preserved) in the fungus. MATERIAL EXAMINED. 54 ex., 4 pupae. Lectotype (?)$> (of boletella) (abdomen missing), [Austria or Sweden]: labelled 'boleti' in Fabricius'hand (Sehested & Tender Lund collection, ZM). Syntypes (of relicta), U.S.S.R.: Siberia, Irkutsk, Minussinsk dist., Sajan Mts, Tiberkul Lake: 1 cf,21.vii., 1 $, 17.vii.1920 (Koshantschikov) (ZI). 37 ex., 4 pupae (ZI) and 15 ex. (BMNH), various localities (see 'Distribution' and Zagulajev (1973)). REMARKS. Esper expressly proposed the name polypori as a replacement name for 'Tinea Boleti' which heconsidered preoccupied by Noctua boleti F. , 1777.This species is one of the largest Tineidae and has a wide but sporadic boreo-alpine/cold temperate 92 GADEN S. ROBINSON distribution in the western Palaearctic region, occurring in montane localities as far south as Italy andYugoslavia. It may be separated from all other Scardia by the genitalia, characteristic in both sexes but verysimilar indeed to those of caucasica; the latter species represents an isolated geographical race of boletellaonly arguably deserving of specific status. With caucasica, boletella may be separated from all other Scardiaby the elongate and pointed lobes of the uncus in the male and by the triangular membranous lobe thatoverlies the ostium of the female and terminates in a laterally invaginated sclerite. The only other Scardiawith pointed uncus lobes is amurensis (although they may also occur in alleni when the male of that speciesis discovered) in which the tegumen forms a shallow triangular hood above the uncus lobes (the tegumen isemarginate in boletella and caucasica) and in which the apex of the aedeagus is conspicuously explanate. S.alleni has a similar membranous lobe overyling the ostium in the female but the lobe is not sclerotized at itsapex. The pronounced allopatry of the Scardia species is helpful in identification although furthercollecting may show the presently disjunct distributions to be artefactual. Scardia caucasica Zagulajev (Figs 39, 100, 156) Scardia caucasica Zagulajev, 1965: 412, figs 2, 3. Holotype C? , U.S.S.R. (ZI) [examined].ADULT (Zagulajev, 1973: fig. 70; Fig. 39). $ , 44 mm. Coloration and external structure similar to boletella. GENITALIA cf (Zagulajev, 1965: figs 3, 19; 1968: fig. 16; 1973: fig. 72; Fig. 100). Similar to boletella butdorsal margins of uncus lobes not extended as far caudally (in lateral view); dorsal region of tegumennarrow, with deep lateral indentations; valva with costal flap hardly developed, apical margin onlyshallowly concave. GENITALIA $ (Zagulajev, 1965: fig. 2; 1968: fig. 17; 1973: fig. 73; Fig. 156). Similar to boletella but eighthtergite with only three pairs of large setae and two or three pairs of small setae; ventral margin of ostiumdeeper, V-shaped; lobes either side of ostium more elongate, each with four strong setae arranged in adiagonal line at one-half, smaller setae concentrated at apex. Lobe dorsal to ostium broader and shorter,with dome-shaped terminal sclerite with small, deep lateral invaginations. DISTRIBUTION. U.S.S.R. - Caucasus. BIOLOGY. Zagulajev (1973) collected a male of this species on a fungus-infested ash trunk. MATERIAL EXAMINED. 10 ex., 4 pupae. Holotype cf , U.S.S.R.: Caucasus, Georgia, Lagodekhi, 8.viii.l961 (Zagulajev) (ZI). U.S.S.R.: Caucasus: 1 cf, Tbilisi dist., Manglis, 13.ix.1882 (Christoph) (ZI); 1 <j>, 4.viii.l891 (Hede-mann) (ZI); 1 $, Lagodekhi, 1888 (Mlokosevitch) (ZI); 2 cf , 3 $, Lagodekhi, wet forest in lower zone ofreserve, 27-30. vii and 8.viii.l961 (Zagulajev) (ZI; 1 $ in BMNH) (all paratypes); 1 cf, Armenia,Idzfrevanskiy dist., 30 km from Sevkar, Kerants Monastery, on fungus-infested ash trunk at dusk,31.vii.1960 (Zagulajev) (ZI). REMARKS. For differentiation of this species from boletella, see above; see also 'Remarks' for boletella. PERILICMETIS Meyrick Perilicmetis Meyrick, 1932ft: 323. Type-species: Perilicmetis diplaca Meyrick, 1932ft: 324, by monotypy. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia shorter than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus with fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width ofhead. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3 and R 4separate; M 3 and CuA\ stalked or very closely approximated at base; light grey speckled with dark brown,with broad, dark brown, oblique subterminal fascia. Male with coremata in eighth abdominal segment;coremata without associated apodemes. Male genitalia with complex uncus fused with tegumen; tegumenunbroken, completely sclerotized dorsally; valva lacking basal setose lobe on inner surface; apex of valvanot forming ventral hook or hooks, without spines; valvae separate, not fused together ventrally, withoutlongitudinal cleft; saccus longer than wide; juxta simple, entire, not divided medially; vesica lackingspicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Forewing with oblique dark brown subterminal fascia constricted inmiddle to form a pair of large, narrowly connected spots; remainder of forewing light grey, finely speckledwith dark brown. FUNGUS MOTHS 93 DISTRIBUTION. Neotropical region - Brazil.BIOLOGY. Unknown. Perilicmetis diplaca Meyrick (Figs 40, 102, 180)Perilicmetis diplaca Meyrick, 1932ft: 324. LECTOTYPE cf , BRAZIL (NM), here designated [examined]. ADULT (Figs 40, 180). cf, 17-22 mm. Vertex and frons chocolate brown. Labial palpus chocolate brown,outer surface of third segment darker. Maxillary palpus buff, dark brown above, 4-segmented, short,reaching only basal one-sixth of second segment of labial palpus. Antennal scape buff, large dark brownspot above; pedicel dark brown; flagellum pale buff; cilia 0-8 x flagellar diameter. Thorax and tegulagreyish white flecked with dark brown anteriorly. Forewing greyish white, strongly flecked with darkbrown; conspicuous oblique subterminal fascia formed from pair of large dark brown spots. Hindwing verypale greyish brown, whitish towards base. Legs pale buff; foreleg and mid-leg strongly marked with darkbrown above and on sides but pale at articulations; hind tarsus banded with brown; outer mid-tibial spur0-45 length of inner; outer proximal hind tibial spur 0-5 length of inner spur. GENITALIA cf (Fig. 102). Saccus triangular; uncus lobes separated, strongly fused with tegumen, taperedand divergent apically, well-sclerotized, setose. Subscaphium not developed. Juxta represented by onlyslight ventral thickening of anellus; transtilla not developed. Valva elongate, broad, simple, apex formingslight dorsal hook. Aedeagus 8 x as long as broad, somewhat flattened dorsoventrally, with dense spinosecarinae on ventral surface; vesica without cornuti. GENITALIA $. Unknown.DISTRIBUTION. Brazil.BIOLOGY. Unknown. MATERIAL EXAMINED. 7 ex. Lectotype cf , Brazil: Santa Catharina, Neu Bremen, 23. ii. 1931 (Hoffmann) (NM). Brazil: 2 cf , (paralectotypes) Santa Catharina, Jaragua, 24.viii.1929, 16.ix.1930 (Hoffmann) (BMNH;NM); 1 cf, Sao Paulo (Jones) (BMNH); 1 cf, Parana, Castro, x.1895 (Jones) (BMNH); 1 cf, SantaCatharina, Neu Bremen, 2.ix.l936 (Hoffmann) (genitalia slide no. 6966; BMNH); 1 cf , Santa Catharina,18.iii.1936 (Hoffmann) (BMNH). REMARKS. The external appearance of this species is very distinctive, the wing-pattern being unlike that ofany other tineid known to me. Although the specimens examined are old and probably faded, the forewingground-colour is distinctly greyish. The only other scardiines that could be thought of as greyish are theAmorophaga species. The genitalia of Perilicmetis resemble superficially those of Amorophaga but thevalva is much more simple and the tegumen is complete dorsally. MOSCARDIA gen. n. Type-species: Myrmecozela renitens Meyrick, 1922ft: 591. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia shorter than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus 5-segmented in renitens but probably with fewer (four or three) segments in varna; pilifers present;second segment of labial palpus shorter than width of head. Outer mid and proximal hind tibial spurs >0-4length of inner spurs. Forewing with /? 3 and R 4 separate; M 2 , M 3 and CuA\ separate; pattern consisting ofpale termen and dorsum on darker ground-colour. Male with coremata in eighth abdominal segment;coremata associated with short, lobe-like apodemes at anterior corners of eighth sternite. Male genitaliawith complex uncus fused with tegumen; tegumen unbroken, completely sclerotized dorsally (but brokendorsally by a membranous suture line in renitens) ; valva lacking basal setose lobe on inner surface; apex ofvalva forming ventral hook or hooks, or with spines; valvae separate, not fused together ventrally, withoutlongitudinal cleft; saccus longer than wide; juxta simple, entire, not divided medially; vesica lackingspicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. None; however, males ofMoscardia species may be recognized externallyby their brown forewing with pale apex and dorsum and completely scaled antenna with short cilia; 94 GADEN S. ROBINSON exclusive microscopic characters are the short apodemes of the eighth sternite in combination with minutemicrotrichia on the second sternite. DISTRIBUTION. Neotropical region -Brazil, Bolivia.BIOLOGY. Unknown. Key to species of Moscardia (only males are known) 1 Tegumen broken by dorsal line of membrane, without processes; uncus lobes strongly sclero- tized and with serrate subapical ridge renitens (p. 94) Tegumen complete dorsally, strongly sclerotized, with pair of small lanceolate caudal processes;uncus lobes almost membranous, apices rounded (Fig. 104) varna (p. 95) Moscardia renitens (Meyrick) comb. n. (Fig. 41)Myrmecozela renitens Meyrick, 19226: 591. Holotype cf , BRAZIL (BMNH) [examined]. ADULT (Clarke, 1970: pi. 32, fig. 3; Fig. 41). d", 22 mm. Vertex and frons pale whitish ochre, frons stronglytinted reddish brown. Labial palpus very dark brown on outer surface but cream at articulation betweensecond and third segments and at tip; inner surface light brown on second segment, cream on third.Maxillary palpus short, not as long as second segment of labial palpus, but probably 5-segmented, darkbrown. Antennal scape and pedicel cream, flecked with light brown; flagellum brownish cream, darkerbrown on basal few segments; cilia 0-8 x (cf ) flagellar diameter. Thorax and tegula medium brown, whitishposteriorly. Forewing badly rubbed; traces of pattern present only at margins, brownish cream flecked withlight brown, strongly spotted with dark brown at costa and with strong dark brown spot on posterior marginat two-thirds. Hindwing light greyish brown with slight purple iridescence. Legs cream flecked with brown;foreleg and mid-leg strongly marked with dark brown above and on sides but pale at articulations, outersurface of mid-tibia with broad oblique whitish band; outer mid-tibial spur 0-5 length of inner spur [hindlegs missing]. GENITALIA cf (Clarke, 1970: pi. 32, figs. 3a, 3b). Saccus elongate, triangular, almost as long as valva; uncuslobes triangular, strongly sclerotized, fused with tegumen but with incomplete membranous sutureforming line of flexion, with serrate subapical ridge. Subscaphium apparently not developed (butpreparation poor). Juxta apparently large and shield-shaped (but preparation damaged in this area anddifficult to interpret); transtilla apparently not developed. Valva simple, with small setose lobe on costa atthree-quarters and with three small peg-like apical processes; membranous subapical suture on outersurface. Aedeagus stout, 5 x as long as broad, asymmetrical, with three strong, subapical horn-shapedcarinae directed caudally, and with three similar but smaller anteriorly-directed carinae closer to apex;vesica without cornuti. GENITALIA $. Unknown.DISTRIBUTION. Brazil.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype cf , Brazil: Teffe, i.1920 (Parish) (genitalia slide no. JFGC 6653; BMNH). REMARKS. The condition of the holotype of this species, and of the slide of its genitalia, is such thatdescription of the forewing pattern and of the juxta/transtilla/subscaphium region is not possible. Despitethis, the genitalia are characterized simply by the structure of the uncus lobes and the valva. M. renitensdiffers from varna in that the maxillary palpus of the latter, while as long as that of renitens, has apparentlyonly three segments. In varna the tegumen is strongly sclerotized and unbroken dorsally and the uncuslobes are almost membranous; the tegumen is produced caudally to form a pair of small, rhomboidalprocesses. In renitens the tegumen is broken dorsally by a zone of membrane, the uncus lobes are stronglysclerotized, and there are no processes from the tegumen. There is no trace of a pattern of pale apex and dorsum on the forewings of the holotype of renitens.Although the specimen is very rubbed, I would have expected to find traces of such a pattern, had onebeen present. It is possible, therefore, that this species and varna have quite different forewing patterns. FUNGUS MOTHS 95 Moscardia varna sp. n. (Figs 42, 104)[Phycis luctuosa Walsingham, 1914: 358, partim- 1 cf only. Misidentification.] ADULT (Fig. 42). cf , 17 mm. Vertex and frons brownish cream. Labial palpus dark brown on outer surface,paler on inner surface and at apex. Maxillary palpus short, not as long as second segment of labial palpus,probably 4- or 3-segmented, dark brown. Antennal scape and pedicel dark brown above; flagellum lightgrey-brown; cilia 1-0 x (cf) flagellar diameter; (thorax obscured by glue). Tegula medium brown.Forewing deep purple-brown, with continuous golden brown posterior and terminal fascia occupying morethan half of total wing area; scales close to junction with deep brown anterior fascia pale gold, increasingthe contrast between the two. Hindwing dark grey. Legs greyish-ochreous (worn and damaged). GENITALIA cf (Fig. 104). Saccus elongate, triangular; uncus lobes small, inturned ventrad, almostmembranous, setose, surmounted by strongly sclerotized tegumen with pair of small rhomboidalmediocaudal processes. Subscaphium not developed. Juxta represented only by slight thickening ofanellus; transtilla not developed. Valva elongate, twice length of saccus, extending well beyond apices oftegumen lobes; simple, with small and shallow triangular lobe bearing 6 small setae at two-thirds of costa;apex of valva rounded, with spinose setae. Aedeagus stout, 6 x as long as broad, with, either side,subapical group of two triangular basally-directed carinae; vesica without cornuti. GENITALIA $. Unknown.DISTRIBUTION. Bolivia.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype cf , Bolivia: Yungas de la Paz, 1908 (Seebold) (genitalia slide no. 13122; BMNH) (paralecto-type of Phycis luctuosa Walsingham). REMARKS. See 'Remarks' for renitens for differentiation from that species. The external appearance ofvarna, with strongly developed, continuous and extensive distal and posterior fasciae on the forewing, ischaracteristic: the posterior pale fascia extends further anteriorly than in any other species dealt with here.The aedeagus is only lightly sclerotized and has suffered slight damage during preparation. The anellus hasbeen torn and has peeled back the wall of the aedeagus from close to the carinae . Interpretation is thereforedifficult but there appear to be a pair of elongate, sinuate processes that originally commenced as ridges atthe base of the aedeagus and reached the apices of the (backward-directed) carinae. These are now foldedbackward from close to the base of the aedeagus and lie within a fold of the anellus. GENTINGIA gen. n. Type-species: Gentingia hollowayi sp. n. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus with fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width ofhead. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3 and R 4 stalkedor very closely approximated at base; M 2 , A/ 3 and CuA^ separate; ground-colour very pale bronze, withpattern of small purple-brown marginal spots and larger costal blotch. Male with coremata in eighthabdominal segment; coremata without associated apodemes. Male genitalia with complex uncus fused withtegumen; tegumen unbroken, completely sclerotized dorsally; valva lacking basal setose lobe on innersurface; apex of valva not forming ventral hook or hooks, without spines; valvae fused ventrally, withoutlongitudinal cleft; saccus longer than wide; juxta, if present, fused with valvae and not recognizable assuch; vesica with spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Male with tegumen produced laterocaudally into pair of hooked processes;female with scobinate signum surrounding inception of ductus bursae in corpus bursae; interior surface ofductus bursae denticulate anteriorly, posteriorly with overlapping scale-like plaques. DISTRIBUTION. Oriental region- Malay Peninsula, Borneo.BIOLOGY. Unknown. 96 GADEN S. ROBINSON Gentingia hollowayisp. n. (Figs 43, 103, 157, 184) ADULT (Figs 43, 184). O"$, 16-19 mm. Vertex and frons pale yellow-brown. Labial palpus ochreous,strongly suffused with brown on outer surface but pale at apex and articulations. Maxillary palpus whitish,flecked with dark brown above, short, apparently 3-segmented. Antennal scape and pedicel ochreous,flecked above with dark brown ; flagellum light greyish ochreous , with darker scales basidorsally ; cilia 1 5 x(O") orO-5 x (5) flagellar diameter. Thorax and tegula cream, strongly flecked anteriorly with dark brown.Forewing pale golden, strongly spotted with dark purple-brown along costa and termen; strong wedge-shaped dark blotch from costa across end of cell to CuP; conspicuous (but frequently abraded) dark spot onmiddle of dorsum. Hindwing off-white. Legs ochreous cream, foreleg and mid-leg dark brown above;outer mid-tibial spur and outer hind proximal tibial spur 0-6 length of inner spurs. GENITALIA cf (Fig. 103). Saccus triangular, strongly tapered anteriorly; uncus lobes short, small, stronglysclerotized, curved ventrad, burin-shaped, setose, entirely fused with tegumen; tegumen with pair oflateral hook-shaped processes. Subscaphium not developed. Juxta, if present, entirely fused with valvaeand not recognizable ; transtilla not developed . Valvae strongly fused ventrally to form single movable unit ;medioventrally with an m-shaped projecting caudal margin, overlying this a medial thimble-shapedprocess; apices of valvae rounded, dorsal to the medial structures and extending caudally beyond them;inner margins of valvae emarginate at level of m-shaped process; ventral surface of valva smooth, apicallyand dorsally with numerous setae, dorsal margin (costa) hardly sclerotized but with line of about six strongsetae. Aedeagus tapered, 12 x as long as broad in middle, with lateral bands of thorn-like carinae fromone-half to apex; vesica with minute blunt lepidote cornuti (?microtrichia). GENITALIA $ (Fig. 157). Eighth tergite considerably longer than eighth sternite, with two subapical pairs ofelongate strong setae, three or four pairs of smaller setae at posterior corners, and with scattered pits incaudal half; eighth sternite triangular, deeply pocketed laterally, apex with V-shaped emarginationinterrupted by keel-like mediocaudal process with bilobed tip, each lobe bearing pair of elongate strongsetae and a few small setae. Antrum broad, funnel-shaped posteriorly then tubular and slightly asymmet-rical, meeting ductus seminalis at two-thirds anteriorly, reaching 0-75 length of apophyses anteriores,anterior end strongly microtrichiate, microtrichia extending into ductus bursae. Ductus bursae verythick-walled, internal surface strongly lepidote, the scale-like surface projections arranged in a regulartransverse pattern giving the appearance of very fine regular constrictions under low-powered magnifica-tion; lepidote projections highly developed anteriorly, resembling sharks' teeth, and continuous withdenticles of signum. Corpus bursae thin-walled, with U-shaped denticular signum surrounding inception ofductus bursae. DISTRIBUTION. Malay Peninsula; Borneo (Fig. 200).BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Holotype cf , West Malaysia: W. Pahang, Genting Highlands, ca 4400', 23. xi. 1981 (Tuck) (genitaliaslide no. 9479; BMNH). Paratypes. Sarawak: 1 cf , MtDulit, R. Koyan, 2500', primary forest, river-bank vegetation, 18. xi. 1932(Hobby & Moore) (BMNH); 1 $ , Gunung Mulu National Park, G. Mulu site 14 - camp 2.5, G.R. 413461,1000 m, lower montane forest, at m.v. light in canopy or understorey, ii.1978 (Holloway et al.) (genitaliaslide no. 9482; BMNH). REMARKS. This taxon is distinctively patterned and structurally atypical. The wing-pattern stronglyresembles that of Semeoloncha from West Africa and there are genital similarities in the male, notably inthe structure of the tegumen and uncus lobes. The valvae of the two genera are very different, however,Semeoloncha having a pair of separated soft, lobate, strongly setose valvae with an extraordinary processfrom the modified apodeme. Both genera have an atypically-shaped saccus and both have a taperedaedeagus; however, the aedeagus is smooth-surfaced in Semeoloncha but with numerous thorn-likecarinae in Gentingia. The female genitalia of Semeoloncha are unknown; however, the genitalia ofGentingia differ from those of all other Scardiinae in the peculiar shape of the eighth sternite and in thepresence of a single U-shaped denticulate signum at the point of inception of the ductus bursae on thecorpus bursae. FUNGUS MOTHS 97 SEMEOLONCHA Gozmany Semeoloncha Gozmany, 1968: 332. Type-species: Semeoloncha penicillata Gozmany, 1968: 324, byoriginal designation and monotypy. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia shorter than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus 5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outermid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with /? 3 and R 4 separate; A/ 3 andCuA\ stalked or very closely approximated at base; ground-colour very pale bronze, with pattern of small,purple-brown marginal spots and larger costal blotch. Male with coremata in eighth abdominal segment;coremata associated with short, lobe-like apodemes at anterior corners of eighth sternite. Male genitaliawith complex uncus fused with tegumen; tegumen unbroken, completely sclerotized dorsally; valva lackingbasal setose lobe on inner surface; apex of valva not forming ventral hook or hooks, without spines; valvaeseparate, not fused together ventrally, without longitudinal cleft; saccus longer than wide; juxta simple,entire, not divided medially; vesica with spicular cornuti; aedeagus smooth-surfaced, without spicularcarinae. CONSPICUOUS AUTAPOMORPHIES. Male with eighth tergite reduced, narrow and rod-like, broadeninganteriorly, thus T-shaped. Coremata very large, recessed as far as anterior margin of seventh segment;apodeme of valva with strong, arcuate dorsal process arising at one-third from apodeme base and directedcaudally. DISTRIBUTION. Afrotropical region - Sierra Leone.BIOLOGY. Unknown. Semeoloncha penicillata Gozmany (Fig. 44) Semeoloncha penicillata Gozmany, 1968: 334. Holotype cf , SIERRA LEONE (BMNH) [examined].Semeoloncha penicillata Gozmany; Gozmany & Vari, 1973: 152, fig. 460. ADULT (Fig. 44). cf , 20 mm. Vertex and frons whitish. Labial palpus whitish, strongly flecked with brownon outer and upper surfaces of first and base of second segment. Maxillary palpus cream, flecked abovewith brown, short, 5-segmented, not reaching one-half length of second segment of labial palpus. Antennacream; cilia 0-25 x (cf ) flagellar diameter. Thorax and tegula cream, slightly darker anteriorly. Forewingvery pale golden or straw-coloured (probably very faded) with small light brown marginal spots, thesemerged to form a terminal line between /? 4 and M 3 ; larger conspicuous spots just beyond one-half on costaand subtornally. Hindwing pale greyish cream with inconspicuous brown blotches on apical margin. Legscream; fore-tibia and fore-tarsus marked above with brown but conspicuously pale at articulations; outermid-tibial spur and outer proximal hind tibial spur 0-7 length of inner spurs. GENITALIA cf (Gozmany, 1968: figs 46-48; Gozmany & Vari, 1973: fig. 460). (For description of the caudalregion of the abdomen, see 'Remarks'.) Saccus diamond-shaped, almost as long as tegumen + uncus;uncus lobes entirely fused with and subsumed into tegumen, possibly forming the setose caudal margin;tegumen + uncus cowl-shaped, with strongly melanized horn-like lateral processes. Subscaphium notdeveloped. Juxta not developed but anellus swollen to form irregular and microtrichiate lobe at base ofeach valva; transtilla not developed. Valvae closely appressed (but not fused) basally, lobate, setose;apodemes extraordinarily elongate, each with long and arcuate caudally-directed spike arising from dorsalsurface two-thirds from apex. Aedeagus tapered, stout, 7 x as long as broad in middle; apex extended as anarrow tapering band of sclerotization; vesica with minute spicular cornuti (?microtrichia). GENITALIA $ . Unknown.DISTRIBUTION. Sierra Leone.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype cf , Sierra Leone: vii.1895 (Clements) (genitalia slide no. 14997; BMNH). Sierra Leone: 1 cf (paratype), data as holotype but v.1895 (genitalia slide no. 15008; BMNH). REMARKS. When examined originally, both genitalia preparations of this species (only two specimens areknown) were very poor. The basal three abdominal segments of the holotype remained attached to the 98 GADEN S. ROBINSON specimen but the remainder of the abdomen had been thrown away, leaving only the badly compressedgenitalia and aedeagus on the slide. In the case of the paratype, the terminal two abdominal segments onlyhad been retained; these were damaged and contorted, making the structure impossible to interpret. Thegenital armature was so badly crushed that it had split longitudinally down the tegumen. Like the holotype,it was grossly overstained. It was thus impossible to determine the structure of the subscaphium, juxta ortranstilla, or to code enough information for inclusion of Semeoloncha in the numerical analyses.Remounting of both preparations after rehydration, cleaning and differentiating clarified the structure ofthe diaphragma and details of the seventh and eighth segment. The eighth sternite is broad and thecoremata are set in the sternite rather than in the pleural membrane: the dorsal margin of the corematalinvagination is strengthened, forming the lateral margin of a dorsal triangular complex consisting ofsternite (lateral and outer anterior margins), tergite (medial rod-shaped structure with expanded anteriorend) and pleural membrane (region with small spiracle either side of rod-like tergite). This extraordinarystructure and the extreme modification of the valval apodeme serve to distinguish Semeoloncha from allother scardiine taxa. CRANAODES MeyrickCranaodes Meyrick, 1919: 238. Type-species: Cranaodes stereopa Meyrick, 1919: 239, by monotypy. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpus5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outer mid andproximal hind tibial spurs >O4 length of inner spurs. Forewing with /? 3 and /? 4 separate; M 2 , A/ 3 and CuA\separate; ground-colour pale bronze, with pattern formed of large purple-brown blotches (but ground-colour paler and blotches more extensive in oroya). Male with coremata in eighth abdominal segment;coremata associated with short, lobe-like apodemes at anterior corners of eighth sternite. Male genitaliawith complex uncus fused with tegumen; tegumen broken dorsally by at least a membranous suture line;valva lacking basal setose lobe on inner surface; apex of valva not forming ventral hook or hooks, withoutspines; valvae separate, not fused together ventrally, without longitudinal cleft; saccus longer than wide;juxta simple, entire, not divided medially; vesica lacking spicular cornuti; aedeagus smooth-surfaced,without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Forewing pattern consisting of bold purple-brown patches on a pale bronzeground-colour (except in oroya). DISTRIBUTION. Neotropical region - Colombia, Panama, Peru; Oriental region - Borneo.BIOLOGY. Unknown. Key to species of Cranaodes Males (males of sequestrata are unknown) 1 Dark markings of forewing divided by broad antemedial band of ground-colour (Fig. 45) ; valvawith densely setose medial digitate process on inner surface; costa and ventral margin of valva simple stereopa (p. 98) Dark markings of forewing continuous through antemedial region (Fig. 46); inner surface ofvalva without process; costa and ventral margin of valva strongly folded (Fig. 106) .... oroya (p. 99) Females (females of oroya are unknown) 1 Dark costal blotch on forewing broad, extending posteriorly to middle of cell (Fig. 47); ventral lip of ostium convex (Fig. 159) . (Borneo) sequestrata (p. 100) Dark costal blotch on forewing narrow, not extending further posteriorly than base of R$ (Fig.45); ventral lip of ostium with U-shaped emargination (Fig. 158). (Neotropical region) stereopa (p. 98) Cranaodes stereopa Meyrick (Figs 45, 158)Cranaodes stereopa Meyrick, 1919: 239. Holotype d", COLOMBIA (BMNH) [examined]. ADULT (Clarke, 1970: pi. 17, figs 2, 2a, 2b; Fig. 45). C?, 27 mm; $, 40 mm. Vertex and frons brownishyellow. Labial palpus cream, brownish on outer surface, slender, not strongly tufted on second segment. FUNGUS MOTHS 99 Maxillary palpus cream, flecked with brown above, relatively short, reaching only about one-half length ofsecond segment of labial palpus. Antennal scape and pedicel light grey-brown; cilia 1-7 x (c?) or 0-5 x (9)flagellar diameter. Thorax and tegula light yellow-brown, strongly flecked with dark brown anteriorly.Forewing pale bronze marked with strong purple-brown blotches forming a broken irregular postmedialband; quadrate basal blotch continuous with strong basicostal blotch; numerous smaller costal spots.Hindwing light brownish grey. Legs buff, foreleg and mid-leg dark brown above but paler at articulationsand across middle of mid-tibia; outer mid-tibial spur 0-6 length of inner; outer hind proximal tibial spur 0-7length of inner spur. GENITALIA c? (Clarke, 1970: pi. 17, figs 2c, 2d). Saccus triangular; uncus lobes entirely fused with outercorners of tegumen, each forming a setose and strongly sclerotized knob-like protuberance with a pair ofsmall apical spines. Subscaphium not developed. Juxta quadrate, strongly sclerotized, transtilla notdeveloped. Valva plain, with strong and setose medial digitate process, apodeme with shallow dorsally-directed flap. Aedeagus stout, 10 x as long as broad in middle, sclerotization of apex a thin tapered processonly, ventrally with distinctive subbasal longitudinal 'keel'. GENITALIA 9 (Fig. 158). Eighth tergite longer than eighth sternite, with three pairs of strong epimarginalsetae and numerous scattered (?)microsetae; eighth sternite with broad terminal ostium with strongV-shaped emargination of ventral lip; margin of eighth sternite either side of ostium with compact group offour elongate setae and numerous smaller setae. Antrum short, broadly funnel-shaped, inception of ductusseminalis posterior to anterior margin of eighth sternite. Ductus bursae thick- walled, inner surface rugose,rugosity with distinct transverse pattern that becomes transverse wrinkling further anteriorly. Corpusbursae thin-walled, elongately ovoid; signa absent. DISTRIBUTION. Panama; Colombia.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype O", Colombia: San Antonio, 5800', xi.1907 (genitalia slide no. JFGC 6645; BMNH).Panama: 1 $, Chiriqui (genitalia slide no. 13109; BMNH). REMARKS. This is one of the largest and most striking Tineidae, its characteristic wing-pattern serving todistinguish it from all other Neotropical species. Similarities to sequestrata may be entirely the result ofconvergence; however, in the absence of further material of sequestrata (particularly males) any discussionas to whether or not the two species are really congeneric would be conjectural. They differ in the length ofthe (female) antennal cilia, in the shape and size of the forewing markings and in the genitalia, stereopahaving a strong U-shaped emargination of the ventral lip of the ostium (= posterior margin of eighthsternite) and sequestrata having a slightly convex margin with considerably more strong and elongate setaethan stereopa. In both species the ductus bursae has fine transverse wrinkling (= regular constrictions) butin stereopa this is represented posteriorly by rugosity with a distinctive transverse pattern. This latterfeature may represent the unexpanded state of the ductus prior to copulation and the ductus may assume aregularly constricted form after being expanded. There is a marked difference between the two species inthe form of the wall of the ovipositor dorsal to the ostium: in sequestrata this wall is ballooned ventrally toform a lobe large enough to close off the ostium, whereas in stereopa the ovipositor wall is quite smooth.Similar swellings of the ovipositor wall occur in most Scardiinae but they are not always conspicuous nor, intheory, capable of closing over the ostium (there is no evidence that this is the function of these lobes). The function and homology of the flap from the valval apodeme of the male of stereopa is uncertain.Similar but much more elongate processes are found in Semeoloncha. Apodemal processes also occur inthe poorly-known genus Leptozanda (Robinson, 1976); this is probably also a scardiine taxon but thematerial available is in such condition as to make precise placement impossible. The apodemal processes ofthese groups are analogous to some of the modifications of the labides that occur in Tinissa (Robinson,1976) although probably not homologous. Cranaodes oroya sp. n. (Figs 46, 106) ADULT (Fig. 46). cf , 26 mm. Coloration and external structure of head similar to stereopa. Thoraxbrownish cream, a few darker scales anteriorly; tegula grey-brown, a few pale scales at margins. Forewingpale bronze (but badly faded) strongly marked with dark brown to form a possibly continuous anteriorfascia similar to that of Daviscardia species (but specimen very worn); paler spots along costa; conspicuouspale round spots at end of cell and at end of fold. Hindwing light grey-brown. Legs buff, foreleg and mid-leg 100 GADEN S. ROBINSON darker above (very worn); outer mid-tibial spur 0-7 length of inner; outer proximal hind tibial spur 0-75length of inner spur. GENITALIA cf (Fig. 106). Saccus elongately triangular; uncus lobes similar to those oistereopa but formingonly a very small pair of hook-shaped processes on the outer corners of the tegumen. Subscaphium notdeveloped. Juxta represented only by ill-defined thickening of anellus; transtilla not developed. Valva withstrongly folded costa and ventral margin, otherwise plain, apodeme without flap (but flap otstereopa mayrepresent initial development or reduction of costal fold of this species). Aedeagus almost cylindrical, 10 xas long as broad in middle, with triangular membranous zone from base to almost one-third and withsimilar but shorter zone from apex. GENITALIA $. Unknown.DISTRIBUTION. Peru.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype cf , Peru: R. Inambari, La Oroya, 3100', dry season, ix.1904 (Ockenderi) (genitalia slide no.12394; BMNH). REMARKS. Smaller and with much more extensive dark forewing markings than stereopa, oroya may also bedistinguished by the male genitalia in which the costa and ventral margin of the valva are strongly foldedinward. The flap from the valval apodeme ofstereopa is absent in oroya but may form a part of the costalfold. The middle of the valva bears a conspicuous setose digitate process mstereopa but is smooth in oroya.Although the wing pattern of the two species is strongly divergent, the similarities of the male genitalia(notably in the complex of the uncus lobes plus tegumen) suggest that they are congeneric. Cranaodes sequestrata Meyrick(Figs 47, 159,200) Craneodes [sic] sequestrata Meyrick, 1926: 164. Holotype cf, SARAWAK (SM, Kuching) [not found -probably destroyed]. ADULT (Robinson, 1984: pi. 6, fig. 4; Fig. 47). $, 32 mm. Vertex and frons yellow-brown. Labial palpuspale buff, strongly speckled with dark brown on outer surface, apex pale. Maxillary palpus grey-brown,reaching only about one-half length of second segment of labial palpus. Antennal scape and pedicel palebuff, flecked with dark brown; flagellum pale buff; cilia 1-0 x flagellar diameter. Thorax and tegulaoff-white, strongly flecked with grey-brown anteriorly, thorax almost entirely dark. Forewing lightgreenish bronze flecked with pale brown; large quadrate dark brown spot from costa to posterior margin ofcell at one-half; basal half of costa flecked with dark brown; dorsum strongly flecked with dark brown (butworn); a few small dark spots at apex and termen. Hindwing pale greyish buff. Legs pale buff; foreleg andmid-leg darker above, mid-tibia with pair of large diffuse blackish spots above; outer mid-tibial spur 0-5length of inner; outer proximal hind tibial spur 0-6 length of inner spur. GENITALIA cf Unknown. GENITALIA 9 (Fig. 159). Eighth tergite longer than eighth sternite, with four pairs of strong terminal setaeand numerous scattered smaller lateral setae; eighth sternite forming a protruding ledge that accommo-dates ostium behind its posterior margin; sternite with 10 pairs of elongate setae and a few smaller setaeclose to caudal margin. Antrum narrow and cylindrical, ostium closed by bulbous protrusion of ovipositorwall; inception of ductus seminalis just posterior to anterior margin of eighth sternite. Ductus bursaeelongate, thin-walled, but with fine and close-set transverse wrinkles, extending beyond apices ofapophyses anteriores. Corpus bursae thin- walled, elongately ovoid; signa absent. DISTRIBUTION. Borneo (Fig. 200).BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Brunei: 1 $, Bukit Pagon, LP 308, 5520', upper montane forest, 15-20. ii. 1982 (Robinson) (genitaliaslide no. 6981; BMNH). REMARKS. The female specimen described here agrees well with Meyrick's original description of the male:discrepancies are in the colour of the head ('greyish ochreous'), the ground-colour of the forewing FUNGUS MOTHS 101 ('prismatic white') and the absence of any mention of dark markings on the dorsum. It is unlikely thatMeyrick's holotype of this species, deposited originally in the Sarawak Museum, Kuching, still exists.Exhaustive searches at my request by the present curators (and by workers in the 1950s - Diakonoff , pers.comm.) have failed to turn up any of the material described in Meyrick's 1926 paper. I am reasonablyconfident that the female described here is of Meyrick's species. On the basis of the characters availablefrom this female there is no reason to exclude sequestrata from Cranaodes. However, the genitalia do differsubstantially from those otstereopa (q.v.) and the similarities may be entirely due to convergence. PECTINISCARDIA gen. n. Type-species: Cranaodes prosty lias Meyrick, 1927: 327. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpus5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outer mid andproximal hind tibial spurs >0-4 length of inner spurs. Forewing with R^ and R 4 separate; A/ 2 , M 3 and CuA\separate; mottled coloration forming cryptic, coarse 'moss' pattern. Male lacking coremata in eighthabdominal segment. Male genitalia with simple uncus - a pair of setose lobes - fused with tegumen;tegumen broken dorsally by at least a membranous suture line; valva lacking basal setose lobe on innersurface; apex of valva forming ventral hook or hooks, or with spines; valvae separate, not fused togetherventrally, without longitudinal cleft; saccus longer than wide; juxta simple, entire, not divided medially;vesica with spicular cornuti; aedeagus damaged - presence of carinae uncertain. CONSPICUOUS AUTAPOMORPHIES. Ventrocaudal margin of valva with strong pectinifer of about 12 spines(this character is paralleled in Morophaga formosana and iriomotensis) . DISTRIBUTION. Neotropical region - Colombia.BIOLOGY. Unknown. Pectiniscardia prostylias (Meyrick) comb. n. (Fig. 48)Cranaodes prostylias Meyrick, 1927: 327. Holotype cf , COLOMBIA (BMNH) [examined]. ADULT (Clarke, 1970: pi. 17, figs 2b, 3; Fig. 48). cf , 27 mm. Vertex and frons cream. Labial palpus cream,flecked on outer surface with brown (but very worn). Maxillary palpus cream flecked with brown,5-segmented, short, reaching only one-fifth length of second segment of labial palpus. Antennal scape andpedicel pale buff, scape flecked strongly above with brown but distal third cream; pedicel and basal flagellarsegments dark brown above; cilia 3-0 x flagellar diameter. Thorax and tegula worn and damaged.Forewing very worn, golden cream patterned with purple-brown; costa and fringe with conspicuouschequering. Hindwing pale cream (worn) with a few scattered darker scales at apex and termen. Legs(worn) pale buff; foreleg and mid-leg marked strongly above and on sides with brown but pale atarticulations; mid-tibia with pale oblique medial bar on outer surface; outer mid-tibial spur 0-6 length- ofinner spur; hind tibial spurs broken. GENITALIA cf (Clarke, 1970: pi. 17, figs3a, 3b). Saccus elongate, triangular, longer than tegumen + uncus;uncus lobes broad, square-ended. Subscaphium not developed. Juxta shield-shaped, hardly sclerotized;transtilla not developed. Valva an obliquely truncated rectangle, simple except for well-developedpectinifer of about 12 stout spines. Aedeagus 9 x as long as broad; apex damaged but with strongsickle-shaped subapical carina and at least a pair of smaller corniform carinae; vesica with minute spicularcornuti (microtrichia). GENITALIA $ . Unknown.DISTRIBUTION. Colombia.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype cf , Colombia: Central Cordilleras, 11,800', xi. 1920 (genitalia slide no. JFGC 6646; BMNH). REMARKS. The holotype of this species is so worn that it is impossible to provide good externalcharacteristics for its recognition. However, the alternated dark and light spots down the costa give it achequered appearance and this feature is distinctive: it is not nearly so strongly developed in any other New 102 GADEN S. ROBINSON World scardiine. The forewing pattern is otherwise hard to discern. The apical fascia is certainly pale: thedorsum has lost almost all scales but was probably also pale and the remainder of the wing appears to havebeen mottled with dark brown. Fresh specimens might resemble Morophagoides species such as iulina. Themale genitalia of Pectiniscardia are quite distinctive. A pectinifer is developed otherwise in the Scardiinaeonly in Morophagaformosana and iriomotensis and I believe this to be a clear case of convergence. HORMANTRIS Meyrick Hormantris Meyrick, 1927: 327. Type-species: Hormantris astragalopa Meyrick, 1927: 327, by monotypy. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpus5-segmented; pilifers absent; second segment of labial palpus longer than width of head. Legs damaged -relative lengths of tibial spurs unknown. Forewing with /? 3 and R 4 separate; M 2 , M 3 and CuA\ separate;mottled coloration forming cryptic, coarse 'moss' pattern. Male lacking coremata in eighth abdominalsegment. Male genitalia with simple uncus - a pair of setose lobes - separated from tegumen by narrowband of membrane; tegumen unbroken, completely sclerotized dorsally; valva lacking basal setose lobe oninner surface; apex of valva not forming ventral hook or hooks, without spines; valvae separate, not fusedtogether ventrally, without longitudinal cleft; saccus longer than wide; juxta simple, entire, not dividedmedially; vesica with spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Ventral half of inner surface of valva with dense field of thorn-like setae.DISTRIBUTION. Neotropical region - Colombia.BIOLOGY. Unknown. Hormantris astragalopa Meyrick (Fig. 49)Hormantris astragalopa Meyrick, 1927: 327. Holotype cf , COLOMBIA (BMNH) [examined]. ADULT (Clarke, 1970: pi. 29, figs 1, la, Ib; Fig. 49). 0", 34 mm. Head and thorax almost entirely denudedbut some white and brown scales on labial palpus. Maxillary palpus whitish, elongate, 5-segmented.Antennal scape and pedicel whitish with some brown scales; flagellum pale buff-cream; cilia about 8 xflagellar diameter. Forewing almost entirely denuded, pale buff-cream with scattered small brownspeckles; slightly larger brown spots on costa; large subapical dark brown spot. Hindwing off-whitemottled with pale grey. Foreleg buff strongly flecked with dark brown; other legs lost. GENITALIA d" (Clarke, 1970: pi. 29, figs Ic, Id). Saccus broad, elongate and strongly sclerotized; uncuslobes small, digitiform, poorly sclerotized, sparsely setose. Subscaphium not developed. Juxta, if present,represented by only slight swelling of anellus; transtilla not developed. Valva almost circular, dorsal halfstrongly spined on internal surface; ventral margin with rounded triangular flap at one-half and with short,stout basal setose lobe. Aedeagus dumb-bell-shaped with dense spinose carinae on dorsal surface of apicalhalf; vesica with group of eight spine-like cornuti. GENITALIA $. Unknown.DISTRIBUTION. Colombia.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype cf , Colombia: Mt Tolima, 12,500', x.1920 (genitalia slide no. JFGC 6644; BMNH). REMARKS. Hormantris differs from Cnismorectis (q.v.) in having a more elongate maxillary palpus; theantennal cilia are considerably longer, the longest of any scardiine known. Unlike Cnismorectis, the ventralsurface of the antenna is scaled and the pecten is sparse with, apparently, fewer than 15 bristles (but thespecimen is worn and the bristle-sockets have not been checked in a microscopic preparation). Theholotype of astragalopa is so worn as to make impossible any comparison of wing pattern except theobservation that the subapical dark brown spot does not occur in Cnismorectis. CNISMORECTIS MeyrickCnismorectis Meyrick, 1936: 109. Type-species: Cnismorectis choritica Meyrick, 1936: 109, by monotypy. FUNGUS MOTHS 103 DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia longer than 1-5 xflagellar diameter. Scape with more than 15 pecten bristles. Interocular index (male) greater than 1-0.Maxillary palpus with fewer than 5 segments; pilifers absent; second segment of labial palpus longer thanwidth of head. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R^ andR 4 separate; M 2 , M 3 and CuA\ separate; mottled coloration forming cryptic, coarse 'moss' pattern. Malelacking coremata in eighth abdominal segment. Male genitalia with simple uncus - a pair of setose lobes -separated from tegumen by narrow band of membrane; tegumen unbroken, completely sclerotizeddorsally; valva lacking basal setose lobe on inner surface; apex of valva not forming ventral hook or hooks,without spines; valvae separate, not fused together ventrally, without longitudinal cleft; saccus longer thanwide; juxta complex, entire, not divided medially; vesica with spicular cornuti; aedeagus smooth-surfaced,without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Ventral margins of valvae invaginated, contiguous with juxta which isinfolded anteriorly, the whole complex forming a deep, broad pocket; vesica with elongate oval of small,thorn-like cornuti. DISTRIBUTION. Neotropical region- Peru, Bolivia.BIOLOGY. Unknown. Cnismorectis choritica Meyrick(Figs 50, 105,160, 182) Cnismorectis choritica Meyrick, 1936: 109. LECTOTYPE $, BOLIVIA (BMNH), here designated [ex-amined]. ADULT (Figs 50, 182). cf $, 26-36 mm. Vertex and frons very pale buff with a few dark scales. Labialpalpus very pale buff flecked with brown. Maxillary palpus whitish, 4-segmented, very short, not reachingbase of second segment of labial palpus. Antennal scape and pedicel ochreous cream flecked with brown,flagellum ochreous cream; cilia 3.0 x (cf ) or 0.3 x ($) flagellar diameter. Thorax and tegula very pale buffflecked with brown. Forewing cream flecked and finely strigulated with orange-brown (particularly alongveins) and dark brown, dark markings concentrated to leave ill-defined paler ante- and postmedial fasciae.Hindwing dirty cream strigulated with pale grey. Legs pale buff strongly flecked with brown, particularlyabove on fore-leg and mid-leg; mid-tibia conspicuously rough-scaled; tarsi ringed with brown; outermid-tibial spur 0.8 length of inner; outer proximal hind tibial spur 0.6 length of inner spur. GENITALIA cf (Fig. 105). Saccus slender, elongate; uncus lobes slender, small, only slightly sclerotized,setose. Subscaphium not developed. Juxta apparently forming ventral wall of pocket that extendsanteriorly behind saccus and extends laterally into ventral margins of valvae; transtilla not developed.Valva almost rectangular with dorsal corner produced into a spatulate lobe; ventral margin invaginated toform conspicuous pocket; internal surface with coarse microtrichia and two or three small peg-likeprocesses (?sensillae) at four-fifths. Aedeagus slender, elongate, 20 x as long as broad in middle, withoutcarinae; base of vesica with elongate oval of small, thorn-like cornuti. GENITALIA $ (Fig. 160). Seventh sternite with corethrogyne of fine, elongate hairs. Eighth tergite slightlylonger than eighth sternite, deeply emarginate medially, with two pairs of elongate terminal setae and fivepairs of smaller terminal setae. Eighth sternite with deep U-shaped medial emargination accommodatingostium, with dense, stiff, elongate bristles. Antrum narrow, cylindrical, one-third length of apophysesanteriores. Ductus bursae thin-walled, as long as antrum; inception of ductus seminalis close to junctionwith antrum. Corpus bursae thin-walled, pyriform, twice length of antrum; signa absent. DISTRIBUTION. Bolivia, Peru.BIOLOGY. Unknown. MATERIAL EXAMINED. 5 ex. Lectotype $ , Bolivia: Santa Cruz, Samaipata, 1500 m, iii (Steinbach) (genitalia slide no. 6970; BMNH). Bolivia: 1 <j> (paralectotype), 1934 (Staudinger) (BMNH); 1 cf , 1 $, Rio Tanampaya, 1894 (Garlepp)(genitalia slide no. 6969; BMNH). Peru: 1 $, Huanuco, Cushi, 1900 m (Hoffmann) (BMNH). REMARKS. Cnismorectis is the sister-group of Hormantris. The two are conspicuous in that they have adistinctive mottled pattern on the hindwings and exceptionally elongate labial palpi: both have lost thepilifers. In the male genitalia of both, the uncus lobes are very small and digitiform and the vesica hasthorn-like cornuti. The microtrichia on the valva of Cnismorectis are, however, replaced functionally by a 104 GADEN S. ROBINSON field of very small thorn-like spines covering the dorsal half of the valva in Hormantris. The relationship of these genera to Dorata Busck, a genus of five greyish cream species with plainhindwings from northern Mexico and south-western U.S.A., requires further investigation. Superficially,Dorata does not look like a scardiine but, like Cnismorectis and Hormantris, has elongate labial palpi andlacks pilifers. The uncus lobes are similar to those of Cnismorectis but bear strong, thorn-like spines.Similar spines occur on the dorsal lobe of the strongly bifurcated valva which also carries a setose digitateprocess from the ventral margin in the one species examined (Dorata atomophora Meyrick). The ventrallobe of the valva of Dorata may represent the juxta: a pouch superficially similar to that of Cnismorectisruns anteriorly from the ventral base of the valva. Davis (in prep.) is including Dorata in a revision of theNorth American Tineidae. MINISCARDIA gen. n. Type-species: Scardia minimella Busck, 1914: 65. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus with fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width ofhead. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R^ and R 4separate: M 3 and CuAi stalked or very closely approximated at base; mottled coloration forming cryptic,coarse 'moss' pattern. Male with coremata in eighth abdominal segment; coremata not associated withapodemes. Male genitalia with simple uncus - a pair of setose lobes - fused with tegumen; tegumenunbroken, completely sclerotized dorsally; valva lacking basal setose lobe on inner surface; apex of valvanot forming ventral hook or hooks, without spines; valvae separate, not fused together ventrally, withoutlongitudinal cleft; saccus longer than wide; juxta, if present, simple, entire, not divided medially; vesicawith spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. None; however, may be recognized by the exclusive combination ofmoss-like forewing pattern with veins M 3 and CuA\ stalked or connate - see 'remarks' for minimella. DISTRIBUTION. Neotropical region - Panama, Costa Rica, Guatemala, Brazil; Nearctic region - U.S.A.(Arizona). BIOLOGY. Unknown. Miniscardia minimella (Busck) comb. n. (Figs 55, 56, 111,165)Scardia minimella Busck, 1914: 65. Holotype cf , PANAMA (NMNH) [examined]. ADULT (Figs 55, 56). cf , 15-19 mm. $ , 20-25 mm. Vertex and frons orange-brown, with dark brown tuftsclose to eyes. Labial palpus orange-brown, strongly marked with deep purple-brown on outer surface.Maxillary palpus dull buff, deep purple-brown on upper and outer surfaces, probably 3-segmented,reaching only one-half length of second segment of labial palpus. Antennal scape and pedicel orange-brown, dark purple-brown above; flagellum scales light grey-brown but darker at base of flagellum; cilia1-5 x (cf ) or 0-2 x ($) flagellar diameter. Thorax and tegula deep purple-brown, thorax with buff-creamtransverse band, tegula tipped with buff-cream. Forewing buff-cream, strongly speckled with deeppurple-brown, speckling concentrated into basal, medial and subterminal fasciae. All deep purple-brownmarkings (especially on underside) with pronounced blue-green iridescence when observed at an acuteangle. Hindwing light brownish grey. Legs buff, foreleg and mid-leg heavily marked above and at sideswith deep purple-brown, but pale at articulations; mid-tibia with pair of conspicuous oblique dark bars onouter surface; hindleg greyish above; outer mid-tibial spur 0-5 length of inner; outer hind proximal tibialspur 0-75 length of inner spur. GENITALIA cf (Fig. 111). Saccus elongate, triangular, more than half length of genital armature; uncuslobes triangular, fused with each other and with tegumen, sclerotized, setose, apices level with apices ofvalvae. Subscaphium not developed. Juxta not developed or represented only by ill-defined thickening ofbase of anellus; transtilla not developed. Valva slender, triangular, simple. Aedeagus 7 x as long as broad,with subapical dorsal transverse band of small, thorn-like carinae; vesica with minute spicular cornuti(?microtrichia). GENITALIA $ (Fig. 165). (Description based on examples from French Guiana and Brazil.) Eighth tergite FUNGUS MOTHS 105 slightly longer than eighth sternite, with about 8 strong setae close to caudal margin and pair of large medialpatches of microtrichia; eighth sternite with shallowly m-shaped caudal margin, strongly folded medially toaccommodate broad, transverse ostium with convex or slightly concave ventral lip; apex of eighth sternitewith two or three pairs of strong setae and numerous short, small setae. Antrum with broad posteriorchamber, dorsal surface of chamber with dome-shaped thickening surrounding inception of ductusseminalis; anterior region of antrum a short, smooth-walled, narrow tube reaching less than 0-25 length ofapophyses anteriores. Ductus bursae thick-walled posteriorly, with about 15 irregular transverse constric-tions, becoming very thin-walled anteriorly. Corpus bursae globular, very thin-walled, extending to 1-25length of apophyses anteriores; signa absent. DISTRIBUTION. Guatemala; Costa Rica; Panama; French Guiana; Brazil.BIOLOGY. Unknown. MATERIAL EXAMINED. 7 ex. Holotype cf , Panama: Porto Bello, v.1912 (Busck) (genitalia slide no. 20156; NMNH, Washington). Guatemala: 1 cf, Poptum, Peten, 15-16. ix. 1973 (Becker) (coll. Becker, Brasilia). Costa Rica: 1 cf , SanJose, 1922 (genitalia slide no. 12689; BMNH); 1 $, 1935 (H. S.) (genitalia slide no. 12680; BMNH). FrenchGuiana: 1 $ , Nouvelle Chantier, x. (Le Moult) (genitalia slide no. 12684; BMNH); 1 $ , St Jean du Maroni(Le Moult) (BMNH). Brazil: 1 , Rio Brilhante, Mato Grosso, 23-27. x. 1970 (Becker) (coll. Becker,Brasilia). REMARKS. This species is one of the few Neotropical taxa of Scardiinae that has an olivaceous 'moss' patternresembling that of the Morophaga choragella-group. Other Neotropical taxa with this pattern are Diataga(with M 2 and M 3 stalked in the forewing and with very short outer tibial spurs) and Morophagoides (allveins free , pattern generally more variegated) . In the forewing A/ 3 and CuA } are always stalked or connate ;this characteristic is also found in the smaller and darker Bythocrates and in Perilicmetis with itscharacteristic dark brown subterminal fascia. The male genitalia are remarkable for their simplicity ofstructure, the female genitalia for the internal thickened and sclerotized ridge that is the posteriorcontinuation of the apophyses anteriores extending and remaining conspicuous as far posteriorly as thesetose apical region of the eighth sternite. The females listed here may not be conspecific with each other or with the males. A single female fromArizona (see below) appears to be a congener but not conspecific. The female from Brazil has similargenitalia to those of the female from French Guiana but the ventral lip of the ostium is concave in theformer and convex in the latter. The broad posterior region of the antrum is much less strongly sclerotizedin the Brazilian specimen and the transverse constrictions of the ductus bursae are more numerous, aboutdouble the number present in the French Guiana specimen. The female from Costa Rica, with a 5 mmlonger wingspan than the other specimens examined, has only 5 pairs of apical setae on the eighth sterniteand a broader, shorter antrum lacking the narrow anterior region of the other two examples; the ductusbursae lacks transverse constrictions and the bursa copulatrix, although collapsed in preparation, does notextend beyond the apophyses anteriores. Miniscardia species A (Fig. 166) ADULT. $ , 17 mm. Coloration and external structure similar to minimella.GENITALIA cf . Unknown. GENITALIA $ (Fig. 166). Similar to those of minimella but ventral lip of ostium not extended as farposteriorly, and with distinct marginal bulbosity; antrum short, not extending anteriorly beyond inceptionof ductus seminalis; posterior region of ductus bursae distinctly scobinate on external surface; apex ofeighth s'ternite with three pairs of strong setae. DISTRIBUTION. U.S.A. (Arizona).BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. U.S.A.: 1 9, Arizona, CochiseCo., HuachucaMts, Miller Canyon, 7. viii. 1974 (Powell) (genitalia slide;UC). REMARKS. This specimen probably represents a species different from minimella but, with only a singlefemale example in poor condition available for examination, its status remains uncertain. The matter is 106 GADEN S. ROBINSON complicated by the apparent variability of females of minimella. The example of minimella from CostaRica has genitalia very similar to those of this specimen from Arizona; both lack sclerotization of the ductusbursae to form an antrum anterior to the inception of the ductus seminalis, and in both the ventral lip of theostium is almost straight, rather than being convex as in examples from Brazil and French Guiana. NECROSCARDIA gen. n. Type-species: Tirtea funeratella Zeller, 1863: 144. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus with fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width ofhead. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with R 3 and R 4separate; M 2 , M 3 and CuA\ separate; pattern consisting of pale termen and dorsum on darker ground-colour. Male with coremata in eighth abdominal segment; coremata associated with elongate, rod-likeapodemes at anterior corners of eighth sternite. Male genitalia with complex uncus separated fromtegumen by narrow band of membrane; tegumen broken dorsally by at least a membranous suture line;valva lacking basal setose lobe on inner surface; apex of valva not forming ventral hook or hooks, withoutspines; valvae separate, not fused together ventrally, without longitudinal cleft; saccus wider than long;juxta complex, divided medially; vesica with spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Apex of tegumen forming a single prong-like or bifid process dorsal to theuncus lobes; juxta enormously enlarged, bipartite, closely appressed to and forming a functional part of thevalvae. DISTRIBUTION. Neotropical region- Venezuela, Colombia, Bolivia.BIOLOGY. Unknown. Key to species of Necroscardia 1 Pair of cream triangular marks on posterior margin of forewing equal in size (Fig. 54); malewithout large phylliform setae on eighth sternite; male with dorsal process from tegumenbifid, extending caudally well beyond lobes of uncus (Fig. 109); female seventh sternite withposterior margin strongly emarginated medially, with scattered field of strong, needle-like setae (Fig. 163) morticina(p. 107) Proximal cream triangular mark on posterior margin of forewing considerably larger than distalmark which is reduced to a minute spot (Fig. 53); male with two pairs of enormous phylliformsetae on posterior corners of eighth sternite (Fig. 110); male with dorsal process from tegumena single spine, not extending beyond lobes of uncus (Fig. 108); female seventh sternite withposterior margin transverse, lacking needle-like setae funeratella(p. 106) Necroscardia funeratella (Zeller) comb. n. (Figs 53, 108, 110, 164)Tinea funeratella Zeller, 1863: 144. LECTOTYPE cf , VENEZUELA, here designated [examined]. ADULT (Zeller, 1863: pi. 2, fig. 6; Fig. 53). cf , 20 mm; $ , 23 mm. Vertex and frons brownish cream, reddishbrown near eyes. Labial palpus cream flecked with brown, purple-brown on outer surface but pale at apex,sparsely scaled. Maxillary palpus greyish brown above, paler beneath, short, not as long as second segmentof labial palpus, probably 3-segmented. Antennal scape and pedicel greyish brown; flagellum greyishbrown cilia 1-7 x (cf) or 0-7 x ($) flagellar diameter. Thorax and tegula cream, anterior third brown.Forewing purple-brown with cream terminal fascia containing brown speckling and small brown apical andmedioterminal spots; costa with a few pale flecks; posterior margin with broad cream spot, large triangularcream spot almost at mid-point, and small cream spot at three-quarters. Hindwing light brownish grey.Legs cream, finely flecked with grey-brown; foreleg and mid-leg grey-brown above and on outer surfaces,tibiae traversed by cream band; articulations pale; outer mid-tibial spur 0-7 length of inner; outer proximalhind tibial spur 0-8 length of inner spur. GENITALIA cf (Figs 10B , 1 10) . Eighth sternite with elongate apodemes reaching anterior margin of seventhsternite, at each posterior corner with one large and one small strong phylliform seta. Saccus broadlytriangular, shorter than valvae; uncus lobes minute, setose, but sclerotized and lacking setae at apices, FUNGUS MOTHS 107 surmounted by spike-shaped caudal process from vinculum almost as long as uncus lobes. Subscaphium notdeveloped. Juxta strongly developed, bipartite, forming pair of tuberculate setose lobes surmounting andappressed basally to valvae; transtilla not developed. Valva short, plain, pyramidal, bearing lobe of juxtaon inner surface. Aedeagus small, strongly tapered apically, and with scattered minute thorn-like carinae;vesica with very few minute spicular cornuti. GENITALIA $ (Fig. 164). Seventh sternite unmodified. Eighth tergite longer than eighth sternite, with lineof minute setae at posterior margin, five or six pairs of very strong submarginal setae and four or five pairsof conspicuous but minute pits scattered in posterior one-quarter. Eighth sternite and genital tractpathologically deformed in specimen examined: illustrated but not described further here. DISTRIBUTION. Venezuela; Colombia.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Lectotype cf , Venezuela: (genitalia slide no. 12379; BMNH).Colombia: 1 $, Sierra del Libano, 6000', v. 1899 (Smith) (genitalia slide no. 12396; BMNH). REMARKS. For differentiation ot funeratella and morticina, see the key above. The female described aboveresembles the lectotype remarkably closely and I am confident that they are conspecific. Walsinghamidentified the female as funeratella - the specimen bears his determination label. Necroscardia morticina sp. n. (Figs 54, 109, 162, 163) ADULT (Fig. 54). cf , 21 mm. $ , 17 mm. Coloration and external structure similar to funeratella but apicesof second and third segments of labial palpus cream, third segment of cf entirely cream; cilia ($) 3 xflagellar diameter; posterior margin of forewing with pale spots coalesced to form continuous band frombase to tornus, spots triangular, of equal size; posterior and terminal cream fascia strongly speckled withpurple-brown; outer mid-tibial spur 0-6 length of inner; outer proximal hind tibial spur 0-7 length of innerspur. GENITALIA cf (Fig. 109). Eighth sternite with short, rod-like apodemes reaching posterior margin ofseventh sternite. Saccus broadly triangular, shorter than valvae; uncus lobes small, outer corners producedinto shallow, setose process, inner posterior margin sclerotized and lacking setae, surmounted byV-shaped process from tegumen that extends well beyond tips of uncus lobes. Subscaphium not developed.Juxta strongly developed, bipartite, forming pair of strong, triangular setose lobes with shallowly serratedorsocaudal margin appressed to and set in apices of valvae; transtilla not developed. Valva short, plain,half barrel-shaped, with juxta lobe set in top of 'barrel'. Aedeagus slender, apex sclerotized only on oneside and duck's bill-shaped, with fine thorn-like carinae at apex; vesica with minute spicular cornuti nearinception of ductus ejaculatorius. GENITALIA $ (Figs 162, 163). Seventh sternite rounded caudally, with shallow medial U-shaped emargina-tion, strongly sclerotized, with scattered strong setae. Eighth tergite slightly longer than eighth sternite,with narrowly V-shaped mediocaudal emargination; posterior margin with numerous small setae, five pairsof strong submarginal setae; four or five pairs of scattered pits at four-fifths posteriorly; a few small,scattered setae at one-half. Eighth sternite wrinkled laterally, short, forming strong keel-shaped sterigmawith trilobed apex (= ventral lip of ostium); apical one-third of sterigma with 10-12 pairs of strong,elongate setae. Antrum elongate, broad but tapering anteriorly, one-half length of apophyses anteriores,lined with elongate microtrichia; inception of ductus seminalis at posterior extremity of antrum (!). Ductusbursae as long as antrum, with fine transverse constrictions. Corpus bursae thin- walled, ovoid; signaabsent. DISTRIBUTION. Bolivia.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype cf , Bolivia: Cochabamba, Yungas del Espiritu Santo, 1888-9 (Germain) (genitalia slide no.12395; BMNH). Paratype. 1 $ , data as holotype (genitalia slide no. 12406; BMNH). REMARKS. For differentiation of morticina and funeratella, see the key above. The female paratype of 108 GADEN S. ROBINSON morticina is the only specimen of Necroscardia that could be considered to be in even fair conditions. Likethe other three specimens, however, the second segment of the labial palpus is not densely scaled as inother scardiine genera with a similar wing-pattern (Moscardia, Scardia, Daviscardid). TINISSA Walker Tinissa Walker, 1864: 780; Robinson, 1976 (revision); 1981 (reclassification and additional species). Type-species: Tinissa torvella Walker, 1864: 780, by monotypy.Polymnestra Meyrick, 1927: 331. Type-species: Polymnestra perilithias Meyrick, 1927: 331, by monotypy. [Synonymized by Gozmany & Vari, 1973: 85.] DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface scaled; cilia longer or shorter than flagellardiameter. Scape with more than 15 pecten bristles. Interocular index (male) greater than 1-0. Maxillarypalpus short, 3-segmented (but a few species with 5 segments) (Fig. 181); pilifers absent; second segment oflabial palpus shorter than width of head. Outer mid and proximal hind tibial spurs >0-4 length of innerspurs. Forewing with Rj, and R 4 separate; M 2 , M 3 and CuA\ separate; pattern consisting of pale spots orspeckles on darker ground-colour. Male with or without coremata in eighth abdominal segment; corematawithout associated apodemes. Male genitalia with simple or complex uncus separated from tegumen bynarrow band of membrane; valva lacking basal setose lobe on inner surface; apex of valva not formingventral hook or hooks, without spines; tegumen broken dorsally by at least a membranous suture line;valvae separate, not fused together ventrally, without longitudinal cleft; saccus wider or narrower thanlong; juxta complex, entire in most species but narrowed ventrally almost to the point of division; vesicalacking spicular cornuti; aedeagus with or without spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Hind tibia with apical and subapical tufts of elongate, dark-tipped scales;juxta of male complex, strongly modified and incorporated with valvae to form a functional valvae-juxtacomplex (as in Necroscardia); females of most species with segments VII markedly reduced in length butwith extended intersegmental membrane between segments VII and VIII bearing a strong corethrogyne offine, elongate scales. DISTRIBUTION. Afrotropical region; Oriental region; Australian region as far east as the Solomon Is and asfar south as Queensland. BIOLOGY. One species has been reared from fungus on bamboo (see Robinson, 1976: 257). Typical habitatsare shown in Figs 199 and 200. REMARKS. Further treatment of Tinissa is not provided here: descriptions and illustrations of the 34included species have been provided by Robinson (1976; 1981). SCARDIELLA gen. n. Type-species: Scardia approximated Dietz, 1905: 27. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with fewer than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpus5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outer mid andproximal hind tibial spurs >0-4 length of inner spurs. Forewing with 7? 3 and R 4 separate; M 2 , M 3 and CuAiseparate; mottled coloration forming cryptic, coarse 'moss' pattern. Male lacking coremata in eighthabdominal segment. Male genitalia with complex uncus separated from tegumen by narrow band ofmembrane; tegumen broken dorsally by at least a membranous suture line; valva lacking basal setose lobeon inner surface; apex of valva not forming ventral hook or hooks, without spines; valvae separate, notfused together ventrally, without longitudinal cleft; saccus longer than wide; juxta simple, entire, notdivided medially; vesica with spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Bulbus ejaculatorius of male very large, almost 3 x length of aedeagus;ductus bursae of female with spicular inner surface from two-thirds to nine-tenths posteriorly (this alsooccurs, probably by convergence, in Montescardiafuscofascielld). DISTRIBUTION. Nearctic region.BIOLOGY. Unknown. FUNGUS MOTHS 109 Scardiella approximately (Dietz) comb. n.(Figs 52, 107, 161) Scardia approximated Dietz, 1905: 27. LECTOTYPE cf , U.S.A. (NMNH), here designated [examinedon author's behalf by D. R. Davis]. ADULT (Dietz, 1905: pi. 1, fig. 8; Fig. 52). O"$, 13-16 mm. Vertex and frons cream, tinted with brownaround eyes. Labial palpus whitish, strongly flecked with brown on outer surface of second segment andmiddle of outer surface of third segment. Maxillary palpus whitish, flecked with light brown, 5-segmented,extending to three-quarters length of second segment of labial palpus. Antennal scape, pedicel andflagellum greyish buff, dark brown above, flagellum dark brown above only on basal five or six segments;cilia 2-0-2-5 x (cf) or 0-4 x () flagellar diameter. Thorax and tegula cream, tegula strongly fleckedanteriorly with brown, thorax flecked laterally and far anteriorly with brown. Forewing whitish, stronglyflecked with light orange-brown, particularly along veins, and patterned with dark brown to form strongmedial and basal fasciae and dense spotting over remainder of wing; fringe with medial grey-brown line andirregular chequering. Hindwing pale greyish cream, fringe lighter. Legs pale buff-cream, foreleg andmid-leg strongly marked with dark brown above and on sides (including tarsi) but pale at articulations andwith strong whitish bar across mid-tibia; hind tarsus strongly marked above with brown; outer mid-tibialspur 0-6 length of inner; outer proximal hind tibial spur 0-7 length of inner spur. GENITALIA cf (Fig. 107). Saccus elongate, triangular; uncus lobes a pair of digitiform processes, widelyseparated, strongly sclerotized. Subscaphium broad but represented only by slight thickening of dia-phragma. Juxta represented only by almost imperceptible thickening of anellus; transtilla not developed.Valva short, rounded, with strong basicostal digitate process. Aedeagus stout, 7 x as long as broad, withshallowly triangular subapical carinae and numerous minute scattered thorn-like carinae; vesica withminute spicular cornuti (microtrichia); bulbus ejaculatorius very long, almost 3 x length of aedeagus. GENITALIA $ (Fig. 161). Eighth tergite longer than eighth sternite, with three pairs of strong subapicalsetae; eighth sternite with strongly developed sterigma accommodating short, cylindrical antrum; eighthsternite quadrangular caudal to sterigma, hardly sclerotized, with two pairs of strong setae and 12-15 pairsof scattered smaller setae; ventral lip of ostium strongly concave. Antrum not extending beyond anteriormargin of eighth sternite. Ductus bursae elongate, 1-5 x length of apophyses anteriores, lined with stoutmicrotrichia anterior to inception of ductus seminalis at level of one-half length of apophyses anteriores(i.e., short length of ductus bursae posterior to ductus seminalis is without microtrichia). Corpus bursaepyriform, continuous with ductus, thin-walled; signa absent. DISTRIBUTION. U.S.A.: Pennsylvania, New Jersey, District of Columbia, Maryland, Ohio (Dietz, 1905);New York; Massachusetts, Georgia (Forbes, 1923). Canada: Toronto. BIOLOGY. The larva has been found in a rotten sycamore log (Forbes, 1923). MATERIAL EXAMINED. 5 ex. Lectotype cf , U.S.A.: Maryland, Plummer's Island, vii.1901 (Busck) (NMNH) [examined on author'sbehalf by D.R.Davis]. U.S.A.: 1 $ (paralectotype), Pennsylvania, Hazleton, 15.vii.1896 (Dietz) (BMNH); 1 $, New York,Fishers, 21.vii.1933 (Klots) (genitalia slide no. 12403; BMNH). Canada: 3 cf, Toronto, vi.1922 (Parish)(genitalia slide no. 12402; BMNH). REMARKS. This small species is very distinctive. Its pattern is lighter than that of all other North Americanscardiines and resembles more that of a nemapogonine. The antennal cilia of the male are extraordinarilylong: few other Scardiinae have cilia more than twice as long as the diameter of the flagellar segments. Thestrongly patterned hind tarsi are also characteristic: they are pale in most other scardiines. Scardiellaappears to be a 'primitive' taxon. It has only six apomorphies (Table 1) and its affinities are obscure. Itsapomorphic features are its elongate antennal cilia, absence of coremata, complex uncus, dorsallydiscontinuous tegumen, sparse setal patch on the second abdominal sternite, and its basicostal process onthe valva. This last has been scored as a 'basal setose lobe' in Table 1, homologous with that ofMorophaga,Amorophaga and Diataga. However, I believe it is an independent development and it is not included inthe generic diagnosis. Afroscardia also has a setose lobe and this has been scored similarly; the lobe issubcostal and situated far distad on the valva . It seems unlikely that it is homologous with that of Scardiella .The affinity of the two taxa suggested by numerical analysis is, in all probability, due to convergence. 110 GADEN S. ROBINSON AFROSCARDIA gen. n. Type-species: Polymnestra capnochalca Meyrick, 1932ft: 207. DIAGNOSIS. Antenna (male) with dorsal cilia, ventral surface scaled; cilia longer than 1-5 x flagellardiameter. Scape with more than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillary palpuswith fewer than 5 segments; pilifers present; second segment of labial palpus shorter than width of head.Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with /? 3 and R 4 separate ; M 2 ,A/3 and CuAi separate; mottled coloration forming cryptic, coarse 'moss' pattern. Male lacking corematain eighth abdominal segment. Male genitalia with simple uncus - a pair of setose lobes - fused withtegumen; tegumen broken dorsally by broad, membranous region; valva lacking basal setose lobe on innersurface; apex of valva forming ventral hook or hooks, or with spines; valvae separate, not fused togetherventrally, without longitudinal cleft; saccus longer than wide; juxta simple, entire, not divided medially;vesica lacking spicular cornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Sclerotization of tegumen interrupted dorsally by broad membranousregion between lateral arms of tegumen; lobes of uncus fused with tegumen so that genital armatureterminates dorsocaudally in a pair of widely separated lobes. DISTRIBUTION. Afrotropical region - Uganda.BIOLOGY. Unknown. Afroscardia capnochalca (Meyrick) comb. n. (Fig. 51)Polymnestra capnochalca Meyrick, 1932ft: 207. Holotype cf , UGANDA (BMNH) [examined]. ADULT (Fig. 51). Cf $, 18-20 mm. Vertex and frons purple-brown. Labial palpus purple-brown, secondsegment white on inner surface and on apex of ventral tuft. Maxillary palpus whitish, very short,3-segmented. Antennal scape, pedicel and flagellum dark brown, paler beneath and on pecten; cilia 2-0 x(cf) or 0-8 x (9) flagellar diameter. Thorax and tegula purple-brown. Forewing uniformly purple-brownwith violet iridescence, apex slightly paler; fringe yellow. Hindwing greyish brown, fringe yellowish. Legsochreous buff, dark brown above and on sides; outer mid-tibial spur 0-6 length of inner; outer proximalhind tibial spur 0-75 length of inner spur. GENITALIA cf (Gozmany & Vari, 1973: fig. 451). Saccus elongate, tapered, longer than tegumen + uncus;uncus lobes flap-like, small, widely separated, only weakly sclerotized; transtilla not developed. Valvaelongately triangular with slightly hooked apex, with small subcostal setose lobe at two-thirds. Aedeagusslender, elongate , more than 15 x as long as broad, with very fine spicular subapical carinae ; vesica withoutcornuti. GENITALIA $. Unknown.DISTRIBUTION. Uganda (Ruwenzori Mts).BIOLOGY. Unknown. MATERIAL EXAMINED. 5 ex. Holotype cf , Uganda: [Ruwenzori Mts], Bujubis, 12,000', 16.viii.1931 (Hancock) (genitalia slide no.10264; BMNH). Uganda: 1 9 (paratype) (abdomen missing), Ruwenzori Mts, 10,000', viii.1931 (Hancock) (BMNH); 1Cf , Ruwenzori Range, Bigo, 11,400', 29. vii. 1952 (Fletcher) (BMNH); 2 cf , Ruwenzori Range, heath zoneabove Nyamgasani Valley, 12-13,000', xii.1934-i.1935 (Buxton) (BMNH). REMARKS. This is the only scardiine that has an entirely plain-coloured forewing. It resembles superficiallyBythocrates from the New World, but in that genus the fringes are concolorous with the forewing and thereis a row of pale spots along the costa. Afroscardia is distinctive also in that, despite its having a completevenation (although R 4 and R 5 are connate), it is very narrow-winged, the forewing more than four times aslong as broad. The only other scardiine genus with the wings so slender is Diataga. The male genitalia ofAfroscardia are distinctive in that the tegumen is broken dorsally by a membranous zone and the uncuslobes are widely separated: the terminal arms of the tegumen are strongly sclerotized and rod-like. FUNGUS MOTHS 111 AMOROPHAGA Zagulajev ^Amorophaga Zagulajev, 1966: 634. [Nomen nudum.] Amorophaga Zagulajev, 1968: 329. Type-species: Amorophaga hyrcanica Zagulajev, 1968: 331, figs 1, 2,by original designation and monotypy. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia longer than 1-5 xflagellar diameter (but shorter in hyrcanica). Scape with more than 15 pecten bristles. Interocular index(male) 1-0 or less. Maxillary palpus 5-segmented; pilifers present; second segment of labial palpus shorterthan width of head. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with /? 3and /? 4 stalked or very closely approximated at base; M 2 , M 3 and CuA\ separate; pattern elementsconsisting of greyish or olivaceous longitudinal streaks on a paler background. Male with coremata ineighth abdominal segment; coremata without associated apodemes. Male genitalia with complex uncusseparated from tegumen by narrow band of membrane ; tegumen broken dorsally by at least a membranoussuture line; valva with setose basal lobe on inner surface; apex of valva forming ventral hook or hooks, orwith spines; valvae separate, not fused together ventrally, with deep emargination forming longitudinalcleft; saccus wider than long; juxta complex, divided medially; however, juxta simple and undivided incryptophori, vesica lacking spicular cornuti; aedeagus with spinose carinae (but not in cryptophori). CONSPICUOUS AUTAPOMORPHIES. Wing pattern composed of longitudinal streaks. DISTRIBUTION. Holarctic-U.S.S.R. (Azerbaidjan), Nepal, Japan, U.S.A. (Idaho, California). BIOLOGY. See Amorophaga cryptophori (Clarke). Key to species of Amorophaga Males (males of rosemariae are unknown) 1 Antennal cilia shorter than 1-5 x flagellar diameter; uncus broad, short, with pair of small lateral ventrally-directed hooks hyrcanica(p. 113) - Antennal cilia longer than 1-5 x flagellar diameter; uncus narrow, elongate, without lateral hooks 2 2 Uncus with pair of strong, curved processes directed ventrally and caudally and resembling a gnathos. (western U.S.A.) cryptophori (p. 112) - Uncus lacking any ventrally-directed processes (Fig. 112). (Japan) japonica(p. 114) Females (females of hyrcanica and japonica are unknown) 1 Forewing ground-colour cream (with olivaceous tint when fresh), strongly marked with orange-buff and dark brown, overall appearance olivaceous (Fig. 57) ; outer mid tibial spur only about0-3 length of inner spur; frenulum with two bristles; seventh sternite with posterior marginstrongly sclerotized and serrated (Fig. 168) . (Nepal) rosemar/ae(p. Ill) - Forewing ground-colour very pale grey, lightly marked with brownish grey, overall appearance greyish (Figs 58, 59); outer mid tibial spur one-half length of inner spur; frenulum with aboutsix bristles; seventh sternite with smooth and only lightly sclerotized posterior margin,(western U.S. A.) cryptophori (p. 112) Amorophaga rosemariae sp. n. (Figs 57, 167,168, 198) ADULT (Fig. 57). $, 23, 29 mm. Vertex and frons whitish, scales greyish brown basally. Labial palpuswhitish, most scales of second segment tipped with blackish brown; third segment with elongate brown spoton outer surface. Maxillary palpus whitish, second and third segment with dark brown scales above.Antennal scape and pedicel pale buff, scape with a few dark dorsal scales; flagellum pale buff; cilia 1-0 xflagellar diameter. Thorax and tegula whitish, strongly flecked with brown. Forewing cream marked withorange-buff (particularly along veins) and dark brown; overall appearance olivaceous in fresh specimenbut rapidly becoming brownish; pattern similar to that of Morophaga choragella but disrupted bylongitudinal streaking; conspicuous dark spots at two-thirds of costa and in middle of posterior margin.Legs dull buff speckled with brown; foreleg and mid-leg strongly speckled with brown above, but all legspaler at articulations; outer mid-tibial spur 0-3 length of inner; outer proximal hind tibial spur 0-6 length ofinner spur. GENITALIA cf . Unknown. 112 GADEN S. ROBINSON GENITALIA $ (Figs 167, 168). Seventh sternite with strongly serrated and sclerotized posterior margin andwith small V-shaped medial emargination. Eighth tergite narrow, shallowly bilobed posteriorly, withabout 10 strong marginal setae and numerous smaller scattered marginal and submarginal setae; eighthsternite almost square in ventral view but extended laterally dorsad to almost meet the narrow tergite;ostium transverse, sternite transversely ridged to either side and extended caudally and inwardly to formpair of terminal lobes posterior to ostium, each bearing three strong setae and numerous smaller setae.Antrum smooth-walled to one-half length of sternite then strongly folded longitudinally anterior toinception of ductus seminalis, reaching two-thirds length of apophyses anteriores. Ductus bursae as long asantrum, very thin-walled, contiguous with corpus bursae which is elongately ovoid and thin-walled; signaabsent. DISTRIBUTION. Nepal. BIOLOGY. Unknown. The holotype was collected at mercury-vapour light at about 2100 hrs in primarymontane Himalayan oak forest (Fig. 198). The paratype was collected some 800 m lower down the sameridge, in the Schima-Castanopsis zone. MATERIAL EXAMINED. 2 ex. Holotype $, Nepal: Kathmandu district, Phulchoki, 2700 m (8800'), 28.V.1983 (Robinson et al.)(genitalia slide no. 6994; BMNH). Paratype. Nepal: 1 $ , data as holotype but 1950 m, mixed Schima-Castanopsis forest, 19. vi. 1984 (Allen)(BMNH). REMARKS. The generic placement of this species must be considered somewhat doubtful. A. rosemariaeexhibits similarities to species of the Morophaga choragella-group (wing pattern; eighth sternite extendeddorsally; antrum elongate and strongly folded) but differs sufficiently (in disruption of pattern and inlaterocaudal extension of the eighth sternite) to make doubtful its attribution to that group. In placing it inAmorophaga, considerable weight is given to the wing-pattern. Such streaking of a recognizable patterndoes not occur elsewhere in the Scardiinae and may represent a stage in the transformation of theMorophaga-type pattern to the markedly derived streaked pattern of the other three Amorophaga species.This is the only species of scardiine in which the caudal margin of the seventh sternite in the female isserrated. The remaining three species of Amorophaga (hyrcanica, cryptophori andjaponica) have a similar pale,streaked wing-pattern but that of hyrcanica is the palest and the most noticeably brown-tinted. The patternof cryptophori is distinctly grey, that ofjaponica greyish brown (although both specimens ofjaponica arefaded). The four species of Amorophaga are allopatric. A tentative phylogeny for the genus would havehyrcanica andjaponica as sister-groups (synapomorphies: pale wing-pattern, ventrocaudal hook on valva,short, stout aedeagus), cryptophori the sister-group of hyrcanica + japonica (synapomorphy: wing-pattern) and rosemariae the sister-group of these three. Amorophaga cryptophori (Clarke) comb. n. (Fig. 58)Morophaga cryptophori Clarke, 1940: 114. Holotype cf , U.S.A. (NMNH) [not examined]. ADULT (Clarke, 1940: figs 2 [head], 3-3b [venation]; Fig. 58). cf$, 23-29 mm. Vertex and frons lightgreyish brown, all scales tipped with white. Labial and maxillary palpus coloured similarly. Antennal scapeand pedicel light greyish brown, all scales tipped with white; pecten bristles darker brown, tipped withwhite; flagellum greyish brown, scales tipped with white; cilia 3 x (cf) or 1-0 x ($) flagellar diameter.Thorax and tegula as head. Forewing very pale grey; strong speckles of brownish grey forming irregularlongitudinal streaks, some darker marks at end of cell, and conspicuously barred termen and fringe.Hindwing pale grey. Legs brownish grey (but hind femur and tibia paler), paler beneath and atarticulations above; fore- and mid-tibiae banded above with white at base and middle; outer mid-tibial spur0-5 length of inner; outer proximal hind tibial spur 0-7 length of inner spur. GENITALIA cf (Clarke, 1940: figs 1-lb, 5). Saccus very broad, 1-3 x as wide as long, and shorter than valvaeor uncus; uncus lobes slender, digitiform, setose, anteriorly infolded ventrad to form pair of caudally-directed hooks resembling the two halves of a gnathos. Subscaphium slender, well-sclerotized, rod-like.Juxta elongately ovate, transversely wrinkled, rugose, moulded round anellus; transtilla not developed.Valva divided by longitudinal cleft into spatulate dorsal lobe and shorter, broader ventral lobe withirregularly dentate ventral margin and similarly dentate mediolongitudinal ridge converging with ventralmargin close to base; base of valva with broad, setose ventral lobe. Aedeagus stout, 8 x as long as broad at FUNGUS MOTHS 113 one-half, tapered and only sclerotized on dorsal surface from one-half, lacking carinae; vesica withoutcornuti. GENITALIA $ (Clarke, 1940: figs 4, 4a). Eighth tergite with strongly trilobed posterior margin, with about12 strong marginal setae and similar number of smaller setae; eighth sternite strongly sclerotized, ostium ata little beyond one-half, flanked by pair of short lobes, each with four or five strong setae and similarnumber of smaller setae ; sternite produced into large , conspicuous caudal flap to side of each lobe . Antrum1-5 x length of eighth sternite, transversely ridged close to ostium just anterior to inception of ductusseminalis at four-figths posteriorly; sides of antrum weakly folded ventrad in anterior third. Ductus bursaeas long as antrum, very thin-walled. Corpus bursae almost spherical, very thin-walled; signa absent. DISTRIBUTION. U.S.A. (Idaho, California). BIOLOGY. The type-series was reared from Cryptophorus (= Polyporus) volvatus (Clarke, 1940). Lawrence& Powell (1969: 43) recorded cryptophori from the same fungus associated with Abies and Pinus in severalscattered localities in California. They considered this to be the only Californian scardiine that ishost-specific. MATERIAL EXAMINED. 4 ex. U.S.A.: 1 cf , 3 $, California, El Dorado Co., Blodgett Forest, 13 m. E. of Georgetown, reared fromPolyporus volvatus coll. 9.vii.l967, em. 23-26. vii. 1967 (Turner} (BMNH). REMARKS. The wing-pattern of this species is similar only to those of the Old World species hyrcanica andjaponica. The male genitalia differ conspicuously in the structure of the uncus lobes which are extendedventrally and caudally to form a striking imitation of a gnathos (a structure absent in the Scardiinae). Amorophaga hyrcanica ZagulajevAmorophaga hyrcanica Zagulajev, 1968: 331, figs 1, 2. Holotype cf , U.S.S.R. (ZI) [examined]. ADULT (Zagulajev, 1973: fig. 98). $, 21, 23 mm. (For full description, see Zagulajev (1968: 331, or 1973:118).) Antenna! cilia 1-0 x (cf) flagellar diameter. Forewing with R 3 and R 4 only approximated at base,cream patterned with diffuse longitudinal streaks of light purplish brown, conspicuous dark brown costaland terminal spots, strong basal streak and more diffuse subtornal blotch at three-quarters of posteriormargin. Hindwing cream, darker marginal spots at ends of veins. Outer mid-tibial and outer proximal hindtibial spurs 0-5 length of inner spurs. GENITALIA cf (Zagulajev, 1968: fig. 2; 1973: fig. 99). Saccus broad, as wide as or wider than long, as long astegumen + uncus; uncus lobes almost fused in dorsal mid-line, inturned ventrad and strongly sclerotizedlaterally to form pair of blunt, ventrally-directed hooks. Valva with elongate, digitiform costal processextending well beyond uncus or ventral region of valva; ventral region bilobed, ventral lobe inturned,blunt, hook-like; base of valva with broad, setose membranous lobe; small, digitate process from middle ofvalva. Aedeagus about 6 x as long as broad at mid-length, as long as genital armature (excluding costalprocess of valva). GENITALIA $ . Unknown.DISTRIBUTION. U.S.S.R. (Azerbaidjan).BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype cf , U.S.S.R.: Azerbaidjan, Talysh, Astara dist., R. Lyakar, district of Mamagon Village (22km W. of Pensar), 2000 m, 10.viii.1962 (Zagulajev) (ZI). 1 cf (paratype), data as holotype but Alashya Village, ix.1962 (ZI). REMARKS. The antennal cilia of male hyrcanica are shorter than those of cryptophori or japonica (males ofrosemariae are unknown) and the genitalia are distinctive; the other two species have the costal lobe of thevalva flattened and smaller, and the uncus lobes are of different form (produced ventrally and caudally toform a passable copy of a gnathos - cryptophori - or tapered with subapical 'thorns' - japonica). See'Remarks' for A. rosemariae for further comments. 114 GADEN S. ROBINSON Amorophagajaponicasp. n. (Figs 59, 112) ADULT (Fig. 59). d", 22 mm - holotype, 23 mm - paratype. (Both specimens faded.) Vertex and frons verylight brown, scales strongly tipped with white. Labial palpus and maxillary palpus coloured similarly.Antennal scape and pedicel with scales light brown basally, otherwise whitish; flagellum whitish; cilia 2-5 xflagellar diameter. Thorax and tegula light brown, scales tipped with white. Forewing white, stronglyspeckled with light brown to form irregular longitudinal streaks and conspicuously barred termen andfringe. Hindwing light brownish grey. Legs light brown; foreleg and mid-leg darker above; all legs whitishat articulations; mid-tibia entirely whitish except for oblique brown streak on outer surface; outermid-tibial spur and outer proximal hind tibial spur 0-7 length of inner spurs. GENITALIA C? (Fig. 112). Saccus short, 1-5 x as broad as long; uncus lobes triangular, strongly sclerotized,setose, each with pair of small, subapical thorn-like projections. Subscaphium broad, tapered caudally.Juxta ill-defined, weakly sclerotized, ovate, moulded round anellus; transtilla not developed. Valva withshort, setose costal lobe (probably homologous with dorsal lobe of cryptophori and costal process ofhyrcanicd) not reaching apex of valva; ventral (conspicuous) lobe of valva tapered and slightly hookedapically , with three small subapical thorn-like processes; ventral margin with strong dorsally-directed spineat two-thirds; base of valva with broad, setose, membranous lobe. Aedeagus stout, 7 x as long as broad,with elongate membranous channel extending dorsally from apex almost to base; apical quarter with fine,spicular carinae; vesica without cornuti. GENITALIA 9- Unknown.DISTRIBUTION. Japan.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype cf , Japan: 1886 (Pryer) (genitalia slide no. 13354; BMNH).Paratype. Japan: 1 d", Tottori, Daisen, 9-11. vii. 1964 (Takahama) (coll. S. Moriuti, Osaka). REMARKS. This species is superficially similar to hyrcanica and cryptophori. It has a finer and more greyishwing-pattern then hyrcanica but, from the limited material available, is not separable by externalcharacteristics from cryptophori. The male genitalia differ in the extremely short costal process (dorsallobe) of the valva being shorter than the ventral lobe of the valva: it is longer in the other two species. Theuncus lobes are characteristic in all three species - modified to form a gnathos-like pair of processes incryptophori, square-tipped and inturned to form a pair of small hooks in hyrcanica and elongatelytriangular with subapical 'thorns' injaponica. DIATAGA Walsingham Diataga Walsingham, 1914: 374. Type-species: Diataga leptosceles Walsingham, 1914: 375, pi. 10, fig. 26,by original designation and monotypy. DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia longer than 1-5 xflagellar diameter. Scape with more than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillarypalpus 5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outermid and proximal hind tibial spurs <0-4 length of inner spurs. Forewing with Rj and R 4 stalked or veryclosely approximated at base; M 2 and A/ 3 stalked or very closely approximated at base; mottled colorationforming cryptic, coarse 'moss' pattern. Male with coremata in eighth abdominal segment; coremataassociated with elongate, rod-like apodemes at anterior corners of eighth sternite. Male genitalia withsimple uncus - a pair of setose lobes - separated from tegumen by narrow band of membrane; tegumenunbroken, completely sclerotized dorsally; valva with setose basal lobe on inner surface; apex of valvaforming ventral hook or hooks, or with spines; valvae separate, not fused together ventrally, withoutlongitudinal cleft; saccus longer than wide; juxta complex, divided medially; vesica with spicular cornuti;aedeagus smooth-surfaced, without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. Two pairs of veins fused, stalked or connate in forewing; M 2 and M 3stalked or very closely approximated at base in forewing; aedeagus bifurcated from close to base, dorsalarm with apex swollen; juxta bifid, forming pair of strongly defined lobes either side of anellus, and closelyassociated with basal setose lobes of valvae. DISTRIBUTION. Neotropical region; Nearctic region - Arizona only. FUNGUS MOTHS 115 BIOLOGY. See Diataga leptosceles Walsingham. Key to species of Diataga Males (males of direpta are unknown) 1 Uncus less than half as long as vinculum + tegumen, with pair of laterobasal spines (Fig. 120); saccus quadrangular, with concave anterior margin mercennaria (p. 1 19) - Uncus more than half as long as vinculum + tegumen, without spines; saccus triangular, with sharply convex anterior margin 2 2 Valva cleft to form hook-tipped ventral lobe (Fig. 115) frustraminis(p. 116) Valva entire 3 3 Dorsal process of aedeagus markedly shorter than ventral arm (only two thirds-length of corresponding section of ventral arm) (Fig. 123) brasiliensis(p. 1 17) - Dorsal process of aedeagus almost as long as or longer than ventral arm (more than 0-85 length of corresponding section of ventral arm) 4 4 Apex of dorsal process of aedeagus bifid (Fig. 124) compsacma (p. 118) - Apex of dorsal process of aedeagus rounded 5 5 Valva with single elongate digitate process tipped with short fine spines (Fig. 118) levidensis (p. 118)Valva with three shallow thorn-like protuberances without spines (Figs 113, 114) leptosceles(p. 115) Females (females of compsacma, brasiliensis , frustraminis and levidensis are unknown). 