er erece ee Vets ve be virgetete . “ale gte A I) eo ‘ het arnnitgre Feta eit plaretete> 136 a ee + ay he x eet) * on Bs oH: erppheaotenspitnaes Sete retreat tee Nye ad Git axe He-t54 vith bckoete ‘fete! bh oh oe tet ste iS 4, "er tees Zt ie biti Pata intels ois faa 4 Mi t if] j afl i * j ’ i * s P { Fi an ‘ - > ) ~ nia \ Va y @ Ls eo fei ye waa vie hn” y ae Waa et J) ca a2 A REVISION OF THE LEUCOSPIDAE ~~ (HYMENOPTERA : CHALCIDOIDEA) OF THE WORLD Z. BOUCEK BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Supplement 23 LONDON : 1974 A REVISION OF THE LEUCOSPIDAE (HYMENOPTERA : CHALCIDOIDEA) OF THE WORLD BY ZDENEK BOUCEK Commonwealth Institute of Entomology Pp 1-241; 272 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY LONDON : 1974 Supplement 23 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), wstituted im 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Paris will appear at wregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Depariment. This paper is Supplement 23 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) Suppl. © Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 16 May, 1974 Price £11.30 A REVISION OF THE LEUCOSPIDAE (HYMENOPTERA : CHALCIDOIDEA) OF THE. WORLD By Z. BOUCEK CONTENTS Page SYNOPSIS 3 INTRODUCTION 4 MORPHOLOGICAL TERMS AND MEASUREMENTS 5 ACKNOWLEDGEMENTS 7 ABBREVIATIONS OF DEPOSITORIES 7 LEUCOSPIDAE WALKER 9 Biology : : : : ‘ ; : : : , 14 Zoogeography : ; : : - - . 16 Classification within the family 16 Key to the genera 16 Polistomorpha Westwood 18 Leucospis Fabricius . ¢ : : : : é : : 28 The New World species . A : : : : 33 The African, Madagascan and Mediterranean species : : - IOI The West Palaearctic species . , : : : - : : 140 The Asiatic and Australian species . : : : ; : : 155 Neleucospis gen. n. . . : ‘ ' . . ; ; P 210 Micrapion Kriechbaumer . F P ; ” : ; : . 211 MISPLACED TAXA . : : . : : 229 NoMINA NUDA AND OTHER UNAVAILABLE NAMES : - . : 230 Host-PARASITE CATALOGUE ‘ : : : - ; ; ; 230 REFERENCES . ‘ R : , ‘ ‘ , : ; 232 PARASITE INDEX 4 P : : . ; : ‘ : 238 SYNOPSIS In this revision, based on a study of material from institutions all over the world, four genera are recognized: Polistomorpha Westwood (7 tropical American species, of which 3 are new), Leucospis Fabricius (109 species, of which 31 are new, from all the warmer regions of the world), Neleucospis gen. n. (I new West African species) and Micrvapion Kriechbaumer (12 African species, of which 8 are described as new, and 1 Madagascan species). The type-material of more than 150 names was examined (including 57 holotypes), and 91 lectotypes are newly designated. Keys are given to all genera and species; in Leucospis three separate keys are given for the American, African and Asiatic-Australian species, respectively, the Palaearctic species being included in both the keys to the Old World species. Two generic, and 58 specific and subspecific names are newly synonymized, and 5 new combinations are proposed. 4 Z. BOUCEK INTRODUCTION THE Leucospids include most of the largest insects among Chalcidoidea, but in spite of this our knowledge of them has been rather poor. The earlier history of the group was well reviewed by Schletterer (1890), in his excellent monograph. The most important contributions before him were a paper on the European species by Klug (1814), another with descriptions of some species from north-east Africa and Arabia (Klug, 1834), the two reviews of the world species by Westwood (1834; 1839) and later a paper on the North American and Mexican species by Cresson (1872). Most publications, especially the numerous descriptions by Walker (1834-1871), did not include comparisons with the previously known species, or any keys (exception: Cresson, 1872). Schletterer provided keys to the three genera then included (two of them with only one and two species, respectively) and in Leucospis to 36 species, whilst about the same number of further species were quoted from the original descriptions. He studied all the types and material available, mainly from various European museums (but not, for example, from London and Paris), and aptly and critically reviewed the existing knowledge, including the known biological data, and for the first time evaluated also the variation in colour and structure. Partly as a result of this, he dropped many names into synonymy, in most cases rightly so. Another three decades were reviewed by Weld (1922) in a similar way, although to a much lesser extent, as the aim of her work was much more restricted. Against Schletterer she had the advantage of a better knowledge of some American species, the types of which she could examine. Working in the U.S. National Museum, Washington, shortly after the first world war, she had, however, almost no contact with Europe. Weld’s work, although it must not be regarded as another revision, is good, but many of her species also proved to be synonyms. The post-Schletterer authors mostly contributed in smaller papers, with single or few descriptions or other information, but Ducke (1906), for example, revised the Polistomorpha species. Later, particularly after Weld’s work, a few local faunas were worked out. Thus Berland (1934) revised the French species of Leucospis, Mani (1937) the Indian species, Steffan (1948) treated the African Micrapion, Nikolskaya (1952) keyed out the Leucospis of the U.S.S.R. and later on (1960) treated them more comprehensively in the Fauna of U.S.S.R., Erdés (1955) keyed out the Hungarian species, Boutek (1959) the West Palaearctic ones, Ceballos (1959) the Spanish ones, Habu (1962) worked out comprehensively the two Japanese species and Porter (1972) briefly the Floridan species. Of all the papers which include valuable criticism, corrections and other information, perhaps the most important is that of Masi (1935). Otherwise descriptions of the Australian species were provided mainly by Girault, whilst those of the other parts of the world are more scattered and are mentioned with the relevant taxa and more fully under References. These include also some recent catalogues, viz. of the Indian species (Mani, 1938), of the North American ones (Peck, 1963) and of the Argentinian species (De Santis, 1967). The taxonomic aims of the present work have been mainly to reach a better understanding of the existing (described or undescribed) taxonomic units of © REVISION OF LEUCOSPIDAE 5 Leucospidae and to draw up their present classification. The re-evaluation of the old taxa could be achieved only by a review of the existing knowledge and by checking it, at the same time, against all the available rich material. This, together with the biological information (including distribution), was supposed to reveal something of the actual range of variation and thus of the natural limits of various taxa and of the gaps between them. Only then could I be relatively sure to which taxa the types eventually belonged, apart from their nomenclatural value. In a few species, however, the available material seems to be still inadequate for safe conclusions. The limits, relationship, variation and other aspects of taxonomy including the biological data are eventually treated, where necessary, with the individual taxa, including the family taxon as a whole. In a general scheme the valid name is followed by the quotation and the synonymy, then by the eventual information on the type material, including its nomenclatural and taxonomic aspects, problems concerning the intraspecific variation and the interspecific relations. In a few cases several infraspecific forms are recognized, in Leucospis affints and L. histrio on the subspecific level, in which case the discussion is followed by a key to the subspecies and then each subspecies is treated as a separate unit. Biological data and general distribution (mostly in terms of countries) of species (or subspecies) are treated in separate paragraphs, followed by an account of the material examined. In the synonymy only names having some bearing on the nomenclature ie. names available under the International Code of Zoological Nomenclature are mentioned. Misidentifications are therefore referred to only where a name is partly or entirely based on them. MORPHOLOGICAL TERMS AND MEASUREMENTS Some of the morphological terms, including all less common ones, are explained here and in a few figures (mainly Text-figs 1-15), together with the measurements used in descriptions. In spite of the relatively large size of these insects exact measurements are sometimes necessary, although their variation may be greater than known at present. There is no point in giving absolute measurements (e.g. in microns), as their main value