1 Eighth sternite explanate either side of and slightly anterior to ostium; dorsal margin of ostium with deep, U-shaped emargination (Fig. 170) mercennaria (p. 1 19) - Eighth sternite evenly tapered to ostium; dorsal margin of ostium either not emarginate or only shallowly concave 2 2 Membrane of ovipositor dorsal to ostium with strong transverse ridging; eighth tergite extending beyond level of ostium (Fig. 169) leptosceles(p. 115) - Membrane of ovipositor dorsal to ostium smooth; eighth tergite not reaching level of ostium (Fig. 171) direpta(p. 120) Diataga leptosceles Walsingham(Figs 60, 113, 114, 121, 169, 183, 185-188, 193-197) Diataga leptosceles Walsingham, 1914: 375. LECTOTYPE cf, MEXICO (BMNH), here designated[examined]. ADULT (Walsingham, 1914: pi. 10, fig. 26 (colour); Figs 60, 183). cf$ , 17-34 mm. Vertex and frons cream,streaked with brown close to eyes. Labial palpus cream flecked with light brown, whitish on inner surface,densely flecked with brown on upper surface of second segment. Maxillary palpus cream, densely fleckedwith brown above on second segment. Antennal scape and pedicel cream, densely flecked above with lightbrown; flagellum light buff; cilia 2-5 x (cf) flagellar diameter (females examined all have the antennaemissing). Thorax whitish flecked with brown; tegula brown with a few white scales. Forewing denselymottled medium and dark brown, with a few scattered pale scales; termen with black spots edged withcream between veins; costa with rectangular cream spot at one-half, and one at three-quarters, smaller andinconspicuous white spots closer to apex. Hindwing dark grey. Legs cream, strongly flecked with brownabove and at sides, more lightly below, but hind tarsus pale; outer mid-tibial and outer proximal hind tibialspur 0-25 length of inner spurs. GENITALIA d* (Figs 113, 114, 121). Saccus slender, elongate, twice as long as broad; uncus lobes slender,elongate, 0-40 length of genital armature, moderately sclerotized, setose. Subscaphium elongate, ribbon-like. Juxta with collar-shaped base, forming pair of lateral shark's fin-shaped lobes with slightly balloonedmembranous internal surface; transtilla collar-shaped, closely associated with setose and spinose lobes atbases of valvae. Valva with triangular process close to ventral margin, a further rounded or triangularprocess just dorsal to this, and a rounded or triangular process nearer to apex. Aedeagus with conspicuousmushroom-shaped dorsal arm extending well beyond ventral arm (aedeagus proper). GENITALIA (Fig. 169). Eighth tergite narrow, elongate, with pair of subapical groups of setae, eachconsisting of 4-5 large and similar number of small setae; eighth sternite forming strongly developedsterigma, ostium apical and flanked either side by pair of large setae plus 4-5 small setae; membrane of 116 GADEN S. ROBINSON ovipositor dorsal to sterigma with strong transverse wrinkles. Antrum twice length of eighth sternite,inception of ductus seminalis at two-thirds posteriorly. Ductus bursae broadening gradually into pear-shaped corpus bursae that extends to 1-25 length of apophyses anteriores; signa absent. DISTRIBUTION. (?)U.S.A. (Arizona - Lawrence & Powell, 1969); Mexico; Costa Rica; Ecuador; Trinidad;French Guiana; Brazil; Peru. BIOLOGY. Lawrence & Powell (1969: 44) record this species from Daedalea microsticta in Mexico and(identified tentatively) from Polyporus vulpinus on Populus fremontii in Arizona. Becker (pers. comrn.)has bred this species from a 'polyporus' in Costa Rica; larvae and pupae from this rearing are figured here(Figs 185-188, 193-197). For notes on larvae and pupae of this species, see 'Biology' for the Scardiinae,above. MATERIAL EXAMINED. 14 ex. Lectotype cf , Mexico: Vera Cruz, Jalapa, 1895 (Schaus) (genitalia slide no. 12409; BMNH). Mexico: 1 $?, data as lectotype but 1897. Ecuador: 1 cf, (Joannis). Trinidad: 1 cf (abdomen missing),vi.1905 (Busck). No data: 1 cf . (All BMNH; all paralectotypes.) Costa Rica: 1 cf , 1 $, Turrialba, 600 m,6.iv.l975, 5.viii.l971 (Becker} (coll. V. O. Becker, Brasilia); 2 cf , 3 pupae, 3 larvae, Turrialba, 25.V.1972[bred from a fungus] (Becker) (coll. V. O. Becker, Brasilia). French Guiana: 1 cf , Nouvelle Chantier, iv.(Le Moult) (BMNH). Brazil: 1 cf , Amazonas, Prainha, at light, 17.xii.1873 (Trail) (BMNH); 1 cf , Teffe,xii.1919 (Parish) (BMNH); 1 cf , Sta Catarina, Brusque, 6.ix.l970 (Becker) (coll. V. O. Becker, Brasilia).Peru: 1 cf , Pacaya, vi.1912 (M.) (BMNH). REMARKS. One male paralectotype of leptosceles is not of this species but is referable to frustraminis (q.v.),the only other Diataga known that is dark-winged and almost patternless. D. leptosceles may be separatedfrom frustraminis by its pale costal markings on the forewing (absent in frustraminis) and brown tegulae(whitish in frustraminis). The remaining species of Diataga are pale (brasiliensis, compsacma, levidensis,mercennarid) or have a strongly developed pattern (direpta) reminiscent of that of Morophaga choragella. The male genitalia of specimens from French Guiana and Brazil differ slightly from those of thelectotype from Mexico in that the costa of the valva is emarginate close to the apex and the two dorsalprocesses of the valva are pointed rather than rounded. The significance, if any, of this regional variationmay become clearer when further material is available. At present I consider the two genital types torepresent a single species. The dorsal process of the aedeagus, and the form of the valva, differ from thatexhibited by other Diataga species (compare illustrations). The male genitalia of leptosceles differ fromfrustraminis in lacking a ventrally hooked valva (compare illustrations) and in the form of the aedeagus;this has the dorsal process mushroom-shaped in leptosceles but spatulate and hooked in frustraminis . Theapex of the aedeagus proper (the ventral arm of the aedeagus) is obliquely truncated in leptosceles andarum flower-shaped in frustraminis. The female genitalia differ from those of the other two species of which the female is known(mercennaria and direpta) in that the dorsal margin of the ostium is shallowly concave (V-shaped inmercennaria', straight in direpta), the sides of the sterigma are not explanate as in mercennaria, the ostium isbroader than in direpta and the eighth tergite extends caudally beyond the eighth sternite (it does not reachthe tip of the sterigma in direpta). Diataga frustraminis sp. n. (Figs 61, 115, 122)[Diataga leptosceles Walsingham, 1914: 375, partim- 1 cf only. Misidentification.] ADULT (Fig. 61). cf , 21 mm. Coloration and external structure similar to leptosceles but head and palpi notso heavily marked with brown, ground-colour chalky white rather than cream; thorax and tegula chalkywhite, only very slightly flecked with pale brown; legs paler and not as heavily marked with brown as inleptosceles; forewing costa with scattered pale scales but lacking pair of elongately rectangular cream spotspresent in leptosceles. GENITALIA cf (Figs 115, 122). Saccus elongate and broad, 1-3 x as long as broad; uncus lobes elongate butonly 0-25 length of genital armature, moderately sclerotized, setose. Subscaphium ribbon-like, broadenedanteriorly. Juxta similar to that of leptosceles, apex of lobes distinctly membranous and ballooned;transtilla only weakly sclerotized, collar-shaped, inconspicuous, closely associated with spinose lobes atbases of valvae. Valva with strongly hooked ventral margin, separated from dorsal region bearing threetriangular processes by deep emargination . Aedeagus with dorsal arm spatulate , strongly curved ventrad at FUNGUS MOTHS 117 apex; ventral arm (the aedeagus proper) characteristically arum flower-shaped; base of vesica with small,sclerotized plate. GENITALIA $. Unknown.DISTRIBUTION. French Guiana; Brazil.BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Holotype cf , French Giuana: Rio Maroni, St Laurent, xi.1906 (Le Moult) (genitalia slide no. 12411;BMNH) (paralectotype of leptosceles) . Paratypes. Brazil: 1 cf, Teffe, xii.1919 (Parish) (genitalia slide no. 12415; BMNH); 1 cf, LowerAmazon, W. end of Parana de Buyassu, 15. i. 1896 (Austen) (genitalia slide no. 12414; BMNH). REMARKS. For separation of this species from the superficially similar leptosceles, see above and under'Remarks' for leptosceles. Diataga brasiliensis (Zagulajev) comb. n. (Figs 62, 116, 123)Morophaga brasiliensis Zagulajev, 1966: 642. Holotype cf , BRAZIL (MING A) [examined]. ADULT (Fig. 62). cf , 21 mm. Vertex and frons buff -cream. Labial palpus cream flecked with dark brown,third segment appearing very short as base surrounded by elongate scales of second segment. Maxillarypalpus cream with dark brown spot above on second and third segments. Antennal scape and pedicelcream, flecked with brown above; flagellum cream; cilia 4 x flagellar diameter. Thorax and tegula creamflecked with brown. Forewing cream marked with orange-brown along veins and speckled with darkbrown; however, speckling almost absent anterior to R 3 , thus forming a paler costal fascia occupyinganterior third of wing; dark quadrate costal spot at nearly two-thirds, smaller deep brown spots at apex of/?! and at apex of wing between R 2 and R 3 , and between R 4 , R 5 and M l . Hindwing brownish cream withdistinct golden tint, darker towards margin, with grey apical and subapical (costal) spots. Legs all missingexcept for foreleg (on slide) and hind tibia (glued); hind tibia cream, flecked below with dark brown, apicesof both proximal spurs and underside of larger (inner) spur dark brown; outer proximal hind tibial spur0-35 length of inner spur. GENITALIA cf (Zagulajev, 1966: fig. 5; Figs 116, 123). Saccus elongate, twice as long as broad; uncus lobesshort but infolded ventrad and extended anteriorly, 0-25 length of genital armature, quite stronglysclerotized, setose only posteriorly, anteriorly wrinkled and more strongly sclerotized. Subscaphium short,broad, inverted T-shaped. Juxta with collar-shaped base, forming pair of triangular lateral lobes; transtillacollar-shaped, closely associated with triangular setose lobes at bases of valvae. Valva almost triangular,ventral margin inturned and with shallow medial and subterminal triangular processes; middle of valvawith strong fold extending to apex where it terminates in a triangular process; swelling of apical processextended as a shallow ridge to process at middle of ventral margin. Aedeagus dorsal arm 0-65 length ofventral arm (aedeagus proper), tapered from two-thirds to apex; apex of aedeagus with shallow thorn-likecarinae; base of aedeagus with strong caudally-directed lobe. GENITALIA $ . Unknown.DISTRIBUTION. Brazil.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype cf , Brazil: Rio Grande do Sul (genitalia slide [GSR]; MING A). REMARKS. In the original description of this species, Zagulajev (1966) gives a date of capture ('5.vii.') forthe holotype. There is no date given on the data labels and the source of his information is unknown. Thegenitalia of the holotype were dissected and stored in glycerine in a vial by Zagulajev; I have mounted thegenitalia in Euparal on a slide. The external appearance of brasiliensis is distinctive; the hindwing is paler than that of leptosceles,frustraminis or direpta, more yellowish than that of mercennaria, and resembles that of compsacma orlevidensis but is distinctively gold-tinted. The forewing pattern is paler than that of leptosceles orfrustraminis and posteriorly as dark as direpta or mercennaria; however, the anterior third of the wing ispale. This division of the forewing into contrasting anterior and posterior fasciae is unknown in any other 118 GADEN S. ROBINSON scardiine. The genitalia of brasiliensis differ from those of all other Diataga in the form of the aedeagus,with its short dorsal arm, and the characteristic pattern of ridging and processes on the valva. Diataga compsacma Meyrick(Figs 63, 117,124) Diataga compsacma Meyrick, 1919: 270. Lectotype cf , GUYANA (BMNH), designated by Clarke, 1970: 47[examined]. ADULT (Clarke, 1970: pi. 23, fig. 1 ; Fig. 63). cf , 15, 16 mm. Vertex and frons cream. Labial palpus cream,flecked with dark brown on outer and upper surfaces. Maxillary palpus cream flecked with brown above onsecond segment. Antennal scape and pedicel cream with a few brown scales; flagellum very pale buff; cilia 4x flagellar diameter. Thorax and tegula cream flecked with brown. Forewing cream marked withorange-brown along veins and speckled with dark brown; dark speckling concentrated in ill-definedwedge-shaped costal spot at one-half, otherwise sparse; pair of apical dark brown spots between R 4 and R 5and R 5 and M l edged inwardly by pair of cream transverse lines. Hindwing off-white, flecked apically withbrown. Legs cream flecked with brown, foreleg and mid-leg more densely flecked, particularly above;outer mid-tibial spur 0-3 length of inner; outer proximal hind tibial spur about 0-35 length of inner spur(damaged). GENITALIA cf (Clarke, 1970: pi. 23, figs la, Ib; Figs 117, 124). Saccus elongate, about twice as long as broad(but angled in preparation); uncus lobes elongate, broadened apically, 0-25 length of genital armature,moderately sclerotized, setose. Subscaphium strongly sclerotized, broadened into inverted Y-shapeanteriorly. Juxta details not visible in preparation; transtilla, if present, very weakly sclerotized andoccluded in preparation by slightly sclerotized and strongly setose lobes at bases of valvae. Valva withventral margin turned inward and extended into a triangular spine at one-half; beyond this, a nodular,setose excrescence bearing a smaller dorsal spine; 'with wedge-shaped subapical process. Aedeagus withdorsal arm swollen distally, curved ventrad and bifurcate at apex, as long as ventral arm (aedeagus proper). GENITALIA $. Unknown.DISTRIBUTION. Guyana.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Lectotype cf , Guyana: Bartica, xii.1912 (Parish) (genitalia slide no. JFGC 6651; BMNH).Guyana: 1 cf (paralectotype), data as lectotype but i.1913. REMARKS. This species is of similar coloration to levidensis and mercennaria although the latter has a morestrongly developed forewing pattern with elements similar to those of direpta weakly expressed. D.compsacma is smaller than mercennaria although this difference may not be significant when furthermaterial is discovered (see, for example, the size-range of leptosceles) . The wing-pattern of compsacma ismore brownish than that of levidensis and the brown markings on legs and mouthparts are heavier. Themale genital armature of compsacma is, like that of levidensis, very small, only about 1-2 mm long. Bycontrast, the genital armature is more than 1 -8 mm long in mercennaria and longer in the other species. Themale genitalia of mercennaria are quite distinctive - see 'Remarks' for that species. The genital differencesbetween compsacma and levidensis are in the valvae and aedeagus. In compsacma the valva bears a medialtriangular process arising close to the ventral margin with, beyond it, a nodular setose excrescence; inlevidensis the medial triangular process is very shallow and beyond it arises an elongate process with achisel-shaped end. The dorsal arm (process) of the aedeagus is bifid in compsacma but only channelledlongitudinally at the apex in levidensis. Diataga levidensis sp. n. (Figs 64, 118, 125) ADULT (Fig. 64). cf , 16 mm. Coloration similar to compsacma, but brown speckling of labial palpus andlegs very light; second segment of maxillary palpus dark brown above; flagellum white; cilia 3 x flagellardiameter. Forewing flecked with whitish scales, these concentrated at costa to form broken pale costalstreak from one-third. Hindwing off-white, flecked with brown apically and with dark brown apical spot.Outer mid-tibial spur 0-35 length of inner; outer proximal hind tibial spur 0-25 length of inner spur. GENITALIA cf (Figs 118, 125). Saccus triangular, almost twice as long as broad; uncus lobes elongate, FUNGUS MOTHS 119 broadened apically, 0-35 length of genital armature, moderately sclerotized, setose. Subscaphium stronglysclerotized, elongate, inverted T-shaped. Juxta with only weakly sclerotized collar-shaped base, formingpair of ovate lateral lobes; transtilla collar-shaped, closely associated with setose/spinose lobes at bases ofvalvae. Valva with shallowly inturned ventral margin forming shallow triangular process at one-third; fromnear middle of valva an elongate digitate process with chisel-shaped apex bearing four or five stiff bristleson internal bevel of 'chisel'. Aedeagus dorsal arm almost as long as ventral arm (aedeagus proper),explanate at apex, surface nearest ventral arm finely scobinate; base of aedeagus with small, roundmembranous areas appearing as perforations. GENITALIA $. Unknown.DISTRIBUTION. Peru.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex. Holotype G", Peru: Iquitos, hi. 1920 (Parish} (genitalia slide no. 12416; BMNH). REMARKS. This small, pale species resembles superficially compsacma and is of similar size. The wing-pattern resembles also that of mercennaria (which is more variegated) but mercennaria is a larger species(about 20 mm wingspan as opposed to 16 mm in levidensis}; however, this difference may not be significantwhen longer series of both species are available. The forewings of compsacma are more brownish and themouthparts and legs more heavily flecked with brown than in levidensis. The male genitalia of levidensis arecharacterized by the valva bearing a single, elongate process with chisel-shaped apex and by the shortdorsal arm of the aedeagus: only brasiliensis and levidensis have the dorsal arm shorter than the ventral arm(aedeagus proper). In levidensis the ventral swelling at the base of the aedeagus is particularly pronounced. Diataga mercennaria sp. n. (Figs 65, 119, 120, 126, 170) ADULT (Fig. 65). cf $, 19-21 mm. Coloration of head and appendages, thorax and tegulae similar tocompsacma but flagellum scales off-white tipped with brown in holotype (off-white in paratypes); cilia 3 x(C?) or 0-4 x ($)flagellar diameter. Forewing cream, finely speckled with brown; speckles concentrated toform inwardly oblique dark fascia from two-thirds of costa; dark brown pair of apical spots between ^ 4 andR 5 and R 5 and M t edged inwardly with pair of fine white transverse lines. Hindwing light grey-brown withpaler speckles towards apex; dark apical spot; some diffuse, smaller, dark terminal spots between veins.Legs cream speckled with dark brown, speckles concentrated at bases of tarsi, on upper surface of foreleg(solid brown on tibia) and to form pair of ill-defined diagonal streaks on outer surface of mid-tibia; outermid-tibial spur 0-4 length of inner; outer proximal hind tibial spur 0-35 length of inner spur. GENITALIA cf (Figs 119, 120, 126). Posterior margin of eighth sternite with short digitate process either sideof mid-line. Saccus square, anterior margin concave, 1-5 x as long as broad; uncus lobes short, 0-25 lengthof genital armature, fairly strongly sclerotized, setose, with strong laterobasal spine. Subscaphium notdeveloped. Juxta narrow and ill-defined medially, forming pair of contorted, strongly sclerotized lobes;transtilla strongly sclerotized, forming pair of flattened rods overlying and closely associated withsetose/spinose lobes at bases of valvae. Valva with strongly folded ventral margin forming sclerotized lobe;small triangular process from folded surface at one-half. Aedeagus with dorsal arm slightly longer thanventral arm, apex slightly swollen and rounded; ventral arm (aedeagus proper) with membranous dorsalsurface, apex with pair of shallow, sclerotized, dorsally-directed processes each bearing an internal wart;base of vesica (? or end of ductus ejaculatorius) with pair of sclerotized plates. GENITALIA $ (Fig. 170). Eighth tergite elongate, with pair of subapical groups each comprising 3 large and3 or 4 small setae; eighth sternite forming strongly developed sterigma with explanate sides; ostium apical,flanked either side by 6 or 7 strong setae and a few smaller setae, dorsal margin with V-shaped medialemargination. Antrum twice length of eighth sternite, dorsal wall folded into pair of shallow lobes atone-half at inception of ductus seminalis. Ductus bursae broadening gradually into corpus bursae thatextends to 1-25 length of apophyses anteriores; ductus and corpus bursae strongly wrinkled (?unexpanded-virgin specimen); signa absent. DISTRIBUTION. Trinidad; French Guiana.BIOLOGY. Unknown. 120 GADEN S. ROBINSON MATERIAL EXAMINED. 3 ex. Holotype cf , French Guiana: St Jean du Maroni, vii (Le Moult) (genitalia slide no. 12407; BMNH). Paratypes. French Guiana: 1 cf, Godebert-Maroni, v (Le Moult) (BMNH). Trinidad: 1 $, Caparo(Birch) (genitalia slide no. 12410; BMNH). REMARKS. This is a large, pale species with some of the wing-pattern elements of the much darker direpta.While mercennaria is larger than known specimens of compsacma or levidensis, this size difference may notbe significant when longer series are available for comparison. The male genitalia of mercennaria are quitedistinct from those of other Diataga: the eighth sternite bears caudal processes, the saccus is truncatedanteriorly with a slightly concave anterior margin, the uncus has a pair of laterobasal spines, the transtillaforms a pair of rods, the juxta lobes are contorted and the subscaphium is not developed. The femalegenitalia are distinctive in that the sterigma is laterally explanate, strongly setose to either side of theostium, and the dorsal margin of the ostium is strongly emarginate. Diataga direpta sp. n. (Figs 66, 171) ADULT (Fig. 66). $, 20, 23 mm. Coloration of head and appendages, thorax and tegulae similar tocompsacma but head tufts with some brown scales admixed; flagellum scales off-white, some tipped withbrown; cilia 0-3 x flagellar diameter. Forewing predominantly light brown, speckled with darker brown toform ill-defined oblique basal and postmedial fasciae; some whitish scales forming medial and subterminalblotches on costa, subterminal blotch broadly U-shaped. Hindwing light grey-brown with a few paler flecksat apex. Legs cream flecked with brown; foreleg more heavily flecked above, mid-tibia with pair ofill-defined oblique brown streaks on outer surface ; outer mid-tibial spur 0-3 length of inner; outer proximalhind tibial spur 0-35 length of inner spur. GENITALIA cf . Unknown. GENITALIA $ (Fig. 171). Eighth tergite shorter than eighth sternite, with about 20 pairs of strong setaescattered along full length of tergite close to lateral margins; eighth sternite forming strongly developedsterigma bearing scattered setae of moderate size over posterior half; ostium apical, narrow, dorsal margintransverse, straight. Antrum twice length of eighth sternite, with slight longitudinal fold (?artefact),inception of ductus seminalis at two-thirds posteriorly. Ductus bursae broadening gradually into ovoid,thin-walled corpus bursae; signa absent. DISTRIBUTION. Argentina.BIOLOGY. Unknown. MATERIAL EXAMINED. 2 ex. Holotype $, Argentina: Alta Gracia, ii.1934 (C.B.) (genitalia slide no. 12408; BMNH).Paratype. 1 $ , same data as holotype (BMNH). REMARKS. This species resembles superficially Morophaga choragella (q.v.), so much so that Meyrick hadidentified the two specimens above as 'Scardia boleti' in his collection. While the wing-pattern of direpta isfar more variegated than that of either leptosceles or frustraminis , it is nevertheless a large , dark species andmore likely to be confused with one of these than with one of the smaller, paler species (brasiliensis ,compsacma, levidensis, mercennaria). The female genitalia differ from those of mercennaria and leptos-celes in that the sterigma is strongly setose in the posterior half, is not laterally explanate, and the ostialdiameter is only about 0-06 mm (about 0- 1 mm in leptosceles, 0-15 mm in mercennaria). MOROPHAGA Herrich-Schaffer Morophaga Herrich-Schaffer, 1853: (7), (22). Type-species: Euplocamus morellus Duponchel, 1838: 79, pi. 288, fig. 5, by subsequent monotypy (Herrich-Schaffer, 1854: 78).Atabyria Snellen, 1884: 164. Type-species: Atabyria bucephala Snellen, 1884: 166, pi. 9, fig. 1, by monotypy. [Synonymized by Petersen (1959: 577).]Osphretica Meyrick, 1910: 475. Type-species: Osphretica chomatias Meyrick, 1910: 475, by monotypy. [Synonymized by Gozmany (1966: 499).]Microscardia Amsel, 1952: 139. Type-species: Noctua boleti F., 1777: 282, by original designation and monotypy. [Synonymized by Petersen (1959: 577).] DIAGNOSIS. Antenna (male) lacking dorsal cilia, ventral surface without scales; cilia longer than 1-5 x FUNGUS MOTHS 121 flagellar diameter. Scape with more than 15 pecten bristles. Interocular index (male) 1-0 or less. Maxillarypalpus 5-segmented; pilifers present; second segment of labial palpus shorter than width of head. Outermid and proximal hind tibial spurs >0-4 length of inner spurs, but less in borneensis. Forewing with R 3 andR 4 stalked or approximated at base; M 2 , M 3 and CuA\ separate; coloration usually mottled, formingcryptic, coarse 'moss' pattern, but conspicuously different - bold purple-brown markings on white - inbucephala-group. Male with or without coremata in eighth abdominal segment; coremata associated withelongate, rod-like apodemes at anterior corners of eighth sternite in choragella-group (but not infasciculata) . Male genitalia with simple uncus - a pair of setose lobes - separated from tegumen by narrowband of membrane ; tegumen completely sclerotized dorsally or broken by a membranous suture line ; valvawith setose basal lobe on inner surface; apex of valva forming ventral hook or hooks, or with spines; valvaeseparate, not fused together ventrally, with deep emargination forming longitudinal cleft; cleft not present,however, in bucephala-group; saccus longer than wide; juxta simple (but complex in donodes), entire, notdivided medially; vesica with or without spicular cornuti; aedeagus with or without carinae. CONSPICUOUS AUTAPOMORPHIES. None; may be recognized, however, by combination of 'moss'-patternedforewing with R 3 and R 4 stalked (but note aberrant pattern of bucephala-group) and deeply cleft valva (notin bucephala-group). DISTRIBUTION. Western and eastern Palaearctic region; Afrotropical region; Oriental region; Australianregion. Absent from the New World. BIOLOGY. See under entries for individual species. REMARKS. Preliminary work on the Scardiinae in 1975 suggested that the type-species of Scardia was Tineamediella Hubner, 1796, a result of the designation by Walsingham (November, 1914) of 'Noctua boleti F. 'as type-species. Noctua boleti F., 1777, is not eligible for designation as type-species of Scardia Treitschkeas it is not an included species. However, Walsingham included Tinea mediella Hubner, 1796, in hissynonymy of boleti and, as mediella is one of the species included by Treitschke, its designation astype-species by Walsingham would have been valid (Int. Code zool. Norn. , Article 69(a) (v)) had it beenthe earliest designation. Tinea mediella Hubner, 1796, is currently considered to be a junior subjectivesynonym of Tinea choragella [Denis & Schiffermiiller], 1775: 137 (Charpentier, 1821: 127). This finding, which would have made Scardia the valid generic name for the species placed here inMorophaga, and Fernaldia the valid name for those in Scardia, was communicated to several workers inlitteris. Generic combinations and synonymy based on the Walsingham designation were published byDavis (1983). My colleagues I. W. B. NyeandD. S. Fletcher have recently found an earlier designation of atype-species for Scardia by Busck (April, 1914). This designation (see below) maintains the usage ofMorophaga and Scardia by authors with the exception of Davis (1983). Morophaga could well be paraphyletic with respect to Diataga and Amorophaga - see the discussionfollowing numerical analysis. The bucephala-group DIAGNOSIS. Outer and mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing pattern"conspicuous, bold purple-brown markings on a white ground-colour. Male with coremata in eighthabdominal segment; coremata without associated apodemes. Tegumen unbroken, completely sclerotizeddorsally; valva without longitudinal cleft; juxta simple, entire, not divided medially; vesica lacking spicularcornuti; aedeagus with spicular or spinose carinae. CONSPICUOUS AUTAPOMORPHIES. Wing pattern of bold purple-brown blotches on a white or off-whitebackground; valva with ventral margin extended caudally so that valva is rudder-shaped. DISTRIBUTION. Eastern Palaearctic region; Oriental region as far east as New Guinea; Afrotropical region.Absent from the western Palaearctic region and Australia. Key to species of the 6ucep/ia/a-group 1 Distal margin of large spot on posterior margin of forewing angled towards apex of wing (Fig.67) ; angle between extended ventral margin and dorsal region of valva with triangular process(Fig. 127) ; antrum less than one-half length of apophyses anteriores (Fig. 173) cremnarcha (p. 122)Distal margin of large spot on posterior margin of forewing angled towards middle of costa;angle between extended ventral margin and dorsal region of valva without triangular process;antrum at least four-fifths length of apophyses anteriores 2 122 GADEN S. ROBINSON 2 Inner surface of labial palpus cream; maxillary palpus cream with a few darker scales on second and third segments; upper surface of hindleg ochreous, like under-surface; dorsal region ofvalva rounded apically (Fig. 128); antrum + ductus bursae as long as apophyses anteriores, corpus bursae only slightly offset to the left (Fig. 174) bucephala(p. 123) Inner surface of labial palpus brown; maxillary palpus brown; upper surface of hindleg grey,paler at articuatlions; dorsal region of valva truncated apically; antrum + ductus bursaeconsiderably longer than apophyses anteriores, ductus strongly looped to the left (Figs 175,176) 3 3 Distal margin of large spot on posterior margin of forewing slightly concave or sinuate (Fig . 69) ; caudal extension of ventral region of valva rounded in cross-section, with subapical process (Fig. 129) ; microtrichia in antrum larger anteriorly than posteriorly soror(p. 124) Distal margin of purple-brown spot on posterior margin of forewing strongly 'stepped' (Fig. 70);caudal extension of ventral region of valva flattened, truncated at apex, without subapicalprocess (Fig. 130); microtrichia in antrum of uniform size vadonella(p. 124) Morophaga cremnarcha (Meyrick) comb. n.(Figs 67, 127, 173) Scardia cremnarcha Meyrick, 1932ft: 323. LECTOTYPE cf , KASHMIR (BMNH), here designated [ex-amined]. Morophaga nigrocapitella Petersen, 1959: 571, pi. 32, fig. 1; fig. 19. Holotype cf , AFGHANISTAN [SW.]:Arghandab R. (LN) [not examined]. Syn. n. ADULT (Petersen, 1959: pi. 32, fig. 1; Fig. 67). cf, 12-25 mm. $, 18-26 mm. Coloration and externalstructure similar to bucephala but cf antennal cilia 4 x flagellar diameter. Distal margin of large spot onposterior margin of forewing angled towards apex. GENITALIA cf (Petersen, 1959: fig. 19; Zagulajev, 1973: fig. 62; Fig. 127). Similar to bucephala but valvawith triangular process present in angle between caudal extension of ventral region of valva and dorsalregion; bilobate process close to costa with distal lobe enlarged and tapered; aedeagus not as slender as inbucephala, only 15 x as long as broad, apical one-sixth with about six minute thorn-like carinae. GENITALIA $ (Fig. 173). Similar to bucephala, but medial emargination of eighth sternite broader,V-shaped. Antrum short, half length of apophyses anteriores, not as thick-walled as in bucephala, innersurface evenly lined with fine microtrichia anteriorly from inception of ductus seminalis. Ductus bursaewith four or five regular transverse constrictions, only slightly offset to the left. Corpus bursae withposterior diagonal ribbing. DISTRIBUTION. Afghanistan; Kashmir; India (Uttar Pradesh, Himachal Pradesh, Bihar, Madras); Nepal. BIOLOGY. Three of Meyrick's syntypes were found on 'old tree-stumps on which Polyporus was growing,but I could find no trace of larvae, pupae or cocoons in the fungus' (Fletcher - quoted by Meyrick, 1932ft:323). The specimen from the Shevaroy Hills referred to as 'Atabyria bucephala' by Fletcher ([1921]: 189)was bred 'from cocoons about a fungus growth on a tree'. Fletcher (1933: 73), referring to this species as'Scardia bucephala', records adults (presumably the syntypes of cremnarcha) resting on rotten tree-stumpsat Gulmarg, Kashmir. MATERIAL EXAMINED. 12 ex. Lectotype Cf , Kashmir: Gulmarg, 8800', vii. 1931 (Fletcher} (BMNH). Kashmir: 1 cf , 2 $ (paralectotypes) , data as lectotype (BMNH) . India: 1 9 , Uttar Pradesh, Mukteswar,19.vii.1927; 1 $ , Himachal Pradesh, Dharmsala, 1879 (Hocking); 1 cf , Himachal Pradesh, Kangra Valley,4500', ix. 1899 (Dudgeon); 2 cf, Bihar, Pusa, 12.viii. 1909 & 30. vi. 1919 (H. L. D ., Fletcher); 1 $ with pupa,Madras, Shevaroy Hills, Yercaud, from cocoons about a fungus growth on a tree, 21. iv. 4. v. 1913 (Y. R.).Nepal: 1 $, Kathmandu District, Godaveri, 1700m, vii. 1982 (Allen). (All in BMNH.) REMARKS. The four species of the bucephala-group (bucephala, soror, vadonella and cremnarcha) areallopatric but restricted to the Old World; however, there may well be overlap of the ranges of cremnarchaand bucephala in NE. India and possibly Nepal. The wing patterns of three species are almost identical butthat of cremnarcha is different, the large purple-brown spot on the posterior margin of the forewing beingextended towards the wing-apex. The valva of cremnarcha bears a characteristic triangular flap betweenthe dorsal lobe and the caudal extension, and the aedeagus is somewhat stouter (length = 15 x width, asopposed to 20-25 x in the other species); in the female the antrum is only half the length of the apophyses FUNGUS MOTHS 123 anteriores as opposed to roughly the same length in the other species. The wing-patterns of bucephala,soror and vadonella are similar, but the distal margin of the large spot on the posterior margin of theforewing is 'stepped' in vadonella (more so in the two females examined than in the rather worn maleholotype) and in examples of bucephala from New Guinea; bucephala from elsewhere, and soror, have themargin slightly concave or sinuate. Both vadonella and soror differ from bucephala in having the whole ofthe labial palpus and the maxillary palpus brown - in bucephala the inner surface of the labial palpus iscream, as is the maxillary palpus except for some darker scaling on the second and third segments. Inbucephala the dorsal region of the valva is rounded distally but it is truncated in soror and vadonella:microtrichia are restricted to the anterior one-quarter of the antrum in bucephala but evenly scattered inthe other two species. The major difference between soror and vadonella lies in the form of the valva: thecaudal extension of the ventral margin is rounded in cross-section in soror and bears a short subapicalprocess. This process is absent in vadonella and the caudal extension of the valva is flattened and truncated.Differences in the female genitalia are minimal - the eighth sternite of soror is marginally narrower thanthat of vadonella and the microtrichia in the antrum are larger anteriorly whereas in vadonella they are ofuniform size. The bursa copulatrix is not as elongate in vadonella as in soror (but still longer than inbucephala). A tentative phylogenetic sequence for the group would have soror and vadonella as sister-groups(synapomorphies: colour of palpi, truncation of dorsal region of valva, strongly looped and elongate ductusbursae), bucephala as their sister-group (synapomorphies: forewing pattern, very slender aedeagus), andcremnarcha as the sister-group of the other three. Morophaga bucephala (Snellen)(Figs 68, 128, 174) Atabyria bucephala Snellen, 1884: 166, pi. 9, fig. 1. LECTOTYPE cf , U.S.S.R. (ZI), here designated [examined].Osphretica chomatias Meyrick, 1910: 475. Holotype cf , SARAWAK (BMNH) [examined]. [Synonymized by Bradley, 1965: 116.]Depressaria rotundata Matsumura, 1931: 1091. Lectotype $, JAPAN (EIHU), designated by Ridout, 1981: 36 [not examined]. [Synonymized by Bradley, 1965: 116.] ADULT (Snellen, 1884: pi. 9, fig. la (colour); Issiki, 1957: fig. 49 (colour); Zagulajev, 1973: pi. 2, fig. 5(colour); Fig. 68). cf $, 11-23 mm. Vertex and frons purple-brown, paler towards mouth. Labial palpusbrown on outer surface, cream on inner surface. Maxillary palpus cream, some darker scaling on secondand third segments. Antennal scape and pedicel dark brown; flagellum ochreous, some darker scalesdorsally; cilia 3 x (cf ) or 0-3 x () flagellar diameter. Thorax and tegula purple-brown anteriorly, creamposteriorly. Forewing cream with a brownish tint, patterned with bold purple-brown spots; distal margin oflarge posterior spot weakly concave or sinuate (but strongly stepped in examples from New Guinea and inone of two specimens known from Assam). Hindwing grey. Legs ochreous cream; foreleg and mid-leg darkbrown above but pale at articulations; outer mid-tibial spur 0-4 length of inner; outer proximal hind tibialspur 0-5 length of inner. GENITALIA cf (Petersen, 1959: fig. 20; Zagulajev, 1973: fig. 59; Fig. 128). Saccus elongate, as long astegumen + uncus; uncus lobes short, infolded ventrad, hardly sclerotized, setose. Subscaphium elongate,ribbon-like. Juxta weakly developed, shield-shaped; transtilla inverted U-shaped, closely associated withsetose lobes at bases of valvae. Valva with rounded ventral apex, with m-shaped bilobate process close tocosta. Aedeagus slender, 25 x as long as wide, apical fifth with about 15 minute, thorn-like carinae; vesicawithout cornuti. GENITALIA $ (Petersen, 1959: fig. 21; Zagulajev, 1973: figs 60, 61; Fig. 174). Eighth tergite slightly longerthan eighth sternite, with subapical row of about 8 strong setae and apical line of numerous smaller setae;eighth sternite only weakly sclerotized , m-shaped , posterior margin forming ventral lip of ostium . Antrumtapered anteriorly, then slightly swollen posterior to inception of ductus seminalis at level of anteriormargin of eighth sternite; anterior to inception of ductus seminalis only weakly sclerotized, but thick-walled, almost reaching apices of apophyses anteriores; with fine microtrichia on inner surface of anteriorone-quarter (and also medially in a specimen from New Guinea). Ductus bursae very short, weakly turnedto the left. Corpus bursae thin-walled, ovoid; signa absent. DISTRIBUTION. U.S.S.R. (Amur, Chabarovsk,Primorsk- Zagulajev, 1973); China (Kwangtung- Meyrick,1934; Kiangsu - Meyrick, 1935; Yunnan - Meyrick, 1938); Japan; Korea; India (Assam); Burma; Malaya;Borneo; Sulawesi; New Guinea. 124 GADEN S. ROBINSON BIOLOGY. See Zagulajev (1973: 81) for an account of the habitat and dates of appearance of this specieswhich does not appear to have been reared. MATERIAL EXAMINED. 69 ex. Lectotype cf (of Atabyria bucephala}, U.S.S.R.: Primorsk, Chabarovsk, Suifun, 14.viii.1887 (ZI).Holotype cf (of Osphretica chomatias), Sarawak: Kuching, x.1907 (Hewitt) (BMNH). 67 ex., various localities (see 'Distribution') (BMNH, ZI). REMARKS. See 'Remarks' for cremnarcha, above. The distribution of bucephala is surprisingly broad.However, only females are known from New Guinea and their conspecificity with material from theeastern U.S.S.R. is uncertain. Morophaga soror Gozmany (Figs 69, 129, 176)Morophaga soror Gozmany, 1965: 281, fig. 33. Holotype cf , CONGO (HNHM) [not examined]. ADULT (Fig. 69). cf $ , 14-23 mm. Coloration and external structure similar to bucephala but labial palpusbrown on inner surface; maxillary palpus brown; antennal cilia (cf) 2-0-2-5 x flagellar diameter; uppersurface of hindlegs greyish, paler at articulations. Forewing with distal margin of large posterior spotusually slightly concave. GENITALIA cf (Gozmany, 1965: fig. 33; Gozmany & Vari, 1973: fig. 449; Fig. 129). Similar to bucephala butvalva with dorsal region markedly truncated; caudal extension of ventral region thus proportionatelylonger, with blunt dorsally-directed subapical process; aedeagus slender, about 20 x as long as broad,apical one-quarter with about 8 minute thorn-like carinae (carinae more numerous - about 30 - in aspecimen from Uganda, Ruwenzori). GENITALIA $ (Gozmany & Vari, 1973: fig. 450; Fig, 176). Similar to bucephala but strong setae mixed withsmaller setae at posterior margin of eighth tergite. Antrum with inner surface evenly lined withmicrotrichia anteriorly from inception of ductus seminalis (but posterior microtrichia only one-third lengthof those in anterior region of antrum). Ductus bursae elongate, with regular transverse constrictions, andstrongly looped to the left. DISTRIBUTION. Sierra Leone; Ivory Coast; Ghana; Cameroon; Fernando Poo (Equatorial Guinea);Uganda; Kenya. Reliable literature records also exist for Congo, Zaire and Zambia (Gozmany & Vari,1973: 148). BIOLOGY. Unknown. MATERIAL EXAMINED. 33 ex., various localities (see 'Distribution') (BMNH). REMARKS. See 'Remarks' for bucephala, above. Morophaga vadonella (Viette)(Figs 70, 130, 175) Sporadartha [sic] vadonella Viette, 1954: 78, fig. 4. Holotype cf , MADAGASCAR (MNHN) [examined].Morophaga vadonella (Viette) Gozmany, 1969: 295. ADULT (Fig. 70). cf, 15 mm. $, 17, 19 mm. Coloration and external structure similar to bucephala but.labial palpus brown on inner surface; maxillary palpus brown; cf antennal cilia 2-5 x flagellar diameter;upper surface of hindleg greyish, paler at articulations. Distal margin of large spot on posterior margin offorewing 'stepped' or strongly concave. GENITALIA cf (Viette, 1954: fig. 4; Fig. 130). Similar to bucephala but dorsal region of valva markedlytruncated apically; caudal extension of ventral region thus proportionately longer, square-ended, withoutblunt subapical process; aedeagus slender, about 25 x as long as broad, with only one ill-defined andminute subapical thorn-like carina. GENITALIA $ (Gozmany, 1969: fig. 11; Fig. 175). Similar to bucephala but differing in the same details assoror; eighth sternite slightly broader than in soror, bursa copulatrix not as elongate, microtrichia ofantrum of even size. DISTRIBUTION. Madagascar. FUNGUS MOTHS 125 BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Holotype cf , Madagascar [NE.]: Maroantsetra, x.1952 (Vadon) (MNHN). Madagascar: 1 $, Betroka, 1955 (Diehl); 1 $, Ambinanindrano, 50 km W. of Mohanoro, i.1913(Kestell-Cornish) (BMNH). REMARKS. See 'Remarks' for bucephala, above. The more//us-group DIAGNOSIS. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with mottledcoloration forming cryptic, coarse 'moss' pattern. Male with coremata in eighth abdominal segment;coremata not associated with apodemes. Tegumen unbroken, completely sclerotized dorsally; valva withdeep emargination forming longitudinal cleft; juxta simple, entire, not divided medially; vesica lackingspicular cornuti; aedeagus smooth-surfaced, without spicular carinae. CONSPICUOUS AUTAPOMORPHIES. See species' description.DISTRIBUTION. Western Palaearctic region. Morophaga morellus (Duponchel)(Fig. 71) Euplocamus morellus Duponchel, 1838: 79, pi. 288, fig. 5. Syntypes, 1 cf , 1 ?, [FRANCE] (7MNHN) [not examined].Morophaga morella t.fungicolella Dumont, 1930: 286. Syntypes, 1 cf , 3 $ , TUNISIA (BMNH) [examined]. Syn. n. ADULT (Duponchel, 1838: pi. 288, fig. 5 (colour); Zagulajev, 1973: pi. 1, fig. 1 (colour); Fig. 71). cf$,21-28 mm. Vertex and frons greyish brown, scales tipped with white, pair of paler lateral tufts arising closeto tentorial pits. Labial palpus greyish brown, scales tipped with white, but inner surface buff; thirdsegment diffusedly banded with black basally and subapically. Maxillary palpus cream, dark scales aboveon second and third segments. Antennal scape and pedicel ochre-brown, with darker brown ventral scales;flagellum medium brown; cilia 2-5 x (cf) or 0-4 x (2) flagellar diameter. Thorax and tegula dark brownanteriorly, light grey-brown posteriorly. Forewing ground-colour light brown, strongly speckled andmottled with dark brown, dark spots concentrated medially and subterminally. Hind wing light grey-brownwith some ill-defined darker speckles towards apex. Legs buff but foreleg and mid-leg dark brown above,banded with buff at articulations; hind tibia light greyish above; outer mid-tibial spur 0-4 length of inner;outer proximal hind tibial spur 0-5 length of inner. GENITALIA cf (Petersen, 1957: fig. 244; Zagulajev, 1968: fig. 7, 1973: fig. 48). Saccus broadly triangular butas long as tegumen + uncus; uncus lobes short, square-ended, infolded both mesad and ventrad,moderately sclerotized, setose. Subscaphium ill-defined, broadening posteriorly. Juxta large, shield-shaped and conspicuously wrinkled; transtilla not developed. Valva with emargination separating ventrallobe bearing three peg-like processes from smaller dorsal lobe bearing subapical ventral flap. Aedeagusstout, S-shaped, 5 x as long as wide, without carinae; vesica lacking cornuti. GENITALIA $ (Petersen, 1957: fig. 245; Zagulajev, 1968: fig. 8, 1973: figs 49, 50). Eighth tergite as long aseighth sternite, with subapical row of about eight strong setae; eighth sternite strongly sclerotized, mediallyemarginate anteriorly, extended posteriorly into pair of digitate processes each bearing three strong setaeand overlying ostium. Antrum cylindrical, 0-25 length of apophyses anteriores. Ductus bursae short, 0-5length of antrum. Corpus bursae globular, only reaching two-thirds length of apophyses anteriores; signaabsent. DISTRIBUTION. S. France; Morocco; Tunisia; Sardinia; Sicily; Malta. Also reliably recorded in the literaturefrom Spain, Algeria (Dumont, 1930); Asia Minor (Petersen, 1957); Italy, Greece (Rhodes) (Petersen &Gaedike, 1979); U.S.S.R. (Crimea, Caucasus) (Zagulajev, 1973). BIOLOGY. This species has been bred from an excrescence on Morus (Duponchel, 1838), bred fromXanthochisma plorans on Populus and from fungus in hollow Pistacia atlantica (Dumont, 1930), bred froma polypore on Quercus suber (Martelli & Arru, [1959]) and bred from dead wood of Quercus (Staudinger,1880: 270). The biology of this species has been discussed by Dumont (1930) who described the egg, larvaand pupa. 126 GADEN S. ROBINSON MATERIAL EXAMINED. 25 ex. Syntypes (of Morophaga morella f. fungicolella Dumont), 1 cf, 3 9, Tunisia: Maknassy, bred fromfungus in Pistacia, various dates 1929-30 (Dumont) (BMNH). 21 ex., various localities (see 'Distribution') (BMNH, ZI). REMARKS. The only scardiine with a circum-Mediterranean distribution, morellus may be recognized by itswing-pattern which is brownish, rather than olivaceous as in members of the choragella-group. Thecharacteristic short, square-ended uncus lobes of the male serve to differentiate it from all otherMorophaga species. Its habitat is probably rather different from that of most other Scardiinae; whereasmost species inhabit moist forest and woodland, the distributional records of morellus suggest that it is aninhabitant of much drier environments with open sclerophyll forest. The sistrata-group DIAGNOSIS. Outer mid and proximal hind tibial spurs <0-4 length of inner spurs. Forewing with mottledcoloration forming cryptic, coarse 'moss' pattern. Male lacking coremata in eighth abdominal segment.However, coremata present (not associated with apodemes) in borneensis. Tegumen broken dorsally by atleast a membranous suture line; valva with deep emargination forming longitudinal cleft; juxta simple,entire, not divided medially; vesica with spicular cornuti; aedeagus smooth-surfaced, without spicularcarinae. CONSPICUOUS AUTAPOMORPHIES. Uncus lobes fused apically and forming a small hook; aedeagus and vesicaelongate, vesica with line of strong cornuti (but not in borneensis). DISTRIBUTION. Oriental region. Key to species of the Morophaga sistrata-group (females are known only ofsistrato) Males 1 Coremata present in eighth abdominal segment; aedeagus short, less than 1 -0 mm long, lacking cornuti (Fig. 135) borneensis(p. 126) Coremata absent from eighth abdominal segment; aedeagus elongate, more than 1-25 mm long,with many stout cornuti (Figs 136, 137) 2 2 Saccus elongate, 2-0-2-2 x as long as wide; apex of dorsal half of valva in the shape of a horse's head; ventral margin of ventral half of valva smooth (Fig. 132) sistrata(p. 127) Saccus exceptionally elongate, 2-5 x or more as long as wide; apex of dorsal half of valva evenlyrounded ; ventral margin of ventral half of valva with strong pectinifer 3 3 Pectinifer on ventral margin of valva regular; strong hook-shaped process between pectinifer and base of valva (Fig. 133) formosana(p. 128) Pectinifer on ventral margin of valva irregular; no process between pectinifer and base of valva(Fig. 134) iriomotensis(p. 128) Morophaga borneensis sp. n. (Figs 72, 131,135) ADULT (Fig. 72). cf , 13 mm. Vertex and frons cream, darker scales near eyes. Labial palpus cream fleckedwith brown, pale on inner surface, at apex of second segment and on outer surface of third segment exceptfor brownish basal and subapical spots. Maxillary palpus cream, dark brown above on second and thirdsegments. Antennal scape cream; pedicel dark brown; flagellum ochreous, darker brown above on basalthree segments; cilia 2-0-2-5 x (cf ) flagellar diameter. Thorax and tegula brown, a little paler posteriorly.Forewing (very worn) cream, marked with orange-brown on veins and strongly mottled with dark brown.Hindwing light brownish grey. Legs ochreous cream; foreleg and mid-leg dark brown above but pale atarticulations and in middle of tibiae. (Hindlegs missing.) Outer mid-tibial spur 0-3 length of inner spur. GENITALIA cf (Figs 131, 135). Small coremata present in eighth segment, without associated apodemes.Saccus only as long as tegumen -I- uncus; uncus lobes extending just beyond apices of valvae, fused caudallyand slightly hooked at apex. Subscaphium strongly developed, broad, ribbon-like. Juxta shield-shaped,only weakly sclerotized; transtilla inverted U-shaped, closely associated with small, digitate setose lobes atbases of valvae. Valva cleft (emarginate) for half its length, dorsal lobe with inwardly-directed triangularflap at apex; ventral lobe with hooked apex, pair of small, triangular sclerotizations extending dorsad fromventral margin across membranous inner face. Aedeagus short and stout, 6 x as long as broad atmid-length, without carinae; vesica without cornuti. FUNGUS MOTHS 127 GENITALIA $. Unknown.DISTRIBUTION. Borneo.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 cf. Holotype cf , Sabah: Ulu Dusun, 30 miles W. of Sandakan, lowland dipterocarp forest, 28-31.1.1976(Classey) (genitalia slide no. 13197; BMNH). REMARKS. This species is the smallest member of the sistrata-group. It may be separated from the threeothers also by the characteristic form of the valva, the presence of coremata, and the short, stubbyaedeagus which lacks cornuti. The strata-group is Oriental and the distributions of its four constituent species are allopatric. Thefemale of only sistrata is known, so the diagnostic comments here are confined to males. Externally, all fourspecies are remarkably similar although borneensis is very small (13 mm as opposed to 16-20 mm in theother species) and is matched in size only by a single dwarf (12 mm) example of sistrata. Rather smaller anddarker than members of the choragella-group, members of the sistrata-group cannot be reliably separatedexternally. Males of two species possess pectinifers on the ventrodistal margin of the valva: the pectinifer isneat and regular informosana but proximally irregular in iriomotensis . Of the other two species, borneensispossesses a pair of small coremata in the eighth segment but sistrata does not. Other differences arehighlighted below. A phylogenetic arrangement of the species would have formosana and iriomotensis assister-groups (synapomorphy: valval pectinifer), sistrata as the sister-group of this pair (synapomorphy:loss of coremata) and borneensis as the sister-group of the other three. Morophaga sistrata (Meyrick) comb. n.(Figs 73, 132, 136, 179) Scardia sistrata Meyrick, 1916: 618. LECTOTYPE cf , SRI LANKA (Ceylon) (BMNH), here designated[examined]. ADULT (Fig. 73). cf $, (12 mm dwarf), normally 16-20 mm. Vertex and frons light brown, scales tippedwith cream. Labial palpus cream flecked with brown, third segment with basal and subapical brown rings.Maxillary palpus whitish, flecked with brown on upper surface. Antennal scape cream, marked with brownabove, pecten bristles light brown tipped with cream; pedicel brown; flagellum creamy brown; cilia 3 x(cf) or 0-5 x () flagellar diameter. Thorax and tegula cream, densely flecked with brown anteriorly.Forewing cream, marked with orange-brown along veins and densely mottled with dark brown. Hindwinglight brownish grey, some darker and lighter speckling at apex. Legs ochreous cream flecked with brown;foreleg and mid-leg brown above but pale at articulations, mid-tibia with pale band at one-half, hind tarsusbrown basally; outer mid-tibial and outer proximal hind tibial spurs 0-3 length of inner spurs. GENITALIA cf (Figs 132, 136). Coremata absent. Saccus elongate, longer than tegumen + uncus; uncuslobes extending well beyond apices of valvae, fused caudally, forming slight apical hook, weaklysclerotized, setose. Subscaphium well-defined, ribbon-like. Juxta moderately sclerotized, U-shaped;transtilla inverted U-shaped, closely associated with small, digitate setose lobes at bases of valvae. Valvadeeply cleft (emarginate) , dorsal lobe with horse's head-shaped apex; ventral lobe with hooked apex andtriangular basal lobe. Aedeagus slender, elongate, 12 x as long as wide at mid-length, lacking carinae;vesica with broken line of about 80 small, strong cornuti. GENITALIA $ (Fig. 179). Eighth tergite longer than eighth sternite, with 7 or 8 strong marginal setaetowards apex and similar number of smaller setae; eighth sternite almost triangular, irregularly margined,with medial ostium flanked by pair of stout digitate processes, each with 3 strong apical setae and numeroussmaller setae. Antrum twice length of eighth sternite, not extending beyond inception of ductus seminalis.Ductus bursae relatively thick-walled, contiguous with corpus bursae, with ventral scobinate band (notmicrotrichia). Corpus bursae thin-walled, extending to almost twice length of apophyses anteriores; signaabsent (but pathological sclerotization of wall of ductus bursae present in one example examined). DISTRIBUTION. Sri Lanka; India; Thailand; Malaya; Sulawesi; Philippines. BIOLOGY. Larvae have been found feeding in decayed Polyporus (Meyrick, 1916: 618) in Ceylon and havebeen reared at Dehra Dun, India, from a fungus (Fletcher, 1933). The biology of this species, found inFomes sp. at Pusa, Bihar, India, is described in detail by Fletcher ([1921]: 186). 128 GADEN S. ROBINSON MATERIAL EXAMINED. 51 ex. Lectotype cf , Sri Lanka (Ceylon): Puttalam, x.1904 (Pole) (BMNH). Sri Lanka (Ceylon): 1 cf, 1 $, Puttalam, v., x.1904 (Pole); 3 $, Wellawaya, xi.1905 (Alston); 1 $,Peradeniya, vii.1905 (Green); 1 cf, 1 ?, 1904 (Pole) (paralectotypes; all BMNH); 18 ex., Colombo,Puttalam, Peradeniya, Nawalapitiya (BMNH). India: 2 cf, 1 $, Bihar, Pusa, 20.vii.1907, 2.ix.l908,6.xi.l908 (Fletcher) (paralectotypes; BMNH); 12 ex. , Lucknow ('ex 1. in fungus'), Pusa, Karwar, N. Arcot- Sangarambadi ('from a fungus') (BMNH). Thailand: 2 cf , without detailed locality (BMNH). Malaya: 1Cf , Prov. Wellesley, 'a fungus on coconut stump', 22.vi.1920 (Corbett) (BMNH). Sulawesi (Celebes): 1 $ ,Sangihe I. ('Sanguir'), 1892 (Doherty) (BMNH); 2 cf , Sulawesi Utara, Dumoga-Bone National Park,alluvial forest, ii.1985 (Barlow; Holloway) (BMNH). Philippines: 1 cf, 1 $, Palawan, Brookes Point,Uring Uring, 22.viii., 27. ix. 1961 (Noona Dan Exp.] (ZM); 1 cf, Mindanao, Sapamoro, Curuan dist.,16.xii.1961 (Noona Dan Exp.) (ZM). REMARKS. There are minor racial differences between the male genitalia of sistrata from Sri Lanka andexamples from the Philippines. The horse's head-shaped dorsal apex of the divided valva is broader andslightly more inturned in Philippine specimens. The triangular flap-like process on the ventral margin of thevalva is basal in Sri Lankan examples but somewhat larger and situated further distad, almost at one-halfthe length of the ventral margin of the valva, in specimens from the Philippines. Joannis (1930: 742) has recorded this species from Vietnam (Hanoi): the record is unconfirmed but is notunlikely. Morophaga formosana sp. n. (Figs 74, 133,137) ADULT (Fig. 74). cf , 18 mm; $ , 16 mm. Vertex and frons cream, tufts close to tentorial pits basally dark.Labial palpus cream flecked with brown, brown scales concentrated at base and middle of third segment.Maxillary palpus cream flecked with brown, dark brown scales above on second segment. Antennal scapecream (worn); pedicel dark brown; flagellum light ochre; cilia 3 x (cf) or 0-5 x ($) flagellar diameter.Thorax and tegula whitish, brown anteriorly. Forewing whitish (?faded) marked with orange-brown onveins and strongly mottled with dark brown. Hindwing light brownish grey. Legs cream flecked withbrown. Foreleg and mid-leg strongly marked with blackish brown above but pale at articulations and inmiddle of tibia; outer mid-tibial and outer proximal hind tibial spurs 0-3 length of inner spurs. GENITALIA cf (Figs 133, 137). Coremata absent. Saccus elongate, 1-2 x length of tegumen + uncus; uncuslobes slender, extending a little beyond apices of valvae, fused caudally, forming shallow and inconspi-cuous ventral hook, moderately sclerotized, setose. Subscaphium well-defined, ribbon-like. Juxta narrow-ly U-shaped, well-sclerotized; transtilla strongly developed, inverted U-shaped, closely associated withsmall, digitate setose lobes at bases of valvae. Valva cleft (emarginate) for one-third its length, costa withthorn-like process at three-quarters; ventral lobe with larger thorn-like process from ventral margin atone-half and with well-ordered apical pectinifer of about 14 spines. Aedeagus elongate, slender, 18 x aslong as wide at mid-length, lacking carinae; vesica with broken line of about 80 small, strong cornuti. GENITALIA 9- Unknown.DISTRIBUTION. China; Taiwan.BIOLOGY. Unknown. MATERIAL EXAMINED. 3 ex. Holotype cf , Taiwan: Taihoku, vi.1935 (Issiki) (genitalia slide no. 13119; BMNH).Paratype. Taiwan: 1 $, Tainan, 10. iv. 1906 (Wileman) (abdomen missing) (BMNH).Excluded from paratype series. China: 1 cf , Fu-chou ('Foochow'), 1935-6 (Yang) (BMNH). REMARKS. Males of this species differ from both sistrata and borneensls in possessing a conspicuouspectinifer at the apex of the ventral lobe of the valva; a pectinifer is also present in iriomotensis (see'Remarks' for that species). Like sistrata (but unlike borneensis) the aedeagus is elongate, the vesica bearsa row of about 80 small, strong cornuti and the eighth segment lacks coremata. Morophaga iriomotensis sp. n. (Fig. 134)ADULT, cf , 19 mm. Similarly patterned to sistrata and formosana (see above). FUNGUS MOTHS 129 GENITALIA cf (Fig. 134). Coremata absent. Saccus narrow, 2-5 x as long as wide, elongate, 1-3 x length oftegumen + uncus; uncus lobes slender, extending a little beyond apices of valvae, fused caudally, formingshallow and inconspicuous ventral hook, moderately sclerotized, setose. Subscaphium weakly sclerotized,ribbon-like. Juxta weakly sclerotized, heart-shaped, contiguous with M-shaped posteriorly-directedextension of medioventral margin of vinculum ; transtilla inverted U-shaped , closely associated with small ,digitate setose lobes at bases of valvae. Valva short, deeply cleft (emarginate); costa with shallow, lobe-likesubapical process and broad, thorn-shaped process at one-half; ventral lobe with strongly dentate ventralmargin, dentations regular and digitate at apex, forming a pectinifer with about 10 spines, but irregular andshallow in basal two-thirds. Aedeagus elongate, about 20 x as long as broad at mid-length, lacking carinae;vesica with line of 70-80 small, strong cornuti from base to apex. GENITALIA $. Unknown.DISTRIBUTION. Ryukyu Is.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 cf . Holotype cf , Ryukyu Is.: Iriomote, Maire-gawa, 8.ix.l965 (Arita) (genitalia slide no. 1060 [Robinson];coll. S. Moriuti, Osaka). REMARKS. Externally practically identical with sistrata and formosana, this species has, like formosana, amore slender saccus than sistrata (in which the saccus is only about twice as long as wide); unlikeformosana, the medioventral margin of the vinculum has an M-shaped process protruding between thevalvae and contiguous with the weak juxta. The valva of iriomotensis resembles that of formosana in havinga pectinifer (unlike sistrata), but the more proximal processes of the ventral margin of the valva are lessordered and there is no strong spine from the mid-point of the margin as in formosana. The costa of thevalva has a stronger thorn-like process situated further proximad than that of formosana. The c/onodes-group DIAGNOSIS. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with mottledcoloration forming cryptic, coarse 'moss' pattern. Male lacking coremata in eighth abdominal segment.Tegumen unbroken, completely sclerotized dorsally; valva with deep emargination forming longitudinalcleft; juxta complex, entire, not divided medially; vesica with spicular cornuti; aedeagus with spicular orspinose carinae. CONSPICUOUS AUTAPOMORPHIES. See species' description.DISTRIBUTION. Australian region. Morophaga clonodes (Meyrick) comb. n. (Figs 75, 138, 172) Scardia clonodes Meyrick, 1893: 523. Holotype cf , AUSTRALIA (BMNH) [examined].Scar dia porphyrea Lower, 1903: 74. Holotype $, AUSTRALIA (SAM) [examined]. Syn. n.Scardia maculosa Diakonoff, 1949: 317. Holotype $, BURU (RNH) [examined]. Syn. n. ADULT (Fig. 75). cf $ , 18-26 mm (9 from Emirau I., 15 mm). Vertex and frons cream mixed with brown.Labial palpus cream flecked with brown, brown spot on outer surface of third segment. Maxillary palpuswhitish, flecked with brown above on second and third segments. Antennal scape whitish, base browndorsally, pecten bristles cream; pedicel ochreous, dark brown dorsally; flagellum dull buff, some darkerdorsal scales towards base; cilia 1-5-2-0 x (cf)orO-4 x ($) flagellar diameter. Thorax and tegula ochreouscream, strongly speckled with brown anteriorly. Forewing cream marked with orange-brown on veins andstrongly mottled with dark brown. Hindwing ochreous grey with violet iridescence, some paler apicalmottling. Legs ochreous cream, strongly flecked with dark brown; foreleg and mid-leg dark brown abovebut pale at articulations and across middle of mid-tibia; scaling on upper surface of hind tibia not flecked,distinctively ochreous; outer mid-tibial spur 0-4 length of inner spur; outer proximal hind tibial spur 0-5length of inner spur. GENITALIA cf (Fig. 138). Coremata absent. Saccus very broad, 1-25 x as long as wide, elongate, 1-3 xlength of tegumen + uncus; uncus lobes small, infolded ventrad, moderately sclerotized, setose. Sub-scaphium not developed. Juxta a small plate lying dorsal to large m-shaped ventrocaudal extension ofvinculum which broadly separates valvae; transtilla not developed. Valva highly modified, complex, with 130 GADEN S. ROBINSON pair of strongly sclerotized, blade-shaped ventral lobes, strong mediocostal spine and elongate, setosebasicostal lobe extending almost twice length of remainder of valva; setose lobes at bases of valvaeprobably represented by extensive setose basidorsal area between basicostal lobe and anellus. Aedeagusslender, 15 x as long as broad at mid-length, with minute subapical thorn-like carinae; vesica with finespicularcornuti. GENITALIA $ (Fig. 172). Intersegmental membrane between seventh and eighth segments with pair oflateroventral pockets, each with conspicuously scobinate lining. Eighth tergite quadrate, shorter thaneighth sternite, with row of about 8 strong subterminal setae and 4 smaller terminal setae; eighth sternitevery short laterally, produced mediocaudally to form a sterigma terminating in a pair of lobes, each withthree strong terminal setae and numerous smaller setae; ostium at one-half length of sterigma. Antrumalmost one-half length of apophyses anteriores, strongly sclerotized and longitudinally ridged; inception ofductus seminalis just posterior to anterior margin of eighth sternite. Ductus bursae and corpus bursaecontiguous, pear-shaped, extremely thin- walled, not reaching apices of apophyses anteriores; signaabsent. DISTRIBUTION. (?)Moluccas (Buru); Australia; New Guinea (Emirau I., Dyaul I.); Norfolk I. BIOLOGY. Specimens in the BMNH collection have been bred from fungus on the roots of Acaciaaulacocarpa and from dead wood of Prunus persica. MATERIAL EXAMINED. 72 ex. Holotype O" (of Scardia clonodes), Australia: New South Wales, South Creek, 2.iii.l878 (Meyrick)(BMNH). Holotype $ (of Scardia porphyred) (more than 20 fragments in gelatine capsule), Australia:Queensland, Cooktown (SAM). Holotype $ (of Scardia maculosd) (head and thorax on pin; about 15fragments in two gelatine capsules), Moluccas: Buru, Station 1, 1922 (Toxopeus) (RNH). Australia: 51 ex. , Queensland, Townsville, ex fungus on roots of Acacia aulacocarpa, various emergencedates, in, iv.1901 (Dodd) (BMNH); 16 ex., Queensland, Townsville, Toowong, Dawson R., Rockhamp-ton, Herberton, Peak Downs, Huberton Hills, Kuranda, various dates (Dodd, Barnard}, including 1 cffrom Toowong, ex 'dead wood of Peach tree' (BMNH). New Guinea: 1 $, St Matthias Group, Emirau('Squally') I., viii.1923 (Eichhorn) (BMNH); 1 cf , Dyaul I., Sumuna, 7.iii.l962 (Noona Dan Exp.) (ZM).Norfolk I.: 1 $, in house J.E./Red Road, 170 m, 18.x. 1977 (Jowett & Jowett) (BMNH). REMARKS. While the general appearance of clonodes is typical of that of other Morophaga (with theexception of members of the bucephala-group), well-preserved specimens are distinctive in that thehindwing has violet iridescence and the erect scales on the upper surface of the hind tibia are brightochreous. The genitalia of both sexes are distinctive. In the female, the scobinate intersegmental pocketsare a conspicuous autapomorphy; however, shallow and smooth-surfaced pouches (no more thanaccentuated infolding of the membrane) are found also in the choragella-group. The highly modified malegenitalia are peculiar within the genus in that coremata are absent (as in three of the four species of thesistrata-group) and the subscaphium and transtilla are not developed. The form of the valva is also highlycharacteristic. Scardia maculosa Diakonoff was described from a single specimen without abdomen. In the originaldescription the specimen is said to be a male. Upon receipt of the holotype, it was found that the specimen,mounted on elder pith (and already, apparently, stuck together with glue), had broken loose in transit andwas very badly damaged. The fragments revealed, however, the typical venation of a Morophaga with /? 3and R 4 stalked at one-half, wing-pattern (rubbed) typical of the sistrata-group or clonodes, the outermid-tibial spur 0-4 the length of the inner spur and the outer proximal hind tibial spur 0-5 the length of theinner spur. The antennae are those of a female, with a cilia length of about 0-6 the flagellar diameter. Thecomparative length of the tibial spurs probably precludes this species from the sistrata-group. Diakonoff 'sreference to the iridescence of the hindwing is suggestive of affinity to clonodes. Accordingly, maculosa istentatively synonymized with clonodes although this requires a dramatic extension of the geographicalrange of the latter species. The choragella-group DIAGNOSIS. Outer mid and proximal hind tibial spurs >0-4 length of inner spurs. Forewing with mottledcoloration forming cryptic, coarse 'moss' pattern. Male with coremata in eighth abdominal segment;coremata associated with elongate, rod-like apodemes at anterior corners of eighth sternite but apodemeslacking in fasciculata. Tegumen broken dorsally by at least a membranous suture line; valva with deepemargination forming longitudinal cleft; juxta simple, entire, not divided medially; vesica with spicularcornuti; aedeagus with spicular or spinose carinae. FUNGUS MOTHS 131 CONSPICUOUS AUTAPOMORPHIES. Only group within Morophaga with rod-like apodemes from anteriorcorners of eighth segment (but see Diataga and Necroscardia); however, apodemes absent in fasciculata;valva with large, hook-shaped articulated process (absent in fasciculata but the large apical depression inthe valva suggests that the ancestor of this species may have had this process). DISTRIBUTION. Western and eastern Palaearctic region. Keys to species of the Morophaga choragella-groupMales (males of kobella are unknown) 1 Valva lacking sickle-shaped articulated process; ventral margin of valva with dense bundle of rod-like setae (Fig. 139). (Japan) fasciculata (p. 133) Valva with large sickle-shaped articulated process ; ventral apex of valva lacking rod-like setae ... 2 2 Ventral apex of valva forming a single strong hook. (Caucasus) hyrcanella(p. 132) - Ventral apex of valva rounded, with marginal line of small, shallowly dentate processes. (western Palaearctic region, east to Novosibirsk) choragella (p. 131) Females 1 Antrum elongate, extending anteriorly beyond apices of apophyses anteriores (Fig. 177) kobella(p. 133) - Antrum short , not reaching one-half length of apophyses anteriores 2 2 Eighth tergite separating dorsolateral extensions of eighth sternite; ventral margin of ostium transverse (Fig. 178) . (Japan) fasciculata (p. 133) - Eighth tergite posterior to dorsolateral extensions of eighth sternite which almost meet dorsally ; ventral margin of ostium V-shaped, deeply emarginate. (Western Palaearctic region; Cau-casus, Siberia) 3 3 Lateral sclerotization of eighth sternite extending dorsally and caudally beyond lobes that form lateral margins of ostium. (Caucasus) hyrcanella(p. 132) - Lateral sclerotization of eighth sternite not extending beyond lobes that form lateral margins of ostium. (Western Palaearctic region east to Novosibirsk) choragella (p. 131) Morophaga choragella ([Denis & Schiffermuller])(Figs 76, 189-192) Tinea choragella [Denis & Schiffermuller], 1775: 137. [AUSTRIA.] No type-material extant. Noctua boletiF., 1777: 282. [EUROPE.] No type-material extant. [Synonymized by Charpentier, 1821: 127.] Tinea fungella Thunberg, 1794: 93. [SWEDEN.] No type-material extant. [Synonymized by Karsholt & Nielsen, 1976: 21.]Tinea mediella Hiibner, 1796: 19, pi. 3, fig. 19. [EUROPE.] No type-material extant. [Synonymized by Charpentier, 1821: 127.] ADULT (Curtis, 1836: pi. 591 (colour); Duponchel, 1838: pi. 288, fig. 3 (colour); Spuler, 1910: pi. 91, fig. 31(colour); Wood, 1839: pi. 49, no. 1565 (colour); Zagulajev, 1973: fig. 51; Fig. 76). C?$, 21-30 mm. Vertexand frons cream. Labial palpus cream mixed with brown, apical segment with ill-defined subapical andbasal brown rings. Maxillary palpus cream mixed with light brown, dark brown scales above on thirdsegment. Antennal scape cream, basally mixed with brown on dorsal surface; pedicel dark brown above;flagellum ochreous, scales tipped with grey; cilia 3 x (a") or 0-7 x ($) flagellar diameter. Thorax andtegula cream, dark brown anteriorly. Forewing cream marked with orange-brown (particularly alongveins), medium brown and dark brown (overall appearance distinctly olivaceous, particularly in freshspecimens). Hindwing grey, paler flecks at apex; bases of fringe scales paler. Legs cream; foreleg andmid-leg strongly marked with dark brown above in basal half of each segment; hindleg similarly markedwith light grey; outer mid-tibial spur 0-5 length of inner; outer proximal hind tibial spur 0-7 length of inner. GENITALIA cf (Petersen, 1957: fig. 240; Zagulajev, 1968: fig. 9, 1973: fig. 52). Coremata present in eighthabdominal segment, associated with elongate, rod-like apodemes from eighth sternite. Saccus 0-6 length oftegumen + uncus; uncus lobes elongate, digitiform, moderately sclerotized, setose. Subscaphium ill-defined , ribbon-like . Juxta plate-shaped , contiguous with vinculum ; transtilla inverted elongate U-shaped ,closely associated with setose lobes at bases of valvae. Valva divided by deep emargination into dorsal andventral lobes; ventral lobe rounded, ventral margin with row of shallowly dentate processes; dorsal lobe 132 GADEN S. ROBINSON with deep recess accommodating mobile and strongly sclerotized hook-shaped process. Aedeagus curved,9 x as long as broad, apex tapered gently, with fine spicular carinae; vesica with fine spicular cornuti. GENITALIA $ (Petersen, 1957: fig. 241; Zagulajev, 1968: fig. 10, 1973: fig. 53). Eighth tergite displacedcaudally, shield-shaped, with slight posteromedial emargination; caudal margin with about 10 strong setae.Eighth sternite broad, lateral margins extended dorsally to almost meet anterior to the displaced tergite;ventrally a broad plate, extended caudally into pair of broad lobes with narrow emargination betweenthem, overlying ostium. Antrum funnel-shaped and smooth-walled to inception of ductus seminalis at 0-6ventral length of eighth sternite anteriorly, then cylindrical with longitudinal ribbing, extending 0-4 lengthof apophyses anteriores. Ductus bursae short, 0-4 length of antrum. Corpus bursae very thin-walled,ovoid, extending slightly beyond apices of apophyses anteriores; signa absent. DISTRIBUTION. Probably all European countries north of the Mediterranean, including Scandinavia andFinland; Bulgaria and Rumania; U.S.S.R. east to Novosibirsk Province (Zagulajev, 1973); Asia Minor.The identity of the specimen recorded from China (Yunnan) by Meyrick (1938) has not been confirmed. BIOLOGY. Described by a number of European authors - see Zagulajev (1973) for a bibliography. The larvahas been described in detail by Hinton (1956: 258, figs 1-14). This is a widespread and common species thathas been bred from many species of bracket-fungi and from dead wood permeated by fungal hyphae. MATERIAL EXAMINED. 346 ex., larvae and pupae, from various localities (see 'Distribution') (BMNH, ZI). REMARKS. Males of choragella and hyrcanella may be recognized by the conspicuous articulated hook set inthe dorsal lobe of the valva; however, this hook is absent infasciculata although the depression in the valvathat accommodates the hook in the other species is present. The male ofkobella is unknown. M. choragellais smaller than hyrcanella (which is restricted to the Caucasus) and the ventral lobe of the valva (formedinto a hook in hyrcanella) has a marginal row of shallowly dentate processes. Females are characterized bythe short antrum and dorsally extended eighth sternite of which the ventral caudally-directed lobes are theposteriormost part. Records of this species from Japan (e.g. Inoue, 1954: 17) are erroneous and may refer instead to eitherkobella (of which I have seen only a single female, in BMNH) or fasciculata or, more improbably, toScardla amurensis (Zagulajev) (q. v.). Morophaga hyrcanella Zagulajev(Fig. 77) Morophaga hyrcanella Zagulajev, 1966: 639, fig. 3. Holotype $, U.S.S.R. (ZI) [examined].Morophaga talyshensis Zagulajev, 1966: 641, fig. 4. Holotype cf, U.S.S.R. (ZI) [examined]. [Synony-mized by Zagulajev, 1973: 76.] ADULT (Zagulajev, 1973: pi. 1, fig. 2 (colour); Fig. 77). cf, 28 mm; $, 32 mm. Similarly patterned tochoragella but frons and vertex more brownish; antennal cilia 4-0 x (cf) or 1-0 x (9) flagellar diameter;thorax and tegula brown; forewing with dark brown scaling more extensive, forming a V-shaped medialfascia; hindwing grey-brown; legs with dark markings more intense and extensive. GENITALIA cf (Zagulajev, 1966: fig. 4; 1973: fig. 56). Similar to choragella but ventral lobe of valva withhook-shaped ventral extension; aedeagus not as strongly curved. GENITALIA $ (Zagulajev, 1966: fig. 3; 1968: fig. 12; 1973: figs 19b, 57). Similar to choragella butsclerotization of eighth sternite extended caudally beyond lobes that form lateral margins of ostium. DISTRIBUTION. U.S.S.R. (Transcaucasus).BIOLOGY. Described by Zagulajev (1966; 1973: 78). MATERIAL EXAMINED. 45 ex., pupae. Holotype $ (of Morophaga hyrcanella), U.S.S.R.: Transcaucasia, Talysh, Lenkoran dist., Sarakh,19.vii.1964 (Zagulajev) (ZI). Holotype cf (of Morophaga talyshensis), U.S.S.R.: Transcaucasia, Talysh,R. Lyakar, nr Mamagon (22 km W. of Pensar), Astara dist., 2100 m, 10.viii.1962 (Zagulajev) (ZI). U.S.S.R.: 1 Cf, 1 $, 1 pupa, Talysh, L. Girkanskiy, 20. ix. 1967 (Zagulajev) (BMNH). Also a further 41ex., 30 pupae, various Transcaucasian localities (see Zagulajev, 1973: 78) (ZI). REMARKS. More strongly marked than choragella, hyrcanella is also distinguished externally by its largersize and more elongate antennal cilia in both sexes. Differences in genitalia are outlined above. FUNGUS MOTHS 133 Morophaga fasciculata sp. n. (Figs 78, 139, 178) ADULT (Fig. 78). cf, 15-18 mm; $, 21 mm. Vertex and frons cream. Labial palpus with apical segmentuniformly pale , outer surface of second segment evenly flecked with brown . Maxillary palpus cream , somedarker scaling above on second segment. Antennal scape whitish; pedicel brown above; flagellumochreous; cilia3 x (cf)orO-5 x ( ^) flagellar diameter. Thorax and tegula cream marked with brown, darkbrown anteriorly. Forewing cream, patterned with brown (specimens worn and slightly faded), patternsimilar to kobella and to species of the s /strata-group. Hindwing grey. Legs ochreous cream; foreleg andmid-leg brownish above, paler at articulations, mid-tibia with cream medial band; outer mid-tibial spur 0-4length of inner; outer proximal hind tibial spur 0-5 length of inner. GENITALIA cf (Fig. 139). Eighth segment with small coremata but no associated apodemes; eighth sternitewith broadly U-shaped posterior emargination. Saccus broad, U-shaped, as long as tegumen + uncus;uncus lobes shorter than saccus, simple, only moderately sclerotized, setose. Subscaphium well-sclerotized, rod-like. Juxta small, quadrate, weakly sclerotized; transtilla not developed. Valva withbroad, deep emargination; with dense dorsoventral band of slender, flattened scales on outer surface;separate tuft of close-set similar scales on ventral margin; apices of ventral and dorsal lobes serrate, ashallowly dentate ridge running anteriorly down internal face of valva; line of 5 or 6 peg-like spines at apexof costa, smaller peg-like spines on internal face of dorsal lobe. Membranous lobe at base of valva elongate,densely hirsute. Aedeagus short, 7 x as long as broad, with three lobate anteriorly-directed carinae; vesicawith minute, inconspicuous spicular cornuti. GENITALIA $ (Fig. 178). Eighth tergite slightly longer than eighth sternite; eighth sternite short, withlateral membranous pouches in intersegmental membrane, caudal margin produced into pair of shortlateral processes either side of ostium, each process with about 6 strong apical setae. Antrum broad,thick-walled, almost twice length of eighth sternite, with ill-defined longitudinal ridging. Ductus bursae +corpus bursae thin-walled, contiguous, elongately pear-shaped. DISTRIBUTION. Japan. BIOLOGY. The type-species was bred from the fungus Trametes kusanoana Imazeki, adults emerging in lateJune and early July. MATERIAL EXAMINED. 4 ex. Holotype cf, Japan: Honshu, Nara, bred from Trametes kusanoana Imazeki, coll. 29. vi. 1965, em.4.vii.l965 (Arltd) (genitalia slide no. 1048 [Robinson]; coll. S. Moriuti, Osaka). Paratypes. 2 cf , 1 9,dataasholotypebutem. 2.vii. and 30. vi. 1965 (genitalia slide no. 1049 [Robinson];coll. S. Moriuti, Osaka; BMNH). REMARKS. Similarities between the male genitalia of fasciculata, choragella and hyrcanella are marked, butfasciculata is distinctive in lacking the strong, articulated hook-like process set in the valva of the last twospecies. The deep emargination of the valva that accommodates the hook in the other two species is alsopresent in fasciculata. Neither choragella nor hyrcanella possess the distinctive bundle of close-set flattenedscales on the ventral margin of the valva that is present in fasciculata. Females of fasciculata are very similarexternally to kobella. Separation of the two species, while simple enough using genitalic characters (theantrum of kobella is longer than the apophyses anteriores, but considerably shorter in fasciculata), is unsafeusing external characteristics only, to judge from the material presently available. Comparison of freshmaterial of the two species may, however, reveal perfectly good pattern differences. Morophaga kobella sp. n. (Figs 79, 177) ADULT (Fig. 79). $ , 23 mm. (Specimen slightly faded and badly discoloured by fine, dark dust.) Vertex andfrons cream. Labial palpus cream flecked with dark brown, but whitish on inner surface. Maxillary palpuscream flecked with brown; dark brown scales above on second segment. Antennal scape cream fleckedwith brown; pedicel similar; flagellum medium brown; cilia 0-5 x flagellar diameter. Thorax and tegulacream flecked with brown, dark brown anteriorly. Forewing cream marked with orange-brown, mediumbrown and dark brown, pattern similar to that of choragella but dark costal markings denser, better-defined, antemedial costal spot not as elongate. Hindwing charcoal grey. Legs cream flecked with brown;foreleg and mid-leg strongly marked with dark brown above but cream at articulations; mid-tibia with 134 GADEN S. ROBINSON cream medial band; hind spurs strongly marked with dark brown; outer mid-tibial spur 0-4 length of inner;outer proximal hind tibial spur 0-5 length of inner. GENITALIA cf . Unknown. GENITALIA $ (Fig. 177). Eighth tergite broad, shield-shaped, slightly longer than eighth sternite, withsubapical row of 4 strong setae surrounded by numerous scattered smaller setae. Eighth sternite broad,strongly sclerotized, with rectangular medial emargination posteriorly; lobes either side of emarginationeach with 3 aprcal setae; anteriorly, surface of sternite strongly ridged and folded to form strong ventralkeel. Antrum smooth-surfaced posteriorly, further anteriorly thick-walled with strong longitudinal ridgeson inner surface, extending beyond apices of apophyses anteriores. Ductus bursae short, convoluted,offset to the left. Corpus bursae elongately ovoid; signa absent. DISTRIBUTION. Japan.BIOLOGY. Unknown. MATERIAL EXAMINED. 1 ex.Holotype $, Japan: [Honshu], Kobe, 6.vii.l927 (Lewis) (genitalia slide no. 13116; BMNH). REMARKS. The external apperance of kobella is similar to that of members of the choragella-group(choragella,fasciculata) and the sistrata-group (sistrata,formosana, iriomotensis) . The bursa copulatrix isas elongate as in members of the sistrata-group but the antrum is internally ridged, as in the choragella-group. The tibial spurs are of a length comparable with that of members of the choragella-group rather thanthe sistrata-group. However, the present placement of kobella should be considered provisional until themale is discovered. This species may be separated from all other Morophaga species by its enormouslyelongate antrum. Taxa incertae sedisLEPTOZANCLA Meyrick Leptozancla Meyrick, 1920: 107. Type-species: Leptozancla talaroscia Meyrick, 1920: 108, by originaldesignation and monotypy. This genus was redescribed by Gozmany & Vari (1973) and by Robinson (1976a). The single species hereincluded, L. talaroscia Meyrick, was described by Meyrick (1920: 107) from two males from Mt Kenya,both in poor condition. No further specimens of Leptozancla have been identified since its originaldescription. In the absence of further material, the taxonomic position of Leptozancla must remain indoubt. However, it is probably a scardiine. Gozmany & Vari (1973) and Robinson (1976a) treatedPhilagrias (see below) as a synonym of Leptozancla. However, in the light of the morphological differencesbetween other scardiine genera this synonymy is not appropriate. L. talaroscia has densely ciliate antennae, the cilia as long as the flagellar diameter. The genitalia arecharacterized by the valvae having been apparently entirely supplanted by the juxta as no typical valvalapodeme is evident: the juxta (if, indeed, the structure is the juxta) forms a ventral complex resembling apair of entirely fused valvae. The transtilla (and, possibly, remnants of the valval apodemes) forms astrongly sclerotized complex with four elongate and posteriorly-directed spines (labides) (Robinson,1976a: fig. 65). PHILAGRIAS Meyrick gen. rev.Philagrias Meyrick, 1932a: 119. Type-species: Philagrias zelotica Meyrick, 1932a: 119, by monotypy. The single species included here, P. zelotica, has been redescribed by Gozmany & Vari (1973) and byRobinson (1976a). It was described from a single male example in poor condition from the highlands ofEthiopia. No further specimens of Philagrias have been identified since its original description. In theabsence of further material, its taxonomic position must remain in doubt. However, it is probably ascardiine. Philagrias is distinguished by the antennae having very short cilia (0-5 x the flagellar diameter) which liealmost parallel to the axis of the flagellum. Previous authors have recorded that Philagrias lacks antennalcilia. The 'valva' bears extraordinarily large and complicated processes (?labides) arising from themodified valval apodemes (Robinson, 1976a: fig. 66). It is not certain whether the valva is present andforms part of a valve-juxta complex or whether it has been supplanted entirely by the juxta. In previouspapers (1976a; 1981) I have assumed the structure to be the juxta on the grounds that no recognizablyfunctional remnant of the valval apodeme remains. f FUNGUS MOTHS 135 Scardia tholerodes MeyrickScardia tholerodes Meyrick, 1894: 27. Syntypes, 3 ex., BURMA (7BMNH) [not found, not examined]. Other specimens collected at Koni in the Shan States by Manders and described by Meyrick in the samepaper as tholerodes are in the BMNH collection (ex Meyrick collection). No specimens of tholerodes havebeen found, however. From Meyrick's original description, this taxon may be a member of the Morophagasistrata-group. Scardia pharetrodes MeyrickScardia pharetrodes Meyrick, 1934: 42. Holotype $, CHINA (7MNHU) [not found, not examined]. The whereabouts of the holotype of this species are in doubt: according to the original description it shouldbe in MNHU, Berlin. However, it has not been mentioned by Petersen who has had access to the Berlincollections. Neither has it turned up in the Caradja collection of MINGA, Bucharest, despite a search onmy behalf by Dr A. Popescu-Gorj. The possible affinities of this taxon are not evident from Meyrick'soriginal description. Scardia isthmiella BusckScardia isthmiella Busck, 1914: 65. Holotype $, PANAMA (NMNH) [examined]. The holotype of this species is in poor condition. It resembles superficially a Daviscardia species but thepale scaling of the terminal fascia does not extend along the dorsum. Davis (in prep.) will publish adescription and illustration of it. The genitalia are nondescript, the eighth sternite hardly sclerotized, theventral margin of the ostium lined with small setae and with a doubly crescentic outline (very shallowlym-shaped). The ductus bursae has irregular transverse constrictions in its posterior two-thirds and extendsto two-thirds the length of the apophyses anteriores. The corpus bursae is small and ovate and lacks signa. 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New species of Pyrales and Microlepidoptera from the Deutsches Entomologisches Institut.Arbeiten uber morphologische und taxonomische Entomologie aus Berlin-Dahlem 3: 94-109. 1937-1938. Microlepidoptera excl. Pyralidae. Pp. 169-182, 1-29. In Caradja, A. & Meyrick, E., Materialien zu einer Mikrolepidopterenfauna des Yulingshanmassivs (Provinz Yunnan). DeutscheEntomologische Zeitschrift Iris 51: 137-182 (1937); 52: 1-29 (1938). Mitterberger, K. 1910. Beitrag zur Biologic von Scardia boletella F. Zeitschrift fur wissenschaftlicheInsektenbiologie 6: 171-173. - 1911. Beitrag zur Kenntnis der Lebenweise der Raupe von Scardia boletella F. EntomologischesJahrbuch 20: 126-128. Moriuti, S. 1976. [Morophagoides ussuriensis (Car.) (Lep.: Tineidae), a pest of the shiitake fungus,Lentinus edodes (Berkeley) Singer, in Japan.] [In Japanese.] Forest Pests 25: 87-92, figs 1-21. - 1982. Tineidae. In Inoue, H. et al., Moths of Japan. 1: 966 pp.; 2: 552 pp., 392 pis. Tokyo.Petersen, G. 1957-1958. Die Genitalien der palaarktischen Tineiden. Beitrage zur Entomologie 7: 55-176, 338-379, 557-595; 8: 111-118, 398-430. - 1959. Tineiden aus Afghanistan mit einer Revision der palaarktischen Scardiinen. Beitrage zurEntomologie 9: 558-579, figs 1-27, pi. 32. - 1960. Zwei neue palaarktische Tineiden aus dem Iran (Lepidoptera). Stuttgarter Beitrage zurNaturkunde 34: 1-3, figs 1-2. 1965. Beitrag zur Kenntnis der Tineiden der Tschechoslowakei. Acta Faunistica Entomologica MuseiNationalis Prague 11: 165-194. - 1968. Beitrag zur Kenntnis der Tineiden Westdeutschlands. Acta Faunistica Entomologica MuseiNationalis Prague 13: 87-107. 1969. Beitrage zur Insekten-Fauna der DDR: Lepidoptera - Tineidae. Beitrage zur Entomologie 19: 311-388, figs 1-205, col. pi. figs 1-44.Petersen, G. & Gaedike, R. 1979. Beitrag zur Kenntnis der Tineiden-Fauna des Mittelmeerraumes. Beitrage zur Entomologie 29: 383-412, figs 1-29.Powell, J. A. [1968]. Taxonomic status and descriptions of some fungus feeding Tineidae. Pan-Pacific Entomologist 43: 292-307.Rawlins, J. E. 1984. Mycophagy in Lepidoptera. Pp. 382-423. In Wheeler, Q. & Blackwell, M. (eds.), Fungus-Insect Relationships, xiii + 514 pp., Columbia University Press, New York.Rebel, H. 1901. In Staudinger, O. & Rebel, H., Catalog der Lepidopteren des palaearctischen Faunenge- bietes. 2. 368 pp., Berlin.Ridout, B. V. 1981. Species described within the genus Depressaria by Matsumura (Lepidoptera). Insecta Matsumurana 24: 29-47, figs 1-22.Robinson, G. S. 1975. Macrolepidoptera of Fiji and Rotuma: a taxonomic and biogeographic study, vi + 362 pp., maps, 10 pis, 530 figs. Faringdon.1976a. A taxonomic revision of the Tinissinae of the world. Bulletin of the British Museum (Natural History) (Entomology) 32: 253-300, 16 pis, 10 figs. 19766. The preparation of slides of lepidoptera genitalia with special reference to the Microlepidop- tera. Entomologist's Gazette 27: 127-132, 2 figs. FUNGUS MOTHS 139 - 1981. Remarks on the classification of the fungivorous Tineidae with special reference to theTinissinae (Lepidoptera). Entomologica Scandinavica 12: 363-380, figs 1-6. 1984. Microlepidoptera in Brunei. The fourth UluTemburong Expedition. Brunei Museum Journal 5 (4): 146-177, figs 1-3, col. pis 1-6.Sneath, P. H. & Sokal, R. R. 1973. Numerical taxonomy, xv + 573 pp. San Francisco.Snellen, P. C. T. 1884. Nieuwe of weinig bekende Microlepidoptera van Noord-Azie. Tijdschrift voor EntomologieTJ: 151-196.Spuler, A. 1903-1910. Die Schmetterlinge Europas. (3rd edn of Hofmann, E. Die Gross-Schmetterlinge Europas.)2. [vi] + 523 pp., 238 figs. Stuttgart.Staudinger, O. 1880. Lepidopteren-Fauna Kleinasien's. Horae Societatis entomologicae rossicae 15: 159-435. Thunberg, C. P. 1794. D.D. Dissertatio entomologica sistens Insecta svecica 7: 83-98, 1 pi. Upsaliae.Treitschke, F. 1830. Die Schmetterlinge von Europa. 8: 312 pp. Leipzig.Viette, P. E. L. 1954. Description de nouveaux Tineides Malgaches. Memoires de I'Institut Scientifique de Madagascar (E)5: 1-37.Walker, F. 1864. List of the specimens of the lepidopterous insects in the collections of the British Museum. 29. Tineites. Pp. iv + 564-835. London.Walsingham, T. de G., Lord 1882. Notes on Tineidae of North America. Transactions of the American Entomological Society 10: 165-204. 1897. Revision of the West-Indian Micro-Lepidoptera, with descriptions of new species. Proceedings of the Zoological Society of London 1897: 54-183. 1914. Family 20. Tineidae. Pp. 344-375. In Biologia Centrali- Americana. Insecta. Lepidoptera Heterocera. 4. Wood, W. 1839. Index Entomologicus; or a complete illustrated catalogue consisting of 1944 figures of thelepidopterous insects of Great Britain, xii + 253 pp. London. Zagulajev, A. K. 1965. [New species of the genus Scardia.} [In Russian.] Entomologicheskoe O bo z rente 44:411-413, figs 1-3. [Translation in Entomological Review, Washington 44: 234-235.]1966. [The subfamily Scardiinae (Lep. , Tineidae) and its new species.] [In Russian.] Entomologisches-koe Obozrenie 45: 634-644, figs 1-5. [Translation in Entomological Review, Washington 45: 359-364.]- 1968. [The new and little-known species of Tineidae (Lepidoptera) in the Caucasian fauna.] [InRussian.] Trudy Vsesoyuznogo Entomologischeskogo Obshchestva 52: 326-366, figs 1-34. 1973. [Tineidae; part 4 - subfamily Scardiinae.] [In Russian.] Fauna SSSR 104: 1-126, 99 figs, 2 col. pis. Zeller, P. C. 1846. Euplocamus boleti und Eupl. tessulatellus. StettinerentomologischeZeitungl: 178-182.1863. Zwolf amerikanische Nachtfalter. Stettiner entomologische Zeitung 24: 136-155, pi. 2. 140 GADEN S. ROBINSON 16 Figs 9-16 Morophagoides species. 9, iranensis, cf; 10, ussuriensis, cf; 11, moriutii, cf paratype; 12,berkeleyella, cf (Burney Mt.); 13, burkerella, cf; 14, montium, $ holotype; 15, pythium, $ holotype;16, nimbiferum, 9 holotype. FUNGUS MOTHS 141 17 19 24 Figs 17-24 17, Morophagoides iulina, cf holotype. 18, 19, Montescardia species. 18, tessulatellus, $ ; 19,fuscofasciella, 9 (North Carolina). 20, Bythocrates drosocycla, 9- 21-24, Daviscardia species. 21,coloradella, $; 22, radulella, cf holotype; 23, bimendella, cf lectotype; 24, beckeri, cf holotype. 142 GADEN S. ROBINSON 26 27 28 29 ^ 31 30 32 Figs 25-32 Daviscardia species. 25, beckeri, $ paratype; 26, luctuosa, cf lectotype; 27, mackiei, cfholotype; 28, mackiei, cf paratype; 29, mackiei, $; 30, bicolorella, cf paratype; 31, lupulella, $holotype; 32, hypocritella, 9 holotype. FUNGUS MOTHS 143 34 35 36 40 Figs 33-40 33-39, Scardia species. 33, anatomella, $ lectotype; 34, anatomella, cf (Venezuela); 35,assamensis, cf holotype; 36, amurensis, $; 37, alleni, $ holotype; 38, boletella, $; 39, caucasica, $paratype. 40, Perilicmetis diplaca, cf paratype. 144 GADEN S. ROBINSON i*CJf 3 S 45 46 47 48 Figs 41^48 41, 42, Moscardia species. 41, renitens, cf holotype; 42, varna, cf holotype. 43, Gentingiahollo way i, cf holotype. 44, Semeoloncha penicillata, cf holotype. 45-47, Cranaodes species. 45,stereopa, $; 46, oroya, cf holotype; 47, sequestrata, $. 48, Pectiniscardia prostylias, cf holotype. FUNGUS MOTHS 145 I *m -JP 49 61 50 52 56 Figs 49-56 49, Hormantris astragalopa, cf holotype. 50, Cnismorectis choritica, cf. 51, Afroscardiacapnochalca, cf holotype. 52, Scardiella approximated, $ . 53, 54, Necroscardia species. 53,funeratella,Cf lectotype; 54, morticina, cf holotype. 55, 56, Miniscardia species. 55, minimella, cf (Costa Rica); 56,minimella, 9 (Costa Rica). 146 GADEN S. ROBINSON 58 61 62 63 64 Figs 57-64 57-59 , Amorophaga species . 57 , rosemariae, $ holotype ; 58 , cryptophori, cf ; 59 , japonica, tfholotype. 60-64, Diataga species. 60, leptosceles, <3 lectotype; 61, frustraminis, <3 paratype; 62,brasiliensis , cf holotype; 63, compsacma, C? paralectotype; 64, levidensis, cf holotype. FUNGUS MOTHS 147 66 Figs 65-72 65, 66, Diataga species. 65, mercennaria, O* holotype; 66, direpta, $ paratype. 67-72,Morophaga species. 67, cremnarcha, 0" lectotype; 68, bucephala, $ (Japan); 69, soror, cf (SierraLeone); 70, vadonella, $?; ll,morellus, 9 (syntype offungicolella); 72, borneensis, cf holotype. 148 GADEN S. ROBINSON 78 , m mm 79 Figs 73-79 Morophaga species. 73, sistrata, cf lectotype; 74,formosana, cT holotype; 75, donodes,76, choragella, $; 77, hyrcanella, tf;l%,fasciculata, cf holotype; 79, kobella, 9 holotype. FUNGUS MOTHS 149 Figs 80-83 Male genitalia of Morophagoides species. 80, ussuriensis; 81, moriutii, paratype (Japan); 82,moriutii (Taiwan); 83, berkeleyella-uncus. Scale line = 1 mm. 150 GADEN S. ROBINSON 85 88 Figs 84-88 Male genitalia. 84, 85, Morophagoides species. 84, burkerella (note reduced scale); 85, iulina,holotype. 86-88, Daviscardia coloradella. 86, uncus and tegumen; 87, valvae; 88, aedeagus. Scale line =1 mm. FUNGUS MOTHS 151 Figs 89-92 Male genitalia of Daviscardia species (fused valvae detached). 89, luctuosa, lectotype; 90,bimendella, lectotype; 91, raduldla, holotype; 92, beckeri, holotype. Scale line = 1 mm. 152 GADEN S. ROBINSON Figs 93-95 Male genitalia of Daviscardia species (fused valvae detached). 93, mackiei, paratype; 94,bicolorella, holotype; 95, unnamed species (Mexico). Scale line = 1 mm. FUNGUS MOTHS 153 96 100 Figs 96-100 Male genitalia - dorsal view of uncus lobes and tegumen of Scardia species. 96, anatomella;97, assamensis, holotype; 98, amurensis; 99, boletella; 100, caucasica (after Zagulajev, 1973). Scale line= 1 mm. 154 GADEN S. ROBINSON Figs 101, 102 Malegenitalia. Wl,Scardiaassamensis, holotype (f used valvae detached); 102, Perilicmetisdiplaca. Scale line = 1 mm. FUNGUS MOTHS 155 103 104 Figs 103, 104 Male genitalia. 103, Gentingia hollowayi, holotype (fused valvae detached); 104, Moscar-dia varna, holotype. Scale line = 1 mm. 156 GADEN S. ROBINSON Figs 105, 106 Male genitalia. 105, Cnismorectis choritica; 106, Cranaodes oroya, holotype. Scale line = 1mm. FUNGUS MOTHS 157 Figs 107-109 Male genitalia. 107, Scardiella approximated. 108, 109, Necroscardia species. 108,funeratella, lectotype; 109, morticina, holotype. Scale line = 1 mm. 158 GADEN S. ROBINSON Figs 110-112 Male genitalia. 110, Necroscardia funeratella, lectotype - modified eighth abdominalsegment; 111, Miniscardia minimella (Costa Rica); 112, Amorophaga japonica, holotype. Scale line = 1 mm. FUNGUS MOTHS 159 119 120 Figs 113-120 Male genitalia of Diataga species (113-119, right valva with juxta-transtilla complex; 120,ventral view of uncus lobes). 113, leptosceles, lectotype; 114, leptosceles (Brazil); 115, frustraminis,holotype; 116, brasiliensis, holotype; 117, compsacma, lectotype (juxta-transtilla region destroyed);118, levidensis, holotype; 119, 120, mercennaria, holotype. Mixed scales, 117 greatly enlarged. 160 GADEN S. ROBINSON 126 122 127 123 128 125 Figs 121-130 Male genitalia (121-126, aedeagus; 127-130, right valva). 121-126, Diataga species. 121,leptosceles, lectotype; 122,frustraminis, holotype; 123, brasiliensis, holotype; 124, compsacma, lecto-type; 125, levidensis, holotype; 126, mercennaria, holotype. 127-130, Morophaga species. 127, crem-narcha; 128, bucephala; 129, soror; 130, vadonella, holotype. Scale line = 1 mm. FUNGUS MOTHS 161 133 134 135 132 Figs 131-137 Male genitalia of Morophaga species (131, 133, 134- right valva; 135-137 -aedeagus). 131,borneensis, holotype; I32,sistrata; 133,formosana, holotype; 134, iriomotensis , holotype; 135, borneen-sls, holotype; 12>6,sistrata; I37,formosana, holotype. Scale line = 1 mm. 162 GADEN S. ROBINSON 138 139 140 Figs 138-140 138, 139, male genitalia of Morophaga species. 138, clonodes; I39,fasciculata, holotype.140, female genitalia of Morophagoides burkerella. Scale line = 1 mm. FUNGUS MOTHS 163 141 Figs 141-144 Female genitalia of Morophagoides species. 141, moriutii (Taiwan); 142, nimbiferum,holotype; I43,pythium, holotype; 144, montium, holotype. Scale line = 1 mm. 164 GADEN S. ROBINSON Figs 145-147 Female genitalia. 145, Bythocrates drosocycla; 146, Montescardiafuscofasciella, lectotype;147, Daviscardia coloradella. Scale line = 1 mm. FUNGUS MOTHS 165 148 150 Figs 148-150 Female genitalia of Daviscardia species. 148, beckeri; 149, lupulella, holotype; 150,mackiei. Scale line = 1 mm. 166 GADEN S. ROBINSON Figs 151, 152 Female genitalia. 