is in relation to the measurements of the other parts. The normal position of the head is taken as that with the mandibles and other mouth parts directed downwards. Consequently the length of head is its maximum thickness in antero-posterior direction (in dorsal view; Text-fig. 2), its breadth (or width) the distance between the outer margins of the eyes (less pubescence) and its height is measured from the uppermost point, usually on the occipital carina, down to the lowermost point of the lower clypeal margin (Text-fig. 1). The head usually has dorsally an area in front of the occipital carina delimited anteriorly by the frontal protuberances. As the part in front of the ocelli is called frons, this area is called fronto-vertex; its breadth is measured as the minimum distance between the inner margins (orbits) of the eyes, at about the level of the median ocellus. Ocellar triangle: the width means the distance between the outer margins 6 Z. BOUCEK of the lateral ocelli, the height is the distance between the anterior margin of the median ocellus and a line through the posterior margins of the lateral ocelli. POL, or post-ocellar length, is the distance between the inner margins of the lateral ocelli, whilst OOL, or ocell-ocular length, is measured between the outer margin of the lateral ocellus and the eye margin. Measurements of the eye give the maximum and minimum diameters, in an antero-lateral view. The scrobes (united antennal or supra-antennal pits) are well delimited by the scrobal carina; the width is the maximum distance between the outer scrobal carinae, and the height is the distance between the upper scrobal carina (at the ocellus) and the lower edge of the antennal forwli (the torulus is the actual hole in which the antennal radicula is inserted). Lower face: the height is measured between the lower margins of the toruli and the lowermost point of the clypeus; the breadth is the minimum distance between the eyes below the antennal insertions. The malar space is measured between the lower extremity of the eye and the mouth margin in a vertical line, i.e. in front of a trace of the malar (genal) suture, not more posteriorly on the gena, where the mouth margin sometimes curves slightly upwards. The mouth is measured between its lateral corners, usually easy to see in a ventro-facial view, outside of the mandibles. The latter always have a well separated lower tooth (less distinct only in Micrapion), by a triangular notch or by a semicircular gap; their inner margin above the notch may be straight or emarginate to form another two teeth. The well developed labio-maxillary complex is rarely used in descriptions, although it shows a few good characters, mainly in the subdivision of the apical part of maxilla, beyond stipes and the membranous part to which the maxillary palpi are attached (I am not sure whether the apical part seen for example in Text-fig. 11 is actually homological with lacinia). The thorax is treated together with the propodeum, i.e. originally the first abdominal segment (mesosoma of some authors). The pronotum often bears transverse keels, carinae; the complete set (Text-fig. 10) includes the carinate hind margin, a premarginal carina and a discal carina. Sometimes a distinct arcuate swelling is developed anteriorly connecting the anterior corners (shoulders). The mesoscutum sometimes shows traces, in form of shallow and broad longitudinal depressions on either side of the middle, diverging forwards; these are the notaular furrows or notauli, whilst the pavapsidal furrows are indicated by short, often slot- like vestiges inside the hind corners of the mesoscutum. The breadth of the scutellum is measured about in its middle and does not include the axillae (Text-fig. 8). The dorsellum is the central raised part of the metanotum. The propodeum usually has the median carina and sub-lateral carinae, called plicae, inside the postspiracular furrow. The thoracic pleurum (I have previously used ‘pleura’ as singular and ‘pleurae’ as plural) is subdivided as shown in Text-fig. 4. The stigmal vein of the fore wing emits a branch subparallel to the anterior margin and this branch is called uncus, whilst the apex of the vein, beyond the uncus, where present (absent in Text-fig. 20) is called the terminal processus. The gaster of the female is shown in Text-fig. 4. It has a strongly reduced petiole and fergites counted as 1-6, the sixth bearing a good landmark in the spiracles and followed by the epipygium (the following two tergites fused together) bearing REVISION OF LEUCOSPIDAE 7 cerci. Some of the tergites in the female are reduced and more or less hidden (but shown in Text-figs 4 and 252), as well as the basal sternites, of which only the last, called the hypopygiwm is always conspicuous. In the male the tergites and sternites can be better observed, but tergites 3-6 are more or less fused into a carapace; its hind corners often protrude as teeth or auricles. The sternites are seven in number, i.e. the last is always counted as the seventh. ACKNOWLEDGEMENTS My work has been facilitated by my recent new position with the Commonwealth Institute of Entomology, close to the collections and the library of the Department of Entomology of the British Museum (Natural History), the Keeper and staff of which I wish to thank for all the facilities offered. Also many other colleagues from many institutions all over the world (see also the list of depositories below) very kindly assisted me in submitting the types and other material for study, or in various other ways, in particular the following: Dr D. P. Annecke (Pretoria), Dr F. Bachmaier (Munich), Dr B. D. Burks (Washington), Dr E. McC. Callan (Canberra), Mr E. C. Dahms (Brisbane), Prof. H. V. Daly (Berkeley), Prof. H. E. Evans (Cambridge, U.S.A.), Mr M. J. Gijswijt (s’Graveland, Netherlands), Dr M. W. R. de V. Graham (Oxford), Dr E. Kénigsmann (Berlin), Prof. M. S. Mani (Agra), Dr L. Masner (Ottawa), Prof. O. W. Richards (London), Dr E. F. Riek (Canberra), Rev. A. Watsham (Salisbury, Rhodesia; also for his kind linguistic help) and Prof. J. T. Wiebes (Leiden). ABBREVIATIONS OF DEPOSITORIES AM, Grahamstown Albany Museum, Grahamstown, Cape Province, South Africa (C. F. Jacot-Guillarmod) AM, Sydney Australian Museum, Sydney, N.S.W., Australia (G. A. Holloway) ANS, Philadelphia Academy of Natural Sciences, Philadelphia, Pennsylvania, U.S.A. (Dr D. C. Rentz) BBM, Honolulu Bernice Bishop Museum, Honolulu, Hawaii, U.S.A. (Dr J. L. Gressitt) BMNH British Museum (Natural History), London CAS, San Francisco California Academy of Sciences, San Francisco, California, U.S.A. (P. H. Arnaud) CIS, Berkeley California Insect Survey, University of California, Berkeley, California, U.S.A. CM, Pittsburgh Carnegie Museum, Pittsburgh, Pennsylvania, U.S.A. (G. E. Wallace) CSIRO, Canberra Division of Entomology, Commonwealth Scientific and Industrial Research Organisation, Canberra City, A.C.T., Australia CU, Ithaca Cornell University, Department of Entomology, Ithaca, New York, U.S.A. (Dr L. L. Pechuman) DEI, Eberswalde Deutsches Entomologisches Institut, now Abteilung Taxonomie der Insekten, Institut fiir Pflanzenschutzforschung, Eberswalde, East Germany (Dr J. Oehlke) DE, Davis Department of Entomology, University of California, Davis, California, U.S.A. (Prof. R. M. Bohart, E. E. Grissell) DE, Riverside Department of Entomology, University of California, Riverside, California, U.S.A. (S. Frommer) 8 EI, Zurich EIHU, Sapporo ELKU, Fukuoka EM, East Lansing ERI, Ottawa EU, Matsuyama FCNM, La Plata IBUR, Rio de Janeiro IEA, Portici IML, Tucuman IRSNB, Brussels IZU, Naples LE, Wageningen MCSN, Genoa MCZ, Cambridge MIZS, Turin MHN, Geneva MNHN, Paris MNHU, Berlin MP, Belem MRAC, Tervuren MZU, Florence NCI, Pretoria NIAS, Tokyo NM, Bulawayo NM, Pietermaritzburg NM, Prague NM, Vienna NR, Stockholm QM, Brisbane RNH, Leiden SAM, Cape Town Z. BOUCEK Entomologisches Institut der E.T.H., Ziirich, Switzerland (Prof. W. Sauter) Entomological Institute, Faculty of Agriculture, Hokkaido University, Sapporo, Japan Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan Department of Entomology, Michigan State University, East Lansing, Michigan, U.S.A. (Dr R. L. Fischer) Entomology Research Institute, Ottawa, Ontario, Canada (Dr C. M. Yoshimoto) Entomological Laboratory, College of Agriculture, Ehime University, Matsuyama, Japan (H. Taguchi) Facultad de Ciencias Naturales y Museo, Universidad Nacional, La Plata, Argentina (Prof. L. De Santis) Instituto de Biologia, Universidade Federal Rural da Rio de Janeiro, Guanabara, Brazil (Prof. C. R. Gongalves). Istituto di Entomologia Agraria, Portici, Italy (Dr G. Viggiani) Istituto