151, Daviscardia hypocritella, holotype; 152, Scardia alleni, holotype.Scale line = 1 mm. FUNGUS MOTHS 167 153 154 156 155 157 Figs 153-157 153-156, female genitalia ofScardia species (detail of eighth sternite). 153, anatomella; 154,amurensis; 155, boletella; 156, caucasica, paratype. 157, female genitalia of Gentingia hollowayi,paratype. Mixed scales. 168 GADEN S. ROBINSON 158 159 Figs 158, 159 Female genitalia of Cranaodes species. 158, stereopa; 159, sequestrata. Scale line = 1 mm. FUNGUS MOTHS 169 163 Figs 160-164 Female genitalia. 160, Cnismorectis choritica\ 161, Scardiella approximatella. 162, 163,Necroscardia species. 162, mortidna, paratype, with (163) seventh sternite; 164,funeratella, lateral view(pathological specimen). Scale line = 1 mm. 170 GADEN S. ROBINSON Figs 165-169 Female genitalia. 165, Miniscardia minimella (Brazil); 166, Miniscardia sp. (Arizona) (tohalf scale); 167-168, Amorophaga rosemariae, holotype, with (168) seventh sternite; 169, Diatagaleptosceles. Scale line = 1 mm. FUNGUS MOTHS 171 170 \ \ 172 Figs 170-172 Female genitalia. 170, 171, Diataga species. 170, mercennaria, paratype; 171, direpta,holotype. 172, Morophaga clonodes. Scale line = 1 mm. 172 GADEN S. ROBINSON 173 Figs 173-176 Female genitalia (outline) of Morophaga species. 173, cremnarcha; 174, bucephala (NewGuinea); 175, vadonella; 176, soror (Angola). Scale line = 1 mm. FUNGUS MOTHS 173 Figs 177-179 Female genitalia of Morophaga species. 177, kobella, holotype; 118,fasciculata, paratype;179, sistrata (Sri Lanka). Scale line = 1 mm. 174 GADEN S. ROBINSON CDD Figs 180-182 Denuded heads. 180, Perilicmetis diplaca; 181, Tinissa insularia; 182, Cnismorectis choriti-ca. Scale line = 1 mm. FUNGUS MOTHS 175 183 R3 184 Figs 183, 184 Venation. 183, Diataga leptosceles; 184, Gentingia hollowayi, paratype. 176 GADEN S. ROBINSON 185 186 187 Figs 185-187 Female pupa of Diataga leptosceles (Costa Rica). 185, dorsal; 186, lateral; 187, ventralview. Scale line = 1 mm. FUNGUS MOTHS 177 188 189 T*VtvT* 'vrrj** 190 192 Figs 188-192 Pupae. 188, Diataga leptosceles (Costa Rica), male, dorsal view. 189, 190, Morophagachoragella, male, dorsal and ventral views. 191, 192, Morophaga choragella, female, dorsal and ventralviews. Scale line = 1 mm. 178 GADEN S. ROBINSON 193 S03 03 194 01 D2 O^-s 196 Figs 193-197 Final instar larva of Diataga leptoscdes (Costa Rica). 193, head, frontal view; 194, head,lateral view; 195, head (part), ventral view; 196, setal map of (P) prothorax, (M) mesothorax, andabdominal segments 1, 6, 8 and 9; 197, anal plate. FUNGUS MOTHS 179 Figs 198-200 Habitats of Scardiinae. 198, montane Himalayan oak forest in Nepal at 2700 m istype-locality of Amorophaga rosemariae. 199, tall mixed dipterocarp forest at 330 m in Brunei is habitatof Tinissa species (foreground clearing is helicopter landing-point: figures at edge of clearing give scalefor 25-metre forest canopy). 200, comparatively stunted upper montane forest on steep 1700 m ridges inBrunei is habitat of Cranaodes sequestrata and an undescribed Tinissa species; lower montane forest onGunong Mulu (distant ridge beyond low-lying cloud) is habitat of Gentingia hollowayi. 180 Abies 113 Acacia aulocarpa 130 Acer pseudoplatanus 109 agrifolia, Quercus 70 ash 92 atlantica, Pistacia 125 aulocarpa, Acacia 130 bamboo 108Betula 91birch 91bracket-fungi 37, 132 Castanopsis 112Coriolus 67Cryptophorus volvatus 113 Daedalea microsticta 116dead wood 75, 125, 132 edodes, Lentinus 69 Fagus 75 fomentarius, Fomes 90 Fomes67,91,127 Fomes fomentarius 90 Fraxinus 92 fremontii, Populus 116 fungus 39, 80, 90, 92, 108, 122, 130 GADEN S. ROBINSON Index to hostplants and fungi Ganoderma 67, 80, 91gilvus, Polyporus 70 himalayan oak 112kusanoana, Trametes 133 Lentinus edodes 69Lithocarpus 70Lupinus 70 microsticta, Daedalea 116Morus 125 oak, himalayan 112 Parrotia 67persica, Prunus 130Picea 75pine 88PinusSO, 113Pistacia atlantica 125plorans, Xanthoschisma 125Polyporaceae 71polypore 125 Polyporus 38, 67, 78, 80, 91, 116,122, 127 Polyporus gilvus 70Polyporus vulpinus 116Populus 80 Populus fremontii 116pore-fungi 75Prunus persica 130pseudoplatanus, Acer 109Pseudotsuga 80 Quercus 67, 80, 125Quercus agrifolia 70Quercus suber 125 rotten tree stumps 122 Schima 112shiitake 69suber, Quercus 125sycamore 109 Trametes kusanoana 133tree-stumps, rotten 122 volvatus, Cryptophorus 113vulpinus, Polyporus 116 Xanthoschisma plorans 125Zelcova 67 Index to Lepidoptera Invalid names are in italics; principal references are in bold. Acrolophus Poey 43Afrocelestis Gozmany 39, 43Afroscardia gen. n. 110Agarica Sodoffsky 86albipuncta Robinson 42alleni sp. n. 90amboinensis Robinson 42Amorophaga Zagulajev 111amurensis Zagulajev 89anatomella Grote 87angulatella Walsingham 43approximatella Dietz 109araucariae Robinson 42assamensis sp. n. 88astragalopa Meyrick 102Atabyria Snellen 120 bakeri Robinson 42baliomicta Meyrick 42beckeri sp. n. 81berkeleyella Powell 69bicolorella sp. n. 84bimendella Zeller 81boletella F. 89, 90boleti., 1777 131 boletiF.,179891borneensis sp. n. 126brasiliensis Zagulajev 117bucephala Snellen 123bucephala-group 121burkerella Busck 70Bythocrates Meyrick 77 capnochalca Meyrick 110caryophylella Busck 70caucasica Zagulajev 92chalcites Robinson 42chaotica Robinson 42chloroplocama Meyrick 42chomatias Meyrick 123choragella Denis & Schiffermiiller 131 choragella-group 130choritica Meyrick 103cinerascens Meyrick 42classeyi Robinson 42clonodes Meyrick 129clonodes-group 129Cnismorectics Meyrick 102coloradella Dietz 79 compsacma Meyrick 118conglomerata Meyrick 43convoluta Robinson 42Cranaodes Meyrick 98cremnarcha Meyrick 122cryptophori Clarke 112cultellata Gozmany & Vari 42 Daviscardiagen. n. 78Diataga Walsingham 114diplaca Meyrick 93direpta sp. n. 120distracta Meyrick 42dohertyi Robinson 42Dorata Busck 104drosocycla Meyrick 77Duomitella Koshantschikov 86 errandella Busck 70errantia Robinson 42eumetrota Meyrick 42 fasciculata sp. n. 133Fernaldia Busck 86fiskeella Busck 87 FUNGUS MOTHS 181 formosanasp. n. 128frustraminis sp. n. 116funeratella Zeller 106fungella Thunberg 131fungicolella Dumont 125fuscofasciella Chambers 76 Gentingia gen. n. 95Glessoscardia Kuznetzov 39goliath Robinson 42gracilis Walsingham 70 heterograpta Meyrick 42hirsutevestita Walsingham 43hollowayi sp. n. 96Hormantris Meyrick 102hypocritella sp. n. 86hyrcanella Zagulajev 132hyrcanica Zagulajev 113 indica Robinson 42insignis Zagulajev 42insularia Robinson 42iranensis Petersen 66iriomotensis sp. n. 128isthmiella Busck 135iulina Walsingham 73 japonicasp. n. 114 kidukaroka Robinson 42kobella sp. n. 133krakatoa Robinson 42kurenzovi Zagulajev 75 leptosceles Walsingham 115, 116 Leptozancla Meyrick 134 levidensis sp. n. 118 lochaea Meyrick 43 luctuosa Walsingham 81, 82, 83, 85, 95lupulella sp. n. 85 mackiei sp. n. 83 maculosa Diakonoff 129mediella Hiibner 131mercennaria sp. n. 119Microscardia Amsel 120minimella Busck 104Miniscardia gen. n. 104Montescardia Amsel 74montium Walsingham 71, 72morellus Duponchel 125morellus-group 125moriutii sp. n. 68Morophaga Herrich-Schaffer 120Morophagoides Petersen 65morticinasp. n. 107Moscardia gen. n. 93 Narycia Stephens 43Necroscardia gen. n. 106nigrocapitella Petersen 122nimbiferum sp. n. 72 oroyasp. n. 99Osphretica Meyrick 120 Palaeoscardiites Kuznetzov 39palmodes Meyrick 42Paraclystis Meyrick 43parallela Robinson 42Pectiniscardia gen. n. 101penicillata Gozmany 97Perilicmetis Meyrick 92perilithias Meyrick 42pharetrodes Meyrick 135Philagrias Meyrick 134philippinensis Robinson 42phrictodes Meyrick 42poliophasma Bradley 42Polymnestra Meyrick 108poly port Esper 91polysema Zagulajev 42polystacta Meyrick 42porphyrea Lower 129pravatella Busck 76Proscardiites Kuznetzov 39 prostylias Meyrick 101pythium sp. n. 72 radulella sp. n. 80relicta Koshantschikov 91renitens Meyrick 94rigida Meyrick 42rosemariae sp. n. Illrotundata Matsumura 123ruwenzorica Gozmany 42 saccharata Meyrick 43Scardia Treitschke 86Scardia sensu lato 135Scardiella gen. n. 108Scardiites Kuznetzov 39Semeoloncha Gozmany 39, 97sequestrata Meyrick 100sistrata Meyrick 127sistrata-group 126soror Gozmany 124spaniastra Meyrick 42sp. (Daviscardia) 84sp. (Miniscardia) 105stereopa Meyrick 98 talaroscia Meyrick 134talyshensis Zagulajev 132tessulatellus Zeller 74tholerodes Meyrick 135Tinissa Walker 108torvella mysorensis Robinson 42torvella torvella Walker 42transversella Walker 42 ussuriensis Caradja 67, 68 vadonella Viette 124varna sp. n. 95 yaloma Robinson 42zelotica Meyrick 134 British Museum (Natural History) The Generic Namesof Moths of the World The aim of this series is to provide a complete list of all thegeneric names, their type-species, designations, and appropriatereferences, for each of the families of moths treated. By ensuringthat each generic name has its type-species fixed in accordancewith the International Code of Zoological Nomenclature, thisseries should rectify present confusion and form a basis for a morestable nomenclature. All generic names have been checked forhomonymy, and if necessary replaced. The authors are all specialists on the staff of the Department ofEntomology, British Museum (Natural History) and have theunrivalled collections and resources of the Museum on which tobase their work. Volume 1; Noctuoidea (part): Noctuidae, Agaristidae and Nolidae. by I. W. B.Nye. 1975 Pp.viii + 568 frontispiece 38.00 Volume 2; Noctuoidea (part): Arctiidae, Ctenuchidae, Dioptidae,Lymantriidae, Notodontidae, Thaumetopoeidae and Thyretidae.by A. Watson, D. S. Fletcher and I. W. B. Nye.1980 Pp.xiv + 228 frontispiece 26.50 Volume 3; Geometroidea: Apoprogonidae, Axiidae, Callidulidae, Cyclidiidae, Drepanidae, Epicopeiidae, Epiplemidae, Geometridae, Pterothysanidae, Sematuridae, Thyatiridae and Uraniidae. by D.S.Fletcher. 1979 Pp.xx + 243 frontispiece 26.50 Volume 4; Bombycoidea, Castnioidea, Cossoidea, Zygaenoidea,Sesioidea. Additions and amendments to Vols 1-3.by D. S. Fletcher and I. W. B. Nye.1982 Pp.xiv + 192 frontispiece 26.50 Volume 5; Pyraloidea. by D. S. Fletcher & I. W. B. Nye. 1984 Pp.xv + 185 25.00 To be published: Volume VI. Microlepidoptera and additions and corrections to Volumes I-V. This volume will complete the series. The set of 5 volumes published so far 125.00 Titles to be published in Volume 52 The sandflies of Egypt (Diptera: Phlebotominae)By R. P. Lane Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae) By G.S.Robinson A revision of the European Agathidinae (Hymenoptera: Braconidae) By G. E. J. Nixon A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revision of certain genera By J. Quinlan Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester (NATURAL 25A' PRESENTED GENERAL Bulletin of the British Museum (Natural History) A revision of the European Agathidinae(Hymenoptera: Braconidae) G. E. J. Nixon Entomology series Vol 52 No 3 24 April 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History),Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1986 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06017 1 ISSN 0524-6431 Entomology series Vol 52 No 3 pp 183-242British Museum (Natural History)Cromwell RoadLondon SW7 5BD Issued 24 April 1986 A revision of the European Agathidinae(Hymenoptera: Braconidae) G. E. J. Nixon c/o Department of Entomology, British Museum (Natural History), Cromwell Road, LondonSW7 5BD I GENERAL LIBContents Synopsis 183 Introduction 183 Depositories 184 Taxonomic characters 184 Biology 185 Host records 186 Agathidinae 187 Key to European genera 188 Synonymic list of species 188 Disophrys Forster 190 Cremnops Forster 191 Agathis Latreille 192 Rhamphagathis Tobias 214 Microdus Nees von Esenbeck 215 Earinus Wesmael 226 Baeognatha Kokujev 229 Species inquirendae 230 Acknowledgements 230 References 230 Index 242 Synopsis Seven genera of the braconid subfamily Agathidinae are revised for the European zoogeographical region .Keys to the genera and species are provided. Fifty-four species are recognized as valid, 11 are newlysynonymized, and 11, including an extralimital species from Jordan, are described as new. Lectotypes aredesignated for 13 nominal species. Authentic host records are included together with an account of thebiology of the subfamily. Introduction Although the braconid subfamily Agathidinae has a world-wide distribution, it is poorlyrepresented in the western Palaearctic region. The present revision deals with the fauna ofEurope, including European U.S.S.R., and Turkey; one species from Jordan is included forcomparative purposes. Earlier contributions to the taxonomy of European Agathidinae are those of Nees vonEsenbeck (1834), Wesmael (1837), Reinhard (1867), Thomson (1895), Marshall (1885; 1888)and Fahringer (1937). The more recent, notable taxonomic revisions on Palaearctic Agathidinaeare those of Telenga (1955) and Tobias (1976<z), the former concentrating on the Europeanspecies, the latter on the Central Asian species. Their combined research reveals an agathidinefauna that hardly differs numerically from that covered in the present work. Even Tobias's paper(19766) on the Far Eastern species of Microdus includes only four that are not represented inEurope. The Agathidinae is well represented in the tropics and Shenefelt's catalogue (1970) lists Bull. Br. Mus. not. Hist. (Ent.) 52 (3): 183-242 Issued 24 April 1986 184 G. E. J. NIXON numerous species; however, most of these will require investigation before their status can beproperly assessed. Some attempt to sort out the chaos of names in the Oriental species has beenmade by Bhat & Gupta (1977), but other than this no major taxonomic research has been doneon the tropical fauna. The classified, tropical material in the British Museum (Natural History)has been examined briefly; although quantitatively small and unreliably determined it providessome information on evolutionary trends within the Agathidinae. It seems that the subfamilydivides into two main sections, the Agathis-Microdus group of genera which have lobed orsimple claws and are restricted mainly to the Northern Hemisphere, and the Cremnops-Disophrys group of genera which are predominantly tropical in distribution. Shenefelt (1970)includes most of the described species in these two genus groups. Microdus is Holarctic but alsooccurs in the Southern Hemisphere. Two mainly tropical genera, Cremnops and Disophrys, areeach represented by a single species in the western Palaearctic region. Little can be said here about the North American Agathidinae, but Agathirsia Westwoodexhibits a line of descent different from the two genera just mentioned. Its wing venation istypical of the subfamily but its body-form and especially the exaggerated length of the labiumisolate it from the Old World genera. The aim of the present review is to enable the reader to identify more easily the Europeanspecies of Agathidinae; to postulate evolutionary trends within the subfamily is beyond itsscope. Depositories AA A. A. Allen collection, Reigate, Surrey. AS Zoological Institute, Leningrad. AZ A. Zaykov collection, Plovdiv. BMNH British Museum (Natural History), London. EH Erasmus Haeselbarth collection, Munich. HNHM Termeszettudomanyi Muzeum, Budapest. IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels. LC Linnaean collection, Linnean Society, London. MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin. MS Mark Shaw collection, Edinburgh. NM Naturhistorisches Museum, Vienna. RNH Rijksmuseum van Natuurlijke Historic, Leiden. UM University Museum, Oxford. USNM National Museum of Natural History, Washington, D.C. ZH Zoological Museum, Helsinki. ZI Zoologiska Institutionen, Lund. ZM Zoologisk Museum, Copenhagen. ZSBS Zoologische Sammlung des Bayerischen Staates, Munich. Taxonomic characters The agathidine taxa studied in this revision are variable in colour, sculpture and ovipositorlength, and Agathis and Microdus contain groups of what at first sight appear to be valid speciesbut which subsequently prove to be inseparable because of intermediate forms. There are twoalternatives: either the variants are regarded as different species, or a wide range of variation isaccepted. The latter alternative has been chosen here since the concept of a species-aggregate,even if it offers no more than a provisional solution, at least avoids an increase in the number ofdubious taxa. Nevertheless, species of Agathidinae have some characters that are reasonablyconstant and therefore useful for identification purposes. The elongation of the head in facial view, though slightly variable, is frequently used inseparating species of Agathis. It is defined as the length of the malar space measured against thelonger diameter of the eye. The surface of the frons is also important, especially in Agathis inwhich it is either smooth, or provided with a deepened cavity or trough margined by raised keels.The conformation of this structure is diagnostic in such species as breviseta and assimilis. A REVISION OF THE EUROPEAN AGATHIDINAE Indispensable for at least a partial breakdown of species of Agathis are the mouth-parts. Theseare modified to form a beak, certain parts of which, especially the prementum, are muchlengthened. Since the whole structure is, to some degree, retractile (and thus hidden), only thegalea is used; this is attached to the base of the maxillary palpus, is constant in length and isgenerally visible. The thorax is uniform in shape though in Agathis it is subject to slightelongation. The notaulices are usually deeply impressed; when weak as in Agathis nigra andMicrodus lugubrator, they may sometimes be absent in some specimens. Propodeal sculpture isvariable and of limited value; within a species, such as Agathis glabricula, it can be entirelyrugose or, more frequently, it has smooth, polished lateral panels. The number of teeth on theouter side of the middle tibia of Agathis is fairly constant; sometimes there are sufficient teeth(8-10, as in fulmeki) to be of diagnostic importance. In Agathis it is essential to examine theclaws for the presence or absence of a lobe or tooth. Venation is used very little to separate species. In Microdus the radius (Rs) is sometimescharacteristically curved on the side towards the stigma, and in one species, cingulipes, thecurvature is so pronounced as to be diagnostic. The shape of the second submarginal cell(referred to herein as the areolet) is virtually uniform in Microdus, but in Agathis it is variable,and can be intraspecifically either triangular or distinctly four-sided; rarely, as in Agathisanglica, the four-sided condition is so pronounced that it always retains its distinctiveness. Lessvariable and more reliable in separating species of Agathis is the ratio between the two abscissaeof the postmarginalis (Fig. 67). This vein is the distal extension of the stigma along the edge ofthe wing known as Rl; sometimes it continues beyond the point where it is joined by the radius(Rs) and thus becomes composed of two abscissae. The distal abscissa is sometimes so long as tobe diagnostic, as in Agathis minuta. In Agathis, the gaster rarely provides characters of taxonomic value, being in greater partsmooth and polished. In Microdus, however, it frequently has sculpture that is specific, thoughvariable in degree of intensity and definition. Earinus gloriatorius demonstrates the unreliabilityof gastral sculpture in the Agathidinae; in this species tergites 2+3 are sometimes smooth andpolished, sometimes conspicuously rugose, with every intermediate condition. Ovipositor length is fairly uniform and of little use in the identification of species. There isnearly always considerable distortion in the apical tergites of dried specimens so that the actuallength of the ovipositor is difficult to determine; the length of the ovipositor sheaths is moreeasily assessed and it is this length compared with that of the whole body that is mentioned in thedescriptions. The morphological terms used in this revision are based on those of Richards (1977) and Eady(1974). The fused second and third tergites of the gaster are referred to as 'tergites 2+3' because,in the majority of braconids, these two tergites have the appearance of a single segment,morphologically bipartite only by the presence of two lateral spiracles. Biology As far as is known species of Agathidinae are internal parasites of lepidopterous larvae, but theirbiology has received little attention. There are a few exceptions, however. Simmonds (1947)described the life history of Agathis vulgaris (Cresson), a parasite of Loxostege sticticalis (L.),the sugar-beet web- worm in North America, and gave a detailed account of the morphology ofthe early stages of the parasite and an assessment of its value in biological control. Simmondsconsidered Agathis vulgaris to be an important parasite of its host and in fact found it to be themost frequently bred parasite in his experiments. Dondale (1954) described the biology of Microdus dimidiator Nees von Esenbeck under thename Agathis laticinctus Cresson (Krombein et al. , 1979: 193). It is one of the parasites attackingthe eye-spotted bud moth, Spilonota ocellana (Denis & Schiffermuller) , an orchard pest intemperate North America. According to Dondale, the eggs of the parasite are deposited withinthe host during August and by early September have given rise to polypod, first-stage larvaewhich then enter a winter diapause. The overwintering larvae can be found within thehost-caterpillars until the following July. The host-caterpillar reaches maturity and spins its 186 G. E. J. NIXON cocoon in late June or early July; the larva of the parasite then bores its way out and feedsexternally on the juices of the host during a period of three or four hours, finally destroying it.Dondale does not consider Microdus dimidiator to be a particularly successful parasite but hispaper, with its excellent illustrations of larval structures, is a notable contribution to the biologyof Microdus. Thorpe (1933) gave a brief account of the biology of Microdus pumilus Ratzeburg, a parasiteof Coleophora laricella (Hiibner), the larch case-bearer. He figured the first and second instarlarvae of the parasite but had no material of the later stages. He found Microdus pumilus to bevery rare in England and , in spite of breeding a wide range of parasites from the host , obtained itonly from the Forest of Dean, Gloucestershire. Host-material from the south of France, on theother hand, produced pumilus in abundance, the parasite reaching a level of 12 per cent, theadults emerging in June. The first instar of the parasite diapauses through the winter within itshost and in this respect resembles Microdus dimidiator. According to Tobias (1964), one of themost serious pests of agriculture in Armenia is the Mallow moth, Pexicopia (Pectinophora)malvella (Hiibner). Over a number of years parasites of this moth were bred by A. S. Avetyan,among them Agathis malvacearum Latreille. Tobias does not assess the significance of mal-vacearum in the control of Pexicopia malvella and used Avetyan's bred material mainly to studythe parasite's seasonal variability in size and colour. The Agathidinae, apart from Agathis vulgaris (mentioned above), have not been used inbiological control in the tropics, consequently the literature provides only casual information onlife-cycles. Gupta (1964), however, bred specimens of an Agathis sp. from the larvae ofHolcocera pulverea Meyrick, the Lac predator. These were sent to C. F. W. Muesebeck whoidentified them as Agathis f estiva Muesebeck, a parasite of the Oriental fruit moth, Grapholitamolesta Busck, in China. Gupta states that nothing much is known about the biology of/estiva,and suggested that it is hardly more than an occasional parasite of pulverea but that it may alsoparasitise other lepidopterous larvae occupying similar habitats. Host records HOST PARASITE LEPIDOPTERA BLASTOBASIDAE Blastobasis lignea Walsingham Microdus rugulosus COLEOPHORIDAE Coleophora granulatella Zeller Agathis artemesiana Coleophora flavipennella (Duponchel) Microdus lugubrator Coleophora frischella (L . ) Microdus cingulipes Coleophora glaucicolella Wood Agathis minuta Coleophora halophyella Zimmerman Agathis asteris Coleophora hemerobiella (Scopoli) Microdus lugubrator Coleophora laricella (Hiibner) Microdus pumilus Coleophora lutipennella (Zeller) Microdus lugubrator Coleophora meridionella Klimesch Agathis meridionellae Coleophora troglodytella (Duponchel) Agathis glabricula GELECHIIDAE Anarsia eleagnella Kuznetzov Baeognatha armeniaca Apodia bifractella (Duponchel) Agathis tibialis Caryocolum fraternella (Douglas) Microdus cingulipes Chrysoesthia hermannella (Fabricius) Agathis meridionellae Chrysoesthia sexguttella (Thunberg) Agathis meridionellae Metzneria lappella (L.) Agathis varipes Pexicopia malvella (Hiibner) Agathis malvacearum Ptocheuusa paupella (Zeller) Agathis tibialis Recurvaria nanella (Denis & Schiffermiiller) Baeognatha armeniaca A REVISION OF THE EUROPEAN AGATHIDINAE 187 NOCTUIDAE Agrochola drcellaris (Hufnagel)Agrochola lota (Clerck)Atethmia centrago (Haworth) OECOPHORIDAE Agonopterix ciliella (Stainton)Agonopterix heradiana (L.) PYRALIDAE Eurrhypara hortulata (L.)Myelois drrigerella (Zincken)Pyrausta aurata (Scopoli) SESIIDAE Conopia sphedformis (Denis & Schiffermiiller) TINEIDAE Morophaga boleti (Fabricius)Triaxomera parasitella (Hiibner) TORTRICIDAE Cydia molesta (Busck) Cydia pomonella (L.) Cydia funebrana (Treitschke) Dichrorampha acuminatana (Lienig & Zeller) Epinotia mercuriana (Frolich) Epiblema scutulana (Denis & Schiffermiiller) Gypsonoma dealbana (Frolich) Hedya nubiferana (Haworth) Pammene regiana (Zeller) Rhopobota ustomaculana (Curtis) Rhyadonia buoliana (Denis & Schiffermiiller) Spilonota ocellana (Denis & Schiffermiiller) Tortrix viridana (L.) Earinus nitidulusEarinus nitidulusEarinus nitidulus Earinus gloriatoriusEarinus gloriatorius Cremnops desertorAgathis varipesAgathis griseifrons Cremnops desertor Microdus calculatorMicrodus calculator Agathis f estivaMicrodus rufipesMicrodus conspicuusCremnops desertorMicrodus conspicuusBaeognatha armeniacaMicrodus tumidulusAgathis anglicaMicrodus clausthalianusMicrodus rufipesMicrodus rufipesMicrodus conspicuusMicrodus conspicuusMicrodus rufipesMicrodus dimidiatorMicrodus fortipesMicrodus rufipesMicrodus dimidiator Most of the above names are based on Kloet & Hincks (1972). AGATHIDINAE DIAGNOSIS. Disco-cubital vein (Rs+M) mostly undeveloped; hence 1st discoidal cell and 1st submarginalcell confluent, except in Earinus Wesmael; 2nd submarginal cell usually very small, triangular orsubtriangular, forming an areolet (Fig. 50); when large (not much!) then 4-sided and often with accessoryvein arising from outer side of 2nd transverse cubitus (2rm) (Disophrys-Cremnops genus-group); 2ndtransverse cubitus absent in Baeognatha Kokujev; hence no closed areolet in this genus; radial cell alwaysshort, mostly narrow, terminating far proximal to apex of fore-wing (Figs 50, 55, 60). Head in facial view elongate (Disophrys-Cremnops genus-group); less frequently transversely elliptical(Microdus) . Mouth-parts mostly lengthened and drawn out in form of beak ; maxillary palpus 5-segmented ;labial palpus 4-segmented; in Microdus segment 3 of labial palpus sometimes so small that palpus appearsto be 3-segmented. Thorax of generalised form except in highly specialised western Palaearctic Rhamphagathis Tobias.Pronotum with deep pit on each side, separated internally by fenestra. Propodeum variable; strongly,almost symmetrically areolated by raised keels (Disophrys-Cremnops genus-group) and then spiracleelongate; smooth to rugose with at most two longitudinal keels and spiracle circular (Agathis-Microdusgenus-group). Gaster usually highly polished, smooth but tergites 2+3 often sculptured in Microdus; in the Afrotropic- 188 G. E. J. NIXON al Braunsia Kriechbaumer, related to Microdus, the gaster shows a highly characteristic sculpture of strongstriation. Spiracles of tergite 1 situated on dorsal plate. Ovipositor variable in length. Based on knowledge of a relatively small sample of the subfamily, the above description has necessarily arestricted value. A study of the virtually unknown fauna of Australia and South America would certainlycall for modifications. The position of the Agathidinae with regard to the other braconid subfamilies has been defined by vanAchterberg (1976). So far there is no evidence to contest the assertion that the Agathidinae can beseparated from all other subfamilies by the venation of the fore-wing: the short radial cell in combinationwith the small second submarginal cell provides a character not found elsewhere in the Braconidae. Thischaracter is in general supported by the elongation of the head. Van Achterberg (1984) has given a more up-to-date review of the phylogeny of the Braconidae basedupon a comprehensive cladistic analysis. Key to European genera 1 Hind claw cleft; frons with sharp keel or rounded ridge between external side of antennal insertion and lateral ocellus; inner spur of hind tibia very long, reaching middle of basalsegment of hind tarsus 2 - Hind claw not cleft; frons unarmed or much more often with basal lobe or tooth; inner spur of hind tibia shorter, not reaching middle of basal segment of hind tarsus 3 2 Body entirely fulvous ; fore- wing having dappled appearance , with broad , hyaline fascia beneath half yellow stigma and paler at base and apex; maxillary palpus in female with long bristlesthat are longer than palpal segments; ovipositor about as long as body CREMVOPSF6rster(p. 191) - Body variable in colour but always partly blackened ; fore- wing without dappled appearance and at most with pale, hyaline streak beneath stigma; maxillary palpus of female with only veryshort hairs; ovipositor very short, projecting only slightly beyond apex of gaster DISOPHRYSF6rster(p. 190) 3 First submarginal cell separated from 1st discoidal cell by a fully sclerotised vein (Fig. 67); sternaulus absent EARINUS Wesmael(p. 226) - First submarginal cell more or less fused with 1st discoidal cell; sternaulus present (rarely reduced to a fine groove or virtually absent and then head in facial view elongate) 4 4 Fore-wing without a closed areolet (2nd submarginal cell) BAEOGNATHA Kokujev(p. 229) - Fore- wing always with a closed areolet (2nd submarginal cell) 5 5 Middle lobe of mesoscutum deeply hollowed out to form a wide trough , each side of which shows anteriorly as a blunt projection or shoulder; clypeus in profile in form of a short snout;mandible broadened, flattened, slightly concave on external side RHAMPHAGATHIS Tobias (p. 214) - Middle lobe of mesoscutum of ordinary form, evenly convex in front; clypeus in profile not or only very slightly projecting; mandible not thus broadened or concave externally 6 6 Head in facial view always elongate; mouth parts almost always lengthened in form of beak, the galea at least 1.5 times longer than wide. (In minuta, which has a very short galea, the head infacial view is subtriangular.) AGATHISLatrei\\e(p. 192) - Head in facial view strongly transverse, rarely slightly lengthened as in linguarius; mouth parts never lengthened in form of beak, galea not longer than wide MICRODUS Nees von Esenbeck(p. 215) Synonymic list of species AGATmSLatreilleachterbergisp. n.anchisiades sp. n.anglica Marshall albanica Fischer syn. n.ariadnesp. n.artemesiana Fischerassimilis Kokujevasteris Fischerbreviseta Nees von Esenbeck A REVISION OF THE EUROPEAN AGATHIDINAE 189 fulmeki Fischerglabricula Thomson albicostellae Fischer syn. n.glaucoptera Nees von Esenbeckgracilipes Hellengriseifrons Thomson laticarpa Telenga syn. n.malvacearum Latreillemelpomene sp. n.meridionellae Fischerminuta Niezabitowskimontana Shestakovnigra Nees von Esenbeck testaceipes Fischer syn. n.pappe/sp. n.pedias sp. n.persephone sp. n.polita sp. n.rostrata Tobiasrufipalpis Nees von Esenbecksemiaciculata Ivanovsyngenesiae Nees von Esenbecktaurica Telengatibialis Nees von Esenbeckumbellatarum Nees von Esenbeck kolazyi Fischer syn. n.varipes Thomsonzaykovisp. n.BAEOGNATHA Kokujevnigra Telengaarmeniaca TelengaCUEM/VOPSForsterdesertor (L.) deflagrator (Spinola)DISOPHRYSForstercaesa Klug anthracina Kriechbaumer syn. n.EAU/JVl/S Wesmaelelator (Fabricius) nitidulus Nees von Esenbeck syn. n. thoracicus Nees von Esenbeck syn. n.gloriatorius (Panzer) affinis Wesmael varicoxis Wesmael syn. n. delusor Wesmael syn. n. tuberculatus Wesmael syn. n. bicingulatus Thomson syn. n. ochropes Lyle syn. n.transversus Lyle MICRODUSNees von Esenbeckcalculator (Fabricius) abscissus Ratzeburgcingulipes Nees von Esenbeckclausthalianus (Ratzeburg)conspicuus Wesmael arcuatus Reinhard syn. n. zonatus (Marshall)dimidiator Nees von Esenbeckeriphyle sp. n. 190 G. E. J. NIXON fortipes Reinhardlinguarius Nees von Esenbecklugubrator Ratzeburgnugax Reinhardpumilus Ratzeburgrufipes Nees von Esenbeckrugulosus Nees von Esenbecktumidulus Nees von Esenbecktegularis Thomson syn. n.zaykovisp. n. RHAMPHAGA THIS Tobiasnasicornis Telenga DISOPHR YS Forster Disophrys Forster, 1862: 246. Type-species: Ichneumon inculcator Linnaeus sensu Forster, 1862 (= Agathis caesa Klug).Diophrys Kriechbaumer, 1898: 181. [Unjustified emendation.] I have examined the original specimen of Ichneumon inculcator Linnaeus which is in the LC, London. It isan ichneumonid and as such has been dealt with by Fitton (1978) and van Rossem (1969). Forster makes no mention of what material he had but merely cites Ichneumon inculcator L. as thetype-species of Disophrys, adding a reference to Nees von Esenbeck: 'Agathis inc. Hym.ichn.aff.Mon. 1:138'. In view of the above, Disophrys must either be considered a genus in the Ichneumonidae, or interpretedas actually described by Forster. The latter course has been followed here but ratification will be requiredby the International Commission for Zoological Nomenclature. The oldest available name for the species misidentified by Forster is caesa Klug, 1835, described from afemale taken by Waltl at Puerto Real in Andalucia, Spain. This specimen has been examined; it bears alabel, presumably in Klug's handwriting: 'Andalusien, Waltl d.' and another, red, marked 'type'. I havelabelled this specimen as 'Agathis caesa Klug. Holotype $, G. E. J. Nixon, 1984', this being the nameunder which it was originally described. DIAGNOSIS. Head in facial view elongate. Galea of maxillary palpus fully 3 times longer than greatest width;labial palpus 4-segmented; two apical segments subequal. Ocelli in high triangle; posterior tangent toanterior ocellus not touching posterior pair. Antennal sockets on inner side each with raised, sublamelli-form tubercle; in frontal view of head, space between antennal sockets shows as bilobed projection. Fronson each side with raised ridge . Notaulices deeply impressed . Areolet of fore-wing 4-sided ; outer side of 2ndtransverse cubitus (r-m) with stub of vein; radius leaving stigma in basal third; distal abscissa ofpostmarginalis about half as long as proximal abscissa. Propodeum with strongly raised ridges enclosingareas of uneven size; spiracle large, subellipitical. Inner spur of hind tibia reaching middle of basal segmentof hind tarsus; outer side of middle tibia without teeth; claws cleft. Gaster smooth, polished. Ovipositorprojecting only slightly beyond apex of gaster. The above diagnosis is based on the type-species and cannot therefore encompass the range of structurethat may later be found to be acceptable for a more realistic definition of the genus. Although numerous species of Disophrys have been described from the tropics (Shenefelt, 1970), onlyfour are known from the Palaearctic region. As well as caesa, these are calcaratrix Telenga (1955), dissorsKokujev (1903) and manifesto Kokujev (1903), all from Central Asia and all represented by at least oneexample in the BMNH. Examination of these three species suggests that modifications of the genericconcept of Disophrys will include variation in the surface sculpture of the hind tibia (manifesto), thethickness and vestiture of the longer hind tibial spur (calcaratrix) , and the strength and sharpness of thefrontal keel (manifesto). Disophrys caesa (Klug) Agathis caesa Klug, 1835: 89. Holotype <j>, SPAIN (MNHU) [examined]. [Ichneumon inculcator Linnaeus; Forster, 1862: 246. Misidentification.] Disophrys caesus (Klug) Marshall, 1890: 574. Diophrys [sic] anthracina Kriechbaumer, 1898: 185. LECTOTYPE cf , SPAIN (ZSBS), here designated [examined]. Syn. n.Disophrys anthracina Kriechbaumer; Szepligeti, 1904: 124. A REVISION OF THE EUROPEAN AGATHIDINAE 191 Cf $ , 8-11 mm long (excluding ovipositor). Body varying in colour from almost entirely red (holotype ofcaesd) to entirely black (anthracind) . Hind tibia varying from black and red to entirely black. Basal half offore-wing either entirely infumate or with median and submedian cells almost hyaline (lectotype ofanthracind). J. Head from above strongly transverse; in facial view somewhat variable; in nominate form decidedlyelongate but less so in central European examples. Face strongly raised along middle line, covered withmuch coarse punctuation or, towards antennal insertions, rugose-punctation; face also much deepened atsite of toruli. Frontal ridge strongly raised, sharp. Galea about 0.75 times longer than malar space.Antenna with 40-48 segments; flagellum tapering apically; middle segments almost transverse. Meso-scutum in profile sloping very gradually towards pronotum in Spanish specimens; in examples from furthereastwards, the slope becomes steeper; this is correlated with smaller size and increase of red colour.Prepectal keel strong , well defined . Sternaulus in form of a wide trough , crossed by very strong rugae . Hindcoxa above very coarsely rugose-reticulate; rugose area often separated from the weak oblong troughalongside it by a raised ridge; hind femur densely covered with shallow pits; hind tibia much roughenedthrough dense covering of tiny, oblong tubercles; inner spur of hind tibia unmodified, about 1.5 timeslonger than outer one and reaching only slightly beyond middle of basal segment of hind tarsus. Gastersmooth, polished. Ovipositor sheath about 3 times longer than basal segment of hind tarsus. O". Like female except for sexual differences. MATERIAL EXAMINED (from the Iberian Peninsula) Portugal: 1 Cf, near Coimbra, Beira littoral, 22.vii.1970 (Pronk) (RNH). Spain: 1 $, Albarracin(RNH); 1 $, Ost Peja, ll.vii.1894 (RNH); 1 $ (holotype of caesd), Andalucia, Puerto Real (Waltt)(MNHU); 5 cT, 3 $, Barcelona, Calella d. Costa, vi.1971 (Boucek) (BMNH); 1 $, Burgos, 26.vi.1961(RNH); 1 Cf , 1 $i Jaen, Las Correderas (RNH); 2 d" (lectotype and paralectotype of anthracind), Castille,Cuenca, 1896 (Korb) (ZSBS).Other material examined 8 d", 34 $. Austria, Greece, Hungary, Turkey, Yugoslavia (all BMNH). COMMENTS. Two males of the original material collected by Korb in Castille, Cuenca, and sent by him toKriechbaumer, are in ZSBS; they agree with the original description in all respects except sex (Kriech-baumer stated that he had three females but found difficulty in sexing specimens of Disophrys). A malelabelled 'Castilien, Cuenca, Korb 1896' is here designated as lectotype; the second specimen is labelledparalectotype. CflEMJVOPS Forster Cremnops Forster, 1862: 246. Type-species: Ichneumon deserter Linnaeus, by monotypy. Achterberg (1982: 136) has caused confusion concerning the correct identity of the type-species ofCremnops and questions whether the latter is valid or a synonym of Vipio Latreille (1804: 173). Hisargument hinges on whether Ichneumon desertor L. , 1758 and what he calls Ichneumon desertor F. (recte /.desertor L. sensu F. , 1775) refer to the same species. He considers that they represent a single species as isclear from his statement 'Ichneumon desertor Fabricius is not a separate species, since Fabricius cited in hisfirst description only the original description of Ichneumon desertor Linnaeus, 1758, together with thereference to Linnaeus' description. Therefore there cannot be any doubt that there is no separate speciesnamed Ichneumon desertor Fabricius. This fact was overlooked by Bradley (1919: 59) when he stated that"the type of Vipio Latreille is Ichneumon desertor Fabricius, not of Linnaeus" '. Achterberg continues'Because nomenclatorily Ichneumon desertor Fabricius is actually /. desertor Linnaeus, 1758, the genusVipio Latreille . . . becomes ... a senior synonym of Cremnops Forster, 1862'. Obviously the truth of thisstatement depends on the validity of his previous statement. It can be shown that Achterberg's reasoning isfalse and that he misinterpreted Bradley (1919) as well as several earlier workers. He weakens hisargument by stating that 'At present in the Fabricius collection there are specimens under desertor whichare not conspecific (or congeneric) with /. desertor Linnaeus and this indicated only that Fabricius haddifficulties in interpreting /. desertor Linnaeus'. This remark does not only indicate that Fabricius hadtrouble in interpreting /. desertor Linnaeus but implies also that he misidentified the Linnaean species andthat /. desertor L. sensu F. , 1775, could in fact represent a different taxon. Earlier workers such as Latreille(1804), Nees von Esenbeck (1834) and Forster (1862) also considered that Fabricius had misidentifieddesertor L. and that the species he had belonged to a different group of Braconidae. Latreille (1804) placedwhat he called desertor F. (recte desertor L. sensu F.) with two other species in his genus Vipio. TheFabrician interpretation of desertor L. has been associated with this generic name ever since, having beencited as the type-species of Vipio by Forster (1862). Ichneumon desertor L., on the other hand, has long 192 G. E. J. NIXON been recognised as an agathidine and is the type-species of Cremnops by monotypy. Since it is obvious that/. deserter L. and /. desertor L. sensu F. refer to radically different species, Achterberg's (1982) contentionthat Vipio and Cremnops are synonyms because they have the same type-species is incorrect. The critical references in the taxonomic history of the name desertor are as follows.Linnaeus, 1758: 563. Original description of Ichneumon desertor.Fabricius, 1775: 334. Ichneumon desertor L. sensu F., 1775. Latreille ? 1804: 173. Ichneumon desertorL. sensu F., 1775. Transferred to genus Vipio Latreille.Forster, 1862: 235. Ichneumon desertorL. sensu F. , 1775. Cited as type-species of Vipio Latreille.Spinola, 1808: 101. Bracon deflagrator, erected (unnecessarily) as replacement name for /. desertor L.Nees von Esenbeck, 1834: 125. Bracon deflagrator Spinola transferred to genus Agathis, with /. desertor L. wrongly placed as junior synonym.Forster, 1862: 246. Agathis deflagrator (wrongly attributed to Nees von Esenbeck) cited as type-species of Cremnops. Correct citation should have been 'Agathis deflagrator (Spinola) sensu Nees, 1834'. DIAGNOSIS, cf $ . Head in facial view elongate. Mouth parts lengthened; galea fully twice as long as wide. Ablunt keel, an extension of the outer rim of antennal socket, extends obliquely across frons half waytowards posterior ocellus. Frons bilobed between antennal insertions. Propodeum almost symmetricallyareolated by sharply raised keels; propodeal spiracle narrowly oval. Outer side of middle tibia withouttrace of teeth (cf . Agathis and Microdus). First abscissa of mediella of hind wing much shorter than second;areolet of fore-wing 4-sided with stub of vein arising from outer side of 2nd transverse cubitus. Gasterpolished, smooth. Cremnops desertor (Linnaeus) Ichneumon desertor Linnaeus, 1758: 563. Holotype $, EUROPE (LC) [examined]. Bracon deflagrator Spinola, 1808: 101. [Unnecessary replacement name for /. desertor Linnaeus.] Agathis deflagrator (Spinola) Forster, 1862: 246. [Attributed to Nees von Esenbeck, 1834: 139.] Cf $, 6-8 mm long (excluding ovipositor). Entirely bright fulvous. Wings with a broad hyaline fascia atmiddle and two large, almost confluent spots at apex of wing. $. Bristles of palpi upstanding, fully as long as palpal segments. Ovipositor sheath fully 0-66 times aslong as gaster. Cf . Bristles of palpi of ordinary form, much shorter than palpal segments. MATERIAL EXAMINED9 cf , 20 $. All from southern Europe. Tobias (1976) gives the range as Palaearctic. HOST. I have seen no bred material. Tobias (1976) gives Cydia (Grapholita) pomonella L. (Tortricidae),Eurrhypara hortulata (L.) (Pyralidae) and Conopia [Syanthedon] spheciformis (Denis & Schiffermiiller)(Sesiidae) as hosts, which include such a wide range of lepidopterous families that I doubt if all can becorrect. COMMENTS. The single European species can be easily recognised by its colour and by the long palpalbristles of the female. AGATHIS Latreille Agathis Latreille, 1804: 173 [no included species] ; 1805: 175. Type-species: Agathis malvacearum Latreille,by subsequent monotypy. DIAGNOSIS. Mostly black species, rarely entirely or partly marked with red. Head in facial view almostalways elongate; if not markedly so then still clearly triangular. Ocelli always in a high triangle, theposterior tangent to the anterior ocellus not touching or cutting posterior pair. Antenna in the speciesunder review never with more than 40 segments. Mouth parts characteristically lengthened, forming abeak. Notaulices almost always present. Propodeum without areolation but with two more or less distinctmedial longitudinal keels; surface on each side of these keels usually smooth, polished; rarely surface ofpropodeum rugose all over. Spurs of hind tibia short, always less than half as long as basal segment of hindtarsus; middle tibia with at least two teeth on its outer side in about apical third. Hind claw with or without alobe or tooth but never cleft; each of the posterior metasternal foramena that receive the insertion of thehind coxae open on its inner side. First discoidal cell and first cubital cell confluent; 2nd cubital cell always A REVISION OF THE EUROPEAN AGATHIDINAE 193 small, most triangular, rarely obviously 4-sided. First tergite varying from entirely smooth to rugose-striate; tergites 2+3 rarely striate all over (semiaciculata Ivanov); sometimes with variable amount ofbroken rugose-striation distal to, and to sides of, faint, tranverse, blister-like swelling. Ovipositor at leastas long as gaster. Key to species (females) 1 Antennal sockets separated on inner side by distance equal to diameter of anterior ocellus. Species strongly marked with red; galea short, 1-5 times longer than wide syngenesiae Nees von Esenbeck(p. 196) - Antennal sockets united on inner side to form a single keel 2 2 At least mesoscutum in greater part, or entirely, red 3 - Mesoscutum black 5 3 Head in dorsal view strongly produced backwards behind eyes; temples strongly swollen (Fig. 26) malvacearum Latreille (p. 202) - Head in dorsal view less produced backwards behind eyes ; temples not , or hardly , swollen 4 4 Head entirely black; galea 3-5 times longer than wide; frons in front of anterior ocellus jutting forwards and showing as a V- or U-shaped cavity; malar space slightly shorter than longerdiameter of eye; areolet of fore- wing 4-sided; ovipositor sheath as long as body umbellatarum Nees von Esenbeck (p. 197) - Head at least almost entirely red; galea 3 times longer than wide; frons in front of anterior ocellus not jutting forwards and at most with shallow dimple here; malar space distinctlylonger than longer diameter of eye, 4:3; areolet of fore- wing stalked; ovipositor sheathabout equal to length of gaster glaucoptera Nees von Esenbeck (p. 195) 5 Mouth-parts much lengthened ; galea at least 4 times longer than its basal width 6 - Mouth-parts less lengthened ; galea at most 3 5 times longer than its basal width 8 6 Hind claw with strong lobe (Fig. 47); notaulices deeply impressed; at most a shallow dimple in front of anterior ocellus 7 - Hind claw without a lobe though a little, and abruptly, widened at base (Fig. 45). Head in facial view weakly elongate (Fig. 11); galea 4-5 times longer than basal width;notaulices often weak, sometimes almost obliterated nigra Nees von Esenbeck (p. 203) 7 Galea 6 times longer than its middle width; outer side of middle tibia with about 30 teeth and appearing densely spinose taurica Telenga (p. 204) - Galea 4 times longer than its middle width; outer side of middle tibia with 6-8 teeth, some of them in pairs zaykovi sp. n. (p. 204) 8 Hind femur greatly lengthened, about 4-5 times longer than wide. Galea very short, only a little longer than wide gracilipes Hellen (p. 212) - Hind femur not more than 3 times longer than wide 9 9 In front of anterior ocellus a V- or U-shaped, or parallel-sided, cavity, bounded by a ridge and prolonged below to form a keel between antennal sockets 10 - In front of anterior ocellus at most a shallow, subtriangular impression or dimple, or surface virtually flat; virtually no keel between antennal sockets or, if one is weakly indicated, then it is not continuous with any ante-ocellar impression that may be present 23 10 Galea very short , inconspicuous, not more than 1-33-1-50 times longer than wide 11 - Galea longer, at least twice as long as wide 13 11 Small species, 2-8-3-0 mm; head in facial view subtriangular (Fig. 29). Impression in front of anterior ocellus shallow; distal abscissa of postmarginalis hardlyshorter than proximal abscissa minuta Niezabitowski (p. 208) - Larger species, at least 4-0 mm; head in facial view markedly elongate. Hind claw with conspicuous lobe; distal abscissa of postmarginalis about 0-75 times longerthan proximal abscissa 12 12 Ovipositor sheath not longer than gaster, about 1-5 times longer than hind tibia; flagellum somewhat thickened medially and thence tapering to apex, not markedly bristly breviseta Nees von Esenbeck (p. 197) Ovipositor sheath very distinctly longer than gaster and about twice as long as hind tibia;flagellum thinner, markedly bristly. Cavity in front of anterior ocellus deeper and more nearly parallel-sided than in breviseta assimilis Kokujev(p. 198) 194 G. E. J. NIXON 13 Enclosed cavity in front of anterior ocellus U-shaped, almost jutting forwards as though on a prominence; stigma longer; radial cell longer 14 Enclosed cavity in front of anterior ocellus only vaguely V- or U-shaped, not jutting forwardsbut lying in same plane as keel below it; stigma shorter; radial cell shorter. Distal abscissa of postmarginalis at least 0-66 times longer than proximal abscissa; hindclaw with conspicuous lobe 15 14 Head in dorsal view strongly produced backwards behind eyes; temples strongly swollen (Fig. 26); head in facial view much lengthened (Fig. 4); hind claw with basal tooth malvacearum Latreille(p. 202) Head in dorsal view less produced backwards behind eyes; temples hardly swollen; head infacial view much less lengthened, subtriangular (Fig. 16); hind claw with large basal lobe varipes Thomson (p. 199) 15 Notaulices virtually obliterated, at most a hardly visible line marking their course. Hind claw with distinct lobe 16 Notaulices clearly defined 17 16 Galea distinctly longer than malar space; a hardly visible line indicates course of notaulices; outer side of middle tibia with 8-9 irregularly spaced, thick teeth; thorax more elongate in profile polita sp. n. (p. 206) Galea shorter, hardly as long as malar space; notaulices absent; outer side of middle tibiawith 4-5 teeth arranged more or less in row; thorax less elongate in profile persephone sp. n. (p. 206) 17 Segment 3 of middle tarsus very short, about 1-25 times longer than wide. Tergite 1 sculptured right to apex 18 - Segment 3 of middle tarsus at least 1-5 times longer than wide 19 18 Ovipositor sheath about as long as gaster; head in facial view not strikingly lengthened achterbergi sp. n. (p. 208) Ovipositor sheath nearly 1-5 times longer than gaster; head in fcial view strikingly lengthened(Fig. 6) anchisiades sp. n. (p. 207) 19 V-shaped cavity in front of anterior ocellus narrow, deep. Outer side of middle tibia with 7-8 teeth rufipalpis Nees (p. 199) V-shaped cavity in front of anterior ocellus less narrow, somewhat shallow 20 20 Thorax in profile more elongate (Fig. 42); outer side of middle tibia with 6-10 closely spaced teeth. Malar space very distinctly shorter than longer diameter of eye 21 Thorax in profile much less elongate; outer side of middle tibia with 4-5 teeth arranged more orless in a row. Distal abscissa of postmarginalis obviously shorter than proximal abscissa 22 21 Galea clearly longer than malar space, polished and virtually smooth; hind claw with conspi- cuous lobe and with deep cleft between lobe and claw proper; 3-4 distal segments ofmaxillary palpus markedly yellowish; distal abscissa of postmarginalis virtually equal to proximal abscissa fulmeki Fischer (p. 198) Galea clearly shorter than malar space, dull, coriaceous; hind claw with only weak basalprojection and no cleft between this and claw proper; maxillary palpus blackish throughout;distal abscissa of postmarginalis a little shorter than proximal abscissa pedias sp. n. (p. 211) 22 Head in facial view wide across clypeus (Fig. 1), longer; hind femur less swollen; antenna with 26 segments; tergite 1 with clearer indication of sculpture over apical two-thirds ariadne sp. n. (p. 206) Head in facial view much less wide across clypeus, shorter, somewhat triangular; tergite 1highly polished over apical two-thirds; hind femur more swollen (Fig. 33); antenna with atmost 23 segments tibialis Nees von Esenbeck (p. 201) 23 Head in facial view hardly lengthened below eyes (Fig. 12); clypeus in lateral view of head markedly protuberant. Wings hyaline; mandibles of powerful build; hind claw without lobe pappeisp. n.(p. 212) Head in facial view always lengthened below eyes (sometimes nearly subtriangular); clypeus inlateral view of head not protuberant 24 24 Sternaulus reaching both anterior and posterior margin of mesopleurum. Galea about 3 times longer than its widest part, somewhat abruptly narrowed in apicalhalf; hind claw virtually without lobe rostrata Tobias (p. 211) - Sternaulus, if present, not reaching both anterior and posterior margin of mesopleurum or, if it A REVISION OF THE EUROPEAN AGATHIDINAE 195 almost reaches anterior margin, then galea shorter (semiaciculatd) or hind claw withconspicuous lobe (melpomene) 25 25 Hind claw virtually without lobe, at most a denticle at apex of thickened, basal part. Distal abscissa of postmarginalis very short ; tergite 1 sculptured right to apex 26 Hind claw with lobe; if weak (anglicd), then areolet markedly 4-sided 29 26 Sternaulus absent or represented by a hardly impressed furrow; thorax elongate, almost twice as long as wide, 44 : 25, as seen in profile.Species at most 2-5 mm excluding ovipositor; galea very distinctly longer than malar space, 7 : 4; flagellum distinctly thickened beyond middle asteris Fischer (p. 209) Sternaulus clearly defined; thorax less elongate, at most 1-66 times longer than wide as seen inprofile 27 27 Galea shorter than malar space 28 - Galea as long as, or slightly longer than, malar space. Face rather sharply narrowed below eyes in facial view (Fig. 28); ovipositor sheath aboutas long as propodeum plus gaster; basal, thickened part of hind claw sometimes with denticleat apex glabricula Thomson (p. 209) 28 Flagellum distinctly thickened distal to middle and thence tapered to apex; segment 10 from apex about 1-25 times longer than wide; head seen from above, with face just out of view,longer.Ovipositor sheath short, about as long as gaster artemesiana Fischer (p. 210) - Flagellum not thus thickened distal to middle and not tapered to apex; segment 10 from apex fully 1-5 times longer than wide; head seen from above, with face just out of view, shorter,more transverse meridionellae Fischer (p. 210) 29 Face densely pubescent. Hind femur entirely, or in greater part, reddened; hind claw with well-developed tooth(Fig. 44); virtually no dimple in front of anterior ocellus and no keel between antennalsockets griseifrons Thomson (p. 202) - Face with ordinary pubescence 30 30 Length at most 3-5 mm excluding ovipositor 31 - Length at least 4-0 mm excluding ovipositor 32 31 Thorax less elongate (Fig. 41); hind femur blackish; lobe of hind claw in form of weak projection or angulation ; antenna with 27-29 segments glabricula Thomson (p. 209) - Thorax more elongate; hind femur entirely reddish; lobe of hind claw in form of free projection; antenna with 23-24 segments montana Shestakov (p. 213) 32 Basal half of tergites 2+3 closely striate all over semiaciculata Ivanov (p. 205) - Basal half of tergites 2+3 with at most very irregular striate-rugosity 33 33 Hind claw with conspicuous lobe; radial cell longer (Fig. 50) 34 Hind claw with at most a weak, basal angulation; radial cell shorter (Fig. 49) Head in facial view much narrowed below eyes (Fig. 18); tergite 1 sculptured right to apex;tergites 2+3 usually with conspicuous, very irregular striate rugosity distal to basal,transverse swelling anglica Marshall (p. 200) 34 Head in facial view subtriangular (Fig. 16); galea about as long as malar space varipes Thomson (p. 199) - Head in facial view much narrowed below eyes; galea 1-33 to 1-50 times longer than malar space melpomene sp. n. (p. 213) Agathis glaucoptera Nees von Esenbeck (Fig. 3)Agathis glaucoptera Nees von Esenbeck, 1834: 128. Holotype ?$ , ITALY (lost). $, 7-8 mm long (excluding ovipositor). Almost entirely reddish yellow; ocellar region and scrobesblackish; mesosternum, mesopleurum and propodeal area black. Hind coxa varying from black to brightreddish yellow, like gaster; hind femur entirely reddish yellow. Head in facial view markedly elongate; in this aspect differing from all other included species in that theface is only weakly narrowed towards the mouth (Fig. 3). Malar space about 1-25 times as long as longerdiameter of eye. Inner rims of antennal sockets joined to form a keel that unites with a shallow, V-shapedimpression in front of anterior ocellus. Anterior ocellus raised as though on a prominence. Antenna 31-33segmented. Galea hardly twice as long as wide. Thorax in profile of generalised form. Notaulices deep but 196 G. E. J. NIXON not costate. Sternaulus deep, wide, strongly costate. Propodeum without clearly defined, longitudinal keelbut with several coarse rugae in their place. Areolet of fore-wing small, stalked; stigma relatively elongate;distal abscissa of postmarginalis at least 0-66 times longer than proximal abscissa; anellus of hind wingjoining anal cell above middle . Hind claw with large , conspicuous lobe ; outer side of middle tibia with 1-2 ,or without , teeth ; inner spur of hind tibia powerful , reaching middle of basal segment of hind tarsus . Gasterrather short. Tergite 1 about 1-33 times longer than wide at apex, smooth, polished. Ovipositor sheathabout as long as gaster. MATERIAL EXAMINED France: 1 $, Perpignan, 10.vi.1976 (G. D. Slob) (RNH). Turkey: 1 $, Konya, ll.vi.1971 (Kl. Warnke)(RHN). Yugoslavia: 1 $, Macedonia, R. Reshik, (/. Waterston) (BMNH). HOST. Unknown. COMMENTS. Marshall (1890) redescribed glaucoptera from a single female in poor condition but beyonddoubt recognised the species correctly. Both Telenga (1955) and Tobias (1954; 1963) were in agreementwith Marshall and these three authors have been followed here. A. glaucoptera is distinguished from allother species from the region by the shape of the head in facial view, in combination with the stalkedareolet and the short ovipositor sheath. The point of emission of the anal vein in the hind wing isnoteworthy and suggests that glaucoptera occupies a somewhat isolated position among the Europeanspecies ofAgathis. Agathis syngenesiae Nees von Esenbeck (Figs 5, 20)Agathis syngenesiae Nees von Esenbeck, 1814: 192, 194. Syntypes, GERMANY (lost). Cf $ , 4-5-5-6 mm (excluding ovipositor). $, Body much marked with red; head varying from almost black to almost entirely red. Prothorax,mesoscutum, propodeum and mesosternum entirely black. Gaster almost entirely red. Head in facial view markedly elongate (Fig. 5); seen from above, strongly transverse and only shallowlyemarginate behind (Fig. 20). Malar space equal to, or slightly longer than, longer diameter of eye. Mouthparts short for genus; galea hardly 1-5 times longer than wide. Antennal sockets not united on inner side toform a single keel but separated by a distance fully equal to diameter of anterior ocellus. Antenna with25-27 segments. Thorax moderately elongate, in profile as 13 : 8. Notaulices generally rather shallow.Sternaulus usually distinct but sometimes almost wanting, not extending beyond middle of mesopleurum.Sculpture of propodeum varying from coarsely reticulate all over to vaguely punctate-reticulate; the twolongitudinal keels ofAgathis (s.str.) virtually wanting. Areolet of fore-wing triangular, almost petiolate;stigma more elongate than that of other species of region ; radial cell very narrow ; radius sometimes slightlycurved inwards towards stigma; distal abscissa of postmarginalis as long as proximal abscissa. Hind clawwith somewhat short, dentiform lobe; outer side of middle tibia with 2-3 widely spaced teeth; inner spur ofhind tibia falling far short of middle of basal segment of hind tarsus. Tergite 1 about 1-25 times longer thanits apical width, polished, smooth. Ovipositor sheath 1-33-1-5 times longer than body. Hypopygium veryshallowly, broadly emarginate at apex. Cf. Inner and outer orbits pale marked (2 ex., Turkey), otherwise generally black. Posterior half ofgaster blackened. Antenna with 25-26 segments. MATERIAL EXAMINED France: 1 $, Carpentras (Teunissen) (RNH); 2 $, 1 cf, Landes, St Girons Plage, on Helychrysumstoechas L. (P. Pronk) (RNH). Germany: 3 $ , Ruthe Coll. (BMNH). Holland: 1 $ , Terschelling (RNH); 1$ , Texel De Koog, pine forest, 6.ix. 1969 (L. Oosterweghel) (RNH). Portugal: 1 $ , Algarve, Val do Lobo,vii. 1967 (P. M. F. Verhoeff) (RNH). Spain: 1 $ , Cadiz, Jerez de la Frontera, viii. 1967 (P. M. F. Verhoeff)(RNH). Turkey: 1 cf , Uludag, viii. 1962 (Guichard & Harvey) (BMNH); 2 $ , 1 cf , Bursa, near Karacabey,viii.1962 (Guichard & Harvey) (BMNH). HOST. Unknown. COMMENTS. This species closely resembles umbellatarum in colour but the two species are very different.Whereas umbellatarum is a typical Agathis, syngenesiae is in some respects like species of Microdus. Theform of the keel between the antennal insertions approaches the condition found in Microdus. So also doesthe strongly transverse head as seen from above. Nevertheless, the elongate head and the lengthened A REVISION OF THE EUROPEAN AGATHIDINAE 197 mouth-parts relate syngenesiae more closely to Agathis. The form of the frons between the antennalinsertions together with the short galea make the species easy to recognise. Agathis umbellatarum Nees von Esenbeck Agathis umbellatarum Nees von Esenbeck, 1814: 195. Syntypes, GERMANY (lost).Agathis kolazyi Fischer, 1959: 2. Holotype cf , YUGOSLAVIA (ZM) [examined]. Syn. n. Cf 9 , 4-5-5-0 mm long (excluding ovipositor). In all material available the body is conspicuously markedwith red. Head black. Mesoscutum, and at least side of pronotum, red. Gaster varying from entirely red tored with blackened apex. Wings strongly darkened. 9- Head in facial view moderately lengthened; from above, strongly transverse, not producedbackwards behind eyes as in malvacearum (cf. Fig. 26). Ocelli in a low, rather wide triangle. A moderatelywell-defined keel between antennal insertions; in front of anterior ocellus a well-defined excavation thattends to jut forwards. Malar space a little shorter than longer diameter of the eye, 11:14. Galeaconsiderably longer than malar space, 14 : 11. Antenna with 23-26 segments: 23 (1), 24(2), 25 (5), 26 (3);flagellum somewhat tapered towards apex. Thorax in profile decidedly elongate. Notaulices sharplydefined. The two longitudinal keels of propodeum sharply defined; side panels polished, virtually smoothall over. Radial cell decidedly long; areolet varying from triangular to distinctly 4-sided; distal abscissa ofpostmarginalis hardly 0-66 times as long as proximal abscissa. Inner spur of hind tibia falling considerablyshort of middle of basal segment of hind tarsus; outer side of middle tibia with very variable number ofteeth, 2-14; hind claw with well-defined lobe; cleft between lobe and claw proper fully half as long as clawitself. Tergite 1 a little longer than apically wide, smooth, highly polished. Ovipositor sheath very slightlylonger than body. Cf . Thorax sometimes entirely black; sometimes mesoscutum red. Antenna with 23-24 segments; twopreapical segments about 1-33 times longer than wide. MATERIAL EXAMINED Bulgaria: 4 cf , 1 $, v.-vi. (Zaykov) (ZC). Cyprus: 21 cf , 5 $, iv.-vi. (G. Mavromoustakis) (BMNH).France: 2 $, Basses Alpes, Digne, vi. (Slob) (RNH). Greece: 1 $, Corfu, Acharavi, v. (Oogstrom)(RNH); 1 , Kerfissos, vi. (Mavromoustakis) (BMNH); 1 9, Drosla, vi. (Mavromoustakis} (BMNH); 19, Ilia, vii. (Day & Else) (BMNH); 1 9, 2 cf , Thessalia, Kalambaka, vii. (Day & Else) (BMNH). Turkey:1 9, Ankara, Temelli, vii. (Guichard & Harvey) (BMNH); 1 9, Nigde, Kocas, vi. (Guichard & Harvey)(BMNH); 10 cf, Istanbul, Yakuplu, vi. (Oorshot & Wiering) (RNH). Yugoslavia: 1 <j>, Macedonia,Gorica, x. (van Achterberg (RNH); 1 cf (holotype of kolazyi), Dalmatia, Ragusa (Kolazy) (ZM). HOST. Unknown. COMMENTS. This brightly coloured species has a Mediterranean distribution. It is very like the northernvaripes Thomson and, beyond colour, there is very little to separate the two species. Both are characterisedby the form of the ante-ocellar impression and the shape of the head in facial view. Thus seen, however, theface of varipes is slightly wider; varipes also has a shorter galea. Agathis breviseta Nees von Esenbeck (Fig. 52)Agathis breviseta Nees von Esenbeck, 1814: 194. Syntypes, GERMANY (lost). The type of breviseta is presumably lost. My interpretation of the species is based on that of Wesmael, thefirst reviser; I have examined two females (IRSNB), both bearing Wesmael's identification label but withno indication of locality. 9, ca 3-5 mm long (excluding ovipositor). Black. Hind femur black; hind tibia blackish on apicalthree-fifths, with dark ring towards base. Head in facial view considerably elongate. Malar space shorter than longer diameter of eye, 5 : 7. Galeadull, coriaceous, twice as long as wide. Between antennal sockets a short, knife-edged keel that projectsabove as a right angle before joining lowest point of deep cavity in front of anterior ocellus; this cavity deep,narrow, V-shaped. Ocelli in high triangle, the posterior tangent of front ocellus passing far in front ofposterior ocelli. Antenna distinctly tapering from middle to apex, weakly bristly and with the two preapicalsegments about 1-33 times longer than wide; 29-30 segmented. Thorax in profile short, high. Notaulicesdeeply impressed. Sternaulus strongly impressed, reaching both anterior and posterior corner of meso-pleurum. Lateral panels of propodeum polished. Fore- wing with areolet almost always distinctly 4-sided 198 G. E. J. NIXON (17 ex.); rarely almost triangular (4 ex.); apical abscissa of postmarginalis slightly more than 0-5 timeslonger than proximal abscissa. Hind femur decidedly thick, 2-5 times longer than widest part; outer side ofmiddle tibia with 6-7 teeth; inner spur of hind tibia just reaching middle of basal segment of hind tarsus;hind claw with well-developed, somewhat pointed lobe. Gaster rather short and broad; segment 1 markedly triangular, as long as wide apically , irregularly striateto striate-rugose all over. Ovipositor sheath short, not longer than gaster; bristle-like hairs of apical third ofovipositor sheath not, or hardly, different from those of middle third (Fig. 52). MATERIAL EXAMINED Belgium: 2 $, Wesmael coll. (IRSNB). Bulgaria: 23 $, Sh. poljana, 6-22.V.1976 (Zaykov) (ZC). GreatBritain: 2 $, England, Buckinghamshire, vi-vii. (Benson) (BMNH). Ireland: 1 $, Antrim, Loch Neagh,26.vi.1967 (Stelfox) (USNM); 1 cf , 1 $, Dublin, Glenasmole, 24.vi.1938 (Stelfox) (USNM). Turkey: 1 $,Edirne, S.v.1960 (Guichard & Harvey) (BMNH). Yugoslavia: 1 $, Slovenia, Kropa (Ward) (BMNH). HOST. Unknown. COMMENTS. This species is fairly easy to recognise on the combination of short galea and unusually shortovipositor sheath. In his description Nees draws special attention to the latter feature. Agathis assimilis Kokujev (Figs 2, 19, 25)Agathis assimilis Kokujev, 1895: 387. Holotype $, U.S.S.R. (AS) [not examined]. Tobias (1963: 878) examined the type of assimilis which, he says, is badly damaged, lacking wings, legs(except one hind leg) and antennal tips. Specimens determined by him as assimilis have been received onloan at BMNH; his interpretation of Kokujev's species is accordingly accepted. $ . Closely related to breviseta on the structure of the frons (Fig. 19) and the shortness of the galea. It maybe compared with that species as follows. Colour similar. Head in facial view (Fig. 2); seen from above, slightly more transverse, less producedbackwards behind eyes (Fig. 25); this is correlated with a slightly lower ocellar triangle. Antenna thinner,filiform; two apical segments relatively a little longer; 28-31-segmented; flagellum considerably morebristly. Sternaulus reaching posterior margin of mesopleurum but not anterior margin. Areolet offore- wing variable to same degree; apical abscissa of postmarginalis slightly longer. Legs more slender;hind femur about 3 times longer than wide; segment 3 of hind tarsus relatively a little longer; outer side ofmiddle tibia with 4-5 teeth, arranged more or less in row; inner spur of hind tibia slightly shorter, relative tolength of basal segment of hind tarsus. Tergite 1 slightly narrower, more shiny because of reduction instrength of sculpture; this vaguely coriaceous, with some weak striation on basal half. Ovipositor sheathmuch longer, about equal to propodeum plus gaster; as in breviseta, the apical, bristle-like hairs of thesheath are not differentiated from the rest. MATERIAL EXAMINEDBulgaria, Great Britain. HOST. Unknown. COMMENTS. This species and breviseta evidently form a small species-group characterised by the curiousformation of the inter-antennal keel, combined with the deep, narrow cavity in front of the anteriorocellus. The long, filiform antenna and the longer ovipositor readily distinguish assimilis from breviseta. One $ examined (Great Britain: Scotland, The Birks, Aberfeldy, 26.vii.1954 (Stelfox) (BMNH)) hasthe propodeum rugulose and dull almost all over. Agathis fulmeki Fischer (Figs 10, 35, 42, 51)Agathis fulmeki Fischer, 19576: 6. Holotype 9 AUSTRIA (NM) [examined]. $, ca 3-5 mm long (excluding ovipositor). Black. Apical four segments of maxillary palpus almost alwaysyellowish and in contrast with black labial palpus. Hind femur black; hind tibia without, or virtuallywithout, a dark basal ring. Head in facial view short, subtriangular (Fig. 10). Longer diameter of eye 2-5 times longer than malarspace. Surface between antennal insertions raised and ridge-like where it joins depression in front of A REVISION OF THE EUROPEAN AGATHIDINAE 199 anterior ocellus; this depressed area V-shaped, not deep, slightly domed at middle. Galea smooth, shiny,2-5 times longer than wide; labial palpus unusually long (Fig. 51). Clypeus largely polished, bare. Antennawith 25-27 segments; two preapical segments of flagellum hardly longer than wide. Thorax in profileelongate (Fig. 42), about twice as long as wide. Mesoscutum strongly shining, not densely hairy and oftenwith vague punctuation, mostly on middle lobe. Notaulices sharply defined but not deep, distinctlyfoveate. Sternaulus short, shallow, sometimes almost obliterated and not reaching either posterior oranterior margin of mesopleurum. Areolet of fore-wing triangular or almost so; distal abscissa ofpost-marginalis hardly shorter than proximal abscissa. Legs rather thick; outer side of middle tibia with 6-7teeth (Fig. 35) ; hind femur about 2-5 times longer than wide ; inner spur of hind tibia just reaching middle ofbasal segment of hind tarsus; hind claw with conspicuous lobe, large, strongly bent and with deep cleftbetween itself and claw proper. Tergite 1 slightly longer than wide at apex, shiny, with faint traces ofrugosity and weak indication of striation laterally. Ovipositor sheath about as long as gaster pluspropodeum; seen from above, hairs of distal, slightly widened part of sheath considerably shorter and lessnoticeable than those proximal to it. MATERIAL EXAMINED Austria: 1 $ (holotype), Modling-Vorderbriihl (NM). Bulgaria: 48 $, Rhodopi, vi-vii. (Zaykov) (ZC;17$ in BMNH). COMMENTS. A most distinctive species, recognisable on combination of narrow thorax, short head andteeth on outer side of middle tibia. It differs from breviseta and assimilis in that the galea is longer and thedecoration of the frons, though basically similar, has no knife-like edge between the antennal insertions. Agathis rufipalpis Nees von Esenbeck (Figs 17, 36)Agathis rufipalpis Nees von Esenbeck, 1814: 192. Syntypes, GERMANY (lost). Wesmael (1837: 24) was the first reviser of Nees von Esenbeck's species; I have based my interpretation ofrufipalpis on specimens in Wesmael's collection bearing his handwritten identification label. This species is extremely likefulmeki, differing from it only in the following characters.$ . Four apical segments of maxillary palpus sometimes paler than labial palpus. Head in facial view slightly more elongate (Fig. 17). Longer diameter of eye about 1-75 times longer thanmalar space. Clypeus hairy all over. Depression in front of anterior ocellus more narrowly V-shaped.Antenna with 24-28 segments. Thorax in profile slightly less elongate (Fig. 36). Sternaulus equally shortand weak. Outer side of middle tibia with 7-9 similarly thick teeth; inner spur of hind tibia distinctly lessthan half as long as basal segment of hind tarsus. Tergite 1 more distinctly sculptured and with somewhatbroken striation almost all over. Ovipositor sheath about as long as gaster plus thorax. MATERIAL EXAMINED Belgium: 5 $, Brussels (1 $ BMNH, rest IRSNB). Bulgaria: 10 $, Rhodopi, vi.-vii. (A. Zaykov) (ZC).Ireland: 1 cf, Wexford, Curracloe, vii. (BMNH). Sweden: 1 $, Solna, viii.1976 (T. Huddleston &J. Quinlan) (BMNH). HOST. Unknown. COMMENTS. This species is very much likefulmeki and the two should always be considered together. Itdiffers from breviseta and assimilis by the same characters as those which separate these species fromfulmeki. Agathis varipes Thomson (Figs 16, 50)Agathis varipes Thomson, 1895: 2228. LECTOTYPE $, SWEDEN (ZI), here designated [examined]. Cf $ , 5-0-5-5 mm long (excluding ovipositor). $ . Black. Gaster tending to be brownish. Hind femur varying from mostly black to entirely brownish red(1$, Italy). Head in facial view subtriangular (Fig. 16). Impression in front of anterior ocellus variable in definition;sometimes sharply U-shaped and jutting forwards as though on a prominence; sometimes showing merelyas a U-shaped, margined depression (in the related tibialis, this depression tends to be V-shaped). Ocelli ina rather low triangle. Malar space 0-66 times as long as longer diameter of eye. Galea rather short, shorter 200 G. E. J. NIXON than in related tibialis, 2-25-2-33 times longer than wide, not longer than malar space. Antenna with 23-26segments: 23(2), 24(5), 25(1), 26(1). Thorax in profile somewhat elongate, fully 1-5 times longer than wideseen in profile. Notaulices somewhat variable in definition; in one bred male (Surrey, Effingham), they arehardly indicated. Sternaulus usually not reaching posterior corner of mesopleurum. Medial, longitudinalkeels of propodeum sharply defined; lateral panels extensively polished. Areolet of fore wing usuallysharply triangular; radial cell rather long (Fig. 50); distal abscissa of postmarginal is usually distinctly morethan half as long as proximal abscissa. Outer side of middle tibia with 2-5 teeth arranged more or less in arow; hind claw with strong, well-defined lobe; inner spur of hind tibia not quite reaching middle of basalsegment of hind tarsus. Gaster on the whole having a smooth, highly polished appearance. Basal half oftergite 1 sometimes with indication of weak rugosity. No trace of sculpture posterior to the hardly indicatedbasal swelling on tergites 2+3. Ovipositor sheath about as long as body. Cf . Two preapical segments of antenna fully 1-5 times longer than wide; whole antenna considerablylonger than in tibialis. Hind femur much less thickened (cf. Fig. 33), infuscate but sometimes reddishtowards apex. MATERIAL EXAMINED Germany: 2 $, 2 cf, no locality, ex Metzneria lappella (L.) (BMNH); 1 9, Goslar, Grauhofer,10.viii.1944 (Bauer) (EH). Great Britain: 1 $, Devon, E. Bovey, viii (Perkins) (BMNH); 2 $, 1 cf,Hampshire, Farley Mount, vi-vii. , ex Myelois cirrigerella (Zincken) (Fassnidge) (BMNH) ; 1 $ , Hertford-shire, Tring, vii (Benson) (BMNH); 1 $, Kent, Erith, swept from Achillea (Ford) (BMNH); 1 cf,Cambridgeshire, Wicken Fen, 9. vii. 1982, ex microlepidopteron collected 16. iv. 1982 on Salix (Pitkin)(BMNH); 2 $ , 2 cf , Surrey, Coulsdon, vii. 1944, ex stems ofArctium medius (Arctium llappa L.) collectediii.1944 (Britten) (BMNH); 1 <j>, 3 cf, Surrey, Effingham Common, vii. 1946, ex Metzneria lappella (L.)collected iii.1946 (Niblett) (BMNH); 1 $, Surrey, Oxshott, vi. (Billups) (BMNH). Sweden: 1 $ (lecto-type), Skane, 'Fg' (= Fogelsang) (Thomson) (LI); 1 $, Ringsjo (Perkins) (BMNH). HOSTS. Metzneria lappella (L.) (Gelechiidae). Myelois cirrigerella (Zincken) (Pyralidae). COMMENTS. This species and tibialis are very much alike, the only reliable difference being the shape of theantero-ocellar impression and the thickness of the hind tibia. A. varipes is slightly larger than tibialis butthis may have little significance in the identification of single specimens. There is some overlap in thenumber of antennal segments. Agathis anglica Marshall(Figs 18, 49) Agathis anglica Marshall, 1885: 265. Holotype $, GREAT BRITAIN (BMNH) [examined].Agathis albanica Fischer, I951b: 3. Holotype $, ALBANIA (NM) [examined]. Syn. n. Cf $ , 4-5-5-0 mm long (excluding ovipositor). $ . Black. Hind femur entirely black; hind tibia yellowish on basal two-thirds, with dark basal band andblackened on apical two-fifths. Medius of fore-wing usually colourless throughout. Head in facial view markedly elongate and considerably narrowed below eyes (Fig. 18) ; seen from abovewith face just out of view strongly transverse. Hardly a trace of a keel between antennal sockets and with atmost a weakly impressed dimple in front of anterior ocellus. Malar space almost equal to longer diameter ofeye, 13 : 15. Galea about 3-5 times longer than wide and about 1-5 times longer than malar space. Antennawith 26-29 segments, slightly tapered towards apex and with segment 4 from apex about 1-25 times longerthan wide. Thorax in profile somewhat short and high. Notaulices deeply impressed. Propodeum with theusual two longitudinal keels and with polished, lateral panels. Sternaulus well defined, reaching posteriorcorner of mesopleurum. Radial cell short, broad (Fig. 49); aerolet markedly, characteristically 4-sided;distal abscissa of postmarginalis very short, 1-25-1-33 times as long as proximal abscissa. Outer side ofmiddle tibia with 3-5 teeth arranged in a row; hind claw with at most a weakly angled projection at base.Tergite. 1 rugose-striate all over. Tergites 2+3 with well-defined, transverse, basal, blister-like swelling;surface posterior to this swelling with a considerable amount of striation or rugose-striation. Ovipositorsheath nearly as long as body. Cf . Like female except for sexual differences. In nine examples from Turkey there is considerablevariation in the definition of the impression in front of the anterior ocellus; sometimes it is relativelydistinct; sometimes virtually absent; in all these examples the areolet of the fore- wing is very obviously4-sided. Tergites 2+3 show much less rugosity than in the female and are sometimes virtually smooth. A REVISION OF THE EUROPEAN AGATHIDINAE 201 MATERIAL EXAMINED Albania: 1 $ (holotype ofalbanica), Kula Ljums (NM). Austria: 1 $, Achenkirch (Haeselbarth) (EH); 1$ , Fliess (Haeselbarth) (EH); 1 $ , Nordkette, 3. vii. 1947, ex Epinotia mercuriana (Frolich) (Bauer) (EH).Bulgaria: 1 $, Rhodopi, Aida, 5.vi.l976 (Zaykov) (ZC); 1 $, Popsko, 21. vi. 1977 (Zaykov) (ZC); 1 $, Sh.poljana, 18.vi.1976 (Zaykov) (ZC); 1 $ , Bojuo, 24.vii.1975 (Zaykov) (ZC). Cyprus: 27 $ , Mavromousta-kiscoll. (BMNH); 1 $,Platres, 16.vi. 1970 (Gallagher) (BMNH). Great Britain: 1 $ (holotype of anglica),Wales, Pembrokeshire, Milford Haven (BMNH); 1 $, England, Surrey, Hackhurst Downs, 28.viii.1982(Allen} (AA). Greece: 1 $, 2 cf, Rhodes, Ixia (Day) (BMNH). Italy: 1 $, Partschins, vii.1966(Haeselbarth) (EH). HOST. Epinotia mercuriana (Frolich) (Tortricidae). COMMENTS. It is possible that the range of specific variation accepted here for anglica may be too wide andthe material examined may form an aggregate. Nevertheless, all the specimens provisionally accepted asanglica have four rather striking features in common: the narrow head as seen from in front, theconspicuously 4-sided areolet of the fore- wing, the unlobed claws and the rugosity of the gaster. Agathis f/Wa/JsNees von Esenbeck (Figs 33, 48, 53)Agathis tibialis Nees von Esenbeck, 1814: 194. Syntypes, FRANCE (lost). Cf $ , ca 4 mm long (excluding ovipositor). $. Black with gaster usually brownish, especially tergites 2+3. Hind femur almost always brightbrownish red but becoming flushed with darker colouring on basal half. Wings strongly darkened; mediusof fore wing heavily brown throughout. Head in facial view, short, subtriangular. Impression in front of anterior ocellus narrow, well definedand, when head seen in profile, not projecting beyond anterior ocellus; this elongate, margined impressionunited in front with sharp, strongly raised keel between antennal insertions. Galea nearly 3 times as long aswide, shining, polished, distinctly a little longer than malar space. Malar space shorter than longerdiameter of eye. Antenna characteristically short, 21-22-segmented: 21 (8), 22 (3); four preapicalsegments of antenna only slightly longer than wide. Thorax in profile somewhat elongate, less than twice aslong as wide, 20: 13. Notaulices well defined. Sternaulus becoming somewhat indistinct before reachingposterior margin of mesopleurum. Propodeum highly polished on each side of the two longitudinal keels.Radial cell rather short, 5 times as long as wide; areolet of fore wing triangular to subtriangular, neverobviously 4-sided; distal abscissa of postmarginalis fully 0-66 times as long as proximal abscissa; medius offore-wing heavily sclerotised throughout. Hind femur much swollen (Fig. 33) ; inner spur of hind tibia muchless than half as long as basal segment of hind tarsus (Fig. 53); hind claw with conspicuous lobe. Tergite 1polished and virtually smooth over most of apical half; vaguely rugose-striate on posterior half; sometimesfaint scaly-reticulation present on apical, polished surface. Rest of gaster highly polished and shining.Ovipositor distinctly a little longer than body. Cf . Like female in having thickened, predominantly reddish hind femur. Antenna characteristicallyshorter than body, 21-23-segmented: 21 (6), 22 (7), 23 (1); two preapical segments about 1-33 times as longas wide. MATERIAL EXAMINED Great Britain: 13 $ , 28 cf , Devon, Dawlish Warren (Allen) (AA); 1 $ , Devon, Braunton Burrows, vii(Allen) (AA); 1 $ , Devon, Heathfield, viii (Allen) (AA); 1 $ , Dorset, Portland, collected from seed-headof Centaurea nigra L. (Hall) (BMNH); 2 $, Cambridgeshire, Wicken Fen, ex seed-head of Pulicariadysenterica L. , collected 29.xiii.1975, emerged 1976 (host either Apodia bifractella Douglas or Ptocheuusapaupella Zeller) (Emmet) (BMNH); 4 $, 3 cf, Surrey, Banstead, ex Aristotelia bifractella Douglas inflower-heads of Inula squarrosa L. , collected ix.1946, emerged viii. 1947 (Niblett) (BMNH); 25 $,25 cf,Surrey, Salfords, viii (Allen) (AA); 4 $, 21 Cf, Sussex, Hailsham (Ford) (BMNH). Holland: 1 $,Terschelling, near Rijsplak, 14. vii. 1967 (Pronk) (RNH); 2 $ , Terschelling, dunes near Doodemauskisten,6.vii.l967 (Pronk) (RNH). HOSTS. Apodia bifractella (Duponchel) (Gelechiidae) and possibly Ptocheuusa paupella (Zeller)(Gelechiidae), both feeding in flower-heads of Pulicaria dysenterica L. COMMENTS. The data indicate that this species flies from July to September, but mainly in August, andapparently is not uncommon where it occurs. What characterises the species essentially are the much 202 G. E. J. NIXON thickened, red-flushed hind femur, the short antenna in both sexes, the narrow, rather low, marginedimpression in front of the anterior ocellus, and the sharp interantennal keel. Agathis malvacearum Latreille (Figs 4, 26)Agathis malvacearum Latreille, 1805: 175. Holotype $, FRANCE (lost). Cf , $ , 5-7 mm long (excluding ovipositor). 9. Black with tergites 1 and 2+3 largely bright yellowish red. Hind femur reddish yellow; middle andfront femora reddish yellow but touched with black at base. Head in facial view strongly elongate (Fig. 4), eyes strongly bulging; seen from above, strongly producedbackwards behind eyes (Fig. 26). In profile a conspicuous bulge in region of temples. Malar space veryslightly shorter than longer diameter of eye, 16 : 17. Galea about 2-25 times longer than wide, distinctly alittle longer than malar space. Antenna with 21-32 segments. Between antennal insertions a fairly sharpkeel that unites above with a deep U-shaped cavity in front of anterior ocellus. Thorax of ordinary form.Notaulices deeply impressed. Sternaulus strongly defined , reaching posterior corner of mesopleurum. Sideof pronotum anterior to oblique trough polished and with hardly a trace of sculpture. Radial cell ratherlong; areolet distinctly 4-sided in material available; distal abscissa of postmarginalis slightly more than 0-5times as long as proximal abscissa. Outer side of middle tibia with 2-5 teeth arranged more or less in a row;inner spur of hind tibia not quite reaching middle of basal segment of hind tarsus ; hind claw with dentiform ,basal lobe; cleft between lobe and claw proper not particularly deep. Tergite 1 becoming smooth, polishedon about apical one-third; elsewhere with fairly distinct striation. Tergites 2+3 polished, unsculptured.Ovipositor sheath about as long as body. Cf . Like female in having tergites 2+3 brightly coloured; a single example from Corsica (BMNH) has themesoscutum as brightly reddish as the base of the gaster but each of the three lobes is suffused withinfuscation; a single male from Bulgaria, Rhodopi, (ZC) has the mesoscutum entirely reddish. MATERIAL EXAMINED Bulgaria: 1 9> Rhodopi, Petelovo, vii (Zaykov) (ZC). France: 1 cf, Basses Alpes, Digne, vi. (Slob)(RNH); 2 cf , 2 $ , Bretagne, Pres'ile de Rhuys, Kerfontaine, 1976, on Malva sp. (Doesberg) (RNH); 1 cf ,Corsica, vii. (BMNH). Greece: 1 $, Drosia, 4.vi.l957 (Mavromoustakis) (BMNH); 1 $, Kerfissos,8.vi.l957 (Mavromoustakis) (BMNH); 1 $, Epidavros, vi. (Mound) (BMNH). Turkey: 1 $, Mersin,Servatul Gecidi (Guichard & Harvey) (BMNH). HOST. Pexicopia malvella (Hiibner) (Gelechiidae). COMMENTS. Tobias (1964) emphasises the colour variability of this species on the basis of series bred fromthe Mallow moth in Armenia. He points out that autumnal specimens are darkest in colour while summerforms emerging in August and September are lighter with tergites 1-3 red and the hind femur alsoreddened. He also considers (1963: 876) that darker examples of malvacearum agree with the originaldescription of tibialis Nees von Esenbeck as well as with the interpretation of this latter species accepted byearlier authors. He accordingly suggests that tibialis Nees von Esenbeck may be a synonym of malvacearumLatreille. But in the absence of formal synonymisation I prefer to use the name 'tibialis Nees von Esenbeck'for a species that agrees better with Nees von Esenbeck's original description than any variety ofmalvacearum. The bulging temples of malvacearum are its most characteristic feature and the head in consequenceappears strongly produced backwards behind the eyes when seen from above. Agathis griseifrons Thomson(Fig. 44) Agathis griseifrons Thomson, 1895: 2227. LECTOTYPE $, SWEDEN: (ZI), here designated [examined].Agathis laticarpa Telenga, 1955: 255. Holotype $>, U.S.S.R.: Ukraine, Chernogavskaya Oblast (AS) [notexamined]. Syn. n. Cf $, 4-8-5-5 mm long (excluding ovipositor). Black. Legs predominantly reddish but colour variable;hind femur often entirely reddish but sometimes darkened on basal half, more rarely with infuscationextending along whole of upper surface; hind tibia often markedly yellowish, infuscate on about apicalthird, with a darkened basal ring. 9 Head in facial view obviously elongate. Malar space fully two-thirds as long as longer diameter of eye, A REVISION OF THE EUROPEAN AGATHIDINAE 203 11 : 15. Galea as long as malar space. Face densely clothed with greyish or pale brownish pubescence. Notrace of a keel between antennal insertions and virtually no impression in front of anterior ocellus. Antennawith 31-33 segments: 31 (1), 32 (8), 33 (3); flagellum rather finely tapered towards apex. Thorax in profileshort and of generalised appearance. Side of pronotum anterior to the oblique trough usually very coarselyrugose. Notaulices deeply impressed. Sternaulus strongly developed, reaching posterior corner of meso-pleurum. Areolet of fore-wing distinctly 4-sided; distal abscissa of postmarginalis about 0-33 as long asproximal abscissa. Hind claw with pale, dentiform lobe at base (Fig. 44); outer side of middle tibia with 1-3teeth arranged in a row. Gaster of ordinary form. Tergite 1 about as long as apically wide, striate-rugose allover. Frequently a certain amount of striation to side of, and apical to, transverse, blister-like, posteriorswelling. Ovipositor sheath about as long as gaster plus thorax. Cf . Like female except for sexual differences. Antenna very long, tapered apically, 29-32-segmented;segment 8 from apex twice as long as wide. Usually more brightly coloured than female with apical foursegments of maxillary palpus entirely yellowish. Tooth at base of hind claw often hardly developed. MATERIAL EXAMINED Bulgaria: 1 $, Rhodopi, Gabrovo, viii. (Zaykov) (ZC). France: 1 cf, Beaune, ex Pyrausta aurataScopoli on Mentha aquatica L. (BMNH); 1 $, Nantua, Marshall coll. (BMNH). Great Britain: 1 <j>, 1 cf ,Kent, Bexley, bred viii. 1952 ex Pyrausta aurata Scopoli (Ford) (BMNH); 3 $, Surrey, Bletchworth,23.viii.1983 (Allen) (AA); 1 $, Clandon Downs, vi. (Perkins) (BMNH); 1 cf , 1 $, Boxhill, vi. (Perkins)(BMNH). Greece: 1 $ , Philippi, v. (Guichard & Harvey) (BMNH). Holland: 1 $ , Maastricht, vi. (Lefebe)(RNH); 2 $ , Asperen, vi.-viii. (Zwakhals) (RNH). Ireland: 1 cf , 1 $ , Woodbrock, vi. (Stelfox) (USNM); 1 $ , Wicklow, Athdown, vii. (Stelfox) (USNM). Italy: 2 cf , 1 $ , Naples, viii. (Osborne) (BMNH). Poland: 2 $, Tatra Mts, Zakopane, vi. (Aubertin) (BMNH). Sweden: 6 <j>, Skane, 'sand-dunes' (ZI); 1 $(lectotype), Skane, labelled 'griseifrons' and 'coll. L-gh' (ZI). HOST. Pyrausta aurata (Scopoli) (Pyralidae). COMMENTS. In one of the two females from Holland, Asperen, tergite 1 is almost smooth over most of theapical half, and the dark ring at the base of the hind tibia is joined to the infuscate area at the apex of thetibia by a broad band of infuscation. Agathis griseifrons is largely characterised by the absence of a keel between the antennal insertions, thevirtual absence of any impression in front of the anterior ocellus, the dense pubescence of the face and theusually predominantly red-marked hind femur. Having a rugose first tergite, it is at once different fromthe superficially similar tibialis and varipes. Agathis nigra Nees von Esenbeck(Figs 11, 38, 45) Agathis nigra Nees von Esenbeck, 1814: 191. Syntypes, GERMANY (lost). Agathis testaceipes Fischer, 19576: 8. Holotype $, AUSTRIA (NM) [examined]. Syn. n. Wesmael (1837: 23) was the first reviser of Nees von Esenbeck's species; I have based my interpretation ofnigra on specimens in Wesmael's collection bearing his handwritten identification label. Cf $ , 4-5-4-8 mm (excluding ovipositor). Black, rarely tergites 2+3 reddish (1 9 , Yugoslavia). Hind femurreddish but flushed with infuscation on basal half; almost entirely reddish in 2 $ from Yugoslavia. $. Head in facial view not strongly elongate (Fig. 11). Malar space distinctly shorter than longerdiameter of eye, 9: 13. A weak or vestigial keel between antennal insertions. A more or less distinctV-shaped impression in front of anterior ocellus. Mouth parts very long; when fully extruded distinctlylonger than head; galea fully 4 times as long as wide, tapering to a fine point. Antenna 23-25-segmented: 23(4), 24(2), 25 (4); two preapical segments about 1-33 times longer than wide. Thorax in profile markedlyelongate (Fig. 38). Mesoscutum having a black, shiny appearance owing to sparseness of pubescence.Notaulices varying much in definition but usually weakly defined, at least anteriorly; virtually absent in 1 $(Yugoslavia, Korab Mts); frequently obliterated in posterior half in other females. Sternaulus usuallyreaching posterior corner of mesopleurum, rarely fading out before this. Propodeum showing a slightpeculiarity in that each lateral panel shows on its dorsal surface a broad, transverse band of very coarserugosities. Areolet of fore-wing always very obviously triangular; in two females even with short stalk;distal abscissa of postmarginalis almost as long as proximal abscissa. Inner spur of hind tibia falling far shortof middle of basal segment of hind tarsus ; outer side of middle tibia with 1-3 teeth arranged more or less in arow; in one female without teeth; hind claw with at most a small projection at base (Fig. 45). Gaster highlypolished; tergite 1 virtually without trace of sculpture. Ovipositor sheath about as long as body. 204 G. E. J. NIXON Cf. Like female except for sexual differences. Antenna with 23-24 segments; flagellum very thin,thread-like, the two preapical segments almost twice as long as wide. MATERIAL EXAMINED Austria: 1 $ (holotype oftestaceipes), Burgenland, Schiitzen, 10.vii.1941 (Fulmek) (NM). Belgium: 7 $,6 cf , viii. (Crevecoeur) (IRSNB); 1 cT, Brussels, Wesmael coll. (IRSNB). Germany: 1 $, Mecklenburg(Konow) (BMNH). Yugoslavia: 1 <j>, Korab Mts, Stirovica, viii. (Martina) (BMNH); 1 $, Nicpur, vii.(Martina] (BMNH). HOST. Unknown. COMMENTS. This is one of the more distinct species, largely characterised by the much lengthened galea,absence of lobe on hind claw and especially by the almost obliterated notaulices. The two other species withlong galea included in this revision are taurica and zaykovi, but both can easily be separated from nigra. A single female from Spain: Navarra, Peralta (Achterberg) (RNH) may possibly represent a furtherspecies. In all important respects it resembles nigra - short head and absence of lobe on hind claw - but thegalea is longer, nearer to six times longer than basal width, and the notaulices are sharply defined. I havelabelled this specimen as '? nigra Nees or sp. n.'. Agathis zaykovi s$. n. (Figs 7, 34, 47) O" $ , ca 4 mm (excluding ovipositor). Black. Hind femur black throughout. $. Head in facial view considerably lengthened (Fig. 7). Malar space 0-66 times as long as longerdiameter of eye. Between antennal insertions virtually no trace of a keel. A weak, vague V-shapedimpression, sometimes almost obliterated, in front of anterior ocellus. Galea tapered towards apex but notso evenly as in nigra, about 3-5 times longer than its basal width (Fig. 34). Antenna with 26-27 segments; 26(8), 27 (4); apical 5-6 segments of flagellum somewhat tapered; two preapical segments hardly longer thanwide. Thorax in profile somewhat elongate but less so than in nigra (cf . Fig. 38). Notaulices costate, deeplyimpressed throughout. Sternaulus strongly defined, costate throughout and reaching posterior corner ofmesopleurum. Areolet of fore- wing usually slightly narrowed above but always distinctly 4-sided; distalabscissa of postmarginalis hardly shorter than proximal abscissa. Hind claw with large lobe (Fig. 47) ; outerside of middle tibia with 6-9 teeth, some of then paired, and forming an irregular row; inner spur of hindtibia not quite reaching middle of basal segment of hind tarsus. Gaster of generalised form and broaderthan in nigra (Fig. 40). Tergite 1 with vague traces of rugosity across brow, lacking the highly polishedappearance of nigra. Ovipositor sheath as long as gaster plus thorax. Cf . Antenna almost as long as that of nigra from which it is distinguished by the structure of the claws. MATERIAL EXAMINED Holotype $, Bulgaria: Rhodopi, Theigovsharu, 9.viii.l979 (A. Zaykov) (ZC). Paratypes. 