de Miguel Lillo, Miguel Lillo, Prov. Tucuman, Argentina (Prof. A. Willink) Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (Dr P. Dessart) Istituto di Zoologia dell’Universita di Napoli, Naples, Italy Laboratorium voor Entomologie van de Landbouwhogeschool, Wagen- ingen, Netherlands (Drs K. W. Zwaart) Museo Civico di Storia Naturale, Genoa, Italy (Prof. E. Tortonese) Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. (Prof. H. E. Evans) Museo ed Istituto di Zoologia Sistematica, Universita di Torino, Turin, Italy (Prof. U. Parenti) Muséum d’Histoire Naturelle, Geneva, Switzerland (Dr C. Besuchet, Dr I. Lobl) Muséum National d’Histoire Naturelle, Paris, France (Mme S. Kellner- Pilault & Dr J. R. Steffan) Museum fiir Naturkunde der Humboldt-Universitat, Berlin, East Germany Museu Paraense ‘Emilio Goeldi’, Belem, Para, Brazil (Dr R. Arlé) Musée Royal d’Afrique Centrale, Tervuren, Belgium (Dr J. Decelle) Museo Zoologico della Specola, Universita degli Studi, Florence, Italy National Collection of Insects, Institute of Plant Protection, Pretoria, Transvaal, South Africa Nationa] Institute of Agricultural Sciences, Nishigahara, Tokyo, Japan (Dr A. Habu) National Museum of Rhodesia, Bulawayo, Rhodesia (Dr E. C. G. Pinhey & Mr F. C. de Moor) Natal Museum, Pietermaritzburg, Natal, South Africa (Dr M. E. Irwin) Entomologické oddéleni, Narodni Museum, Praha-Kunratice, Czecho- slovakia Naturhistorisches Museum, Vienna, Austria (Dr M. Fischer) Naturhistoriska Riksmuseet, Stockholm, Sweden (Dr K. J. Hedqvist) Queensland Museum, Fortitude Valley, near Brisbane, Queensland, Australia (E. C. Dahms) Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands (P. J. van Helsdingen) South African Museum, Cape Town, South Africa (A. J. Hesse) REVISION OF LEUCOSPIDAE 9 SM, Lawrence Snow Entomological Museum, University of Kansas, Lawrence, ’ Kansas, U.S.A. (Prof. C. D. Michener) SMT, Dresden Staatliches Museum fiir Tierkunde, Dresden, East Germany (Frau R. Eck) TM, Budapest Termeszéttudomanyi Mizeum Allattara, Budapest, Hungary (Dr J. Papp, Prof. G. Szelényi) TM, Pretoria Transvaal Museum, Pretoria, South Africa (J. van Reenen) Townes H. & M. Townes Collection, American Entomological Institute, Ann Arbor, Michigan, U.S.A. (Dr H. Townes) UM, Oxford Hope Department of Entomology, University Museum, Oxford, England USNM United States National Museum, Washington, D.C., U.S.A. (Dr P. D. Hurd) UZM, Copenhagen Universitetets Zoologiske Museum, Copenhagen, Denmark (Dr B. Petersen) Watsham Rev. A. Watsham, Salisbury, Rhodesia ZI, Leningrad Zoological Institute, Academy of Sciences of U.S.S.R., Leningrad, U.S.S.R. (Dr V. A. Trjapitzin) ZM, Amsterdam Zodlogisch Museum, now Instituut voor Taxonomische Zoologie, Amsterdam, Netherlands ZS, Munich Zoologische Staatssammlung, Munich, West Germany (E. Diller) ZSI, Calcutta Zoological Survey of India, Calcutta, India (Dr A. P. Kapur) LEUCOSPIDAE Walker Leucopsidae Walker, 1834 : 13. Type-genus: Leucopsis Dumeril (= Leucospis Fabricius). Leucospidae Walker; Haliday, 1839 : ii. [Justified emendation.]} Leucaspoidae Forster, 1856 : 18, 20. Type-genus: Leucaspis Burmeister (= Leucospis Fabri- cius). Leucospidinae Cameron, 1883 : 76. [Unjustified emendation.]} Leucospinae Walker; Howard, 1886 : 197. Leucospididae Cameron; Brues & Melander, 1932 : 485. In the past both spellings Leucospidae and Leucospididae were used. As there is no definite proof that the ending of the name Leucospis is derived from the Greek aspis, aspidis or ops, opos (cf. Schletterer’s comments, 18go0 : 144), the shorter form is preferred. The family-group name was first used by Walker (1834 : 13) as ‘Leucopsidae’ (from the unjustified emendation of Leucospis to Leucopsis, see generic synonymy) and the group was regarded for some time as a family (for example by Forster, 1856 : 18, 20), but later on it was lowered to subfamily rank. A slight taxonomic change was introduced by Ashmead (1899 : 247), who, while regarding the other major groups of the Chalcidoidea as families, divided Chalcididae into two subfamilies: Leucospidinae and Chalcidinae, thus stressing the similarity of the two groups. The major groups of Chalcidoidea were treated again as subfamilies by Schmiedeknecht (1909), who again levelled Leucospinae with them, although he largely followed Ashmead. This status has been retained by the more recent authors, except that most major groups, including the Leucospids, have been regarded as families. The question of whether this is justified is partly outside the scope of this paper, as it largely depends on the relative weighting of various 10 Z. BOUCEK characters, the gaps between the relevant groups and the measure which is taken in comparing the families. I feel that family rank for Leucospidae is much more justified than in, for example, some groups of bees, quite apart from higher animals such as mammals. There are no fossil records to offer any lead as to how old the group may be. Some characters, for example the unspecialized dense pilosity of the wings with many veins indicated by darker shades or folds (cf. also Burks, 1938), seem to point to a relative primitiveness among the Chalcid flies. Some other characters, often regarded as specialized (apomorphic), for example the ovipositor bent over the gaster, may not be of such value; the latter position of the ovipositor is normal in the parasitic Hymenoptera with a long ovipositor in the pupal stage and in Leucospidae it is retained into the adult stage. Body of relatively large size (2-3-16-5 mm), heavily sclerotized, mainly black or brownish, often partly to extensively red (varying extent of rufinism), as a rule with yellow or whitish markings, and in the Americas and the Indo-Australian region often with metallic tinge; in- cluding the gaster relatively coarsely punctured and pubescent, though hairs usually not long. Head densely punctured or rugulose-punctured, face with rather dense short pubescence. Eyes large, pubescent, inner margin more or less emarginate in upper third. Ocelli normal, usually large. Occipital carina mostly developed, often sharp, but temples at lower part of eye and genae terete and strongly receding to conspicuous hypostomal carina; malar sulcus fine or indistinct. Antennal scrobes very deep, carinate at margin, reaching near to median ocellus. Frons above more or less elevated (frontal protuberances) ; flat or slightly convex interantennal area triangular, often with median keel, in almost same plane as its lower part called the supraclypeal area, which is usually well delimited. Clypeus always large, more or less trapezoidal or subquadrangular (Text-figs 1, 7, 15); tentorial pits indistinct; lower margin of clypeus free, usually produced. Labrum not traceable. Mandibles with upper edge hidden (when closed) behind mouth margin (or clypeus), generally with two teeth, the upper tooth then often broad and eventually double. Labio-maxillary complex well developed (Text-figs 11, 17, 18), with long glossa emarginate at apex; labial palpus 3-segmented, maxillary palpus usually rather long and 4-segmented, rarely rudimentary (Polistomorpha). Antennae 13- segmented (Text-fig. 9); scapus at most about 3-5 times as long as broad; pedicellus short; flagellar segments with basal one not reduced to anellus but narrowed at base; funicle counted therefore as 8-segmented and remaining three segments regarded as clava (its apical segment very short, indistinct and often appearing double), although the first segment is well separated, its suture being almost as conspicuous as those between preceding segments. The antennal segmentation was discussed by Habu (1961 : 85-86). Antennae of both sexes subequal, with very short dense pubescence, sensilla minute and not conspicuous. Thorax with large pronotum, often with transverse carinae; sides of collar (pronotal dorsum) subparallel or slightly converging forwards, sometimes concave in middle; anterior corners always conspicuous; lateral panel not high (Text-fig. 4), convex or with subhorizontal depression, posteriorly often with adspiracular emargination. Mesoscutum only rarely with vague notaular depressions, these never groove-like; parapsidal furrows reduced to short (often slit- like) but distinct vestiges posteriorly at lateral corners. Scutellum not subdivided, its hind margin low, rounded or subtruncate; frenum not developed. Axillae very short, their hind part vertical, outer corner more or less tooth-like. Metanotum with well differentiated dorsellum, latter often of characteristic form, sometimes carinate or even toothed at margin. Propodeum punctured or rugose-punctured, always pilose, mostly with distinct median carina and simple