12 $ , same data (ZC; 4 in BMNH). Non-paratypic material Bulgaria: 3 cf , Rhodopi, Theigovsharu, 9. viii. 1979; 3 cf , Stointe, 4. viii. 1978 (A.Zaykov) (ZC). Greece: 6 $, 8 cf , Macedonia, Korab Mts, Stirovica, 27. vii. -3. viii. (V. Martina) (BMNH). HOST. Unknown. COMMENTS. The length of the galea together with the relatively large number of teeth on the outer side ofthe middle tibia make zaykovi distinct from all other included species that have a generalised gastral shape.The strongly lobed claws at once separate it from nigra to which it may not be closely related in spite of thelengthened galea. Agathis taurica TelengaAgathis taurica Telenga, 1955: 260. Lectotype $, U.S.S.R. (AS) designated by Tobias [not examined]. My interpretation of this species is based on a paralectotype cf received on loan from AS. I have beenunable to discover where, or indeed whether, the lectotype designation was published. $, 5 mm (excluding ovipositor). Like zaykovi in having hind claw with strong lobe though this is slightlyless well developed than in zaykovi; taurica may be compared with zaykovi as follows. Antenna with 25 segments (Telenga gives 24 segments for the female). Head in facial view like zaykovi(cf. Fig. 7). Mouth-parts longer; galea tapering to fine point, six times as long as basally wide. Head fromabove slightly less transverse and less emarginate behind. Thorax in profile slightly less elongate. A REVISION OF THE EUROPEAN AGATHIDINAE 205 Notaulices deeply impressed but smooth throughout. Virtually no trace of a sternaulus. Areolet offore-wing distinctly 4-sided but slightly narrower; distal abscissa of postmarginalis slightly shorter inrelation to proximal abscissa. Outer side of middle tibia with broad band of close, densely placed teeth,about 23 (only 1 ex.!); inner spur of hind tibia about 0-33 times as long as basal segment of hind tarsus.Gaster a little longer and narrower. Tergite 1 a little longer than its apical width, shining and with vaguetraces of sculpture across brow. Ovipositor sheath about as long as gaster plus thorax. MATERIAL EXAMINED Turkey: 1 $, Amasya, 1400 ft, 6.vi.l959 (Guichard) (BMNH). U.S.S.R.: 1 cf (paralectotype), Tauria,Simferopol (Pliginskij) (AS). HOST. Unknown. COMMENTS. As far as is known this species is not part of the main European fauna and has an easternMediterranean distribution. Among the species under review it is the third with greatly lengthenedmouth-parts. It is distinct from the other two species - nigra and zaykovi- with much lengthened galea (andindeed from all other included species) on the dense band of teeth on the outer side of the middle tibia.Telenga (1955) records taurica from Simferapol and Sevastopol; Tobias (1976) records it from the Crimeaand Armenia. Agathis semiaciculata Ivanov (Figs 13, 43)Agathis semiaciculata Ivanov, 1899: 364. Syntypes, U.S.S.R. (lost). Cf $,4-5-5-5 mm (excluding ovipositor). Black. Hind femur black throughout; middle and front femorablackish on at least basal half. 9 . Head in facial view only moderately elongate (Fig. 13). Malar space very slightly shorter than longerdiameter of eye, 13 : 15. Galea rather short, distinctly shorter than malar space, slightly more than 2-5 timeslonger than wide. Antenna thin, of even thickness throughout, 26-28-segmented: 26(2), 27 (1), 28 (1).Face almost as thickly pubescent as in griseifrons and in two out of four females with distinct traces ofstriate-punctation. Virtually no keel between antennal insertions and no depression in front of anteriorocellus. Thorax in profile of generalised shape. Notaulices deeply impressed. Side of pronotum anterior tooblique trough as coarsely rugose as in griseifrons. Sternaulus strongly developed, reaching posteriorcorner of mesopleurum. Areolet of fore- wing a little narrowed above but distinctly 4-sided; distal abscissaof postmarginalis hardly half as long as proximal abscissa. Propodeum with the usual, two longitudinalkeels, sometimes obscured by surrounding rugosities. Outer side of middle tibia with 1-2 teeth, difficult tosee because of infuscation of tibia; hind claw with small tooth at base but no deep cleft between it and clawproper (Fig. 43); hind claws longer and more powerfully built than in griseifrons; inner spur of hind tibiawell developed, two-fifths as long as basal segment of hind tarsus. Tergite 1 hardly longer than its apicalwidth, closely, evenly striate all over; striation tending to be broken and confused at middle of segment.Basal half of tergites 2+3 closely striate all over, the striation tending to be concentric around basal,transverse swelling. Ovipositor sheath almost as long as body. Cf . Like female except for sexual characters. Antenna with 26-28 segments: 26 (1), 27 (2), 28 (2).Definition of basal tooth of hind claw variable; sometimes represented by a mere projection or angulation.Sculpture of basal half of tergites 2+3 tending to be broken down to intricate rugose-striation. MATERIAL EXAMINED Italy: 1 $ , Bolzano, Seiser-Alm., 1900m, 24.vi.1976 (Zwakhals) (RNH). Switzerland: 2 cf , 3 $ , Arolla,7000-8000 ft, 29-30.vi.1935 (Benson) (BMNH); 1 9, Valais, Les Hauderes, vi.1935 (Benson) (BMNH); 1Cf , 1 $, Miistairtal, 2400ft, vi.-vii.1960 (Benson) (BMNH); 1 cf , 1 $, Saas-Fee, 7000-8000 ft, 25.vi.1962(Benson) (BMNH). HOST. Unknown. COMMENTS. This species has a fairly close affinity with griseifrons from which it can at once be separated onthe sculpture of the gaster. On the basis of the available material, it seems to be an alpine species.According to Telenga (1955) it is wide-spread in south-western regions of the U.S.S.R. with a rangeextending to northern Mongolia. No other species in this revision has such extensive sculpture on tergites 2+3 though there is sometimes aweak approach to this condition in anglica. The head in facial view is hardly different from that ofgriseifrons though slightly wider towards the mouth; also, griseifrons has a distinctly tapered flagellum. 206 G. E. J. NIXON Agathis polita sp. n. (Figs 8, 23) $ , 4-5 mm (excluding ovipositor) . Black with no trace of paler colouring on gaster . Wings almost hyaline .Head in facial view markedly elongate and considerably constricted beneath eyes (Fig. 8); seen fromabove, considerably prolonged backwards behind eyes (Fig. 23). A sharp keel between antennal insertionsunites above with a fairly distinct, V-shaped cavity in front of anterior ocellus. Antenna long, thin with26-27 segments. Galea rather broad, not tapered to apex and distinctly a little longer than malar space,5:4, and 2-5 times longer than wide. Thorax in profile markedly elongate. Notaulices at first sight wanting,indicated by barely visible, linear impressions. Sternaulus represented by short, hardly impressed line.Propodeum appearing highly polished, its twin keels virtually obliterated. Areolet of forewing triangular inholotype, distinctly 4-sided in paratype; distal abscissa of postmarginalis hardly shorter than proximalabscissa. Outer side of middle tibia with 8-9 irregularly spaced teeth; inner spur of hind tibia much less thanhalf as long as basal segment of hind tarsus; hind claw with prominent lobe. Gaster decidedly narrow.Tergite 1 about 1-33 times longer than apically wide, smooth, polished. Ovipositor sheath a little shorterthan head plus thorax. MATERIAL EXAMINED Holotype $, Jordan: Zerkatal, b. Romana, 200m, ll.iv.1959 (J. Klapperich) (HNHM).Paratype. 1 $, same data (BMNH). HOST. Unknown. COMMENTS. This is an extra-limital species and strictly has no place in this revision. It is included, however,because being virtually without notaulices it allows the following species, persephone, to be defined withmore accuracy. Agathis persephone sp. n. Cf 9, 5-5-6-0 mm (excluding ovipositor). Black with no hint of paler colouring on gaster. Hind femurblack. 9- Head in facial view short, subtriangular; from above, hardly distinguishable from that of polita (cf.Fig. 23). Keel between antennal insertions weak to almost absent. In front of anterior ocellus a fairlywell-defined, somewhat projecting, V-shaped cavity. Galea rather short, twice as long as wide, hardly aslong as malar space, 10: 11. Antenna long, somewhat tapered towards apex, 25-segmented. Thorax inprofile considerably elongate, almost twice as long as wide, 9:5. Mesoscutum without trace of notaulices.Mesopleurum with very short, weak sternaulus, separated from posterior corner of mesopleurum by fullyits own length. Areolet of fore-wing triangular, with distinct basal stalk (1 ex.!); distal abscissa ofpostmarginalis fully 0-66 times as long as proximal abscissa. Outer side of middle tibia with 4 teeth, the twoapical ones not easy to see because of infuscation of tibia; inner spur of hind tibia much less than half as longas basal segment of hind tarsus; hind claw with pale, angular projection rather than lobe. Side panels ofpropodeum in far greater part smooth, polished. Gaster beyond tergite 1 decidedly long, narrow; tergite 1slightly longer than apically wide with vague rugosity towards sides but virtually smooth across apical third.Ovipositor sheath fully as long as body. Cf . Like female but head, seen from above, slightly more produced backwards behind eyes. Antenna26-27-segmented. Sternaulus absent in one example, hardly indicated in the other. Areolet of fore-wingtriangular. MATERIAL EXAMINED Holotype $, France: Aspin, 680 m, 5.vii.l951 (H. Teunissen) (RNH). Paratypes. France: 1 cf, Dordogne, Sarlat, 4.vi.l975 (D. C. Geijsked) (RNH); 1 cf, Figeac, vi.1977(H. Teunissen) (BMNH). HOST. Unknown. COMMENTS. This species is distinct because it lacks notaulices. The short head, in facial view, together withthe ornamentation of the frons, suggests a probable relationship with varipes. Agathis ariadne sp. n. (Fig. 1)Cf $, co 4-5 mm (excluding ovipositor). Black. Gaster with no trace of red colour on tergites 2+3. A REVISION OF THE EUROPEAN AGATHIDINAE 207 Mandibles deep yellow; in the single female (holotype) labrum also deep yellow. Except for weak, basalinfuscation, hind femur bright reddish yellow in holotype and two out of three males. $. Head in facial view moderately elongate, wide across clypeus (Fig. 1), appearing very black andshiny, especially across clypeus owing to sparsity of pubescence. A well-defined keel between antennalinsertions and a fairly deep, V-shaped impression in front of anterior ocellus. Galea about 3 times as long asits basal width and 1-5 times longer than malar space. Antenna thin, rather short, not tapering towardsapex, 26-segmented. Thorax in profile somewhat elongate. Notaulices well defined. Sternaulus short,deep, not reaching posterior corner of mesopleurum. Side panels of propodeum polished except towardsantero-lateral corner. Areolet of fore-wing almost triangular; distal abscissa of postmarginalis about 0-5times as long as proximal abscissa; radial cell rather long, much as in varipes (cf. Fig. 50). Outer side ofmiddle tibia with 5 teeth arranged more or less in row; inner spur of hind tibia reaching virtually middle ofbasal segment of hind tarsus; hind claw with strong lobe. Gaster of generalised form. Tergite 1 only slightlylonger than its apical width and with only a very weak indication of sculpture. Ovipositor sheath about aslong as gaster plus propodeum; hairs of sheath very bristly, those in apical fifth hardly different from thoseproximal to it. d". Like female except for sexual differences. Antenna with 23-24 segments, long, rather thin. In onemale, tergite 6 is fully extruded and shows at base a specialised area (? sex gland), from which arise twopencils of hairs. MATERIAL EXAMINED Holotype $, Czechoslovakia: Szlovak, Erchs, Selmecbanya, 600 m, 22.vii.1976 (/. Papp) (HNHM).Paratypes. 3 cf , same data (HNHM, one in BMNH). HOST. Unknown. COMMENTS. This species certainly has some affinity with varipes but differs in the ornamentation of thefrons and in having face much widened below. The virtually undifferentiated bristles towards the apex ofthe ovipositor sheath provide an additional, useful character for the recognition of ariadne. The existence of what could be a sex gland at the base of the 6th tergite in the male needs to beinvestigated in detail; no similar structure is apparent in any of the males of varipes. Agathisanchisiadessp. n. (Figs 6, 22) $, ca 4-5 mm long (excluding ovipositor). Black. Gaster with no hint of paler colouring on tergites 2+3.Hind femur blackish with a small suffusion of paler colouring at extreme apex; hind tibia obscurely reddishyellow, infuscate in apical two-fifths and virtually with no trace of a darkened prebasal ring. Head in facial view characteristically elongate (Fig. 6); from above rather strongly produced backwardsbehind the rather large eyes (Fig. 22). A sharp keel between antennal insertions and a distinct, narrow,V-shaped cavity in front of anterior ocellus. Antenna long, tapering, 29-segmented. Galea a little shorterthan malar space, slightly more than 2-5 times longer than wide. Malar space two-thirds as long as longerdiameter of eye. Thorax in profile rather short, of generalised form. Notaulices well defined but not sharplycostate. Sternaulus sharply defined, reaching posterior corner of mesopleurum. Dorsal surface of sidepanels of propodeum showing much intricate rugosity. Areolet of fore- wing almost triangular; distalabscissa of postmarginalis almost as long as proximal abscissa. Segments 3-4 of middle tarsus very short, 4being not longer than wide; segment 4 of hind tarsus about 1-33 times longer than wide; inner spur of hindtibia reaching middle of basal segment of hind tarsus; this spur is margined with a particularly distinct rowof short bristles on its inner side; hind claw with strong lobe and deep cleft between lobe and claw proper;outer side of middle tibia with 7 thick teeth, very irregularly spaced. Gaster of generalised form. Tergite 1about as long as its apical width, markedly triangular, striate all over. Ovipositor sheath about as long aspropodeum plus gaster and, seen from above, as bristly at apex as at middle. MATERIAL EXAMINEDHolotype $, Hungary: Hortobargy, Zam, 2-23.V.1975 (Kaszab) (HNHM). HOST. Unknown. COMMENTS. A. anchisiades is remarkable for the shortness of certain tarsal segments; because of thisfeature it is distinct from all the other species included in this revision, with the exception ofachterbergi. Itis also to some extent characterised by the appearance of the head in facial view and also by the bristly apexof the ovipositor sheaths. 208 G. E. J. NIXON Agathis achterbergi sp. n. (Fig. 14) $ , ca 4-5 mm long (excluding ovipositor). Black. Hind femur entirely black; hind tibia obscurely yellowishon apical half but with heavily blackened apex; a distinct prebasal, infuscate band present. Wings brownishbut less so than in the related anchisiades. Head in facial view only moderately elongate (Fig. 14). Malar space about 0-66 times as long as longerdiameter of eye. Galea short, slightly less than twice as long as wide, dull, coriaceous. Between antennalinsertions a rather sharp keel that is sharply angled before it extends upwards, on a lower level, and uniteswith a deep, narrowly V-shaped cavity in front of anterior ocellus. Antenna long, thin, tapering towardsapex, 31-segmented; flagellum distinctly more bristly than in anchisiades; segment 4 from apex about 1-66times longer than wide. Thorax in profile like that of anchisiades . Notaulices deeply impressed. Sternaulusreaching posterior corner of mesopleurum. Side of pronotum anterior to oblique trough coarselyrugose-reticulate, especially on upper half. The two longitudinal keels of propodeum obscured bylongitudinal rugosities; side panels in greater part smooth, polished. Areolet of fore- wing markedly4-sided; radial cell rather long; distal abscissa of postmarginalis fully 0-75 times as long as proximalabscissa. Middle tarsus short, segment 4 hardly longer than wide; outer side of middle tibia with 5 ratherweak teeth arranged more or less in a row ; inner spur of hind tibia reaching middle of basal segment of hindtarsus; hind claw with strong, pointed lobe. Gaster of generalised form. Tergite 1 about as long as apicallywide, rugulose but becoming smooth across apical quarter; rest of gaster highly polished. Ovipositorsheath hardly longer than gaster, bristly at apex as in anchisiades; ovipositor itself straight, rather thick. MATERIAL EXAMINEDHolotype $, Holland: Waarder (Z-H), Oostende, 18.V.1970 (C. van Achterberg) (RNH). HOST. Unknown. COMMENTS. This species, as noted under anchisiades, is characterised by the shortness of the middle tarsus.The two species are in fact remarkably alike but the difference in the shape of the head in a facial view andin the length of the galea is too strong to fall within the range of specific variation. It is probable thatanchisiades and achterbergi are representatives of a species-group which is characterised essentially by theshortness of the middle tarsus and the strong lobe of the hind claw. Agathis minute Niezabitowski (Fig. 29)Agathis minuta Niezabitowski, 1910: 81. Syntypes, POLAND (lost). Cf $, 2-5-3-0 mm long (excluding ovipositor). Black. Hind femur infuscate more or less throughout,sometimes flushed with paler colouring towards apex; hind tibia pale brownish to pale brownish yellow butpaler parts of legs dingy. $. Head in facial view hardly lengthened, subtriangular (Fig. 29). Malar space very short, slightly lessthan half longer diameter of eye, 5:11. Mouth-parts very short; galea hardly longer than wide; preapicalsegment of labial palpus hardly twice as long as wide. A keel present between antennal insertions andalmost always a rather well-defined V-shaped impression in front of anterior ocellus. Antenna with 23-25segments. Thorax of generalised form, much like that of glabricula as seen in profile (cf. Fig. 41).Notaulices sharply defined. Sternaulus very short, isolated, not reaching posterior corner of mesopleurum.Propodeum tending to be rugose all over, the two medial, longitudinal keels sometimes obscured byrugosities. Areolet of fore- wing usually sharply triangular; radial cell rather small, distal abscissa ofpostmarginalis fully 0-66 times as long as proximal abscissa. Hind claw without a lobe; inner spur of hindtibia not reaching middle of basal segment of hind tarsus; outer side of middle tibia with 4-5 teeth arrangedin a row (difficult to see because of infuscation of tibia). Gaster rather short and broad. Tergite 1 about aslong as wide at apex and vaguely striate to rugose-striate all over. Tergites 2+3 with the usual, transverse,blisterrlike swelling at base and sometimes with traces of rugosity to sides of, and apical to, this swelling.Ovipositor sheath about as long as body but markedly variable in length. Cf . Like female in all essential details. Antenna thin, the preapical segment about twice as long as wide. MATERIAL EXAMINED 42 Cf , 84 $, Germany, Great Britain, Ireland, Sweden. A REVISION OF THE EUROPEAN AGATHIDINAE 209 HOST. Coleophora glaucicolella Wood (Coleophoridae) on Juncus inflexus L. (England: Worcester, nrRedditch, 3 $, 1 C? (BMNH)). COMMENTS. This is the most commonly represented species in all the collections examined and is widelydistributed. Easy to recognise on three main features: the shape of the head in facial view, the shortness ofthe galea and the long distal abscissa of the postmarginalis. Agathis asteris Fischer (Figs 31, 37)Agathis asteris Fischer, 1966: 185. Holotype $, AUSTRIA (NM) [examined]. C? $, 2-8-3-5 mm long (excluding ovipositor). Black. Hind femur blackish or at least considerablyinfuscate; general leg-colour much as in minuta. $. Head in facial view not much lengthened (Fig. 31). Malar space a little longer than half longerdiameter of eye, 1:12. Mouth-parts much lengthened in comparison with minuta; galea about 1-5 timeslonger than malar space. Almost no keel between antennal insertions and virtually no impression in frontof anterior ocellus. Head in dorsal view rather deeply hollowed out behind to receive the somewhatstrongly narrowed, anterior part of thorax. Antenna with 21-23 segments (2 ex.); flagellum distinctlythickened beyond middle; segments at this thickened part varying from being almost square in outline to1-25 times longer than wide. Thorax markedly elongate (Fig. 37); in profile almost twice as long as wide; indorsal view considerably narrowed in front, much more so than in minuta. Sternaulus long, groove-like,sometimes almost absent. Propodeum with some sort of rugosity almost everywhere. Areolet of fore-wingdistinctly 4-sided, narrowed above; distal abscissa of postmarginalis virtually absent. Hind claw withoutlobe; outer side of middle tibia with 3-5 teeth, difficult to see in two out of three females because tibia ingreater part infuscate; hind spurs rather weak; inner one not reaching middle of basal segment of hindtarsus. Tergite 1 slightly longer than wide, striate-rugose all over. Basal half of tergites 2+3 withoutrugosity in the three females available. Ovipositor sheath about twice as long as gaster, distinctly a littlewidened at extreme apex; hairs on outer side of expanded part distinctly shorter and less obvious than thoseproximal to it. C?. Like female. Antenna with 23 segments, thread-like; preapical segment twice as long as wide.Sternaulus virtually absent in the single example. MATERIAL EXAMINED Austria: 1 $ (holotype), Burgenland, Zitzmanns-dorfer Wiesen (Kasy) (NM); 3 $, 1 cf, Illmitz,Einsetzlacke, 7.ix.l971, ex Coleophora halophyella Zimmerman (Jackh) (EH). HOST. Coleophora halophyella Zimmerman (Coleophoridae). COMMENTS. This species is largely characterised by the long mouth-parts (cf. minuta, also a parasite ofColeophoridae), undecorated frons, elongate thorax and very short distal abscissa of the postmarginalis.Of the three last-mentioned characters it is mainly the elongate thorax that separates asteris from theglabricula-complex with its much more generalised thoracic form. Agathis glabricula Thomson (Figs 28, 41, 46) Agathis glabricula Thomson, 1895: 2228. LECTOTYPE $, SWEDEN (ZI), here designated [examined].Agathis albicostellae Fischer, 1966: 399. Holotype $, AUSTRIA (NM) [examined]. Syn. n. Cf $ , 3-0-3-2 mm long (excluding ovipositor). $. Black. Hind femur black; hind tibia blackish on apical third with rather conspicuous, blackish, basalring. Head in facial view rather sharply narrowed below eyes (Fig. 28), strongly transverse in dorsal view.Virtually no trace of a keel between antennal insertions, and at most a shallow dimple in front of anteriorocellus. Malar space about 0-66 times as long as longer diameter of eye, 7 : 12. Galea 2-0-2-5 times longerthan wide, about as long as malar space or sometimes a little shorter. Antenna with 25-28 segments;flagellum decidedly thin with the three preapical segments considerably longer than wide. Thorax in profileof generalised form (Fig. 41). Notaulices deeply impressed. Sternaulus deeply impressed, reachingposterior corner of mesopleurum . Sculpture of propodeum variable ; sometimes the normally polished sidepanels (as in lectotype) covered with an intricately rugose sculpture. Areolet of fore- wing variable; mostly 210 G. E. J. NIXON 4-sided, more rarely triangular; radial cell short; distal abscissa of postmarginalis hardly 0-33 times as longas proximal abscissa; proximal abscissa sometimes distinctly thickened. Outer side of middle tibia with 2-3teeth; inner spur of hind tibia not quite reaching middle of basal segment of hind tarsus; hind tarsal clawmostly without trace of free lobe but sometimes distal end of weak, basal thickening produced to form aslight angulation or tooth (as in lectotype, Fig. 46). Tergite 1 about as long as apically wide, sculptured allover, the sculpture varying between even striation and rugose-striation. Basal half of tergites 2+3 withtransverse, blister-like swelling and apical to this with a variable amount of broken, rugose-striation.Ovipositor sheath 1-33-1-50 times longer than gaster.Cf . Like female except for sexual differences. MATERIAL EXAMINED Austria: 1 $ (holotype), Braunsborg bei Hainburg, ex Coleophora albicostella Duponchel (NM).Bulgaria: 2 <j>, Rhodopi, Costinbrod, vi. (Zaykov) (ZC); 2 $, Valtshe, vii. (Zaykov) (ZC); 1 $, Plovdiv,ix. (Zaykov) (ZC). Great Britain: 3 9, 7 cf, Dorset, Portland Bill, bred viii.1959 ex Coleophoratroglodytella Duponchel, collected v.1959 (Ford) (BMNH). Ireland: 3 9, 1 d", Co. Clare, The Burren, exColeophora troglodytella Duponchel (Bradley) (BMNH). Sweden: 1 cf (lectotype), Skane, Ilstorp(Thomson) (ZI). HOST. Coleophora troglodytella (Duponchel) (Coleophoridae). COMMENTS. Fischer (I951a: 11) examined the syntypes of glabricula and labelled one of them 'Agathisglabricula Th.Type' but did not publish the designation; this specimen is here designated lectotype and islabelled accordingly. This species and the two that follow present a difficult taxonomic problem and only a provisional solutionis suggested. In certain respects, such as length of ovipositor and perhaps also of galea, the three speciesappear to overlap. They may be forms of a single species whose variation in structure could be due to theeffect of extrinsic factors. However, since names exist for the three segregates, they are kept separate untilmore material or biological information becomes available. Agathis meridionellae Fischer (Fig. 27)Agathis meridionellae Fischer, 1957&: 1. Holotype $, AUSTRIA: Karawanken (ZSBS) [not examined]. 9 This segregate differs from glabricula only in the following details. Galea a little shorter. Head in facial view triangular, less narrowed below eyes (Fig. 27). In two females(paratypes), ovipositor sheath clearly longer than gaster; in five females from England, Worcester City, itis hardly longer, and in one the propodeum is rugose all over, as in some of the females of glabricularecorded from England, Portland Bill. The specimens from Sweden have the ovipositor sheath hardlylonger than the gaster. MATERIAL EXAMINED Austria: 2 $ , 1 d" (paratypes), Karawanken, Loibtal, ex Coleophora meridionella Klimesch (EH). GreatBritain: 5 $, Worcester, ex Chrysoesthia hermannella F. (BMNH). Sweden: 17 $, 7 cf, Lund, exChrysoesthia (Microsetia on label) sexguttella (Thunberg) (ZI, 2 cf , 6 $, in BMNH). HOSTS. Coleophora meridionella Klimesch (Coleophoridae). Chrysoesthia sexguttella (Thunberg); Chry-soesthia hermannella (F.) (Gelechiidae). Agathis artemesiana Fischer (Fig. 24)Agathis artemesiana Fischer, 1966: 397. Holotype $, AUSTRIA (NM) [examined]. This species, which may belong to the glabricula-aggregate, differs from glabricula and meridionellae asfollows. $ , ca 3-5 mm long (excluding ovipositor); slightly larger than the other two species. Wings slightly moredarkened. Head in facial view slightly more elongate than in meridionellae, more like that of glabricula (cf.fig. 28); in dorsal view (Fig. 24). Flagellum distinctly thickened distal to middle, the thickest segmentsbeing no more than 1-25 times longer than wide; antenna 24-26-segmented. Areolet triangular or almostso. Ovipositor sheath apparently always short, about as long as gaster. A REVISION OF THE EUROPEAN AGATHIDINAE 211 MATERIAL EXAMINED Austria: 1 $ (holotype), Schlossofer Platte, Marchfeld (Kazy) (NM). Bulgaria: 2 <j>, Rhodopi,Nikolovo, viii.1976, on Medicago sativa L. (Zaykov) (ZC); 1 $, Velingrad, viii.1977 (Zaykov) (ZC); 1 9,Plovdiv, ix.1976 )Zaykov) (ZC). Great Britain: 1 $, Cambridgeshire, Fleam Dyke, 20.vii.1958 (Ford)(BMNH); Cornwall, St Osyth, 3.viii.l914 (Norwood) (BMNH). Ireland: 3 $, Co. Louth, Baltray,23.vii.1941 (Stelfox) (BMNH); 1 $, Co. Kildare, Royal Canal, 30.vi.1950 (Stelfox) (BMNH). Sweden:1 9, Skane, Degaberga, 14.vii.1938 (Perkins) (BMNH). HOST. Coleophora granulatella Zeller (Coleophoridae). COMMENTS. What characterises this species, if indeed it is distinct, is the shape of the middle segments of theflagellum. Agathis pedias sp. n. (Figs 15, 39) $, 4-5-4-8 mm (excluding ovipositor). Black. Hind tibia pale yellowish, almost whitish, with dark basalring and apical infuscation occupying about apical two-fifths. Wings pale compared with other species,almost hyaline. Head in facial view weakly elongate (Fig. 15). Virtually no keel between antennal insertions and only avery feeble impression in front of anterior ocellus. Malar space about 0-66 times as long as longer diameterof eye. Mouth-parts rather short; galea as long as malar space, markedly coriaceous. Antenna with 24-25segments. Thorax decidedly lengthened, twice as long as wide in profile; hardly different from that offulmeki (cf. Fig. 42). Sternaulus fairly sharply defined but remote from both anterior and posterior marginof mesopleurum. Panels of propodeum not extensively polished, sometimes almost obscured by en-croaching lateral rugosities. Outer side of middle tibia with 8-10 teeth arranged in a row, some of thempaired; inner spur of hind tibia hardly more than 0-33 times as long as basal segment of hind tarsus; legslong, slender; 3rd segment of hind tarsus very slightly longer than apical segment; hind claw with weak,angular pro j ection at base . Areolet of f orewing almost 4-sided ; distal abscissa of postmarginalis about 75times as long as proximal abscissa. Gaster somewhat long, narrow (Fig. 39); tergite 1 about 1-33 timeslonger than apically wide, rugose-striate all over. Ovipositor sheath as long as body. MATERIAL EXAMINED Holotype $, Spain: Cadiz, Villamartin, ll.vi.1960 (RNH). Paratypes. Spain: 2 $, same data as holotype (RNH, BMNH); 1 $, Alicante, Puerto de Confrides,(H. Teunissen) (RNH). Portugal: 1 9 , Caldes da Rainha, 14.V.1958 (RNH); 2 9 , Trajouce, San Domingosde Rana, 15.V.1958 (RNH). Greece: 1 $, Athens, Imittos, 26.iv.1980 (BMNH). HOST. Unknown. COMMENTS. The relatively large number of teeth on the outer side of the middle tibia suggests that pediashas an affinity with fulmeki, but the latter has shorter legs, a much more clearly defined impression in frontof the anterior ocellus and a longer galea. A decidedly slender habitus is somewhat characteristic of pedias. Agathis rostrata TobiasAgathis rostrata Tobias, 1963: 881. Holotype $, U.S.S.R. (AS) [not examined]. My interpretation of this species is based on an examination of a single female determined by Tobias. Thisspecimen bears the label 'W. Russland, Jurburg (Winogradoff- Nikitin)' and is probably that which Tobiasrecords from Lithuania, Jurbakas, in his original description. Although the data agree, Tobias does notindicate that it is a paratype, merely labelling it as 'Agathis rostrata sp. n., Tobias det.'. 9, 3-5-4-0 mm long (excluding ovipositor). Similar to anglica with which it may be compared as follows.On the whole darker in colour (this may have little significance) . Paler parts of hind tibia dingy yellow ; darkring at base of hind tibia sometimes united with infuscate, apical part by a dark band along upper surface.Head in facial view more gradually narrowed towards mouth. Mouth-parts longer; galea 1-25-1-33 timeslonger than malar space and rather abruptly narrowed in apical half. No trace of keel between antennalinsertions and at most the merest trace of a dimple in front of anterior ocellus. Antenna with 24-26segments: 24 (1), 25(2), 26 (1). Thorax slightly shallower than in anglica, seen in profile. Side of pronotumanterior to oblique trough more distinctly rugose-striate. Sternaulus different in that it reaches anteriormargin of mesopleurum as foveate rugosity or row of foveae. Sculpture of propodeum variable as in 212 G. E. J. NIXON anglica; sometimes rugose almost all over. Radial cell short as in angllca and areolet equally 4-sided; distalabscissa of postmarginalis slightly longer. Gaster as in anglica but ovipositor sheath hardly as long as gasterplus propodeum. MATERIAL EXAMINED Germany: 1 $,Ruthecoll. (BMNH). Great Britain: 1 $, Kent, Eynsford,24.vii. 1932 (Nixon) (BMNH).Italy: 1 $, Siidtirol, Algund, 1800 m, 30.viii.1967 (Haeselbarth) (EH); 1 $, St Peter/ Ahrntal, 1350 m(Haeselbarth) (EH). Sweden: 1 $, Skane, Degaberga, 12.vii.1938 (Perkins) (BMNH); 1 $, Silvakra,viii.1976 (Huddleston & Quintan) (BMNH). U.S.S.R.: 1 $, Lithuania, Jurbakas (Winogradoff-Nikitin)(AS). HOST. Unknown. COMMENTS. This species is essentially characterised by its long sternaulus, which reaches the anteriormargin of the mesopleurum, and the long, rather subtly distinctive galea. Agathis pappeisp. n. (Fig. 12) $, 3-0-3-5 mm (excluding ovipositor). Black. Legs obscurely yellowish where they are pale; hind femurinfuscate but yellowish at apex; in two out of three examples, including type, pale colour at apex of hindfemur extends as a pale band along dorsal surface as far as base. Wings almost hyaline. Head in facial view only weakly lengthened (Fig. 12). Face very shiny, with variable amount ofsomewhat indistinct punctation, more clearly evident in holotype. Anterior margin of clypeus somewhatprotuberant. Mouth opening wide and mandibles of powerful build. Malar space 0-66 times as long aslonger diameter of eye. Galea about twice as long as wide and as long as malar space. No trace of animpression or dimple in front of anterior ocellus. Antenna short, thin, with 22-24 segments. Mesoscutum alittle flattened, its middle lobe, and to a less extent, lateral lobes, with some vague, rather sparsepunctation. Notaulices deeply impressed, foveate, their outer margin bordered with a row of indistinctpunctures. Scutellum slightly depressed behind and rugulose here. Areolet of forewing weakly 4-sided;stigma rather broad; apical abscissa of postmarginalis almost absent. Hind femur slightly more than twiceas long as wide, the legs being rather short and thick; inner spur of hind tibia hardly reaching middle ofbasal segment of hind tarsus ; claws rather long , without basal lobe or tooth . Sternaulus in the form of a finegroove, not reaching anterior margin of mesopleurum. Gaster somewhat short. Tergite 1 about 1-25 timeslonger than apical width, indistinctly striated more or less all over; rest of gaster smooth, shining.Ovipositor sheath as long as gaster plus propodeum. MATERIAL EXAMINED Holotype ?, Hungary: Or, Sz.Miklos, l.xi.1917 (HNHM).Paratypes. Hungary: 2 $, Tompa, 11. ix. 1962 (Solymosne) (HNHM, BMNH). HOSTS. Unknown. COMMENTS. This aberrant species cannot be confused with any other from the region; it is characterisedmainly by the shape of the head as seen from in front, the wide mouth-opening and the correspondinglylarge, powerful mandibles. Agathis gradlipes Hellen Agathis gracilipes Hellen, 1956: 122. Lectotype $, FINLAND (ZH), designated by Tobias [examined].I have not been able to discover where or even whether the lectotype designation was published. Cf $, 4-0-4-5 mm (excluding ovipositor). Black. Hind femur infuscate but paler in apical half; hind tibiaobscurely yellowish, darkened over apical two-fifths, with a broad but faint infuscate band at base. $. Head in facial view not very elongate. Malar space slightly shorter than longer diameter of eye,10: 13. Galea very short, hardly longer than wide. No trace of keel between antennal insertions; a veryweak, V-shaped impression in front of anterior ocellus. Head in dorsal view strongly transverse, evenlyrounded behind eyes. Antenna long, thin, with at least 26 segments. Thorax of generalised form, 1-5 timeslonger than wide in profile. Notaulices deeply impressed. Sternaulus well defined, rugose, reachingposterior margin of mesopleurum and also, through a line of rugose-punctation, anterior margin.Propodeum with two weak, longitudinal keels; lateral panels polished medially. Radial cell rather long;areolet sharply triangular; distal abscissa of postmarginalis fully 0-5 times as long as proximal abscissa. Legs A REVISION OF THE EUROPEAN AGATHIDINAE 213 long, thinner than in any other included species; 3rd segment of middle tarsus fully 2-5 times longer thanwide; hind femur 4-5 times longer than wide; hind claw without lobe but with slight, basal thickening; outerside of middle tibia with one, indistinct tooth. Tergite 1 about 1-33 times longer than apically wide. Basalhalf of tergites 2+3 with some vague, rugose-striation towards sides. Ovipositor sheath about as long asbody. Cf . Antenna with 26 segments, long, thin, with apical segment fully 1-5 times longer than wide. Areoletof fore-wing with short stalk. Otherwise like female. MATERIAL EXAMINED Finland: 1 $ (lectotype), Tvarminne, Henriksberg (ZH); 1 cf , 1 $, Kexholm, 17.vii.1923 (Krogerus)(Cf in ZH, $ in BMNH). HOSTS. Unknown. COMMENTS. This species occupies a marginal position within Agathis. Because of its long, slender legs andvery short mouth-parts it could not be confused with any other species under review. Agathis melpomenesp. n. Cf $, 4-8-5-0 mm long (excluding ovipositor). Black. Hind femur yellowish, contrastingly darkened onapical one-third to two-fifths. $ . Head in facial view elongate and considerably narrowed below eyes; in lateral view, line of face andthat of clypeus appear virtually in one plane. No keel between antennal insertions and virtually noimpression in front of anterior ocellus. Galea 1-35-1-50 times longer than malar space, faintly coriaceousand rather abruptly narrowed from middle to apex. Antenna with 28 segments; flagellum virtually filiform.Thorax of generalised form. Notaulices deeply impressed throughout. Sternaulus strongly developed,reaching posterior corner of mesopleurum , and in one paratype (Markovo) extending also as far as anteriormargin of mesopleurum. Propodeum with usual, two longitudinal keels; lateral panels towards anteriordorsal surface covered with coarse shiny rugosity. Radial cell rather long; distal abscissa of postmarginalisfully 0-75 times longer than proximal abscissa; areolet 4-sided but distinctly narrowed towards edge ofwing. Outer side of middle tibia (holotype) with a row of 8 rather broad-based teeth, some of themoverlapping; 6-7 teeth in the two paratypes; these slightly smaller and less close together than in holotype;inner spur of hind tibia about two-fifths as long as basal segment of hind tarsus ; hind claw with conspicuous,dentiform lobe and wide cleft between this and claw proper. Tergite 1 hardly longer than apically wide;almost smooth in the two paratypes but with vague rugosity over middle part in holotype. Tergites 2+3polished, smooth (cf. anglicd). Ovipositor sheath hardly as long as gaster plus thorax. Cf . Antenna with 25-27 segments; flagellum very thin. In profile clypeus rises slightly above line efface.Outer side of middle tibia with 7 teeth. MATERIAL EXAMINED Holotype , Hungary: Taratovar6s, 28.V.1959 (Bajdri) (HNHM). Paratypes. Bulgaria: 1 $, Rhodopi; 1 $, Markovo, 23.vi.1978 (A. Zaykov) (HNHM); 1 $, S. poljana,24.vi.1975 (A. Zaykov) (BMNH). Nonparatypic material. Hungary: 3 cf , same data as holotype (HNHM; 1 in BMNH). HOST. Unknown. COMMENTS. This species is at once separable from anglica by the size of the radial cell, the stronglydeveloped, dentiform lobe of the hind claws and the reduction of sculpture on tergite 1. The presence ofmore numerous teeth on the outer side of the middle tibia is also of value though this character is likely tovary. The general conformation of the radial cell is like that of varipes, but melpomene is distinguished bythe shape of the head in facial view. The males are variable, consequently they are excluded from thetype-series. Agathis montana Shestakov (Figs 9, 21) Agathis montana Shestakov, 1932: 261. Holotype <j>, U.S.S.R. (AS) [not examined].The present interpretation of this species is based on specimens compared with the holotype by Tobias.Cf $,3-5 mm (excluding ovipositor). Black. Hind femur infuscate throughout in one female, tipped with 214 G. E. J. NIXON reddish in another (Kokcengir Hills) and entirely red in a third (Hungary: Vacz Csorog). Wings variablyinfuscate. $ . Head in facial view moderately elongate (Fig. 9); in lateral view of head, clypeus and face lie in thesame plane (Fig. 21). Malar space a little shorter than longer diameter of eye, 10 : 13. Galea slightly longerthan malar space, 13 : 10. No trace of a keel between antennal insertions. Virtually no trace of animpression or dimple in front of anterior ocellus. Antenna with 22-23 segments. Thorax in profilemarkedly elongate, nearly twice as long as wide, 25 : 14. Notaulices sharply defined but not deep so thatlobes lack a markedly convex appearance. Sternaulus in form of fine groove, only weakly rugose, notreaching anterior margin of mesopleurum. Propodeum sculptured all over in one female (Kokcengir Hills)but with polished area within lateral panels in the others. Areolet of fore-wing clearly 4-sided, thoughsometimes triangular according to Tobias; distal abscissa of postmarginalis hardly 0-33 times as long asproximal abscissa. Claws rather long, without basal tooth or basal thickening; outer side of middle tibiawith a row of 4-6 rather thick teeth; hind femur somewhat thick, 2-5 times as long as wide; inner spur ofhind tibia a little less than 0-5 times as long as basal segment of hind tarsus. Tergite 1 about as long asapically wide, finely rugose-striate all over. Ovipositor sheath about as long as gaster plus propodeum. Cf . Like female except for sexual differences. Antenna short, very thin; segment 4 from apex fully twiceas long as wide. MATERIAL EXAMINED Hungary: 1 <j>, Vacz Csorog (Biro) (HNHM); 1 $, Gyula, Veszelycsarda, 19.xi.1963 (Moczar)(HNHM); 1 $, Apajpuszta (Moczar) (HNHM). U.S.S.R.: 1 $, 1 cf, Kazachstan, Zhana-Arka, Kara-gand, viii-ix.1959 (Tobias) (AS); 1 $, Moldavia, Kishenev, S.x.1966 (Goncharenko) (AS). HOST. Unknown. COMMENTS. This somewhat squat-looking species is largely characterised by the shortness of the antennaand the elongate thorax. In respect of leg coloration and the shape of the areolet of the fore- wing montanais, as Tobias points out, extremely variable. There is also a subtle distinctiveness in the shape of the head asseen in profile, but it is based on features that are not easily described. RHAMPHAGA THIS TobiasRhamphagathis Tobias, 1962: 1195. Type-species: Agathis nasicornis Telenga, by monotypy. DIAGNOSIS. Face very broad across clypeus; latter in profile projecting as a snout. Labrum small,transverse. Mandible seen from in front, broadened, flattened, with two blunt, short teeth at apex; thusseen, mandible almost lamelliform. Mesoscutum in dorsal view flattened, with strongly produced anteriorcorners. Propleura much flattened in dorso-ventral direction, their posterior margin reflexed to accommo-date base of anterior coxae. Wing-venation like that of Agathis. Legs rather short; hind femur very stout. Rhamphagathis nasicornis (Telenga) Agathis nasicornis Telenga, 1955: 253. Lectotype <J>, U.S.S.R.: Kharkov (AS), designated by Tobias,1962: 1195 [not examined]. Cf $ , 3-5-4-0 mm long (excluding ovipositor). 9 Black. Hind femur deeply infuscate; hind tibia obscurely yellowish on basal half with faint dark bandnear base. Head in facial view as long as broad; seen from above with occiput deeply emarginate, the margin fallingaway steeply, almost angularly where it reaches posterior ocelli. Galea very slightly longer than malarspace. In front of anterior ocellus hardly a trace of a dimple but with short keel extending downwardsbetween antennal insertions. Antenna thin, thread-like, with 27-28 segments. Thorax in profile markedlyelongate. Propleura curiously flattened, dull, rugulose all over, densely pubescent and contrasting withsmooth, highly polished side of pronotum. Middle lobe of mesoscutum projecting far forwards on each sideto form a conspicuous shoulder; these shoulders coarsely rugose where they join notaulices; in profile,flattened mesoscutum falls perpendicularly to pronotum. Notaulices distinct throughout, slightly widenedin front and here rugose-foveate. Sternaulus in form of thin groove reaching posterior corner ofmesopleurum but not anterior margin of this. Legs rather short, thick; hind femur twice as long as wide;outer side of middle tibia with 6-10 thick, spine-like teeth on apical half; front coxa rather small, flattenedon lower side and here with sculpture similar to that of propleura and in sharp contrast with smooth, shinysurface of middle and posterior coxae; claws with a lobe. Radial cell rather narrow; areolet in femalealmost triangular; distal abscissa of postmarginalis very short, reduced almost to a point. Gaster somewhat A REVISION OF THE EUROPEAN AGATHIDINAE 215 short; tergite 1 about 1-25 times longer than apically wide, much widened towards apex; surface finely,rather weakly striated. Ovipositor sheath very slightly longer than gaster. Cf . Antenna with 26 segments (2 ex.); flagellum very slender; two preapical segments fully twice as longas wide. Otherwise like female except for sexual differences. MATERIAL EXAMINEDGermany: 2 cf, 2 $, Ruthe coll. (BMNH). U.S.S.R.: 1 $, Melito (Kokujev) (AS). HOST. Unknown. COMMENTS. This is a highly peculiar, quite isolated species. Apart from its generic features, it resembles thegenus Agathis in general habitus. The curious appearance of the propleura suggests an adaptation to aspecialised mode of existence. Among the species from the region Rhamphagathis nasicornis is easilyrecognised by its protuberant clypeus and prominent mesoscutal shoulders. MICRODUS Nees von Esenbeck Microdus Nees von Esenbeck, 1814: 184. Type-species: Ichneumon calculator Fabricius, by subsequentdesignation (Haliday in Westwood, 1840: 63, synopsis). DIAGNOSIS, cf $. Head in facial view transversely elliptical. Frons in front of anterior ocellus withoutdeepened and margined pit or furrow (cf. Agathis). Mouth-parts unmodified, never lengthened to form abeak-like structure; galea not longer than wide; maxillary palpus often appearing 3-segmented becausepreapical segment (3rd) sometimes so short as to be hardly noticeable; this segment always conspicuouslyshorter than apical segment. Sternaulus always present. Venation much as in Agathis (cf. Fig. 50); 1stdiscoidal cell and 1st cubital cell not separated by a vein; 2nd cubital cell always in form of small triangularor subtriangular areolet; distal abscissa of postmarginalis fully as long as proximal abscissa. Outer side ofmiddle tibia with 2-8 teeth, usually 3-4. Ovipositor at least 0-66 times as long as gaster. Holarctic in distribution. Muesebeck (1927) considered that Microdus could not be retained as a genus separate from Agathis andaccordingly synonymised them. His opinion was based on a study of North American material and found itssupport in the breakdown of the single character - the shape of the head - traditionally used to separate thetwo genera. Telenga (1955: 273) reinstated Microdus without giving reasons for rejecting Muesebeck's opinion. Hehas been followed by Tobias (1976ft: 97) who recognised Microdus on biological rather than morphologicalgrounds. Tobias argues that Agathis species, because of their lengthened mouth-parts, tend to inhabit arid,steppe-like regions where flowers conceal their nectar in deep corollas, a floral adaptation, Tobias adds,that makes the nectar available to long tongued insects inhabiting regions where they are exposed tomoisture deficiency. Microdus, on the other hand, is considered to be largely dependent on Lepidopterafeeding on forest trees and in consequence is well represented in the richly forested regions of the Far East. Tobias' observations are probably pertinent to the biology of the Agathidinae but it is arguable whetherthey provide a foundation for validating genera in a group with so unstable a classification as theBraconidae. Nevertheless, the two genera can be separated on sound morphological characters as follows. 1 Mouth-parts lengthened in the form of a beak, galea rarely as short as 1-33 times longer thanwide, nearly always much longer; head in facial view lengthened (Figs 1-6); ocelli in a hightriangle, the posterior tangent to the anterior ocellus passing freely in front of posterior pair;foramen of metasternum accommodating hind coxa not closed on inner side by a sclerotizedbridge. AGATHIS Latreille Mouth-parts not lengthened in the form of a beak, the galea not longer than wide; head in facialview transversely elliptical; ocelli in a low triangle, the posterior tangent to the anteriorocellus usually cutting, or at least touching, posterior pair; foramen of metasternum accom-modating hind coxa closed on inner side by a sclerotized bridge. MICRODUS Nees von Esenbeck Key to species (females) 1 Claws unarmed, without basal lobe; whole of hind leg blackish, except for yellowish band atextreme base of tibia.Thorax marked with black and red calculator F. (p. 217) 216 G. E. J. NIXON Claws armed with conspicuous basal lobe ; at least hind tibia predominantly pale 2 2 Inner margins of antennal sockets uniting to form a smooth knob that may be prolonged behind to form a short, blunt keel; tergites 2+3 with at least basal half with a well-defined sculpture that extends as far as the middle transverse suture or beyond 3 Inner margins of antennal sockets widely separated (except in pumilus but this species hasovipositor sheath much shorter than gaster), the surface between them either flat, smooth,with weak medial groove or (rarely) rugose; tergites 2+3 at most with some vague sculptureon basal half 7 3 Sculpture of tergites 2+3 tending to cease abruptly at second suture; at most faint traces of scaly-reticulation on apical half of tergite 4 Sculpture of tergites 2+3 extending almost to apex 6 4 Basal half of tergites 2+3 with sharp, shiny, longitudinal striation; ovipositor sheath much longer than gaster dimidiator Nees von Esenbeck (p. 219) Basal half of tergites 2+3 with somewhat confused rugosity, the longitudinal elements varyingmuch in intensity ; ovipositor sheath at most slightly longer than gaster 5 5 Notaulices almost fading out on disc; sculpture of basal half of tergites 2+3 consisting of vague scaly-reticulate rugosity with only weak longitudinal elements lugubrator Ratzeburg(p. 219) Notaulices sharply defined throughout; sculpture of basal half of tergites 2+3 with a predomi-nant sculpture of striation fortipes Reinhard (p. 220) 6 Tergites 2+3 evenly, longitudinally striate; hind tibia, except for darkened apex, reddish yellow rufipcs Nees von Esenbeck (p. 218) Tergites 2+3 not evenly striate, being in part rugulose to scaly-reticulate; hind tibia whitishwith darkened apex and darkened, basal ring rugulosus Nees von Esenbeck (p. 217) 7 Radius distinctly bent inwards at middle towards stigma; gaster often conspicuously pale- marked 8 Radius straight or almost so; gaster virtually always black 11 8 Small species not exceeding 3 2 mm ; gaster entirely black or blackish ; hind femur blackish or at least deeply infuscate 9 Larger species not less than 4-5 mm; gaster with at least basal half of tergites 2+3 reddish oryellowish (rarely gaster entirely dark) ; hind femur red 10 9 Radius very strongly bent inwards towards stigma, markedly colourless in middle third; notaulices deeply impressed throughout; hind tibia whitish, infuscate on rather less than apical half and usually with dark basal ring cingulipes Wesmael (p. 221) Radius less bent inwards towards stigma, darker at middle; notaulices almost absent on disc;hind tibia dull reddish with less apical infuscation and with hardly a trace of a basal band nugax Reinhard (p. 222) 10 Head in facial view slightly elongate (Fig. 32); internal rim of antennal sockets with distinct dentiform projection; hypopygium emarginate at apex, appearing strongly truncate in profile linguarius Nees von Esenbeck (p. 223) Head in facial view strongly transverse (Fig. 30); internal rim of antennal sockets without such aprojection; hypopygium not emarginate at apex, appearing pointed in profile. Usually entire gaster, except tergite 1 , reddish yellow conspicuus Wesmael (p. 225) 1 1 Head in facial view somewhat elongate 12 Head in facial view not elongate 13 12 Internal rim of antennal socket raised to form a distinct, dentiform projection; in profile, head behind lowest point of eye only weakly concave linguarius Nees von Esenbeck (p. 223) Internal rim of antennal socket without such a dentiform projection; in profile, head behindlowest point of eye rather deeply concave. Gaster beyond tergite 1 entirely reddish yellow, wings, strongly infumate zaykovi sp. n. (p. 226) 13 Ovipositor sheath hardly 0-66 times as long as gaster pumilus Ratzeburg (p. 221) Ovipositor sheath much longer than gaster 14 14 Tergite 1 with a sculpture of fine scaly-reticulation with at most traces of striation towards sides eriphyle sp. n. (p. 222)Tergite 1 without scaly-reticulation, the surface either almost smooth or with longitudinal striation varying in intensity 15 15 Hind femur stout, 3 times as long as wide; gaster shorter and broader; antenna with 29-31 A REVISION OF THE EUROPEAN AGATHIDINAE 217 segments; always an elongate, pale mark against posterior margin of eye tumidulus Nees von Esenbeck(p. 224) Hind femur not so stout, at least 3-5 times longer than wide; antenna with not less than 33segments ; gaster longer, narrower clausthalianus Ratzeburg (p. 224) Microdus calculator (Fabricius) Ichneumon calcula tor Fabricius, 1798: 225. Holotype $, ITALY (ZM) [not examined]. Microdus calculator (Fabricius) Nees von Esenbeck, 1834: 144. Microdus abscissas Ratzeburg, 18446: 58. Holotype cf , GERMANY (lost). [Synonymised by Ratzeburg, 1852: 45.] Agathis calculator (Fabricius) Shenefelt, 1970: 323.Microdus calculator (Fabricius) Tobias, 1971: 260. There never has been any doubt about the identity of this species. O" 9, 6-0-7-5 mm long (excluding ovipositor). At least head, propodeum and gaster black; sometimesmesosternum also black; rest of thorax red. Whole of hind leg blackish except for a pale, yellowish band atextreme base of tibia. Wings more or less hyaline. 9 . Head in facial view like that oilinguarius (cf. Fig. 32), less transverse than in other species; seen fromabove, very short, strongly transverse. A single blunt, almost knob-like keel between antennal sockets.Ocelli in a low triangle, the posterior tangent to anterior ocellus just touching posterior pair. Preapicalsegment (3rd) of labial palpus well developed, fully 2-5 times longer than wide. Antenna with 34-35segments. Notaulices deeply impressed throughout. Sternaulus broad, costate. Prepectal margin clearlydefined. Propodeum very coarsely reticulate but without discrete, longitudinal keel or keels. Areolet offore- wing triangular, usually with short stalk; distal abscissa of postmarginalis about as long as proximalabscissa; anal cell of hind-wing without anellus. Outer side of middle tibia with 3-4 isolated teeth; claw ofhind tarsus without trace of either lobe or tooth. Tergite 1 1-33-1-50 times longer than its apical width,strongly, evenly striate all over. Basal half of tergites 2+3 with sculpture like that of tergite 1 but sculptureless regular and not reaching apical corners of this half of tergites 2+3; sculpture tends to be confined totransverse, blister-like basal area. Ovipositor sheath about as long as thorax plus gaster. Cf. Like female but frequently darker; mesopleurum sometimes completely blackened. MATERIAL EXAMINED Great Britain: 1 9> Hampshire, New Forest, Bishop's Dyke, emerged 21.V.1963 from lepidopterouslarva pupating under dead bark of beech (Fagus) (BMNH); 4 cf , Winchester, flying around dead beechtree, viii.1979 (Else) (BMNH); 2 cf , 3 ?, Surrey, Mickleham, ex Scardia boleti, v.1933 (Ford) (BMNH);2 Cf , 3 $ , Box Hill, vii.1938, ex Scardia boleti (Ford) (BMNH). HOSTS. Morophaga boleti (Fabricius) (Tineidae) and probably also Triaxomera parasitella (Hiibner)(Tineidae). According to records quoted by Shenefelt (1970), various people have bred calculator fromthese two lepidopterous hosts. COMMENTS. This is an unmistakeable species, at least in the region under consideration, and is readilycharacterised by its red and black colour, large size and, most significantly, by the unique simple claws. Microdus rugulosus Nees von Esenbeck comb. rev. Microdus rugulosus Nees von Esenbeck, 1834: 148; Reinhard, 1867: 357. Holotype $, GERMANY (lost).Braunsia rugulosa (Nees von Esenbeck) Telenga, 1955: 278; Shenefelt, 1970: 376. Reinhard was the first reviser of this species and from his excellent treatment of it, the identity of rugulosusis not in doubt. $, 4-5 mm long (excluding ovipositor). Head and thorax black. Gaster usually entirely blackened abovebut frequently pale-marked towards apical corners of tergites 2+3; in such cases, the segments posterior tothe pale area tend to be entirely pale yellowish brown or reddish brown; in one example (Surrey,Mickleham), basal half of tergites 2+3 is bright reddish yellow. Hind femur varying from entirely infuscateto entirely reddish yellow; when hind femur is pale, the hind tibia is contrastingly whitish with apical thirdand a basal band infuscate; in examples with darkened hind femur, the tibia is correspondingly darkened;hind coxa reddish yellow over most of basal half or entirely infuscate. Head in facial view transverse, as in majority of species (cf. Fig. 30). Frons between antennal socketsshowing as a smooth, weakly rounded boss, laterally compressed to form a single, short, rounded ridge. 218 G. E. J. NIXON Segment 3 of labial palpus fully 0-66 times as long as apical segment. In front of anterior ocellus anobsolescent, V-shaped depression. Ocelli in a low triangle, the posterior tangent to anterior ocellusvirtually touching posterior pair. Antenna with 30-34 segments: 30 (5), 31(2), 34(1). Notaulices sharplydefined but shallow, finely foveate throughout. Sculpture of propodeum variable, tending to fade out oneach side of convexity, leaving an almost smooth area. Areolet of fore- wing almost always sharplytriangular; rarely with short stalk. Inner spur of hind tibia not reaching middle of basal segment of hindtarsus; hind claw with conspicuous, angular lobe; outer side of middle tibia with 2-3 well-separated teeth.Tergite 1 about 1-66 times longer than apically wide, with fine, mostly broken rugose-striation all over;where the striation becomes weak, towards apex of tergite, surface shows a considerable amount of weakscaly-reticulation. Tergites 2+3 almost 1-5 times longer than apically wide, sculptured almost all over;distal to 2nd suture, which shows as a shallow furrow, the surface is weakly scaly-reticulate, the sculpturetending to fade out towards apical corners; basal to the suture (the area probably corresponding to the truesecond tergite), a weakly rugose furrow sets off the usual, transverse, slightly raised area; whole of basalhalf of tergites 2+3 vaguely rugose-striate with either the rugose or striate element predominating.Hypopygium sharply pointed at apex. Ovipositor sheath about equal to gaster plus thorax. MATERIAL EXAMINED Germany: 3 $, Ruthe coll. (BMNH). Great Britain: 2 $, Bucks., vii., on dead Quercus sp. (BMNH);1 $, Essex, Epping Forest, viii.1954 (BMNH); 3 $, Hants., Romsey, vii.-viii.1980, from wood of deadLarix (C. Vardy) (BMNH); 1 $, Southsea, coll. 19.iii.1983, em. vi.1983, ex Blastobasis lignea in deadtwigs of Ulex (BMNH); 1 $, Surrey, Mickleham, vii.1935 (BMNH). Greece: 1 $, Ilia, Olympia, vii.1979(Day, Else & Morgan) (BMNH). Ireland: 4 $ , Glengariff , Lodge Woods, vii.1935, on Quercus sp. (A. W.Stelfox) (BMNH). Switzerland: 1 $, Wallis, Brig, 2.vii.l959 (E. Bauer) (EH). HOST. Blastobasis lignea Walsingham (Blastobasidae). According to Meyrick (1968), the larva of this mothfeeds on dead leaf-refuse but this may also include dead wood which would explain the data given above. COMMENTS. Though variable both in colour and sculpture, this species is highly distinctive on the sculptureof the gaster; it could not be confused with any other Microdus from the region. Microdus rufipes Nees von Esenbeck Microdus rufipes Nees von Esenbeck, 1814: 189. Syntypes, GERMANY (lost).Braunsia rufipes (Nees von Esenbeck) Telenga, 1955: 277; Shenefelt, 1970: 375.Microdus rufipes Nees von Esenbeck; Tobias, 1971: 260. C? $, 4-5 mm long (excluding ovipositor). Black; rarely basal third of tergites 2+3 reddish yellow (1 $,England: Suffolk, Sudbury). Hind coxa and hind femur bright reddish yellow; hind tibia slightly duskierthan its femur and darkened on about apical third. A single female from Jugoslavia: Ristovaca has the face,mesoscutum and sides of thorax richly marked with paler colouring. 9. Head like that of rugulosus. Structure of frons between antennal sockets like that of rugulosus butthickened keel tapering behind and uniting with a distinct V-shaped depression in front of anterior ocellus.Ocelli in a low triangle, the tangent to anterior ocellus cutting posterior pair. Antenna with 31-32segments. Notaulices deeply impressed, much more so than in rugulosus. Propodeum covered with coarse,confused rugosities. Wings like those of rugulosus but distal abscissa of postmarginalis not longer thanproximal abscissa. Outer side of middle tibia with 6-7 closely spaced teeth near apex; inner spur of hindtibia relatively longer than in rugulosus and almost reaching middle of basal segment of hind tarsus. Tergite1 considerably widened towards apex, about 1-25 times longer than apically wide, strongly, evenly striateall over. Tergites 2+3 showing three transverse furrows of variable definition, its entire surface, exceptacross surface distal to third furrow, with shiny, on the whole even, longitudinal striation. Tergites 4 and 5sometimes with faint traces of scaly-reticulation. Ovipositor sheath about equal to length of thorax plusgaster. Cf . Like female. Rarely the striation at base of tergites 2+3 somewhat broken and transverse indirection. MATERIAL EXAMINEDBulgaria, Great Britain, Sweden, Turkey, Yugoslavia. HOSTS. Cydia pomonella (L.) (Tortricidae); Spilonota ocellana (Denis & Schiffermiiller) (Tortricidae);Hedya nubiferana (Haworth) (Tortricidae); Gypsonoma dealbana (Frolich) (Tortricidae). The foregoinghosts are from specimens personally examined. Tobias (1971) cites the second two and in addition givesRhyacionia buoliana (Denis & Schiffermiiller) (Tortricidae). A REVISION OF THE EUROPEAN AGATHIDINAE 219 COMMENTS. This is probably the most distinct of all European species of Microdus and is easily recognisedby the even striation of the gaster. It is because of this highly characteristic striation that Telenga (1955)transferred rufipes to Braunsia Kriechbaumer but this genus is almost exclusively Afrotropical and thoughcharacterised by a striated gaster, it has no close affinity with Microdus. Microdus dimidiator Nees von Esenbeck Microdus dimidiator Nees von Esenbeck, 1834: 146. Holotype $, POLAND (lost).Agathis dimidiator (Nees von Esenbeck) Shenefelt, 1970: 329.Microdus dimidiator Nees von Esenbeck; Tobias, 1971: 260. Cf 9> 4-5 mm (excluding ovipositor). Black; anterior corners of tergites 2+3 somewhat pale, the palecolouring sometimes forming a complete band across base of segment. Hind coxae usually entirely reddishyellow but sometimes becoming infuscate towards base; hind femur reddish yellow; hind tibia whitishyellow but infuscate on about apical third and with faint to absent darker ring near base. Wings almosthyaline. 9- Head as in rugulosus. Between antennal sockets a gently rounded hump as in other species withsculptured gaster. Segment 3 of labial palpus poorly developed, almost moniliform and much shorter thanapical segment. Posterior tangent to anterior ocellus virtually touching posterior pair. Antenna with 33-35segments: 22 (2), 33 (1), 34 (2), 35 (2). Notaulices deeply impressed. Propodeum more coarsely andintricately rugose than in rufipes. Lobes of mesoscutum sometimes showing a weak, sparse punctation.Areolet of fore wing smaller than in rufipes and more markedly stalked. Hind spurs more powerfully builtthan in either rugulosus or rufipes, the inner one just reaching middle of basal segment of hind tarsus; outerside of middle tibia with 3-4 teeth arranged in a row and well separated from small cluster of teeth atextreme apex. Tergite 1 about 1-5 times longer than apically wide, longitudinally striate all over; striationclose, dense, shiny. Tergites 2+3 with similar striation extending as far as second suture and here endingabruptly; distal to second suture, surface (corresponding to area of tergite 3) highly polished andsometimes showing the faintest trace of a transverse furrow. Ovipositor sheath about as long as body. O". Like female except for sexual differences. Tergite 1 tending to be a little longer than in female andstriation of tergites 2+3 sometimes broken or obsolescent. MATERIAL EXAMINED France: 8 O" , 15 9 , Vaucluse, Mont Ventoux, bred vi.-vii. 1970, 1980 ex Tortrix viridana L. , on Quercussp. (P. du Merle) (PdMC, 2 cT, 5 9, BMNH). Germany: 1 $, Ruthe coll. (BMNH); 1 9, Saxony (Reinh. . .) (RNH). Greece: 1 C?, Krausia, Oros, 4.vi.l973 (Schach) (RNH). Holland: 1 9 (on label 'v.Voll.Holl.') (RNH). Poland: 2 cf, 3 9, Skierniewice, vi.-vii. 1957 (1 9, labelled ex 'ocellana') (Wiackowski)(BMNH). Yugoslavia: 1 9, Ristovaca, 8.vi.l982 (Mihajlovic) (BMNH). HOSTS. Tortrix viridana L. (Tortricidae); Spilonota ocellana (Denis & Schiffermuller) (Tortricidae). COMMENTS. The female from Yugoslavia has the hind coxa blackish throughout, and the hind femurdarkened with the apex almost infuscate. The traditional interpretation of dimidiator is certainly correct, for Nees von Esenbeck has carefullydrawn attention to its salient features in his description. The essential character for recognising this speciesis the abrupt termination, at the second suture, of whatever sculpture exists on the basal half of tergites2+3, i.e. the sculpture is sharply restricted to the surface of what is morphologically the second tergite.Further, M. dimidiator is more slenderly built than either rugulosus or rufipes and has noticeably longerlegs. Microdus lugubrator Ratzeburg comb. rev.(Figs 57, 59, 63) Microdus lugubrator Ratzeburg, 1852: 45. Syntypes, GERMANY (lost).Agathis lugubrator (Ratzeburg) Fulmek, 1962: 103; Shenefelt, 1970: 341. Cf 9 , 3-5-4-0 mm long (excluding ovipositor). 9 . Black, sometimes with paler colouring on gaster. Hind coxa deeply infuscate; hind femur dull reddishyellow; hind tibia much the same colour but faintly darkened at apex. Wings considerably darkened. Head not strongly transverse (Fig. 63). Frons between antennal sockets weakly raised to form a single,blunt, almost knob-like keel as in rugulosus and rufipes. Antenna with 28-31 segments: 28(2), 30(2), 31(1);scape rather short (Fig. 59). Ocelli in rather a high triangle, the posterior tangent to anterior ocellus not 220 G. E. J. NIXON touching posterior pair. Notaulices poorly defined, sometimes almost absent on disc. Propodeum coarsely,intricately rugose all over; sometimes with 2 fairly distinct, longitudinal keels. Legs rather thick; outer sideof middle tibia with 3-4 well-spaced teeth in a row; inner spur of hind tibia not quite reaching middle ofbasal segment of hind tarsus. Areolet of fore- wing varying from distinctly triangular to distinctly 4-sided;distal abscissa of radius virtually straight, widely separated from stigma (Fig. 57). Gaster shorter andbroader than in other species of the rugulosus-complex. Tergite 1 much widened to apex, about 1-25 timeslonger than apically wide, densely rugose-striate all over and usually with a clear indication of superim-posed scaly-reticulation. Tergites 2+3 with a distinct 2nd suture and usually a curved furrow proximal tothis; tergites 2+3 proximal to 2nd suture with a fine variable sculpture all over; within the slightly raised,basal area, sculpture usually shows distinct striate elements, sometimes transverse in direction; elsewheresculpture a mixture of scaly-reticulation and fine striation; distal to 2nd suture, surface shows traces,sometimes very indistinct, of scaly-reticulation. Ovipositor sheath 0-66-0-75 times length of gaster. Cf . Gaster sometimes with dull reddish tinge. Antenna with 26-28 segments: 26(2), 28(1). In the scantymaterial available, gaster rather more coarsely sculptured than in female. MATERIAL EXAMINED France: 1 $, Gironde, 1936, ex Coleophora sp. (/. Suire) (BMNH); 1 $, Fontainebleu, viii.1936 (A.Alston) (BMNH). Germany: 1 $, Ruthe Coll. (BMNH). Great Britain: 1 $, Essex, Epping Forest, exColeophora case on oak bark (Quercus sp.), coll. 3.vii.l983, em. 25.vii.1983 (P. J. Johnson) (MS); 2 $,Hampshire, New Forest, ex Coleophora lutipennella Zeller or C.flavipennella Duponchel on Quercus sp.,coll.28.v.l983, em.1983 (P. H. Sterling) (MS); 1 $, 1912, 1 $, 22.vii.1908 (G. T. Lyle) (BMNH); 1 $,Kent, Bexley, vii.1937, ex Coleophora hemerobiella Scopoli (R. L. E. Ford) (BMNH); 2 $ , SE. London,Norwood, viii.1928 (/. H. Lowe) (BMNH); 1 $>, Surrey, Ashtead, viii.1928 (G. E. J. Nixon) (BMNH); 1$, Worcester, Trench Wood, ex Coleophora flavipennella Duponchel, on Quercus sp., coll. vi.1983, em.vii.1983 (A. N. Simpson) (MS). Hungary: 1 $, Budapest, Kamaraerdo, 31.vii.1976 (J. Papp) (HNHM); 1Cf , Nagykovacsi, 30. vi. 1973, ex Coleophora sp. (SzocsJ.) (HNHM). HOSTS. Evidently a parasite of various species of Coleophora: Coleophora flavipennella (Duponchel),Coleophora hemerobiella (Scopoli) and possibly Coleophora lutipennella (Zeller). COMMENTS. This species is mainly characterised by its weak notaulices, sculpture of gaster and shortovipositor. Its nearest relative seems to be fortipes (see below). Microdus fortipes Reinhard (Fig. 65)Microdus fortipes Reinhard, 1867: 356. Syntype $. GERMANY (MNHU) [examined]. Cf $ , 4-5 mm long (excluding ovipositor). $ . Hind femur entirely deep reddish in two females (Munich) but much darkened in single female fromWitzenhausen; hind tibia much the same colour as hind femur but infuscate at extreme apex. Wingsmarkedly infuscate, as in lugubrator. Head from above much more transverse (Fig. 65). Antenna shorter,thicker, with 23-25 segments; three preapical segments hardly longer than wide; in lugubrator, they arefully 1-25 times longer than wide; scape slightly longer. Thorax in profile much less elongate. Notaulicesdeeply impressed throughout. Propodeum with much coarser reticulate rugosity. Anella of hind wing notemitting an anal vein; in this respect markedly different from lugubrator; areolet of fore-wing (3 9 , 2 C?)always sharply triangular and sometimes with short stalk. Legs thicker than in lugubrator; outer side ofmiddle tibia with a cluster of 8-10 teeth close to apex. Gaster as in lugubrator but tergite 1 and basal half oftergites 2+3 as far as second suture with irregular sculpture of shiny, longitudinal striation; on basal part oftergites 2+3 the striation tends to be concentric around a mid-basal area and towards sides shows weaklysuperimposed scaly-reticulation; distal to second suture, tergites 2+3 mainly polished with faint traces ofscaly-reticulation and a transverse linear impression across middle. Ovipositor sheath almost as long asgaster plus propodeum; seen from above, hairs of apical fifth of ovipositor sheath shorter and lessnoticeable than in lugubrator. Cf . Like female except for sexual differences. Sculpture of gaster reduced; that on tergites 2+3 almostobliterated. MATERIAL EXAMINED Germany: 2 $, Munchen (Buchecker) (BMNH); 2 cf, 1 $, Witzenhausen, viii.-ix.1965 (Haeselbarth)(EH). A REVISION OF THE EUROPEAN AGATHIDINAE 221 HOST. According to Reinhard this species was bred frequently by Bouche from Spilonota ocellana (Denis &Schiffermuller) (Tortricidae). COMMENTS. Microdus fortipes is perhaps more closely related to lugubrator than to any other species fromthe region. From lugubrator it is distinguished mainly by the more transverse head, the sculpture of thegaster and the much more deeply impressed notaulices. In lugubrator the sculpture of the basal half oftergites 2+3 presents a dull, vaguely scaly-reticulate appearance in which longitudinal elements are poorlydefined; in fortipes, on the other hand, the sculpture here is predominantly strong, longitudinal striation. Microdus pumilus Ratzeburg Microdus pumilus Ratzeburg, 1844ft: 57. Syntypes, GERMANY (lost).Agathis pumila (Ratzeburg) Shenefelt, 1970: 350.Microdus pumilus Ratzeburg: Tobias, 1971: 260. Cf $ , ca 2 mm (excluding ovipositor). $. Black. Hind femur deeply infuscate; hind tibia whitish on basal half with darkened subbasal band,infuscate on apical half. Head rather deep from back to front, as seen from above; much as in lugubrator (cf. Fig. 63). Ocelli in ahigh triangle, the posterior tangent to anterior ocellus passing far in front of posterior pair. Face having ashiny, convex, polished appearance. Frons between antennal sockets raised at middle to form a knob that isslightly lengthened in direction of anterior ocellus; in this respect similar to the rugulosus-group. Palpishort; segment 4 of maxillary palpus much shorter than 5. Notaulices shallow, sometimes fading out beforereaching posterior fovea. Sternaulus shallow, broken, isolated and not reaching back as far as posteriorcorner of mesopleurum . Hind spurs weak , subequal , the inner one not reaching middle of basal segment ofhind tarsus. Stigma rather broad; apical abscissa of postmarginalis considerably shorter than proximalabscissa. Outer side of middle tibia with 2-3 teeth, difficult to see because of infuscation of tibia; hind clawsmall but with distinct lobe. Gaster somewhat short; tergite 1 triangularly widened to apex, about 1-33times longer than its apical width; delicately, somewhat brokenly, longitudinally striate. Ovipositor sheathabout 0-66 times as long as gaster, shorter than in any other species from the region. Cf . Like female except for sexual differences. Antenna with 27 segments. MATERIAL EXAMINED Germany: 1 cf , 6 , S.-Holstein, Segeberg, ex Coleophora laricella Hiibner (T. Weber) (EH). Holland:1 $, Bergentheim, 9.vii.l975 (B. v. Aartsen) (RNH). HOST. Coleophora laricella (Hiibner) (Coleophoridae). This host, according to Meyrick (1968), isrestricted to Larix (see discussion under 'Biology'). COMMENTS. This is the smallest species from the region and is most distinctive on account of its very shortovipositor. It stands apart from the other species with polished tergites (except tergite 1) because of thestructure of the frons between the antennal insertions. Microdus cingulipes Nees von Esenbeck (Fig. 55)Microdus cingulipes Nees von Esenbeck, 1814: 189. Syntypes, GERMANY (lost). Wesmael (1837: 18) was the first reviser of cingulipes; I have based my interpretation of the species on aspecimen in Wesmael's collection bearing his handwritten identification label. $, 3-5-4-0 mm (excluding ovipositor). Black. Hind femur black; hind tibia obscurely whitish yellow atmiddle; deeply infuscate on apical two-fifths and with ring of infuscation near base. Head in facial view as transverse as in majority of species; seen from above strongly transverse. Antennalsockets on their inner side joined to form a weakly bipartite projection. Ocelli in low triangle, the posteriortangent to anterior ocellus virtually touching posterior pair. Labial palpus with only 3 clearly visiblesegments. Antenna with 29-31 segments. Notaulices deeply impressed throughout, sometimes formingpatch of rugosity where they unite behind. Sternaulus distinct throughout but narrow, linear and at mostfinely rugose. Propodeum rugose all over but showing no special feature. Hind spurs rather weak; innerspur of hind tibia falling far short of middle of basal segment of hind tarsus; outer side of middle tibia with2-3 teeth just proximal to apical cluster of 3-4 teeth. Radius colourless over middle third, strongly bentinwards towards stigma and, where closest to this, separated from it by about thickness of radius itself (Fig. 222 G. E. J. NIXON 55); stigma shorter and less wedge-shaped than in tumidulus and close allies. Tergite 1 1-50-1-66 timeslonger than its apical width, somewhat weakly striate all over. Tergites 2+3 basal to 2nd suture with usual,curved furrow, delimiting a transversely elliptical area; rarely this furrow finely costate; gaster otherwisesmooth, polished. Ovipositor sheath about as long as thorax plus gaster. MATERIAL EXAMINED Belgium: 1 $, Wesmael coll. (IRSNB). Germany: 2 cf, Ruthe coll. (BMNH); 5 $, Oberbayern,Hochstadt, vii.-viii. (Haeselbarth) (EH); 1 $, Oberbiberg, 2.vii.l969 (Haeselbarth) (EH); 1 $, Wessling,7.viii.l972 (Haeselbarth) (EH); 1 $, Glonn, 14.vii.1968 (Haeselbarth) (EH). Great Britain: 1 $,Hampshire, Havant Thicket, ex Epinotia fraternella Douglas on Abies grandis, collected 10. xi. 1977,emerged iii.1978 (Langmaid) (MS); 1 $, Essex, Saffron Walden, ix.1966, ex Coleophora frischella L.(Emmet) (AA); 1 9 , Yorkshire, Langdale, on Pinus (Day) (BMNH). Ireland: 2 $ , Co. Down, TollymorePark, vii.1961 (Stelfox) (USNM); 1 $>, Dublin, Glenasmole, viii.1944 (Stelfox) (USNM). Italy: 1 $>,Sudtirol, St Peter Ahrntal, viii.1967 (Haeselbarth) (EH). HOSTS. Gary ocolum fraternella Douglas (Gelechiidae) ; Coleophora frischella (L.) (Coleophoridae). COMMENTS. This is a very distinctive species, largely characterised by the deeply incurved radius. It seemsto be closely related to the following species but there is no possibility of confusing the two. Microdus nugax Reinhard (Fig. 54) Microdus nugox Reinhard, 1867: 354. Syntype $, GERMANY (MNHU) [examined].Agathis nugax (Reinhard) Shenefelt, 1970: 347.Microdus nugax Reinhard; Tobias, 1971: 260. Cf $ , 3-5-4-0 mm (excluding ovipositor). $ . Very close to cingulipes with which it may be compared as follows. Paler part of hind tibia obscurelybrownish red with virtually no trace of a darkened, basal ring; hind femur varying from obscurely brownishred to entirely infuscate; hind spurs darker. Head from above slightly less transverse, deeper from back to front. Face polished, convex, virtuallyimpunctate. Ocelli slightly smaller; posterior tangent to anterior ocellus passing freely in front of posteriorpair. Preapical segment (3rd) of labial palpus so small as to be virtually absent. Antenna with fewersegments: 23-26; 23 (1), 24 (3), 25 (4), 26 (1). Thorax more elongate. Notaulices characteristically weak,sometimes not or hardly indicated posteriorly. Hind spurs less well developed. Radius only slightly bentbut pale along middle stretch (Fig. 54) as in cingulipes. Tergite 1 a little less elongate, its sculptureconsiderably weaker. Ovipositor sheath a little shorter; seen from above, with denser, more closely sethairs that on apical third of sheath do not noticeably project. Cf . Like female except for sexual differences. MATERIAL EXAMINED Italy: 4 cT, 3 ?, Sudtirol, Algund, 1900 m, 24.vii.1966 (Haeselbarth) (EH; 1 cf , 1 $ in BMNH); 2 cf , 19, Karthaus, 1200 m, 14.vii.1976 (Haeselbarth) (EH); 3 cf, 2 $, Marling, 1200 m, 12.vii.1966 (Haesel-barth) (EH; 1 Cf , 1 $ in BMNH); 2 $ , St Peter/ Ahrntal, vii.-viii. (Haeselbarth) (EH); 1 $ , Campi, Riva s.Garda, 1400 m, 7.vii.l976 (Haeselbarth) (EH). HOST. Unknown. COMMENTS. A small, obscurely coloured species, essentially characterised by the weakly developednotaulices and the shape of the radial cell. Microdus eriphyle sp. n. (Figs 56, 62, 64) $ , 4 mm (excluding ovipositor). Black. Hind femur black; hind tibia obscurely whitish yellow, infuscate inapical third and with a broad infuscate prebasal band. Head from in front as transverse as in majority of species; seen from above as in Fig. 64. Antennalsockets unusually deep; seen from above frons between sockets very slightly concave. Behind frontal ridgeand towards anterior ocellus a very short, almost tuberculiform keel. Ocelli in a high triangle, the posteriortangent to anterior ocellus passing in front of posterior pair. Preapical segment of labial palpus (3rd) welldeveloped, clearly longer than wide. Notaulices deeply impressed throughout, smooth, uniting posteriorly A REVISION OF THE EUROPEAN AGATHIDINAE 223 to form an elongate, smooth furrow. Propodeum appearing smooth, shiny, its sculpture very weak and inone female (paratype) almost absent. Radius of fore-wing strongly incurved but not so much as indngulipes (cf. Fig. 55), evenly pigmented throughout. Outer side of middle tibia with a row of 4 teeth,widely spaced and linking up with 2 at extreme apex of tibia; inner spur of hind tibia not quite reachingmiddle of basal segment of hind tarsus; claws of all legs powerful; lobe almost as long as claw proper, a verydeep cleft between it and claw proper (Fig. 54). Caster unusually long and narrow (Fig. 62). Tergite 1 abouttwice as long as its apical width, shiny, with a weak, almost shagreened sculpture in which longitudinalelements are virtually absent. Basal half of tergites 2+3 with faint traces of scaly-reticulation distal to thehardly indicated curved furrow common to all species. Ovipositor sheath about as long as gaster pluspropodeum; bristle-like hairs towards apex of sheath not different from those elsewhere. Hypopygiumlong, sharply pointed. MATERIAL EXAMINED Holotype $, Greece: Ilia, Olympia, 4-11. vii. 1979 (Day, Else & Morgan) (BMNH).Paratype. 1 $, same data. HOST. Unknown. COMMENTS. This is an elegant species, differing in several significant characters from the other Microdusspecies dealt with in this revision. Particularly characteristic is the sculpture of the first gastral tergite. This,and the bend of the radius, suggest some affinity with Baeognatha Kokujev but this genus differs in nothaving a closed areolet in the fore- wing. It is possible that the loss of the second transverse vein inBaeognatha is apomorphic and in itself does not preclude a relationship with Microdus eriphyle. Microdus linguarius Nees von Esenbeck(Figs 32, 66) Microdus linguarius Nees von Esenbeck, 1814: 190. Syntypes, GERMANY (lost).Agathis linguarius (Nees von Esenbeck), Shenefelt, 1970: 340.Microdus linguarius Nees von Esenbeck, Tobias, 1971: 260. Marshall (1885: 270) was the first reviser of linguarius and I have no doubt that his interpretation is correct.Nees von Esenbeck 's description is also explicit on the details that define linguarius. Cf $,5-0-5-5 mm (excluding ovipositor). $ . Black. Basal half of tergites 2+3 frequently marked with reddish; sometimes whole surface reddishexcept for a large, isolated, elliptical black patch at base; more often pale colouring restricted to a lateralmark on each margin and sometimes whole of tergites 2+3 black. Hind femur red; hind tibia reddish butinfuscate on about apical quarter. Head from in front distinctly lengthened (Fig. 32). Inner margin of each antennal socket produced toform an angular tooth. Antenna with 28-31 segments, usually 30. Preapical segment of labial palpus (3rd)well developed, slightly longer than wide. Notaulices well defined throughout. Sternaulus of usual form.Propodeum coarsely rugose-reticulate all over; in this respect, somewhat uncharacteristic of genus;frequently with 2 indistinct, longitudinal keels. Radius of fore- wing slightly bent inwards towards stigma;areolet triangular or even stalked; stigma shorter than in clausthalianus but hardly different from that oftumidulus. Prepectal margin strongly raised, with knife-like edge. Outer side of middle tibia with 1-3isolated teeth in about apical third. Gaster less elongate than in clausthalianus, much as in tumidulus.Tergite 1 only a little longer than apically wide, finely striated to within about apical third where striationsfade and give way to a broad, transverse band of setiferous punctures. Hairs of tergites somewhatcharacteristically short so that gaster has an almost bare, highly polished appearance. Ovipositor sheathfully 1-25 times longer than body. Hypopygium widely, moderately deeply emarginate at apex; in profileappearing markedly truncate (Fig. 66), rather sparsely hairy. Cf . Antenna with 28-29 segments. Gaster sometimes entirely black. Striation of tergite 1 more variablein intensity than in female, sometimes almost wanting. MATERIAL EXAMINED Great Britain: 1 $>, Berkshire, Boxmoor, 29-30.vii.1933 (R. B. Benson) (BMNH); 1 cf, Buckingham-shire, Gog Magog Hills, 27.vii.1937 (R. L. E. Ford) (BMNH); 1 $, Devon, Chudleigh, 14.viii.1935 (/. F.Perkins) (BMNH); 1 $, Hampshire, Portsdown, 4.viii.l956 (/. Clark) (BMNH); 1 $, I. of Wight, Cowes,20.xi.1938 (R. C. L. Perkins) (BMNH); 1 cf , Kent, Shoreham, 15.viii.1958 (R. L. E. Ford) (BMNH); 1 cf ,1 $, Surrey, Banstead (D. Clark) (BMNH); 49 , Hackhurst Downs, 27-28. viii. 1982 (A. A. Allen} (AA, 224 G. E. J. NIXON 12 in BMNH); 2 cf, Sussex, Stonecross, 26.vii.1946 (R. L. E. Ford) (BMNH). Hungary: 1 cf, 1 ,Baranya-megya, Nagyharsane, vii.1963 (L. Zombori) (HNHM); 1 $, Somberek, 22.vii.1964 (L. Zom-bori) (HNHM); 10 $, 3 cT, Kam, Jeli arboretum (J. Papp) (HNHM); 2 , Keszthely, vii.1977 (J. Papp);1 Cf , 1 ?, Bajansenye, 16.viii.1977 (/. Papp) (HNHM); 1 $, F6t.Ret, 16.viii.1960 (Mihalyi) (HNHM). HOST. Unknown. COMMENTS. This is an easily recognised species and is characterised by the relatively elongate head, as seenfrom in front, the angular projection of the inner margin of the antennal socket and, in the female,especially by the emarginate hypopygium. This last feature is diagnostic for linguarius and it is unknown inother species from the region. Dates of capture for the numerous examples examined indicate that linguarius occurs only in July andAugust; it is probably single brooded. Microdus clausthalianus (Ratzeburg) Ichneumon (Microdus) clausthalianus Ratzeburg, 1844a: 25. Holotype $, GERMANY (lost).Microdus clausthalianus (Ratzeburg) Ratzeburg, 18446: 58. Agathis clausthalianus (Ratzeburg) Kloet & Hincks, 1945: 234: Shenefelt, 1970: 326.Microdus clausthalianus (Ratzeburg) Tobias, 1971: 260. Cf $, 6-0-6-2 mm (excluding ovipositor). 9 Black, rarely with a pale mark against and behind eye. Hind femur always entirely reddish; hind tibiausually same colour except for weak infuscation in apical quarter; but sometimes hind tibia faintlyyellowish and contrasting with red femur and, when thus pale, then with faint, prebasal band ofinfuscation; a pale-coloured hind tibia always correlated with an entirely red hind coxa; hind coxa varyingfrom entirely red to entirely black. Labrum and mandible varying from entirely yellowish to obscurebrownish red, or even black (exs. from Sweden: Skane, Fjellfota sjo). Face punctate but punctation variable in intensity; when weak, face more shiny. Antennal socketsalmost completely margined behind. Ocelli somewhat variable in size; when largest, they are in a lowtriangle with posterior tangent to the anterior ocellus touching posterior pair and distance between anteriorand posterior ocellus less than diameter of ocellus. Prepapical segment of labial palpus (3rd) about as longas wide. Antenna with 37-40 segments in examples with red hind coxa: 37 (4), 38 (7), 39 (4), 40 (2); with33-36 segments in examples with blackened hind coxa; 33 (1), 34 (1), 35 (5), 36 (1). Mesoscutum somewhatelongate. Notaulices deeply impressed throughout. Sternaulus extending forwards almost exactly tomiddle of mesopleurum. Radial cell markedly elongate. Outer side of middle tibia with 3-4 teeth in a row.Gaster elongate, more so than in the tumidulus-aggregate. Tergite 1 with weak, longitudinal striationwhich is frequently so weak as to be almost absent. Ovipositor sheath fully as long as body. Cf . Like female except for sexual differences. MATERIAL EXAMINEDGermany, Great Britain, Holland, Ireland. HOST. Epiblema scutulana (Denis & Schiffermuller) (Tortricidae) (1 $, ex stem of Cirsium, 17. iv. 1979(P. R. Sokoloff) (MS); 1 $, Berwick, Oxton, ex stem of thistle sp., 10.iii.1979 (K. Bland) (MS)). COMMENTS. Most specimens of clausthalianus are fairly easily recognised by their slender form and longovipositor. Smaller examples are sometimes hard to separate from tumidulus (s.l.). A prolonged study ofboth species has failed to reveal the limits of the range of variation in either. Microdus tumidulus Nees von Esenbeck(Fig. 58) Microdus tumidulus Nees von Esenbeck, 1814: 189. Syntypes, GERMANY (lost). Microdus tegularis Thomson, 1895: 2231. LECTOTYPE $, SWEDEN: Palsjo (ZI), here designated [examined]. Syn. n. Agathis tumidulus (Nees von Esenbeck) Shenefelt, 1970: 362.Microdus tumidulus Nees von Esenbeck; Tobias, 1971: 260. This species is very like clausthalianus. The main differences between the two species have been given inthe key; there is very little to add. $. Hind coxa mostly black, rarely entirely red. Antennal sockets unmargined behind; surface betweensockets virtually smooth except for a weakly impressed, median, longitudinal groove. Antenna with 27-30 A REVISION OF THE EUROPEAN AGATHIDINAE 225 segments. Sternaulus ending at middle of mesopleurum. Hind femur stouter than in clausthalianus , 3 timesas long as greatest width; outer side of middle tibia with 2-4 teeth towards apex, in addition to small clusterof teeth at extreme apex. Fore-wing (Fig. 58). Ovipositor sheath equal to length of gaster plus propodeum.Cf . Like female except for sexual differences. MATERIAL EXAMINEDFrance, Germany, Great Britain, Greece, Ireland, Italy, Yugoslavia. HOST. Dichrorhampha acuminatana (Lienig & Zeller) (Tortricidae). Reinhard (1867: 354) recordstumidulus as having been bred from the pupae of this moth (under the name Phthoroblastis acuminatana)in August on Chrysanthemum leucanthemum L. COMMENTS. This species, which may be an aggregate, has been correctly interpreted (s.l.) by all majorworkers on Microdus. It is easily recognised by the pale mark behind the eyes - a striking example ofconstancy in a colour-character. The notes given above cover the majority of the specimens examined, including Thomson's tegularis.There is, however, a series of 13 specimens in BMNH that, in certain respects, are intermediate betweentumidulus and clausthalianus. On the whole the habitus is more slender than that of typical tumidulus andthe ovipositor is longer, almost as long as the whole body. The antennal sockets are sometimes completelymargined behind, sometimes smooth and evenly hollowed out; the surface between the sockets is usuallyraised in the form of an evenly rugose platform but more often the surface here is weakly bipartite, beingdivided by a feeble channel. Antenna with 29-33 segments. The material is from the following localities.Germany: 4 $, Ruthe coll. (BMNH). Great Britain: 1 $, Surrey, Banstead, 29.vi.1952 (BMNH); 1 $,Hampshire, Stockbridge, 15.vi.1953 (R. L. E. Ford) (BMNH). France: 1 $, Pyrenees-Orientales,Vernet-les-Bains, 20.vi.1963 (BMNH). Sweden: 1 $, Skane, Silvakra, viii.1976 (T. Huddleston & J.Quintan) (BMNH); 4 $, Upsala, Solna, viii.1976 (T. H. & J. Q.) (BMNH); these last mentioned 4specimens have the hind tarsus markedly blackened and the area between the antennal sockets showingmaximum rugosity. Switzerland: 1 $, St Gall, Lake Zurich, ix.1951 (BMNH). Microdus conspicuus Wesmael(Figs 30, 61) Microdus conspicuus Wesmael, 1837: 17. LECTOTYPE $, BELGIUM: Brussels, Wesmael coll. (IRSNB), here designated [examined] . Microdus arcuatus Reinhard, 1867: 353. Syntype cf , GERMANY (MNHU) [examined]. Syn. n.Earinus zonatus Marshall, 1885: 268. Syntypes, GREAT BRITAIN (lost). [Synonymised by Lyle, 1920: 184.]Agathis conspicua (Wesmael) Muesebeck & Walkley, 1951: 119.Microdus conspicuus Wesmael; Tobias, 19766: 98. Cf $, 5-0-5-5 mm long (excluding ovipositor). Black; greater part of gaster bright reddish yellow. Legsalmost entirely reddish yellow; hind coxa black; hind femur entirely reddish yellow. Antenna of 9 palebeneath on more than basal half. In the male, base of tergites 2+3 sometimes with a blackened patch, andthree apical tergites may also be darkened. 9- Head in facial view strongly transverse. Third segment of labial palpus hardly discernible. Antennawith 34-35 segments. Notaulices deeply impressed throughout. Sternaulus rugose. Propodeum usuallywith two discernible keels but these usually obscured by coarse rugosities. Radius of fore- wing markedlybent inwards towards stigma and here tending to be colourless (Fig. 61); areolet sharply triangular andoften with conspicuous stalk. Outer side of middle tibia in apical half with group of 5-8 thick teeth; innerspur of hind tibia not quite reaching middle of basal segment of hind tarsus. Gaster as in tumidulus. Tergite1 about 1-33 times longer than its apical width; finely, evenly striate all over, the surface appearing veryshiny; basal half of tergites 2+3 sometimes with traces of fine rugose-striation tending to be concentricaround the feebly raised, transverse, basal swelling. Apical sternite (hypopygium) evenly pointed at apex.Ovipositor sheaths a little shorter than body. Cf . Gaster somewhat darker than in female. Sometimes eye-orbits, both on inner side and temples, palerin colour. MATERIAL EXAMINEDBelgium, Finland, France, Great Britain, Hungary, Ireland, Italy, Sweden, Switzerland. HOSTS. Shenefelt (1970) lists the following: Conchy Us nitidulana Zeller (?? Gypsonoma nitidulana Lienig& Zeller) (Tortricidae); Pammene regiana (Zeller) (Tortricidae); Cydia molesta Busck (Tortricidae), a 226 G. E. J. NIXON species which, according to Meyrick (1968: 590) has been introduced into North America, Australia andEurope. Bred specimens from Cydia pomonella (L.), Pammene regiana (Zeller) and Rhopobota ustomaculana(Curtis) (all Tortricidae) have been examined. It seems that Tortricidae are the main, if not the only, hosts. COMMENTS. This species has some economic importance because of its parasitism on Cydia pomonella theCodling moth. As a species it is chiefly characterised by its bright colour and bent radius. Nevertheless, itbears a strong superficial resemblance to zaykovi but the latter has a differently shaped head, both in facialview and profile. Microdus zaykovisp. n. Cf 9, 4-5-5-0 mm (excluding ovipositor). Black. Gaster, except for 1st tergite, reddish yellow but surfacebecoming slightly infuscate on posterior half in the paratype. Hind coxa black; hind femur red; hind tibiahardly paler than hind femur and faintly darkened only at extreme apex. Wings deeply infuscate. 9. Head in facial view like that of linguarius (cf. Fig. 32); in lateral view, hind margin of head deeplyhollowed out at a point level with lowest point of eye; in same view of head, cheeks behind markedlyangled. Surface between antennal sockets virtually flat; no trace of a projection on inner side of antennalsocket as in linguarius. Antennal scrobes broader than in linguarius, reaching close to inner eye-margin andmuch closer to posterior ocellus. Posterior tangent to the anterior ocellus passing far in front of posteriorpair in the paratype; almost touching them in the holotype. Mesoscutum and notaulices of ordinary form.Sternaulus reaching middle of mesopleurum. Radius of fore-wing slightly more strongly curved inwardstowards stigma than in linguarius, with stigma more narrowed towards apex and radial cell slightly longerthan in that species. Outer side of middle tibia with one tooth at extreme apex in holotype, just behind the2-3 closely spaced teeth at the edge; no preapical teeth in the paratype; hind tarsus with apical lobe smallerthan in linguarius, the claw itself hence appearing longer and less curved. Gaster rather short and broad;tergite 1 considerably widened towards apex, about 1-25 times longer than its apical width; on posteriorhalf almost smooth and polished in the two females available. Hypopygium not emarginate at apex and inprofile angled at about 60 degrees. Ovipositor sheath much shorter than in linguarius, hardly as long asthorax and gaster combined. Cf . Like the female in all essential details but apical half of gaster blackened. In one of the two paratypes,there is a large pale oblong spot behind the eye (much as in tumidulus). Colour probably variable. MATERIAL EXAMINED Holotype $, Bulgaria: Rhodope, Nikolovo, 19.viii.1976 (Zaykov) (ZC). Paratypes. Bulgaria: 1 $, Rhodopi, 'h.Ruen', 29.vii.1960 (A. Germanov) (BMNH); 1 cf, Nikolovo,15.viii.1976 (Zaykov) (ZC); 1 cf , Bojno, 24.vii.1975 (Zaykov) (ZC). COMMENTS. The most characteristic feature of this very dark-winged species is the hollowing out of the backof the head when this is seen in profile. In colour it bears a superficial resemblance to conspicuus underwhich the differences are pointed out. EAjR/M/SWesmael Earinus Wesmael, 1837: 8. Type-species: Microdus nitidulus Nees von Esenbeck (= elator (Fabricius)), bysubsequent designation (Muesebeck & Walkley, 1951: 116). DIAGNOSIS. Head in facial view not lengthened, like that of Microdus. Third segment of labial palpusvirtually twice as long as wide. Ocelli in low triangle; posterior tangent to anterior ocellus cutting posteriorpair. Notaulices absent but mesoscutum behind with elongate furrow. Sternaulus absent. First discoidalcell separated from 1st cubital cell by a fully pigmented, sclerotised vein; areolet of fore- wing 4-sided.Outer side of middle tibia with row of 4-6 teeth; outer side of front tibia with at least one tooth.Metasternum behind not completely surrounding hind coxal insertion. Ovipositor in the three speciesunder consideration not longer than gaster plus propodeum. Key to species (females) 1 Head and thorax densely hairy; face strongly, conspicuously punctate, somewhat flattened oneach side of a broad, longitudinal ridge; mesoscutum usually black but sometimes reddened elator (Fabricius) (p. 227) - Head and thorax with ordinary pubescence; face shiny and at most with fine punctation; a longitudinal, facial ridge hardly indicated 2 A REVISION OF THE EUROPEAN AGATHIDINAE 227 2 Ovipositor sheath two-thirds as long as gaster; hairs of ovipositor sheath longer, thicker, more bristly trans versus Lyle (p. 228) Ovipositor sheath about as long as gaster plus propodeum; hairs of ovipositor sheath shorter,thinner, less bristly gloriatorius (Panzer) (p. 227) Earinus elator (Fabricius) Banchus elator Fabricius, 1804: 128. Lectotype d" , AUSTRIA (ZM), designated by Fitton, 1985 [examined].Microdus nitidulus Nees von Esenbeck, 1814: 187. Syntypes, GERMANY (lost). Syn. n.Microdus thoracicus Nees von Esenbeck, 1834: 143. Syntypes, GERMANY (lost). Syn. n.Earinus nitidulus (Nees von Esenbeck) Reinhard, 1867: 351.Earinus thoracicus (Nees von Esenbeck) Shenefelt, 1970: 407. O" $ , 7-8 mm long (excluding ovipositor). $ . Black but mesoscutum sometimes reddened. Legs entirely red except for infuscate hind tarsi. At leastthe two apical segments of maxillary palpus yellowish. Head characteristically clothed with long pubescence. Face conspicuously punctate, the puncturestending to be separated by slightly less than their diameter. Antennal sockets united on inner side to form araised, hump-like area, densely punctate. Scrobes with some rugose-punctation. Antenna with 36-40segments. Thorax densely hairy. Mesoscutum sharply, distinctly punctate. Mesopleurum closely, sharplypunctate, thickly pubescent over almost entire surface. Propodeum at middle with 2-3 very irregular,longitudinal keels; surface on each side of keels tends to be smooth and polished. Areolet of fore-wingsubquadrate, large, frequently with extra vein in form of stub arising from middle of outer side of 2ndcubital vein; distal abscissa of postmarginalis about 0-5 times as long as proximal abscissa. Outer side ofmiddle tibia with 3-4 teeth arranged in row; front tibia with 4-5 teeth along outer side; inner spur of hindtibia slightly less than half as long as basal segment of hind tarsus. Gaster, except for tergite 1 which isvaguely rugose-punctate to rugose-striate, highly polished and only very sparsely hairy; ventral surface andlaterotergites by contrast thickly hairy. Ovipositor sheaths about as long as gaster plus propodeum, denselybeset with thick, bristle-like setae. Cf . Like female except for sexual differences. MATERIAL EXAMINED Austria: 2 d 1 (including lectotype), Fabricius coll. (ZM), Germany: 3 d", 2 <j>, Ruthe coll. (BMNH); 2 $,Celle, 23. v. 1942 (BMNH). Great Britain: 1 $, Berkshire, Woolhampton, ex Agrochola lota Clerck, coll.30.V.1978, em. 22.iv.1979 (M. Shaw) (MS); 1 $, Devon, Dartmoor, 23.iv.1924 (J. F. Perkins) (BMNH); 2$, Newton Abbott, 28.iv.1928 (R. C. L. Perkins) (BMNH); 1 $, Tiverton (F. H. Lyon) (BMNH); 2 $,Hertfordshire, Brickett Wood, 24.iv.1948 (R. B. Benson) (BMNH), 1 cf, Aldbury, 2.iv.l949 (R. B.Benson) (BMNH) ; 2 $ , Lancashire , nr Nutford , Hawes Wood , ex Agrochola circellaris (Hufnagel) in Salixcatkins, coll. 7.iv.l977, em. 19.iii.1978 (M. Shaw) (MS); 1 cf, Shropshire, Church Stretton (D. K.Mckevan) (BMNH); 8 cf , 5 $ , Surrey, Salfords, beaten from privet (Ligustrum), 15.iii.1981 (A. A. Allen)(AA; BMNH); 1 Cf , 3 $ , Sussex, Plaistow, King's Park Wood, ex Agrochola lota Clerk on Salix caprea,coll. 20.V.1978, em. 19.iv.1979 (M. Shaw) (MS); 1 <j>, Sussex (no other loc.), ex Orthosia xerampelinaHiibner (BMNH); 2 $, Lothian, Edinburgh, 16.iv.1981 (BMNH). Ireland: 1 cf, Dublin, Glenasmole,5.iv.l936 (Stelfox) (USNM); 1 cf, Wicklow, Glencullen, 25.iii.1932 (Stelfox) (USNM). Sweden: 2 $,Skane, 28-30.iv.1938 (Perkins) (BMNH). HOSTS. Agrochola circellaris (Hufnagel); Agrochola lota Clerck on Salix caprea; Atethmia centrago(Haworth) (all Noctuidae). The stout cocoon of the parasite is barrel-shaped, white, papery. All the aboverecords indicate that elator is single-brooded and occurs from March to May. COMMENTS. This is a very distinct species on account of its unusual hairiness and the thick setae of thefemale ovipositor sheaths. Females with a red mesoscutum are particularly easy to recognise. Earinus gloriatorius (Panzer)(Figs 67, 68) Bassus gloriatorius Panzer, 1809: 102. Syntypes, GERMANY (lost).Microdus ochropes Curtis, 1829: 105. Nomen nudum. [See ochropes Lyle, 1920.]Microdus gloriatorius (Panzer) Haliday, 1833; 263. [First reviser.]Earinus gloriatorius (Panzer) Haliday in Westwood, 1839: 63. Microdus affinis Wesmael, 1837: 11. LECTOTYPE , BELGIUM: Brussels, Wesmael coll. (IRSNB), heredesignated [examined]. [Synonymised by Marshall, 1885: 267.] 228 G. E. J. NIXON Microdus varicoxis Wesmael, 1837: 10. LECTOTYPE $, BELGIUM: Brussels, Wesmael coll. (IRSNB), here designated [examined]. Syn. n.Microdus delusor Wesmael, 1837: 12. LECTOTYPE $, BELGIUM: Brussels, Wesmael coll. (IRSNB), here designated [examined]. Syn. n.Microdus tuberculatus Wesmael, 1837: 13. LECTOTYPE $ , BELGIUM: Brussels, Wesmael coll. (IRSNB), here designated [examined]. Syn. n.Agathis bicingulatus Thomson, 1895: 2234. LECTOTYPE $, SWEDEN: Ortofta, Thomson coll. (ZI), here designated [examined]. Syn. n.Earinus ochropes Lyle, 1920: 248. LECTOTYPE $, GREAT BRITAIN (Dale) (UM), here designated [examined]. [Attributed to Curtis, 1829.] Syn. n. C? $ , ca 5 mm (excluding ovipositor). $. Black. Hind coxa almost always red, sometimes bicoloured, rarely entirely black; hind tibiasometimes yellowish with infuscate apex and infuscate basal ring; more often reddish throughout. Facesmooth, shining, rarely with traces of punctation towards antennal insertions. Antennal scrobes polished,smooth. Antenna with 32-36 segments. Mesoscutum with only moderately thick pubescence. Virtually notrace of notaulices but usually mesoscutum with a longitudinal furrow posteriorly. Prepectal margin almostobliterated medially. Propodeum on the whole smooth, shining and with two longitudinal keels of variablestrength; lateral panels sparsely hairy. Areolet of fore-wing always distinctly 4-sided; usually somewhatnarrowed towards stigma (Figs 67, 68); rarely more obviously quadrate and with an accessory vein arisingfrom middle of outside of 2nd transverse cubitus as in elator. Claws of hind tarsus more strongly bent thanin elator and with deeper lobe; middle tibia with 3-4 teeth on outer side; fore tibia with 3-4 teeth on outerside . Tergite 1 about 1-33 times longer than its apical width ; vaguely sculptured on apical half but sculpturevarying much in intensity. Tergites 2+3 with variable sculpture; at base usually with a more or less distinct,transversely triangular area that is slightly raised and generally smooth and shining; distal to this area andas far as second suture, the amount of sculpture is highly variable; sometimes it appears as fine rugosity,sometimes as rugose-striation; sometimes whole of tergites 2+3 entirely smooth. Ovipositor sheath aboutas long as gaster plus propodeum; its setae much less thickened than in elator and, towards apex, shorter,finer and more adpressed. Cf . Like female except for sexual differences. MATERIAL EXAMINEDBelgium, Germany, Great Britain, Holland, Hungary, Ireland, Sweden, Switzerland. HOST. Agonopteryx ciliella (Stainton) or Agonopteryx heradiana (L.) (Oecophoridae) on AngelicasylvestrisL. (MS). COMMENTS. Earinus gloriatorius is an abundant species but shows a confusing variation in the colour of thehind legs and the rugosity of the gaster. Like elator, with which it could not be confused, it occurs in spring.Wesmael's three species synonymised above and about whose separate validity he had some misgivings arerecorded by him as having been captured towards the end of April. Earinus transversus Lyle Earinus transversus Lyle, 1920: 249. LECTOTYPE $; GREAT BRITAIN (Dale) (UM), here designated[examined] . $, 6-5 mm long (excluding ovipositor). Black. Hind coxa and hind femur red; hind tibia bright yellow,deeply infuscate in apical third and with faint post-basal infuscate spot; hind tarsus deeply infuscatethroughout. Face smooth, shining, finely punctate. Antennal scrobes polished. Antenna with 38 segments. Pubesc-ence of mesoscutum thicker than in gloriatorius but less so than in elator. Mesoscutum with some vague,indistinct punctation. No trace of notaulices. Propodeum shorter than in gloriatorius, with denserpubescence, more even and somewhat characteristically directed backwards; lateral panels appearingduller because of fine punctation. Areolet of fore-wing markedly 4-sided, narrowed towards stigma. Fronttibia with a single tooth on outer side; middle tibia with 3 teeth on outer side; hind claw much as in elator,less bent than in gloriatorius . Tergite 1 about 1-25 times longer than its apical width, with shining, indefinitesculpture much as in gloriatorius. Tergites 2+3 shining, with rather coarse, variable rugosity extendingvirtually to apex of segment; at base with weakly defined, transverse, subtriangular area. Ovipositor sheath0-66 times as long as gaster, beset with bristly hairs, thicker than in gloriatorius and almost as thick as thoseof elator. A REVISION OF THE EUROPEAN AGATHIDINAE 229 MATERIAL EXAMINED Great Britain: 1 $ (lectotype), 'type', 'exPolycommata', Dale coll. (UM); 1 cf , 'type', Dale coll. (UM);1 $, 'cotype', '1899' Dale coll. (UM); 1 cf , 'cotype', Dale coll. (UM). HOST. The label 'ex Polycommata' may refer to Trichopteryx polycommata (Denis & Schiffermiiller)(Geometridae). BAEOGNATHA Kokujev Baeognatha Kokujev, 1903: 243. Type-species: Baeognatha turanica Kokujev, by monotypy.CamptothlipsisEnderlein, 1920: 166. Type-species: Camp tothlipsiscostalisEnder\ein, by original designa-tion. [Synonymised by Tobias, 1976a: 214.] DIAGNOSIS. Head in facial view like that of Microdus, transversely elliptical. Surface between antennalinsertions slightly raised in the form of a weak hump or tubercle in longitudinal direction or simply roundedfrom back to front. Mouth parts not lengthened. Notaulices distinct throughout. Sternaulus very distinct.Outer side of middle tibia with at least 2 teeth remote from apical margin; inner spur of hind tibia notreaching middle of basal segment of hind tarsus; claws lobed. Areolet of fore-wing open (Fig. 60). Tergite 1and basal half of tergites 2+3 with or without sculpture. Ovipositor at least as long as gaster. The genus is typically Microdus-like in habitus but differs essentially in that the 2nd submarginal cell isnot closed externally by a vein (r-rri) so that an areolet is absent. Microdus eriphyle approaches Baeognathain the kind of sculpture on the basal tergites of the gaster. Key to species (females) 1 Ovipositor sheath about as long as gaster; basal segment of hind tarsus slightly shorter than following segments together; hind femur blackish nigra Telenga (p. 230) - Ovipositor sheath about as long as body; basal segment of hind tarsus distinctly longer than following segments together; hind femur bright yellow armeniaca Telenga (p. 229) Baeognatha armeniaca Telenga (Fig. 60)Baeognatha armeniaca Telenga, 1955: 300. Holotype $, U.S.S.R.: Armenia (AS) [not examined]. $, 4 mm long (excluding ovipositor). Head mainly bright yellowish. Thorax variable in colour but withmuch pale marking; sometimes only mesoscutum and pronotum pale with rest of thorax infuscate. Gastervarying from entirely reddish yellow (except tergite 1) to almost black. Legs bright reddish yellow exceptfor hind coxa which may show darkening at base. Wings virtually hyaline; medius colourless proximal tobasalis. Head in facial view transverse, only very weakly narrowed towards mouth. Face smooth, shining.Between antennal sockets a smooth, blunt keel. Preapical segment (3rd) of labial palpus very small, hardlylonger than wide. Ocelli in rather low triangle with posterior tangent to anterior ocellus virtually touchingposterior pair. Notaulices deeply impressed throughtout. Sternaulus deeply impressed, reaching posteriorcorner of mesopleurum. Propodeum coarsely rugose-reticulate with only the faintest indication of twobroken, longitudinal keels. Distal abscissa of postmarginalis about 1-5 times longer than proximal abscissa(Fig. 60). Outer side of middle tibia with 1-2 teeth at middle and two close teeth at apex; inner spur of hindtibia not quite reaching middle of basal segment of hind tarsus; basal segment of hind tarsus distinctlylonger than remaining segments together, 6:5; claws small, with large, basal lobe. Tergite 1 about twice aslong as apically wide, with a dull, even sculpture of scaly-reticulation, the surface having almost ashagreened appearance with only a few weak, longitudinal ridges towards sides. Basal half of tergites 2+3with a sculpture similar to that of tergite 1 but weaker and more vague; rest of gaster polished, shining.Ovipositor sheath about as long as body, less head. MATERIAL EXAMINED Austria: 1 $, Neusiedlersee, 20.viii.1960 (G. J. Kerrich) (BMNH). Turkey: 1 $, Ankara, Kavaklidere,6.viii.l960 (Guichard & Harvey) (BMNH). U.S.S.R.: 3 $, ? Armenia, exAnarsia eleagnella Kuznetzov(AS; one in BMNH). HOSTS. Anarsia eleagnella Kuznetzov (Gelechiidae); Cydiafunebrana Treitschke (Tortricidae) ; Recurvar-ia nanella (Denis & Schiffermiiller) (Gelechiidae). These records from Tobias (1976: 214).COMMENTS. This species is included on the strength of the single female recorded above from Austria. Itdiffers from the more eastern specimens in being a little larger and much darker in colour, the head beingmainly reddish yellow with darkened occiput and a dark patch in the middle of the face; the thorax and 230 G. E. J. NIXON gaster above are entirely blackened and the hind coxa is pale only on the apical third; the relation betweenlength of basal segment of hind tarsus and combined length of remaining segments has not been checkedbecause the legs are broken.The specific differences between armeniaca and nigra have been given in the key. Baeognatha nigra Telenga Baeognatha nigra Telenga, 1955: 300. Syntypes <j>, U.S.S.R. (AS) [not examined].The interpretation of this species is based on a single female , identified by Tobias and borrowed from him. $, 3 mm long (excluding ovipositor). Black. Inner orbits brownish yellow, the yellow colour reachingantennal sockets. Hind femur infuscate; hind tibia infuscate on apical third, with faint, dark ring near base;hind coxa blackish. Head in facial view slightly more narrowed below than in armeniaca. Face smooth, shining. Betweenantennal sockets a smooth, blunt ridge as in armeniaca. In dorsal view head slightly less transverse than inarmeniaca. Antenna broken in both examples. Notaulices and sternaulus as in armeniaca. Radial cellslightly narrower than in armeniaca; distal abscissa of postmarginalis only slightly longer than proximalabscissa. Teeth of middle tibia and spur of hind tibia as in armeniaca; basal segment of hind tarsus slightlyshorter than following segments together, 9:11. Gaster slightly less elongate than in armeniaca butsculpture of tergite 1 and basal half of tergites 2+3 similar, though that on tergites 2+3 is sharper and betterdefined than in armeniaca and the curved furrow is slightly deeper and more distinct. Ovipositor as long asgaster. MATERIAL EXAMINED U.S.S.R.: 1 $, Kazachstan, Urals, Ganvartsjevo (AS). France: 2 $, Mulhouse, Bois de Nonnenbruch,vi.1977, ex larvae of Coleophora ? flavipennella (Duponchel) on Quercus (S. E. Whitbread) (BMNH). HOSTS. Coleophora ? flavipennella (Duponchel). The cocoon of the parasite is spun within the case of thehost. The distributional data on this species are scant but suggest that Baeognatha nigra is widespread. Species inquirendae Type-material of the following species is not available for examination.Microdus abbreviator Ratzeburg, 1852: 45.Microdus brevicaudis Reinhard, 1867: 356.Microdus cingulator Ratzeburg, 1852: 46.Agathis initiator Fonscolombe, 1846: 40.Vipio insularis Snellen van Vollenhoven, 1873: 192.Agathis major Fonscolombe, 1846: 39.Ichneumon purgatorFabricius, 1793: 156. Acknowledgements I thank the following for the loan of material: Dr C. van Achterberg (Leiden); Dr A. A. Allen (Reigate);Dr R. Danielsson (Lund); Dr P. Dessart (Brussels); Dr Max Fischer (Vienna); Dr E. Haeselbarth(Munich); Dr F. Koch (Berlin); Dr Paul Marsh (Washington); Dr J. Papp (Hungary); Dr Atti Pekkarinen(Helsinki); Dr B. Petersen (Copenhagen); Dr Mark Shaw (Edinburgh); Dr V. I. Tobias (Leningrad); DrA. Zaykov (Bulgaria). I am grateful to Barry Bolton and Michael Day, Department of Entomology, British Museum (NaturalHistory), for invaluable help in matters of nomenclature. Very special thanks go to Tom Huddleston of thesame department for painstakingly checking the typescript and making whatever corrections werenecessary to bring it into line with editorial requirements. I am grateful also to Dr Laurence Mound,Keeper of Entomology, for providing me with facilities to work in his department. Finally, I acknowledge my indebtedness to the Royal Society and the Leverhulme Trust for the financialsupport that has enabled me to accomplish this project. References Achterberg, C. van 1976. A preliminary key to the subfamilies of the Braconidae (Hymenoptera).Tijdschrift voor Entomologie 119: 33-78. A REVISION OF THE EUROPEAN AGATHIDINAE 231 1982. Notes on some type-species described by Fabricius of the subfamilies Braconinae, Rogadinae,Microgasterinae and Agathidinae (Hymenoptera: Braconidae) Entomologische Berichten 42: 133-139.1984. Essay on the phylogeny of Braconidae (Hymenoptera: Ichneumonoidea). Entomologisk Tidskrift 105: 41-58.Bhat, Shama & Gupta, V. K. 1977. The Subfamily Agathidinae (Hymenoptera: Braconidae). Oriental Insects no. 6: 353 pp.Bradley, J. C. 1919. The synonymy and types of certain genera of Hymenoptera, especially of those discussed by the Rev. F. D. Morice and Mr Jno. Hartley Durrant in connection with the long forgotten 'Erlangen list' of Panzer and Jurine. Transactions of the Entomological Society of London 1909: 50-75.Curtis, J. 1829. A guide to an arrangement of British insects vi pp + 256 columns. London.Dondale, C. D. 1954. Biology ofAgathis laticinctus (Cress.) (Hymenoptera: Braconidae) a parasite of the Eye-spotted Bud Moth, in Nova Scotia. The Canadian Entomologist 86: 40-44.Eady, R. D. 1974. The present state of nomenclature of wing venation in the Braconidae (Hymenoptera): its origins and comparison with related groups. Journal of Entomology (B) 43: 63-72.Enderlein, G. 1920. Zur Kenntniss ausser europaischer Braconiden. Archiv fur Naturgeschichte. 84(A) (1918) (11): 51-224.Fabricius, J. C. 1775. Systema Entomologicae. 1-832 pp. Flensburg & Leipzig. 1798. Supplementum Entomologiae Systematicae. 225 pp. Copenhagen. 1804. Systema Piezatorum. 439 pp. Brunsvigae. Fahringer, J. 1937. Opuscula braconologica IV. Palaearktische Region 3 (4-6): 402-510. Fischer, M. 1957a. Zur Kenntniss der Thomson'schen Braconiden- Arten. Entomologisches Nachrichten- blatt Oesterreichischer und Schweizer Entomologen 9: 10-11.19576. Beitrage zur Kenntniss palaarktischen Braconiden (Hymenopteren). Mitteilungen derMiinch- ner Entomologischen Gesellschaft 41 ': 1-21. 1966. Geziichtete Braconiden aus Niederosterreich und aus dem Burgenland (Hymenoptera). Zeitschrift fur angewandte Zoologie 53: 385-402.Fitton, M. G. 1978. The species of 'Ichneumon' (Hymenoptera) described by Linnaeus. Biological Journal of the Linnean Society 10: 361-383.1985. The Ichneumon-fly genus Banchus (Hymenoptera) in the Old World. Bulletin of the British Museum (Natural History) (Entomology) 51: 1-59.Forster, A. 1862. Synopsis der Familien und Gattungen der Braconen. Verhandlungen des naturhistoris- chen Vereins des Preussischen Rheinlandes 19: 225-288. Fulmek, L. 1962. Parasitinsekten der Blattminierer Europas. 203 pp. Den Haag.Gupta, V. K. 1964. Agathis festiva Muesebeck, a new braconid parasite of the Lac predator Holcocera pulverea, in India (Insecta, Hymenoptera, Braconidae). Current Science 33: 220.Haliday, A. H. 1833. An essay on the classification of the Parasitic Hymenoptera of Britain which corresponds with the Ichneumones minuti of Linnaeus. Entomological Magazine 1: 259-276.Hellen, W. 1956. Zur Kenntniss der Agathidinen Finnlands (Hym., Brae.). Notulae Entomologicae 36: 116-125.Ivanov, P. 1899. Braconides cryptogastres et areolaires des environs de Koupiansk avec tableaux synoptiques des genres et des especes de ces insectes. Trudy Obshchestva Ispytatelei Prirody pri Imperatorskom Khar'kovskom Universitete 33: 273-382.Kloet, G. S. & Hincks, W. D. 1945. A check list of British insects, lix + 483 pp. Stockport. 1972. Handbooks for the identification of British insects 9 (2): Lepidoptera 1-153.Kokujev, N. 1895. Fragments Braconologiques IV-V. Horae Societatis Entomologicae Rossicae 29: 363-392.- 1903. New Transcaspian species of the subfamily Agathidinae (Hymenoptera, Braconidae). [In Russian.] Trudy russkago entomologicheskago Obshchestva 36: 240-247.Kriechbaumer, J. 1898. Ueber Diophrys caesa Klg. und inculcatrix auct. nebst einer neuen Art dieser Gattung. Entomologische Nachrichten 24: 181-185.Krombein, K., Hurd, P., Smith, D. & Burks, B. (Eds) 1979. Catalog of Hymenoptera in America North of Mexico. 2735 pp.Latreille, P. A. 1804. Nouveau dictionnaire d'histoire naturelle. 258 pp. Paris. 1805. Histoire naturelle generate et particuliaire des crustaces et des insectes. 13: 175. Paris. Linnaeus, C. 1758. Systema naturae. Edn 10, 1: 563. Stockholm. Lyle, G. T. 1920. Contributions to our knowledge of British Braconidae. No. 6. , - Agathidae. Entomolo-gist 53: 177-186, 248-250.Marsh, P. M. 1961. A taxonomic study of the genus Cremnops Forster in America north of Mexico (Hymenoptera, Braconidae). Annals of the Entomological Society of America 54: 851-861. 232 G. E. J. NIXON Marshall, T. A. 1885. Monograph of British Braconidae. Part I. Transactions of the Entomological Society of London 1885: 1-280. 1888. In Andre, E., Species des Hymenopteres d' Europe et d'Algerie 4: 609 pp. Beaune.Meyrick, E. 1928 [facsimile reprint, 1968] A revised handbook of British Lepidoptera. 914 pp. London.Muesebeck, C. F. W. 1927. A revision of the parasitic wasps of the subfamily Braconinae occurring in America north of Mexico. Proceedings of the United States National Museum 69: 1-73.Muesebeck, C. F. W. & Walkley, L. M. 1951. In Muesebeck et al. (Eds), Hymenoptera of America north of Mexico. Agriculture Monograph no. 2: 1420 pp.Nees von Esenbeck, C. G. 1814. Ichneumonides adsciti, in genera et familias divisi. Magazin der Gesellschaft Naturforschender Freunde zu Berlin 6 (1812): 183-221.1834. Hymenopterorum Ichneumonibus affinium monographiae, genera Europaea et species illus- trantes. 1: 320 pp. Stuttgart & Tubingen.Niezabitowski, E. 1910. Materyaly do Fauny Brakonidow Polski. Braconidae, zebrane w Galicyi. Sprawozdania Akademii umiejetnosci w Krakowie 44: 47-105.Panzer, G. F. W. 1809. Fauna Insectorum Germanicae 9: 102.Ratzeburg, J. T. C. 1844a. Die Forst-Insecten oder Abbildung und Beschreibung der in den Wdldern Preussens und der Nachbarstaaten als schadlich oder nutzlich bekannt gewordenen Insecten; In systema- tischer Folge und mil besonderer Rucksicht aufdie Vertilgung der Schadlichen 3: 314 pp. Berlin.1844ft. Die Ichneumonen der Forstinsecten in forstlicher und entomologischer Beziehung 1: 224 pp. Berlin. 1852. Die Ichneumonen der Forstinsecten in forstlicher und entomologischer Beziehung 3: 272 pp. Berlin.Reinhard, H. 1867. Beitrage zur Kenntniss einiger Braconiden-Gattungen. Berliner Entomologischer Zeitschrift 11: 351-374.Richards, O. W. 1977. Hymenoptera. Introduction and key to families. 2nd edn. Handbooks for the Identification of British Insects 6 (1): 100 pp.Rossem, G. van 1969. A revision of the genus Cryptus Fabricius s.str. in the western Palaearctic region, with keys to genera of Cryptina and species of Cryptus (Hymenoptera, Ichneumonidae). Tijdschrift voor Entomologie 112: 299-374.Shenefelt, R. D. 1970. Hymenopterorum Catalogus (nov. ed.) Part 6. Braconidae 3: 307-428. S'Gravenhage. Shestakov, A. 1932. Zur Kenntniss der asiatischen Braconiden. Zoologischer Anzeiger 99: 255-263.Simmonds, F. J. 1947. The biology of the parasites of Loxostege sticticalis L. in North America - Bracon vulgaris (Cress.) (Braconidae, Agathinae). Bulletin of Entomological Research 38: 145-155.Spinola, M. 1808. Insectorum Liguriae species novae out rariores, quas in agro Ligustico nuper detexit, descripsit et iconibus illustravit (Hymenoptera) 2: 262 pp. Genuae. Szepligeti, G. V. 1904. Hymenoptera, Fam-Braconidae. In Wytsmann, P., Genera Insectorum 22: 253 pp.Telenga, N. A. 1955. Fam. Braconidae, subfamilies Microgasterinae and Agathinae. [In Russian.] Fauna SSSR (Hymenoptera) 5 (4): 312 pp. Thomson, C. G. 1895. Bidrag till Braconidernas Kannedom. Opuscula Entomologica 20: 2141-2339.Thorpe, W. H. 1933. Notes on the natural control of Coleophora laricella, the larch case-bearer. Bulletin of Entomological Research 24: 271-291.Tobias, V. 1. 1962. New genera of Braconids in the fauna of the USSR. (Hymenoptera, Braconidae). [In Russian with English summary.] Zoologischeskii Zhurnal4l: 1190-1197. - 1963. The species of the genus Agathis Latr. (Hymenoptera, Braconidae) from Kazakhstan andMiddle Asia. Entomologicheskoe Obozrenie 42: 864-883. 1964. On two new species of the genus Agathis Latr. (Hymenoptera, Braconidae) from the Caucasus. [In Russian.] Izvestiya Akademii Nauk Armyanskoi SSR 17 (3): 59-66.1971. Review of the Braconidae (Hymenoptera). [In Russian.] Trudy Vsesoyuznogo Entomo- logicheskogo Obshchestva 54: 156-268. - 1976a. Braconidae of the Caucasus. [In Russian.] Opredeliteli po Faune SSSR 110: 1-286. 1976ft. Contribution to the knowledge of the Far Eastern Braconids of the genus Microdus Nees (Hymenoptera, Braconidae). [In Russian.] Trudy biologo- pochvennogo Instituta AN SSSR 43: 96-106.Waltl, J. 1835. Reise durch Tirol, Oberitalien und Piedmont nach dem sudlichen Spanien. 120 pp. Passau.Wesmael, C. 1837. Monographie des Braconides de Belgique. Nouveaux Memoires de V Academic Royale des Sciences et Belles-Lettres de Bruxelles 10: 1-70.Westwood, J. O. 1840. An introduction to the modern classification of insects 2: 587 pp.; Synopsis 158 pp. London. A REVISION OF THE EUROPEAN AGATHIDINAE 233 Figs 1-6 Agathis species, heads in facial view. 1, ariadnemalvacearum $ ; 5, syngenesiae $ ; 6, anchisiades $. 2, assimilis $; 3, glaucoptera $; 4, 234 G. E. J. NIXON 12 Figs 7-12 Agathis species, heads in facial view. 7, zaykovi $ ; S,polita $ ; 9, montana $ ; I0,fulmeki $ ; 11,nigra $ ; 12, pappei $ . A REVISION OF THE EUROPEAN AGATHIDINAE 235 15 16 18 Figs 13-18 Agathis species, heads in facial view. 13, semiaciculata $ ; 14, achterbergi $ ; I5,pedias $ ; 16,varipes $ ; 17, rufipalpis $ ; 18, anglica $ . 236 Figs 19-26 Agathis species. 19, assimilis $ , head, lateral; 20, syngenesiae $ , head, dorsal; 21 , montana $ ,head, lateral; 22, anchisiades $, head, dorsal; 23, polita $, head, dorsal; 24, artemesiana $, head,dorsal; 25, assimilis $, head, dorsal; 26, malvacearum $, head, dorsal. A REVISION OF THE EUROPEAN AGATHIDINAE 237 29 Figs 27-32 27-29, Agathis species, heads in facial view. (27) meridionellae <j>; (28) glabricula $; (29)minuta $. 30, Microdus conspicuus $, head, facial; 31, Agathis asteris $, head, facial; 32, Microduslinguarius , head facial. 238 G. E. J. NIXON 33 36 40 Figs 33-42 Agathis species. 33, tibialis $ , hind femur; 34, zaykovi 9 , galea; 35,fulmeki 9 , mid tibia; 36,rufipalpis 9, thorax, lateral; 37, asteris 9, thorax, lateral; 38, nigra 9, thorax, lateral; 39, pedias $,gaster, dorsal; 40, zaykovi 9, gaster, dorsal; 41, glabricula $, thorax, lateral; 42,fulmeki 9, thorax,lateral. 43 52 A REVISION OF THE EUROPEAN AGATHIDINAE 44 \ 45 239 46 47 49 53 50 54 Figs 43-55 43-53, Agathis species. (43) semiaciculata $, hind claw; (44) griseifrons $, hind claw; (45)nigra $, hind claw; (46) glabricula $, hind claw; (47) zaykovi $, hind claw; (48) tibialis $, hind claw;(49) anglica $, part of forewing; (50) varipes $, part of forewing; (51) fulmeki $, mouthparts; (52)breviseta $ , ovipositor sheath, dorsal; (53) tibialis 9 , hind tibial spurs and basal segment of hind tarsus.54, 55. Microdus species. (54) nugax $, part of forewing; (55) cingulipes $, part of forewing. 240 G. E. J. NIXON 56 57 61 62 Figs 56-62 56-59 Microdus species. (56) eriphyle $ , hind claw; (57) lugubrator $ , part of forewing; (58)tumidulus $ , part of forewing; (59) lugubrator $ , base of antenna. 60, Baeognatha armeniaca $ , part offorewing. 61, 62, Microdus species. (61) conspicuus $ , part of forewing; (62) eriphyle $ , gaster, dorsal. A REVISION OF THE EUROPEAN AGATHIDINAE 241 64 67 68 Figs 63-68 63-66, Microdus species. (63) lugubrator $ , head, dorsal; (64) eriphyle $ , head, dorsal; (65)fortipes 9, head, dorsal; (66) linguarius 9, apex of gaster, lateral. 67, 68, Earinus gloriatorius $. (67)part of forewing; (68) part of hind wing. 242 G. E. J. NIXON Index Invalid names and species inquirendae are in italics. abbreviator 230abscissus 217achterbergi 208a/finis 227Agathis 192albicostellae 209anchisiades 207anglica 200anthradna 190arcuatus 225ariadne 206armeniaca 229artemesiana 210assimilis 198asteris 209 Baeognatha 229bicingulatus 228brevicaudis 230breviseta 197 caesa 190calculator 217Camptothlipsis 229cingulator 230cingulipes 221clausthalianus 224conspicuus 225Cremnops 191 deftagrator 192delusor 228deserter 192dimidiator 219Disophrys 190 Earinus 226elator 227eriphyle 222 fortipes 220fulmeki 198 glabricula 209glaucoptera 195gloriatorius 227gracilipes 212griseifrons 202 initiator 230insularis 230 kolazyi 197 laticarpa 202linguarius 223lugubrator 219 major 230malvacearum 202melpomene 213meridionellae 210Microdus 215minuta 208montana 213 nasicornis 214nigra (Agathis) 203nigra (Baeognatha) 230nitidulus 227nugax 222 ochropes 228 pappei 212pedias211persephone 206polita 206pumilus 221purgator 230 Rhamphagathis 214rostrata 211rufipalpis 199rufipes 218rugulosus217 semiaciculata 205syngenesiae 196 taurica 204tegularis 224testaceipes 203thoracicus 227tibialis 201transversus 228tuberculatus 228tumidulus 224 umbellatarum 197 varicoxis 228varipes 199 zaykovi (Agathis) 204zaykovi (Microdus) 226zonatus 225 Occasional Papers on Systematic Entomology New Series The economic importance of insects, and the enormous number of species, have resulted in avast literature written in many languages; that which is of direct economic importance andrecently published can, to an increasing extent, be searched by using computerized data bases,but a great amount of more general information is unlikely to be available so readily in thenear future. The objective of this new occasional series is to make available in hard copy someof the basic data that is essential to the preparation of comprehensive accounts of the worldinsect fauna. The papers have been fully researched bibliographically and consist of checklistsof nominal taxa, and faunal lists with information on host plants and localities, based mainlyon the collections and libraries of the British Museum (Natural History). No. 1. Catalogue of the Neotropical Tiger-moths. A. Watson & D. T. Goodger 72pp. inc. 4 colour plates 27 February 1986 No. 2. An annotated checklist of the Carabidae (including Cicindelinae, Rhysodinae and Paussinae) recorded from Borneo. N. E. Stork 26pp . , 1 map 27 March 1 986 Catalogues of BM (NH) Bulletins and Books free on request to Publication Sales. Titles to be published in Volume 52 The sand Hies of Egypt (Diptera: Phlebotominae) By R. P. Lane Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae) By G. S. Robinson A revision of the European Agathidinae (Hymenoptera: Braconidae) By G. E. J. Nixon A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revision of certain generaBy J. Quinlan Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester 27 Bulletin of the British Museum (Natural History) A key to the Afrotropical genera ofEucoilidae (Hymenoptera) , with arevision of certain genera J. Quinlan Entomology series Vol 52 No 4 26 June 1986 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History),Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1986 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06018 XISSN 0524-6431 British Museum (Natural History)Cromwell RoadLondon SW7 5BD _ Us ^-tr.cs .^^ - ^ Entomology seriesVol 52 No 4 pp 243-366 Issued 26 June 1986 A key to the Afrotropical genera of Eucoilidae(Hymenoptera), with a revision of certain genera J. Quintan Department of Entomology, British Museum (Natural History), Cromwell Road, LondonSW7 5BD Contents Synopsis 243 Introduction 243 Material examined and terminology 244 Depositories 244 Affinities of Eucoilidae with other Cynipoidea 245 Taxonomic characters 246 The generic classification of Eucoilidae and discussion of relationships 247 Checklist of the Afrotropical Eucoilidae included in this paper 249 Eucoilidae Thomson 250 Key to the genera of Afrotropical Eucoilidae 250 Afrodontasp Weld 253 Cothonaspis Hartig 255 Ealatagen.n 257 Eucoilidea Ashmead 259 Hexacola Foerster 272 Kleidotoma Westwood 280 Nordlanderia gen . n 288 Rhoptromeris Foerster 290 Stentorceps Quinlan 309 TrichoplastaEenoit 310 Acknowledgements 322 References 322 Index... 365 Synopsis A key to the 19 genera of Afrotropical Eucoilidae is provided, together with keys to the species of tengenera. Two genera (Ealata, Nordlanderia) and 98 species are newly described; the primary types of eightnominal species have been examined, of which two are newly synonymized. A checklist of the ten generaand 111 species dealt with is included. Introduction The virtually cosmopolitan family Eucoilidae is one of the largest and most distinctive groups ofCynipoidea. These small, mostly blackish insects are easily recognizable by the presence of ahighly modified scutellum (Fig. 2) . They occur wherever their dipteran hosts are found. As far asis known (and the biology of many species has never been investigated) eucoilids are proteleanendoparasites of the larvae and puparia of cyclorrhaphous Diptera. A number of species attackvarious phytophagous Diptera in the families Agromyzidae, Anthomyiidae, Chloropidae,Drosophilidae and Tephritidae (Thompson, 1955), families that include a variety of seriousagricultural pests. Consequently several eucoilids are of considerable economic importance aspotential biological control agents. A number of species in the genera Hexacola, Trybliographa,Rhoptromeris, Kleidotoma and Ganaspidium are of interest since they parasitize species of Bull. Br. Mus. not. Hist. (Ent.) 52 (4): 243-366 Issued 26 June 1986 244 J. QUINLAN Liriomyza, Oscinella and Hylemyia which attack cereal and related crops. Trybliographa dadWeld is parasitic on the oriental fruit-fly (Dacus dorsalis Hendel) which is a well-known pest of avariety of fruit and vegetable crops. Probably the most extensively studied eucoilid is Tryblio-grapha rapae, a common parasite of Hylemyia species which attack brassicas (Wishart &Monteith, 1954). In Europe Rhoptromeris heptoma (Hartig) is an important parasite of the fritfly (Oscinella frit (L.)) (Nordlander, 19786). A variety of other species of Eucoilidae attack dipterous larvae inhabiting dung and carcasses,including species of the families Calliphoridae, Muscidae, Sepsidae and Sphaeroceridae(Thompson, 1955). For example, the European eucoilid Kleidotoma marshalli Cameron isknown to parasitize larvae of a variety of Calliphora, Lucilia and Musca species, whilst Eutriastritoma (Thomson) has been reared in the U.S.A. from sepsid puparia in cow dung (Weld,1952). Various authors have published papers on the Afrotropical Eucoilidae: Benoit (1956),Bridwell (1919), Cameron (1904), Hedicke (19280, 19286), Kieffer (1919, 1913), Masner (I960)and Risbec (1950, 1956). Some have described new genera and species, others only species.Apart from Kieffer (1910) and Weld (1952), who give keys to genera on a world basis, no one hasattempted to key genera and species of the Afrotropical Eucoilidae. During the preparation ofthis paper all available type-material has been examined and redescribed. When type-materialof a taxon was not available its presumed taxonomic position is discussed in the text. A key to thegenera of Afrotropical Eucoilidae is given and ten of these genera are revised with keys tospecies; two genera and 98 species are described as new. One genus, Stentorceps, is included in ageneric key for the first time. A further nine genera remain to be revised. Material examined and terminology I have been fortunate to have on loan large amounts of material from Dr J. Decelle, MuseeRoyal de 1'Afrique Centrale, Tervuren. This material, together with R. E. Turner's collectionand other more recently obtained specimens (principally from Malaise traps) in the BritishMuseum (Natural History), has formed the basis for this revision. Very large areas of theAfrotropical region remain from which material has not been available for examination. In veryfew instances are host data given and then only from the labels attached to the specimens. Theterminology follows Richards (1977) but some older names are used, particularly in reference towing venation. I have, where pertinent, related older terminology to the more recentlyintroduced terminology of Nordlander (19826). The term Afrotropical Region used in thispaper was coined by Crosskey & White (1977) but was not employed in my earlier paper of 1979.Figures 1 and 2 have their respective parts labelled in abbreviated form, as follows: anterior parallel lines - apl; hairy ring - hr; hypopygium - hy; lateral bar - Ib; lateral lines - 11;lateral ridges - Ir; mesoscutum - m; mesopleurae - mp; mesopleural suture - mps; malar ridge -mr; metapleurae - mt; notaulices - nt; nucha - nu; propodeum - pd; pronotal plate - pp;pronotum - pr; scutellum - sc; scutellar cup - sec; scutellar disc - scd; scutellar foveae - sf;tegulae - tg. The following terms are used for the lines or furrows on the mesoscutum, anterior parallel lines,notaulices and lateral lines. Nordlander (1982) used the terms antero-admedian lines, notaulicesand parapsidial furrows respectively. Depositories Type depositories are abbreviated in the text as follows. BMNH British Museum (Natural History), London. GERDAT Groupement D'Etudes et de Recherches pour le developpment de L' Agronomic Tropi- cale, Montpellier. MNHN Museum National d'Histoire Naturelle, Paris. MRAC Musee Royale de 1'Afrique Centrale, Tervuren. NCISA National Collection of Insects, Division of Entomology, Pretoria. AFROTROPICAL EUCOILIDAE 245 USNM United States National Museum of Natural History, Washington, D.C. ZMC Zoologisk Museum, Copenhagen. ZSBS Zoologische Sammlung des Bayerischen Staates, Munich. Affinities of Eucoilidae with other Cynipoidea Amongst the Cynipoidea many of the higher taxa are definable in terms of biological as well asstructural differences. Known biologies may be summarized thus: FAMILY-GROUP TAXON HOST Ibaliidae Siricoidea (Chrystal, 1930; Spradberry, 1970) Liopteridae Oberthuerellinae Unknown Liopterinae Unknown Mesocynipinae ? Cerambycidae (Diaz, 1973) Eucoilidae Cyclorrhaphous DipteraFigitidae Figitinae Cyclorrhaphous Diptera (Askew, 1971) Aspicerinae Syrphidae (Weld, 1952) Anacharitinae Hemerobioidea (Selhime & Kanavel, 1968) Himalocynipinae Unknown Cynipidae Cynipinae Phytophagous gall-causers and gall inquilines (Eady & Quinlan, 1963) Austrocynipinae Reared from seeds ofAraucaria (Araucariaceae) (Riek, 1971)Charipidae Alloxystinae Aphidiinae and Aphelinidae (as hyperparasites through Homoptera)(Quinlan & Evenhuis, 1980) Charipinae Psylloidea (Kierych, 1979) The Eucoilidae is distinguished from the five other families of Cynipoidea by the highly modifiedscutellum. This comprises an outer disc surmounted by a cup- or tear-drop-shaped elevation inthe centre with a foveolate depression posteriorly. The family is apparently most closely relatedto the Figitidae which, except for the form of the scutellum, they resemble in structure. The twofamilies have rather similar life-histories, in that most species are endoparasites of larvae andpuparia of Cyclorrhaphous Diptera, although the morphologically distinctive Anacharitinaeparasitize hemerobioid lacewing larvae (Selhime & Kanavel, 1968; Lipkow, 1969). TheEucoilidae and Figitidae, together with the Cynipidae and Charipidae, have segments 2 and/or 3of the gaster enlarged, whilst the more posterior segments (4+) are the largest in other cynipoids(Quinlan, 1979). Tergite 1 is variable in shape from a long petiolate structure to an almostobscured crescent-shape, wider than long. The elongate petiolate form does not occur in theCynipidae or Charipidae, but is found in the Eucoilidae and Figitidae. Males of most familieshave much less specialized gasters. Whilst the Eucoilidae are easily separated from otherfamilies by the form of the scutellum, some eucoilid genera do show apomorphic featuresexhibited by genera in other families. For example, a strongly indicated ring of hairs is present atthe base of tergite 2 in many eucoilid genera (e.g., Trybliographa, Rhoptromeris and Kleido-toma), whilst a less conspicuous ring of hairs is found in certain genera of Cynipidae (e.g.,Aulacidea and the related genera Ay lax and Isocolus). A sparser ring of pubescence is also foundin some Charipidae. The plant-feeding habits of the Cynipidae are in contrast to the endoparasitic entomophagy ofthe Eucoilidae and Figitidae. At first sight, therefore, the shared characteristics of the gaster ofthese taxa would appear to be the result of convergence, though it is possible that the Cynipidaeadapted to their herbivorous mode of life from a figitid ancestor consuming the larvae of leaf- orstem-mining Diptera. 246 J. QUINLAN Taxonomic characters Head The female antenna is 13-segmented (occasionally 11, 12 or 14) and the overall shape is ofimportance, both at generic and specific level, but must be used in combination with othercharacters. Some species have filiform antenna, others are distinctly clavate. In those specieswith filiform antenna the term 'club' refers specifically to those apical segments of the antennathat bear rhinaria (Figs 229, 230), even though they may not appear swollen. In those specieswith clavate antenna the club is very clearly swollen as well as each segment having rhinaria (Figs237, 256). The male antenna is 15-segmented, filiform and in some species the third or fourthsegment is modified. This is of major importance at generic and specific level. The characters ofthe head have not been used to any great extent at the generic level although the sculpture of theocciput and the presence of striations around the malar area, the size and shape of the compoundeye (the distance between them in relation to the height of an eye), the variation in the shape ofthe vertex viewed posteriorly, and the occipital carina are important, particularly at specieslevel. Thorax The thorax (including the first abdominal segment of the propodeum, vide Richards, 1977) hasmany important characters. As mentioned above, the pronotal plate is extremely diverse inshape and form. The mesoscutum is invariably polished; notaulices, anterior parallel lines andlateral lines are either present, partially present or completely absent. Sometimes a line of hairsis present and replaces the notaulices. These structures are referred to under different names byNordlander (19826). The mesopleuron exhibits a wide variety of shapes and in the majority ofgenera the mesopleural carina or suture is present, either partly or completely. This structurehas also been referred to as the mesopleural furrow (Eady & Quinlan, 1963). In some genera thesubalar pit is very distinct whilst it is absent in others. The metapleuron varies in the number oftransverse ridges present and in some species an anteroventral cavity is present, either with orwithout setae. The scutellum viewed dorsally comprises a central plate and an outer margin,'cup' and 'disc' of earlier authors, termed scutellar plate and dorsal surface of the scutellumrespectively by Nordlander (19826). As indicated earlier, the cup varies in size and shape and inthe number of foveae (if any) present around the outer margins. In most genera a single largefovea is present medially on the apical half of the cup. The scutellar disc is variable in both dorsaland, in some genera, lateral views, its surface is sometimes smooth and polished or in others avariety of sculpture is present. The shape of the scutellar disc at the apex is either rounded,conical or spine-shaped; in a few genera the apex is bicuspid. Many earlier authors have placedthose species with a conical or spine-shaped scutellar disc in the same genera without regard tomore fundamental characters. The propodeum in dorsal view invariably has carinae varying inshape and alignment to each other, with varying densities of sculpture medially, and pubescenceon either side of the carinae. The nucha at the posterior end of the propodeum is obscure butgenerally ridged. Gaster The gaster comprises eight segments and has spiracles on segments 1 and 8 (Richards, 1977).The number of segments visible in lateral view varies from genus to genus. Segment 1 of thegaster is short, about as broad as long, normally smooth or crenulate, except for a fewNeotropical genera in which it is long and narrow, in some cases longer than the combinedlengths of the remaining segments. The second segment is always the largest, generally as largeas the remaining visible segments combined. In some genera the base of the second tergite has aring of pubescence obscuring the first tergite, either in lateral view or completely on the dorsalsurface as well. The overall shape of the gaster is variable within genera and it is laterallycompressed to a lesser or greater extent. The hypopygium of the female is moderately to AFROTROPICAL EUCOILIDAE 247 strongly projecting. The legs vary little within the Eucqilidae except that the size ratio of coxae,femora, tibiae and tarsi varies from species to species. In some genera the presence of hairpatches on the mid-coxae has proved to be of value at species level (Nordlander, 1982a). Wings The venation of the fore wing of Cynipoidea has been used extensively at family, generic andspecific level. Earlier authors gave much emphasis, particularly with eucoilids, to the radial cell,placing great reliance on whether it was closed or open on the wing margin. In many instances itis hard to decide one way or the other, the venation being vestigial (Day, 1984), having linearconvexity in dorsal view, being concave ventrally, thus showing no clear or distinct longitudinalboundary when viewed with transmitted light, being only pigmented and not sclerotised. Size The overall size of Eucoilidae varies from genus to genus, ranging from 1-5-5-0 mm. The generic classification of Eucoilidae and discussion of relationships Earlier authors (Dalla Torre & Kieffer, 1910; Weld, 1952) tried to delimit genera on superficialcharacters, such as the radial cell being either open or closed, the presence or absence of a ring ofhairs at the base of tergite two of the gaster, the scutellum either with or without a spine on theapical margin or the apex rounded, truncate, "conical or emarginate. This resulted in anassemblage of genera containing species not necessarily related to each other. Nordlander (19826) in his phylogenetic classification of the Eucoilidae, recognised 28 validgenera which he divided into the following monophyletic groups. 1. Gronotoma-group comprising Zaeucoila, Gronotoma, Diglyphosema, Disorygma, Micro- stilba. 2. Trybliographa-group comprising Trybliographa, Eucoila, Bothrochacis. 3. Rhoptromeris-group comprising Leptopilina, Cothonaspis, Rhoptromeris, Trichoplasta. 4. Chrestosema-group comprising Chrestosema, Glauraspidia, Pseudopsichacra, Odon- teucoila, Dieucoila, Leptolamina. 5. Ganaspis-group comprising Ganaspis, Tetramerocera, Paramiomoea, Pentamerocera, Coneucoela, Didyctium, Hexacola, Hypodiranchis. 6. Kleidotoma-group comprising Eutrias, Kleidotoma. At present I recognise 19 of these genera as occurring in the Afrotropical region. It is impossibleto elucidate Nordlander's generic parameters from his five papers sufficiently well to recognisegeneric relationships precisely. Quinlan (1978) established the significance of the pronotal plateat species level. In the present work further emphasis is placed on this feature at generic level,particularly as to whether or not the pronotal plate projects forward when viewed dorsally andlaterally, and to the presence of either enclosed or open lateral foveae either side of the medialarea of the plate. The present key to genera attempts to indicate the phylogenetic relationships.The genera Nordlanderia, Disorygma, Diglyphosema, Ealata and Eucoilidea all have a plesio-morphic form of pronotal plate, i.e. it does not project forward, being similar in manyrespects to that found in the Figitinae and the cynipine genera Aulacidea and Ay lax. Coupledwith this form of pronotal plate, the base of tergite two of the gaster never has a ring of densepubescence, at the most only a few sparse hairs on the lateral margins. The remaining genera,the majority, form a monophyletic group characterized by the apomorphic form of pronotalplate: In these genera the plate projects forward and is visible in dorsal and lateral view. Withinthis group, the genera Trichoplasta, Rhoptromeris and Stentorceps have the plate furthermodified, in that there is lateral fusion of the anterior and posterior parts so that the fovea eitherside of the pronotal plate are closed on the lateral margins (Figs 246, 351). Nordlander (1982a)was the first to draw attention to this shared character of Trichoplasta and Rhoptromeris. Insome species of this group the medial bridge between the foveae is obscure or in the form of a 248 J. QUINLAN transverse groove (Fig. 250). Stentorceps differs remarkably from Trichoplasta and Rhop-tromeris in having extraordinary protuberances on the head, a trumpet-shaped protrusion on thesupraclypeal area of the face (Figs 31, 32) and elongate mandibles (Fig. 290). Trichoplasta isprimarily separated from Rhoptromeris and Stentorceps by the elongated, almost spine-shapedapex of the scutellum. Rhoptromeris is further distinguished from Trichoplasta by the forewingvenation. All other genera with a projecting pronotal plate have the foveae either side of themedial area of the pronotal plate open and form separate groups of genera. Nordlander (1982a)published a cladistic analysis of Trybliographa, Leptopilina, Cothonaspis, Rhoptromeris andTrichoplasta using a matrix of 16 characters; the characters were divided into plesiomorphic(primitive) and apomorphic (derived) states and the character polarity determined throughoutgroup comparisons using the operational rule of Watrous & Wheeler (1981). Nordlanderconcluded that Leptopilina, Cothonaspis, Rhoptromeris and Trichoplasta had seven synapomor-phies in common and constitute a monophyletic group. I believe that the form of the pronotalplate clearly separates Rhoptromeris and Trichoplasta from Leptopilina and Cothonaspis.Masner (1958), in discussing the genus Ganaspis, relates it to Leptopilina, Rhoptromeris,Pseudeucoila (now a synonym of Trybliographa) and Odonteucoila. Masner (1960) included inhis key to Odonteucoila three species now recognised as Trichoplasta from the AfrotropicalRegion, the significance of the pronotal plate not being noted. I have included both Odon-teucoila and Coneucoela in the generic key although both genera appear to be restricted to theNeotropical Region; species originally described in both genera from the Afrotropical Regionhave recently been transferred to Trichoplasta (Nordlander, 19820). Weld (1952), as didMasner, characterised Ganaspis by the flat, mirror-like, elliptical scutellar cup. Quinlan (1978)erroneously based his interpretation of the genus on one of the species included in Masner's key,the holotype of the type-species, Ganaspis mundata Foerster not being available. Nordlander(1980), however, placed G. subnuda with Leptopilina heterotoma (Thomson) as a juniorsynonym. A female of the type-species was redescribed by Weld (1952). Ganaspis is a complexgenus and needs more research at species level before deciding its relationship to other genera.At present it is defined by the mirror-like, almost flat scutellar cup and the weakly conical apex ofthe scutellum. Leptopilina shares a number of characters with Cothonaspis, Rhoptromeris andTrichoplasta, and this is illustrated in Nordlander's cladogram referred to above. Prior toNordlander (1980), Leptopilina had hardly been referred to in the literature. Weld (1952) haddoubts about the placement of Leptopilina, the holotype of the type-species being a male, andsuggested that it could be a Ganaspis species. However, Leptopilina, although dissimilar,appears to be closest to Cothonaspis in the shape of the thorax and in the petiole (first segment ofthe gaster) which is widened posteriorly although it has a denser ring of hairs (sometimes thin) atthe base of tergite two. Cothonaspis (one of the earliest described genera in the Eucoilidae) hasmany characters in common with Rhoptromeris and Trichoplasta. It differs (apart from thepronotal plate character referred to above) primarily by the form of the first segment of thegaster, which is virtually hairless and carinate, and the base of tergite two which has only verysparse pubescence on the lower lateral margins. Hexacola and Kleidotoma have a number ofcharacters in common, such as, for example, the shape and sculpture of the scutellum. In bothgenera the cup is small and the scutellar disc is either longitudinally striate or reticulate-striate,but exceptionally it can be almost smooth. They differ mainly in the wing characters. InKleidotoma the apex of the wing is either incised or arcuate, and the radial cell of the forewing isopen on the margin and distinctively shaped (Fig. 8). In Hexacola the apex of the wing isrounded and the radial cell is decidedly closed on the margin (Fig. 164). Nordlander (19826)regards Hexacola as being in the Ganaspis group of genera (p. 247). Afrodontaspis has a numberof unique characters that do not indicate a clear relationship with any other group of genera, i.e. ,the scutellar cup is declined posteriorly so that it is not completely visible in dorsal view (Fig. 50) .The sides of the pronotum, mesoscutum, lateral bars of the scutellum and scutellar disc arestriate (Fig. 50). The scutellum is sharply pointed apically and the wing surfaces are dotted withhair bases similar to those found in Bothrochacis and Eucoila. In Weld's key (1952) it would runclose to Trissodontaspis , a Neotropical genus. It is distinguished from it, however, by the radialcell of the forewing being open and by the structure of the scutellum. Eucoila and Bothrochacis AFROTROPICAL EUCOILIDAE 249 both have almost bare wing surfaces (only a few hairs are present) and are separated from eachother by the striking apomorphic cup of Bothrochacis in which the scutellar cup itself is sharplydeclined apically . The surface of the cup is elevated in front of the posterior fovea so that, whenviewed dorsally, it appears declined apically. Trybliographa has world-wide distribution and isextremely rich in numbers of species; they are slightly larger in size than those of other generaexcept Bothrochacis and Eucoila. Nordlander (1981) synonomized eight Foerster genera,previously regarded as distinct, with Trybliographa. No keys are given to genera but from hisremarks on the generic relationships, Bothrochacis and Eucoila are related by a series ofcharacters regarded as symplesiomorphic. In common with Eucoila and Bothrochacis, thescutellar plate is large, almost circular in some species and well elevated, depressed in the centrewith a large apical fovea. The scutellar disc varies in degree and intensity of sculpture, frompunctate-reticulate-rugose. In a large number of species the lateral bars of the scutellum arestriate in part. The ring of pubescence at the base of tergite two of the gaster is dense andcomplete on the dorsal surface. The radial cell of the forewing can be open or closed.Nordlander (1982a) places Glauraspidia in the Chrestosema-group. Glauraspidia itself ischaracterized and distinguished from related genera by the absence of a mesopleural suture,although in some species it may be weakly indicated. The dense pubescence either side of thepronotal plate and on the entire metapleuron, and the density of the pubescence on thepropodeum together with a dense felt-like ring of pubescence at the base of tergite two of thegaster further distinguish this genus. Benoit (1956) described a monotypic genus Daruna,relating it to Rhoptromeris but distinguishing it by the rather unusual form of the scutellar cupwhich has two large foveae, one centrally and almost round, the other on the apex andtransverse. The surface of the scutellar disc is smooth and shining, the apex rounded. Thepronotal plate projects forward and the foveae are open on the lateral margins. The systematicposition of Daruna is unclear, but the form of the pronotal plate suggests that it is not closelyrelated to Rhoptromeris. Checklist of the Afrotropical Eucoilidae included in this paper EUCOILIDAE Thomson, 1862AFRODONTASPISWeld, 1961 lanatussp. n. striatissima Benoit, 1956COTHONASPIS Hartig, 1840 (Ink-is sp. n. ealissp. n. pentatoma Hartig, 1840EALATA gen. n. clava sp. n. marica sp. n. saha sp. n. EUCOILIDEA Ashmead, 1887Afrostilba Benoit, 1956 syn. n. ad vena sp. n. bucca sp. n. compressa sp. n. conversa sp. n. duhia sp. n. extraria sp. n. fetura sp. n. furcula sp. n. lacerta sp. n. /ana sp. n. leptissp. n. marce//ussp. n. maurisp. n. nitida (Benoit, 1956) comb. n. pallida sp. n. perangusta sp. n. trull a sp. n. tyrus sp. n. urundiensis Benoit, 1956HEXACOLA Foerster, 1869 absensa sp. n. amantia sp. n. atropos sp. n. hit'aria sp. n. compacts sp. n. fringa sp. n. hexatoma (Hartig, 1841) octoclava sp. n. pallida sp. n. quinqueclavata sp. n. quisnama sp. n. septemius sp. n. zama sp. n.KLEIDOTOMA Westwood, 1833 arbitra sp. n. bifurcate sp. n. cojiica sp. n. distends sp. n. eala sp. n. elongula sp. n. 250 J. QUINLAN erebussp. n. favussp. n. fimbriata sp. n. montana Kieffer, 1910 morsum sp. n. nigranssp. n. nitidiuscula sp. n. norms sp. n. strigosa sp. n. vcntosussp. n.NORDLANDERIA gen. n. acissp. n. pallida sp. n. plowa sp. n.RHOPTROMERISFoerster, 1869 abba sp. n. a/ersp. n. agissp. n. attissp. n. bicolorsp. n. bupalussp. n.p cepheus sp. n. connatus sp. n. crito sp. n. cubitalissp. n. diversa sp. n. enna sp. n. equalissp. n. nebe sp. n. heptoma (Hartig, 1840) iiaviu.vsp.n. na.vuvsp. n. oe-fa sp. n. pagasa sp. n. pallidus sp. n. persiussp. n. punctata sp. n. rii/ii/u.v sp. n. rutshurissp. n. rwanA/sp. n. sinissp. n. temesa sp. n. t hales sp. n. vt7/a sp. n. zetessp. n. zeus sp. n.SrEJVrOUCEPS Quinlan, 1984 fubicen Quinlan, 1984TRICHOPLASTA Benoit, 1956 hicolorsp.n. brevispina (Masner, 1960) conJca sp. n. contrasts sp. n. equalissp. n. f.vft'n.vussp. n. filiforinissp.n. gracilicornis (Kieffer, 1910) longispina (Masner, 1960) medlia sp. n. narrate sp. n. novetna sp. n. octonariussp. n. quinclava sp. n. ru/u.vsp. n. tanganyikensis(WQ\d, 1944)basilewskyi Benoit, 1956 testacea sp. n. unicolora sp. n. zeussp. n. EUCOILIDAE Thomson Eucoilidae Thomson, 1862: 397; Eady & Quinlan, 1963: 7. Type-genus: Eucoila Westwood [emended].Eucoilinae Thomson; Foerster, 1869: 186. The Eucoilidae is the largest and most widely distributed of the parasitic families in the Cynipoidea. Thespecies form a distinctive group, distinguished from other Cynipoidea by the presence of a raised cup orplate (Fig. 359), which varies in shape and sculpture. They are internal parasites of dipterous larvae,emerging from the puparia (Quinlan, 1978). A number of them are parasites of cereal crop pests and are ofsufficient economic importance to warrant close studies of their life histories (James, 1928; Imms, 1930,1932; Jenni, 1951; Nostvik, 1954; Kerrich & Quinlan, 1960; Masner, 1958, 1960; Carton, 1977; Carton,Roualt & Kitano, 1977; Nordlander, 19786; Barbotin, Carton & Kelner Pillault, 1979; Tsacas, 1979).Nineteen genera are at present known to occur in the Afrotropical Region. Key to the genera of Afrotropical Eucoilidae 1 Pronotal plate either distinct in anterodorsal view but not strongly projected forward (Fig. 18) ,or indistinct viewed dorsally , see Nordlanderia (Fig. 11), Eucoilidea (Fig. 83) . First tergite ofgaster, if visible, in form of a narrow ring, sometimes obscured by tergite 2 which never has ahairy ring at base, at most only a few sparse hairs on lower lateral margins; mesopleuron witha distinct suture, or weakly striate (Microstilba); mesoscutum shiny, notaulices present though sometimes aberrant or indicated by lines of hairs 2 Pronotal plate distinct both in anterodorsal and lateral views, i.e. projected forward frompronotum (Fig. 2). First tergite of gaster, if visible, sometimes in the form of a crenulate ring,without a dense ring of hairs (visible in Cothonaspis, Leptopilina etc.), or tergite 2 with a AFROTROPICAL EUCOILIDAE 251 dense ring of pubescence at base, not always complete on dorsal surface or sometimes withonly a few sparse hairs in place of hairy ring; mesopleuron generally with a suture though thiscari be aberrant or absent; mesoscutum smooth and shiny, satiny or granulate; notaulicesgenerally absent 6 First tergite of gaster not wholly visible in dry-mounted specimens either in dorsal or lateralview, at most in the form of a crescent, obscured by tergite 2 (Fig. 220); notaulices eitherfaintly indicated or with a row of hairs in their place, if visible, at junction with scutellumeither converging or parallel (Figs 17, 217) 3 First tergite of gaster usually visible in dorsal and lateral views, distinctly crenulate (Fig. 22);notaulices distinct, sometimes converging sharply on approaching scutellum (Figs 217, 218) 5 Malar space area and supraclypeal area with short protrusions (Fig. 16); area below mesopleu-ral suture coriaceous (Fig. 214); scutellar cup large, oval, excavate on lower half with anumber of smaller fovea round outer edge (Fig. 15), scutellar disc reticulate, scutellar foveaeaberrant; notaulices distinct, radial cell closed, almost as deep as wide; $ antenna clavate, cfantenna filiform, segment 3 curved, swollen distally NORDLANDERIA gen. n. (p. 288) Malar space and supraclypeal area without protrusions; area below mesopleural suturesmooth, polished; scutellar cup variable in shape, scutellar disc punctate-reticulate (Figs 17,21) ; notaulices distinct or aberrant; radial cell open or closed 4 Scutellar cup large, oval, with a pale rim, concave, punctate within the rim, scutellar discreticulate-rugose, malar space and supraclypeal area without protrusions; notaulices obso-lete anteriorly, widely separated at juncture with scutellar suture (Fig. 17). Radial cell of forewing generally open, as wide as deep (specimens of this genus with aclosed radial have been seen from Trinidad); 9 antenna weakly clavate, cf with segment 3weakly curved on outer margins DISORYGMA Foerster Scutellar cup small, polished; scutellar disc reticulate-punctate (Fig. 69); notaulices aberrant,at most represented by a row of hairs, pronotal plate adpressed but distinct, with a largelateral fovea on either side (Fig. 20) EALATA gen. n.(p. 257) Notaulices broadened apically, sculptured (Fig. 21); scutellar cup very large, concave, extend-ing to apex of scutellar disc, almost obscuring lateral margins of disc, basal margin of cup witha fringe of long setae, rim of cup edged with small fovea. First tergite of gaster distinct, in theform of a crenulate ring or collar, base of tergite 2 with a few scattered hairs (Fig. 22); $antenna subclavate, cf with segment 3 curved and swollen distally DIGLYPHOSEMA Foerster Notaulices not generally broadened apically, always smooth; scutellar cup not obscuringscutellar disc laterally (Fig. 74) though sometimes obscuring disc apically, scutellar cup witha large central fovea, sometimes with a ring of minute foveae around lower half and close toedge of cup. First tergite of gaster crenulate, tergite 2 generally the largest viewed laterally.$ antenna variable, from filiform to subclavate, cf antenna filiform, the 4th segment curved,swollen apically; malar space with a distinct groove, without striations on either side EUCOILIDEA Ashmead(p. 259) Scutellum produced apically to form a distinct spine or viewed dorsally and laterally conical(Figs 10, 350) and overhanging metanotum; pronotal plate with fovea on either side ofmedian bridge, enclosed or open (Figs 30, 228) 7 Scutellum not produced apically to form a spine or cone, either rounded, emarginate ortruncate, sometimes with a tooth on each side but not tridentate, if triangular, or conicalviewed dorsally, cup narrow, disc striate and wings emarginate 10 Scutellar cup narrow, declined posteriorly, almost reaching apex of scutellar disc (Fig. 10);radial cell of forewing open at base and apex (Fig. 9). Side margins of pronotum, mesoscutum and lateral bars striate; pronotal plate with lateralfovea open (Fig. 24) AFRODONTASPIS Weld (p. 253) Scutellar cup variable in shape, not reaching apex of disc nor declined; radial cell of forewingopen or closed on wing margin 8 Pronotal plate, viewed frontally, with a narrow or obsolete medial bridge, fovea on either sideenclosed laterally (Fig. 323). Scutellar disc sharply conical, sometimes almost spine- or beak-shaped, overhangingpropodeum (Fig. 292); $ antenna clavate-subclavate or with segment 4 of antenna longerthan 3, weakly curved on inner margin, sometimes swollen distally; cf antenna with segment3 shorter than 4, 4 longer than each of following segments; radial cell of forewing open orclosed (Figs 294, 299) TRICHOPLASTA Benoit(p. 310) 252 J. QUINLAN Pronotal plate, viewed frontally, with a wide medial bridge, lateral fovea between anterior and posterior parts open, not bounded by a lateral fusion (Fig. 26) 9 9 Head either aciculate, striate or matt on temples and/or vertex; scutellar cup small, occasion-ally raised; fovea above propodeal spiracle with hairs (Neotropical) ODONTEVCOILA Ashmead - Head smooth and shining; scutellar cup large, not raised; fovea above propodeal spiracle shallow and hairless. (Neotropical) CONEVCOELA Kieffer 10 Wing surface very lightly pubescent, at most with a few sparse hairs, most prominent on veins, apical half on both surfaces bare but dotted with hair bases on under surface, almost alwayswith short apical hair fringe on posterior margins; radial cell of forewing closed, R\ thinneron wing margin than Rs 2 (Fig. 27); pronotal plate, viewed dorsally, in form of collar, viewedanterodorsally , rounded , medial bridge large with lateral fovea open (Fig . 28) 11 - Wing surfaces densely pubescent, ciliate, with distinct hair fringe on apical margins; pronotal plate with lateral fovea open or closed , medial bridge variable in breadth (Fig. 26) 12 11 Scutellar cup narrow, declined posteriorly (the usual large posterior fovea not visible in dorsal view) (Fig. 29); base of forewing smoky, radial cell partially closed on wing margin;pronotum either side of pronotal plate canaliculate (Fig. 30); pronotal plate (Fig. 30); $antenna subclavate-clavate, cf antennal segments 3 and 4 modified BOTHROCHACIS Cameron - Scutellar cup large, rounded, elevated, not declined posteriorly (cf. Fig. 43); base of forewing clear, radial cell nearly always closed on wing margin; pronotal plate in the form of a collarviewed dorsally (Fig. 28); $ antenna moniliform, cf antennal segment 4 swollen distally EUCOILA Westwood 12 Pronotal plate with lateral fovea between anterior and posterior parts closed laterally (Fig. 246) 13Pronotal plate with lateral fovea open (Fig. 34) 15 13 Head with pyriform protuberances on inner orbits of face (Figs 31, 32), supraclypeal area of face with a large trumpet-shaped protrusion STENTORCEPS Quinlan (p. 309) Head without the above characters 14 14 Scutellar cup large, with a large central fovea and a small transverse fovea below it, scutellar disc smooth, polished; $ antenna filiform DARUNA Benoit - Scutellar cup small , long and narrow with minute pits , scutellar disc reticulate to almost smooth and polished; $ antenna invariably clavate, rarely filiform, club segments usually darkerthan basal segments. Anteroventral cavity of mesopleura distinct, some times reduced; propodeum denselypubescent ; radial cell of forewing closed on wing margin RHOPTROMERIS Foerster (p . 290) 15 First tergite of gaster distinct, in the form of a crenulate or furrowed ring, abruptly widened posteriorly. Second tergite of gaster with a few sparse hairs at base, never with a dense ring ofhairs (Fig. 33); scutellar cup elliptical, scutellar disc rounded behind, surface with weakreticulate sculpture, sides of propodeum lacking pubescence; wings narrow, roundedapically, radial cell of forewing closed on wing margin though sometimes indistinctly so,cubitus (M) not or barely indicated. $ antenna with or without a distinct club, cf antenna filiform, 4th segment sinuate, longerthan either 3rd or 5th (Fig. 360); pronotal plate with lateral fovea open (Fig. 60); anteroven-tral cavity of metapleura absent COTHONASPIS Hartig (p. 255) - First tergite of gaster not always visible , if visible , tergite 2 with a dense ring of hairs at base and wings broad; or base of tergite 2 with a distinct ring of hairs, though neither dense norcomplete on dorsal surface; scutellar cup variously shaped, disc sculpture variable, sides ofpropodeum with dense pubescence; wings broad, never narrow, radial cell of forewing openor closed, apex of forewing rounded or incised 16 16 Brachypterous or fully winged, apex of wings either incised, arcuate or truncate, cubitus (M) not distinct, radial cell of forewing distinctly open on wing margin, vein RI thickened at apexnear margin of forewing, 2rm thickened, areolet absent, Rs+M usually absent (Figs 8, 177)or wings short, not extending to apex of gaster; scutellar cup small, not extending to apex oflogitudinally striate scutellar disc which is either rounded, conical or in some species spine- orbeak-shaped. Tergite 2 of gaster with a hairy ring at base, not usually complete on dorsal surface (Fig.36); $ antenna clavate, cf antenna filiform, segment 3 elongate, curved, viewed dorsally,outer margin flattened (Fig. 225) KLEIDOTOMA Westwood (p. 280) - Fully winged , never brachypterous , apex of forewing rounded , rarely truncate , venation not as AFROTROPICAL EUCOILIDAE 253 above, radial cell open or closed on wing margin, scutellar cup and disc variable in size, shape and sculpture, disc rarely striate 17 17 Mesopleural suture either absent or indistinct (Fig. 37); frons prominent in cf, less so in $; wings in $ of normal length (generally shortened in British specimens); scutellar cup raisedabove level of scutellar disc which is striate to reticulate-rugose; sides of pronotal plate andpropodeum with dense tufts of pubescence (usually yellowish white); tergite 2 of gaster witha dense felt-like ring of pubescence; antenna of $ filiform, segment 3 of cf antenna in typespecies equal to 4+5 (see note) (in African species 3 is shorter than 4+5; radial cell offorewing open or closed) GLA URASPIDIA Thomson - Without the above combination of characters ; mesopleural suture generally present ; pronotum and propodeum without dense felt-like tufts; tergite 2 of gaster with hairy ring variable indensity of pubescence; $ antenna usually clavate, cf antenna filiform, 3rd or 4th segmentsometimes modified 18 18 Scutellar cup narrow, elliptical, disc reticulate-striate, rounded at apex; radial cell of forewing open on front margin (Ri projecting slightly on the margin but not joining Rs 2 to form aclosed radial cell) (Fig. 38); tergite 2 of gaster with a ring of pubescence, generally completeon dorsal surface. $ antenna clavate, cf antenna with segment 3 longer than 4, sharply curved on outermargin (Fig. 39), swollen distally; pronotal plate with lateral pits or fovea open (Fig. 40),with tufts of pubescence on either side (in some species mesopleural suture aberrant orabsent) HEXACOLA Foerster(p. 272) - Scutellar cup variable in shape, sculpture of scutellar disc variable, sometimes smooth, never striated as in Kleidotoma; radial cell of forewing open or closed on wing margin; tergite 2 ofgaster with ring of pubescence at base dense or weak , sometimes complete on dorsal surface. 19 19 Tergite 2 of gaster with ring of hairs at base present but rather thin or sparse, not usually complete on dorsal surface, not obscuring tergite 1 which is crenulate to reticulate-crenulate(Fig. 41), abruptly widened posteriorly (anteroventral cavity of mesopleura present). Cf antenna filiform, segments 3 and 4 subequal in length, 4th slightly flattened on outermargins (Fig. 361); $ antenna long, slender, with club segments sometimes only indicated bypresence of rhinaria (Fig. 362) LEPTOPILINA Foerster - Tergite 2 of gaster with a dense ring of pubescence at base, obscuring tergite 1 completely, complete on dorsal surface, not woolly in appearance. Scutellar cup elevated, variable in size and shape from oval to almost circular, scutellardisc sculpture variable, from reticulate to reticulate-punctate, lateral bars of scutellumusually striate, especially on lateral margins; pronotal plate variable in size and shape, lateralfovea between anterior and posterior parts open 20 20 Scutellar cup almost flat, mirror-like, elliptical, if raised above level of disc then only very slightly (Fig. 44), scutellar disc closely punctate-reticulate, apical margin weakly conical(Fig. 44); lateral bars of scutellum weakly striated on lateral margins, polished dorsally;pronotal plate with lateral fovea between anterior and posterior parts small, not sharplydelineated; apex of wing rather blunt, radial cell of forewing usually closed; $ antennaclavate or filiform, cf antenna filiform, 4th segment modified GANASPIS Foerster - Scutellar cup almost circular, well elevated above scutellar disc, rugose to rugose-fovelate, apical margin rounded (Fig. 359), lateral bars of scutellum usually striate on dorsal andlateral surfaces; pronotal plate large, lateral fovea open, clearly delineated (Fig. 12); apex offorewing rounded, radial cell of forewing open or closed; $ antenna usually clavate, cfantenna filiform , segment 4 usually swollen (Fig . 46) TR YBLIOGRAPHA Foerster AFRODONTASPIS Weld Afrodontaspis Weld, 1961: 280. Type-species: Coneucoela striatissima Benoit, by original designation. DIAGNOSIS. $ antenna 13-segmented, subclavate, cf antenna 15-segmented. Pronotum, mesoscutum,mesopleura and lateral bars striate (Figs 10, 49, 50, 51). Scutellum with a long narrow cup extending almostto apex of scutellar disc, the usual large basal fovea on cup declined (not visible in dorsal view), scutellardisc falling sharply away from cup, with crenulate sculpture (Fig. 10); either side of pronotal plate with atuft of dense yellow-white pubescence; base of tergite 2 of gaster with dense hairy ring obscuring tergite 1.Wings clear, not ciliate, dotted with hair bases, lower margins with a weak fringe of hairs. DISTRIBUTION. Zaire; Uganda. 254 J. QUINLAN Key to the species of Afrodontaspis Weld 1 Head behind eye (vertex) strigose, viewed frontally, strongly coriaceous, face coriaceous- reticulate. Pronotal plate (Fig. 47); mesoscutum with fine longitudinal striations (Fig. 50);apical segment of antenna 2x as long as wide; antenna yellowish, thorax and gaster reddish brown ; segments 2-5 of gaster visible in lateral view (Fig. 48) striatissima (Benoit)(p. 254) - Head behind eye (vertex) strongly aciculate, viewed frontally, face with weak radiating striae.Pronotal plate (Fig. 49); mesoscutum with broad longitudinal striations (Fig. 50); apicalsegment of antenna 2 5 x as long as wide ; antenna brown , thorax black , gaster reddish brown ;segments 2-3 of gaster visible in lateral view lanatus sp. n. (p. 254) Afrodontaspis lanatus sp. n. (Figs 10, 49, 51, 52, 53, 54) DESCRIPTION. $ . Antenna 13-segmented, clavate, 7-13 with rhinaria forming a weak club, 3-6 subequal inlength and breadth, 7-13 each weakly swollen, 7-10 each same length as segments 3-6, 11-13 each longerthan 7 (Fig. 52). Head, viewed frontally, longer than wide, vertex coriaceous, eyes further apart measuredmedially than height of an eye, face with closely spaced radiating striate sculpture converging towardsfrontal line area which is coriaceous (Fig. 53), malar groove not conspicuous in the striate sculpture,clypeus coriaceous, mandibles tri-dentate, occiput and vertex strigose-coriaceous. Pronotal plate pro-duced forward, medial bridge between anterior and posterior parts narrow, fovea or lateral cavities openlaterally (Fig. 49), these cavities filled with dense pubescence extending out from and on sides of fovea.Pronotum, viewed dorsally, striated; mesoscutum longitudinally striated, weakly so medially (Fig. 51);lateral bars of scutellum striate, scutellar fovea large, deep, separated by a long thin ridge with scutellar cupdeclined posteriorly, scutellar disc coriaceous in dorsal view, tongue-shaped apically, lateral margins ofapex of disc crenulate (Fig. 10); carinae of propodeum bowed medially, pubescent on outer margins.Mesopleura completely transversely striate. Segment 1 of gaster completely obscured by dense ring ofpubescence at base of tergite 2, woolly in appearance, tergites 2-3 of gaster visible in lateral view, segment 2 the largest, occupying most of visible area, ventral spine and hypoygium barely visible; coxae and femorafinely coriaceous, tibiae and tarsi finely and densely strigose, pubescent, tibiae with two apical setae. Wingsnot ciliate, without apical hair fringe, surface of wings dotted with hair bases, radial cell of forewing openon wing margin (vein RI not reaching margin of wing, but indicated by pigmentation), Rs+M and M veryweakly indicated by a trace of pigmentation (Fig. 54). Colour: antenna yellowish brown, head and thoraxblackish, gaster chestnut-red. Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kaziba, affl.g. Senze s. affi. dr. Lufira, 1140 m, 15-27.ii.1948 (G. F. de Witte)(MRAC). Paratypes. Zaire: 4 $ (MRAC). Kenya: 1 $ (BMNH). REMARKS. This species is closely related to striatissima but separated by the different sculpturing on thehead and mesoscutum. A larger and darker species. Size 3 mm. Afrodontaspis striatissima (Benoit)(Figs 10, 50, 56) Coneucoela striatissima Benoit, 1956: 533. Holotype <j>, ZAIRE (MRAC) [examined].Afrodontaspis striatissima (Benoit) Weld, 1961: 280. DESCRIPTION. $. Antenna 13-segmented with 3-segmented club, club segments with rhinaria, remainderwith granulate sculpture, segments 3-10 subequal in length, each wider apically than at base. Head, viewedfrontally, strongly coriaceous, frontal area raised with weak impressed sculpture extending to base ofclypeus, mandibles tridentate, head, viewed dorsally, finely coriaceous, back of head strigose. Pronotalplate large; pronotum longitudinally striate (with a dense tuft of pubescence on medial margins) (Fig. 50).Mesopleura strongly and broadly striated on lateral margins, weakly strigose to smooth medially. Lateralbars of scutellum striate. Scutellar disc ending in a blunt spine, scutellar fovea large, half as long as entirescutellum, scutellar cup very narrow, extending to base of scutellar spine, the usual large basal fovea on cupdeclined, dorsal surface of scutellar disc coriaceous, lateral margins sharply declined with large adpressedcrenulations (cf. Fig. 10); carinae of propodeum bowed. Base of tergite 2 of gaster with dense ring ofpubescence obscuring tergite 1 from view, tergites 2-5 visible in lateral view, the apical margins finely AFROTROPICAL EUCOILIDAE 255 punctate, tergite 2 the largest (Fig. 48). Femur and tibia with reticulate-coriaceous sculpture, tibia withstriated sculpture, tarsus 5-segmented. Wings with hair fringe on hind margin only, surface dotted with hairbases, not ciliate, radial cell of forewing open basally (vein RI not reaching margin of wing), Rs+M and Mnot indicated (Fig. 56). Colour: antenna orange-yellow, head blackish red, thorax and gaster chestnut-red,legs orange-yellow. Cf . Antenna 15-segmented, segment 3 3-5 x length of 4, 4-15 subequal in length, narrower at apex;sculpture of head and thorax similar to $ . MATERIAL EXAMINED Zaire: 1 $ (holotype), Rutshuro, xii.1937 (/. Ghesquiere) (MRAC).Uganda: 1 $, 20 cf (BMNH); Zaire: 44 $ (MRAC). REMARKS. This species is separated from lanatus by the sculpture of the head and thorax. A much smallerspecies. Size 2 mm. COTHONASPIS Hartig Cothonaspis Hartig, 1840: 201. Type-species: Cothonaspis pentatoma Hartig, by original designation. P-silosema Keiffer, 1901: 160. Type-species: Psilosema giraudi Dalla Torre & Keiffer, by subsequentdesignation, Rowher & Pagan, 1917: 374. [Synonomy by Weld, 1952: 243.] DIAGNOSIS. $ antenna 13-segmented with apical segments forming a club (Fig. 57), cf antenna 15-segmented, the fourth segment swollen, longer than third or fifth. Head, viewed frontally, spherical, aswide almost as thorax, genae broad, eyes round, further apart measured medially than height of an eye.Pronotal plate projected forward, anterior and posterior parts not fused laterally, i.e. fovea or cavitiesopen (Fig. 34). Mesoscutum smooth, polished, notaulices absent, in some species long scattered hairspresent; scutellar cup elongate, narrow, broader basally, scutellar disc with radiating striae, lateral barssmooth, polished, scutellar foveae ovoid, smooth. Propodeal carinae parallel, propodeum produced toform a neck-shaped nucha, hairless laterally. Segment 1 of gaster short, broad, in form of a crenulate ring,base of tergite 1 without a ring of pubescence, at most a few sparse hairs present, gaster compressedlaterally. Wings narrow, rounded apically, surface normally pubescent, with a longish hair fringe on apicalmargins, radial cell of forewing closed on distal margin though in a few species it appears open due toabsence of a complete vein, i.e. pigmentation only present. DISTRIBUTION. Europe; South Africa; North and South America. DISCUSSION. Cothonaspis is separated from the closely related Leptopilina by the form of the thorax andthe lateral margins of the propodeum. In Cothonaspis the thorax is elongate and the lateral margins of thepropodeum are not noticeably hairy. Two Afrotropical species have at one time or another been assignedto the supposed subgenera of Cothonaspis. Erisphagia mahensis Kieffer was placed in Cothonaspis(Erisphagia) by Weld (1952: 244), and Nordlander (1980) transferred it to Leptopilina Foerster. Psilosemafenerivae Kieffer, a species based on a male from Madagascar but not located, is not accounted for in thispaper. Two further species are described and compared with Cothonaspis pentatoma, the type-species ofthe genus. BIOLOGY. The European species of Cothonaspis are associated with Sepsidae in cow dung (Nordlander,1976; Quinlan, 1978). Key to the species of Cothonaspis Hartig Females 1 Antennal club 5-segmented, apical segments not contrasted to basal or medial segments; scutellar cup not extending to apex of disc, apex of disc rounded (Fig. 57) 2 - Antennal club 7-segmented , apical 3-4 segments orange-yellow in contrast to the brownish basaland medial flagellar segments (Fig. 59); scutellar cup elongate, extending almost to apex ofthe almost truncate scutellar disc (Fig . 71 ) ; apex of gaster finely punctate dulcis sp . n . (p . 256) 2 Tergites 1-2 of gaster visible in lateral view , gaster without a dense hair patch , scutellar disc with radiating striate sculpture; mesopleura strigose above mesopleural suture; lateral bars of scutellum striate on dorsal surface ; hind coxa clearly strigose pentatoma Hartig (p . 256) Tergites 1-4 of gaster visible in lateral view, base of tergite 2 with a patch of short pubescence onlower lateral margins; scutellar disc with broken areolate sculpture; mesopleura smooth andpolished above mesopleural suture; lateral bars of scutellum polished on dorsal surface; hindcoxa smooth, polished ealis sp. n. (p. 256) 256 J. QUINLAN Cothonaspis dulcis sp. n. (Figs 59, 60, 71) DESCRIPTION. $. Antenna 13-segmented, clavate, segments 3 and 4 subequal, 5 shorter than 4, 6 shorterthan 5, 7-13 forming a distinct club, all club segments with rhinaria (Fig. 59). Head, viewed frontally, witheyes further apart than height of an eye measured medially, cheeks converging towards clypeal area, malargroove distinct. Pronotal plate projected forward, anterior and posterior parts bridged medially, lateralfovea open on margins (Fig. 60). Mesoscutum smooth, polished, scutellar fovea smooth, shallow, lateralbars polished, scutellar disc almost truncate viewed dorsally, surface reticulate-rugose, scutellar cup long,narrow, narrower apically than medially, extending almost to apex of disc (Fig. 71). Propodeal carinaeparallel, weakly pubescent on lateral margins, mesopleurae smooth, polished, mesopleural suturecomplete, a few rugae present below suture adjacent to lateral margins of pronotal plate, metapleurapolished. Segment 1 of gaster short, broad, crenulate, in form of a ring, tergite 2 the largest, tergites 3 and 4visible in lateral view, apex of tergite 2 and whole of visible parts of 3 and 4 finely punctate, tergite 2 with asmall tuft of hairs on lateral margins not prominent or obscuring tergite 1, hypopygium not pronounced,ventral spine not visible. Legs long, slender. Wings long, narrow, rounded apically, surface pubescent withapical hair fringe, radial cell of forewing closed on wing margin. Colour: antenna dark yellow basally,brownish medially, apical 3-4 segments light yellow, head reddish black, thorax and gaster chestnut-brown, legs orange-brown.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kibati, 1000 m, 10-12.U934 (G. F. de Witte) (MRAC).Paratypes. Zaire: 21 $ (MRAC). Madagascar: 1 $ (MNHN). Cothonaspis ealis sp. n. (Figs 61, 62) DESCRIPTION. $ . Antenna 13-segmented, with 5-segmented club, segment 3 longer than 4, 4 longer than 5,6-8 subequal, each slightly longer than 5 (Fig. 61). Head, viewed frontally, rounded, eyes oval, facesmooth, shining, with sparse scattered hairs, malar groove percurrent, vertex smooth. Pronotal plateproduced forward, anterior and posterior parts connected medially, lateral margins, i.e. fovea, openlaterally (Fig. 62); pronotum weakly striated on dorsal surface adjacent to mesoscutum; mesoscutumsmooth, polished, with a few scattered hairs; lateral bars of scutellum smooth, polished, scutellar foveashallow, almost as broad as long, scutellar disc polished, with trace of broken striae, apex rounded, cupsmall, long, narrow. Propodeal carinae parallel, without pubescence on lateral margins, propodeumelongate, carinate, hairless; mesopleural suture complete, mesopleurae smooth, polished; metapleuraridged, an tero ventral cavity distinct, with a few setae. Segment 1 of gaster short, broad, in the form of acrenulate ring, tergite 2 the largest, smooth, polished, without a ring of pubescence at base, impuncate,tergites 3 and 4 partially visible in lateral view, hypopygium not pronounced, ventral spine not visible. Legsshort, robust, tarsal segments longer than tibia. Wings pubescent, margins ciliate, radial cell of forewingclosed on wing margin. Colour: antenna light brown, head, thorax and gaster brown-black, legs yellowish.Cf unknown. MATERIAL EXAMINEDHolotype $, South Africa: Cape Town, Milnerton, i.1926 (R. E. Turner) (BMNH). Cothonaspis fenerivae (Kieffer) Psilosema fenerivae Kieffer, 1910a: 534; Weld, 1952: 243. Holotype cf , MADAGASCAR: Fenerive, vii.(Voeltzkow) (depository unknown). Kieffer 's description is based on a male and could fit a number of species. Cothonaspis pentatoma Hartig(Figs 57, 58, 63) Cothonaspis pentatoma Hartig, 1840: 201. Holotype $, GERMANY (MNHN) [examined].Eucoila (Pentamerocera) pentatoma (Hartig) Kieffer, 1901: 175.Cothonaspis (Cothonaspis) pentatoma Hartig Weld, 1952: 242. AFROTROPICAL EUCOILIDAE 257 DESCRIPTION. $. Antenna 13-segmented with a 5-segmented club, segment 3 subequal to 4+5, 4 longerthan 5 (8th segment weakly swollen but not regarded as a club segment, rhinaria are not present as in thefive apical segments) (Fig. 57). Head, viewed frontally, rounded, eyes normally rounded, smallish, malarspace area striated, face with sparse hairs, maxillary palp 4-segmented, labial palp 2-segmented, vertexstrigose. Pronotal plate projected forward, anterior and posterior parts connected medially, lateralmargins open, i.e. fovea open laterally (Fig. 58); mesoscutum smooth, polished, with sparse hairs,notaulices absent. Lateral bars of scutellum striate dorsally, polished, scutellar fovea smooth, shallow, asbroad as long, scutellar disc with longitudinal or radiating ridges, cup small, slightly longer than broad.Propodeal carinae pronounced, parallel, propodeum elongated in lateral view, a few sparse hairs present(Fig. 63). Mesoscutum, in lateral view, long, slender. Mesopleural suture complete, strigose above suture,metapleura ridged, antero ventral cavity indicated. Segment 1 of gaster broad, short, canaliculate, tergite 2the largest in lateral view, with a few long hairs at its base. Legs long, slender, fore and mid-legs of similarsize and shape, hind-leg larger, tarsi short, hind coxa strigose. Wings narrow, rounded apically, surfacepubescent with hair fringe on apical margin, radial cell of forewing closed (sometimes indistinctly so, due topresence only of pigmentation, the vein becoming vestigial). Colour: antenna brownish, head and thoraxblack, gaster dark brownish black, chestnut-red basally. Legs brownish basally, yellowish apically. REMARKS. Although pentatoma appears to be known only from Europe it is included in the key to species tofacilitate identification and comparison with the two Afrotropical species. EALATA gen. n. (Figs 19, 20, 69)Type-species: Ealata clava sp. n. DIAGNOSIS. 9 antenna 13-segmented, clavate, c? antenna 15-segmented, filiform, segment 3 curved,subequal in length to segment 4. Pronotal plate not projected forward, anterior part wider than posteriorpart, fovea large, not enclosed on lateral margin (Fig. 20). Mesoscutum smooth, polished, notaulicesweakly indicated basally, foveae broader than long, kidney-shaped; scutellar disc reticulate-rugose,weakly rounded apically, scutellar cup oval, slightly longer than broad, raised, with a weak depressionapically, with a pale-edged rim (Fig. 69). Propodeum with bowed carinae, pubescent on either side.Mesopleura smooth, polished, suture complete. Segment 1 of gaster partially visible, sculptured, in form ofa crescent, tergite 2 the largest in lateral view, without a ring of pubescence at base, at most with a fewscattered hairs on lateral margins. Wings pubescent, apical margins with a hair fringe, radial cell offorewing closed on wing margin (Fig. 19). Legs of normal size and shape, coxae with a fringe of hairs onmargins viewed laterally, dark brown, femora, tibiae and tarsi orange-yellow with usual pubescence. DISTRIBUTION. Afrotropical Region. DISCUSSION. This genus superficially resembles Leptopilina. However, the pronotal plate is not projectedforward, and the notaulices, though aberrant, are indicated at least on the anterior of the mesoscutum.Tergite 2 of the gaster has a few hairs present laterally, never with a ring of pubescence, however thin.Ealata is closely related to Nordlanderia, Disorygma and Diglyphosema. It is separated from Disorygma bythe mesoscutum and the form of the scutellar cup. Key to the species of Ealata gen. n. 1 Antenna of $ with 6-segmented club ; hypopygium not protruding (Fig. 67) ........................... 2 - Antenna of $ with 7-segmented club; hypopygium protruding (exceptionally long) (Fig. 64) marica sp. n. (p. 258) 2 Antennal segments 8-13 forming a distinct club, rhinaria very distinct on apical three segments, all club segments wider apically than basally in contrast to basal segments (Fig. 65) saba sp. n. (p. 258) - Antennal segments 8-13 forming a very weak club, rhinaria present, all segments at least 2 x as long as wide, not wider apically than basally (Fig. 66) ................................. clava sp. n. (p. 257) Ealata clava sp. n. (Figs 20, 66, 67, 69, 72) DESCRIPTION. $. Antenna 13-segmented, clavate, segment 3 1-5 x length of 4, 4, 5 and 6 subequal inlength, 7 almost twice length of 6, 8-13 forming a club though segment 7 could be mistaken for a club 258 J. QUINLAN segment (see note), apical segment shorter than 11 + 12 (Fig. 66). cf antenna 15-segmented, filiform, 3 and4 subequal in length, 3, viewed dorsally, weakly curved. Face, viewed frontally, smooth, polished, withscattered hairs, malar suture distinct, eyes weakly converging. Pronotum pubescent either side of pronotalplate, plate not projected forward, anterior part much wider than posterior part, lateral foveae large, notenclosed (Fig. 20). Mesoscutum on lateral margin of pronotum from tegulae to pronotal plate channelled,notaulices very weakly indicated anteriorly (best seen viewed laterally), lateral bars of scutellum polished,scutellar foveae shallow, polished, scutellar cup small, convex, polished, with apical depression, discreticulate-rugose (Fig. 69), rounded apically. Propodeal carinae bowed, densely pubescent on either side.Mesopleural suture complete. Segment 1 of gaster partially visible, in form of crenulate ring, obscured inpart by tergite 2, lateral margins of tergite 2 with a few sparse hairs at base; tergite 2 the largest in lateralview, remaining segments not always visible, surface smooth, shining, hypopygium pronounced (Fig. 67).Wing surface pubescent, apical margins with hair fringe, radial cell of forewing closed on the margin,cubitus (M) not indicated (Fig. 72). Legs long, slender. Colour: femora brownish yellow, head, thorax andgaster blackish, base of gaster orange, legs orange-yellow. Cf . Antenna 15-segmented, segment 3 weakly curved, same length as 4. MATERIAL EXAMINED Holotype $, Zaire: Eala, ii.1935 (/. Ghesquiere) (MRAC). Paratypes. Cameroun: 2 $ (BMNH). Kenya: 1 $ (BMNH). Mauritius: 1 cf (BMNH). South Africa: 2 $(BMNH). Uganda: 2 $ (BMNH). Principe Is.: 1 $ (BMNH). Zaire: 34 $, 19 cf (MRAC). DISCUSSION. This species is separated from others in the genus by the club segments; these can be easilycounted and comprise seven segments, but rhinaria are not present on the seventh. The sixth segment,although very similar in shape to the other club segments, does not have rhinaria and is therefore notregarded as a true club segment. Ealata marica sp. n. (Figs 19, 20, 64, 69) DESCRIPTION. $. Antenna 13-segmented, subclavate, segment 3 1-3 x length of 4, 4, 5 and 6 subequal inlength, 7-13 forming a club, each club segment with rhinaria, weakly swollen medially, apical segmentshorter than 11 + 12 (Fig. 66). Face, viewed frontally, smooth, polished, with scattered hairs, malar suturedistinct, eyes as far apart measured medially as height of an eye. Pronotal plate not projected forward,anterior part wider than posterior part, the lateral foveae large, not enclosed (cf. Fig. 20). Mesoscutumchannelled from lateral margins to pronotal plate, notaulices not indicated, sparse hairs present in theirplace, lateral bars of scutellum polished, scutellar foveae shallow, wider than long, scutellar cup small,slightly longer than wide, not extending to apex of disc, scutellar disc finely reticulate-rugose, apexrounded (cf. Fig. 69); propodeum obscured by pubescence, mesopleural suture complete, metapleuradensely pubescent, propodeum extended. Segment 1 of gaster in form of a crenulate ring, obscured by baseof tergite 2, tergite 2 the largest, with two or three hairs basally on lateral margins, remaining segments notvisible, hypopygium long, narrow (Fig. 64). Wing surface densely pubescent, margins with apical hairfringe, radial cell of forewing closed on margin, long, narrow (Fig. 19). Legs short, stout, hind coxa swollenmedially, tibia as long as tarsus. Colour: antenna orange-yellow, head, thorax and gaster dark brownishblack, legs, coxa brownish black, femora, tibiae and tarsi orange-yellow.Cf unknown. MATERIAL EXAMINEDHolotype $, Zaire: Kivu, Rutshuru, 1285 m, ll.vii.1935 (G. F. de Witte) (MRAC). REMARKS. This species is easily distinguished by the antenna and extremely long, narrow hypopygium. Ealata saba sp. n. (Figs 19, 20, 65) DESCRIPTION. $. Antenna 13-segmented, clavate, with distinct 6-segmented club, club segments exceptapical slightly longer than broad, clearly wider apically than basally, apical segment as long as 11 + 12 (Fig.65), segment 3 longer than 4, 4-7 subequal in length, 7 shorter than 8. Face, viewed frontally, smooth,polished, with scattered hairs, malar groove distinct, eyes weakly converging, mandibles tridentate.Pronotum sparsely pubescent either side of pronotal plate , plate not pro j ected forward , anterior part muchwider than posterior part, lateral foveae large, not enclosed (cf. Fig. 20). Mesoscutum smooth, polished,notaulices absent, a row of hairs in their place, lateral bars of scutellum polished, scutellar foveae wider AFROTROPICAL EUCOILIDAE 259 than long, smooth, shallow, scutellar cup small, almost oval, scutellar disc reticulate-rugose with scatteredpubescence, apex weakly conical viewed dorsally, propodeum elongate in lateral view, propodeal carinaebowed basally, inner surface sculptured. Mesopleural suture complete, metapleura ridged on upper half,anteroventral cavity with tuft of hairs. Segment 1 of gaster partially visible, in form of a crenulate ring,tergite 2 the largest, occupying whole of visible area in lateral view, obscuring tergite 1, base of tergite 2without hairs, hypopygium not pronounced, with a few hairs on ventral surface. Wing surface pubescentwith apical fringe of hairs, radial cell of forewing closed on wing margin, elongate (cf. Fig. 19); legs shortand stout, hind coxa swollen, elongate, mid- and fore coxae smaller, femora short, swollen medially, tibiaeand tarsi slender, fore and mid-tibiae shorter than tarsus, hind tibia and tarsus subequal in length. Colour:antenna brownish yellow, head and thorax brownish, gaster chestnut-red; legs: coxae orange, femora,tibiae and tarsi yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Nigeria: Ibadan (B. R. Critchley) (BMNH). Paratypes. Nigeria: 4 $ (BMNH). Rhodesia: 1 $ (BMNH). South Africa: 2 $ (BMNH). Uganda: 2 $(BMNH). Zaire: 6 $ (MRAC). REMARKS. This species is separated from clava by the distinctive antennal club (Fig. 65). EUCOILIDEA Ashmead(Figs 84, 86, 93, 101) Eucoilidea Ashmead, 1887: 154. Type-species: Eucoilidea canadensis Ashmead, by subsequent designa-tion, Ashmead, 1903: 60.Afrostilba Benoit, 1956: 544. Type-species: Afrostilba nitida Benoit, by monotypy. Syn. n. DIAGNOSIS. $ antenna usually 13-segmented, clavate to almost filiform, club segments distinguished bypresence of rhinaria, cf antenna 15-segmented. Face smooth, shining, malar space with a distinctepistomal suture (malar ridge), mandibles tridentate. Pronotal plate weakly protruding viewed dorsally,cavities either side of median bridge open laterally (Fig. 101). Pronotum either side of pronotal platepolished to strongly aciculate. Mesoscutum smooth, polished, notaulices complete, distinct, converging onapproach to scutellum where medial distance between notaulices can vary (Figs 86, 93); lateral bars ofscutellum smooth, scutellar fovea deep, scutellar cup variable, from oval to round to elongate, scutellardisc usually visible laterally and apically though this is sometimes overlapped apically by scutellar cup, apexof disc rounded (Figs 86, 93, 97). Segment 1 of gaster sulcate, short, as broad as long, segment 2 occupyingalmost whole of visible part of gaster in lateral view (Fig. 14), surface without punctures at x45magnification. Wings with apical fringe of hairs, surface pubescent, radial cell of forewing closed on wingmargin (Fig. 84). DISCUSSION. The holotype of Eucoilidea canadensis Ashmead is a $ and has its antenna missing and isbadly obscured by glue. Nineteen species, including seventeen newly described, are recognized from theAfrotropical region. Weld (1952) states that the antenna in the $ of canadensis has the third segmentslightly longer than the fourth, without a distinct club, cf with segment three twice the length of four andstrongly bent. In a $ from the West Indies the third segment is slightly shorter than four. The holotypewould run in the present key to dubia but differs in the ratios of the antennal segments, the shape of the cupand the acuteness of the notaulices on approaching the scutellum. Afrostilba is here synonymised withEucoilidea after comparison with related genera. First examination of Afrostilba, using Weld's key (1952)based on scutellar characters, distinguishes it from Eucoilidea by the absence of striations in the malar areaof the face together with scutellar differences which also separate it from Microstilba. However, theemphasis given to the scutellar characters after comparison with the holotype of Eucoilidea canadensis andthe species described here do not merit a separate genus. Eucoilidea has been recorded by Weld (1952)from the U.S.A., Mexico, Hawaii and the Philippines. The type-species is from Canada. Undescribedspecies of Eucoilidea from India and Singapore are in the BMNH. Afrostilba nitida Benoit is transferred toEucoilidea. DISTRIBUTION. North and South America, Hawaii, Philippines, South Africa. Key to the Afrotropical species of Eucoilidea Ashmead Females 1 Antenna 14-segmented; notaulices converging sharply towards scutellum (cf. Fig. 74); scutel- 260 J. QUINLAN lar disc reticulate-rugose, scutellar cup large, extending to apex of disc, obscuring apical margins; tergites 2-4 of gaster visible in lateral view, finely punctate extraria sp. n. (p. 265) Antenna 13-segmented; notaulices either sharply converging or gently converging towardsscutellum (Figs 85, 97); scutellar disc visible laterally and sometimes apically; tergite 4 ofgaster sometimes visible in lateral view, gaster either impunctate or punctate on apicalmargins (Figs 94, 100) 2 2 Apical margins of either tergite 2 of gaster , or 3 and 4 when visible , with dense fine punctures on apical margins , or apical tergites not strongly compressed (Fig .77) 3 Apical margins of gastral tergites without dense punctation or apical tergites of gaster stronglycompressed laterally, with very sparse punctation on the apical margins (Figs 94, 116, 119). ... 