sublateral plicae but without further carinae or regular alveolae; spiracles narrowly reniform, situated in anterior part of large sublateral furrows. Prepectus showing as small movable sclerite below anterior end of the unusually elongate spatulate tegula. Mesopleurum with deep depression beneath, to accept mid femur and tibia; upper part punctured, distinctly REVISION OF LEUCOSPIDAE II subdivided in subalar area, upper episternum and upper epimerum; anterior aspect of mesopleurum without any shelf in front of mid coxae but with deep elongate depression on either side above, to accept the postero-lateral edge of pronotum. Metapleurum sub-triangular, often produced above at hind wing. Fore coxa longer than half of femur, anteriorly pubescent, without oblique carina. Fore tibia (Text-figs 92, 93) with distinct tooth at apex, spur curved, its apex shortly (sometimes indistinctly) cleft. Mid coxa relatively short; mid tibia slender, apical spur not long. Hind coxa unusually large, with distinct depression externally between dorsal edge and blunter lateral edge; more or less punctured, at least between lateral and meso-ventral edges; often with dorsal tooth. Hind femur greatly enlarged, its ventral edge, except at base, toothed. Hind tibia arcuate, dorsally rounded, ventrally with percurrent carina and another externo-ventral carina usually not reaching apex; apex of tibia with two spurs (Text-fig. 6) but sometimes produced into a spine which bears only a rudiment of the outer spur on its top (Text-figs 5, 40, 131, 144), whilst the inner spur is normally developed, microscopically pubescent. ‘Tarsi 5-segmented, normal; claws of fore and mid legs pectinate, teeth often different in outer and inner claw (Text-fig. 62); hind claws simple or with very short comb at broadened base. Wings densely pubescent, including lower surface of costal cell, and if pubescence slightly reduced in proximal part of wing, then hairs not forming any special formations; fore wing with long and smoothly curved submarginal vein, very long postmarginal vein, but marginal vein very short, shorter than stigmal vein which often bears a distinct uncus; sclerotized spot present outside of lower corner of basal cell. Hind wing with rather long marginal vein, but this even apically slightly removed from the margin, with three subequal hamuli. Gaster rather broadly sessile, the second abdominal segment (petiole) strongly reduced, mostly hidden but sometimes more apparent, often with transverse carina dorsally. First tergite (postpetiolar tergite) in both sexes large. In female second tergite strongly reduced, mostly hidden under the first (Text-figs 4, 252), consisting of lateral sclerotized discs, connected medially with a broad membrane (which extends greatly at oviposition) ; third tergite very short, its basal impunctate part mostly not exposed, punctured part very narrow; fourth tergite always at least partly exposed, punctured, not long. Fifth tergite the largest, forming broadest part of gaster. Sixth tergite (landmarked well by spiracles) normal only in Polistomorpha, in other genera its exposed part divided in two, showing at sides in front of epipygium; the latter extending along ovipositor. Cerci always present, but very low, disc-like, their setae short. Sheaths of ovipositor often long and then bent upwards and forwards; in that case an ovipositorial furrow (to accept ovipositor) of corresponding length developed on dorsal side of thorax (Text-figs 8, 55). Sternites strongly reduced except the last (hypopygium) which projects far back along ventral side of gaster. In the male, second tergite much shorter than the first but distinct (e.g. Text-figs 31, 80, 120), punctured, well separated from the following tergites which are more or less fused into a carapace. Carapace at base with sublateral keels marking off subvertical epipleura which show better margins of fused tergites; segmentation of carapace often indicated by changing density of puncturation (e.g. Text-figs 114, 115) or by yellow bands (Text-figs 169, 171, 197, etc.), sixth tergite again bearing spiracles. Epipygium readily separated, more or less trans- verse, usually with cross-depression; cerci low. Exposed parts of sternites with strong scleroti- zation, punctured; all well developed, the first often with tooth-like projection ventrad.