11 3 Antennal segments filiform or subclavate, segments 3 and 4 subequal in length (Fig. 81), or segment 4 longer than 3 ; Rs and M of forewing present or absent 6 Antenna subclavate or filiform, segment 3 at least 1-2 x length of 4; segments 1-4 of gasternormally visible in lateral view 4 4 Antenna filiform, segments 4-13 with rhinaria (Fig. 129); veins Rs+M and M of forewing indicated (Fig. 130) 5 - Antenna clavate, club 8-segmented, segment 3 weakly curved (Figs 79, 144); veins Rs+M and M of forewing not indicated, radial cell of forewing 2 x as long as broad (Fig. 147); gasterfinely punctate, tergites 1-4 normally visible in lateral view, hypopygium prominent (Fig.145) trullasp. n.(p. 271) 5 Tergites 1-2 of gaster visible in lateral view, tergite 2 densely, coarsely punctate on apical half; radial cell of forewing 2-6 x as long as wide (Fig. 78); scutellar cup extending to andobscuring apex of scutellar disc (cf. Fig. 132); legs yellowish marcellus sp. n. (p. 268) - Tergites 1-3 of gaster visible in lateral view, tergites 2 and 3 finely punctate; radial cell of forewing 3-6 x as long as wide (Fig. 80); scutellar cup elongate, scutellar disc visible laterally and apically, legs bright orange-yellow tyrus sp. n. (p. 271) 6 Segment 4 of antenna clearly longer than 3, at least 1-2 x as long as wide (Fig. 81); notaulices converging sharply towards scutellum ; pronotum strongly aciculate (Fig .83) 7 - Segment 3 longer than or subequal in length to 4 (Fig. 127); notaulices converging sharply or gently towards scutellum 8 7 Antenna subclavate, segments 5-7 equal in length, 8-12 each progressively shorter than preceding segment; pronotum with long scattered pubescence; radial cell 2 x as long as wide (Fig. 82) ; segments 2-4 of gaster visible in lateral view lacerta sp. n. (p. 266) - Antenna filiform, blackish; scutellar disc punctate-reticulate; radial cell 3 x as long as wide (Fig. 84) /anasp. n. (p. 267) 8 Notaulices not converging sharply towards juncture with scutellum (Fig. 86) 9 - Notaulices converging sharply towards juncture with scutellum (Fig. 85) 10 9 Antenna clavate; scutellar cup not extending to apex of scutellar disc (Fig. 85) nitida Benoit (p. 269)Antenna filiform; scutellar cup extending to apex of scutellar disc (Fig. 86) leptis sp. n. (p. 267) 10 Antenna filiform, clearly longer than head-apex of gaster, segments 5-13 each progressively shorter than the preceding one, of equal width, two apical segments of equal length (Fig. 87),all much longer than broad; radial cell of forewing elongate, Rs+M and M of forewingindicated, not pigmented; scutellar disc punctate-reticulate-rugose, visible viewed dorsallyon lateral and apical margins (cf. Fig. 114); side margins of pronotum weakly sculptured withlong scattered hairs, distance between notaulices at juncture with scutellum as wide as a notaulix (cf. Fig. 114) ; segment 3 of gaster not visible in lateral view con versa sp. n. (p. 264) Antenna subclavate, at most slightly longer than head-apex of gaster; segments 6-12 eachshorter than its preceding segment but each broader, apical segments weakly expanded togive a weak clavate appearance (Fig. 88); radial cell of forewing (Fig. 140), Rs+M and M notindicated; scutellar disc punctate-reticulate, visible on lateral and apical margins (Fig. 139);pronotum either side of pronotal plate aciculate (cf. Fig. 83); segment 3 of gaster visible inlateral view (cf. Fig. 77) parma sp. n.(p. 270) 11 Antennal segments 3 and 4 subequal in length, never with segment 4 longer than 3 (Fig. 89); scutellar cup elongate or oval 12 Antennal segments 3 and 4 not subequal, either clearly longer or shorter than each other (Figs105 , 120) ; scutellar cup elongate 18 AFROTROPICAL EUCOILIDAE 261 12 Antennal segment 4 at most 1-2 x length of 5, antenna subclavate (Figs 89, 90); radial cell of forewing elongate, at least 2 x as long as wide (Fig. 112), Rs+M and M indicated, base ofwing fumate or yellowish; scutellar cup elongate; gaster not conspicuously compressed laterally 13 Antennal segments 4 and 5 subequal in length (Fig. 142), antenna filiform or subclavate; radialcell of forewing variable in length to width ratio, Rs+M and M present or absent, base offorewing rarely fumate; scutellar disc sculpture variable, from rugose-punctate to punctate-reticulate, scutellar cup elongate or oval; notaulices converging towards scutellum, some-times sharply; gaster sometimes strongly laterally compressed 14 13 Notaulices converging gently towards scutellum (Fig. 92), scutellar disc punctate-reticulate, visible laterally and apically, scutellar cup narrow (Fig. 92); antenna yellowish basally,brownish apically, subclavate (Fig. 89) fetura sp. n. (p. 265) - Notaulices sharply converging towards scutellum (Fig. 93) ; scutellar disc strigose, scutellar cup obscuring apical surface of disc (Figs 93, 141); antenna bright yellow. Radial cell of forewing appearing open on anterior margin due to very weakpigmentation pattida sp. n. (p. 269) 14 Gaster very strongly compressed laterally, tergites 3 and 4 visible in lateral view (dorsally they appear knife-edged and acutely angled to tergite 2) (Fig. 94).Scutellar cup large , oval (Fig. 102) 15 - Gaster not conspicuously compressed laterally, swollen medially, tergites 3 and 4 not clearly visible in lateral view (Fig. 100) 16 15 Gastral segments, viewed laterally, impunctate (Fig. 94); antennal segments 3-6 subequal in length, apical antennal segments weakly swollen medially (Fig. 91), rhinaria present onsegments 7-13 compressa sp. n.(p. 263) - Gastral segment 3 finely but densely punctate (Fig. 100); antenna subclavate, segment 5 very slightly shorter than 4, 4-13 each progressively broader than preceding segment to form aclavate antenna (Fig. 98) , rhinaria present on segments 5-13 urundiensis Benoit (p. 272) 16 Radial cell of forewing long, narrow (Fig. 95), 3-6 x as long as wide; antennal segments 3-8 subequal in length; notaulices converging gently towards scutellum; scutellar cup oval, disccoriaceous, visible laterally.Tergite 2 of gaster only visible in lateral view, bright orange-yellow . . perangusta sp. n. (p. 270) - Radial cell of forewing at most 3 x as long as wide; antennal segments 3-6 subequal in length; notaulices converging sharply or gently towards scutellum , scutellar cup variable in shape .... 17 17 Notaulices converging sharply towards scutellum; scutellar cup oval, scutellar disc visible on lateral margins only, viewed dorsally (Fig. 97); antenna subclavate, as long as head-apex ofgaster (Fig. 103); tergites 2-4 of gaster visible in lateral view, hypopygium pronounced (cfFig. 77) bucca sp. n. (p. 263) - Notaulices converging gently towards scutellum; scutellar cup elongate, scutellar disc visible laterally and apically in dorsal view (Fig. 104); antenna filiform, longer than head-apex ofgaster (Fig. 113); tergite 4 of gaster not visible, ovipositor extended (cf. Fig. 94) advena sp. n. (p. 262) 18 Antenna clavate, segment 3 clearly longer than 4 (Fig. 120) 19 - Antenna filiform, segment 3 clearly shorter than 4 (Fig. 105). Scutellar disc punctate, visible laterally and apically, scutellar cup elongate (cf. Fig. 132);notaulices converging gradually towards scutellum; radial cell of forewing short, broad (cf.Fig. 80) duWasp. n. (p. 264) 19 Notaulices converging sharply towards scutellum (cf. Fig. 1 14) ; scutellar cup elongate , narrow , scutellar disc visible laterally and apically ; radial cell of forewing lightly pigmented (Fig . 1 1 1 ) furcula sp. n. (p. 266) Notaulices widely separated at juncture with scutellum (Fig. 132); scutellar cup large, oval,scutellar disc only visible viewed dorsally on lateral margins, radial cell of forewing heavilypigmented (cf. Fig. 130) maurisp. n.(p. 268) Males 1 Apical margins of tergite 3 (4 whenvisible) of gaster with fine dense punctures (Fig. 106) Apical margins of tergite 3 without dense punctures 6 2 Segments 2-6 of gaster clearly visible in lateral view (Fig. 106); legs bright orange-yellow; 262 J. QUINLAN segment 3 of antenna curved, expanded apically, antenna black except for basal threesegments brownish trulla sp. n. (p. 271) - Segment 6 of gaster not visible in lateral view; legs never bright orange-yellow; segment 3 of antenna curved , sometimes expanded apically 3 3 Radial cell of forewing at most 2-5 x as long as broad (Fig. 135); antenna yellowish, segment 3 less than 1-2 x as long as segment 4 (Fig. 109), weakly curved 4 - Radial cell of forewing 3 x as long as broad (Fig. 130); antenna brownish black, segment 3 strongly curved , swollen apically (Fig. 110) 5 4 Rs+M and M of forewing clearly pigmented, base of wing infuscate or weakly fumate, radial cell 3 x as long as broad; pronotum either side of pronotal plate weakly sculptured; segment3 of antenna strongly curved, weakly swollen apically (Fig. 108); notaulices sharply converg-ing towards scutellum (cf. Fig. 114) conversa sp. n.(p. 263) - Rs+M and M of forewing not pigmented, base of wing sometimes infuscate, radial cell 4 x as long as broad; pronotum either side of pronotal plate smooth; segment 3 of antenna curved,strongly swollen apically (Fig. 115); notaulices converging gently towards scutellum (Fig. 85) iiitidaBenoit(p. 269) 5 Segment 3 of antenna at most 1-2 x length of 4, weakly curved (Fig. 109); wings weakly fuscous basally lana sp. n. (p. 267) - Segment 3 of antenna 1-6 x length of 4, strongly curved, swollen apically (Fig. 110); wings hyaline marcel/us sp. n. (p. 268) 6 Segment 3 of antenna 1 -5 x length of 4, weakly curved on outer margins, swollen distally (Fig. 107), 4-10 each decreasing in length (each segment shorter than preceding one), 11-13subequal, shorter than 10, all segments black; forewing with vein RI thinner and less stronglypigmented than Rsi or Rs 2 , Rs+M and M not indicated, forewing not infuscate basally (Fig.Ill); scutellar disc reticulate-rugose, scutellar cup elongate (longer than wide), not extend-ing to apex of disc (Fig . 1 1 4) ; sides of pronotum not sculptured furcula sp . n . (p . 266) - Segment 3 of antenna at most 1-2 x length of 4, weakly curved on outer margin, sometimes swollen distally (Fig. 143), 4-5 subequal, 6-8 subequal, each shorter than 5, 9-10 subequal,each shorter than 8, 11-14 shorter than 8, each progressively shorter than the precedingsegment, basal segments brownish yellow, apical segments darker; forewing with veins RI,Rsi and Rs 2 of equal thickness, base of wing sometimes infuscate; side margins of pronotumwith or without sculpture; scutellar disc either punctate-reticulate or rugose, scutellar cupeither as wide as long or longer than wide 7 7 Notaulices converging gradually towards scutellum (Fig. 92); scutellar cup elongate, narrow, scutellar disc punctate-reticulate, visible in lateral and apical view; side margins of pronotumwith canaliculate sculpture; wings with conspicuous fuscous or smoky base (Fig. 112)(difficult to separate from advena) fetura sp. n. (p. 265) - Notaulices converging sharply at a point two-thirds of way towards scutellum (cf. Fig. 114); scutellar cup elongate or oval, lateral margins only of scutellar disc visible in dorsal view,apex of disc obscured by cup (cf. Fig. 1 14) ; side margins of pronotum without sculpture ; wingnot fumate basally 8 8 Radial cell of forewing at least 3 x as long as wide; scutellar cup almost as wide as long, truncated apically (cf. Fig. 114), scutellar disc rugose; pronotum with scattered pubescence on later al margins ; tergites 1-4 visible in lateral view perangusta sp. n.(p.270) - Radial cell of forewing at most 2 x as long as wide (Fig. 95); scutellar cup longer than wide, oval, rounded apically (Fig. 104); scutellar disc punctate-reticulate; pronotum sculptured onlateral margins advena (p. 262) ; ftucca-group; compressa-group Eucoilidea advena sp. n. (Figs 94, 101,112, 113) DESCRIPTION. $ . Antenna 13-segmented, filiform, segments 3-5 subequal, 6-10 each progressively shorterthan preceding segment (Fig. 113). Head, viewed frontally, smooth, shining, frontal line not pronounced,malar space with epistomal suture, mandibles bidentate, cheeks converging sharply, eyes further apartmeasured medially than height of an eye. Pronotal plate not conspicuous, lateral foveae not enclosed, i.e.open (cf. Fig. 101); mesoscutum with distinct notaulices, converging gently on approach to scutellum butnot merging, scutellar disc rugose-reticulate, visible laterally and posteriorly, scutellar cup longer than AFROTROPICAL EUCOILIDAE 263 broad, apical half with a large fovea (Fig. 104), scutellar foveae deep, lateral bars of scutellum broad andpolished; propodeum with parallel carinae, mesopleura smooth, shining, with a distinct suture. Segment 1of gaster short, crenulate, as wide as long, segment 2 the largest, occupying most of visible area in lateralview, segment 3 partially visible in lateral view, gaster impunctate, hypopygium with long basal setae,ovipositor sheath long (cf. Fig. 94). Legs yellowish orange, tibiae with two apical spurs, claws simple.Wings pubescent on surface, with apical fringe of hairs, radial cell of forewing closed on wing margin,elongate, Rs+M and M indicated, pigmented basally, base of wing with a smoky appearance (cf. Fig. 112).Colour: antenna yellow basally, apical segments dark, head and thorax black, gaster chestnut-brown-black, legs yellowish orange. Cf . Antenna 15-segmented, filiform, segment 3 the largest, weakly curved on inner margin, twisted andexcavate medially on outer margin (included by association). MATERIAL EXAMINED Holotype $, Zaire: Rutshuru (riv. Kanzarue), 1200 m, 16.vii.1935 (G. F. de Witte) (MRAC). Paratypes. Cameroun: 4 $ (BMNH). Kenya: 5 cf (BMNH). Rhodesia: 2 $ , 2 cf (BMNH). South Africa:5 , 4 cf (BMNH). Southern Yemen: 1 $ (BMNH). Uganda: 2 $ , 5 cf (BMNH), 1 cf (ZMC). Zaire: 198$, 35 Cf (MRAC). Zambia: 1 $, 2 cf (BMNH). Zimbabwe: 1 $ (BMNH). REMARKS. This species is closely related to bucca but is distinguished by the length of the scutellar cup, theshape of the pronotal plate and the convergence of the notaulices. Eucoilidea bucca sp. n. (Figs 97, 103) DESCRIPTION. 9 Antenna 13-segmented, subclavate, segments 3-6 subequal in length, 9-11 equal in length(Fig. 103). Face, viewed frontally, smooth, polished, malar space with epistomal suture, eyes further apart,measured medially, than height of an eye. Pronotum either side of pronotal plate weakly aciculate.Mesoscutum smooth, polished, notaulices distinct, converging sharply towards scutellum two- thirds ofway from posterior (Fig. 97); scutellar fovea deep, smooth, shining, scutellar cup large, with a large centralfovea, the periphery with a ring of smaller fovea; scutellar disc reticulate-rugose with long hairs on its apex;carinae of propodeum diverging at base , either side of carinae with sparse hairs , medial area with scatteredhairs; mesopleural suture distinct, surface smooth, polished. Segment 1 of gaster short, wider than long,strongly crenulate, tergite 2 the largest in lateral view, tergites 3 and 4 partially visible, hypopygium viewedlaterally, broad, ventral surface with long hairs, ventral spine as long as hypopygium, legs orange-yellow,hind metatarsus as long as tarsal segments 2-4 combined, tibia with two apical spurs, tarsal claws simple.Wings pubescent on surface, apical margins with a hair fringe, radial cell of forewing closed on margin, 2 xas long as broad, Rs+M and M weakly indicated, base of wing with a smoky hue. Colour: antenna withbasal segments yellowish, apical segments blackish, head and thorax black, gaster dark chestnut-red, legsyellowish. Cf . Antenna 15-segmented (included by association). MATERIAL EXAMINED Holotype $, Zaire: St Edouard, Kamanda, 8.iv.l936 (L. Lippens) (MRAC). Paratypes. Ethiopia: 1 cf (BMNH). Kenya: 8 $ (BMNH). South Africa: 3 $ (BMNH). Uganda: 2 $(BMNH). Yemen: 1 $ (BMNH). Zaire: 121 $, 7 cf (MRAC). Zambia: 1 cf (BMNH). HOST DATA. 1 $ from South Africa is labelled ex Munroella myiopitina Bezzi (Trypetidae).REMARKS. Superficially very similar to advena but differing in the shape of the pronotal plate. Eucoilidea compressa sp. n. (Figs 91, 94, 97, 121,128) DESCRIPTION. $. 13-segmented, filiform, segments 3-6 subequal, 7-13 with rhinaria, weakly swollenmedially (Fig. 91). Face, viewed frontally, smooth, polished, with dense pubescence either side of frontalarea, eyes weakly converging towards clypeus, eyes further apart than height of an eye measured medially.Pronotal plate very weakly produced forward, anterior and posterior parts fused medially by a bridge,fovea on either side open (cf. Fig. 128); pronotum either side of pronotal plate with scattered setae.Mesoscutum smooth, polished, notaulices distinct, converging sharply towards scutellum at a pointtwo-thirds of way towards scutellum but not meeting at juncture; lateral bars of scutellum smooth,polished, scutellar fovea smooth, polished, shallow, scutellar disc reticulate-rugose, rounded apically, 264 J. QUINLAN visible on lateral margins, scutellar cup large, broad, extending almost to apex of disc, rim of cup lighterthan central area which has large fovea, outer margins of cup with small pits or fovea on lower half adjacentto rim (cf. Fig. 97). Mesopleural suture distinct, surface smooth, polished, metapleura with a dense tuft ofpubescence on anteroventral margin; propodeal carinae weakly bowed medially, obscured by densepubescence. Segment 1 of gaster short, as wide as long, crenulate, tergites 1-4 visible in lateral view (Fig.94), tergite 2 the largest with a few sparse hairs on lateral margins at base, 2-4 impunctate, lenticular inshape, hypopygium not pronounced, ovipositor short. Legs orange, coxa elongate, swollen basally,trochanters elongate, femur and tibia normal, hind metatarsus as long as tarsal segments 2-4 combined.Wing surface pubescent, apical fringe of hairs short, radial cell of forewing closed on wing margin, twice aslong as wide measured medially (Fig. 121). Rs+M and M indicated, weakly pigmented. Colour: antennaorange-yellow, head and thorax black, gaster orange-yellow. d". Antenna 15-segmented, 3rd segment curved, swollen distally (included by association). MATERIAL EXAMINED Holotype $, Zaire: P. N. U. Lusifiga (1700 m, ll-18.vii.1947 (Mis. G. F. de Witte} (MRAC). Paratypes. Kenya: 4 $, 3 cf (BMNH). South Africa: 3 $ (BMNH). Uganda: 1 cf (BMNH). Zaire: 35 $(MRAC). Zimbabwe: 4 $ (BMNH). REMARKS. This species is closely related to urundiensis but is distinguished by the shape of the antenna. Eucoilidea converse sp. n. (Figs 85, 87, 108) DESCRIPTION. $ . Antenna 13-segmented, almost filiform, segment 3 longer than 4, all flagellar segments atleast twice as long as wide, segments 5-13 each progressively shorter than preceding segment, all longerthan segment 4 (Fig. 87), all segments of equal width. Face, viewed frontally, smooth, shining; notaulicesdistinct, converging sharply towards scutellum two-thirds of distance from it, scutellar foveae smooth,polished, deep, scutellar disc reticulate-rugose with long scattered hairs not obscured by scutellar cup,scutellar cup elongate, longer than broad, with small pits or fovea around rim of cup, medial area with alarge fovea (cf. Fig. 85); carinae of propodeum indistinct, sculptured medially. Mesopleura smooth,polished, suture complete. Segment 1 of gaster wider than long, crenulate, segment 2 the largest in lateralview, apical half with dense punctures, remaining segments not visible, hypopygium short, not generallyvisible. Legs orange-yellow, metatarsus as long as tarsal segments 2-4 combined, tibia with two apicalspurs, tarsal claws simple. Wings pubescent on surface, apical hair fringe distinct, radial cell of forewingclosed on wing margin, veins Rs+M and M indicated, not pigmented, radial cell almost 3 x as long 3sbroad, 2rm as broad as long. Colour: antenna yellowish basally, brownish apically (entirely yellow inMauritian specimen), head and thorax blackish, gaster chestnut-brown, legs yellow.Cf . Antenna 15-segmented, filiform, segment 3 curved, longer than 4 (Fig. 108). MATERIAL EXAMINED Holotype $, Zaire: 3.L1952 (H. de Saeger) (MRAC). Paratypes. Cameroun: 1 cf (BMNH). Gold Coast: 20 $ , 3 cf (BMNH). Kenya: 6 $ , 4 cf (BMNH); 1 $(ZMC). Madagascar: 8 $, 3 cf (BMNH); 10 $, 6 cf (MRAC); 1 cf (MNHN). Mauritius: 1 $, 1 cf(BMNH). Nigeria: 7 $, 10 cf (BMNH). South Africa: 3 $, 1 cf (BMNH). Tanzania: 1 $ (BMNH).Uganda: 20 $ (BMNH). Zimbabwe: 5 $, 2 cf (BMNH). REMARKS. This species is similar toparma but the antennae have an overall filiform appearance; it is alsovery close to nitida but is distinguished by the shape of the antenna and the notaulices. Eucoilidea dubia sp. n. (Fig. 116) DESCRIPTION. $. Antenna 13-segmented, filiform, 5 shorter than 4, 4 and 5 subequal in length, 6 and 7subequal, each shorter than 5, 8 shorter than 7, 9 and 10 each shorter than 8, subequal in length, 11 shorterthan 10, 12 shorter than 11, 13 longer than 12. Head, viewed frontally, smooth, polished, malar space witha distinct ridge with scattered setae either side of supraclypeal area, mandibles dentate, eyes measuredmedially as far apart as height of an eye. Pronotum weakly pubescent, aciculate either side of pronotalplate. Pronotal plate small, not protruding. Mesoscutum smooth, polished, notaulices distinct, convergingapically, but not meeting, scutellar fovea deep, lateral bars of scutellum smooth, polished, scutellar cupelliptical, longer than wide, not extending to apex of disc, scutellar disc reticulate-rugose with apical fringe AFROTROPICAL EUCOILIDAE 265 of long hairs; propodeal carinae parallel. Segment 1 of gaster as wide as long, short, crenulate, segment 2 ofgaster the largest in lateral view, smooth, impunctate, other segments not visible in lateral or dorsal view,hypopygium not projecting, with long hairs on ventral surface, ovipositor usually projecting, gaster as inFig. 116. Legs yellowish orange, tibiae with two apical spurs, tarsal claws simple. Wing surface pubescent,apical margins with a hair fringe, radial cell of forewing closed, longer than broad, Rs+M and M indicated,base of wing pigmented. Colour: antenna orange-yellow basally, apical segments brownish, head andthorax black, gaster reddish basally, blackish apically.Cf unknown. MATERIAL EXAMINED Holotype <j>, South Africa: E. Cape Province, Katberg, 4,000 ft (1,220 m), xii.1932 (R. E. Turner)(BMNH). Paratypes. South Africa: 8 $ (BMNH). Zaire: 17 $ (MRAC). REMARKS. This species is related to mauri but differs in the form of antennae and the notaulices. Eucoilidea extraria sp. n. (Figs 76, 77, 97, 123, 128) DESCRIPTION. $ . Antenna 14-segmented, subclavate, 8-14 forming a weak club, segments 3-8 subequal inlength, 9-11 subequal, shorter than preceding segments, 12 and 13 each shorter than segment 11, apicalsegment as long as segment 3 (Fig. 76). Head, viewed frontally, smooth, polished, frontal area raised,malar space with a distinct carina, mandibles tridentate, eyes diverging at top and bottom. Pronotal plateweakly protruding, pronotum either side of plate smooth, weakly pubescent, pronotal plate foveaelongate, open on lateral margins (cf. Fig. 128). Mesoscutum smooth, shining, with weak crenulatesculpture on margins, juncture with pronotum crimped, notaulices distinct, sharply converging at a pointtwo-thirds from pronotum (in outline similar to a wine glass) (cf. Fig. 97) ; scutellar disc punctate-reticulate,scutellar cup very large, almost as wide as long, with a large central fovea, the rim of cup with a ring ofsmaller fovea, scutellar disc viewed dorsally, visible only on lateral margins. Mesopleura smooth, shining,with a distinct suture; propodeal carina diverging basally. Segment 1 of gaster viewed dorsally, wider thanlong, strongly crenulate (cf. Fig. 77), segment 2, viewed laterally, the largest, apical quarter with widelyscattered punctures, segment 3 partially visible, remaining segments not visible, hypopygium distinct,ovipositor as long as visible part of hypopygium. Legs orange-yellow, tibiae with two apical spurs, clawssimple. Wing surface pubescent, apical margins with a fringe of hairs, radial cell of forewing closed, longerthan broad, Rs+M and M weakly indicated (Fig. 123). Colour: antenna dark orange-yellow except apicalsegments dark brown, head and thorax black, gaster dark chestnut-red-black.Cf unknown. MATERIAL EXAMINED Holotype $, Madagascar: La Mandrake, ii.1944 (A. Seyrig) (MRAC).Paratype. 1 9 , same data as holotype (MRAC). REMARKS. This species is easily distinguished by the extra segment in the antenna, the very large scutellarcup and the almost completely orange antenna. Eucoilidea fetura sp. n. (Figs 89, 92, 112, 117, 118) DESCRIPTION. $. Antenna 13-segmented, segments 3 and 4 equal in length, 5-12 each shorter than 4,overall shape filiform (Fig. 89). Head, viewed frontally, smooth, shining, face with scattered hairs,mandibles tridentate, eyes further apart at top of eyes than medially (cf. Fig. 117), malar suture distinct.Pronotal plate not protruding, with elongate fovea on either side of medial bridge (cf. Fig. 128). Pronotumwith sparse pubescence either side of plate; mesoscutum smooth, polished, notaulices distinct, convergingin apical third towards scutellum, lateral bars of scutellum smooth, scutellar fovea large, scutellar cupelliptical with a large fovea basally, scutellar disc punctate-reticulate, disc visible laterally and apically (Fig.92), apex of disc with a fringe of long hairs; mesopleural suture distinct; propodeum sculptured, carinaeindistinct. Segment 1 of gaster short, as broad as long, crenulate, segment 2 the largest, occupying most ofvisible area in lateral view, segment 3 partially visible, gaster impunctate, hypopygium short with sparsehairs ventrally, ovipositor protruding (Fig. 118). Legs yellowish orange, tibiae with two apical spurs, clawssimple. Wings pubescent, margins of wings with a hair fringe, radial cell of forewing closed on wing margin, 266 J. QUINLAN longer than wide, Rs+M and M indicated, base of wings infuscate (Fig. 112). Colour: antenna yellowbasally, with apical four segments blackish, head and thorax black, gaster blackish, chestnut-red basally.CT. Antenna 15-segmented, filiform, segment 3 curved, weakly swollen apically. MATERIAL EXAMINED Holotype , Zimbabwe: Salisbury, Chishawasha, ii.1979 (A. Watsham) (BMNH). Paratypes. Cameroon: 4 $ (BMNH). Nigeria: 2 $ (BMNH). South Africa: 6 $ , 4 cf (BMNH). Senegal:1 $ (BMNH). Uganda: 4 $, 1 cf (BMNH); 1 $ (ZMC). Zaire: 26 $, 38 cf (MRAC). REMARKS. This species is easily confused with advena but in the female the overall shape and the ratios ofthe antennal segments distinguish it. In both sexes the apex of the wings tends to be smoky. Eucoilidea furcula sp. n. (Figs 85, 107, 111,119, 120) DESCRIPTION. 9 . Antenna 13-segmented, clavate, segment 3 longer than 4, 4-10 subequal in length, 11-12subequal, each shorter than 10 (Fig. 120). Head, viewed frontally, smooth, polished, with scattered hairseither side of supraclypeal area, mandibles tridentate, eyes as far apart as the height of an eye measuredmedially, malar suture distinct. Pronotal plate not protruding, with lateral fovea open; pronotum weaklypubescent either side of pronotal plate; mesoscutum smooth, polished, notaulices distinct, convergingsharply on nearing scutellum, not confluent, scutellar foveae large, shallow, lateral bars smooth, scutellarcup large with apical depression, longer than broad measured medially, scutellar disc reticulate-rugose,visible laterally and apically, viewed dorsally (cf. Fig. 85), apex of disc with a fringe of long hairs;mesopleural suture distinct, metapleura weakly ridged, anteroventral cavity with a tuft of hairs; propodealcarinae close, parallel, medially with canaliculate sculpture. Segment 1 of gaster short, as broad as long,crenulate, sometimes obscured by tergite 2. Segment 2 the largest in lateral view, with two or three hairspresent basally on lateral margins, gaster impunctate, remaining segments of gaster not visible, hypopy-gium not pronounced, with sparse hairs present on ventral surface, ovipositor protruding, gaster (Fig.119). Legs yellowish orange, tibiae with two apical spines, claws simple. Wing surface pubescent, apicalmargins with a hair fringe, radial cell of forewing closed, lightly pigmented, as broad as long measuredmedially, Rs+M and M not indicated (Fig. Ill), base of wing weakly infuscate. Colour: antenna palebrownish yellow, head and thorax black, gaster dark chestnut-brown.Cf . Antenna 15-segmented, segment 3 1-5 x length of 4 (Fig. 107). MATERIAL EXAMINED Holotype $, South Africa: Witzenberg Vail, SW. Cape Province, 3,000 ft [915 m], 19.U921 (R. E.Turner) (BMNH). Paratypes. Ethiopia: 1 $, 1 cT (BMNH). Kenya: 2 $, 1 cf (BMNH); 1 $ (ZMC). South Africa: 50 $,15 Cf (BMNH). Uganda: 6 $, 4 cf (BMNH). Zaire: 1 (MRAC). Zimbabwe: 6 $, 1 cf (BMNH). REMARKS. This species is distinguished from others by the shortish clavate antenna, the third segment beingdistinctly longer than the fourth. It is separated from mauri, a closely related species, by the form of thenotaulices and the shape of the scutellar cup. Eucoilidea lacerta sp. n. (Figs 81, 82, 85, 117) DESCRIPTION. $. Antenna 13-segmented, very weakly club-shaped, segment 3 shorter than 4 (Fig. 81),5-13 each progressively shorter than the preceding segment, all with rhinaria. Face smooth, shining, withsparse scattered pubescence, eyes further apart measured at top of eye than at bottom, malar suturedistinct (Fig. 117). Pronotal plate not sharply protruding, pronotum either side of pronotal plate with longscattered hairs, surface aciculate. Mesoscutum smooth, polished, notaulices distinct, converging towardsscutellar suture, anterior parallel lines weakly indicated; lateral bars of scutellum polished, scutellar foveaeshallow, polished, scutellar cup long, wide, extending almost to apex of scutellar disc, apical half with alarge depression, rim with a border of smaller pits along edge of lower margins, scutellar disc rugose-reticulate, apex with a fringe of long hairs, apical margin obscured by scutellar cup (cf. Fig. 85).Mesopleura smooth, polished, suture distinct; propodeum with pronounced parallel carinae. Segment 1 ofgaster short, as broad as long, crenulate, segment 2 the largest viewed laterally, punctate on apical margins,segment 3 partially visible, hypopygium short, ovipositor not protruding. Legs yellowish orange, withscattered setae, tarsal claws simple. Wing surface pubescent, with apical fringe of hairs, radial cell of AFROTROPICAL EUCOILIDAE 267 fore wing closed on margin, elongate, Rs+M and M indicated (Fig. 82). Colour: antenna yellow basally,6-12 dark, head, thorax and first segment of gaster black, second segment of gaster chestnut-brown at base,becoming black towards apex.Cf unknown. MATERIAL EXAMINED Holotype $ , Ivory Coast: Pakobo, 44 km N. Tlassale, x.1962 (/. Decelle) (MRAC).Paratypes. Ivory Coast: 45 $, same data as holotype. Zaire: 4 $ (MRAC). REMARKS. The females of this species and marcellus are distinguished by the 4th segment of the antennabeing clearly longer than the 3rd. Eucoilidea lana sp. n. (Figs 86, 109, 124, 125, 126) DESCRIPTION. $ . Antenna 13-segmented, filiform, segment 4 longer than 3, 7-13 each progressively shorterthan preceding segment, all of equal width, rhinaria present on segments 4-13 (Fig. 124). Face, viewedfrontally, smooth, shining, malar space with a distinct epistomal suture, eyes diverging at top and bottom.Pronotal plate not projected forward, foveae large, open on lateral margins of medial bridge (Fig. 125).Pronotum either side of pronotal plate strongly aciculate (Fig. 125). Mesoscutum smooth, polished,notaulices distinct, converging sharply towards scutellum two-thirds from it but not confluent (Fig. 114),scutellar fovea deep, lateral bars polished, scutellar disc punctate-reticulate, with long scattered hairs onapical margin, scutellar disc visible on lateral margins, viewed dorsally, apex obscured by large scutellarcup, cup with a large apical depression, the rim with small pits along edge (cf. Fig. 86); carinae ofpropodeum weakly bowed medially, pubescent on either side. Mesopleura with a distinct suture. Segment1 of gaster short, as broad as long, crenulate, segment 2 the largest in lateral view, with dense punctures onapical margins, other segments not visible. Legs brownish yellow, metatarsus as long as tarsal segments 2-4combined, tibiae with 2 apical spurs, tarsal claws simple, wing surface pubescent, apical margins with a hairfringe, radial cell of forewing closed on margin, 2 x as long as wide, Rs+M and M weakly indicated.Colour: antenna blackish, head and thorax black, gaster chestnut-red. Cf . Antenna 15-segmented, filiform, segment 3 the largest, strongly curved, swollen apically (Figs 109,126). MATERIAL EXAMINED Holotype $, Zaire: Kivu, Nyyongera (near Rutshuru), Batumba, 1218 m, 17.vii.1935 (MRAC).Paratypes. Zaire: 4 $ (MRAC). REMARKS. This species is very close to lacerta but differs from it on the antennal shape and the sculpture ofthe scutellar disc. The male is similar to marcellus but differs by the length of the antennal segments. Eucoilidea leptis sp. n. (Figs 86, 127, 128) DESCRIPTION. $. Antenna 13-segmented, filiform, without trace of a distinct club, segments 3-13 withrhinaria, segment 3 longer than 4, 5-7 subequal, each slightly shorter than 4, 8-13 each shorter than 5,segment 3 distinctly curved, swollen apically (Fig. 127). Head, viewed frontally, with eyes further apartmeasured medially than height of an eye, face with long scattered setae, malar space with a distinctepistomal suture, mandibles tridentate. Pronotum either side of pronotal plate sculptured, pronotal platenot projected forward, fovea either side of medial bridge open laterally (Fig. 128). Mesoscutum smooth,polished, notaulices distinct, weakly converging on approach to scutellum, further apart at juncture withscutellum than width of a notauli (Fig. 86); scutellar foveae broader than long, lateral bars polished,scutellar disc visible on lateral margins, reticulate-rugose, apex of disc with long setae, scutellar cup longerthan broad, obscuring apex of disc, cup with a large apical depression or fovea, rim of cup with a circle ofminute fovea or pits; carinae of propodeum parallel, densely pubescent on outer margins. Mesopleura witha distinct suture, antero ventral cavity of metapleura with a few hairs. Segment 1 of gaster short, wider thanlong, crenulate, segment 2 the largest in lateral view, apical half densely punctate, segments 3 and 4partially visible, punctate, hypopygium short, setae present on margins. Legs slender, coxae, femora andtibiae brownish, tarsi yellow, coxae short, swollen, femora swollen on basal half, metatarsus as long astarsal segments 2-4 combined, tibiae with two apical spines, tarsal claws simple. Wings densely pubescent,apical margins with a hair fringe, radial cell of forewing closed on margin, 3 x as long as broad measured 268 J. QUINLAN medially, vein 2rm almost square. Colour: antenna dark brownish black, head and thorax black, gasterblack except for brownish apex.Cf unknown. MATERIAL EXAMINEDHolotype $, Cameroon: Nkoemvon, viii-xi.1980 (D. Jackson) (BMNH). REMARKS. Distinguished from nitida by the antennal ratios and the scutellar cup. Eucoilidea marcellus sp. n. (Figs 78, 110, 129, 130) Description. $. Antenna 13-segmented, filiform, segment 3 1-2 x as long as 4, 5-12 each progressivelydecreasing in length (Fig. 129), 4-13 with rhinaria. Face viewed frontally, smooth, shining, with scatteredpubescence, malar space with a distinct suture, mandibles tridentate, maxillary palp 4-segmented, labialpalp 2-segmented, eyes elongate, oval, further apart than height of an eye measured medially; pronotalplate not protruding, viewed fronto-dorsally minute, with lateral fovea open either side of medial bridge,pronotum either side of pronotal plate densely pubescent. Mesoscutum smooth, polished, notaulicesdistinct, converging towards scutellum at a point two-thirds of way towards scutellar suture, not confluentat juncture with scutellum; lateral bars of scutellum polished, scutellar fovea large, deep, scutellar cuplarge, extending to apex of scutellar disc, convex basally, with a large fovea on apical half with a ring ofsmaller punctures or fovea around rim of cup, scutellar disc reticulate-rugose, with long setae on apicalmargins. Propodeal carinae subparallel, converging basally. Mesopleura smooth, polished, carinaecomplete, metapleura with two ridges, anteroventral cavity with a few hairs, ventral margin of meso-pleuron pubescent. Segment 1 of gaster canaliculate, as broad as long, viewed laterally, crescent-shaped(partially obscured by tergite 2), tergite 2 of gaster the largest in lateral view, with a few sparse hairs at baseon lateral margins, apical half densely punctate, remaining segments not visible, hypopygium small,ventral spine long. Legs short, robust, yellow, fore and mid coxae with long hair fringe, femora swollenmedially, tibiae and tarsi densely pubescent, tarsal claws simple. Wings pubescent on surface, with apicalfringe of hairs, radial cell of forewing closed on anterior margin, veins Rs+M and M distinct, weaklypigmented, radial cell of forewing elongate, 2-6 x as long as broad (Figs 78, 130). Colour: antennabrownish black, head and thorax black, gaster dark brown-black, legs yellowish. Cf . Antenna 15-segmented, filiform, segment 3 the largest, strongly curved, swollen apically (Fig. 110). MATERIAL EXAMINED Holotype $, Madagascar: Tan, Perinet, 27.iv-3.v.l983 (J. S. Noyes & M. C. Day) (BMNH). Paratypes. Madagascar: 4 $, 4 cf, same data as holotype. Mauritius: 3 d" (BMNH). Zaire: 1 $(MRAC). REMARKS. This species is separated from tyrus by the shape of the scutellum and the radial cell of theforewing (see key). Eucoilidea maur/sp. n. (Figs 131, 132) DESCRIPTION. $. Antenna 13-segmented, clavate, segment 3 1-2 x length of 4, 5-12 each graduallydecreasing in length, 13 longer than 12, 5-13 with rhinaria forming a weak club (Fig. 131). Head, viewedfrontally, smooth, shining, with sparse pubescence, long setae on anterior tentorial pit, eyes weaklyconverging, as far apart as height of an eye measured medially, malar suture distinct. Pronotal plate notprojected forward, lateral fovea either side of medial bridge open; mesoscutum smooth, polished,notaulices weakly converging towards scutellum, widely spaced at juncture with scutellar suture (Fig. 132);lateral bars of scutellum polished, scutellar foveae large, smooth, shallow, scutellar cup large, oval, with alarge medial fovea or depression, scutellar disc viewed dorsally, visible on lateral margins only, reticulate-rugose. Mesopleura smooth, polished, suture distinct, metapleura polished, weakly ridged on margin,anteroventral cavity pubescent; propodeal carinae bowed medially, sparsely pubescent. Segment 1 ofgaster partially visible, crenulate, as broad as long, segments 1-3 visible in lateral view, impunctate, tergite2 the largest, hypopygium not pronounced, ventral spine with long sparse setae. Legs orange-yellow, coxaeelongate, swollen basally, trochanters normal, femora short, swollen medially, tibiae normal with twoapical spines, metatarsus shorter than tarsal segments 2-5 combined, claws simple. Wing surfacepubescent, apical margins with a hair fringe, radial cell of forewing closed on margin, 2-5 x as long as broad AFROTROPICAL EUCOILIDAE 269 measured medially, veins heavily pigmented, vein 2rm distinct, in form of a spur, Rs+M and M indicated,weakly pigmented. Colour: antenna light yellow-brown basally, dark at apex, head and thorax blackish, gaster chestnut-brown. C? unknown. MATERIAL EXAMINED Holotype $, South Africa: Mossel Bay, Cape Province, iv.1927 (R. E. Turner) (BMNH).Paratype. 1 $, same data as holotype (BMNH). REMARKS. This species closely resembles furcula and could, without close examination, be mistaken for it.The form of the notaulices and the large, almost round cup, distinguish it from related species. Eucoilidea nitida (Benoit) comb. n. (Figs 115, 133, 134, 135,137)Afrostilba nitida Benoit, 1956: 544. Holotype $, ZAIRE (MRAC) [examined]. DESCRIPTION. $. Antenna 13-segmented, subclavate, segments 3 and 4 equal in length, 4-13 forming aweak club, each segment slightly shorter and wider in length than preceding one (Fig. 133), apical segmentnever twice as long as wide. Face, viewed frontally, smooth, shining, malar space with a distinct epistomalsuture, eyes diverging at top and bottom, frontal area pronounced, mandibles tridentate. Pronotum eitherside of pronotal plate smooth, sometimes with weak sculpture on the margins. Pronotal plate not projectedforward, lateral fovea either side of medial bridge open laterally, fovea broad, narrow, separated by anarrow bridge medially (Fig. 134). Mesoscutum smooth, polished, notaulices distinct, gradually converg-ing on approach to scutellar suture, almost as far apart at juncture as half width of scutellar cup, scutellarfoveae smooth, polished, scutellar disc punctate-reticulate, with scattered hairs, scutellar cup almost oval,with a large apical fovea (Fig. 137), rim of cup with small fovea on lateral margin, apex of disc obscured bycup; carinae of propodeum parallel, pubescent on outer margins. Mesopleura with a distinct suture,metapleura smooth, polished. Segment 1 of gaster short, wider than long, crenulate, segment 2 the largestin lateral view, densely punctured on apical half viewed laterally, remaining segments not visible,hypopygium short. Legs yellow, coxae yellow, meta- tarsus as long as tarsal segments 2-4 combined, tibiaewith two apical spurs, tarsal claws simple. Wings pubescent on surface, apical margins with a hair fringe,radial cell of forewing closed on anterior margin, Rs+M and M weakly indicated, radial cell 4 x as long asbroad, elongate (Fig. 135), 2rm as broad as long. Colour: antenna yellowish black basally, black apically,head and thorax black, gaster dark chestnut-red-brown, legs yellowish. Cf . Antenna 15-segmented, filiform, segment 3 the largest, strongly curved, swollen apically, longerthan 4 (Fig. 115). MATERIAL EXAMINED Holotype $, Zaire: Kivu, Rutshuru, ll.v.1936 (L. Lippens) (MRAC). Paratypes. Ghana: 2 $, 5 C? (BMNH). Kenya: 11 $, 1 cf (BMNH). South Africa: 1 $ (BMNH). Zaire:27 $, 2 Cf (MRAC). Zambia: 3 $ (BMNH). Zimbabwe: 1 $ (BMNH). REMARKS. A difficult species to separate from conversa, but distinguished from it by the overall shape of theantenna and the convergence of the notaulices towards the scutellar suture (Figs 133, 137). Eucoilidea pallida sp. n. (Figs 83, 93, 112, 117, 141) DESCRIPTION. $. Antenna 13-segmented, subclavate, pale yellow, segments 9-13 forming a weak club.Face, viewed frontally, smooth, polished, frontal area raised, face with long scattered pubescence, malarspace with a distinct carina, mandibles tridentate, eyes further apart than height of an^ye measuredmedially, diverging at top and bottom (cf. Fig. 117). Pronotum either side of pronotal plate crenulate-foveolate (cf. Fig. 83), pronotal plate not protruding, lateral fovea either side open laterally. Mesoscutumsmooth, shining, the juncture with pronotum weakly channelled, notaulices distinct, converging sharplytowards scutellum at a point two-thirds distance from scutellar suture (Figs 93, 141), scutellar disctransversely strigose, visible on lateral margins, viewed dorsally, apex sometimes obscured by scutellarcup, scutellar cup elongate, with a large apical depression, rim of cup with a ring of smaller fovea or pits.Mesopleura with a distinct suture, carinae of propodeum parallel. Segment 1 of gaster short, as broad aslong, crenulate, segment 2 the largest, obscuring in lateral view remaining tergites of gaster, impunctate, 270 J. QUINLAN hypopygium hardly visible, ovipositor protruding. Legs orange-yellow, tibiae with two apical spines, clawssimple. Wings pubescent on surface, apical margins with a distinct hair fringe, radial cell of forewing closedon margin but very weakly pigmented on anterior margin (cf. Fig. 112), at least twice as long as wide,Rs+M and M indicated but not pigmented, 2rm distinct. Colour: antenna pale yellow, head and thoraxblack, gaster chestnut-red-black.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Mossel Bay, Cape Province, 5-31.vii.1921 (R. E. Turner) (BMNH).Paratypes. South Africa: 10 $ (BMNH). Southern Yemen: 1 $ (BMNH). Zaire: 2 $ (MRAC). REMARKS. This species is close tofetura but can be separated by the very distinctive, bright yellow antennaand the large scutellar cup. It is far more robust thanfetura. A male from Zambia (BMNH) could be thisspecies, but without an associated female I am reluctant to treat it as such. Eucoilidea parma sp. n. (Figs 88, 138, 139, 140) DESCRIPTION. $? Antenna 13-segmented, subclavate, segments 3 and 4 subequal in length, segments 6-13forming a weak club, all with rhinaria, each segment longer than broad, slightly wider apically than at base(Figs 88, 138). Face, viewed frontally, smooth, shining, malar space with epistomal suture, mandiblestridentate. Pronotum with pubescence either side of pronotal plate, surface aciculate. Pronotal plate veryweakly produced, foveae either side open on lateral margins. Mesoscutum smooth, polished, notaulicesdistinct, converging sharply at a point one-quarter from scutellar suture, not confluent (Fig. 139), scutellarfoveae deep basally, scutellar disc punctate-reticulate, with long scattered hairs, scutellar disc visiblelaterally and apically, not obscured by scutellar cup; carinae of propodeum parallel, pubescent on eitherside. Mesopleura with a distinct carina. Segment 1 of gaster short, wider than long, crenulate, segment 2the largest in lateral view, densely punctured on apical third, segment 3 partially visible in lateral view,hypopygium short. Legs yellow, coxae orange-yellow, metatarsus as long as tarsal segments 2-4 combined,tibiae with 2 apical spurs, tarsal claws simple. Wings ciliate or pubescent on surface, with apical hair fringeon margins, radial cell of forewing closed on anterior margin, Rs + M and M not indicated, radial cell 3 x aslong as wide, elongate (Fig. 140), vein 2rm longer than broad. Colour: antenna yellowish without darkenedapical segments, head and thorax black, gaster chestnut-red.Cf unknown. MATERIAL EXAMINED Holotype $, Nigeria: Ile-Ife, W. State, vi.1973 (J. T. Medler) (BMNH).Paratypes. Nigeria: 2 $ (BMNH). Madagascar: 1 $ (BMNH). Zaire: 4 $ (MRAC). REMARKS. Similar to conversa but distinguished by the subclavate antenna. Eucoilidea perangusta sp. n. (Figs 95, 96, 137, 143) DESCRIPTION. $. Antenna 13-segmented, weakly subclavate, segment 3 very slightly shorter than 4, 6-13forming a very weak club, all with rhinaria (Fig. 96). Face, viewed frontally, smooth, shining, malar spacewith a distinct epistomal suture, eyes diverging at top and bottom, frontal area pronounced, mandiblestridentate. Pronotal plate weakly protruding viewed dorsally, lateral fovea open, wide and narrow,pronotum either side of plate with long scattered pubescence, surface with widely spaced aciculatesculpture. Mesoscutum smooth, polished, juncture with pronotum on lateral margins crimped, notaulicesdistinct, converging gently towards scutellum (cf. Fig. 137), scutellar fovea shallow, scutellar disccoriaceous, cup oval with a large central fovea, outer margins of cup without a ring of smaller fovea:propodeal carinae parallel. Mesopleura smooth, polished, with distinct suture. Segment 1 of gaster short,broader than long, crenulate, segment 2 the largest in lateral view, remaining segments not visible,hypopygium short, not protruding. Legs deep yellow, meta-tarsus shorter than tarsal segments 2-4combined, tibiae with two apical spines, tarsal claws simple. Wings pubescent on surface, apical marginswith a hair fringe, radial cell of forewing closed on anterior margin, elongate, 3-6 x as long as wide, Rs+Mand M weakly pigmented, 2rm broader than long (Fig. 95). Colour: antenna pale basally, apical 7 segmentsdark brown, head and thorax black, gaster orange-yellow in $ , brownish in cf .Cf . Antenna filiform, 3rd segment curved, swollen distally (Fig. 143). AFROTROPICAL EUCOILIDAE 271 MATERIAL EXAMINED Holotype $, Zambia: 15 km E. Lusaka, 13-22.xi.1979 (R. A. Beaver) (BMNH).Paratypes. Zimbabwe: 1 $ (BMNH). Zaire: 1 $ (MRAC). Zambia: 2 $, 4 cf (BMNH). Eucoilidea trulla sp. n. (Figs 79, 85, 107, 145, 147) DESCRIPTION. 9- Antenna 13-segmented, subclavate, segment 3 larger than 4, shorter than 4+5, 4 and 5subequal, 5-12 gradually decreasing in length (Fig. 79). Head, viewed frontally, smooth, shining, frontalline (frons) convex, malar space with epistomal suture distinct, mandibles tridentate, cheeks sharplyconverging. Pronotal plate not projecting, lateral fovea open on margin, sparsely pubescent on outermargins. Mesoscutum with notaulices distinct, converging two-thirds of way towards scutellum but notconfluent, scutellar disc punctate-rugose, clearly visible laterally and apically (cf. Fig. 85), scutellar cuplarge with a fovea basally, with traces of small punctures around rim, scutellar fovea large, polished.Mesopleura smooth, polished, suture distinct; carinae of propodeum parallel. Segment 1 of gaster short,wider than long, crenulate, segment 2 the largest in lateral view, impunctate, segments 3 and 4 punctate,hypopygium short with long basal setae, ovipositor sheath short (Fig. 145). Legs yellow-orange, tibiae with2 apical spurs, claws simple. Wings pubescent on surface, with apical fringe of hairs, radial cell of forewingclosed on margin, elongate, Rs+M and M indicated, not pigmented (Fig. 147). Colour: antenna black,head and thorax black, gaster black dorsally, chestnut-red ventrally. Cf . Antenna 15-segmented moniliform, 3rd segment the largest, weakly curved (Fig. 107). MATERIAL EXAMINED Holotype $ , South Africa: Natal, nr Pietermaritzburg, 21.iii.1980 (P. Joubert) (BMNH).Paratypes. South Africa: 1 $, 2 Cf , same data as holotype (BMNH). Zaire: 12 $ , 6 cf (MRAC). REMARKS. Closely related to a group of species in which the third antennal segment of the $ is clearlylonger than the 4th, and the gaster is finely punctate. Eucoilidea tyrus sp. n. (Figs 80, 148) DESCRIPTION. $. Antenna 13-segmented, filiform, segment 3 weakly curved, 1-2 x length of 4, 4-13 withrhinaria, 4-6 subequal in length, 7-13 each progressively shorter than the preceding segment. Head,viewed frontally, smooth, shining, eyes further apart measured medially than height of an eye, weaklyconverging, face with scattered setae, mandibles tridentate, basal tooth blunt, malar suture distinct.Pronotum either side of pronotal plate smooth with scattered pubescence, pronotal plate not conspicu-ously produced forward, medial bridge narrow, fovea on lateral margins open (Fig. 148). Mesoscutumsmooth, polished, with scattered hairs, notaulices complete, converging towards scutellum, further apartat juncture with scutellum than width of one notauli, scutellar cup slightly longer than broad, apical halfwith a large fovea, rim of cup with a number of minute fovea around edge, scutellar disc reticulate-rugose,visible in lateral and apical view. Mesopleura smooth, shining, suture distinct, metapleura ridged on upperpart, anteroventral cavity with a few sparse hairs; propodeal carinae parallel, densely pubescent on lateralmargins. Segment 1 of gaster crenulate, short, wider than long, segment 2 the largest in lateral view, apicalhalf finely punctate, visible part of segment 3 finely punctate, hypopygium pronounced, broad with longbasal setae, ventral spine produced past apex of gaster. Legs bright yellow, coxae short, broad, all femoraswollen medially, tibiae with two apical spurs, slender, as long as tarsi. Wing surface pubescent, apicalmargins with a hair fringe, radial cell of forewing closed on margin, 3-6 x as long as broad measuredmedially (Fig. 80), Rs+M and M indicated by weak pigmentation, base of wing not fumate. Colour:antenna yellow basally, apical segments darker, head and thorax blackish brown, gaster brownish, legsbright orange-yellow.Cf unknown. MATERIAL EXAMINEDHolotype , Cameroun: Nkoemvon, x-xi.1980 (D. Jackson) (BMNH). REMARKS. Closely related to marcellus but separated by the gaster and the form of the scutellum. 272 J. QUINLAN Eucoilidea urundiensis Benoit (Figs 98, 99, 100, 139)Eucoilidea urundiensis Benoit, 1956: 548. Holotype $, ZAIRE (MRAC) [examined]. DESCRIPTION. $. Antenna 13-segmented, subclavate, segment 3 longer than 4, 5 shorter than 4, 5-13 withrhinaria, forming a weak club (Fig. 98). Face, viewed frontally, smooth, shining, with long scatteredpubescence, malar space with epistomal suture, eyes slightly further apart than height of an eye measuredmedially, weakly converging, mandibles tridentate. Pronotal plate not produced forward, foveae eitherside of medial, bridge open. Pronotum either side of plate with sparse scattered hairs on an aciculatesurface. Mesoscutum smooth, polished, notaulices distinct, converging sharply towards scutellar suturetwo-thirds from scutellar suture, scutellar fovea large, smooth, polished, scutellar cup large, almost round,with a large central fovea, rim of cup with a few fovea around edges, scutellar disc rounded apically , surfacereticulate-rugose (cf. Fig. 139) with scattered hairs on apical margin; carinae of propodeum diverging atbase, with dense pubescence on either side; mesopleural suture distinct, metapleura smooth. Segment 1 ofgaster wider than long, short, crenulate, segment 2 the largest in lateral view, segments 3 and 4 partiallyvisible, finely punctate (Fig. 100), hypopygium with basal hairs. Legs orange, with moderately densepubescence, hind meta-tarsus as long as tarsal segments 2-4 combined, tibia with two apical spines, tarsalclaws simple. Wings pubescent on surface, apical margins with a hair fringe, radial cell of forewing closed, 2x as long as wide, Rs+M and M absent, 2rm longer than wide. Colour: antenna yellowish with dark apices,head and thorax blackish, gaster chestnut-brown. Cf . Antenna 15-segmented, segment 3 longer than 4, curved, flattened on outer margins (Fig. 99). MATERIAL EXAMINED Zaire: 1 $ (holotype), Urundi (Mosso), Makoronkwe, 1450 m, 12.iii.1953 (P. Basilewsky) (MRAC).Uganda: 1 cf (BMNH). Zaire: 12 $ (MRAC). Zimbabwe: 1 $ (BMNH). (All paratypes.) HEXACOLA Foerster Hexacola Foerster, 1869: 342. Type-species: Eucoela picicrus Giraud, by monotypy.Hexaplasta Foerster, 1869: 345. Type-species: Cothonaspis hexatoma Hartig, 1841: 357, by originaldesignation. [Synonymy by Rohwer & Pagan, 1917.] DIAGNOSIS. $ antenna generally 13-segmented but in some species 11-12 segmented, clavate, cf antenna15-segmented, segment 3 curved, longer than 4th (Fig. 39) ; wings pubescent, ciliate, radial cell of forewingopen on front margin (vein R\ projecting slightly on margin but not joining Rs 2 ), Rs+M and M (cubitus)indicated; scutellar cup elliptical, not extending to apex of disc, scutellar disc striate on lateral margins. DISCUSSION. Closely related to Kleidotoma but distinguished by the shape of the radial cell of the forewingand the absence of an incision on the apical margin of the wing. In Hexacola the apex of the forewing isnormally rounded, neither truncate nor incised. The scutellar disc is not as distinctly striated as inKleidotoma and the wings are generally narrow. DISTRIBUTION. Europe; South Africa; North and South America. BIOLOGY. The type-species of Hexaplasta is a known parasite of Oscinella frit (L), the frit fly (Imms, 1930;Kerrich & Quinlan, 1960). REMARKS. Nordlander (1981) notes that some earlier authors considered that the nominal speciesdesignated by Foerster as type-species of Hexacola was misidentified and that in fact the species wasKleidotoma hexatoma Thomson, 1862 (Kieffer, 1901 , 1902; Ashmead, 1903; Dalla Torre & Kieffer, 1910).Ashmead (1903) also designated K. hexatoma Thomson as type-species of Hexacola. Rohwer & Fagan(1917) concluded that Eucoela picicrus Giraud, 1860 is the type-species of Hexacola by original designationand monotypy. They made Hexaplasta Foerster a junior synonym of Hexacola, thereby acting as firstrevisers. Nordlander (1981) is submitting a case to the International Commission on Zoological Nomencla-ture to decide on the type-species designation, and states that he has compared a female in the Foerstercollection, labelled 'Hexacola picicrus Giraud', with syntypes of Kleidotoma hexatoma Thomson, andconsiders them to be conspecific. Key to the Afrotropical species of Hexacola Foerster Females 1 Antennaof$ 11-12 segmented, club 5-6 segmented, segment 3 as long as 4 +5 2 AFROTROPICAL EUCOILIDAE 273 Antenna of $ 13-segmented, club 4-8 segmented 5 2 Antenna of $ 11 -segmented, club 5-segmented, sharply defined, together distinctly longer than flagellar segments 1-6, 4-6 quadrate (Fig. 149) absensa sp. n. (p. 273) - Antenna of $ 12-segmented, club 5-6 segmented, flagellar segments longer than broad 3 3 Antennal club 5-segmented, as long as combined lengths of flagellar segments 1-7, segments 4-7 each slightly longer than wide (Fig. 150) ; mesopleural suture absent .... pallida sp. n. (p. 277) - Antennal club 6-segmented, much longer than combined lengths of flagellar segments 1-6, segments 4-6 as wide as long or longer than wide (Figs 151, 152); mesopleural suturecomplete 4 4 Antennal segments 4-6 longer than wide (Fig. 151) atropos sp. n. (p. 274) - Antennal segments 4-6 as wide as long (Fig. 152) zama sp. n. (p. 279) 5 Antennal club 4-5 segmented 6 - Antennal club with 6 or more segments 8 6 Antennal club 4-segmented (Fig. 153) fringa sp. n. (p. 276) Antennal club 5-segmented (Figs 154, 155) 7 Club segments together slightly shorter than flagellar segments 1-8 , 4-8 longer than broad (Fig . 154) quisnama sp. n. (p. 278) - Club segments together much longer than flagellar segments 1-8, as broad or broader than long (Fig. 155) quinqueclavata sp. n. (p. 278) 8 Antennal club 6-7 segmented (Figs 157, 60) 9 - Antennal club 8-segmented (Fig. 156). Segment 3 as long as 4+5, apex of tergite 2 of gaster and tergites 3-5 completely punctate octoclava sp. n. (p. 277) 9 Antennal club 6-segmented (Figs 157, 158) 10 - Antennal club 7-segmented (Fig. 160) 12 10 Antennal club sharply defined (Fig. 158), segments 4-7 of flagellum as wide as long, apex of tergite 2 and segments 3 and 4, when visible, punctate 11 - Antennal club not sharply defined, segments 4-7 of flagellum longer than wide, not compacted (Fig. 157); gaster impunctate hexatoma (Hartig)(p. 276) 11 Flagellar segments 47 of antenna wider than long, compacted, club segments together much longer than segments 1-7 combined (Fig. 158) compacta sp. n. (p. 275) Flagellar segments 4-7 of antenna as wide as long, not compacted, club segments together aslong as segments 1-7 (Fig. 159).Bicoloured; legs bright yellow; gaster punctate bifaria sp. n. (p. 275) 12 Antennal club sharply defined (Fig. 160) septemiussp. n. (p. 279) - Antennal club not sharply defined (Fig. 161). Segments 1-7 of flagellum bright yellow, segments 8-13 blackish brown, gaster im-punctate amantiasp. n.(p. 274) Males 1 Vertex of head striate; segment 3 of antenna the longest, strongly swollen distally, outer margins flattened medially (Fig. 39); radial cell of fore wing 2-5 x as long as wide, vein Rs 2 notextended along wing margin hexatoma Hartig (p. 276) - Vertex of head smooth, polished; segment 3 of antenna the longest, curved, weakly, swollen apically (Fig. 170); radial cell of forewing long and narrow, 3-5-4-0 x as long as broad, Rs 2sometimes extended along wing margin 2 2 Apex of tergite 2 of gaster and following segments punctate; radial cell almost as wide as long (1-25 x as long as wide), Rs 2 weakly extending along wing margin octoclava sp. n.(p. 277) - Gaster impunctate; radial cell elongate, Rs 2 extended along wing margin bifaria- and quinqueclavata-groups Hexacola absensa sp. n. (Figs 40, 149, 168)DESCRIPTION. $. Antenna 11 -segmented, with sharply defined 5-segmented club, the club longer than 274 J. QUINLAN flagellar segments 1-6, 4-6 quadrate (Fig. 149). Head, viewed frontally, a little longer than broad, smooth,shining, eyes almost round, further apart measured medially than height of an eye, malar groove distinct.Pronotal plate projected forward, anterior and posterior parts joined by narrow medial bridge, fovea oneither side open on lateral margins (Fig. 40); pronotum either side of pronotal plate with a tuft ofpubescence. Mesoscutum smooth, polished, notaulices absent, lateral bars of scutellum polished with afew striations apically; scutellar foveae wider than long, angular, scutellar cup long, narrow, polished,weakly excavate, not extending to apex of disc (cf. Fig. 168), scutellar disc with broken longitudinalstriations laterally, apex rounded; propodeal carinae obscured by pubescence. Mesopleura smooth,polished, suture complete [metapleura not entire due to pin on which the specimen is mounted]. Segment 1of gaster obscured by a ring of pubescence at base of tergite 2, not complete on dorsal surface, tergites 3 and4 visible in lateral view, punctate, hypopygium not prominent. Wings short, broad, surface pubescent,apical hair fringe sparse, radial cell of forewing partially open on wing margin but appears closed due topigmentation, vein RI not complete, apex of wing rounded. Legs short, robust, mid and hind coxae thick,femora swollen medially, tibiae broad apically, tarsi slender. Colour: club segments of antenna brownish,flagellar segments orange-brown, head, thorax and gaster dark brown, legs yellow.Cf unknown. MATERIAL EXAMINEDHolotype $, Zaire: Uele, Buta, 450 m, ll.iv.1935 (G. F. de Witte) (MRAC). REMARKS. Distinguished primarily from all other species in the genus by the 11 -segmented antenna. Hexacola amantia sp. n. (Figs 161, 162, 163, 167) DESCRIPTION. $. Antenna 13-segmented with a weakly defined, 7-segmented club, flagellar segments 3-6each longer than wide, 3 longer than 4 (Fig. 161). Head, viewed frontally, smooth, polished, with scatteredhairs, eyes oval, further apart measured medially than height of an eye, malar grooves distinct, mandibleswith scattered setae extending to supraclypeal area, anterior tentorial pits distinct, epistomal suturepresent. Pronotal plate projected forward, anterior and posterior parts connected by a broad medialbridge, lateral fovea either side of bridge open (Fig. 162). Pronotum either side of plate with a tuft of hairs.Mesoscutum smooth, polished, notaulices absent, lateral bars of scutellum smooth, polished, scutellarfovea narrow, angularly opposed to each other, smooth, polished, scutellar cup oval-elongate, lateralmargins of scutellar disc narrow, weakly striated (almost smooth in some specimens), apex of discreticulate-rugose, rounded (cf. Fig. 167), propodeal carinae subparallel, weakly bowed posteriorly atjuncture with nucha, lateral margins of propodeum densely pubescent. Mesopleura and metapleurasmooth, polished, metapleura with traces of ridges, mesopleural carina complete. Segment 1 of gasterobscured by a dense ring of hairs at base of tergite 2, complete on dorsal surface, tergite 2 occupying wholeof visible surface in lateral view, smooth, impunctate, hypopygium weakly protruding, ventral spine notvisible. Wing surface pubescent, apical margins rounded, with a long hair fringe, radial cell of forewingopen on margin, vein RI partially projected along margin. Legs normal, mid and hind coxae robust, 2 x aslong as broad, swollen basally, trochanters longer than broad, fore and mid femora longer than fore andmid tibiae, hind femora shorter than hind tibia, fore and mid-tibiae longer than fore and mid-tarsi, hindtibiae and tarsi subequal in length. Colour: antennal segments 1-7 orange-yellow, 8-13 dark brown, head,thorax and gaster blackish brown, legs yellow. Cf . Antenna 15-segmented, filiform, segment 3 the largest, strongly curved, surface flattened on outermargin (Fig. 163). MATERIAL EXAMINED Holotype $ , Nigeria: Samaru, em. 30.viii.1968, ex Oscinellinae pupa on Scoliophthalmus micantipennisDuda (guinea corn shoot) (J. C. Deeming) (BMNH). Paratypes. Nigeria: 5 $, 3 cf (BMNH). Uganda: 1 9 (BMNH). Zaire: 7 $, 2 cf (MRAC). REMARKS. Closely related to septemius but distinguished by the antennal shape (Fig. 161). Hexacola atropossp. n. (Figs 151, 164) DESCRIPTION. $ . Antenna 13-segmented with a sharply defined 6-segmented club (Fig. 151), the club muchlonger than flagellar segments 1-6, flagellar segments longer than wide. Head, viewed frontally, almost AFROTROPICAL EUCOILIDAE 275 rounded, eyes slightly further apart than height of an eye measured medially, face smooth, polished, with afew scattered setae, malar groove distinct, vertex polished. Prbnotal plate projected forward, anterior andposterior parts joined by a narrow medial bridge, fovea either side open laterally; pronotum either side ofpronotal plate with a few long setae. Mesoscutum smooth, polished, notaulices absent, lateral bars ofscutellum smooth, polished, scutellar foveae small, wider than long, polished, scutellar cup long, narrow,with a small apical fovea, scutellar disc longitudinally striate on lateral margins, apex rounded with a fewlong setae; propodeal carinae weakly bowed, sides of propodeum densely pubescent. Mesopleura andmetapleura smooth, polished, mesopleural carinae distinct. Segment 1 of gaster completely obscured bydense wool-like ring of pubescence at base of tergite 2, complete on dorsal surface, tergite 2 the largest inlateral view, tergites 3 and 4 partially visible, punctate, hypopygium short, broad, ventral spine short. Wingsurface densely pubescent, apical hair fringe long, radial cell of forewing partially closed on wing margin,apex of wing rounded (Fig. 164). Legs normal, mid and hind coxae swollen, femora swollen basally, tibiaebroad apically, tarsi longer than tibiae. Colour: antenna orange-brown, basal segments lighter, headbrownish, thorax orange-yellow, gaster brownish, legs pale yellow.CT unknown. MATERIAL EXAMINED Holotype <j>, Cameroun: Nkoemvon, i-ii.1980 (D. Jackson) (BMNH).Paratypes. Cameroun: 1 $ (BMNH). Zaire: 1 $ (MRAC). REMARKS. Distinguished from zama by the antenna (Fig. 151). Hexacola bifaria sp. n.(Figs 159, 165) DESCRIPTION. $. Antenna 13-segmented with distinct 6-segmented club, together longer than flagellarsegments 1-7, segments 4-7 each as broad as long (Fig. 159). Head, viewed frontally, smooth, polished,with a few hairs medially, anterior tentorial pits prominent, epistomal suture distinct, malar groovescomplete, cheeks weakly converging, eyes almost round, further apart measured medially than height ofan eye, mandibles bright yellow. Pronotal plate viewed dorsally, projected forward, anterior and posteriorparts connected medially by a broad bridge, fovea on each side open laterally, pronotum either side ofpronotal plate with a tuft of pubescence. Mesoscutum smooth, polished, notaulices absent; lateral bars ofscutellum smooth, polished, scutellar foveae shallow, polished, as wide as long, scutellar cup narrow,elliptical, scutellar disc striate on lateral margins, apex rounded with weak radiating striae; carinae ofpropodeum subparallel, converging towards nucha, lateral margins of propodeum densely pubescent,meso- and metapleura smooth, polished, mesopleural carinae complete, metapleura with one or two ridgesin upper region. Segment 1 of gaster obscured by ring of dense pubescence at base of tergite 2, complete ondorsal surface, tergites 2-4 visible in lateral view, apex of tergite 2 and visible parts of 3 and 4 punctate,tergite 2 the largest in lateral view, hypopygium short, broad, ventral spine as long as hypopygium. Wingsurface pubescent, apical margins rounded, with a hair fringe, radial cell of forewing open on wing margin,vein RI partially extended along margin. Legs normal, coxa a little longer than wide, swollen medially,trochanters as long as wide, front femora swollen medially, longer than tibiae, mid femora and tibiaesubequal in length, femora with usual medial swelling, hind femora shorter than hind tibiae, swollenbasally. Colour: club segments of antenna darker than basal flagellar segments, head, thorax and gasterdark chestnut-brown, legs pale yellow. Cf. Antenna 15-segmented, filiform, segment 3 the largest, bent or angled at base, the outer sideflattened (Fig. 165). MATERIAL EXAMINED Holotype $, Zimbabwe: Salisbury, Chishawasha, i.1979 (A. Watsham) (BMNH).Paratypes. Nigeria: 1 $ (BMNH). Zaire: 17 $, 2 cf (MRAC); 3 $ (BMNH). REMARKS. Closely similar to compacta but distinguished by the form of the antenna. Hexacola compacta sp. n. (Figs 158, 166) DESCRIPTION. $. Antenna 13-segmented, with sharply defined 6-segmented club, the club 2 x as long asflagellar segments 1-7, segment 3 subequal in length to 4+5, segments 4-7 each broader than long, segment1 of club (8th flagellar segment) as long as segments 4-7 combined and much wider (Fig. 158). Head,viewed frontally, smooth, shining, with scattered hairs in supraclypeal area and on mandibles, eyes 276 J. QUINLAN semi-circular, further apart measured medially than height of an eye, anterior tentorial pits distinct, malargroove percurrent, cheeks converging, vertex polished. Pronotal plate projected forward, anterior andposterior parts connected medially by a narrow bridge, fovea on either side open laterally (Fig. 166).Pronotum on lateral margins of plate with tufts of pubescence. Mesoscutum smooth, polished, with trace ofnotaulices, lateral bars of scutellum smooth, polished, scutellar foveae as wide as deep, polished, scutellarcup longer than broad, outer rim paler than middle, apex of cup with lateral fovea, scutellar discreticulate-rugose, not striated, apex rounded. Meso- and metapleura smooth, polished, mesopleuralcarinae complete, metapleura without ridges. Propodeal carinae parallel, lateral margins of propodeumpubescent. Segment 1 of gaster obscured by a dense ring of pubescence at base of tergite 2, complete ondorsal surface, tergites 2-3 visible in lateral view, tergite 2 the largest in lateral view, visible part of tergite 3punctate, hypopygium small, ventral spine weakly produced. Wing surface pubescent, apex of wing broadwith apical hair fringe, radial cell of forewing partially open on wing margin, i.e. vein RI weakly producedalong margin, not closing cell. Legs short, robust, hind coxa 2 x as long as wide, swollen basally,trochanters longer than broad, femora swollen basally, shorter than tibiae, tibiae long, narrow basally,wide apically, tibiae and tarsi subequal in length. Colour: antenna dark brown, head brownish, gaster andthorax chestnut-brown, legs yellow-orange.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: 25.viii.1951 (J. Verschureri) (MRAC).Paratypes. Zaire: 2 $ (MRAC). REMARKS. Distinguished from other closely related species by the bright yellow legs. Hexacola fringa sp. n. (Figs 153, 164, 167) DESCRIPTION. $. Antenna 13-segmented with weakly defined, 4-segmented club, the total length of clubless than combined lengths of flagellar segments 1-9 which are each longer than broad (Fig. 153). Head,viewed frontally, as broad as long, eyes round, weakly converging towards clypeus, further apart measuredmedially than height of an eye, face smooth, polished, with a few sparse hairs. Pronotal plate projectedforward, anterior and posterior parts joined by a medial bridge, fovea on either side open laterally,pronotum on either side of pronotal plate with a tuft of pubescence. Mesoscutum smooth, polished,notaulices absent, in their place a row of hair bases, lateral bars of scutellum short, broad basally, polished,scutellar foveae small, almost round, scutellar cup long, narrow, not well defined, scutellar disc withaberrant sculpture, apex rounded (Fig. 167); propodeal carinae parallel, weakly converging basally, sidesof propodeum densely pubescent. Mesopleura and metapleura smooth, polished, mesopleural carinaecomplete, metapleura ridged near posterior margin. Segment 1 of gaster obscured by a ring of densepubescence at base of tergite 2 which is complete on dorsal surface, tergites 34 visible in lateral view,impunctate, tergite 2 the largest visible tergite, hypopygium not pronounced. Wings densely pubescent,apical fringe long, wing margin at apex not sharply rounded, radial cell of forewing open on wing margin,vein RI weakly projected along margin (cf. Fig. 164). Legs with mid and hind coxae short, front femora thelargest, swollen, mid and hind femora short, swollen, tibiae short, broad apically, front tarsi longer thanfront tibiae, mid and hind tibiae subequal in length to their tarsi. Colour: antenna yellow basally, clubsegments darker, head dark brown, thorax and gaster dark chestnut-brown, legs pale yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zimbabwe: Salisbury, Chishawasha, i.1979 (A. Watsham) (BMNH).Paratypes. Zimbabwe: 3 9 same data as holotype (BMNH). REMARKS. Separated primarily from quisnama by the antennal structure (Fig. 153). Hexacola hexatoma (Hartig)(Figs 157, 168, 169) Cothonospis hexatoma Hartig, 1841: 357. Lectotype $. GERMANY, designated by Quinlan in Kerrich &Quinlan, 1960: 191 [examined]. DESCRIPTION. $. Antenna 13-segmented, subclavate, with weakly defined 6-segmented club, flagellarsegments 1-7 each longer than broad (Fig. 157). Head, viewed frontally, smooth, polished, with a few AFROTROPICAL EUCOILIDAE 277 scattered hairs, particularly in region of mandibles, anterior tentorial pits indicated, malar groovescomplete, eyes further apart measured medially than height of an eye, cheeks weakly converging, vertexstrigose. Pronotal plate projected forward, anterior and posterior parts connected by a medial bridge,fovea on either side of pronotal plate open laterally. Pronotum either side of pronotal plate with a tuft ofpubescence. Mesoscutum smooth, polished, with scattered setae, notaulices absent, lateral bars ofscutellum polished, scutellar cup narrow, elliptical with a fovea on apical half, scutellar disc with fineaberrant striate sculpture (Fig. 168) , scutellar fovea polished, wide, narrow, apex of scutellar disc rounded.Mesopleura and metapleura polished, mesopleural carina complete, lateral margins of pronotum denselypubescent, propodeal carinae subparallel. Segment 1 of gaster obscured by dense ring of pubescence atbase of tergite 2, complete on dorsal surface, tergites 2-4 visible in lateral view, impunctate, tergite 2 thelargest, hypopygium broad. Wing surface pubescent with apical fringe of hairs, radial cell of forewing openon wing margin, vein /?i partially projected along wing margin. Legs long, slender, mid and hind coxaepubescent. Colour: antenna, head, thorax and gaster brownish black. Cf . Antenna 15-segmented, filiform, segment 3 curved, swollen at apex, flattened on outer margin (Fig.169). MATERIAL EXAMINEDLectotype $, Germany.England: 30 $, 28 cf (BMNH). Uganda: 1 $, 1 cf (BMNH). REMARKS. Similar to compacta and bifaria but separated initially by the antennal club and the ratio of thebasal flagellar segments, and by the impunctate gaster. Hexacola octoclava sp. n. (Figs 156, 170) DESCRIPTION. $. Antenna 13-segmented with a sharply defined 8-segmented club, much longer thanflagellar segments 1-5 combined, segment 3 as long as 4+5 (Fig. 156). Face, viewed frontally, wider thanlong, eyes converging, slightly further apart than height of an eye measured medially, face smooth,polished, with sparse setae in supraclypeal area and on mandibles, malar grooves distinct. Pronotal platedistinct, projected forward, anterior and posterior parts connected by a medial bridge, fovea on either sideopen on lateral margins; pronotum either side of pronotal plate with tufts of pubescence. Mesoscutumsmooth, polished, notaulices absent, lateral bars of scutellum polished, scutellar foveae kidney-shaped,scutellar cup narrow, elliptical, not extending to apex of scutellar disc, lateral margins of disc with brokenstriate sculpture laterally, reticulate-rugose basally; propodeal carinae subparallel, lateral margins pubes-cent. Mesopleural suture complete, metapleura smooth, polished. Segment 1 of gaster obscured by a densering of pubescence at base of tergite 2. Tergites 2-5 visible in lateral view, tergite 2 the largest, apex oftergite 2 and visible parts of 3-5 densely punctate, hypopygium slender, ventral spine protruding. Wingsurface pubescent, apical margins rounded, with a fringe of hairs, radial cell of forewing open on margin,vein RI partially extended along margin of wing. Legs slender, mid and hind coxae elongate, swollenmedially, trochanters longer than wide, femora short and swollen, tibiae and tarsi long, slender, subequalin length. Colour: antenna brownish apically, basal segments lighter, head, thorax and gaster darkchestnut-brown, legs yellow. Cf . Antenna 15-segmented, segment 3 curved, swollen apically, flattened on outer margin (Fig. 170). MATERIAL EXAMINED Holotype 9, Zaire: 18. ix. 1952 (H. deSaeger) (MRAC).Paratypes. Zaire: 3 $, 3 cf (MRAC). REMARKS. This species is distinguished from all others described here by the 8-segmented club of theantenna. Hexacola pallida sp. n. (Figs 148, 150, 177) DESCRIPTION. . Antenna 13-segmented, with a sharply defined 5-segmented club, the combined lengths ofclub segments little longer than flagellar segments 1-7 (Fig. 150). Head, viewed frontally, almost spherical,eyes rounded, further apart measured medially than height of an eye, face smooth, polished, with a fewscattered setae, malar groove distinct. Pronotal plate projected forwards, anterior and posterior partsconnected by a narrow medial bridge, foveae opon on lateral margins (cf. Fig. 148); pronotum either side 278 J. QUINLAN of plate with tufts of pubescence. Mesoscutum smooth, polished, notaulices absent, lateral bars ofscutellum short, broad basally, polished, smooth, scutellar foveae shallow, polished, as wide as long,scutellar disc rounded apically, surface with aberrant longitudinal striate sculpture, scutellar cup long,narrow, not extending to apex of disc; propodeal carinae weakly converging towards apex, denselypubescent on lateral margins. Mesopleural suture absent, metapleura smooth, polished. Segment 1 ofgaster completely obscured by a ring of dense pubescence at base of tergite 2, sparse on dorsal surface,tergite 3 visible in lateral view, impunctate, hypopygium short, broad. Wings long, narrow, surfacepubescent, with long apical hair fringe (Fig. 171), radial cell open on margin, apex of wing not acutelyrounded. Legs short, robust, tarsi narrow, longer than tibiae. Colour: antenna yellowish, head, thorax,and gaster orange-yellow, legs pale yellow.C? unknown. MATERIAL EXAMINED Holotype $, Zaire: Eala, ix.1935 (/. Ghequiere) (MRAC).Paratypes. Zaire: 2 ty, same data as holotype (MRAC). REMARKS. This species is distinguished from others by the absence of a mesopleural suture, the narrowwings and the shape of the radial cell (Fig. 171). Hexacola quinqueclavata sp. n. (Fig. 155) DESCRIPTION. $. Antenna 13-segmented with sharply defined 5-segmented club 1-3 x length of flagellarsegments 1-8, segment 3 subequal in length to 4+5, segments 4-8 as broad as long (Fig. 155). Face, viewedfrontally, very slightly longer than wide, eyes almost circular, further apart measured medially than heightof an eye, face smooth, polished, with a few sparse hairs, clypeus with sparse hairs, malar groove distinct.Pronotal plate projected forward, anterior and posterior parts connected medially by a bridge, fovea eitherside open laterally; pronotum either side of plate with tufts of pubescence. Mesoscutum smooth, polished,notaulices absent, lateral bars of scutellum smooth, polished, short, broad basally, scutellar foveae widerthan long, polished, scutellar disc with weak broken longitudinal striations, apex weakly conical,reticulate-rugose, scutellar cup small, longer than wide, not extending to apex of scutellar disc; propodealcarinae bowed medially, lateral margins of propodeum densely pubescent, mesopleura smooth, polished,metapleura pubescent on ventral margin, mesopleural suture complete. Segment 1 of gaster obscured by adense ring of pubescence at base of tergite 2, complete on dorsal surface, gaster impunctate, hypopygiumnot projecting. Wing surface pubescent, with apical hair fringe, apex of wing broad, rounded, radial cell offorewing open on margin, vein RI weakly extending along margin. Legs of normal proportions, mid andhind coxae swollen, longer than broad, femora short, swollen, slightly shorter than tibiae, tarsi a littlelonger than tibiae. Colour: antenna yellowish basally, club segments darker, head, thorax and gaster darkbrownish to chestnut-red, legs pale yellow.C? unknown. MATERIAL EXAMINED Holotype $, Zimbabwe: Salisbury, Chishawasha, xii.1978 (A. Watshani) (BMNH). Paratypes. Cameroun: 3 $ (BMNH). Madagascar: 3 $ (BMNH). Nigeria: 1 $ (BMNH). Zaire: 14 $(MRAC). REMARKS. This species is close to quisnama but can be separated by the antenna, the club segments beingmuch longer than the basal flagellar segments (Fig. 155). The specimen from Nigeria is assigned to thisspecies but is extremely light coloured and has a more prominent hypopygium. Hexacola quisnama sp. n. (Figs 154, 172, 173) DESCRIPTION. $. Antenna 13-segmented with sharply defined, 5-segmented club, slightly shorter thanflagellar segments 1-8, segment 3 as long as 4+5, 4-8 each a little longer than wide (Fig. 154). Head, viewedfrontally, a little longer than wide, eyes semi-circular, further apart than height of an eye measuredmedially, weakly converging towards clypeus, vertex polished. Pronotal plate produced forward, anteriorand posterior connected by a medial bridge, the fovea open on lateral margins, pronotum either side ofpronotal plate with tufts of pubescence. Mesoscutum smooth, polished, notaulices absent, lateral bars ofscutellum polished, scutellar foveae kidney-shaped, smooth, polished, scutellar cup small, little longer AFROTROPICAL EUCOILIDAE 279 than broad, not extending to apex of disc, scutellar disc with longitudinally radiating striae on lateralmargins, coriaceous at apex, apex rounded (Fig. 173); propodeal carinae parallel, converging at base,densely woolly on lateral margins. Mesopleura and metapleura smooth, polished, mesopleural carinaedistinct. Segment 1 of gaster obscured by dense ring of pubescence at base of tergite 2, complete on dorsalsurface, tergites 2-4 visible in lateral view, tergite 2 the largest visible segment, apex of 3-5 punctate,hypopygium short, broad basally, ventral spine short. Wings densely pubescent with long apical hairfringe, fore wing short, broad apically, radial cell of forewing open on margin, vein RI weakly indicated inpart (Fig. 172). Legs normal except trochanters long, narrow, mid and hind coxae swollen basally. Colour:antenna yellowish brown basally, club segments darker, head brownish, thorax orange-yellow, gasterchestnut-brown, legs yellow.O" unknown. MATERIAL EXAMINED Holotype $, Zaire: Massif Ruwenzori, riv. Kakalari, affl. Bombi, 1,725 m, 10. iv. 1954 (R. Vanschuyt-broeck & H. Synave) (MRAC). Paratypes. South Africa: 3 $ (BMNH). Uganda: 1 $ (BMNH). Zaire: 11 $ (MRAC). REMARKS. Distinguished from quinqueclavata by the antennal segment ratios (Fig. 154). Hexacola septemius sp. n. (Fig. 160) DESCRIPTION. $. Antenna 13-segmented with sharply defined 7-segmented club, longer than flagellarsegments 1-6 combined, segment 3 equal in length to 4+5 (Fig. 160). Head, viewed frontally, smooth,polished, with one or two hairs above supraclypeal area, cheeks converging, eyes further apart than heightof an eye measured medially between eyes, malar grooves distinct. Pronotal plate projected forward,anterior and posterior parts connected by a narrow medial bridge, fovea on either side open laterally,lateral margins of pronotum on either side of plate without tufts of pubescence, at most a few setae present.Mesoscutum smooth, polished, without trace of notaulices, lateral bars of scutellum polished, scutellarfovea kidney-shaped, wider than long, scutellar cup long, narrow, polished, not extending to apex of disc,lateral margins of disc with broken longitudinal striations, basally reticulate-rugose, rounded apically.Propodeal carinae parallel, pubescent on lateral margins. Mesopleural suture complete, mesopleura andmetapleura smooth, polished. Segment 1 of gaster obscured by a ring of dense pubescence at base of tergite2, ring complete on dorsal surface, tergites 2-4 visible in lateral view, tergite 2 the largest, apex of tergite 2and visible parts of tergites 3+4 punctate, hypopygium not prominent, ventral spine short. Wing surfacepubescent, apical margins with a fringe of hairs, apex rounded, radial cell of forewing open on margin, veinRI projected partially along margin. Colour: antenna yellow basally, club segments darker, head, thoraxand gaster chestnut-brown, legs yellow.C? unknown. MATERIAL EXAMINED Holotype $, Zaire: 10.vii.1952 (H. deSaeger) (MRAC).Paratypes. Zaire: 2 $ (MRAC). REMARKS. Distinguished from amantia, a closely similar species, by the sharply defined club of the antenna(Fig. 160). Hexacola zama sp. n. (Figs 148, 152, 174) DESCRIPTION. $. Antenna 12-segmented with a sharply defined, 6-segmented club, the club much longerthan flagellar segments 1-6, flagellar segments 4-6 as wide as long (Fig. 152). Head, viewed frontally,smooth, polished, supraclypeal area and mandibles with a few scattered hairs, eyes almost spherical,slightly further apart measured medially, than height of an eye, anterior tentorial pits weakly indicated,malar grooves complete, cheeks weakly converging, vertex polished. Pronotal plate projected forward,anterior and posterior parts connected by a narrow medial bridge, fovea on either side open laterally (cf.Fig. 148). Pronotum either side of plate with a few hairs. Mesoscutum smooth, polished, notaulices absent,lateral bars of scutellum striate dorsally, polished laterally, scutellar fovea kidney-shaped, polished,scutellar cup elliptical, longer than wide, the rim paler than inner surfaces, apical quarter with a fovea,lateral margins of scutellar disc with broken reticulate-rugose sculpture appearing striated (Fig. 174), apex 280 J. QUINLAN of disc rounded, surface reticulate-rugose. Propodeal carinae subparallel. Mesopleural carinae complete,meso- and metapleura smooth, polished. Segment 1 of gaster obscured by a ring of dense pubescence atbase of tergite 2, complete on dorsal surface, tergites 3+4 partially visible, weakly punctate, hypopygiumshort, broad basally, ventral spine not visible. Wing surface pubescent, apical margins with a fringe ofhairs, radial cell of forewing open on margin, vein RI extending partially along margin. Legs normal.Colour: antenna brownish yellow, head, thorax and gaster chestnut-brown, legs yellow.Cf unknown. MATERIAL EXAMINEDHolotype $, Zaire: N. Lac, Kivu, Rwanki, xii.1951 (/. V. Leroy) (MRAC). REMARKS. This species is separated from atropos by the relative lengths of the antennal segments. KLEIDOTOMA Westwood Kleidotoma Westwood, 1833: 494. Type-species: Kleidotoma psiloides Westwood, by monotypy. Aphyoptera Foerster, 1869: 343. Type-species: Aphyoptera inustipennis Foerster, by monotypy. [Synony-mised by Hellen, 1960: 23.] Aphiloptera Foerster, 1869: 351. Type-species: Aphiloptera anisomera Foerster, by monotypy. [Synony-mised by Quinlan, 1967: 1.] Agroscopa Foerster, 1869: 352. Type-species: Agroscopa helgolandica Foerster, by monotypy. [Synony-mised by Quinlan, 1967: 1.] Nedinoptera Foerster, 1869: 350. Type-species: Nedinoptera halophila Thomson, by monotypy. [Synony-mised by Quinlan, 1967: 1.] Rhynchacis Foerster, 1869: 349. Type-species: Cothonaspis nigra Hartig, by monotypy. [Synonymised byQuinlan, 1978: 25.] Pentacrita Foerster, 1869: 349. Type-species: Cothonaspis retusa Hartig, by monotypy. [Synonymised byWeld, 1952: 210.] Tetrahoptra Foerster, 1869: 349. Type-species: Clidotoma heterotoma Thomson, by subsequent designa-tion of Ashmead 1903: 62. [Synonymised by Weld, 1952: 207.] Tetratoma Cameron, 1890 preocc. Type-species: Kleidotoma heterotoma Thomson, by subsequent de-signation of Rohwer & Pagan, 1917: 376. Schizosema Kieffer, 1901: 161. Type-species: Cothonaspis emarginatus Hartig, by subsequent designationof Ashmead, 1903: 62. [Synonymised by Weld, 1931: 233.] DIAGNOSIS. $ antenna 13-segmented, clavate, with 3-segmented club inpsiloides though a variable numberof club segments are found within the genus, C? antenna 15-segmented, filiform, segment 3 the largest,curved, swollen at apex. Head, viewed frontally, longer than broad, eyes converging towards malar region,face smooth, polished, with long sparse hairs either side of clypeus, vertex striate. Pronotal plate producedforwards, anterior and posterior parts connected by a medial bridge, foveae on either side of bridge openon lateral margins (Fig. 183). Mesoscutum smooth dorsally, without trace of notaulices; mesopleurasmooth, polished, suture complete; scutellar fovea large, deep, smooth, scutellar disc longitudinallystriate, scutellar cup long, elliptical, not extending to apex of rounded disc (disc in some species conical orbeak-shaped). Segment 1 of gaster obscured by a ring of pubescence at base of tergite 2, not complete ondorsal surface (Fig. 36). Segment 2 the largest in lateral view, segments 3-5 smaller, not always visible,hypopygium with long apical hairs. Wing surface pubescent, radial cell of forewing (Fig. 35), completelyopen on wing margin, apex of wing either incised (Fig. 177) or truncate (Fig. 196). Legs of normal size andshape. Colour: antenna yellowish, head, thorax and gaster brownish black, gaster chestnut-red, legsyellowish. DISTRIBUTION. Old and New World. REMARKS. At present keys to males are difficult to construct. In separating females emphasis has been givento the number of antennal club segments and the relative lengths of the basal flagellar segments. In somespecimens the wings can be brachypterous to almost absent, particularly in the males. K. psiloides has notbeen recognised as occurring in the Afrotropical region. Key to the Afrotropical species of Kleidotoma Westwood Females 1 Antenna with 2-segmented club (Fig. 179) - Antennal club with 3 or more segments 3 AFROTROPICAL EUCOILIDAE 281 2 Antennal segment 2 as long as 3; apex of wing with wedge-shaped incision (Fig. 179); scutellar disc polished, cup long, narrow, not extending to apex of scutellum (Fig. 176); gasterimpunctate morsum sp. n. (p. 286) - Antennal segment 2 shorter than 3; apex of wing strongly arcuate (Fig. 208); scutellar disc striate, scutellar cup long, narrow, extending to apex of disc (Fig. 180); apex of tergite 2 andwhole of tergite 3 strongly punctate (cf. Fig. 201) nigrans sp. n. (p. 286) 3 Antenna with 3-segmented club 4 - Antennal club with 4 or more segments 10 4 Antennal segments 4-10 longer than wide (Fig. 182) ; scutellar disc either normally rounded at apex or truncate 5 - Antennal segments 4-10 of varying length, some apical segments, excluding club segments, only as wide as long (Fig. 200); scutellar disc either rounded or spine-shaped apically (Figs187,205) 8 5 Pronotum laterally and occiput very strongly, densely strigose (Fig. 181); antennal segments 4-10 each progressively shorter, longer than wide, wider apically than basally (Fig. 182);apex of tergite 2 with a broad band of punctures (cf. Fig. 201) ventosus sp. n. (p. 288) - Pronotum either smooth and polished , occiput not strongly strigose , or antenna not as above ... 6 6 Antennal segments 4-10 at least more than twice as long as wide, filiform, segment 3 longer than 4+5 (Fig. 188) ; apex of wing arcuate (Fig. 202) distenda sp. n. (p. 283) - Antennal segments 4-8 at most twice as long as wide, 9-10 not twice as long as wide, segments wider apically than basally 7 7 Occiput finely strigose, including area between ocelli (Fig. 183); sides of pronotum striated; tergites 2-4 of gaster visible in lateral view (Fig. 184); scutellar disc with strong and closelongitudinal striations, apex of disc almost square, scutellar cup extending to apex of disc(Fig. 185) ereftussp. n.(p. 284) - Occiput smooth , polished ; tergites 2-3 of gaster visible in lateral view ; scutellar disc with weak , widely spaced longitudinal striations, apex of disc rounded; scutellar cup not extending to apex of disc (Fig. 186) arbitra sp. n.(p. 282) 8 Scutellar disc strongly produced apically to form a long, blunt, truncated spine. Scutellar cup elliptical, lateral margins of scutellar disc striate; head strongly sculptured infrontal region, occiput strigose; sides of pronotal plate and mesoscutum with striatesculpture conica sp. n. (p. 283) - Scutellar disc rounded apically, not produced to form a spine 9 9 Antennal segments 5-8 almost quadrate (Fig. 189), same width apically as basally strigosa sp. n. (p. 287) - Antennal segments 4-6 longer than wide (Fig. 190), 7-10 as wide as long, conical, wider apically than basally favus sp. n. (p. 285) 10 Antenna with 4-segmented club 11 - Antennal club with 5 or more segments 12 11 Antennal club sharply defined (Fig. 191); apex of segment 2 and whole of visible parts of segments 3 and 4 of gaster densely punctate (cf. Fig. 184); hypopygium strongly protruding nitidiuscula sp. n.(p. 287)Antennal club not sharply defined, almost filiform (Fig. 192); gaster weakly punctate; hypopygium not protruding fimbriata sp. n. (p. 285) 12 Antenna with 5-segmented club, first segment of club distinctly smaller than four apical segments (Fig. 193) eala sp. n.(p. 283) - Antennal club with 6 or more segments 13 13 Antennal club 6-segmented 14 - Antennal club 7-segmented, not sharply defined, almost filiform (Fig. 195). Scutellar disc normally rounded apically, polished, with very weak striations (Fig. 194);apex of wing truncate (Fig. 196) norma sp. n.(p. 287) 14 Scutellar disc bifurcate at apex (Fig. 197), strongly striated laterally, apex of wing sharply incised (Fig. 198) bifurcata sp. n.(p. 282) - Scutellar disc normally rounded at apex, polished, with weak striations laterally; apex of wing weakly arcuate (cf. Fig. 202) elongulasp. n. (p. 284) 282 J. QUINLAN Kleidotoma arbitra sp. n. (Figs 175, 186, 204) DESCRIPTION. 9.. Antenna 13-segmented with a distinct 3-segmented club, club segments with rhinaria,each wider and longer than preceding flagellar segments, segment 3 subequal in length to 4+5, segments4-10 each longer than broad (Fig. 204). Head, viewed frontally, smooth, polished, with a few scatteredhairs medially, eyes small, round, further apart measured medially than height of an eye, malar groovecomplete, anterior and tentorial pits distinct, occiput smooth, shining, pronotal plate projected forward,anterior and posterior parts connected by a medial bridge, foveae either side of bridge open on lateralmargins. Pronotum either side of pronotal plate polished; mesopleura smooth, polished, suture distinct,mesoscutum smooth, polished, notaulices absent, scutellar disc striate on lateral margins, apex of discrounded, scutellar cup long, elliptical, not extending to apex of disc, apex of cup with a large fovea, lateralbars of scutellum smooth on dorsal surface, striate laterally, scutellar foveae large, smooth, shallow,separated by narrow septum of cup (Fig. 186). Segment 1 of gaster obscured by a dense ring of hairs at baseof tergite 2, not complete on dorsal surface, apex of tergite 2 smooth, polished, without punctures, tergite 3visible laterally (Fig. 175). Wings pubescent on surface, radial cell of forewing completely open on margin,vein Rs 2 extended along wing margin, cell triangular, apex of wing arcuate with a long apical hair fringe.Colour: segments 1-10 of antenna dark yellow, 11-13 brownish yellow, head, thorax and gaster darkchestnut-red, coxae yellow with brownish apices, basal half of femora light brown, apex yellow, tibiae andtarsi dark yellow.Cf unknown. MATERIAL EXAMINED Holotype 9. , South Africa: Mossel Bay, Cape Province, x.1921 (R. E. Turner) (BMNH). Paratypes. South Africa: 6 $ (BMNH). Uganda: 1 $ (BMNH). Zaire: 19 9 (MRAC). Zimbabwe: 8 9(BMNH). REMARKS. This species resembles psiloides, the Palaearctic species associated with Leptocera manictaRichards, but can be separated by the polished occiput. Kleidotoma bifurcata sp. n. (Figs 197, 198, 199, 201) DESCRIPTION. 9.. Antenna 13-segmented, clavate with a 6-segmented club, not sharply indicated, clubsegments with rhinaria, segment 3 longer than 4, 4-7 each progressively shorter than each other, clubsegments 8-13 subequal, all longer than segment 7. Head, viewed frontally, with coriaceous sculpture oninner orbits, eyes twice as far apart measured medially than height of an eye, striations extending fromlower margin of eye towards clypeal area, malar grooves obscured by striations, occiput strongly strigose.Pronotal plate, viewed dorsally, sculptured on anterior surface, polished on posterior surface, medialfoveae open on lateral margins (Fig. 199). Pronotum either side of pronotal plate striated. Mesoscutumfinely coriaceous except for a polished area where parallel lines would be, if present, lateral bars ofscutellum strongly striated, scutellar disc strongly striate, apex of disc truncated, with a blunt spine oneither side (Fig. 197), scutellar cup long, elliptical, bulbous apically, with large central fovea, scutellarfoveae large, deep, polished. Mesopleura smooth, polished, suture pronounced (carina-like on lowermargin, suture-like on upper margin), with coriaceous sculpture above and below, area below precoxalcarina pubescent. Lateral margins of propodeum pubescent. Segment 1 of gaster obscured by a dense ringof pubescence, complete on dorsal surface, apex of tergites 2 and 3 with dense punctures, hypopygiumpronounced (cf. Fig. 201). Wing surface with dotted hair bases, with apical hair fringe, apex of wingincised, radial cell of forewing open on margin (Fig. 198). Colour: antenna yellowish brown nasally, clubblackish brown, head and thorax blackish, gaster blackish brown dorsally, reddish yellow laterally, legsyellow brown.O" unknown. MATERIAL EXAMINED Holotype 9., Zaire: Rweru (Vole. Mikeno), 2,400 m (Bambous), 26-27.vii.1934 (G. F. de Witte)(MRAC). Paratype. 1 9. , same data as holotype except 3.vii.l934 (BMNH). REMARKS. This species cannot be confused with any other described in this paper. Apart from the shape ofthe head, the exceedingly distinctive shape of the entire scutellar area (Fig. 197) singles it out from all otherspecies. AFROTROPICAL EUCOILIDAE 283 Kleidotoma conica sp. n. (Figs 187, 200, 201) DESCRIPTION. $. Antenna 13-segmented with the three apical segments forming a distinct club (Fig. 200),segments 4-10 each progressively shorter than preceding segment, apical width of each wider than basalbreadth. Head, viewed frontally, strongly striated in frontal area below ocellus, malar grooves complete,clypeus with long scattered hairs, occiput strongly striated. Pronotal plate with lateral fovea open, striated.Pronotum and side margins of mesoscutum striated. Scutellar disc tapering to a blunt conical beak at apex,striate laterally, reticulate apically, lateral bars of scutellum striated, scutellar foveae large, shallow,polished, scutellar cup obscure (Fig. 187). Mesopleural suture distinct, with tuft of pubescence onantero ventral cavity. Gaster lenticular in shape, segment 1 obscured by a dense ring of pubescence at baseof tergite 2, apex of tergite 3 with scattered punctures, 4 punctate, hypopygium short (Fig. 201). Wingsurface with dotted hair bases, apical margins arcuate, with a hair fringe, radial cell of fore wing open onmargin, Rs 2 weakly extended along margin. Colour: antenna and legs dark yellow, first antennal segmentand coxae dark brown, head and thorax black, gaster dark chestnut-red. MATERIAL EXAMINED Holotype $, Zaire: 2.L1952 (H. de Saeger) (MRAC).Paratype. Zaire: 1 $ (MRAC). REMARKS. A very distinctive species. The very long, truncated, spine-like projection of the scutellar discand the strongly striated upper regions of the face distinguish it from all known species. The radial cell ofthe forewing is typical of Kleidotoma in shape, and the apex of the wing is weakly arcuate. Kleidotoma distenda sp. n. (Figs 188, 199, 202) DESCRIPTION. $ . Antenna 13-segmented with apical three segments forming a distinct club, segments 3-10at least 3 x as long as broad (Fig. 188). Head, viewed frontally, smooth, polished, malar grooves distinct,eyes oval, further apart measured medially than height of an eye, occiput very weakly sculptured; pronotalplate weakly sculptured with a large fovea on either side of medial bridge, open laterally (cf. Fig. 199).Pronotum either side of pronotal plate smooth. Mesopleura smooth, shining, suture distinct; scutellar discwith weak broken sculpture to fine striations on lateral margins, apex rounded, weakly sculptured,scutellar cup long, narrow, not reaching apex of disc, scutellar foveae large, shining, separated by a wideseptum at base of cup, lateral bars of scutellum smooth, shining. Segment 1 of gaster obscured by amoderately hairy ring at base of tergite 2, gaster impunctate. Wings densely pubescent, arcuate apically,with fringe of hairs, radial cell of forewing completely open on margin, cell triangular (Fig. 202). Colour:segments 1-10 of antenna brownish yellow, 10-13 brown, head dark brown, thorax and gaster chestnut-brown, legs yellow, hind coxa with longitudinal sculpture of varying degree.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Mossel Bay, Cape Province, 5-31.vii.1921 (R. E. Turner) (BMNH).Paratypes. Nigeria: 6 $ (BMNH). South Africa: 11 $ (BMNH). Zaire: 28 $ (MRAC). REMARKS. Distinguished from arbitra by antennal segments 4-10, each being twice as long as wide. Kleidotoma eala sp. n. (Figs 193, 194) DESCRIPTION. $. Antenna 13-segmented with a distinct 5-segmented club, segment 2 shorter than 3 butbroader than 3, segment 3 subequal to 4+5 in length, 4 shorter than 5, 5-7 subequal, club segments 9-13distinctly wider than segments 3-8 (Fig. 193). Pronotal plate small with a large fovea on either side ofmedial bridge, open laterally; pronotum either side of pronotal plate with tufts of pubescence, mesoscutumsmooth, polished, scutellar disc weakly conical at apex, lateral margins either side of scutellar cup weaklystriate, scutellar cup long, narrow, elliptical, scutellar foveae large, shallow (cf. Fig. 194), lateral bars ofscutellum smooth, polished, extending past centre of cup. Mesopleura smooth, polished, suture distinct;propodeum with tufts of pubescence on lateral margins. Segment 1 of gaster obscured by dense ring ofpubescence at base of tergite 2, not complete on dorsal surface, 2-4 visible in lateral view, punctate, 284 J. QUINLAN hypopygium clearly visible in lateral view, not extending past apex of sheath. Wings pubescent on surface,apical margin weakly arcuate, with fringe of hairs, radial cell of forewing completely open on wing margin.Colour: antenna pale yellow, head, thorax and gaster dark brownish red, legs pale yellow.Cf unknown. MATERIAL EXAMINED Holotype <J>, Zaire: Kivu, Kalondalac Ndaraga Mokotol, 1,750 m, 22-27. iii. 1934 (G. F. de Witte)(MR AC). REMARKS. Distinguished from other species by the 5-segmented club. Kleidotoma elongula sp. n. (Figs 180, 203, 207) DESCRIPTION. $. Antenna 13-segmented, the apical 6 segments with rhinaria and forming a weak club,segment 2 shorter and broader than 3, 3 longer than 4, 4-7 subequal in length and breadth, cylindrical, 8-13each longer than 7, wider medially than at either end (Fig. 203). Head, viewed frontally, smooth, polished,eyes further apart measured medially, than height of an eye, frontal area with scattered hairs, malargrooves percurrent, anterior tentorial pits distinct, occiput smooth. Pronotal plate small, with foveae oneither side of medial bridge open (cf. Fig. 207). Mesoscutum smooth, polished; scutellar disc rounded atapex, very weakly striated on lateral margins, scutellar cup long, elliptical, with a large fovea on apical half,scutellar foveae large, lateral bars of scutellum smooth, shining, short (cf. Fig. 180). Mesopleural suturedistinct, sterno-pleural suture distinct; propodeum with weak pubescence at base of tergite 2, complete ondorsal surface, tergite 2 the largest in lateral view, segment 3 partially visible in lateral view, gasterimpunctate, hypopygium visible, not extending past apex of sheath. Wing surface densely pubescent, apexof wing arcuate, with a fringe of hairs, radial cell of forewing completely open on wing margin. Colour:antenna pale yellow, head, thorax and gaster dark brownish red, legs pale yellow.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Mossel Bay, Cape Province, x.1921 (R. E. Turner) (BMNH).Paratypes. Zaire: 1 $ (MRAC). Zimbabwe: 1 $ (BMNH). REMARKS. This species is distinguished by the filiform antenna, and the six apical segments which are veryweakly clavate and bear rhinaria (Fig. 203). Kleidotoma erebus sp. n. (Figs 182, 183, 184, 185, 198) DESCRIPTION. $. Antenna 13-segmented, the apical 3 segments forming a distinct club, segment 3 1-5 xlength of 4, 4-7 subequal in length, 8-10 almost as wide apically as long (cf. Fig. 182). Head, viewedfrontally, smooth, polished, eyes oval, as far apart measured medially as height of an eye, inner orbits witha carina partially extending from outer area of antennal scrobe, malar grooves percurrent, anteriortentorial pits distinct, lower face with sparse setae extending to clypeal region, occiput strongly strigose.Pronotal plate projected forward, anterior and posterior parts joined medially by a broad bridge, foveaeither side of bridge open laterally, with sparse pubescence (Fig. 183). Pronotum with striations either sideof pronotal plate. Mesoscutum smooth, polished, lateral bars of scutellum striate, scutellar foveae largeand deep apically, scutellar cup long, narrow, extending to apex of scutellar disc, apex of disc vieweddorsally almost square (Fig. 185). Mesopleura smooth, polished, carina complete, metapleura ridgedabove hind coxa; propodeum pubescent dorsally and laterally. Segment 1 of gaster obscured by a ring ofhairs at base of tergite 2, tergite 2 the largest in lateral view, 3 and 4 partially visible, apex of tergite 2 andvisible parts of 3 and 4 punctate, hypopygium not pronounced (Fig. 184). Wing surface pubescent, radialcell of forewing open on margin, Rs 2 partially extending along margin, apex of wing incised, with a fringe ofhairs (Fig. 198); legs of normal proportions. Colour: antenna yellowish basally, club segments darker,head, thorax and gaster chestnut-red, legs orange-yellow.O" unknown. MATERIAL EXAMINED Holotype $, Zaire: N. Lac Kivu, Rwanki, 15. ii. 1952 (/. V. Leroy) (MRAC).Paratypes. Zaire: 3 $ (MRAC). AFROTROPICAL EUCOILIDAE 285 REMARKS. Similar to arbitra in many respects but differing by the sculpture of the occiput and the shape ofthe scutellar disc. Kleidotoma favussp. n. (Figs 183, 190, 199,205) DESCRIPTION. $. Antenna 13-segmented with a distinct 3-segmented club, club segments with rhinaria,wider and longer than preceding flagellar segments, segment 3 as long as 4+5, segments 4-10 as wideapically as long, some obconical (Fig. 190). Head, viewed frontally, smooth, polished, eyes small, almostround, further apart measured medially than height of an eye, malar grooves percurrent, anterior tentorialpits distinct. Pronotal plate, viewed dorsally, with a large fovea on either side of medial bridge, notpubescent, surface reticulate-rugose (cf. Fig. 199). Pronotum on dorsal and side margins at juncture withmesoscutum canaliculate-striate (cf. Fig. 183). Mesopleura smooth, shining, suture distinct. Mesoscutumsmooth, polished, scutellar disc with strong striate sculpture on lateral margins, apex irregularly rounded,rugose, lateral bars of scutellum strongly striate, scutellar fovea large, shallow, polished, separated byseptum of scutellar cup, scutellar cup long, narrow, not extending to apex of scutellar disc, with a fovea onapical half (Fig. 205). Segment 1 of gaster obscured by a dense ring of pubescence at base of tergite 2,tergite 2 the largest in lateral view, 3 and 4 partially visible, impunctate. Wing surface with dotted hairbases, radial cell of forewing completely open on margin, apex of wings incised, with apical hair fringe.Colour: antenna yellow at apex, becoming darker apically, head and thorax dark reddish black, gasterchestnut-red-brown, legs orange-yellow. Cf . Antenna 15-segmented, filiform, segment 3 curved, swollen distally. MATERIAL EXAMINED Holotype <J>, Cameroun: Nkoemvon, 1980 (D. Jackson) (BMNH). Paratypes. Cameroun: 3 $ , 2 cf (BMNH). Kenya: 12 $ , 2 cf (BMNH). Nigeria: 1 $ (BMNH). Uganda:2 $ (BMNH). Zaire: 44 $ (MRAC). Zimbabwe: 17 $, 15 cf (BMNH). REMARKS. This species closely resembles strigosa but is separated by the flagellar segments of the antennapreceding the club. Kleidotoma fimbriata sp. n. (Figs 192, 205, 206) DESCRIPTION. $. Antenna 13-segmented, almost filiform, club 4-segmented, not sharply defined, all withrhinaria, segment 3 clearly longer than 4 (Fig. 192), 4-9 almost subequal in length, the apical segmentsslightly progressively shorter than each preceding segment, club segments 10-13 subequal to each other,each longer than segment 9 and weakly swollen medially. Head, viewed frontally, smooth, polished, malargrooves distinct, eyes further apart measured medially than height of an eye, clypeus and area above withscattered pubescence, occiput striated. Pronotal plate projected forward, anterior part sculptured,posterior part polished; lateral bars and scutellar disc with coarse striations, scutellar cup elliptical, apex ofdisc rounded (cf. Fig. 205), scutellar foveae large, polished. Mesopleural suture distinct. Segment 1 ofgaster obscured by a ring of dense pubescence at base of tergite 2, apex of tergite 2 and visible part of 3 withsparse punctures, hypopygium not protruding (Fig. 206). Wing surface pubescent, apex of wings incised,with a long fringe of hairs, radial cell of forewing open on wing margin. Colour: antenna brownish yellow,head and thorax black, gaster reddish brown, legs orange-yellow.Cf unknown. MATERIAL EXAMINED Holotype $ , Zaire: Ruanda, Kibga, Vole. Bishoke, 2,400 m, 8-9.xi.1935 (G. F. de Witte) (MRAC).Paratypes. Zaire: 3 9 (MRAC). REMARKS. This species has almost filiform antenna and an impunctate gaster, distinguishing it fromnitidiuscula. Kleidotoma montana Kieffer Kleidotoma montana Kieffer, 1910c: 107. Holotype $ , RWANDA: Vulkan Karishimbi, 2,700 m, xi. (Exped.Herzog Adolf Friedrich zu Mechlenburg). The holotype of this species cannot be traced. In his description Kieffer indicates that the club of the 286 J. QUINLAN antenna is 3-segmented, flagellar segments 4-10 quadrate, head smooth, polished. The scutellar cup is notmentioned in the description. In his figure of the whole insect, the wings have an apical fringe of hairs andare indented apically. Tergite 2 of the gaster has a ring of hairs at the base and appears complete on thedorsal surface; segments 2-5 are visible in lateral view. The drawing does not indicate whether or not anysegments of the gaster are punctate, and the mesopleural suture is not indicated. The radial cell of theforewing is typical for Kleidotoma. A female from Zaire, Ruanda, Contrefort Est. Muhavura, 2,100 m,28. i. 1953 (P. Basilewsky), and determined by Benoit as montana, is in MRAC but cannot be accepted asthis species. From the description and figure I am unable to recognise montana and regard it as incertaesedis. The figure indicates that the antennal segments 4-10 are quadrate as in strigosa. Kleidotoma morsum sp. n. (Figs 176, 177, 179, 207) DESCRIPTION. $. Antenna 13-segmented, segment 2 as long as 3 and twice as wide, segment 3 as long as4+5, 4 and 5 subequal, segments 6-11 subequal, short, wider than long, obconical, 12-13 forming a2-segmented club, both with rhinaria present (Fig. 179). Head, viewed frontally, smooth, polished, eyessmall, round, further apart measured medially than height of an eye, malar grooves weakly indicated.Pronotal plate projecting, weakly crenulate, lateral fovea open (Fig. 207). Mesoscutum smooth, polished,scutellar disc sharply conical at apex, lateral margins smooth, shining, without striations, scutellar cupnarrow, elliptical, with a large fovea at apex (Fig. 176), lateral bars of scutellum smooth, shining, extendingjust past scutellar foveae, scutellar foveae small, deep. Mesopleural suture distinct. Segment 1 of gasterobscured by a ring of dense pubescence at base of tergite 2, hypopygium weakly visible, apex of tergite 2without punctures, gaster lenticulate. Wing surface without pubescence, dotted hair bases present, apex ofwing strongly incised, apical hair fringe long, radial cell of forewing completely open on margin (Fig. 177).Colour: antennal segments 1-11 dark yellow, 12-13 brownish yellow, head, thorax and gaster darkbrownish red, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $ , South Africa: Port St. John, Pondoland, 15-31.V.1923 (R. E. Turner) (BMNH).Paratypes. Zaire: 6 9 (MRAC). REMARKS. Distinguished from the closely related nigrans by the elongate antennal segment 2, thewedge-shaped incision of the apical margin of the forewing, the polished scutellar disc and impunctategaster. Kleidotoma nigrans sp. n. (Figs 178, 180, 207, 208) DESCRIPTION. $ . Antenna 13-segmented, segment 2 shorter than 3 and twice as wide, segment 3 as long as4+5, slightly longer than 5, 5-9 each longer than broad and subequal to each other, 10 and 11 equal inlength, shorter than 9, 12 and 13 forming a distinct club, both with rhinaria, club segments 12-13 togetheras long as segments 6-11 combined (Fig. 178). Head viewed frontally longer than wide, eyes further apartmeasured medially than height of an eye, mandibles large tridentate, with a few hairs in clypeal region,malar grooves distinct, occiput weakly sculptured. Pronotal plate small, projected forwards viewedfrontally, ventral surface sculptured, lateral fovea either side of medial bridge open (cf. Fig. 207).Mesoscutum smooth, polished, suture distinct; scutellar disc rounded at apex, lateral margins withobsolete striate sculpture, shiny, scutellar fovea large, squarish, scutellar cup narrow, elliptical, with alarge fovea apically (Fig. 180), lateral bars of scutellum short, weakly sculptured. Segment 1 of gasterobscured by a ring of pubescence at base of tergite 2, incomplete on dorsal surface, gaster lenticular,punctate, tergite 3 partially visible in lateral view. Wing surface dotted with hair bases, radial cell offorewing completely open on wing margin, apex of wing sharply incised, with apical hair fringe (Fig. 208).Colour: segments 1-11 of antenna dark brownish yellow, club segments 12 and 13 black, head, thorax andgaster chestnut-brown, legs clear yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Terr. Rutshuru, 7.iv.l937 (Mission Prophylactique) (MRAC).Paratype. Zaire: 1 $ , same data as holotype except 4.iv.l934 (MRAC). AFROTROPICAL EUCOILIDAE 287 REMARKS. Closely related to morsum but separated from it by a number of characters, particularly thestriations of the scutellar disc, the elongate cup which extends to the apex of the disc, and the short antennalsegment 2. Kleidotoma nitidiuscula sp. n. (Figs 191, 201, 209, 210) DESCRIPTION. $ . Antenna 13-segmented, segment 2 shorter than 3, 3 twice length of 4, equal to 4+5, 5 and6 subequal in length, 7 and 8 subequal, slightly shorter than 6, 9 longer than 8, broader at base, 10-13forming a 4-segmented club, club segments with rhinaria, segment 10 1-2 x length of 9, 11 and 12 equal inlength, 13 slightly longer than 12 (Fig. 191). Face, viewed frontally, smooth, polished, malar groovespercurrent, eyes further apart measured medially than height of an eye, occiput with strong strigosesculpture. Pronotal plate projected forward, viewed dorsally, sculptured on basal half and hind margin ofapical half, fovea on lateral margins open (Fig. 209); side margins of pronotum adjacent to pronotal platesculptured, scutellar disc truncate at apex, strongly striate on lateral margins, scutellar cup long, narrow,elliptical, extending to apex of disc or beyond, lateral bars of scutellum striate on lateral margins, dorsalsurface smooth, shining, scutellar foveae large, deep, shiny. Mesopleura smooth, polished, suture distinct.Segment 1 of gaster obscured by a dense ring of pubescence at base of tergite 2, tergites 2-4 visible in lateralview, apex of 2 and visible parts of 3 and 4 densely punctate (cf. Fig. 201), hypopygium not prominent.Wing surfaces with dotted hair bases, apex of forewing sharply incised, with apical hair fringe, radial cell offorewing completely open (Fig. 210). Colour: antennal segments 1-9 dark yellow, 10-13 brownish, headand thorax black, gaster reddish basally, black on dorsal and lateral surfaces, legs dark yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: '(Congo Beige PN.G.) Miss.II De Saeger, ii/Fd/17, 15.ix.1951. (Rec.H. de Saeger)'(MR AC). REMARKS. This species is very similar tofimbriata but is distinguished by the sharply defined club segmentsand dense punctuation of the gaster. Kleidotoma norma sp. n. (Figs 194, 195,196,211) DESCRIPTION. $. Antenna 13-segmented, filiform, segment 3 longer than 4, 4-6 subequal in length, 7-13forming a weak club, distinguished by presence of rhinaria and the slight medial swelling of each segment,club segments 7-12 a little longer than preceding flagellar segments, segment 13 1-3 x length of 12 (Fig.195). Face, viewed frontally, smooth, polished, area above clypeus raised, malar grooves percurrent,anterior tentorial pits large, clypeal margin with long sparse setae, occiput smooth, shiny, with brokensculpture. Pronotal plate projected forward, viewed dorsally, sculptured on anterior part, fovea either sideof medial bridge open on lateral margins (Fig. 211). Pronotum either side of pronotal plate smooth,polished. Mesoscutum smooth, polished, with a few long scattered hairs; lateral bars of scutellum andscutellar disc very weakly striated, scutellar foveae smooth, polished, scutellar cup elliptical, apex of discrounded (Fig. 194). Mesopleural suture distinct, lower part of pronotum adjacent to mesopleura striated,area below precoxal carina pubescent. Segment 1 of gaster obscured by a ring of dense pubescence at baseof tergite 2, tergites 2 and 3 impunctate, hypopygium not protruding. Wing surfaces densely pubescent,truncate at apex which has an apical fringe of hairs, radial cell of forewing completely open on margin (Fig.196). Colour: antenna brownish, head and thorax black, gaster blackish brown, legs orange-yellow.C? unknown. MATERIAL EXAMINEDHolotype $, Zimbabwe: Chishawasha, nr Salisbury, x.1979 (A. Watsham) (BMNH). REMARKS. This species is separated by the inconspicuous 7-segmented club, which is almost filiform, andthe presence of rhinaria on each club segment. Kleidotoma strigosa sp. n. (Figs 183, 189,205,211,212)DESCRIPTION. $. Antenna 13-segmented with a distinct 3-segmented club, each segment broader and 288 J. QUINLAN longer than preceding flagellar segments, segment 3 as long as 4+5, segment 4 slightly longer than broad,5-10 quadrate, as wide as long, apical and basal widths equal (Fig. 189). Head viewed frontally smooth,polished, malar grooves percurrent, eyes round, further apart measured medially than height of an eye,occiput with weak strigose sculpture. Pronotal plate, viewed dorsally, large, projected forward, foveaeither side of medial bridge large, open on lateral margins, with tufts of pubescence (cf. Fig. 211).Pronotum on side margins at juncture with mesoscutum and pronotal plate strigose (cf. Fig. 183).Mesoscutum smooth, polished, scutellar disc with widely spaced longitudinal striations, apex elongate,rounded, reticulate, scutellar cup narrow, elliptical, not extending to apex of disc, apex of cup with a largefovea, lateral bars of scutellum short, striate, as long as scutellar foveae which are deep and separated bybroad septum of cup (cf. Fig. 205). Mesopleura smooth, polished, suture distinct. Segment 1 of gasterobscured by a ring of dense pubescence, apex of tergite 2 and visible segments impunctate, hypopygiumpronounced. Wing surface with very short pubescence, apex of wing strongly arcuate, with apical hairfringe, radial cell of forewing completely open on wide margin (Fig. 212). Colour: antenna dark reddishbrown, head and thorax blackish red, coxae brownish red, femora, tibiae and tarsi brownish yellow.Cf . Antenna 15-segmented, flagellar segments 4-10 as broad as long. MATERIAL EXAMINED Holotype $, Cameroun: Nkoemvon, 1980 (D. Jackson) (BMNH). Paratypes. Cameroun: 5 $, 2 cf (BMNH). Kenya: 11 $, 12 cf (BMNH). Nigeria: 1 $ (BMNH).Uganda: 2 $ (BMNH). Zaire: 42 $ (MRAC). Zimbabwe: 17 $ (BMNH). REMARKS. A distinctive species, the quadrate flagellar segments 5-10 separate it from the closely relatedfavus. Kleidotoma ventosus sp. n. (Figs 181, 182, 186,210,211) DESCRIPTION. $ . Antenna 13-segmented with a distinct 3-segmented club, club segments each broader andlonger than the preceding flagellar segments, segment 3 subequal to 4+5, flagellar segments 4-11 eachprogressively shorter than each other, longer than wide, wider apically than basally (Fig. 182). Head,viewed frontally, smooth, polished, malar grooves complete, eyes almost round, further apart measuredmedially than height of an eye. Pronotal plate, viewed dorsally, protruding, with large fovea on either sideof medial bridge, open laterally, smooth, shining (cf. Fig. 211). Pronotum on side margins at junction withmesoscutum and pronotal plate strongly canaliculate (Fig. 181). Mesopleural suture distinct, with finereticulate sculpture below suture. Mesoscutum smooth, polished; scutellar disc striate, apex of discrounded, reticulate, scutellar cup long, narrow, elliptical, not reaching apex of disc, apex of cup with alarge fovea, lateral bars of scutellum strongly striate, scutellar foveae large, polished, with narrow septumof cup separating them (cf. Fig. 186). Segment 1 of gaster obscured by a ring of dense pubescence at base oftergite 2, apex of tergite 2 and visible parts of tergites 3 and 4 punctate. Wing surfaces dotted with hairbases, apex of wing sharply incised with apical hair fringe, radial cell of forewing completely open on wingmargin, vein Rs 2 weakly extending along margin of wing (Fig. 210). Colour: segments 1-10 of antenna darkyellow, club segments 11-13 brown, head dark brown-black, thorax black, except for propodeum andmetapleura which are dark yellow-orange, legs dark yellow.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Port St John, Pondoland, l-5.iv.1923 (R. E. Turner} (BMNH).Paratypes. Ethiopia: 1 $ (BMNH). South Africa: 3 $ (BMNH). Zaire: 4 $ (MRAC). REMARKS. The length of the antennal segments and the densely strigose occiput distinguish this species. NORDLANDERIA gen. n. Type-species: Nordlanderia plowa sp. n. DIAGNOSIS. $ antenna 13-segmented, clavate, with 8-9 club segments, cf antenna 15-segmented, filiform,segment 3 1-5 x length of 4, weakly curved, swollen distally. Head, viewed frontally, with eyes slightlyfurther apart measured medially than height of an eye, inner margins almost parallel, malar space with asubocular suture, a few radiating striae on either side, supraclypeal area and anterior region of face withshort protrusions (Fig. 16). Pronotal plate not protruding, very weakly indicated (Fig. 223), lateral foveaeither side of median bridge open. Mesoscutum with or without notaulices, if present, very weakly AFROTROPICAL EUCOILIDAE 289 indicated, converging sharply towards scutellum, scutellar foveae obsolete. Mesopleural suture complete,area below suture sculptured (Fig. 214). Propodeal carinae bowed, medial area polished, with tufts ofpubescence on either side. Gaster with segment 1 almost obscured, base of tergite 2 partially enclosing it,when visible, short and broad, in the form of a weakly sculptured ring, tergite 2 the largest, occupyingwhole area of gaster viewed laterally, smooth, polished (Fig. 13). Wing surfaces pubescent, with ciliatemargins, longest at apex, forewing with a closed radial cell although this can appear open in somespecimens. Legs short, broad, coxae with scattered hairs, femora, tibiae and tarsi moderately pubescent. DISCUSSION. Nordlanderia has close affinities with Microstilba Foerster, but differs in having pronouncedtriangular projections on the face and the notaulices almost absent. In those species where they are presentthey converge posteriorly towards the scutellum at a point almost mid-way between the pronotum and thescutellum. As with most eucoilids, the males are difficult to key and separate to species. DISTRIBUTION. Afrotropical Region. Key to the species of Nordlanderia gen. n. Females 1 Hypopygium prominent, with long apical hairs (Fig. 222), segments 2-4 of gaster visible in lateral view ads sp. n. (p. 289) - Hypopygium not prominent (Figs 215, 220), hairs sparse, segments 2-3 only of gaster visible in lateral view 2 2 Antennal club 8-segmented, segment 3 longer than 4 (Fig. 216); notaulices complete though very weakly indicated (Fig. 218) ; head, thorax and gaster black plowa sp. n. (p. 290) - Antennal club 9-segmented, segments 3 and 4 of equal length; notaulices aberrant, almost absent anteriorly (Fig. 217) ; head and thorax black, gaster orange-yellow .... pallida sp. n. (p. 289) Nordlanderia acissp. n. (Figs 214, 217, 219, 221, 222, 223) DESCRIPTION. $. Antenna 13-segmented, weakly clavate, club 8-segmented, segment 3 1-2 x length ofsegment 4 (Fig. 219). Head, viewed frontally, smooth, polished, frons raised, malar grooves percurrentwith striations on either side of them, cheeks below grooves extruded to form three tooth-like appendages(Fig. 221), eyes weakly converging, frontal area raised. Pronotal plate not projected, anterior andposterior parts connected by a medial bridge, with fovea on lateral margins open (cf. Fig. 223).Mesoscutum smooth, polished, notaulices very weakly represented, almost absent in some specimens,converging sharply towards scutellum, lateral bars of scutellum polished, scutellar cup slightly longer thanwide, with a large apical fovea, scutellar disc with widely spaced radiating striae or ridges, scutellar foveaeaberrant (Fig. 217). Mesopleural suture complete, area below suture coriaceous (cf. Fig. 214); propodealcarinae parallel, converging at base towards gaster, lateral margins densely pubescent. Segment 1 of gasternot completely visible in lateral view, in the form of a ring, tergites 2-3 visible in lateral view, hypopygiumpronounced (Fig. 222). Wing surface pubescent, apical margins with a hair fringe, radial cell of forewingcompletely closed on wing margin, longer than wide. Legs normal, moderately pubescent. Colour:antenna yellow basally, apical segments darker, legs orange-yellow, head and thorax chestnut-brown,gaster lighter. Cf . Antenna 15-segmented; segment 3 longer than 4, curved, swollen distally. MATERIAL EXAMINED Holotype $, Namibia: Swakopmund, 26-30.L1972 (BMNH).Paratypes. South Africa: 31 $, 45 cf (BMNH). REMARKS. This species is easily distinguished from others by the prominent hypopygium (Fig. 222). Nordlanderia pallida sp. n. (Figs 213, 217, 220, 223) DESCRIPTION. $. Antenna 13-segmented, weakly clavate, with a weak 9-segmented club, segment 3 equalin length to segment 4. Head viewed frontally with scattered hairs between eyes, malar space withsubocular suture distinct, with striae on either side of sutures, supraclypeal area and anterior region effacewith three strongly projecting spine-shaped keels (cf. Fig. 213). Pronotal plate not projected forward,almost aberrant, not prominent, anterior and posterior parts connected by a medial bridge with an open 290 J. QUINLAN fovea on either side of bridge (Fig. 223). Mesoscutum smooth, polished, with only the faintest indication ofnotaulices, sometimes virtually absent (Fig. 217); scutellar cup large, concave basally, rim of cup almostlevel with apex of scutellar disc which has radiating sculpture (Fig. 217) . Mesopleural suture complete , areabelow suture coriaceous. Propodeal carinae weakly bowed, almost parallel, median area smooth,polished, sides of propodeum with tufts of pubescence. Segment 1 of gaster obscured by base of tergite 2,viewed laterally, in form of a thin ring, tergite 2 the largest, occupying whole of lateral surface of gaster(Fig. 220). Wing surface pubescent, with a long apical hair fringe, radial cell of forewing closed on margin.Legs with narrow coxae, femora, tibiae and tarsi moderately pubescent. Colour: antenna yellow, legsyellow, head and thorax black, gaster reddish yellow. Cf . Antenna filiform, segment 3 longer than 4, weakly curved, flattened medially on inner margin orsurface. MATERIAL EXAMINED Holotype $, South Africa: Pretoria, i.1931 (N. E. Munro) (BMNH).Paratypes. Ghana: 1 $, Accra, vi.1921 (/. Ingram) (BMNH), 1 $, 2 cf , Irat (GERDAT). REMARKS. This species is closely related toplowa but can be separated by the antennal characters and theabsence of notaulices. Nordlanderia plowa sp. n. (Figs 213, 216, 218, 223) DESCRIPTION. $. Antenna 13-segmented, clavate, with a weakly defined 8-segmented club, segment 3 1-5x length of 4 (Fig. 216). Head, viewed frontally, with scattered hairs between eyes which are almostparallel and slightly further apart than height of an eye measured medially, malar space with subocularsuture distinct, with weak striae on either side, supraclypeal area and anterior region of face stronglyprotruding with tooth-like appendages (Fig. 213). Pronotal plate not protruding, viewed fronto-dorsally,flattened into pro no turn (cf. Fig. 223), anterior and posterior parts of plate connected by a medial bridgewith a fovea on either side, open laterally. Mesoscutum smooth, polished, with notaulices weaklyindicated, converging towards scutellum, sometimes absent in anterior area (Fig. 218); scutellar cup large,rim of cup overhanging disc, viewed dorsally, basal half concave, apical half convex (Fig. 218); scutellardisc with radiating ridges. Mesopleura with a distinct suture, area below suture coriaceous; propodealcarinae bowed medially, inner area smooth, polished, lateral margins with tufts of pubescence. Segment 1of gaster obscured, in the form of a narrow ring, tergite 2 the largest, tergites 3 and 4 sometimes visible inlateral view . Wing surface pubescent , with apical hair fringe , radial cell of forewing with a closed cell (this issometimes difficult to appreciate). Legs with short broad coxae and scattered pubescence, femora, tibiaeand tarsi moderately pubescent. Colour: antenna yellow basally, brownish apically, coxa and femurbrownish, tarsus yellow, head, thorax and gaster black. Cf . Antenna 15-segmented, filiform, segment 3 longer than 4, curved, swollen distally. MATERIAL EXAMINED Holotype $, Zimbabwe: Salisbury, Chishawasha, x.1979 (A. Watsham) (BMNH). Paratypes. Aldabra: 1 cf, Astove Atoll (BMNH). Nigeria: 4 9, 8 cf (BMNH). Zaire: 17 $, 43 cf(MRAC). Zambia: 5 cf (BMNH). Zimbabwe: 19 $, 13 cf (BMNH). REMARKS. Distinguished frompallida by the presence of notaulices and by the antenna. RHOPTROMERIS Foerster Rhoptromeris Foerster, 1869: 344. Type-species: Cothonaspis eucera Hartig, 1841, by original designation[a junior synonym of Cothonaspis heptoma Hartig, 1840]. Miomoera Foerster, 1869: 352. Type-species: Miomoera aberrans Foerster, 1869, by subsequent designa-tion, Rohwer & Pagan 1917: 371. Hexamerocera Kieffer, 1901: 175. Type-species: Eucoila rufiventris Giraud, 1860, by subsequent designa-tion, Ashmead, 1903: 66. DIAGNOSIS. $ antenna 13-segmented, with 7-segmented club in type-species (this varies from species tospecies), all club segments bear rhinaria and are generally darker than basal segments of flagellum.Segment 3 longer than 4 in type-species, cf antenna 15-segmented, segment 4 longer than 3, curved,swollen distally (Fig. 225). Head, viewed frontally, round, slightly longer than wide, supraclypeal arearaised, malar area with a groove or sulcus extending from bottom of eye to margin of clypeus. Pronotal AFROTROPICAL EUCOILIDAE 291 plate, viewed dorsally, protruding, well developed (Fig. 6), lateral fovea on either side of plate enclosed byouter margins of anterior and posterior parts of plate. Mesoscutum smooth, polished, notaulices absent, insome species a row of hairs present in their place. Lateral bars of scutellum smooth, scutellar discreticulate-rugose, sometimes with radiating striae from edge of narrow scutellar cup which has one or twofovea centrally. Mesopleurae smooth, shiny, mesopleural carina straight, metapleura sometimes withweak strigose sculpture ventrally and posteriorly; thorax with short dense pubescence on ventral parts.Propodeum with almost parallel carinae, either side of carinae with short dense pubescence. Segment 1 ofgaster (petiole) visible in part laterally, crenulate (Fig. 249), base of tergite 2 with a ring of dense yellowishpubescence, usually complete on dorsal surface though usually incomplete in males (cf. Fig. 36), tergites2-5 visible in $, 2-6 in cf in lateral view, hypopygium distinct (Fig. 227). Legs varying in length androbustness between species, fore-coxae slender, mid- and hind-coxae swollen, meta-tarsi as long as tarsalsegments 2-^4 combined, claws simple. Wings densely pubescent on surface, with apical fringe of hairs,radial cell of fore wing long, between 2-3 x as long as width (Fig. 224), closed on front margin. Colour:antenna yellow basally, club segments darker, head and thorax blackish, gaster dark yellow-brown, legsyellowish, coxae and femora darker. Rhoptromeris is related to Trichoplasta and Stentorceps, sharing aderived character in the form of the pronotal plate. In all three genera the pronotal plate has the medialfovea between the anterior and posterior parts fused laterally. In most other genera the foveae or pits areopen (Nordlander, 1982). Rhoptromeris has a characteristic radial cell, the veins 2r and Rs (Nordlander,19826) and Rs 2 and Rsi (Quinlan, 1979; Eady & Quinlan, 1963) of about equal length. The scutellar cup inall three genera is relatively small. In the males the 4th antennal segment is longer than the 3rd andmodified (Fig. 225). DISTRIBUTION. Old and New World. Key to the Afrotropical species of Rhoptromeris Foerster Females 1 Antenna filiform, segment 3 generally subequal to or shorter than 4 (Fig. 229), 4-13 at least 4 x as long as wide, those segments with rhinaria weakly swollen medially, not forming aconspicuous club 2 - Antenna subclavate, apical segments 4-13 less than 4 x as long as wide, segment 3 rarely shorter than 4, either subequal to or longer than 4, club segments with rhinaria, widerapically than basally (Fig. 237) 10 2 Antenna with the 6 apical segments bearing rhinaria (Fig. 229); hypopygium pronounced, ventral spine short; pronotal plate with a transverse groove, anterior and posterior parts of plate fused laterally zeus sp. n. (p. 308) Antenna with segments 7-9 bearing rhinaria (Figs 230, 267, 278), hypopygium not pro-nounced, ventral spine variable in length; pronotal plate either with a transverse groovemedially between anterior and posterior parts of plate or with two fovea enclosed on lateralmargin 3 3 Antenna with the 7 apical segments bearing rhinaria 4 - Antenna with 8-9 apical segments bearing rhinaria 6 4 Pronotal plate with a transverse groove medially between anterior and posterior parts (cf. Fig. 250); antenna with segment 3 shorter than 4 (Fig. 230). Scutellar cup oval, scutellar disc with weak broken sculpture laterally, apex with radiatingsculpture (Fig. 231) oeta sp. n. (p. 302) - Pronotal plate with lateral foveae separated by a medial bridge (cf. Fig. 6); antennal segment 3 shorter than 4 (Fig. 272) 5 5 Apical antennal segments of antenna darker than basal flagellar segments; scutellar cup elliptical , polished , scutellar disc smooth , polished cepheus sp . n . (p . 297) Apical 3 segments of antenna white, segments 3 and 4 subequal, scutellar cup long, narrow,scutellar disc with radiating ridged sculpture laterally (cf. Fig. 238) rwanki sp. n. (p. 305) 6 Antenna with 9 apical segments bearing rhinaria, weakly swollen medially, pronotal plate with lateral fovea separated medially by a bridge crito sp. n. (p. 297) Antenna with 8 apical segments bearing rhinaria; fovea of pronotal plate either separatedmedially or in the form of a furrow 7 7 Pronotal plate with a transverse groove separating anterior and posterior parts (cf. Fig. 250), no medial bridge between lateral pits apparent pagasa sp. n.(p. 302) 292 J. QUINLAN Pronotal plate with two lateral fovea separated by a shallow medial bridge (cf. Fig. 275)connecting anterior and posterior parts , antennal segment 3 shorter than 4 8 8 Scutellar disc with strong radiating striations, apex with strongly radiating ridges (Fig. 234), scutellar cup long, elliptical; the 8 apical segments of antenna with prominent rhinaria,weakly swollen medially (Fig. 282); fovea on pronotal plate separated by a broad medial bridge (cf. Fig. 232) temesa sp. n. (p. 306) Scutellar disc with aberrant sculpture; pronotal plate with a narrow medial bridge connectinganterior and posterior parts (cf. Fig. 275) 9 9 Apical segments of antenna white; visible parts of gastral segments 3 and 4 punctate rutshuris sp. n. (p. 305) Apical segments of antenna darker than yellowish basal segments; segments 3 and 4 of gasterimpunctate agissp. n.(p. 295) 10 Antennal club 6-segmented 11 - Antennal club 7-8-segmented 15 11 Pronotal plate with a transverse groove between anterior and posterior parts, closed laterally (cf. Fig. 250) , medial bridge not clearly defined 12 - Pronotal plate with two fovea between anterior and posterior parts, separated medially by a bridge (cf. Figs 232, 275) 13 12 Antennal club sharply indicated (Fig. 237); segment 3 as long as 4+5, 5-7 a little longer than wide (Fig. 237) afersp. n. (p. 294) - Antennal club weakly indicated, segments 8-13 weakly swollen medially, 3 and 4 subequal in length velia sp. n. (p. 307) 13 Apex of tergite 2 of gaster with scattered punctures, tergites 3 and 4 densely punctate on visible parts. Segment 3 of antenna clearly longer than 4 (Fig. 240); scutellar cup elliptical, discpolished, rounded apically abba sp. n. (p. 294) - Tergites of gaster impunctate, scutellar disc conical 14 14 Apex of scutellar disc weakly conical, surface reticulate-coriaceous (Fig. 244); antenna short, as long as distance between front of head and apex of gaster, club segments a little longer than wide (Fig. 242) diversa sp . n. (p. 299) - Apex of scutellar disc weakly tapered (Fig. 245) , surface reticulate-rugose ; antenna longer than distance between front of head and apex of gaster, club segments of antenna at least 2 x as long as wide (Fig. 241) connatus sp. n.(p. 297) 15 Antennal club 7-segmented 16 - Antennal club 8-segmented 30 16 Pronotal plate with a weak transverse area between anterior and posterior parts, closed on lateral margins (Fig. 250) 17 - Pronotal plate with two fovea separated medially by a bridge between anterior and posterior, closed laterally (cf. Fig. 275) 20 17 Antennal segment 3 longer than 4, club segments sharply delineated (Fig. 247) , at most twice as long as broad; hypopygium deep basally. Ovipositor as long as base of hypopygium; pronotum, mesopleura and gaster yellow-brown, sharply contrasting with the black pronotal plate, thorax and head bicolor sp. n. (p. 296) - Antennal segments 3 and 4 subequal and/or club segments not sharply delineated ; hypopygium not deep basally (Fig. 249) 18 18 Apex of 3rd and 4th segment of gaster finely punctate (Fig. 249). Basal six segments of antenna yellowish, club segments 7-10 black, 11-13 white zetessp. n.(p. 308) - Apex of 3rd segment and visible apical segment of gaster impunctate 19 19 Antennal segments 4-6 subequal in length, the seven apical segments with rhinaria, not forming a distinct club (Fig. 251) persius sp. n. (p. 304) - Antennal segments 3-6 subequal, the seven apical segments with pronounced rhinaria, forming a distinct club (Fig. 253) equalis sp. n.(p. 300) 20 Apex of segment 3 of gaster and visible apical segments finely punctate 21 AFROTROPICAL EUCOILIDAE 293 Apex of segment 3 of gaster and visible apical segments impunctate (if some species weaklypunctate then antennal segments 4-6 quadrate) 25 21 Antennal segment 3 as long as 4+5 22 - Antennal segment 3 shorter than 4+5 23 22 Club segments 7-13 of antenna not pronounced (Fig. 248), segments 4-6 each longer than broad ;scutellar disc smooth, polished punctate sp. n. (p. 304) Club segments 7-13 pronounced (Fig. 259), segments 4-6 each as wide as long; scutellar discreticulate-coriaceous enna sp. n. (p. 299) 23 Antennal segments 3 and 4 equal in length sinis sp. n . (p. 306) Antennal segment 3 longer than 4 24 24 Segments 2-4 of gaster visible in lateral view , densely punctate (Fig . 254) ; metapleura smooth , anteroventral cavity without a tuft of pubescence; club segments 2 x as long as wide (Fig. 255) frupa/ussp. n.(p. 296) Segments 2-3 of gaster visible in lateral view, apex of 2 and visible part of 3 punctate;metapleura ridged, anteroventral cavity polished; club segments 3 x as long as wide (Fig.256) attissp. n. (p. 295) 25 Segment 3 of antenna longer than 4 26 Segment 3 of antenna equal to or shorter than 4 29 26 Anteroventral cavity of metapleuron reduced ; segment 3 of antenna as long as 4 + 5 , 4-6 almost quadrate, club segments sharply defined (Fig. 259) enna sp. n. (p. 299) Anteroventral cavity of metapleuron with a tuft of pubescence; segment 3 of antenna shorterthan 4+5 27 27 Foveae of pronotal plate very weak , broadly separated medially (cf. Fig . 262) ; club segments of antenna slightly longer than wide; whole insect completely pale yellowish orange, clubsegments very pale pallidussp. n. (p. 303) - Foveae of pronotal plate separated by a narrow medial bridge between anterior and posterior parts (Fig. 228); club segments of antenna clearly longer than wide, darker than flagellarsegments except apical segments in some species 28 28 Club segments of antenna 3 x as long as wide, antenna as long as distance between face and apex of gaster; scutellar cup large, oval, scutellar disc with weak radiating ridges, strongeston apical margin; pronotum, mesonotum and gaster light reddish orange, contrasting withblackish head and thorax; antenna yellow basally, brownish medially, apical segment or segments white rufulussp. n. (p. 304) Club segments of antenna 2 x as long as wide, antenna shorter than distance between face andapex of gaster; scutellar cup narrow, elliptical, scutellar disc weakly sculptured to smoothand polished; head, thorax and gaster blackish brown nepfoma (Hartig) (p. 300) 29 Antennal segment 3 as long as 4; radial cell of forewing short, broad (Fig. 261) ; lateral margins of scutellar disc polished, apex with radiating sculpture (Fig. 263), scutellar cup elliptical cubitalis sp. n. (p. 298) Antennal segment 3 shorter than 4; radial cell of forewing elongate (Fig. 265); scutellar discwith radiating striae, scutellar cup pear-shaped (cf. Fig. 263) thales sp. n. (p. 307) 30 Antennal segment 3 longer than 4, 4 and 5 subequal. Scutellar disc reticulate-coriaceous, apex weakly conical, scutellar cup almost oval (Fig.273); pronotal plate with a transverse groove medially connecting anterior and posterior parts, closed on lateral margins naxos sp. n . (p. 302) Antennal segment 3 equal to or shorter than segment 4 31 31 Anterior and posterior parts of pronotal plate with a transverse groove between them (cf. Fig. 250) hebesp. n.(p. 300) - Anterior and posterior parts of pronotal plate with two lateral foveae , one either side of medial bridge (Fig. 275) 32 32 Antennal segment 3 shorter than 4, 5 longer than 4, apical segments of gaster punctate agis sp. n. (p. 295) - Antennal segments 3 and 4 subequal in length, 5 shorter than 4, apical segments 7-12 darker, 13th segment white, basal segments yellowish; lateral margins of scutellar disc weaklystriated, apex strongly rugose nav/us sp. n. (p. 301) 294 J. QUINLAN Rhoptromeris abba sp. n. (Figs 232, 240, 243) DESCRIPTION. $. Antenna 13-segmented, segment 3 longer than 4, subequal to 4+5, 6 longer than 5, 7longer than 6, 8-13 with rhinaria, forming a club (Fig. 240). Head, viewed frontally, with eyes further apartmeasured medially than height ofan eye, face with sparse hairs on lower face, supraclypeal area slightlyraised, malar grooves distinct, extending from bottom of eye to edge of mandibles, mandibles yellowishbrown, maxillary palp 4-segmented, labial palp 2-segmented. Pronotal plate with lateral fovea either sideof medial bridge closed, either side of pronotal plate with sparse pubescence (cf. Fig. 232). Mesoscutumsmooth, polished, notaulices absent (some specimens with hairs in their place). Scutellar disc rounded atapex, lateral margins of disc weakly striated, almost smooth, scutellar cup narrow, minute, scutellar foveaeshallow, polished, axillae minute, polished. Mesopleural suture distinct, almost parallel with ventralmargin, metapleura ridged, anteroventral cavity distinct. Propodeal carinae subparallel, densely pubes-cent. Segment 1 of gaster as broad as long, base of tergite 2 of gaster with ring of dense pubescence at base,not complete on dorsal surface. Tergite 2 the largest in lateral view, apical margin finely punctate, visibleparts of tergites 3 and 4 densely punctate, hypopygium prominent, ovipositor short. Legs of normal shape,coxae robust, mid and hind coxae pubescent apically, tarsi 5-segmented. Forewings relatively narrow,surface densely pubescent, apical margin with a fringe of hairs, radial cell of fore wing closed, vein R\ short,one-third as long as Rs2, veins M and Rs+M obsolete. Colour: antenna yellowish basally, club segmentsdark, head, thorax and gaster dark chestnut-brown, legs orange-yellow. CT. Antenna 15-segmented, 4th segment bent, swollen, outer margin weakly excavate (Fig. 243); gastralsegments 2-6 visible laterally. MATERIAL EXAMINED Holotype $, Kenya: Kisumu, nr Lake Victoria, xi.1979 (M. D. Croft) (BMNH).Paratypes. Kenya: 1 $, 1 C? (BMNH). Zaire: 3 $ (MRAC). REMARKS. Closely related to diversa and connatus but distinguished by the punctate gastral segment. Rhoptromeris afer sp. n. (Figs 237, 238, 239, 250) DESCRIPTION. 9 . Antenna 13-segmented, segment 3 as long as 4+5, 6 longer than 5, 7 longer than 6, 8-13bearing rhinaria, forming a very distinct club (Fig. 237). Head, viewed frontally, with eyes as far apartmeasured medially as height of an eye, supraclypeal area weakly raised, anterior tentorial pits distinct,epistomal suture indicated, malar grooves percurrent, mandibles yellowish, maxillary and labial palpsnormal, i.e. 4- and 2-segmented respectively. Pronotal plate with a transverse groove medially betweenanterior and posterior parts, lateral margins of groove not open (cf. Fig. 250), either side of pronotal platewith a tuft of hairs. Mesoscutum smooth, polished, notaulices absent. Apex of scutellum rounded, lateralmargins of scutellar disc with weak radiating striae, scutellar cup narrow, elliptical (Fig. 238), scutellarfoveae deep, arched basally, axillae polished. Mesopleural suture distinct, metapleura with a few aberrantwrinkles, anteroventral cavity with a tuft of pubescence. Propodeal carinae subparallel, convergingbasally, pubescent on either side. Segment 1 of gaster obscured by a ring of pubescence at base of tergite 2,complete on dorsal surface, tergite 2 the largest segment in lateral view, 3 and 4 partially visible, allsegments smooth, polished, hypopygium short, ovipositor long. Legs normal shape, coxae robust, hindcoxa pubescent apically, tarsi 5-segmented. Wings densely pubescent, apical margin with a hair fringe,radial cell closed, RI short, one-third as long as Rs 2 (Fig. 239), veins M and Rs+M obsolete. Colour:antenna yellowish basally, club segments dark, head, thorax and gaster dark brownish yellow, legsyellowish.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: N. Lac Kivu, Rwanki, 25.xii.1950 (/. V. Leroy) (MRAC).Paratypes. Zaire: 4 $, same data as holotype (MRAC). Uganda: 1 $ (BMNH). REMARKS. This species is one of two which can be separated from closely related species by the form of thepronotal plate. Together with velia, the medial area connecting the anterior and posterior parts is in theform of a shallow furrow. AFROTROPICAL EUCOILIDAE 295 Rhoptromerisagissp. n. (Figs 230, 246) DESCRIPTION. $. Antenna 13-segmented, segment 3 shorter than 4, 5 longer than 4, 6-13 with rhinaria,forming a weak club. Head, viewed frontally , with eyes weakly converging, as far apart as height of an eyemeasured medially, anterior tentorial pits distinct, malar grooves present, face with scattered hairsextending to clypeal area, clypeus and mandibles yellowish, maxillary palp 4-segmented, labial palp2-segmented. Pronotal plate with two fovea separated medially by a narrow bridge between anterior andposterior parts, closed on lateral margins, anterior part protruded (cf. Fig. 246), pronotum either side ofplate with sparse pubescence. Mesoscutum smooth, polished, notaulices absent; scutellar disc roundedapically, scutellar foveae polished, lateral bars polished, scutellar disc with weak reticulate sculpture onlateral margins and long scattered setae, scutellar cup semi-oval, surface with a ring of minute hairs (cf. Fig.230). Mesopleural suture distinct, metapleura weakly ridged, anteroventral cavity bare. Propodeal carinaeweakly converging towards metanotal plate, either side of carinae pubescent. Segment 1 of gaster almostobscured by a ring of hair at base of tergite 2, sparse on dorsal surface, tergite 2 the largest in lateral view,occupying most of visible area, base of tergite 3 visible, impunctate, hypopygium long, narrow, ventralspine short. Legs long, thin, with fine pubescence on femora, tibiae and tarsi, tarsi 5-segmented. Forewingsbroad apically, surface densely pubescent, apex with a fringe of hairs, radial cell of forewing longer thanwide, narrow basally, entirely closed on margin, veins M and Rs+M indicated, not pigmented. Colour:antenna yellowish basally, club segments dark, head, thorax and gaster dark chestnut-brown, legsyellowish.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Massif Ruwenzori, Kalonge, 2,600 m, Ruiss Karumbura, affl. Katauleko,30.i.-21.ii.l953 (P. Vanschuytbeock & J. Kekenbosch) (MRAC). Paratypes. Zaire: 5 $ (MRAC). REMARKS. This species is very similar to rutshuris but can be distinguished by the antennal and gastralcharacters listed in the key. Rhoptromeris attis sp. n. (Figs 246, 256, 257) DESCRIPTION. $ . Antenna 13-segmented, segment 3 longer than 4, 4 longer than 5, 5 and 6 equal in length,7-13 forming a distinct club (Fig. 256). Head, viewed frontally, with eyes as far apart measured medially asheight of an eye, a row of hairs extending from outside antennal sockets to clypeus, malar grooves distinct,maxillary palp 4-segmented, labial palp 2-segmented. Pronotal plate with two fovea, one on either side ofmedial bridge joining anterior and posterior parts of plate, anterior part protruding forward (cf. Fig. 246).Pronotum either side of pronotal plate pubescent. Mesoscutum smooth, polished, notaulices absent, insome specimens a row of hairs in their place; scutellar disc rounded apically, lateral and hind margins ofdisc weakly striate, scutellar cup oval, apex with a large round fovea, base with minute fovea, scutellarfoveae kidney-shaped, polished, lateral bars of scutellum short, broad, polished. Mesopleural suturedistinct, metapleura with three distinct ridges, anteroventral cavity bare, propodeal carinae bowedmedially, either side of carinae pubescent. Segment 1 of gaster in the form of a weakly crenulated ring,obscured by a ring of pubescence at base of tergite 1 , complete though sparse on dorsal surface , tergite 2 thelargest in lateral view, occupying most of surface area of gaster, apex of tergite 2 and whole of visible part of3 punctate, hypopygium weakly swollen medially, coxae, femora, tibiae and tarsi pubescent, tarsi5-segmented. Wing surface pubescent, with apical hair fringe, forewing broad apically, radial cell a littleantenna yellowish basally, apical segments darker, head and thorax chestnut-brown, gaster light reddishbrown, legs yellowish. Cf . Antenna 15-segmented, filiform, segment 4 the largest, narrow basally, swollen and twisted medially(Fig. 257). MATERIAL EXAMINED Holotype $, Zaire: Makpe/8, 5.xi.l951 (H. de Saeger) (MRAC).Paratypes. Zaire: 2 $, 7 cf (MRAC). REMARKS. Very similar to bupalus but separated by the gaster, the anteroventral cavity and the clubsegments. 296 J. QUINLAN Rhoptromerisbicolorsp. n. (Figs 246, 247) DESCRIPTION. $ . Antenna 13-segmented, segment 3 longer than 4, 4 longer than 5, 5 and 6 subequal, clubsegments 7-13 sharply indicated, rhinaria prominent (Fig. 247). Head, viewed frontally, with eyes as farapart measured medially as height of an eye, a row of hairs present where frontal carinae would normallyextend to clypeal area, clypeus and mandibles pubescent, malar grooves distinct, maxillary palp 4-segmented, labial palp 2-segmented, cheeks converging. Pronotal plate, viewed dorsally, protruding,medial bridge between anterior and posterior parts almost aberrant, a weak transverse area in its place,fovea either side closed laterally (Fig. 246) , tufts of pubescence present on either side of plate . Mesoscutumsmooth, polished, notaulices absent, lateral bars of scutellum smooth, scutellar fovea polished, scutellardisc dull with sparse broken radiating sculpture, absent in some specimens, apex rounded, scutellar cupbroad, elliptical, apex with a small fovea and a ring of smaller fovea around rim, scutellar disc with longscattered hairs. Mesopleural suture distinct, metapleura with weak ridges, anteroventral cavity with a fewshort hairs. Propodeal carinae almost parallel, lateral margins of propodeum pubescent. Segment 1 ofgaster short, broad, crenulate, almost obscured by a dense ring of hairs at base of tergite 2, tergite 2 thelargest, occupying whole of visible area in lateral view, tergites 3 and 4 not visible, gaster impunctate,hypopygium prominent, ventral spine long, broad apically in lateral view. Legs long, coxae swollenbasally, femora short, swollen medially, tibiae and tarsi long, legs overall with scattered pubescence, tarsi5-segmented. Wing surface pubescent, with apical fringe of hairs, forewing broad apically, radial cellclosed on wing margin, longer than broad, broader apically than at base, veins M and Rs+M not indicated.Colour: antenna yellow basally, dark apically, head and mesoscutum black, dorsal surface of gasterchestnut-brown, pronotum, including plate, mesopleura and lower half of gaster, bright orange-yellow.CF unknown. MATERIAL EXAMINED Holotype <J>, Zimbabwe: Salisbury, Chishawasha, iii.1980 (A. Watshani) (BMNH).Paratypes. Nigeria: 2 $ (BMNH). Zaire: 10 $ (MRAC). REMARKS. A distinctive and easily recognised species by the sharp colour contrasts of the various thoracicparts. Very closely related to rufulus but the colour pattern of the antenna of rufulus distinguishes it. Rhoptromeris bupalus sp. n. (Figs 250, 255) DESCRIPTION. 9 Antenna 13-segmented, segment 3 longer than 4, 4 longer than 5, 6 longer than 5, shorterthan 7, 7-13 forming a distinct club (Fig. 255), club segments with rhinaria. Head, viewed frontally, withinner margins of eye further apart measured medially than height of an eye, a row of scattered hairsextending from outer margins of antenna to anterior tentorial pits, cheeks and mandibles with scatteredhairs, malar grooves distinct, maxillary palp 4-segmented, labial palp 2-segmented, cheeks weaklyconvergent. Pronotal plate fovea either side of medial bridge between anterior and posterior parts closedon lateral margins (cf. (Fig. 250), pubescent on either side. Mesoscutum smooth, polished, notaulicesabsent, lateral bars polished, broad basally, scutellar foveae kidney-shaped, polished, scutellar discpolished, with scattered hairs, apex rounded, with radiating ridges, scutellar cup semi-oval with atransverse fovea apically. Mesopleural suture distinct, metapleura almost completely smooth, anteroven-tral cavity without hairs. Propodeal carinae converging towards metanotal plate, sparsely pubescent onouter margins, lateral margins of propodeum with tufts of pubescence. Segment 1 of gaster in the form of aring, obscured by ring of hairs at base of 2, ring of hairs sparse on dorsal surface, tergites 2-4 visible inlateral view, 3 and 4 densely punctate, hypopygium not pronounced, with scattered submarginal hairs,ovipositor short. Legs normal shape, coxae elongate, weakly swollen medially, sparsely pubescent. Wingsurface pubescent, with apical hair fringe, radial cell of forewing entirely closed on wing margin, veins Mand Rs+M not indicated. Colour: antenna yellow, gaster chestnut-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zimbabwe: Salisbury, Chishswasha, ii.1980 (A. Watsham) (BMNH). Paratypes. Kenya: 1 $> (BMNH). Uganda: 1 $ (BMNH). Zaire: 3 $ (MRAC). Zimbabwe: 1 $, samedata as holotype (BMNH). Reunion: 4 $ (S. Quilici). REMARKS. Closely related to attis but separated by the antennal segments and the presence of a tuft of hairsin the anteroventral cavity of the metapleuron. AFROTROPICAL EUCOILIDAE 297 Rhoptromeris cepheus sp. n. (Figs 246, 272) DESCRIPTION. $. Antenna 13-segmented, filiform, segment 3 shorter than 4, 5-13 subequal, 7-13 withrhinaria, medially swollen (Fig. 272). Head, viewed frontally, with inner margins of eyes convergent, a rowof widely spaced hairs along margins of supraclypeal area, malar grooves distinct, maxillary palp4-segmented, labial palp 2-segmented. Pronotal plate protruded, anterior and posterior parts joined by amedial bridge, fovea either side of bridge closed on lateral margins (cf. Fig. 246). Pronotum either side ofplate with tufts of pubescence. Mesoscutum smooth, polished, notaulices absent, lateral bars broad-based,smooth, polished, scutellar fovea smooth, polished, kidney-shaped, lateral margins of scutellar discpolished, apex rounded, surface with radiating ridges, scutellar cup semi-oval, apical half with a transversefovea and minute pits on basal and medial areas. Mesopleural suture distinct, parallel with pre-coxalsuture, metapleura ridged, anteroventral cavity bare. Propodeal carinae almost parallel, very weaklybowed medially, pubescent on lateral margins. Segment 1 of gaster crenulate, in form of a ring, segment 2of gaster with a ring of pubescence at base, complete on dorsal surface, tergites 2-4 of gaster visible inlateral view, tergite 2 the largest, smooth, impunctate, hypopygium pronounced, ovipositor short. Legslong, coxae elongate, femora short, swollen basally, tibiae and tarsi densely pubescent, tarsi 5-segmented.Wing surface pubescent, with apical hair fringe, radial cell of forewing entirely closed, veins M and Rs+Mnot indicated. Colour: antenna yellow basally, apical segments blackish, with rhinaria, head, thorax andgaster dark chestnut-brown, legs pale yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Kenya: Kisumu, Lake Victoria, xi.1979 (M. D. Croft) (BMNH).Paratype. Kenya: 1 $, same data as holotype (BMNH). REMARKS. Distinguished from rwanki by the apical segments of the antenna and the polished scutellar disc. Rhoptromeris connatus sp. n. (Figs 241, 245) DESCRIPTION. $. Antenna 13-segmented, segments 3 and 4 equal in length, 5 shorter than 4, 6 longer than5, 6-7 subequal, 8-13 forming a weak club, swollen medially, bearing rhinaria (Fig. 241). Head, viewedfrontally, with inner margins of eyes weakly converging, further apart than height of an eye measuredmedially, a row of hairs extending from outer margins of antenna to clypeal area, malar grooves distinct,clypeus and surface of mandibles with long scattered hairs, maxillary palp 4-segmented, labial palp2-segmented. Pronotal plate with lateral fovea either side of pronotal plate closed. Mesoscutum smooth,polished, notaulices absent, lateral bars of scutellum polished, scutellar fovea semi-circular, sculptured inpart, scutellar disc weakly conical apically, reticulate-rugose, scutellar cup oval with a transverse foveaapically (Fig. 245). Mesopleural suture distinct, bowed medially, metapleura ridged, anteroventral cavityobsolete. Propodeal carinae parallel, densely pubescent on lateral margins. Segment 1 of gaster obscuredby a ring of hairs at base of tergite 2, complete on dorsal surface, segment 1 crescent-shaped in lateral view,crenulate, tergite 2 the largest, occupying whole of visible surface in lateral view, impunctate, hypopygiumweakly produced, ventral spine very short. Legs long, thin, pubescent, coxae elongate, swollen basally,femora short, swollen medially, tibiae and tarsi long. Wing surface pubescent with apical hair fringe, radialcell of forewing entirely closed on wing margin, veins M and Rs+M indicated by folds. Colour: antennacompletely yellow, head, thorax and gaster light chestnut-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Mabanga, 23.ix.1952 (H. de Saeger) (MRAC).Paratypes. Zaire: 9 $ (MRAC). REMARKS. This species could be confused with those species of Trichoplasta with a closed radial cell andconical scutellum, but the sculpture of the scutellar disc and the venation distinguish it as a Rhoptromeris. Rhoptromeris crito sp. n. (Figs 232, 267, 274) DESCRIPTION. Antenna 13-segmented, filiform, segment 3 shorter than 4, 4 longer than 5, 5-8 subequalin length, 9-13 shorter than 8, 11-13 white, 5-13 each with rhinaria, forming a very weak club (Fig. 267), 298 J. QUINLAN segments 3-13 at least 4 x as long as wide. Head, viewed frontally, almost round, slightly longer thanbroad, eyes as far apart as height of an eye measured medially, weakly converging, a few widely spacedhairs present between margins of antenna and clypeus, anterior tentorial pits distinct. Pronotal plateprotruding, medial bridge between anterior and posterior parts distinct, fovea either side enclosedlaterally, either side of plate with tufts of pubescence (Fig. 232). Mesoscutum smooth, polished, anteriormedian lines and notaulices absent; lateral bars of scutellum polished, scutellar foveae kidney-shaped,scutellar cup a little longer than wide, with a transverse fovea on lower half, basal area of cup pitted,scutellar disc reticulate-rugose laterally, weakly ridged apically, rounded (Fig. 274). Mesopleural suturecomplete, metapleura weakly ridged, anteroventral cavity with short abraded hairs; lateral margins ofpropodeum sparsely pubescent, propodeal carinae almost parallel, very weakly bowed medially. Segment1 of gaster short, broad, crenulate, obscured by a ring of dense pubescence at base of tergite 2, complete ondorsal surface, tergite 2 the largest in lateral view, tergites 3 and 4 partially visible, gaster impunctate,hypopygium not pronounced, with long subapical setae. Wing surface pubescent, with apical fringe ofhairs, radial cell of forewing long, narrow basally, closed on wing margin, veins Rs+M weakly pigmentedbasally, obsolete towards apex. Legs long, slender, coxae elongate, weakly swollen medially, femoraweakly swollen basally, tibiae and tarsi long, slender. Colour: antenna dark brown-orange basally, brownmedially, apical segments light orange-yellow.Cf unknown. MATERIAL EXAMINEDHolotype $, Madagascar: Mandrake, ii.1944 (A. Seyrig) (MRAC). REMARKS. This is the only species so far recognised which has rhinaria on 9 apical segments of the antenna(Fig. 267). Rhoptromeris cubitalis sp. n. (Figs 261, 262, 263) DESCRIPTION. $ . Antenna 13-segmented, segments 3 and 4 equal in length, 5 shorter than 4, 5-13 subequalin length, club segments 7-13 with rhinaria, not sharply delineated. Head, viewed frontally, with innermargins of eye further apart measured medially than height of an eye , with a row of scattered hairs betweenouter margins of antenna and clypeus, anterior tentorial pits distinct, with a carina extending to clypeus,malar grooves distinct. Pronotal plate with two lateral foveae, one on either side of medial bridge betweenanterior and posterior parts, closed on lateral margins (Fig. 262), either side of plate with a tuft ofpubescence. Mesoscutum smooth, polished, anterior median lines and notaulices absent though indica-tions present below surface; lateral bars smooth, polished, scutellar fovea deep, scutellar disc with ridgesradiating out from narrow elliptical scutellar cup (Fig. 263) , cup with an apical fovea, disc truncate apically.Mesopleural suture distinct, metapleura polished, anteroventral cavity with a tuft of pubescence, areaclose to propodeum densely pubescent. Propodeal carinae parallel, pubescent. Segment 1 of gaster in theform of a crenulate ring, partially hidden by a dense ring of pubescence at base of tergite 2, complete ondorsal surface, tergites 2-4 visible in lateral view, impunctate, tergite 2 the largest, hypopygium notpronounced, ventral spine long. Legs long, slender, coxae weakly swollen basally, femora strongly swollenbasally, tibiae and tarsi long, slender, coxae and femora with sparse hairs, femora tibiae and tarsipubescent. Wing surface pubescent, with apical hair fringe, fore wings broad apically with radial cell closedon wing margin (Fig. 261), short, little longer than broad, veins M and Rs+M weakly pigmented basally,obsolete apically. Colour: antenna dark orange-yellow, head dark brown, thorax and gaster light brown,legs yellow.cT unknown. MATERIAL EXAMINED Holotype $, Zaire: Secteur Tahisberimu, Talya Nerd, 23.iii.1954 (P. Vanschuythbroeck & M. Synave)(MRAC). Paratype. Zaire: 1 $ (MRAC). REMARKS. This species is close to thales but the length of antennal segment 3 is equal to 4, the lateralmargins of the scutellar disc are polished and the radial cell is shorter than in thales. AFROTROPICAL EUCOILIDAE 299 Rhoptromeris diversa sp. n. (Figs 242, 244, 246) DESCRIPTION. $ . Antenna 13-segmented, segment 3 as long as 4-1-5,5 shorter than 6, 8-13 forming a distinctclub, rhinaria prominent (Fig. 242). Head, viewed frontally, with eyes further apart measured medially,than height of an eye, with scattered hairs in a line extending from outer edge of antennal socket to clypealarea, anterior tentorial pits distinct, malar grooves distinct, clypeus and mandibular area with longscattered hairs. Pronotal plate with two fovea, one either side of medial bridge between anterior andposterior parts, closed on lateral margins (cf. Fig. 246), either side of plate pubescent. Mesoscutumsmooth, polished, notaulices absent, lateral bars of scutellum broad basally, smooth, polished, scutellarfoveae half-moon-shaped, polished, scutellar disc weakly conical apically, surface rugose, scutellar cupsemi-oval, apex with a transverse fovea, base pitted (Fig. 244). Mesopleural suture distinct, metapleuraridged, anteroventral cavity polished. Propodeal carinae parallel, sides of propodeum weakly pubescent.Segment 1 of gaster in the form of a crenulate ring, partially hidden by ring of pubescence at base of tergite2, tergite 2 the largest in lateral view, segment 3 visible, gaster impunctate, hyopygium small, ovipositorshort. Legs short, robust, coxae and femora weakly swollen basally, tibiae stout apically, tarsi short, stout,coxae sparsely pubescent, tibiae and tarsi moderately pubescent. Wing surface pubescent, apical marginwith a fringe of hairs , f orewing narrow , radial cell closed on margin , longer than broad , veins M and Rs+Mnot indicated. Colour: antenna yellow basally, brownish apically, head, thorax and gaster light orange-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Cape Province, Somerset East, 23-3 l.xii. 1930 (R. E. Turner) (BMNH). Paratypes. South Africa: 1 <j>, same data as holotype (BMNH). Uganda: 1 $ (BMNH). Zaire: 7 $(MRAC). REMARKS. The absence of punctures on the gaster and the weakly conical scutellum separate this speciesfrom abba, and the lengths of the individual club segments separate it from connatus, see key. Rhoptromeris enna sp. n. (Figs 246, 259) DESCRIPTION. $. Antenna 13-segmented, segment 3 as long as 4+5, 4-6 subquadrate, 7-13 forming adistinct club (Fig. 259). Face, viewed frontally, with eyes as far apart as height of an eye measured medially,face with a row of scattered hairs extending from outer margins of antenna to clypeus, malar groovesdistinct, lower face with scattered hairs. Pronotal plate with two foveae, one on each side of medial bridgebetween anterior and posterior parts of plate, bridge narrow (cf. Fig. 246) , fovea closed on lateral margins.Pronotum either side of pronotal plate with tufts of pubescence. Mesoscutum smooth, polished, anteriormedian lines and notaulices absent; lateral bars of scutellum polished, scutellar disc reticulate-rugose-coriaceous, rounded apically, scutellar cup narrow, elliptical, apex with a large transverse fovea, basal halfsculptured, rim of cup broad. Mesopleural suture distinct, metapleura with a weak ridge on margins closestto propodeum, anteroventral cavity with a tuft of pubescence. Propodeal carinae weakly bowed medially,either side of propodeum pubescent. Segment 1 of gaster obscured by a ring of dense pubescence at base oftergite 2, tergite 2 the largest in lateral view, 3-4 partially visible, apex of tergite 2 and visible parts of 3 and4 punctate, hypopygium prominent, ventral spine short, subbasal hairs present. Legs normally prop-ortioned, mid and hind coxae swollen basally, femora swollen basally and medially, tibiae widest apically,tarsi long, slender, femora, tibiae and tarsi pubescent. Wing surface pubescent, with apical hair fringe,radial cell of forewings closed, longer than broad, veins M and Rs+M not pigmented, folds indicated.Colour: antenna yellowish basally, darker apically, head, thorax and gaster dark chestnut-brown, legsyellow. Cf unknown. MATERIAL EXAMINEDHolotype $, Zaire: Ruwenzori, Riv. Katunda, 1,600m, 8.1.1954 (//. Synave) (MRAC). REMARKS. This species is closely related to punctata but is distinguished by antennal segments 4-6 beingsubquadrate and by the sculpture of the scutellar disc. 300 J. QUINLAN Rhoptromeris equalis sp. n. (Figs 253, 290) DESCRIPTION. $. Antenna 13-segmented, segments 3-6 subequal in length, 7-13 with rhinaria, forming adistinctive club, each swollen medially (Fig. 253). Head, viewed frontally, with eyes converging weakly,narrowest at a point between inner margins of eyes, and less than height of an eye, with a row of scatteredhairs between outer antennal sockets and epistomal suture, anteroventral pits and malar grooves distinct,clypeus and mandibular area with sparse pubescence. Pronotal plate with a transverse groove betweenanterior and posterior parts of plate, medial bridge not indicated (Fig. 290), lateral margins of grooveclosed. Mesoscutum smooth, polished, notaulices absent, lateral bars of scutellum small, polished,scutellar fovea oval, polished; scutellar disc with radiating ridges, apex rounded. Mesopleural suturedistinct, metapleural ridges on ventral margins, anteroventral cavity bare. Propodeal carinae bowedmedially, densely pubescent on lateral margins. Segment 1 of gaster not visible, obscured by a dense ring ofhairs at base of tergite 2, tergite 2 the largest, 3 and 4 partially visible in lateral view, gaster impunctate,hypopygium small, ventral spine not exposed. Legs normal, coxae elongate, swollen basally, femoraswollen basally, tibiae swollen apically, tarsi slender. Wings long, narrow, surfaces pubescent, marginswith a fringe of hairs, radial cell of forewing closed on margin, slightly longer than broad, veins M andRs+M not pigmented, only folds indicated. Colour: antenna orange-yellow basally, apical segmentsdarker, head and thorax dark chestnut-brown, gaster chestnut-red, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Cameroun: Nkoemvon, 29.vii.1979 (D. Jackson) (BMNH).Paratypes. Cameroun: 2 $, same data as holotype (BMNH). Zaire: 3 $ (MRAC). REMARKS. Distinguished frompersius by the distinct club with pronounced rhinaria on each segment. Rhoptromeris hebe sp. n. (Fig. 250) DESCRIPTION. $. Antenna 13-segmented, segments 3+4 subequal, 5 shorter than 4, 6-13 forming a weakclub, each segment with rhinaria. Head, viewed frontally, with inner margins of eyes further apart thanheight of an eye measured medially, face with scattered hairs, anterior tentorial pits distinct, malar groovesdistinct, clypeus and mandibles sparsely pubescent. Pronotal plate with a transverse groove mediallybetween anterior and posterior parts of plate, medial bridge between both parts not apparent (cf. Fig. 250),groove closed on lateral margins, either side of plate with tufts of pubescence. Mesoscutum smooth,polished, scutellar foveae half-moon-shaped, polished; lateral bars of scutellum narrow, polished;scutellar disc rounded apically, lateral surfaces smooth, with radiating ridges apically. Mesopleural suturedistinct, metapleura weakly ridged, anteroventral cavity with a minute tuft of hair. Propodeal carinaeparallel, almost as far apart as their length, with tufts of pubescence on outer margins. Segment 1 of gastercrescent-shaped, viewed laterally, crenulate, partially obscured by a ring of dense pubescence at base oftergite 2, complete on dorsal surface, tergite 2 the largest, 3 and 4 partially visible in lateral view, gasterimpunctate, hypopygium broad basally, ventral spine short, with sparse subbasal hairs. Legs normal, midand hind coxae swollen, hind coxae elongate, femora swollen basally, tibiae broader apically, allpubescent, tarsi 5-segmented. Wings narrow, surfaces pubescent, apical margins with a fringe of hairs,radial cell of forewing closed on margin, longer than broad, veins M and Rs+M not pigmented, weaklyindicated by folds. Colour: antenna yellowish basally, club segments darker, head, thorax and gasteryellowish brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: 18.ix.1952 (H. deSaeger) (MRAC).Paratypes. Zaire: 10 $ (MRAC). REMARKS. Closely resembles navius but distinguished by the pronotal plate (see key). Rhoptromeris heptoma (Hartig) (Figs 7, 224, 225, 226)Cothonospis heptoma Hartig, 1840: 201. Holotype $, GERMANY (ZSBS). AFROTROPICAL EUCOILIDAE 301 Cothonaspis eucerus Hartig, 1841: 357. Holotype $, GERMANY (ZSBS). [Synonomy by Cameron, 1890:206; Hellen, 1960: 8.] DESCRIPTION. $. Antenna 13-segmented, segment 3 longer than 4, 4 longer than 5, 5 and 6 subequal inlength, 7-13 forming a distinct club (Fig. 7). Face, viewed frontally, with eyes further apart than height ofan eye measured medially, face with a row of hairs extending from outer margins of antennal sockets toclypeus, malar grooves distinct, supraclypeal area and mandibular area with sparse scattered hairs.Pronotal plate with two fovea, one each side of medial bridge between anterior and posterior parts of plate,either side of plate with a tuft of pubescence on lateral margins of pronotum. Mesoscutum smooth,polished, notaulices absent, sparse hairs along margin with pronotum; lateral bars of scutellum polished,scutellar foveae almost round, scutellar disc punctate-reticulate to almost smooth, rounded apically,scutellar cup longer than broad, elliptical (Fig. 226). Mesopleural suture strongly bowed medially,metapleura mainly smooth with a few ridges, anteroventral cavity with a tuft of hairs. Propodeal carinaebowed medially, pubescent laterally. Segment 1 of gaster obscured by a ring of dense pubescence at base oftergite 2, only a crescent-shape apparent, tergites 2-4 visible in lateral view, impunctate, hypopygium notprominent, ventral spine short, subbasal hairs present. Legs normal, mid and hind coxae swollen, slender,with tufts of hair, femora swollen basally, tibiae widest apically, slender, all segments of legs weaklypubescent. Wing surfaces pubescent, margins with apical hair fringe, radial cell of forewing closed onmargin, Rs 2 and Rs^ of equal length, M and Rs+M indicated (Fig. 224). Colour: antenna yellow basally,dark apically, head, thorax and gaster shining black-brown, legs yellowish. Cf . Antenna 15-segmented, segment 3 curved on outer margin, weakly swollen apically, segment 4 twicelength of 3, curved, swollen, 5-15 filiform (Fig. 225). MATERIAL EXAMINEDSouth Africa: 3 $ (BMNH). Zaire: 3 $ (MRAC). REMARKS. R. heptoma is a parasite of Chloropidae (Nordlander, 1978). The specimens from South Africaand Zaire are very similar to European heptoma but differ very slightly in colour and the form of thepronotal plate. Rhoptromerisnaviussp. n. (Figs 275, 276) DESCRIPTION. $. Antenna 13-segmented, segments 3 and 4 subequal in length, 5 shorter than 4, 6-13forming a club, rhinaria prominent. Head, viewed frontally, with eyes weakly convergent, further apartmeasured medially than height of an eye, a row of scattered long hairs extends from outer margin ofantennal sockets to clypeal area, clypeus and mandibles with sparse hairs, anterior tentorial pits and malargrooves distinct; pronotal plate, viewed frontally, angular laterally, anterior and posterior parts separatedmedially by a bridge with a fovea on either side, closed on lateral margins (Fig. 275), lateral margins eitherside of pronotal plate with tufts of pubescence. Mesoscutum smooth, polished, notaulices absent, in theirplace a row of sparse hairs; lateral bars of scutellum polished, scutellar fovea lenticular in shape, smooth,polished, disc reticulate-rugose, scutellar cup elliptical, a little longer than broad, rim distinct, apex with alarge fovea, apical margin of disc rounded. Mesopleural suture distinct, metapleura with longitudinalridges, anteroventral cavity with a tuft of hairs. Propodeal carina parallel, weakly pubescent medially,strongly pubescent laterally. Segment 1 of gaster partially visible in lateral view, in the form of a crenulatering, tergite 2 of gaster with a dense ring of pubescence basally, entire on dorsal surface, tergites 2-4 ofgaster visible in lateral view, tergite 2 the largest, apex of 2 and visible parts of 3 and 4 punctate,hypopygium not pronounced, ventral spine short, subbasal hairs present. Legs long, slender, mid and hindcoxae weakly swollen, femora swollen basally, tibiae broadest apically, tarsi 5-segmented. Wings relativelylong, slender, surface pubescent, apical margins with a fringe of hairs, radial cell of forewing closed onmargin, narrow basally, veins M and Rs+M not indicated (Fig. 276). Colour: antenna yellow basally,median segments blackish brown, apical segments yellowish, head and thorax blackish, gaster chestnut-brown, legs yellow.C? unknown. MATERIAL EXAMINED Holotype $, Zaire: 26.ix.1951 (H. deSaeger) (MRAC).Paratypes. Kenya: 2 $ (BMNH). Zaire: 4 $ (MRAC). Zimbabwe: 2 $ (BMNH). REMARKS. This species has an 8-segmented club with a distinctive arrangement of colour; this and therelative lengths of the antennal segments enable it to be easily separated from other species. 302 J. QUINLAN Rhoptromeris naxos sp. n. (Figs 250, 268, 273) DESCRIPTION. $ . Antenna 13-segmented, segment 3 longer than 4, 4 and 5 subequal, 5 shorter than 6, 6-13forming a club, rhinaria prominent (Fig. 268). Head, viewed frontally, with eyes further apart measuredmedially than height of an eye, face with a row of scattered hairs extending from outer edges of antennalsockets to anterior tentorial pits, malar grooves distinct, mandibles with sparse hairs. Pronotal plate,viewed frontodorsally, without a bridge between anterior and posterior parts, separated medially by atransverse groove closed laterally, lateral margins of plate angular (cf. Fig. 250), either side of plate withtufts of pubescence. Mesoscutum smooth, polished, with a row of hairs in place of notaulices; lateral bars ofscutellum polished, scutellar fovea kidney-shaped, scutellar disc weakly sculptured laterally, apex weaklyrounded, rugose (Fig. 273). Mesopleural suture distinct, metapleura strongly ridged on ventral margin,anteroventral cavity without hairs. Propodeal carinae bowed medially, lateral margins of propodeumweakly pubescent. Segment 1 of gaster, viewed laterally, wider than long, in the form of a crenulate ring,tergites 24 visible in lateral view, tergite 2 the largest with a ring of pubescence at its base, apex of 2 andvisible parts of 3 and 4 punctate, hypopygium not pronounced, ventral spine not projecting. Legspubescent, with long mid and hind coxae, swollen from base to midway to apex, femora short, swollenbasally, tibiae broad apically, tarsi 5-segmented. Wing surfaces pubescent, with apical fringe of hairs,radial cell of forewing closed on margin, veins M and Rs+M absent. Colour: antenna yellow basally, apicalsegments darker, except segment 13 yellow, head, thorax and gaster reddish chestnut-brown. Legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kanayabayonga, Kubasha, 1,760 m, 7.xii.l934 (G. F. de Witte) (MRAC).Paratypes. Uganda: 1 $ (BMNH). Zaire: 13 (MRAC). REMARKS. This species is separated by the antennal segment ratios (Fig. 268). Rhoptromeris oeta sp. n. (Figs 230, 231, 250) DESCRIPTION. $ . Antenna 13-segmented, segment 3 shorter than 4, 4 longer than 5, 5-6 subequal in length,7-13 weakly swollen medially, with rhinaria, the whole filiform, club segments not sharply defined (Fig.230). Head, viewed frontally, with eyes converging, as far apart ventrally as height of an eye measuredmedially, scattered hairs extending from antennal sockets to clypeal margin, malar grooves distinct,anterior tentorial pits distinct. Pronotal plate with a transverse groove between anterior and posteriorparts, closed on lateral margins (cf. Fig. 250). Mesoscutum smooth, polished, notaulices absent; lateralbars of scutellum smooth, polished, scutellar foveae lenticular, lateral margins of disc weakly sculptured,apex with radiating sculpture, apical margin rounded (Fig. 231). Mesopleural suture distinct, metapleuraridged, sparsely pubescent on upper margin, anteroventral cavity with tuft of pubescence; propodealcarinae parallel, sides of propodeum pubescent. Segment 1 of gaster obscured by a ring of densepubescence at base of tergite 2, visible in lateral view, tergite 2 the largest, gaster impunctate, hypopygiumpronounced, ventral spine with subbasal hairs. Legs long, slender, coxae elongate, weakly swollen basally,narrow apically, femora swollen basally, apical half narrow, tibiae as long as tarsi. Wings broad apically,surface pubescent, apical margins with a fringe of hairs, radial cell of forewing closed on margin, veins Mand Rs+M indicated, weakly pigmented. Colour: antenna dark orange-yellow, apical segments samecolour as basal segments, in some specimens yellowish apically, head, thorax and gaster chestnut-brown,gaster orange-yellow, legs yellow.$ unknown. MATERIAL EXAMINED Holotype 9, Zaire: P.N.A., 30.i.-21.xii.l953 (P. Vanschuytbroeck&J. Kekenbosh) (MRAC).Paratypes. Zaire: 10 $ , same data as holotype except dates (MRAC). REMARKS. A very distinctive species with thread-like antenna (Fig. 230). Rhoptromeris pagasasp. n. (Figs 231, 233, 278)DESCRIPTION. $ . Antenna 13-segmented, segment 3 shorter than 4, 4 and 5 equal in length, 6-13 subequal AFROTROPICAL EUCOILIDAE 303 in length, with rhinaria, not forming a distinct clavate shape (Fig. 278). Face, viewed frontally, with eyesmeasured medially as close together as height of an eye, face with sparse scattered hairs, malar groovedistinct, mandibular area with sparse hairs. Pronotal plate with a transverse furrow between anterior andposterior parts, furrow closed on lateral margins (Fig. 233); either side of plate with tufts of pubescence.Mesoscutum smooth, polished, notaulices absent; lateral bars of scutellum polished, scutellar foveakidney-shaped, polished; scutellar disc with aberrant sculpture laterally, almost smooth, rugose apically,scutellar cup elongate, rim thick (cf. Fig. 231), apex of disc rounded. Mesopleural suture distinct,metapleura smooth, anteroventral cavity with a tuft of hairs. Propodeal carinae almost parallel, weaklybowed apically, lateral margins pubescent. Segment 1 of gaster, viewed laterally, crescent-shaped,partially obscured by a ring of dense pubescence at base of tergite 2, tergites 2-4 visible in lateral view,tergite 2 the largest, occupying almost whole visible area, segments 3 and 4 only partially visible, gasterimpunctate, hypopygium pronounced, visible part of ventral spine as long as base of hypopygium. Legsnormal, coxae longer than broad, swollen basally, femora swollen from base for two-thirds of length, tibiaewidest apically, meta-tarsi slender, longer than remaining combined tarsal segments. Wing surfacespubescent, apical margins with a fringe of hairs, radial cell closed on margin, longer than broad, veins Mand Rs+M not indicated. Colour: antenna yellow basally, 8 apical segments darker than basal segments,head, thorax and gaster orange-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype 9, Cameroun: Nkoemvon, malaise trap, ix.1979 (D. Jackson) (BMNH).Paratypes. Zaire: 10 $ (MRAC). Cameroun: 1 $, same locality as holotype, different date (BMNH). REMARKS. This species is closely related to temesa but can be separated by the relative lengths of antennalsegments 3 and 4 and the form of the pronotal plate. Rhoptr omens pallidussp. n. (Figs 246, 264) DESCRIPTION. $. Antenna 13-segmented, segment 3 as long as 4+5, 5 and 6 subequal in length, 7-13forming a distinct club. Head, viewed frontally, with eyes further apart measured medially than height ofeye, eyes diverging posteriorly, face with sparse hairs extending from base of antennal sockets to clypealarea, malar grooves distinct, anterior tentorial pits not clearly defined, apex of clypeus with long scatteredhairs extending over mandibles. Pronotal plate with two medial fovea between anterior and posteriorparts, separated by a broad medial bridge, closed on lateral margins (cf. Fig. 246). Mesoscutum smooth,polished, notaulices absent; lateral bars of scutellum polished basally, broader than long, scutellar foveapolished, scutellar disc rounded apically, surface with sparse weak radiating striae (very weak in somespecimens); scutellar cup raised, long, narrow, rim almost as wide as inner surface of cup. Mesopleuralsuture distinct, metapleura with weak ridges, anteroventral cavity with minute hairs. Propodeal carinaeconverging basally, weakly bowed apically, propodeum pubescent medially, with tufts of pubescence onlateral margins. Segment 1 of gaster obscured by a ring of hairs at base of tergite 2, tergite 2 the largest inlateral view, tergites 3-4 partially visible, apex of 2 and visible parts of 3 and 4 finely punctate, hypopygiumbroad, with subbasal hairs, ventral spine short. Legs normal, mid and hind coxae elongate, swollen basally,shorter than the femora, femora widest apically, shorter than tarsis. Wing surfaces pubescent, with a longapical fringe of hairs on apical margins, radial cell of forewing closed on margin, veins strongly pigmented,M and Rs+M absent. Colour: antenna and legs pale yellow, head, thorax and gaster light orange-yellow (insome specimens apical segments of antenna darker than basal segments). Cf . Antenna 15-segmented, segment 3 shorter than each of following segments, 4th curved and swollenapically (Fig. 264). MATERIAL EXAMINED Holotype $, Nigeria: Ibadan (B. R. Critchley) (MRAC).Paratypes. Nigeria: 5 $, 4 cf same data as holotype (BMNH). Zaire: 1 $ (MRAC). REMARKS. This species is not closely related to any of those described in this paper. The fovea on thepronotal plate appear closed but are not well defined. 3U4 J. QUINLAN Rhoptromeris persius sp. n. (Figs 250, 251, 252) DESCRIPTION. $ . Antenna 13-segmented, segment 3 longer than 4, 4-6 subequal in length, 7-13 forming aweak club, the segments swollen medially (Fig. 251). Face, viewed frontally, with eyes as far apartmeasured medially between eyes as height of an eye, sparse scattered hairs extending from outer marginsof antennal sockets to clypeus, malar grooves distinct, lower face with scattered hairs. Pronotal plate with atransverse groove between anterior and posterior parts, closed on lateral margins (cf. Fig. 250).Mesoscutum smooth, polished, notaulices absent; lateral bars of scutellum polished basally, weaklystriated laterally, scutellar fovea polished, kidney-shaped, scutellar disc with radiating striae, apexrounded, scutellar cup longer than broad, polished, rim light-coloured, apex of cup with a fovea.Mesopleural suture distinct, metapleura weakly ridged, anteroventral cavity with a distinct tuft of hairs.Propodeal carinae weakly bowed apically, lateral margins pubescent. Segment 1 of gaster in the form of acrescent-shaped ring, crenulate, tergite 2 the largest in lateral view, impunctate, with a ring of pubescenceat its base, tergite 3 only just visible, hypopygium not prominent, ventral spine very short. Legs normal,mid and hind coxae swollen basally, longer than broad, trochanters long, femora swollen basally, tibiaewidest at apex. Wing surfaces pubescent, margins with a fringe of hairs, radial cell of forewing closed onmargin, long, narrow (Fig. 252), veins M and Rs+M absent. Colour: antenna orange-yellow basally,brownish apically, head and thorax reddish brown, gaster orange-brown, legs yellow.C? unknown. MATERIAL EXAMINEDHolotype $, Zaire: 25.vi.1951 (J. Verschurch) (MRAC). REMARKS. Only one specimen of this species has been seen. It is close to equalis but separated by theantennal characters, see key. Rhoptromeris punctata sp. n. (Figs 239, 246, 248) DESCRIPTION. $. Antenna 13-segmented, segment 3 as long as 4+5, segment 4 longer than 5, each morethan twice as long as wide, 7-13 forming a distinct club (Fig. 248). Face, viewed frontally, with eyes furtherapart than height of an eye measured medially, face with a row of hairs extending from outer margins ofantenna to clypeus, malar grooves distinct, lower face with scattered hairs. Pronotal plate with two fovea,one either side of medial bridge between anterior and posterior parts of plate , closed on lateral margins (cf.Fig. 246). Pronotum either side of plate with sparse hairs. Mesoscutum smooth, polished, notaulicesabsent, a row of scattered hairs in their place, lateral bars of scutellum smooth, polished, scutellar discpolished, rounded apically, cup narrow, elliptical. Mesopleural suture weak though complete, metapleurawith a few weak impressions, anteroventral cavity with a tuft of hairs. Propodeal carinae parallel,pubescent on lateral margins. Segment 1 of gaster in the form of a short crenulate ring, tergite 2 the largest,with a ring of pubescence at base, not complete on dorsal surface, tergites 2-4 of gaster visible in lateralview, visible parts of 3 and 4 punctate, hypopygium weakly protruding, ventral spine short. Legs of normalproportions, coxae weakly swollen basally, femora medially swollen, tibiae broad apically, tarsi short,equal in length to tibiae; femora, tibiae and tarsi with sparse pubescence. Wing surface pubescent, with along apical fringe of hairs on frontal and apical margins, radial cell of forewing closed on margin, veinsthick, cell small (cf. Fig. 239), veins M and Rs+M not indicated. Colour: antenna yellowish, apicalsegments darker, head, thorax and gaster chestnut-red-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kivu, Sake, v.1938 (/. Ghesquiere) (MRAC).Paratypes. Zaire: 2 $ (MRAC). REMARKS. Very closely related to enna but distinguished by antennal segments 4-6 and the scutellar disc. Rhoptromeris rufulussp. n. (Figs 246, 258, 260) DESCRIPTION. $?. Antenna 13-segmented, segment 3 longer than 4, 4 longer than 5, 5 subequal to 6, 7-13forming a club (Fig. 260). Head, viewed frontally, with eyes converging closer together on approach to AFROTROPICAL EUCOILIDAE 305 lower face, face with a line of scattered hairs extending from outer margins of antennal sockets to clypealarea, malar groove distinct, lower face and frontal area with long scattered hairs. Pronotal platepronounced, anterior and posterior parts connected by a medial bridge with a fovea on either side ofbridge, closed on lateral margins (cf. Fig. 246); lateral margins of pronotum with a tuft of pubescence eitherside of plate. Mesoscutum polished, notaulices absent, a row of scattered hairs in their place, anteriorparallel lines weakly indicated; lateral bars of scutellum polished, scutellar fovea kidney-shaped, scutellardisc with weak radiating ridges laterally, stronger apically, apex rounded, scutellar cup semi-oval, apexwith a small fovea, basal area with minute pits. Mesopleural suture distinct, metapleura weakly ridged,anteroventral cavity with a few hairs. Propodeal carinae weakly bowed, lightly pubescent laterally.Segment 1 of gaster in form of a crenulate ring, partially obscured by a dense ring of hairs at base of tergite2, tergites 2-3 visible in lateral view, tergite 2 the largest, segment 3 punctate, hypopygium not prominent,ventral spine with subequal hairs, ovipositor short. Legs normal, coxae not noticeably swollen, femoraswollen basally, tibiae slender, widest apically, tarsi a little shorter than tibiae. Wing surfaces pubescent,margins with a fringe of hairs, radial cell of forewing closed, veins M and Rs+M not indicated. Colour:antenna yellow basally, darker apically except for segment 13 yellow, head and thorax dark brown, legsyellow. Cf . Antenna 15-segmented, segment 4 twisted, larger than 3 (Fig. 258). MATERIAL EXAMINED Holotype $, Zaire: 26.xi.1952 (H. de Saeger) (MRAC).Paratypes. South Africa: 5 $ (BMNH). Zaire: 49 $, 1 cf (MRAC). REMARKS. This species is distinguished from the closely related bicolor by the colour pattern of the antenna(see p. 296). The contrast of the colour pattern varies slightly, and the number of completely yellow apicalsegments also varies. It is possible that two species are involved. Rhoptromeris rwanki sp. n. (Fig. 272) DESCRIPTION. $ . Antenna 13-segmented, segment 3 shorter than 4, 4 longer than 5, 5 equal in length to 6,segments 7-13 forming a very weak club, each segment swollen medially, with rhinaria (cf. Fig. 272).Head, viewed frontally, with eyes further apart measured medially than height of an eye, a row of hairsextending from outer margin of antennal sockets to clypeal area, malar grooves distinct. Pronotal platewith two fovea, one either side of medial bridge connecting anterior and posterior parts. Pronotum eitherside of plate with tufts of pubescence. Mesoscutum smooth, polished, notaulices absent. Scutellumrounded apically, lateral bars polished dorsally, scutellar fovea shallow, smooth, polished, lateral marginsof scutellar disc with aberrant ridged sculpture, scutellar cup long, narrow, apex with a fovea, with smallpits alongside inner margin of cup rim. Mesopleural suture distinct, metapleura polished, anteroventralcavity with a tuft of pubescence. Propodeal carinae weakly bowed medially, densely pubescent mediallyand laterally. Segment 1 of gaster distinct, a little wider than long, crenulate, tergite 2 the largest in lateralview, remaining segments not visible, base of tergite 2 with a ring of dense pubescence, sparse on dorsalsurface, hypopygium not pronounced, ventral spine short, with subbasal hairs. Legs slender, pubescent,mid and hind coxae elongate, weakly swollen basally, femora weakly swollen basally, tibiae slender, broadapically, tarsi normal. Wing surfaces pubescent, with apical hair fringe, forewing rather blunt apically,radial cell of forewing closed on margin, almost as broad medially as long measured medially, veins M andRs+M indicated by weak pigmentation. Colour: antenna yellow basally, dark medially, apical 2-3segments yellowish, head, thorax and gaster dark chestnut-red, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: N. Lac Kivu, Rwanki, 15.ii.1952 (/. V. Leroy) (MRAC).Paratypes. Zaire: 8 9 (MRAC). REMARKS. This species is very similar to cepheus but the antennae are not white apically and the scutellardisc has weakly ridged sculpture. Rhoptromeris rutshuris sp. n. (Figs 236, 275) DESCRIPTION. $. Antenna 13-segmented, filiform, segment 3 shorter than 4, 4 and 5 subequal in length,6-13 with rhinaria, weakly swollen medially, apical segments white (Fig. 236). Face, viewed frontally, with 306 J. QUINLAN eyes converging towards clypeus, scattered hairs extending from antennal sockets to clypeal margin, malargrooves and anterior tentorial pits distinct, mandibles pubescent. Pronotal plate, viewed frontally, withlateral margins and dorsal margin straight, posterior and anterior parts fused medially by a narrow medialbridge separating two fovea which are closed on lateral margins (cf. Fig. 275). Mesoscutum smooth,polished, notaulices absent; lateral bars of scutellum broader than long, polished on dorsal surface,scutellar fovea semi-oval, polished, scutellar disc with weak radiating ridges laterally and apically, apicalmargin of disc rounded; scutellar cup slightly longer than wide, with a large fovea apically and smaller foveamedially, rim of cup not pronounced. Mesopleural suture distinct, metapleura with distinct ridges,antero ventral cavity with a tuft of pubescence. Mesopleural suture bowed apically, pubescent either side ofpropodeum. Segment 1 of gaster in the form of a crenulate ring, a little longer than wide, tergite 2 thelargest viewed laterally, with a ring of hairs at its base, tergite 3 partially visible, apex of tergite 2 and visibleparts of tergite 3 punctate, hypopygium not prominent, ventral spine short. Legs normal, femora swollen inbasal two-thirds, tibiae widest apically, as long as femora, shorter than tarsi. Wing surfaces pubescent, witha fringe of hairs on front and apical margins, radial cell of forewing closed on margin, narrow basally, veinsM and Rs+M not indicated. Colour: antenna yellowish basally, brownish medially, apical segments paleyellow, head, thorax and gaster orange-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype <j>, Zaire: Rutshura, xi.1937 (J. Ghesquiere) (MRAC).Paratypes. Cameroun: 2 $ (BMNH). Zaire: 38 $ (MRAC). REMARKS. Distinguished from the related species agis by the punctate segments of the gaster and the whiteapical segments of the antenna. Rhoptromeris sinis sp. n. (Fig. 279) DESCRIPTION. $ . Antenna 13-segmented, segments 3-6 subequal in length, 7-13 forming a weak club, eachclub segment with rhinaria (Fig. 279). Face, viewed frontally, with eyes converging, further apart thanheight of an eye, face with long sparse hairs scattered in area between antennal sockets and clypeus, malargrooves distinct. Pronotal plate with two laterally closed fovea, one on either side of medial bridge betweenanterior and posterior parts, outer parts of plate angular, either side of plate with tufts of pubescence.Mesoscutum smooth, polished, notaulices absent, scattered hairs present; lateral bars of scutellumpolished dorsally, scutellar foveae kidney-shaped, polished, scutellar disc rounded apically, surface withweak to strongly radiating sculpture; scutellar cup long, elliptical, apex with a small fovea, the anterior areawith transverse striae. Mesopleural suture distinct, metapleura ridged, anteroventral cavity bare. Pro-podeal carinae parallel, pubescent dorsally and laterally. Segment 1 of gaster broader than long, crenulate,tergite 2 the largest in lateral view, with a ring of dense pubescence at its base, complete on dorsal surface,segment 3 partially visible, gaster weakly punctate though not conspicuously, hypopygium weaklyprojecting, ventral spine with sparse hairs basally. Legs normal, mid and hind coxae elongate, weaklyswollen basally, pubescent medially, femora, tibiae and tarsi subequal in length, femora swollen in basalhalf, tibiae broader apically than basally, tibiae and tarsi pubescent with a hair fringe along margins. Wingsurfaces pubescent, with apical fringe of hairs, radial cell of forewing closed on margin, veins Rs longerthan 2r, veins M and Rs+M indicated as folds, with a weak trace of pigmentation. Colour: antenna paleyellow basally, darker apically, head, thorax and gaster reddish brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: N. Lac Kivu, Rwanki, 15.ii.1952 (/. V. Leroy) (MRAC).Paratypes. Cameroun: 1 (BMNH). Zaire: 17 $ (MRAC). REMARKS. This species is difficult to separate from bupalus and attis; however, the third antennal segment isequal in length to and not longer than the fourth as in the other species. Rhoptromeris tetnesa sp. n. (Figs 275, 282, 283, 284) DESCRIPTION. $. Antenna 13-segmented, segment 3 shorter than 4, 5 a little shorter than 4, 6-13 withrhinaria, the club only discernible by the presence of rhinaria, very weakly swollen medially (Fig. 282). AFROTROPICAL EUCOILIDAE 307 Head , viewed frontally , with eyes as far apart measured medially as the height of an eye , face with scatteredhairs, prominent around malar grooves, clypeus and mandibles. Pronotal plate with two fovea separatedby a medial bridge between anterior and posterior parts, with a tuft of pubescence on either side (cf. Fig.275). Mesoscutum smooth, polished, lateral bars polished, scutellar fovea polished, weakly sculptured;scutellar disc with radiating and ridged sculpture apically, apex rounded; scutellar cup large, elongate,broad, apex with a transverse fovea, basal half polished, with weak sculpture, rim of disc with a few longsetae (Fig. 283). Mesopleural suture distinct, straight, metapleura polished on upper margins, ridgedapically, anteroventral cavity with minute hairs. Propodeal carinae parallel, thick, pubescent laterally.Segment 1 of gaster completely hidden, ring of hairs at base of tergite 2 short and thick, tergite 2 the largestin lateral view, punctate, segments 3 and 4 not visible, hypopygium short, squarish apically, ventral spineshort. Legs slender, normal shape, sparsely pubescent. Wing surfaces pubescent, apical margins with afringe of hairs. Forewing broad apically, radial cell closed on margin, longer than broad (Fig. 284), veins Mand Rs+M not pigmented, visible as folds. Colour: antenna orange-brown basally , apical segments darker,head blackish red, thorax and gaster chestnut-red, legs yellow.Cf unknown. MATERIAL EXAMINEDHolotype $, Zaire: N. Lac Kivu, Rwanki, 15.ii.1952 (J. V. Leroy) (MRAC). REMARKS. A distinctive species which can be separated from rutshuris by the strong radiating sculpture ofthe scutellar disc and the broad median bridge on the pronotal plate. Rhoptromeris thalessp. n. (Figs 228, 265, 266) DESCRIPTION. $ . Antenna 13-segmented, segment 3 shorter than 4, 4 and 5 equal in length, 6 shorter than 5,7-13 with rhinaria, each segment wider apically than basally (Fig. 266). Face, viewed frontally, with eyes asclose together measured medially as height of an eye, face with sparse hairs between outer margins ofantennal sockets and clypeal area, malar grooves and anterior tentorial pits distinct. Pronotal plate withtwo fovea, one either side of medial bridge between anterior and posterior parts, lateral margins of foveaclosed (cf. Fig. 228), either side of plate with sparse hairs. Mesoscutum smooth, polished, notaulicesabsent; lateral bars of scutellum polished, scutellar fovea polished, kidney-shaped, wider than long;scutellar disc rounded apically, lateral margins with widely spaced radiating sculpture, apical surface thesame; scutellar cup long, narrow, with apical fovea. Mesopleural suture distinct, metapleura weaklyridged, anteroventral cavity with minute hairs. Propodeal carinae bowed medially, propodeum pubescenton dorsal and lateral surfaces. Segment 1 of gaster in the form of a crenulate ring, visible part in the form ofa crescent, tergite 2 the largest, base with a dense ring of hairs, tergites 3 and 4 partially visible, gasterimpunctate, hypopygium pronounced, ventral spine short. Legs slender, mid and hind coxae elongate,swollen basally, femora swollen basally, tibiae widest apically, longer than femora, slightly longer thantarsi. Wing surfaces pubescent, with apical fringe of hairs, forewing narrow, radial cell elongate, narrowbasally, weakly rounded (Fig. 265), closed on margin, veins M and Rs+M not indicated, folds only present.Colour: antenna yellow basally, apical 7 segments darker, head, thorax and gaster blackish brown, legsyellow. Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kivu, Rutshuru, Lubiriz, 1,285 m, 13.vii.1935 (G. F. de Witte) (MRAC).Paratypes. South Africa: 2 $ (BMNH). Zaire: 8 $ (MRAC). REMARKS. This species can be confused with those having a thread-like antenna, but the length to breadthratios of the segments are not as great and the club segments are discernible not only by the rhinaria beingdistinct but by the very weak but apparent clavate form of each segment (Fig. 266). Rhoptromeris velia sp. n. (Fig. 250) DESCRIPTION. 9- Antenna 13-segmented, segment 3 very slightly shorter than 4, almost equal in length, 5shorter than 4, equal in length to 6, 8-13 subequal in length, swollen medially, with distinct rhinaria. Head,viewed frontally, with eyes converging, further apart measured medially than height of an eye, with a rowof hairs extending from outer margin of antennal sockets to clypeal area, anterior tentorial pits and malargrooves distinct. Pronotal plate with a transverse groove medially between anterior and posterior parts of 308 J. QUINLAN plate, closed on lateral margins (cf. Fig. 250). Pronotum either side of plate pubescent. Mesoscutumsmooth, polished, notaulices absent. Scutellar fovea transversely long, kidney-shaped, deep, polished;lateral bars of scutellum polished; scutellar disc smooth laterally to weakly reticulate with radiating brokenrugose sculpture apically, scutellar cup almost oval, with a wide rim, apex with a small semi-circular fovea,inner margin of rim with a few small fovea or pits. Mesopleural suture distinct, metapleura weakly ridged,anteroventral cavity bare. Propodeal carinae subparallel, weakly converging apically, lateral margins ofpropodeum with dense tufts of pubescence . Segment 1 of gaster obscured by a ring of dense hairs at base oftergite 2, in the form of a crescent-shaped ring, crenulate; tergite 2 the largest, occupying almost whole ofvisible lateral area, tergites 3 and 4 only partially visible, gaster impunctate, hypopygium weakly visible,ventral spine short, with sparse subbasal hairs present. Legs slender, mid and hind coxae elongate, swollenmedially, femora swollen basally, tibiae widest apically, tarsi longer than tibiae. Wing surfaces pubescent,margins with a fringe of hairs, apex of wings broad, radial cell of forewing closed on margin, veins M andRs+M not indicated. Colour: antenna yellowish orange basally, apical segments darker, head, thorax andgaster chestnut-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, South Africa: Cape Province, Mossel Bay, 18-30.ix.1921 (R. E. Turner).Paratypes. South Africa: 1 $ (BMNH). Zaire: 2 $> (MRAC). REMARKS. This species is distinguished from afer by the shape of the club segments, which are weaklypronounced, whereas in afer the club is sharply defined. Rhoptromeris zetes sp. n. (Figs 249, 250) DESCRIPTION. $ . Antenna 13-segmented, segments 3-6 subequal in length, 7-13 forming a weak club, eachclub segment with rhinaria and clearly wider apically than basally. Face, viewed frontally, with eyes weaklyconverging, further apart measured medially than height of an eye, scattered hairs extending from outermargins of antennal sockets to clypeus. Pronotal plate with a transverse groove between anterior andposterior parts, closed on lateral margins (Fig. 250), lateral margins of pronotum with tufts of pubescenceeither side of pronotal plate. Mesoscutum smooth, polished, notaulices absent; lateral bars of scutellumpolished, scutellar disc smooth laterally, ridged apically, apical margin of disc rounded; scutellar cupelongate, narrow apically, rim of cup paler than inner surface, apex with minute fovea. Propodeal carinaeparallel, lateral margins of propodeum pubescent. Mesopleural suture distinct, metapleura with completetransverse ridges, anteroventral cavity with a tuft of short hairs. Propodeal carinae subparallel, weaklybowed apically, strongly pubescent laterally. Segment 1 of gaster in the form of a crenulate ring, partiallyobscured by a ring of short hairs at base of tergite 2, tergite 2 the largest in lateral view, punctate apically,visible parts of tergites 3 and 4 punctate, hypopygium small. Ventral spine short (Fig. 249). Legs normal,mid and hind coxae elongate, coxae swollen basally, shorter than femora, femora widest apically, tarsilonger than tibiae. Wing surfaces pubescent, apical margin with a hair fringe, radial cell of forewing closed,veins M and Rs+M weakly indicated by pigmentation. Colour: antenna yellow basally, 7-11 brownish,12-13 light yellow, almost white, head and thorax chestnut-brown, gaster dark orange- yellow, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Kivu, Nyongera, near Rutshuri, Butamba, 1,218 m, 17.vii.1935 (C. F. de Witte)(MRAC). Paratypes. Zaire: 9 $ (MRAC). REMARKS. This species is similar topersius but has fine punctures on tergites 2-4 of the gaster and the apicalsegments of the antenna are yellowish. Rhoptromeris zeus sp. n. (Fig. 229) DESCRIPTION. 9 . Antenna 13-segmented, segment 3 shorter than 4, 4 longer than 5, 5-7 each progressivelyshorter than preceding segment, 8-13 with rhinaria, swollen medially (Fig. 229). Face, viewed frontally,with eyes as far apart measured medially than height of an eye, face with a few sparse hairs, malar groovesand anterior tentorial pits distinct. Pronotal plate with a transverse groove medially between anterior and AFROTROPICAL EUCOILIDAE 309 posterior parts, closed on lateral margins, either side of plate with tufts of pubescence. Mesoscutumsmooth, polished, notaulices absent; lateral bars of scutellum polished, broad basally, scutellar foveaekidney-shaped, scutellar disc polished on lateral margins, apex rounded, apical surface with weak rugae;scutellar cup long, narrow, rim pale, apex with a small fovea. Mesopleural suture distinct, metapleura withweak ridges posteriorly, anteroventral cavity with an excrescence. Propodeal carinae parallel, pubescenton lateral margins. Segment 1 of gaster very slightly longer than broad, petiolate, in the form of a crenulatering, tergite 2 the largest in lateral view, with a dense ring of hairs at base, not complete on dorsal surface,tergites 3 and 4 partially visible, gaster impunctate, hypopygium pronounced, long, broad apically inlateral view, ventral spine short. Legs normal, mid and hind coxae elongate, swollen medially, femoraswollen in basal two-thirds, tibiae longer than femora, widest at apex, tarsi slightly longer than tibiae. Wingsurfaces pubescent, with long hair fringe on frontal and apical margins, radial cell of forewing closed onmargin, very narrow basally, veins M and Rs+M weakly indicated by pigmentation. Colour: antennayellowish basally, becoming yellowish brown apically, head, thorax and gaster orange-brown, legs yellow.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Massif Ruwenzori, Kalonge, 260m, Ruiss Karambura, affl. Katauleko, 30.i.-ii.l953(P. Vanschuytbroeck&J. Keknbosch) (MRAC). Paratypes. Zaire: 2 9 (MRAC). REMARKS. Separated from other species by the six-segmented club and the very pronounced hypopygium. STENTORCEPS Quinlan Stentorceps Quinlan, 1984: 479. Type-species: Stentorceps tubicen Quinlan, by original designation andmonotypy. DIAGNOSIS. $ antenna 13-segmented, segment 3 subequal to 4+5, 4 and 5 subequal in length, 7-13 forminga club (Fig. 281), darker than segments 1-6; Q" antenna 15-segmented, 4th segment grossly swollen (Fig.288). Head, viewed frontally, with two pyriform protuberances, one each side of face on inner orbits,occupying whole space between the half of area between eyes (Figs 31, 32) except for a medial furrow and anarrow gap between inner margins of protuberances and antenna. Supraclypeal area of face with a largeprotrusion shaped like the mouth of a trumpet, slightly narrower at its base (Fig. 32), mandibles very long,apically truncate, with scissors-like action (Fig. 289), clypeus long, narrow, with a few sparse setae (Fig.289). Pronotal plate projected forward, posterior and anterior parts separated medially by a transverseridge with two shallow foveae, not open on lateral margins. Segments 1-4 of gaster visible in $ , and 1-6 inC? , apical segments punctate (Fig. 286). Stentorceps tubicen Quinlan(Figs 31, 32, 281, 284, 285, 286, 287, 288, 289)Stentorceps tubicen Quinlan, 1984: 479. Holotype $>, KENYA (BMNH) [examined]. DESCRIPTION. $. Antenna 13-segmented, segment 3 subequal to 4+5, 4 and 5 subequal in length, 7-13forming a club (Fig. 281), darker than flagellar segments 1-6. Head, viewed frontally, as broad as long,eyes as far apart measured medially as height of an eye, the two protuberances occupying inner orbits,being separated medially, the trumpet-shaped protrusion on face with outer rim, viewed dorsally,light-coloured, lateral margins with a ring of setae extending past rim (Fig. 31); lower face, mandibles andclypeus with long scattered setae; pronotum either side of pronotal plate with tufts of pubescence.Mesoscutum smooth, polished, notaulices absent, in their place a row of hairs, scutellar foveae smooth,shallow, lateral bars smooth, scutellar disc polished laterally, apex weakly conical, surface with reticulate-rugose sculpture (Fig. 284), scutellar cup long, elliptical, with a large apical fovea. Propodeal carinaeparallel, converging apically, lateral margins of propodeum with tufts of pubescence. Mesopleura smooth,polished, mesopleural suture complete, metapleura ridged (Fig. 285), anteroventral cavity with a fewminute hairs (Fig. 285), open basally. Segment 1 of gaster in the form of a short strigose ring, partiallyobscured by a ring of hairs at base of tergite 2, tergite 2 the largest, viewed laterally, ring of hairs at baseincomplete on dorsal surface, hypopygium prominent, ovipositor short. Forewing broad, roundedapically, hindwing narrow, wing surfaces densely pubescent, apical margins with a fringe of hairs, radialcell of forewing closed on margin viewed with a Leitz stereo microscope [under the electron microscope itappears open, this is due to the presence of pigmentation (Fig. 287)]. Legs robust, coxae weakly swollen,with scattered pubescence, femora broad medially, margins of mid and hind coxae with a sparse hair fringe, 310 J. QUINLAN tibiae pubescent, tarsi 5-segmented. Colour: head and thorax blackish brown, gaster chestnut-red basally,otherwise brown, mandibles yellowish, antenna yellow basally, club segments brownish. O". Antenna 1 5-segmented, 4th segment grossly swollen (Fig. 288); segments 3-6 of gaster attachedtangentially to tergite 2 (Fig. 286). MATERIAL EXAMINED Holotype $, Kenya: Nairobi, Karen, 6,000m, i.l982-v.!982 (C. F. Dewhurst) (BMNH).Paratypes. Kenya: 7 $ , 5 cT, same data as holotype (BMNH). REMARKS. No other species have been seen that can possibly be assigned to this genus. TRICHOPLASTA Benoit Trichoplasta Benoit, 1956: 537. Type-species: Trichoplasta basilewskyi Benoit, by original designation andmonotypy. DIAGNOSIS. $ antenna 13-segmented, pubescent, with a 6-9-segmented club usually darker than basalflagellar segments, segment 3 generally shorter than 4, C? antenna 15-segmented, segment 3 shorter thanfollowing segments, segment 4 elongate, sometimes swollen distally (Fig. 308). Head, viewed frontally,rounded, frons raised, malar space with a subocular suture with carinae and weak striations on lower side.Pronotal plate similar to that of Rhoptromeris ', i.e. lateral fovea either side of median bridge betweenanterior and posterior parts closed (Fig. 300), medial bridge generally narrow. Scutellar disc produced,overhanging propodeum, scutellar cup with a large fovea on lower half. Segment 1 of gaster visible, notobscured by dense woolly ring of pubescence at base of tergite 2, apical segments usually punctate. Wingsurfaces densely pubescent, usually narrow, hair margins along apical margins long, radial cell of forewingopen or closed on margin, not deep. Segment 1 of gaster visible, not widened at either end, tergite 2 with adense hairy ring basally, apical segments of gaster generally with dense puncturation (Fig. 334). Legsusually slender. DISTRIBUTION World- wide except Neotropical region. REMARKS. This genus is closely related to Rhoptromeris and Stentorceps by the form of the pronotal platebeing common to all three genera, but is separated by the apomorphic scutellar prolongation. Key to the Afrotropical species of Trichoplasta Benoit Females 1 Radial cell of forewing open on front margin (Fig. 294); gaster punctate (Fig. 334); ring of pubescence at base of tergite 2 complete on dorsal surface (Fig . 334) 2 - Radial cell of forewing closed on front margin (Fig . 342) ; gaster impunctate ; ring of pubescence at base of tergite 2 not complete on dorsal surface 7 2 Antenna with a 7-segmented club (Fig. 303) 3 Antenna with 8-9-segmented club (Figs 328, 330) 6 3 Scutellar disc beak-shaped apically, surface punctate-reticulate laterally (Fig. 298), polished apically, long, narrow, scutellar cup large, oval, outer rim pale, lower half of cup with a largecircular fovea (Fig. 298); antennal club sharply defined (Fig. 303); pronotum smooth,polished rufus sp. n. (p. 319) - Scutellar disc beak-shaped or conical, surface reticulate-rugose (Fig. 296); scutellar cup variable in shape, antennal club either sharply or weakly defined 4 4 Antennal club sharply defined (Fig. 296) ,, 5 Antennal club weakly defined (Fig. 301), antenna short, not as long as distance between front of head and apex of gaster, segment 6 clearly shorter than 7; sides of pronotum polished medlia sp. n. (p. 317) 5 Antennal segment 3 shorter than 4+5, all segments at least 2-5 x as long as broad; veins Rs+M of fore wing not indicated ;metapleural region weakly crenulate equalissp. n. (p. 314) - Antennal segment 3 subequal to 4+5, all segments less than 2 x as long as broad, Rs+M distinct; metapleural region strongly ridged (cf. Fig. 334) tanganyikensis (Weld) (p. 319) 6 Antennal club 8-segmented (Fig. 330); ventral border of pronotum strongly striated; pronotal plate rounded on margins of posterior plate (Fig. 325) octonarius sp. n.(p. 318) AFROTROPICAL EUCOILIDAE 311 Antennal club 9-segmented (Fig. 328); ventral border of pronotum weakly sculptured;pronotal plate angled on margins of posterior plate (Fig. 324), rounded on dorsal margin novema sp. n. (p. 318) 7 Antennal segments 7-13 forming a weak 7-segmented club, each segment with rhinaria (Fig. 337).Scutellar disc tapering almost to a point (beak-shaped) (Fig. 338), reticulate-rugose bicolor sp. n. (p. 312)Antenna with 5-6-segmented club 8 8 Antennal club 5-segmented 9 - Antennal club 6-segmented 13 9 Apex of scutellar disc not beak-shaped (Fig. 320), surface with radiating sculpture; club segments of antenna sharply defined brevispina (Masner) (p. 313) - Apex of scutellar disc extended to form a spine or beak-shaped protrusion (Fig. 335), surface reticulate-rugose ; club segments of antenna weakly or sharply defined 10 10 Club segments of antenna sharply defined (Fig. 336) longispina (Masner) (p. 316) - Club segments of antenna weakly defined (Fig. 341) 11 11 Antenna completely pale yellow, segments 4-7 subequal in length, club segments very weakly swollen medially, with rhinaria, segment 4 as long or longer than 5 (Fig. 321) 12 - Antenna with club segments yellow-brown, never completely pale (Fig. 341); scutellum beak-shaped (Fig. 339) Sliformissp. n.(p. 315) 12 Head, thorax and gaster blackish brown; pronotum orange-yellow in sharp contrast; antenna distinctly longer than distance between front of head to apex of gaster; scutellar cup large,oval, scutellar disc rugose, apex acutely spined (Fig. 314); pronotal plate, viewed frontodor-sally, with anterior plate angled dorsally, medial bridge narrow (cf. Fig. 300) contrasts sp. n. (p. 314) - Head, thorax, gaster and pronotum blackish brown, antenna at most as long as head to apex of gaster, scutellar cup longer than broad, scutellar disc reticulate-rugose, apex of disc sharplybut not acutely spined (cf. Fig. 338); pronotal plate, viewed frontodorsally, with anteriorplate rounded on margins, not angled, medial bridge broad (Fig. 344) .... quinclava sp. n. (p. 319) 13 Scutellar disc conical, not spine-shaped at apex 14 - Scutellar disc spine-shaped or in the form of a beak 16 14 Club segments of antenna conspicuous , moniliform (Fig. 306) , apical segments twice as long as wide com'casp. n. (p. 313) - Club segments of antenna inconspicuous (Fig. 346) 15 15 Antennal segments 3 and 4 subequal in length (Fig. 346) gracilicornis (Kieffer) (p. 316) - Antennal segment 3 clearly shorter than 4 (Fig. 358) zeussp. n. (p. 321) 16 Antennal segment 3 as long as 4. Scutellar disc reticulate-rugose (cf. Fig. 338); pronotal plate with a broad medial bridgebetween anterior and posterior parts, posterior part striated (cf. Fig. 344) extensus sp. n. (p. 315) - Antennal segment 3 clearly shorter than 4 17 17 Scutellar disc areolate, cup small, with a small declined apical fovea (Fig. 350). Pronotal plate with a narrow medial bridge between anterior and posterior parts of plate;radial cell of forewing triangular in shape (Fig. 347), head, thorax, gaster and legs orange-yellow testaceasp. n. (p. 320) - Scutellar disc rugose-reticulate , scutellar cup large with apical fovea normal , not declined 18 18 Pronotal plate with a very weak bridge medially between anterior and posterior parts (cf. Fig. 351); apical segments of antenna dark narrate sp. n. (p. 317) - Pronotal plate with a distinct bridge medially between anterior and posterior parts (cf. Fig. 344); antenna completely pale yellow unicolora sp. n. (p. 321) Males 1 Radial cell of forewing open on front margin (Fig. 294) ; gaster punctate (Fig. 334) , with ring of hairs at base of tergite 2 complete on dorsal surface 2 - Radial cell of forewing closed on front margin (Fig. 342); gaster impunctate, ring of hairs at base of tergite 2 incomplete on dorsal surface 4 2 Pronotum striate on lateral margins (Fig. 295); antennal segment 3 subequal to 4, 4 swollen 312 J. QUINLAN distally, curved, strongly flattened medially on outer margin, viewed dorsally (cf. Fig. 291); gaster densely punctate equalis sp. n. (p. 314) Pronotum polished on lateral margins, antennal segment 3 distinctly shorter than 4 (cf. Fig.317) 3 3 Antennal segment 4 weakly swollen, not curved or flattened medially (Fig. 331); scutellar cup elliptical, longer than broad, scutellar disc reticulate-rugose, apex beak-shaped (Fig. 297) medlia sp. n. (p. 317) Antennal segment 3 subequal to 4, weakly swollen medially, flattened on outer medial margin(Fig. 317); scutellar cup oval, scutellar disc finely reticulate-rugose, apex conical (Fig. 298) rufussp. n.(p. 319) 4 Antennal segment 4 2-6 x length of segment 3, with very prominent rhinaria, 4-6 darker than basal segments, narrow basally and apically, weakly swollen medially (Fig. 304); scutellarcup almost oval, scutellar disc with radiating striae below scutellar fovea, apex of disc with a distinct spine narrate sp. n.(p. 317) Antennal segment 4 either curved, swollen medially or basally, or twisted, less than 2-3 xlength of 3; scutellar disc cone- or spine-shaped, viewed apically, surface reticulate-rugose,exceptionally with radiating striations 5 5 Antennal segment 4 of antenna twisted , flattened on outer margin , swollen apically (sometimes very swollen) (Fig. 308); scutellar disc reticulate-rugose, apex almost pointed (Fig. 307); pronotal plate weakly striated on anterior part (cf. Fig. 344) conica sp. n. (p. 313) Segment 4 of antenna swollen medially or basally, or if curved not strongly swollen at apex;scutellar disc conical or beak-shaped 6 6 Segment 4 of antenna, viewed laterally, flattened on outer margin, weakly swollen medially, narrow apically and basally (Fig. 311).Scutellar disc reticulate-rugose laterally, with weak striae, apex with a long narrow blunt spine (Fig. 310) ; scutellar cup longer than wide filiformis sp. n. (p. 315) Segment 4 of antenna swollen basally or curved, or weakly swollen apically; scutellar cupsemi-oval 7 Antennal segment 4, viewed laterally, swollen basally (Fig. 352); scutellar disc with radiating sculpture , apex with a narrow beak-like protuberance (cf. Fig. 350) unicolora sp. n. (p. 321) - Antennal segment 4 curved, not swollen (Fig. 313); scutellar disc reticulate-rugose, apex conical (Fig. 314) contrasta sp. n.(p. 314) Trichoplasta bicolorsp. n. (Figs 337, 338, 344) DESCRIPTION. $. Antenna 13-segmented, very weakly subclavate, the 7-segmented club not sharplydefined (Fig. 337), rhinaria visible on all club segments, each club segment weakly swollen medially. Head,viewed frontally, smooth, polished, with long sparse hairs, antennal sockets pronounced, anterior tentorialpits distinct, subocular sulcus distinct, occipital carina weak. Pronotal plate rounded laterally and dorsallyon posterior margins, anterior rounded on lateral margin, fovea either side of medial bridge closed onlateral margins (cf. Fig. 344), either side of plate with tufts of whitish pubescence. Mesoscutum smooth,polished, lateral bars of scutellum polished dorsally, scutellar fovea shallow, polished; scutellar discreticulate-rugose, tapering almost to a point (similar to a beak), scutellar cup small, elliptical, with a palerim, sculptured basally with a large apical fovea on apical third (Fig. 338). Lateral margins of pronotum,mesopleura and metapleura smooth, polished, mesopleural suture distinct, metapleura ridged, pubescentadjacent to hind coxa, nucha ridged, pubescent. Propodeal carinae weakly bowed, obscured by pubesc-ence. Segment 1 of gaster completely obscured by a dense woolly ring of hairs at base of tergite 2,incomplete on dorsal surface, tergite 2 impunctate, remaining tergites not visible in lateral or dorsal view,hypopygium clearly visible. Legs long, slender, coxae elongate, coxae, femora and tibiae pubescent. Wingsurface pubescent, with apical hair fringe, radial cell of forewing closed on wing margin, venation pallid,vein M (cubitus) not indicated. Colour: head and thorax brownish, gaster chestnut-red, legs orange-yellow, antenna yellow basally, dark apically.Cf unknown. MATERIAL EXAMINEDHolotype $, South Africa: Port St John, Pondoland, 12-30.vi.1923 (R. E. Turner) (BMNH). REMARKS. This species is distinguished by the 7-segmented club (see key). AFROTROPICAL EUCOILIDAE 313 Trichoplasta brevispina (Masner)(Figs 318, 320) Odonteucoila brevispina Masner, 1960: 357. Holotype $, KENYA (MRAC) [examined].Trichoplasta brevispina (Masner) Nordlander, 19820: 273. DESCRIPTION. $. Antenna 13-segmented, with a distinct 5-segmented club, segment 3 shorter than 4, 4longer than 5, 5 shorter than 6, 5-12 subequal in length, 13 as long as 3, apical 5 segments distinctly broaderthan preceding segments. Head, viewed frontally, with sparse scattered hairs, antennal sockets pro-nounced, viewed dorsally, subocular sulcus and malar suture distinct, with striae on either side, occipitalcarinae pronounced dorsally. Pronotal plate rounded posteriorly, viewed frontodorsally, anterior marginsangled laterally, medial bridge between anterior and posterior parts separated by a narrow bridge, foveaeither side of bridge closed on lateral margins. Mesoscutum smooth, polished, with widely scattered hairs,notaulices absent; lateral bars of scutellum polished, weakly sculptured; scutellar disc with radiating,ridged rugose sculpture, apex conical, not elongated to form a beak-like projection (Fig. 320), sparse hairspresent; scutellar cup small, elliptical, rim pronounced, inner surface depressed, sculptured with a smallapical fovea. Pro no turn smooth, polished, with pubescence either side of pronotal plate; mesopleuralsuture distinct, metapleura ridged basally with tufts of hair either side of propodeum. Propodeal carinaeparallel , nucha longitudinally ridged , pubescent . Segment 1 of gaster crenulate , base of tergite 2 with a ringof hairs, incomplete on dorsal surface, apex of tergite 2 impunctate, segments 3 and 4 not visible,hypopygium weakly projecting. Legs long, slender, coxae and femora with long sparse hairs. Wingsnarrow, surfaces pubescent, with apical hair fringe, radial cell of forewing closed on margin (Fig. 318),venation very pallid, vein M (cubitus) indicated, not pigmented. Colour: head and thorax brownish black,gaster chestnut-red, legs yellowish, coxa yellowish brown, antenna yellowish, club segments yellowishbrown. Cf unknown. MATERIAL EXAMINED Kenya: 1 $ (holotype), Molo (Mau Escarpment) 2150-2200 m, ll-12.vi.1957 (Basilewsky & Leleip)(MRAC). Zimbabwe: 1 $ (BMNH). REMARKS . This species is distinguished from gracilicornis by the sculpture of the scutellar disc and the shapeof the radial cell. Trichoplasta conica sp. n. (Figs 300, 306, 307, 308, 312) DESCRIPTION. $. Antenna 13-segmented with a distinct, moniliform, 6-segmented club, segment 3 as longas 4+5, 4 longer than 5, 5 shorter than 6, club segments distinctly wider than preceding flagellar segments(Fig. 306). Head, viewed frontally, with sparse scattered hairs, antennal sockets pronounced, subocularsuture (malar suture) distinct, with weak aberrant sculpture on either side, occipital carina distinct, withlong hairs on lateral margins. Pronotal plate, viewed frontodorsally, with posterior part with a flatteneddorsal margin, laterally rounded, anterior and posterior parts fused by a medial bridge, fovea on either sideclosed (cf. Fig. 300). Mesoscutum smooth, polished, anterior parallel lines indicated, notaulices absent,scattered hairs in their place; lateral bars of scutellum polished, scutellar foveae kidney-shaped; scutellardisc reticulate-rugose, conical at apex, not beak-shaped (Fig. 307), cup elliptical, rim pale, a fovea at baseand apex. Pronotum smooth, polished either side of pronotal plate; mesopleural suture distinct, meta-pleura ridged near junction with hind coxa. Propodeal carinae weakly bowed, strongly pubescent on eitherside, nucha ridged. Segment 1 of gaster crenulate, partially obscured by a ring of hairs at base of tergite 2,apex of tergite 2 with light punctures, tergites 3-5 partially visible in lateral view, punctate, hypopygiumpronounced (Fig. 312). Legs short, coxae and tibiae pubescent. Wings broad, surface pubescent, apicalhair fringe long, radial cell closed on wing margins, venation of radial cell strongly pigmented, vein M(cubitus) indicated, not pigmented. Colour: head, thorax and gaster dark chestnut-red-brown, legsorange-yellow, antenna yellowish basally, apical segments blackish orange-brown. Cf . Antenna 15-segmented, segments 3-15 with rhinaria, segment 3 shorter than 4, 4 shorter than 5,twisted and swollen apically (Fig. 308). MATERIAL EXAMINEDHolotype $, Zaire: Mont Hoyo, 1280 m, 7-15.vii.1955 (P. Vanschuytbroeck) (MRAC). 314 J. QUINLAN Paratypes. Kenya: 6 $ (BMNH). Nigeria: 1 9 (BMNH). Uganda: 4 $ (BMNH). Zaire: 153 $, 1 cf(MR AC). REMARKS. Like gracilicornis this species has a sharply defined 6-segmented club, the apical segments areless than 2 x as long as wide. Trichoplasta contrasts sp. n. (Figs 300, 313, 314, 321) DESCRIPTION. $. Antenna 13-segmented, filiform with a very weak 5-segmented club, segment 3 shorterthan 4, 4 longer than 5, 5-7 subequal in length, club segments 8-13 with rhinaria (Fig. 321). Head, viewedfrontally, with scattered hairs, antennal sockets pronounced, frontal region not raised, malar suturedistinct, not pronounced, occipital carina visible dorsally, obscured laterally by dense pubescence eitherside of pronotal plate. Pronotal plate, viewed dorsally, with anterior margin rectangular, posterior marginrounded, medial bridge between anterior and posterior parts of plate narrow, fovea either side closedlaterally (cf. Fig. 300). Pronotum smooth, polished laterally (orange in colour), mesopleural suturedistinct, metapleura weakly sculptured on lower margin; mesoscutum smooth, polished, with scatteredhairs in place of notaulices; lateral bars of scutellum smooth, scutellar foveae kidney-shaped; scutellar cuppear-shaped, with a large fovea apically, scutellar disc reticulate-rugose, apex acute, beak-shaped (Fig.314). Propodeal carinae parallel, weakly pubescent medially, densely pubescent on lateral margins, nucharidged. Segment 1 of gaster obscured by a ring of pubescence at base of tergite 2, 3 and 4 visible in lateralview, gaster impunctate, hypopygium not protruding. Legs long, slender, coxae elongate, moderatelypubescent. Wings densely pubescent, with apical fringe of hairs, radial cell of forewing closed on wingmargin, narrow basally, wide apically, venation dark yellow. Colour: head brownish black, pronotumorange-yellow, mesoscutum and scutellum blackish, gaster chestnut-brown. Legs yellowish orange,antenna yellowish orange. Cf . Antenna 15-segmented, filiform, segment 3 swollen basally, twisted medially (Fig. 313). MATERIAL EXAMINED Holotype $, Nigeria: Ondu State, 21.iii.1971 (/. T. Medler) (BMNH).Paratypes. Zaire: 16 $, 5 cf (MRAC). REMARKS. This species is closely related to quinclava but is distinguished by the contrasting colour patternof the head, pronotum and scutellum, and the shape of the pronotal plate (cf. Fig. 300). Trichoplasta equalissp. n. (Figs 291, 292, 296, 300) DESCRIPTION. $ . Antenna 13-segmented, clavate, segment 3 longer than 4, 4 longer than 5, 5 shorter than6, 7-13 forming a distinct club (Fig. 296). Head, viewed frontally, with strong frontal ridges in part,subocular sulcus distinct, antennal sockets protruding, occipital carina pronounced. Pronotal plate withanterior and posterior fused laterally and medially with a small fovea either side of medial bridge, plate,viewed fronto-dorsally, with anterior rounded laterally (cf. Fig. 300), posterior transversely striated.Pronotum either side of pronotal plate pubescent, with transverse striations extending to mesopleura,mesopleural suture distinct; metapleura with crenulate ridges, antero ventral cavity pubescent, side ofmetapleura adjacent to nucha densely pubescent; mesoscutum smooth, polished, notaulices absent;scutellar foveae kidney-shaped, smooth, lateral bars of scutellum smooth, polished, septum broad;scutellar disc reticulate-punctate, apex beak-shaped (cf. F\g. 292), scutellar cup large, almost round,viewed laterally, elevated, apical half with a large fovea; propodeal carinae parallel, weakly pubescentmedially, more densely pubescent on lateral margins; nucha ridged, pubescent; base of metapleurapubescent. Segment 1 of gaster not visible, obscured by a dense ring of pubescence at base of tergite 2,apical two-thirds of tergite 2 punctate, tergites 3 and 4 visible in lateral view, punctate; hypopygium notproduced. Legs moderate to short, coxae swollen, elongate, sparsely pubescent. Wing surfaces pubescent,with apical fringe of hairs, radial cell of forewing open on wing margin, deep, not elongate, venation darkyellow, vein M (cubitus) extending almost to apex of wing. Colour: head and thorax blackish, gasterreddish yellow, antenna blackish brown, legs orange-yellow. O" . Antenna 15-segmented, segment 3 shorter than 5 which is curved and swollen distally, flattened onouter side, viewed dorsally (cf. Fig. 291). AFROTROPICAL EUCOILIDAE 315 MATERIAL EXAMINED Holotype $, Zaire: Tshamugussa (Beweza) Bambous, 10.viii.1934 (G. F. de Witte) (MRAC).Paratypes. Nigeria: 1 cf (BMNH). Zaire: 36 $, 26 cf (MRAC). REMARKS. Distinguished from the closely related tanganyikensis by the antennal and sculptural characters,see p. 320. Trichoplasta extensus sp. n. (Figs 338, 344, 348) DESCRIPTION. $. Antenna 13-segmented, clavate, with a distinct 6-segmented club, segments 3-5 sub-equal, segments 8-13 with rhinaria forming a club, each segment clearly broader than apical segments.Head, viewed frontally, elongate, face smooth, polished, with long scattered hairs, subocular suture orsulcus distinct, antennal sockets weakly protruding. Occipital carina pronounced in dorsal view, obscuredlaterally by tufts of pubescence either side of pronotal plate. Posterior part of pronotal plate, vieweddorsally, straight, rounded laterally, anterior half diverging outwards, both parts joined laterally andmedially to enclose two foveae (cf. Fig. 344). Mesoscutum smooth, polished, with a few hairs in place ofnotaulices; lateral bars of scutellum polished in dorsal view, scutellar foveae angular, polished; scutellardisc reticulate-rugose, converging apically to form a spine or beak-shaped apex (cf. Fig. 338). Pronotumand mesopleura smooth, polished; mesopleural suture distinct, metapleura with traces of ridges on loweredge, with sparse hair tufts. Propodeal carinae parallel, weakly pubescent medially, denser on outermargins of carinae, nucha ridged, pubescent. Segment 1 of gaster visible, crenulate, partially obscured by aring of hairs at base of tergite 2, tergites 3 and 4 not visible in lateral view, gaster impunctate, hypopygiumsmall, not projected. Legs long, slender, coxae with lateral fringe of hairs, femora, tibiae and tarsipubescent. Wing surfaces pubescent, with apical fringe of hairs, radial cell of forewing closed on wingmargin, short, broad (Fig. 348). Colour: head and thorax blackish brown, gaster chestnut-brown, legsorange-yellow, antenna yellowish orange basally, apical segments darkened.Cf unknown. MATERIAL EXAMINED Holotype $, Zaire: Secteur Tshiaberimu, Mont Kitwa, 2840 m, 29.viii.-7.ix.1953 (P. Vanschuytbroeck& V. Hendrickx) (MRAC). REMARKS. Separated from narrata by the pronotal plate character and the antennal segment ratios. Trichoplasta filiformis sp. n. (Figs 309, 310, 311, 339) DESCRIPTION. $ . Antenna 13-segmented, filiform, with indistinct 5-segmented club, segment 3 shorter than4, 9-13 with rhinaria, forming a very weak club (Fig. 309). Head, viewed frontally, with frons raised, areaabove clypeus with long scattered hairs, antennal sockets protruding, subocular sulcus distinct. Occipitalcarina pronounced medially, fading laterally. Pronotal plate rounded on dorsal margin, angular on lateralmargin, fovea on either side of medial bridge enclosed laterally, i.e. posterior and anterior parts joinedlaterally (cf. Fig. 324). Pronotum pubescent on either side of pronotal plate, not striated; mesopleuralsuture distinct, ventral margin of mesopleura pubescent, metapleura pubescent on apical margin;mesoscutum smooth, polished, notaulices absent, a few scattered hairs in their place; lateral bars ofscutellum polished, scutellar fovea smooth, polished, scutellar disc reticulate-rugose, conical to beak-shaped viewed dorsally (Figs 310, 339), scutellar cup longer than broad. Propodeal carinae parallel,pubescent on either side, nucha ridged. Segment 1 of gaster obscured by a ring of pubescence at base oftergite 2, ring incomplete on dorsal surface, tergite 2 impunctate, smooth, polished, hypopygiumprotruding. Legs long, slender. Wing surfaces pubescent, with apical hair fringe, radial cell of forewingclosed on wing margin, vein M (cubitus) not indicated. Colour: head, thorax and gaster orange-brown,antenna yellow basally, apical segments brownish black, legs yellow. Cf . Antenna 15-segmented, segment 4 the largest, curved, weakly swollen medially (Fig. 311). MATERIAL EXAMINED Holotype $, South Africa: E. Cape Province, Katberg, 19-22.ii.1924 (R. E. Turner) (BMNH).Paratypes. South Africa: 1 $ (BMNH). Zaire: 13 ?, 15 cf (MRAC). REMARKS. Separated from closely related species by the colour pattern of the antenna and the ratio of thelengths of segments 3 and 4. 316 J. QUINLAN Trichoplasta gracilicornis (Kieffer)(Figs 344, 346) Coneucoela gracilicornis Kieffer, 1910a: 534. Holotype $, MADAGASCAR (ZM).Odonteucoila gracilicornis (Kieffer) Weld, 1944: 63; Masner, 1960: 356 (redescribed).Trichoplasta gracilicornis (Kieffer) Nordlander, 1982a: 272. DESCRIPTION. $ . Antenna 13-segmented filiform, club 6-segmented, segments 3 and 4 subequal in length, 5shorter than 4, segments 6-11 each subequal to 3, subequal to each other, 12 and 13 each fractionallyshorter than 6, equal to 3, club segments very slightly wider than preceding segments, not forming a distinctclub (Fig. 346). Head, viewed frontally, smooth, polished, subocular suture distinct, antennal socketspronounced, frons weakly raised, occipital carina distinct in dorsal view. Pronotal plate, viewed fronto-dorsally, weakly rounded on posterior margin, lateral margins straight, diverging, posterior and anteriorparts of plate bridged laterally and medially to enclose two foveae or pits (cf. Fig. 344), anterior areastriated. Mesoscutum smooth, polished, with a few scattered hairs, lateral lines (parapsidial furrows) veryweakly indicated apically; lateral bars of scutellum polished dorsally, scutellar foveae shallow, lenticular,scutellar disc reticulate-rugose, conical at apex, not spine- or beak-shaped, scutellar cup large, almost oval,with a pale rim, inner margins of rim with a circle of small punctures, surface of cup weakly striated basally,base of cup with a small eliptical fovea. Pronotum, viewed laterally, smooth, shining; mesopleural suturedistinct, metapleura polished. Propodeal carina almost parallel, bowed medially, lateral margins denselypubescent, weakly pubescent medially; nucha ridged, pubescent. Segment 1 of gaster partially obscured bya dense ring of hairs at base of tergite 2 , visible part of tergite 1 crenulate . Segment 2 of gaster the largest inlateral view, segments 3 and 4 partially visible in lateral view, hypopygium produced. Wing surfacespubescent, with apical hair fringe, radial cell of forewing closed on wing margin, vein M (cubitus)indicated, not pigmented. Colour: head, thorax and gaster chestnut-brown, legs yellowish orange, antennayellowish basally, apical segments darker.Cf unknown. MATERIAL EXAMINEDSouth Africa: 1 $ (BMNH). Zaire: 4 $ (MRAC). REMARKS. This species was intended as the basis for the erection of the genus Coneucoela Kieffer, 1910a,but was published subsequent to the description of Coneucoela brasiliensis Kieffer, 1909. Thus brasiliensisis the type-species of Coneucoela. Nordlander (19820) transferred gracilicornis to Trichoplasta. I have notbeen able to see the holotype of gracilicornis but have used Masner's keys (1960) to Odonteucoila, theredescription of gracilicornis, and Kieffer's original description to identify the material listed above onwhich the figures are based. .Trichoplasta longispina (Masner)(Figs 300, 335, 336) Odonteucoila longispina Masner, 1960: 358. Holotype $, SOUTH AFRICA (NCISA).Trichoplasta longispina (Masner) Nordlander, 1982a: 273. DESCRIPTION. $. Antenna 13-segmented, with distinct 5-segmented club, segment 3 shorter than 4, 4-6subequal in length, equal in width, 7-10 equal in length, 1 1 and 12 equal in length, each shorter than 6, 13 aslong as 10 (Fig. 336). Head, viewed frontally, smooth, polished, with sparse hairs, frons raised, subocularsulcus distinct; pronotum polished laterally. Pronotal plate, viewed fronto-dorsally, straight on posteriormargin, angled laterally, anterior and posterior parts of plate joined by a medial bridge, fovea on eitherside of bridge enclosed laterally (cf. Fig. 300). Mesoscutum smooth, polished, with a few scattered hairs;lateral bars of scutellum smooth, polished dorsally; scutellar foveae lenticular, scutellar disc with radiatingreticulate-rugose, apex acute, spine-shaped, scutellar cup large, almost oval, surface raised medially, witha pale outer rim, base with a small elliptical fovea (Fig. 335). Mesopleural suture distinct, metapleurapolished, ridged near juncture with hind coxa. Propodeal carinae weakly bowed, pubescent on outermargins. Segment 1 of gaster sulcate, partially obscured by a ring of hairs at base of tergite 2, complete ondorsal surface though weak, tergite 3 partially visible, hypopygium weakly protruding, gaster impunctate.Legs long, pubescent, mid and hind coxae elongate. Wing surfaces pubescent, with apical hair fringe,radial cell closed on wing margin, vein M (cubitus) not indicated. Colour: head brownish, thorax and gasterdark chestnut-red, legs yellowish orange, antenna yellowish orange basally, apical segments darker.Cf . Not seen, described by Masner (1960). AFROTROPICAL EUCOILIDAE 317 MATERIAL EXAMINEDZaire: 1 $ (MRAC). REMARKS. I have not seen determined material of this species, but the specimen from Zaire referred toabove compares well with Masner's description and figures (1960). The species is included in the key purelyon the basis of the description, but the figures are based on the specimen from Zaire. Trichoplasta medlia sp. n. (Figs 297, 300, 301, 331) DESCRIPTION. $ . Antenna 13-segmented, clavate, segment 3 longer than 4, 4 and 5 subequal, 6 longer than5, 7-13 forming a weak club (Fig. 301). Head, viewed frontally, smooth, polished, elongate, subocularsulcus distinct, antennal sockets protruding, ocelli equidistant, occipital carina distinct. Pronotal plate withanterior and posterior parts fused laterally, medial bridge narrow, fovea either side of ridge elongate,posterior part of plate, viewed frontally, angular, anterior part projected, with weak transverse striations(Fig. 300); pronotum either side of pronotal plate with dense tufts of hair. Pronotum smooth, polishedlaterally; meopleural suture distinct, metapleura with a number of ridges; mesoscutum smooth, polished,without trace of notaulices; scutellar fovea large, polished; lateral bars of scutellum polished; scutellar disccoarsely reticulate-rugose, apex beak-shaped, viewed laterally it appears elevated; scutellar cup large, oval(Fig. 297), concave in apical region. Propodeal carinae almost parallel, medial area weakly pubescent,densely pubescent on outer margins of carinae, nucha ridged, base of metapleura with tufts of pubescence.Segment 1 of gaster not visible, obscured by a dense ring of hairs at base of tergite 2, complete on dorsalsurface, tergite 2 densely punctate apically, segment 3 partially visible in lateral view, punctate, hypo-pygium not protruding. Legs short, stout, coxae swollen, elongate, with a fringe of hairs on side margins.Wing surfaces pubescent, with apical fringe of hairs, radial cell of forewing closed on margin, venationpallid, cell longer than broad (Fig. 299). Colour: head, thorax and gaster blackish to dark chestnut-red,antenna and legs reddish yellow. Cf . Antenna 15-segmented, filiform, segment 3 shorter than 4, 4 weakly swollen medially, very slightlylonger than 5 (Fig. 331). MATERIAL EXAMINED Holotype $, Zaire: Uele, Monga, 18.iv.-8.v.l935 (G. F. de Witte) (MRAC).Paratypes. Nigeria: 1 $, 1 Cf (BMNH). Zaire: 8 $, 4 cf (MRAC). REMARKS. This species is very closely related to rufus but is distinguished by the sculpture of the scutellardisc and the antennal segment ratios. Trichoplasta narrata sp. n. (Figs 304, 305, 324) DESCRIPTION. $. Antenna 13-segmented, weakly clavate, with indistinct 6-segmented club, club segmentswith rhinaria, segment 3 shorter than 4, 4 longer than 5, 5 and 6 subequal, 7 shorter than 6, 8-13 subequal inlength, each as long as 7, weakly swollen medially (Fig. 305), darker than basal segments. Head, viewedfrontally, smooth, polished, antennal sockets protruding, subocular sulcus distinct, occipital carina welldefined. Pronotal plate, viewed frontodorsally, with typically enclosed fovea either side of medial bridgebetween anterior and posterior parts of plate (cf. Fig. 324), posterior part of plate rounded dorsally,anterior part angled laterally. Pronotum either side of plate with tufts of pubescence. Mesoscutum smooth,polished; lateral bars of scutellum smooth, polished, scutellar fovea deep, clearly separated, polished;scutellar disc reticulate-rugose, apex beak-like, acutely pointed, cup large, with apical fovea. Mesopleuralsuture very fine, metapleura polished, weakly ridged, with a tuft of pubescence on anterior margin, nucharidged, with long scattered pubescence. Segment 1 of gaster crenulate, partially obscured by a ring ofpubescence at base of tergite 2 which is impunctate, remaining tergites not visible in lateral view,hypopygium pronounced. Legs slender, coxae with a hair fringe on outer margins. Wing surfacespubescent, with long apical hair fringe, radial cell of forewing closed on wing margin, vein M (cubitus)indicated, not pigmented. Colour: head brownish, thorax and gaster orange-brown, antenna yellowbasally, apical segments brownish. Cf . Antenna 15-segmented, segment 3 twisted medially, not swollen (Fig. 304). MATERIAL EXAMINED Holotype $, Uganda: Kawanda, x.1942 (H. C. Taylor} (BMNH).Paratypes. Uganda: 1 $ , 1 cf , same data as holotype (BMNH). Zaire: 1 cf (MRAC). 318 J. QUINLAN REMARKS. This species is very closely related to unicolora but can be separated by the shape and form of thepronotal plate. Trichoplasta novema sp. n. (Figs 324, 328, 332, 333) DESCRIPTION. $ . Antenna 13-segmented, clavate, segment 3 longer than 4, 4 shorter than 5, 5-13 forming avery distinct club (Fig. 328). Head, viewed frontally, smooth, polished, frons raised, with distinct lateralmargins extending from raised antennal sockets, subocular sulcus (malar suture) distinct, with weak ridgeson either side, occipital carina pronounced dorsally. Pronotal plate, viewed frontodorsally, weaklyrounded, lateral margins more angular, anterior and posterior parts fused medially and laterally to enclosea fovea on either side of medial bridge, medial bridge narrow, fovea elongate, distinct, anterior part ofplate weakly sculptured (Fig. 324). Pronotum pubescent either side of pronotal plate with a fewcanaliculations on lower lateral margins. Mesopleural suture distinct, metapleura ridged on lower margins(Fig. 334). Mesoscutum smooth, polished, notaulices absent; scutellar foveae kidney-shaped, lateral barsof scutellum with weak aberrant sculpture apically, scutellar disc with punctate-reticulate-rugose sculpture(Fig. 332), apex conical, scutellar cup large, almost circular, with a round fovea apically (Fig. 332).Propodeal carinae horseshoe-shaped, weakly pubescent medially, densely pubescent on lateral margins,nucha pubescent. Segment 1 of gaster obscured by a ring of dense pubescence at base of tergite 2 which isvery weakly punctate apically, segment 3 partially visible in lateral view, hypopygium weakly produced.Legs slender, coxae narrow, sparsely pubescent. Wing surfaces pubescent, with apical hair fringe, radialcell of fore wing elongate, open on wing margin, venation yellowish, vein M (cubitus) indicated, weaklypigmented basally (Fig. 333). Colour: head and thorax blackish, gaster chestnut-brown, antenna blackishbrown, except basal segments lighter.C? unknown. MATERIAL EXAMINED Holotype $, Uganda: Butandiga, xii.1938 (H. C. Taylor) (BMNH).Paratypes. Uganda: 1 $, same data as holotype (BMNH). Zaire: 2 $ (MR AC). REMARKS. This species is very similar to octonius but has a 9-segmented antennal club, a weakly sculpturedpronotum and the pronotal plate is angular (Fig. 324). Trichoplasta octonarius sp. n. (Figs 354, 355, 356) DESCRIPTION. $ . Antenna 13-segmented, clavate, segment 3 longer than 4, 4 and 5 subequal, 6-13 forminga distinct club (Fig. 354). Head, viewed frontally, smooth, polished, frontal ridges prominent in proximityto antennal sockets which protrude, occipital carina pronounced, subocular suture distinct. Pronotal platewith anterior and posterior parts fused medially and laterally to enclose a fovea either side of medialbridge, medial bridge narrow, posterior part of plate rounded, anterior part angular on lower margins,anterior part smooth, polished. Pronotum viewed laterally pubescent either side of pronotal plate, withstrong transverse striations extending to mesopleura. Mesopleural suture distinct, metapleura with fiveridges. Mesoscutum smooth, polished, without trace of notaulices; scutellar foveae large, polished, with ahole in basal corner ending under lateral bars of scutellum which are polished; scutellar disc reticulate-rugose, apex conical, scutellar cup large, almost round, basal half with weak striations, apical half with alarge fovea (Fig. 356). Propodeal carina broad, parallel, densely pubescent; nucha ridged; margin ofmetapleura pubescent. Segment 1 of gaster not visible, obscured by a ring of dense pubescence at base oftergite 2, apical two-thirds of tergite 2 punctate, tergites 3 and 4 visible laterally, hypopygium stronglyproduced. Legs short, coxae swollen, elongate, sparsely pubescent. Wing surfaces pubescent, with apicalhair fringe, radial cell of forewing open on wing margin, longer than broad, venation pale, vein M (cubitus)visible, not pigmented (Fig. 355). Colour: head, thorax and gaster blackish, antenna orange-yellowbasally, darker apically, legs orange-yellow.C? unknown. MATERIAL EXAMINEDHolotype $, Zaire: Congo de Lemba, i.1913 (R. Mayne) (MRAC). REMARKS. Closely related to novema but distinguished by the 8-segmented club and the laterally striatepronotum. AFROTROPICAL EUCOILIDAE 319 Trichoplasta quindava sp. n. (Figs 314, 342, 343, 344) DESCRIPTION. $ . Antenna 13-segmented, filiform, with weak 5-segmented club, segment 3 shorter than 4,4-7 subequal in length, 8 and 9 subequal, each shorter than 7, 10-13 each slightly shorter than 9 (Fig. 343).Head, viewed frontally, smooth, polished, with long scattered hairs, cheeks converging, antennal socketsprotruding, frons raised, subocular sulcus distinct, occipital carina distinct on dorsal margin, obscured bytufts of pubescence either side of pronotal plate which, viewed frontodorsally, is rounded posteriorly.Pronotal plate with anterior and posterior parts fused medially and laterally to enclose a fovea either side ofmedial bridge (Fig. 344). Pronotum smooth, polished laterally. Mesopleural suture distinct, metapleurastrongly ridged basally, with tufts of hair either side of propodeum. Mesoscutum smooth, polished,notaulices absent, a few long hairs in their place; lateral bars of scutellum smooth, polished; scutellarfoveae polished, kidney-shaped; scutellar disc reticulate-rugose, apex narrowing sharply to form abeak-shaped apex, scutellar cup elliptical (cf. Fig. 314). Propodeal carinae bowed medially, denselypubescent on outer margins; nucha ridged. Segment 1 of gaster crenulate, tergite 2 with a ring of hairs at itsbase, incomplete on dorsal surface, tergite 3 weakly protruding, hypopygium obscure, gaster impunctate.Legs short, stout, coxae elongate, with sparse hairs apically. Wings pubescent, with apical hair fringe,radial cell of forewing closed on wing margin, vein M (cubitus) indicated basally, not pigmented (Fig. 342).Colour: head and thorax blackish brown, legs orange-yellow, antenna yellow.CT unknown. MATERIAL EXAMINED Holotype $, Zaire: N. Lac Kivu, Rwanki, 15.ii.1952 (/. V. Leroy) (MRAC).Paratypes. Zaire: 3 $ (MRAC). REMARKS. This species is separated from controsta by the antennal coloration and the shape of the pronotalplate. The antennal club is not sharply contrasted to the basal flagellar segments but all club segments haverhinaria. Trichoplasta rufus sp. n. (Figs 298, 302, 303, 315, 317) DESCRIPTION. $ . Antenna 13-segmented, clavate, segment 3 longer than 4, 4 longer than 5, 6 longer than 5,7-13 forming a club (Fig. 303). Head, viewed frontally, elongate, smooth, polished, subocular sulcusdistinct, occipital carina distinct, ocelli weakly elevated, equidistant, antennal sockets protruding,mandibles tridentate, sparsely pubescent. Pronotal plate with anterior and posterior parts fused mediallyand laterally to enclose a fovea either side of medial bridge (Fig. 302). Pronotum either side of pronotalplate smooth, polished, with scattered hairs. Mesoscutum polished, without trace of notaulices; scutellardisc punctate-reticulate, apex beak-shaped (Fig. 298) ; scutellar cup almost oval, with a basal fovea, viewedlaterally, elevated; scutellar foveae shallow, polished, separated by a weak septum; lateral bars ofscutellum polished; propodeal carinae almost parallel, densely pubescent medially and laterally. Meso-pleural carinae distinct, metapleura polished, with a few crenulations at juncture with coxa; nucha ridged,pubescent. Segment 1 of gaster not visible, obscured by a ring of dense pubescence at base of tergite 2,complete on dorsal surface, apex of tergite 2 with sparse scattered punctures, hypopygium weaklyprotruding. Legs stout, short, coxae elongate, pubescent, with a fringe of hairs. Wing surfaces denselypubescent, with apical hair fringe, radial cell of forewing open on wing margin, longer than broad (Fig.315). Colour: head, thorax and gaster reddish brown, antenna and legs reddish yellow. d" . Antenna 15-segmented, filiform, segment 3 shorter than 4, 4 flattened on outer margins (Fig. 317). MATERIAL EXAMINED Holotype 9, Madagascar: Mandraka, ii.1944 (A. Seyrig) (MRAC).Paratype. Madagascar: 3 cf , same data as holotype (MRAC). Zaire: 1 $ (MRAC). Trichoplasta tanganyikensis (Weld)(Figs 292, 293, 324, 329) Coneucoela tanganyikensis Weld, 1944: 63. Holotype $, TANZANIA (USNM) [examined].Trichoplasta basilewskyiBenoit, 1956: 538. Holotype $, RWANDA (MRAC) [examined]. [Synonymy by Nordlander, 1982a: 272.]Trichoplasta tanganyikensis (Weld) Nordlander, 1982a: 272. 320 J. QUINLAN DESCRIPTION. 9 . Antenna 13-segmented, segment 3 subequal to 4+5, 6 longer than 4 or 5, 7-13 subequal inlength, all distinctly wider than preceding flagellar segments and forming a sharply defined club (Fig. 329).Head, viewed frontally, smooth, shining, rounded, frons raised, eyes same distance apart as height of aneye, malar space with a carina and weak striations on lower side, vertex smooth, shining. Pronotum eitherside of pronotal plate with tufts of pubescence. Pronotal plate produced, anterior and posterior parts fusedlaterally, medial bridge between both parts narrow (cf. Fig. 324), fovea either side of bridge wider thanlong. Mesopleural suture distinct, metapleura ridged. Mesoscutum smooth, polished, notaulices repre-sented by a row of hairs; scutellar foveae large, smooth, shining; scutellar disc reticulate-rugose, apexconical, extending out over propodeum, scutellar cup large, a little longer than wide, apical quarter with alarge fovea, lateral margins of cup with a line of small foveae or pits (Fig. 292); lateral bars of scutellumsmooth and polished on dorsal surface; propodeal carinae bowed medially, pubescent on outer margins ofcarinae, less so medially. Segment 1 of gaster rugulose though obscured by a ring of dense pubescence atbase of tergite 2, complete on dorsal surface, apical half of tergite 2 and visible parts of tergites 3 and 4strongly punctate (Fig. 293), hypopygium strongly protruding. Wing surfaces densely pubescent, withapical fringe of hairs, radial cell of forewing open on wing margin, vein M (cubitus) weakly indicated inpart, apex of wing rounded. Legs short, stout, coxae swollen medially, tibiae and tarsi pubescent. Colour:antenna yellowish black, head and thorax black, gaster chestnut reddish brown, legs reddish brown. Cf . Antenna 15-segmented, segment 3 shorter than 4, outer margin flattened, segment 4 longer thanfollowing segments, swollen distally, flattened medially. MATERIAL EXAMINED Holotype $ (tanganyikensis), Tanganyika: Mutresa, 21.ix.1935 (F. Bianchi) (USNM, cat.No. 56814).Holotype $ (basilewskyi), Rwanda: Contrefort est du Muhavara, 2100 m, 28. i. 1953 (P. Basilewsky)(MRAC). Zaire: 1 $, Mt Sesero, near Bitashrmva (Bombous), 2000 m, l-2.viii.1934 (G. F. de Witte) (MRAC).Cameroun: 1 $ (BMNH). REMARKS. Trichoplasta tanganyikensis (Weld) was originally assigned to Coneucoela. Benoit (1956)established the genus Trichoplasta and designated basilewskyi as type-species. Nordlander (1982a)synonymised both species. Masner (1960) and Nordlander (1978a, 1980) had previously regardedOdonteucoila a genus in which Masner had described a number of species now recognised as belonging inTrichoplasta as being closely related to Rhoptromeris . The type-species of Odonteucoila, however, belongsto those groups of genera with the lateral cavities of the pronotal plate open. Trichoplasta testacea sp. n. (Figs 347, 349, 350, 351) DESCRIPTION. 9- Antenna 13-segmented, clavate, with a 6-segmented club, segment 3 shorter than 4, 4longer than 5, 5 shorter than 6, first club segment slightly less broad than others (Fig. 349), all with rhinaria.Head, viewed frontally, smooth, polished, with a line of hairs extending from base of clypeus to antennalsockets, subocular sulcus present, occipital carina distinct viewed dorsally. Pronotal plate, viewedfrontodorsally, with two foveae, one each side of medial bridge between anterior and posterior parts ofplate, fovea closed laterally, sides of plate angular (Fig. 351), pronotum either side of plate with dense tuftsof pubescence. Mesoscutum smooth, polished, notaulices very weakly indicated anteriorly by a fewscattered hairs in their place; lateral bars of scutellum very weakly indicated; scutellar foveae kidney-shaped, smooth, polished; scutellar disc shining, weakly areolate, apex sharply tapered, scutellar cup long,narrow, apex with a small declined fovea (Fig. 350). Pronotum smooth on lateral margins; mesopleuralsuture distinct, metapleura smooth, pubescent on anterior margins; nucha ridged. Segment 1 of gasterobscured by a ring of hairs at base of tergite 2, incomplete on dorsal surface, apex of tergite 2 impunctate,tergite 3 partially visible, hypopygium pronounced. Legs long, slender, coxae and tibiae weakly pubescent.Wing surfaces pubescent, apical margins with a hair fringe, radial cell of forewing closed on wing margin(Fig. 347), vein M (cubitus) not indicated. Colour: head, thorax, gaster and legs orange-yellow.Cf unknown. MATERIAL EXAMINEDHolotype $>, Nigeria: Ibadan (B. R. Critchley}. REMARKS. This species is completely orange-yellow and differs from others by the scutellar cup which has adeclined fovea apically (Fig. 350). AFROTROPICAL EUCOILIDAE 321 Trichoplasta unicolora sp. n. (Figs 319, 320, 351, 352) DESCRIPTION. $ . Antenna 13-segmented, weakly clavate, with a 6-segmented club, segment 3 shorter than4, 5-10 subequal in length, segments 8-13 broader medially than segments 3-7 (Fig. 319). Head, viewedfrontally, smooth, polished, with long scattered hairs on frons and close to clypeal area, sutures weaklypresent below antenna, antennal sockets not prominent, anterior tentorial pits distinct; occipital carinadistinct dorsally, obscured laterally by tufts of pubescence on either side of pronotal plate. Pronotal plateappearing rectangular viewed frontodorsally, anterior and posterior parts bridged medially and laterally toenclose two pits, one on either side of medial bridge, medial bridge broad (cf. Fig. 351). Mesoscutumsmooth, polished, with a few long hairs in place of notaulices; scutellar foveae shallow, weakly separated,polished; scutellar disc reticulate-rugose, tapered to form a blunt point apically; lateral bars of scutellumpolished dorsally; scutellar cup oval, with a pale rim, centre raised, polished, with small pits or foveaearound rim (cf. Fig. 320). Lateral margins of pronotum, mesopleura and metapleura polished. Mesopleu-ral suture distinct, metapleura with weak ridges, lower edges with tufts of hairs. Propodeal carinae parallel,weakly pubescent; nucha ridged, pubescent. Segment 1 of gaster crenulate, partially obscured by a ring ofpubescence at base of tergite 2, incomplete on dorsal surface, gaster impunctate, hypopygium notpronounced. Legs long, slender, pubescent, coxae elongate. Wings narrow, surfaces pubescent, withapical hair fringe, radial cell of forewing closed on wing margin, vein M (cubitus) not indicated, venationweakly pigmented. Colour: head light brown, thorax and gaster orange-brown, legs yellow, antenna paleyellow. Cf antenna 15-segmented, segment 4 curved medially, swollen basally (Fig. 352). MATERIAL EXAMINED Holotype $, Zaire: Rutshuru, xii.1937 (/. Ghesquiere) (MRAC).Paratypes. Zaire: 9 $, 6 cf (MRAC). REMARKS. Closely related to narrata but separated by the form of the pronotal plate and the completelypale antenna. Trichoplasta zeus sp. n. (Figs 344, 356, 358) DESCRIPTION. $ . Antenna 13-segmented, very weakly clavate, segment 3 shorter than 4, 5-13 each shorterthan 4, 4-6 subequal in length and width, 7-13 with rhinaria, weakly swollen medially, subequal in length,forming a 6-segmented club (Fig. 358). Head, viewed frontally, smooth, polished, with scattered hairsextending from antennal region to clypeus, anterior tentorial pits distinct, subocular sulcus fine butdistinct, antennal sockets weakly protruding, occipital carinae, viewed dorsally, not conspicuous, head,viewed dorsally, with hind margin almost straight, ocelli equidistant, viewed frontally, eyes as far apartmeasured medially as height of an eye. Pronotal plate projected forward, anterior and posterior parts fusedlaterally and medially very weakly, to enclose a fovea on either side of medial bridge (cf. Fig. 344); lateralmargins of pronotum with tufts of pubescence on either side of lower margins of pronotal plate, surfacepolished. Mesoscutum smooth, polished, notaulices absent; lateral bars of scutellum smooth, polished;scutellar disc reticulate-rugose, apex conical, beak-shaped in lateral view; scutellar cup large, with apicalfovea (cf. Fig. 356). Mesopleura smooth, polished, suture distinct, metapleura with long hairs on anteriormargin, ante ro ventral cavity open basally, a few hairs present. Propodeal carinae parallel, converging atjuncture with nucha, densely pubescent on lateral margins, with sparse hairs medially. Segment 1 of gasterridged , obscured by a ring of dense pubescence at base of tergite 2 , tergites 3 and 4 partially visible in lateralview, gaster impunctate, hypopygium weakly protruding. Legs long, slender, pubescent. Wings narrow,surface pubescent, apical margins with a hair fringe, radial cell of forewing closed on wing margin, venationpale yellow, vein M (cubitus) not indicated. Colour: head and thorax brownish black, gaster brownishyellow, legs orange-yellow, antenna yellow basally, apical segments darkish.Cf unknown. MATERIAL EXAMINED Holotype $, Cameroun: Mt Cameroon, Mann's Quelle, 7400 ft (2250 m), 30.U932 (M. Steele)(BMNH). Paratypes. Cameroun: 3 $ (BMNH), same data as holotype (one damaged). REMARKS. Distinguished from the closely related species gracilicornis by the antennal segment ratios. 322 J. QUINLAN Acknowledgements I wish to thank my colleagues in the Hymenoptera Section for their advice, particularly Mr T. Huddlestonwho critically examined the introduction. My thanks are extended to my sister-in-law Marina Ellwood fortyping the manuscript. References Ashmead, W. H. 1887. On the cynipidous galls of Florida with descriptions of new species and synopsis of the described species of North America. Transactions of the American Entomological Society 14: 150-154.1903. Classification of the gall-wasps and parasitic Cynipoids, or the super family Cynipoidea. II. Psyche 10: 59-73. Askew, R. R. 1971. Parasitic insects xvii + 316 pp.Barbotin, F., Carton, Y. & Kelner Pillault, S. 1979. Morphologic et biologic de Cothonaspis (Cothonaspis) boulardi n. sp. , parasite de drosophiles. Bulletin de la Societe Entomologique de France 84: 20-26.Benoit, P. L. G. 1956. Contributions a 1'etude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky 1953) GIX. Hymenoptera Cynipidae. Annales du Musee Royal du Congo Beige (Serie 8:zool.) 51: 532-550.Bridwell, J. C. 1919. Descriptions of new species of hymenopterous parasites of muscoid Diptera with notes on their habits. Proceedings of the Hawaiian Entomological Society 4: 166-179.Cameron, P. 1890. Monograph of British phytophagous Hymenoptera 3: 274 pp. London.1904. On the Hymenoptera of the Albany Museum, Grahamstown, South Africa. Record of the Albany Museum 1: 161-175.Carton, Y. 1977. Attraction de Cothonaspis sp. (Hymenoptere Cynipidae) par le milieu trophique de son hole: Drosophila melanogaster. Colloques Internationaux du Centre National de la Recherche Scienti- fique no. 268:285-303. Carton, Y., Rouault, J. & Kitano, H. 1977. Susceptibility of seven sibling species of sub-group melano-gaster infected with a cynipid parasite. Drosophila Information Service 53: 183.Chrystal, R. N. 1930. Studies of the Sirex parasites. The biology and post-embryonic development oflbalia leucospoides Hochenw. Oxford Forestry Memoirs 11: 1-63.Crosskey, R. W. & White, G. B. 1977. The Afrotropical Region, a recommended term in zoogeography. Journal of Natural History 11: 541-544. Dalla Torre, K. W. & Kieffer, J. J. 1910. Cynipidae. Tierreich 24: 1-891.Day, M. C. 1984. The enigmatic genus Heterogyna Nagy (Hymenoptera Sphecidae: Heterogyninae). Systematic Entomology 9: 293-307. Diaz, N. B. 1973. Una familia de Cynpoidea nueva para lar Argentina. Neotropica 19: 141-144.Eady, R. D. E. & Quinlan, J. 1963. Hymenoptera Cynipoidea. Key to families and subfamilies and Cynipinae (including galls). Handbooks for the Identification of British Insects 8 (la): 1-81.Foerster, A. 1869. Ueber die Gallenwespen. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 19: 327-370.Giraud, J. 1860. Enumeration des Figitides de 1'Autriche. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 10: 123-176. Hartig, T. 1840. Uber die Familie der Gallwespen. Zeitschrift fur Entomologie (Germar) 2: 176-209.1841. Erster Nachtrag zur Naturgeshichte der Gallwespen. Zeitschrift fur Entomologie (Germar) 3: 358.Hedicke, H. 1928a. Beitrage zur Kenntnis der Cynipiden (Hym). Verhandlungen des Vereins fur Natur- wissenschaftliche Unterhaltung zu Hamburg 19 (1926-27): 72-96.l92Sb. Beitrage zur Kenntnis der Cynipiden (Hym.) Deutsche Entomologische Zeitschrift, Iris 1: 81-85. Hellen, W. 1960. Die Eucoilen Finnlands. Fauna Fennica 9: 1-31.Imms, A. D. 1930. Observations on some parasites of Oscinella frit L. Part I. Parasitology, Cambridge 22: 11-36.1932. Observations on some parasites of Oscinella frit L. Part II. Parasitology, Cambridge 24: 440-447.James, H. C. 1928. On the life histories and economic status of certain cynipid parasites of dipterous larvae, with descriptions of some new larval forms. Annals of Applied Biology 15: 287-316. AFROTROPICAL EUCOILIDAE 323 Jenni, W. 1951. Beitrag zur Morphologic und Biologic der Cynipide Pseudeucoila bochei Weld, eines Larvenparasiten von Drosophila melanogasterMeig. Acta Zoologica fennica 32: 177-254.Kerrich, G. J. & Quinlan, J. 1960. Studies on eucoiline Cynipoidea (Hym.). Opuscula Entomologica 25: 179-196. Kieffer, J. J. 1901. Revision des Eucoilines. Feuille desJeunes Naturalistes 31: 158-176.1902. Les Cynipides 4 e Tribu-Eucoelines (Eucoelinae Dalla Torre). In Andre, E. Species des Hymenopteres d' Europe & d'Algerie 7 (2): 78-240.1909. Description de nouveaux Cynipides zoophages. Bulletin de la Societe d'Historie Naturelle (du Department) de la Moselle 26: 57-96.1910a. Serphiden und Cynipiden von Madagaskar. Wissenschaftliche Ergebnisse. Reise in Ostafrika in den Jahren 1903-1905 von A. Voeltzhkow 2: 529-534. 19106. Nouveaux Cynipides exotiques. Bollettino del Laboratorio Zoologica Portici4: 329-342. 1910c. Serphidae, Cynipidae, Chalcidae, Evanidae und Stephanidae aus Aequatorial- Africa. Wissen- schaftliche Ergebnisse des Deutschen Zentral-Afrika Expedition 3 (2): 1-6, 97-112, 23-29. 1913. Proctotrupidae, Cynipidae et Evaniidae. In: Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientate (1911-1912). Resultats scientifiques. Hymenoptera 1: 1-198.Kierych, E. 1979. Notes on the genera Dilyta Forster, 1869 and Glyptoxysta Thomson, 1877 Hymenoptera, Cynipoidea, Alloxystidae, part 1. Annales Zoologica fennila 34: 453-460.Lipkow, E. 1969. Cynipoidea und Ichneumonidae (Hym.) als Parasiten von Boriomyia subnebulosa (Neur. Hemerobiidae). Entomophaga 14: 229-241.Masner, P. 1958. Contribution to the knowledge of the genus Ganaspis Foerster, 1869 (Hym. Cynipoidea). Casopis Ceskoslovenske Spolecnosti Entomologicke 55: 264-279.1960. Revision of the genus Odonteucoila Ashmead (Hym., Cynipoidea). Casopis Ceskoslovenske Spolecnosti Entomologicke 57: 348-364.Nordlander, G. 1976. Studies on Eucoilidae (Hym., Cynipoidea) I. A revision of the north-western European species of Cothonaspis Htg. with description of a new species and notes on some other genera. Entomologisk Tidskrift97: 65-77.1978a. Revision of the genus Rhoptromeris Forster, 1869 with reference to north-western European species. Studies on Eucoilidae (Hym., Cynipoidea) II. Entomologica Scandinavica 9: 47-62.19786. Parasitoids of the frit fly, Oscinella frit (L.), on oats, Norwegian Journal of Entomology 25: 89-90.1980. Revision of the genus Leptopilina Forster, 1869, with notes on the status of some other genera (Hymenoptera, Cynipoidea: Eucoilidae). Entomologica Scandinavica 1: 428-453.1981. A review of the genus Trybliographa Forster, 1869 (Hymenoptera, Cynipoidea: Eucoilidae). Entomologica Scandinavica 12: 381-402.1982a. Identities and relationships of the previously confused genera Odonteucoila, Coneucoela and Trichoplasta (Hymenoptera, Cynipoidea: Eucoilidae). Entomologica Scandinavica 13: 269-292. 19826. Systematics and phytogeny of an interelated group of genera within the family Eucoilidae (Insecta: Hymenoptera, Cynipoidea). 32pp. Stockholm.Nostvik, E. 1954. A study of Pseudeucoila bochei Weld (Hymenoptera: Eucoilidae) and its relationship to Drosophila melanogaster Meig. (Diptera: Drosophilidae). Symposia Genetica, Pavia 2 (Genetica ed Entomologia): 139-160.Quinlan, J. 1963. Hymenoptera, Cynipoidea, key to families and subfamilies, and Cynipinae (including galls). Handbooks for the Identification of British Insects 8, 1, (b): 1-81.1967. The brachypterous genera and species of Eucoilidae (Hymenoptera), with descriptions and figures of some type-species. Proceedings of the Royal Entomological Society London (B)36: 1-10.1978. Hymenoptera, Cynipoidea, Eucoilidae. Handbooks for the Identification of British Insects 8, 7 (b): 1-58.1979. A revisionary classification of the Cynipoidea (Hymenoptera) of the Ethiopian Zoogeographi- cal Region. Aspicerinae (Figitidae) and Oberthuerellinae (Liopteridae). Bulletin of the British Museum (Natural History) (Entomology) 39: 85-133. 1984. Stentorceps, a remarkable new genus of eucoilid (Hymenoptera) from Africa. Systematic Entomology 9: 479-485.Quinlan, J. & Evenhuis, H. H. 1980. Status of the subfamily names Charipinae and Alloxystinae (Hymenoptera: Cynipidae). Systematic Entomology 5: 427-430.Rohwer, S. A. & Fagan, M. M. 1917. The type-species of the genera of the Cynipoidea, or the gall wasps and parasitic cynipoids. Proceedings of the United States National Museum 53: 357-380.Richards, O. W. 1977. Hymenoptera. Introduction and keys to families, 2nd edn. Handbooks for the identification of British Insects 6(1): 100 pp. 324 J. QUINLAN Riek, E. F. 1971. A new family of cynipoid wasps (Hymenoptera: Cynipoidea) from Australia. In Syoziro Asuhira (Ed.) Entomological Essays to Commemorate the Retirement of Professor K. Yasumatsu pp. 107-112.Risbec, J. 1950. 1 Lafaune entomologique des Cultres au Senegal etau Soudan Frangais: II. Proctotrupidae. 668 pp. Paris.1956. Hymenopteres parasites du Cameroun (2 et 3 contributions). Bulletin de I'lnstitut Francois d'Afrique Noire 18: 97-164.Selhime, A. G. & Kanavel, R. F. 1968. Life cycle and parasitism of Micromus posticus and M. subanticus in Florida. Annals Entomological Society of America 61: 1212-1215.Spradbury, J. P. 1970. The biology oflbalia drewseni Borries, a parasite of siricid woodwasps. Proceedings of the Royal Entomological Society London (A) 45: 104-113.Thomson, C. G. 1862. Forsok till uppstallning och beskrifning av Sverges Figiter. Ofversigt af Kongl Vetenskaps-Akademiensforhandlingar 18: 395420.Thompson, W. R. 1955. A catalogue of the parasites and predators of insect pests. Section 2, part 3. Hosts of the Hymenoptera (Calliceratid to Evaniid). pp. 191-332. Ottawa.Tsacas, L. 1979. Contribution des donnees africaines a la comprehension de la biogeographie de 1'evolution du sous-genre Drosophila (Sophophord) Sturtevant (Diptera, Drosophilidae). Compte Rendu des Seances de la Societe de Biologic no. 480: 29-51.Watrous, L. E. & Wheeler, Q. D. 1981. The out group comparison method of character analysis. Systematic Zoology 30: 1-11.Weld, L. H. 1931. Additional notes on types with description of a new genus (Hymenoptera: Cynipidae). Proceedings of the Entomological Society of Washington 33(a): 220-227.1944. Descriptions of new Cynipidae including two new genera (Hymenoptera). Proceedings of the Entomological Society of Washington 46: 55-66. 1952. Cynipoidea (Hymenoptera) 1905-1950. 357 pp. Ann Arbor, Michigan (privately printed). 1961. New synonymy in Cynipoidea (Hymenoptera). Proceedings of the Entomological Society of Washington 63: 279-280.Westwood, J. O. 1833. Notice on the habits of a cynipideous insect parasitic upon the rose louse (Aphis rosae); with descriptions of several other parasitic Hymenoptera. Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology and Meteorology 6: 491-495.Wishart, G. & Monteith, E. 1954. Trybliographa rapae (Westw.), a parasite of Hylemya spp. (Diptera: Anthomyiidae). Canadian Entomologist 86: 145-154. AFROTROPICAL EUCOILIDAE sec 325 Figs 1-6 1, thorax and gaster, lateral view of Eucoilidae. 2, thorax, dorsal view, of Eucoilidae. 3,forewing of Eucoilidae. 4, forewing of Anacharitinae. 5, lateral view of gaster of Anacharitinae. 6,pronotal plate of Rhoptromeris sp. 326 J. QUINLAN 13 Figs 7-13 7, $ antenna of Rhoptromeris heptoma (Hartig). 8, forewing of Kleidotoma psiloidesWestwood. 9, 10, Afrodontaspis sp.; (9) forewing; (10) mesonotum. 11, Nordlanderia sp., pronotalplate. 12, Trybliographa sp., pronotal plate. 13, Nordlanderia sp., gaster. AFROTROPICAL EUCOILIDAE 327 22 Figs 14-23 14, Eucoilidea sp., gaster. 15, Nordlanderia sp., scutellar disc. 16, Nordlanderia sp., lateralview of head. 17, Disorygma sp., mesonotum. 18, Diglyphosema sp., pronotal plate. 19, Ealata sp.,radial cell. 20, Ealata clava sp. n. , pronotal plate. 21 , Diglyphosema conjungens Kieffer , mesonotum. 22,Diglyphosema sp., 1st tergite of gaster. 23, Eucoilidea sp., scutellum. 328 J. QUINLAN 29 Figs 24-32 24-26, pronotal plate of (24) Afrodontaspis striatissima Benoit; (25), Trichoplasta medlia sp.n. ; (26) Odonteucoila sp. 27, Eucoila sp., forewing. 28, Eucoila sp. , dorsal view of pronotal plate. 29, 30,Bothrochacis sp.; (29) scutellum; (30) pronotal plate. 31, Stentorceps tubicen Quinlan; (31) head,dorsal view; (32) head, lateral view. AFROTROPICAL EUCOILIDAE 329 Figs 33-42 33, Cothonaspis sp. , 1st tergite of gaster. 34, Cothonaspis sp. , pronotal plate. 35, Kleidotomabifurcata sp. n. , forewing. 36, Kleidotoma sp. , dorsal view of 1st tergite of gaster. 37, Glauraspidia sp. ;mesopleura. 38, Hexacola sp.; radial cell. 39, Hexacola sp.; cf antenna, basal segments. 40, Hexacolasp.; pronotal plate. 41, Leptopilina davipes (Hartig) $, gaster. 42, Trybliographa sp.; pronotal plate. 330 J. QUINLAN 53 Figs 43-53 43, Trybliographa rapae (Westwood), mesonotum. 44, 45, Ganaspis seticornis (Hellen); (44)mesonotum; (45) pronotal plate. 46, Trybliographa rapae (Westwood), C? basal segments of antenna.47, Afrodontaspis sp., pronotal plate. 48, A. striatissima Benoit, gaster. 49, A. lanatus sp. n., pronotalplate. 50, A. striatissima Benoit, mesoscutum. 51 , A. lanatus sp. n. , mesoscutum. 52, A. lanatus sp. n. $ ,antenna. 53, A. lanatus sp. n., frontal view of head. AFROTROPICAL EUCOILIDAE 331 Figs 54-63 54, Afrodontaspis lanatus sp. n. , forewing. 55, 56, A. striatissima Benoit; (55) 9 antenna; (56)forewing. 57, 58, Cothonaspis pentatoma Hartig; (57) $ antenna; (58) pronotal plate. 59, 60, C. dulcissp. n.; (59) $ antenna; (60) pronotal plate. C. eala sp. n.; (61) $ antenna; (62) pronotal plate. 63, C.pentatoma Hartig, lateral view of basal segments of gaster. 332 J. QUINLAN Figs 64-71 64, Ea/ata marica sp. n. , gaster. 65, 66, $ antenna of (65) E. subba sp. n. ; (66) E. clava sp. n.67-69, E. clava sp. n.; (67) gaster; (68) propodeum; (69) mesonotum. 70, 71, mesonotum of (70)Cothonaspis pentatoma Hartig; (71) C. dulcis sp. n., 72, Ealata clava sp. n., forewing. AFROTROPICAL EUCOILIDAE 333 Figs 73-81 73, Ealata marica sp. n., $ antenna. 74, 75, Eucoilidea canadensis Ashmead; (74) mesono-tum; (75) radial cell. 76, E. extraria sp. n., 9 antenna. 77, E. trulla sp. n., gaster. 78, E. marcellus sp. n.,radial cell. 79, E. trulla sp. n., $ antenna. 80, E. tyrus sp. n. radial cell. 81, E. lacerta sp. n., $ 4thantennal segment. 334 J. QUINLAN Figs 82-92 82, radial cell of Eucoilidea lacerta sp. n. 83, 84, E. lana sp. n. ; (83) lateral view of pronotum;(84) radial cell. 85, 86, mesonotum of (85) E. nitida (Benoit); (86) E. leptis sp. n. 87-89, $ antenna of(87) E. conversa sp. n. ; (88) E. parma sp. n. ; (89) E. fetura sp. n. 90, 91 , cf antenna, basal segments, of(90) E. dubia sp. n.; (91) E. compressa sp. n. 92, mesonotum of E. fetura sp. n. AFROTROPICAL EUCOILIDAE 335 Figs 93-101 93, mesonotum of Eucoilidea pallida sp. n. 94, gaster of E. compressa sp. n. 95, radial cell ofE. perangusta sp. n. 96, $ antenna of E. perangusta sp. n. 97, mesonotum of E. bucca sp. n. 98, $antenna of E. urundiensis Benoit. 99, O" basal antennal segments of E. urundiensis Benoit. 100, 101, E.urundiensis Benoit; (100) gaster; (101) pronotal plate. 336 J. QUINLAN 1 1 1 Figs 102-112 102, mesonotum of Eucoilidea urundiensis Benoit. 103, 9 antenna of E. bucca sp. n. 104,mesonotum of E. advena sp. n. 105, basal segments, $ antenna of E. dubia sp. n. 106, O" gaster of E.trulla sp. n. 107-110, d" basal antennal segments of (107) E. furcula sp. n. ; (108) E. conversa sp. n. ; (109)E. lana sp. n. ; (110) E. marcellus sp. n. Ill, 112, fore wing of (111) E. furcula sp. n. ; (112) E.fetura sp. n. AFROTROPICAL EUCOILIDAE 337 1 13 Figs 113-119 113, $ antenna of Eucoilidea advena sp. n. 114, mesonotum of E. lana sp. n. 115, basalsegments, cf antenna of E. nitida(Benoii). 116, $ gasterof E. dubiasp. n. 117, $ face of E. lacertasp. n.118, 119, gaster of (118) E. fetura sp. n.; (119) E. furcula sp. n. 338 J. QUINLAN 120 126 Figs 120-126 120, $ antenna ofEucoilideafurcula sp. n. 121-123, radial cell of (121) E. compressa sp. n. ;(122) E. lacerta sp. n.; (123) E. extraria sp. n. 124, 9 antenna of E. lana sp. n. 125, lateral view ofpronotum of E. lana sp. n. 126, cf basal antennal segments of E. lana sp. n. AFROTROPICAL EUCOILIDAE 339 131 1 27 Figs 127-134 127, 128, Eucoilidea leptis sp. n.; (127) $ antenna; (128) pronotal plate. 129, 130, E.marcellus sp. n.; (129) $ antenna; (130) radial cell, 131, 132, E. mauri sp. n.; (131) $ antenna; (132)mesonotum. 133, 134, E. nitida Benoit; (133), 9 antenna; (134) pronotal plate. 340 J. QUINLAN 142 144 140 Figs 135-144 135-137, Eucoilidea nitida Benoit; (135) radial cell; (136) basal segments of d" antenna;(137) mesonotum. 138-140, E. parma sp. n.; (138) $ antenna; (139) mesonotum; (140) radial cell. 141,mesonotum of E. pallida sp. n. 142, 143, E. perangusta sp. n. ; (142) $ antenna; (143) basal segments ofC? antenna. 144, $ antenna of E. trulla sp. n. AFROTROPICAL EUCOILIDAE 341 145 Figs 145-154 145-147, Eucoilidea trulla sp. n. ; (145) $ gaster; (146) basal segments of cf antenna; (147)radial cell, 148, pronotal plate of E. tyrus sp. n. 149-154, 9, antenna of (149) Hexacola absensa sp. n.;(150) H. pallida sp. n.; (151) ti. atropos sp. n.; (152) H. zama sp. n.; (153) H. fringa sp. n.; (154) H.quisnama sp. n. 342 J. QUINLAN 161 163 164 Figs 155-164 155-161, $ antenna of (155) Hexacola quinquedavata sp. n.; (156) H. octoclava sp. n.;(157) H. hexatoma (Hartig); (158) H. compacta sp. n.; (159) H. bifaria sp. n.; (160) H. septemius sp. n.;(161) H. amantia sp. n. 162, 163, H. amantia sp. n.; (162) pronotal plate; (163) basal segments of C?antenna. 164, fore wing of//, atropos sp. n. AFROTROPICAL EUCOILIDAE 343 Figs 165-176 165, basal segments of c? antenna of Hexacola bifaria sp. n. 166, pronotal plate of H.compacta sp. n. 167, 168, mesonotum of (167) H. fringa sp. n.; (168) H. hexatoma (Hartig). 169, 170,basal segments of C? antenna of (169) H. hexatoma (Hartig); (170) H. octoclava sp. n. 171 , 172, forewingof (171) H. pallida sp. n. ; (172) H. quisnama sp. n. 173, 174, scutellum of (173) H. quisnama sp. n. ; (174)H. zama sp. n. 175, gaster of Kleidotoma arbitra sp. n. 176, scutellum of K. morsum sp. n. 344 J. QUINLAN Figs 177-184 177, forewing of Kleidotoma morsum sp. n. 178, 179, $ , antenna of (178) K. nigrans sp. n.(179) K. morsum sp. n. 180, scutellum of K. nigrans sp. n. 181 , lateral view of head and pronotum of K.ventosus sp. n. 182, $ antenna of K. ventosus sp. n. 183, pronotal plate and occiput of K. erebus sp. n.184, gaster of K. erebus sp. n. 191 AFROTROPICAL EUCOILIDAE186 345 90 Figs 185-194 185-187, scutelium of (185) Kleidotoma erebus sp. n.; (186) K. arbitra sp. n.; (187) K.conica sp. n. 188-193, $ antenna of (188) K. distenda sp. n. ; (189) K. strigosa sp. n. ; (190) K. favus sp. n. ;(191) K. nitidiuscula sp. n.; ;192) K.fimbriata sp. n.; (193) K. eala sp. n. 194, scutellum of AT. norma sp. n. 346 J. QUINLAN 1 95 200 Figs 195-201 195, 196, Kleidotoma norma sp. n.; (195) $ antenna (196) forewing. 197-199, K. bifurcatasp. n. ; (197) scutellum; (198) forewing; (199) pronotal plate. 200, 201, K. conica sp. n. ; (200) $ antenna;(201) $ gaster. AFROTROPICAL EUCOILIDAE 347 Figs 202-208 202, forewing oi Kleidotoma distenda sp. n. 203, $ antenna of K. elongula sp. n. 204, $antenna of K. arbitra sp. n. 205, scutellum of K. favus sp. n. 206, $ gaster of K. fimbriata sp. n. 207,pronotal plate of K. morsum sp. n. 208, forewing of K. nigrans sp. n. 348 J. QUINLAN 214 Figs 209-218 209, 210, Kleidotoma nitidiuscula sp. n. ; (209) pronotal plate; (210) forewing. 211, pronotalplate of-K. norma sp. n. 212, forewing of K. strigosa sp. n. 213, 214, Nordlanderia plowa sp. n.; (213)lateral view efface; (214) mesopleura. 215, 216, N. plowa sp. n.; (215) $ gaster; (216) $ antenna. 217,218, mesonotum of (217) N. pallida sp. n. ; (218) N. plowa sp. n. AFROTROPICAL EUCOILIDAE 349 Figs 219-226 219, $ antenna ol Nordlanderia acis sp. n. 220, gaster of N. pallida sp. n. 221, 222, N. adssp. n.; (221) lateral view of face; (222) $ gaster. 223, pronotal plate of N, pallida sp. n. 224-226,Rhoptromeris heptoma (Hartig); (224) forewing; (225) basal segments of O" antenna; (226) scutellum. 350 J. QUINLAN Figs 227-236 227, 228, Rhoptromeris heptoma (Hartig) ; (227) gaster; (228) pronotal plate. 229, 230, $antenna of (229) R. zeus sp. n.; (230) R. oeta sp. n. 231, scutellum of R. oeta sp. n. 232, 233, pronotalplate of (232) R. crito sp. n.; (233) R. pagasa sp. n. 234, mesonotum of R. temesa sp. n. 235, 236, $filiform antenna of (235) Rhoptrmeris sp.; (236) R. rutshuris sp. n. AFROTROPICAL EUCOILIDAE 351 Figs 237-245 237-239, Rhoptromeris afer sp. n.; (237) $ antenna; (238) scutellum; (239) forewing.240-243, $ antenna of (240) R. abba sp. n.; (241) R. connatus sp. n.; (242) R. diversa sp. n. 243, basalsegments of cf antenna of R. abba sp. n. 244, 245, scutellum of (244) R. diversa sp. n. ; (245) R. connatussp. n. J. QUINLAN 255 Figs 246-256 246, 247, Rhoptromeris bicolor sp. n.; (246) pronotal plate; (247) $ antenna. 248, $antenna of/?, punctata sp. n. 249, 250, R. zetes sp. n.; (249) $ gaster; (250) pronotal plate. 251, 252, R.persius sp. n.; (251) 9 antenna; (252) radial cell. 253, $ antenna of/?, equalis sp. n. 254, 9 gaster of/?.bupalus sp. n. 255, 256, $ antenna of (255) /?. bupalus sp. n. ; (256) /?. attis sp. n. AFROTROPICAL EUCOILIDAE 353 265 Figs 257-266 257, 258, basal segments of d" antenna of (257) Rhoptromeris attis sp. n. ; (258) R. rufulussp. n. 259, 260, $ antenna of (259) R. enna sp. n. ; (260) R. rufulus sp. n. 261-163, R. cubitalis sp. n. ; (161)radial cell; (262) pronotal plate; (263) scutellum. 264, basal segments of c? antenna of R. pallidus sp. n.265, 266, R. thales sp. n.; (265) radial cell; (266) $ antenna. 354 J. QUINLAN 268 Figs 267-274 267-273, $ antenna of (267) Rhoptromeris crito sp. n.; (268) R. naxos sp. n.; (272) R.cepheus sp. n. 273, 274, scutellum; (273) R. naxos sp. n. ; (274) R. crito sp. n. AFROTROPICAL EUCOILIDAE 355 Figs 275-284 275, 276, Rhoptromeris navius sp. n.; (275) pronotal plate; (276) radial cell. 277-279, $antenna of (277) R. oeta sp. n. ; (278) R. pagasa sp. n. ; (279) R. sinis sp. n. 280, radial cell of R. temesa sp.n. 281, 282, $ antenna of (281) Stentorceps tubicen Quinlan; (282) Rhoptromeris temesa sp. n. 283, 284,scutellum of (283) R. temesa sp. n. ; (284) Stentorceps tubicen Quinlan. 356 J. QUINLAN Figs 285-293 285-289, Stentorceps tubicen Quinlan; (285) mesopleura; (286) cf gaster; (287) radial cell;(288) Q" basal segments of antenna; (289) frontal view of face. 290, Rhoptromeris equalis sp. n.; (290)pronotal plate. 291 , Rhoptromeris sp. d" , basal segments of antenna. 292, 293 , Trichoplasta tanganyiken-sis (Weld); (292) mesonotum; (293) $ , gaster. AFROTROPICAL EUCOILIDAE 357 Figs 294-303 294-296, Trichoplasta equalis sp. n. ; (294) radial cell; (295) lateral view of pronotum; (296)$ antenna. 297, 298, scutellum of (297) T. medlia sp. n. ; (298) T. rufus sp. n. 299, 300, T. medlia sp. n. ;(299) radial cell; (300) pronotal plate. 301, $ antenna of T. medlia sp. n. 302, 303, T. rufus sp. n.; (302)pronotal plate; (303) 9 antenna. 358 J. QUINLAN 309 Figs 304-312 304, 305, Trichoplasta narrata sp. n. ; (304) cf , basal segments of antenna; (305) $ antenna.306-308, T. conica sp. n.; (306) $ antenna; (307) scutellum; (308), cf basal segments of antenna.309-311, T.filiformis sp. n.; (309) antenna; (310) scutellum; (311) cf basal segments of antenna. 312,$ gaster of T. conica sp. n. AFROTROPICAL EUCOILIDAE 359 317 Figs 313-322 313, 314, Trichoplasta contrasta sp. n. ; (313) cf basal segments of antenna; (314) scutellum.315-317, T. rufus sp. n.; (315) radial cell; (316) $ antenna; (317) cf basal segments of antenna. 318,radial cell of T. brevispina (Masner). 319, $ antenna of T. unicolora sp. n. 320, scutellum of T.brevispina (Masner). 321, 322, $ antenna of (321) T. contrasta sp. n.; (322) T. medlia sp. n. 360 J. QUINLAN 330 328 Figs 323-331 323-325 , pronotal plate of (323) Trichoplasta sp. ; (324) T. novema sp. n. ; (325) T. octonarussp. n. 326, 327, radial cell of (326) T. medlia sp. n. ; (327) T. octonarius sp. n. 328-330, $ antenna of (328)T. novema sp. n.; (329) T. tanganyikensis (Weld); (330) T. octonarius sp. n. 331, C? basal segments ofantenna of T. medlia sp. n. AFROTROPICAL EUCOILIDAE 361 332 336 337 Figs 332-339 332-334, Trichoplasta novema sp. n. ; (332) scutellum; (333) radial cell; (334) $ lateral viewof thorax and gaster. 335, 336, T. longispina (Masner); (335) scutellum; (336) $ antenna. 337, 338, T.bicolorsp. n.; (337) $ antenna; (338) scutellum. 339, scutellum of T.filiformis sp. n. 362 J. QUINLAN 343 Figs 340-348 340, 341, Trichoplasta filiformis sp. n.; (340) cf basal segments of antenna; (341) $antenna. 342-344, T. quinclava sp. n. ; (342) radial cell; (343) $ antenna; (344) pronotal plate. 345, 346,T. gracilicornis (Kieffer); (345) scutellum; (346) $ antenna. 347, 348, radial cell of (347) T. testacea sp.n.; (348) T. extensus sp. n. AFROTROPICAL EUCOILIDAE 363 358 Figs 349-358 349-351, Trichoplasta testacea sp. n.; (349) $ antenna; (350) scutellum; (351) pronotalplate. 352, 353, T. unicolora sp. n.; (352) C? basal segments of antenna; (353) $ antenna. 354-357, T.octonarius sp. n.; (354) $ antenna; (355) radial cell; (356) scutellum; (357) pronotal plate. 358, $antenna of T. zeus sp. n. 364 J. QUINLAN Fig. 359 Trybliographa rapae Westwood, $ . AFROTROPICAL EUCOILIDAE 365 362 Figs 360-362 360, 361 , basal segments of C? antenna of (360) Cothonaspis sp. ; (361) Leptopilina sp. 362,Leptop ilina heterotoma (Thomson), $ antenna. Index Principal references are in bold, synonyms are in italics. abba 292, 294aberrant 290absensa 273acis 289 advena261,262afer292,294Afrodontaspis 251, 253Afrostilba 259, 269agis 292, 293, 295Agroscopa 280Alloxystinae 245amantia 273, 274Anacharitinae 245anisomera 280Aphiloptera 280Aphyoptera 280arbitra281,282Aspicerinae 245atropos273,274attis293,295Aulacidea 245Austrocynipinae 245Aylax 245 basilewskyi31Q, 319 bicolor, Rhoptromeris 292, 296 bicolor, Trichoplasta 311, 312bifaria273,275bifurcata281,282Bothrochacis 247, 248, 249, 252brevispina 31 1,313bucca261,262,263bupalus 293, 296 canadensis 259 cepheus291,297 Charipidae 245 Chrestosema 247, 249 clava 257 compacta273,275 compressa 261, 262, 263 Coneucoela 247, 248, 252 conica, Kleidotoma 281, 283 conica, Trichoplasta 311, 312, 313 connatus 292, 297 contrasta311,312,314 conversa 260, 262, 264 Cothonaspis 247, 248, 250, 252, 255 crito291,297cubitalis293,298Cynipidae 245 Cynipinae 245 dad 244 Daruna 249, 252Didyctium 247Dieucoila 247Diglyphosema 247, 251Disorygma247,251distenda281,283diversa 292, 299dubia261,264dulcis255,256 eala281,283 Ealata243,247,251,257 ealis255,256 elongula281,284 emarginatus 280 enna 293, 299 equalis, Rhoptromeris 292, 300 equalis, Trichoplasta 311, 312, 314 erebus 28 1,284 eucera 290 Eucoila 247, 248, 249, 252 Eucoilidae245,250 Eucoilidea247,251,259 366 Eucoilinae 250Eutrias 247extensus 31 1,315extraria 260, 265 favus281,285fenervae 256fetura261,262,265Figitidae 245filiformis311,312,315fimbriata281,285fringa273,276furcula261,262,266 Ganaspidium 243Ganaspis247,248,253giraudi 255 Glauraspidia 247, 249, 253gracilicornis 311, 316Gronotoma 247 halophila 280 hebe293,300 helgolandica 280 heptoma293,300 heterotoma248,280 Hexacola 243, 247, 248, 253, 272 Hexamerocera 290 Hexaplasta 272 hexatoma 272, 273, 276 Himalocynipinae 245 Hypodiranchis 247 Ibaliidae 245inustipennis 280Isocolus 245 Kleidotoma 243, 245, 247, 248,252, 280 lacerta 260, 266Iana260,267lanatus 254Ieptis260,267Leptolamina 247Leptopilina 247, 248, 250, 253Liopteridae 245Liopterinae 245longispina 311, 316 J. QUINLAN marcellus260,262,268marica 257, 258marshalli 244mauri 261 , 268Mesocynipinae 245Microstilba 247Miomoera 290montana 285morsum 281 , 286mundata 248 narrata 31 1,312, 317 navius293,301 naxos293,302 Nedinoptera 280 nigra 280 nigrans281,286 nitida259,260,262,269 nitidiuscula 281, 287 Nordlanderia 243, 247, 251, 288 norma281,287 novema311,318 Oberthuerellinae 245octoclava 273, 277octonarius 310, 318Odonteucoila 247, 248, 252oeta291,302pagasa291,302pallida, Eucoilidea 261, 269pallida, Hexacola 273, 277pallida, Nordlandria 289pallidus293,303parma 260, 270Paramiomoea 247Pentacrita 280Pentamerocera 247, 256pentatoma255,256perangusta 261, 262, 270persius 292, 304picicrus 272plowa 288, 290Pseudeucoila 248Psedopsichacra 247psiloides 280punctata 293, 304 quinclava 31 1,319quinqueclavata 273, 278 quisnama273,278 retusa 280 Rhoptromeris 243, 245, 247, 248, 249, 252, 290Rhynchacis 280rufiventris 290rufulus293,304rufus 310, 312, 319rutshuris 292, 305rwanki291,305 saba 257, 258 Schizosema 280 septemius273,279 sinis293,306 Stentorceps 244, 247, 248, 252, 309 striatissima253,254strigosa281,287subnuda 248 tanganyikensis 310, 319 temesa 292, 306 testacea 31 1,320 Tetrahoptra 280 Tetramerocera 247 Tetratoma 280 thales293,307 Trichoplasta 247, 248, 251, 310 Trissodontaspis 248 trulla 260, 271 Trybliographa 243, 245, 247, 248, 249, 253tubicen 309tyrus260,262,271 unicolora311,312,321urundiensis 261, 272 velia292,307ventosus281,288 Zaeucoila 247 zama 273, 279 zetes 292, 308 zeus, Rhoptromeris 291, 308 zeus, Trichoplasta 311, 321 Occasional Papers on Systematic Entomology New Series The economic importance of insects, and the enormous number of species, have resulted in avast literature written in many languages; that which is of direct economic importance andrecently published can, to an increasing extent, be searched by using computerized data bases,but a great amount of more general information is unlikely to be available so readily in thenear future. The objective of this new occasional series is to make available in hard copy someof the basic data that is essential to the preparation of comprehensive accounts of the worldinsect fauna. The papers have been fully researched bibliographically and consist of checklistsof nominal taxa, and faunal lists with information on host plants and localities, based mainlyon the collections and libraries of the British Museum (Natural History). No. 1. A checklist of Neotropical arctiine and pericopine tiger moths. A. Watson & D. T. Goodger 72pp. inc. 4 colour plates 27 February 1986 No. 2. An annotated checklist of the Carabidae (including Cicindelinae, Rhysodinae and Paussinae) recorded from Borneo. N. E. Stork 26pp. , 1 map 24 April 1986 Catalogues of BM (NH) Bulletins and Books free on request to Publication Sales. Titles to be published in Volume 52 The sandflies of Egypt (Diptera: Phlebotominae) By R. P. Lane Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae) ByG. S.Robinson A revision of the European Agathidinae (Hymenoptera: Braconidae) By G. E. J. Nixon A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revision of certain genera By J. Quinlan Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester