Tijdschrift Voor Entomologie

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

DEEL 130 1987

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

Tijdschrift voor Entomologie, deel 130, 1987 NEDERLANDSE ENTOMOLOGISCHE VERENIGING

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TIJDSCHRIFT VOOR ENTOMOLOGIE

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The Journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. Subscription rate: D.Fl. 300,— per year.

Volume 130 appeared on 30.XI.1987

ISSN 0040-7496 INHOUD

Blackith, R. E., & R. M. Blackith. — Tridactyüds and Tetrigids (Orthoptera) from Sulawesi, Indonesia 1 Blommers, L. H. M., zie Mensen, R.V.

Compton, S. G., zie Gardiner, A. J. Gardiner, A. S. G. Compton. — New species of the fig J., & wasp genus Diaziella (Hymenoptera, Chalcidoidea, Sycoecinae) 129 Hensen, R. V,, & L, H. M. Blommers. — Review of the Malagasy species of Belonogaster Saussure (Hymenoptera, Vespidae) 11

Jong, M. R. de. — Taxonomy and Biogeography of Oriental Prasiini. 3. The fatUoqua and parvula groups of the genus Lembeja Distant, 1892 (Homoptera, Tibicinidae) 177 Nijveldt, W. — On four new Palaearctic species of the genus Cecidomyia. (Diptera, Cecidomyiidae) 211 Rossem, G. van. — A revision of western Palaearctic oxytorine genera. Part VI. (Hymenoptera, Ichneumonidae) 49 Russev, B. K. — Ecology, life history and distribution of Palingenia longicauda (Olivier) (Ephemer- optera) 109 Speight, M. C. D. — External morphology of adult Syrphidae (Diptera) 141

Wiederiska, J. — Morphologie der Larven und Puppen einiger Phylidorea- Anen (Diptera, Limoniidae) 33 NOTICE TO CONTRIBUTORS

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INDEX

(Synonyms in italics)

accusator (? Cylloceria) 58 erythropyga (Plectiscidea) 64, 74 Acroblapticus 86 erythrostoma Gmelin (Ichneumon) 98 agitator (Plectiscidea) 64, 75 erythrostoma Gravenhorst (Plectiscus) 98 alpigena (Cylloceria) 54 erythrostoma (Helictes) 93 alpigena (Eusterinx) 94 eurystigma (Plectiscidea) 66, 85 alpigenus (Catomicrus) 94 Eusterinx 88 90 ambulator (Plectiscus) 73 eversorius (Plectiscus) 68 amicalis (Plectiscidea) 71 64, fabularis (Helictes) 98, 102 Apoclima 53 flavicoxis (Plectiscus) 76 aquilonigena (Eusterinx) 89, 90, 97 flavipes (Gnathochorisis) 88 ardentis (Proclitus) 59 60 flavipes (Hemiphanes) 50 argutula (Eusterinx) - 90, 91 92 flavizonus (Plectiscus) 68 armata (Eusterinx) 89 95 foersteri (Plectiscidea) 64 78 armatus (Oxytorus) 52 fracticornis (Lampronota) 58 atteotus (Proclitus) 59 60 fraterna (Plectiscidea) 65 82 binodulus (Plectiscus) 77 Fugatrix 63! 86 bispinosa :'. (Eusterinx) . . . . 89, 94 fulvicornis (Cryptus) (Helictes) 98 bistriata (Plectiscidea) 67 63, fulvicornis (Proclitus) 59 blandita (Plectiscidea) 65 82 fulvipectus (Proclitus) 60 Blapticus Forster 86 fulvus (Plectiscus) 85 Blapticus Thomson 86 fusciventris (Cylloceria) 54, 55 borealis (Cylloceria) 54, 55 gilvus (Plectiscus) 86 borealis (Helictes) 98, 100, 102 Gnathochorisis §6 brachyurus (Plectiscus) 70 gracilis (Pantisarthrus) 61 caligata (Cylloceria) 54, 55 56 gravator (Hemiphanes) 51 canaliculata (Plectiscidea) 64, 74 habilis (Plectiscus) 83 captiosus (Proeliator) IO3 104 haeselbarthi (Apoclima) 53 Catomicrus 88 93 Helictes 93 cinctula (Plectiscidea) 64, 71 helvola (Plectiscidea) 64, 72 circaea (Eusterinx) 89 96 Hemiphanes 50 coUaris (Plectiscidea) 64 77 Holomeristus 88 96 collaris (Plectiscus) 77 hortense (Hemiphanes) 51 comes (Proclitus) 59 hostilis (Plectiscus) 85 communis (Plectiscidea) 86 humeralis (Plectiscidea) 66, 85 conjuncta (Plectiscidea) 64, 76 Idioxenus 93 connexa (Plectiscidea) 65, 81 imperspicua (Cylloceria) 55, 56 conspicuus (Helictes) 98, 100 inaequalis (Eusterinx) 89, 90, 95 coxalis (Idioxenus) lOQ inaequalis (Idioxenus) 98 coxator (Plectiscus) 70 inaequalis (Pantisarthrus) 61 crassicornis (Plectiscidea) 65, 80 incongruens (Helictes) 98, 102 crassulus (Gnathochorisis) 87 88 indomita (Plectiscidea) 63, 68 crenicornis (Lampronota) 56 infirmus (Plectiscus) 86 cruentator (Megastylus) 104 inquilinus (idioxenus) lOO cruentatus (Cryptus) (Helictes) 104 intricator (Idioxenus) IQO curticauda (Plectiscus) 71, 72, 76 84 inusitatum (Hemiphanes) 52 Cylloceria 54 invalidus (Idioxenus) lOO Dallatorrea 88 95 invicta (Cylloceria) 54 56 dentifer (Gnathochorisis) 87, 88 invictus (Proeliator) 103, 104 deterior (Plectiscidea) 65,' 82 Ischyracis 88 94 determinatus (Plectiscus) 71 jugorum (Eusterinx) 90, 91 Dialipsis 62 Laepserus Forster 86 dispar (Pantisarthus) : 61, 62 Laepserus Van Rossem 86 disparilis (Eusterinx) 89, 94 langei (Cylloceria) 54, 55 distinctus (Plectiscus) '74 longicornis (Chalinoceras) 58 Divinatrix 88 95 luridator (Oxytorus) 52 edwardsi (Proclitus) 59, 60 luridator (Oxytorus) f. nigricoxa 52 108 Tijdschrift voor Entomologie, deel 130, 1988

luridus (Pantisarthrus) 61, 62 propinquus (Idioxenus) 100 maderensis (Misoleptus) 105 proprius (Proeliator) 103, 104 mancus (Chalino ceras) 56 proximus (Plectiscus) 73 mediator Forster (Idioxenus) 98 pseudochropus (Pantisarthrus) 61 mediator Schiedte (Megastylus) 98 pseudoligomera (Eusterinx) 90, 91, 92 Megastylus 104 pseudominutus 92 Megastylus (Helictes) Thomson 98 pseudominutus (Hemiteles) var. jugorum 91 Megastylus Holmgren 98 pungens (Plectiscus) 80 melancholica (Cylloceria) 55, 58 pusilla (Eusterinx) 89, 90, 93 melancholica f. denticornis (Cylloceria) 58 refractaria (Eusterinx) 89, 90, 97 melancholica f. marginator (Cylloceria) 58 restrictus (Gnathochorisis) 88 melanocera (Plectiscidea) 64, 73 rudepunctatus (Pantisarthrus) 61, 62 mendica (Plectiscidea) 65, 82 rudis (Proclitus) 60 mesoxantha (Plectiscidea) 65, 81 rugosissima (Phosphoriana) 103 minima (Eusterinx) 90, 97 rugosissimum (Entypoma) 103 moerens (Plectiscidea) 63, 68 rugosissimus (Phosphorus) 103 montanum (Hemiphanes) 52 signaticorne (Apoclima) 54 monticola (Plectiscidea) 64, 74 sodalis (Plectiscus) 71 Myriarthrus 98 striolata (Cylloceria) 57 nava (Plectiscidea) 64, 78 subalpinus (Aniseres) 61 nemorensis (Plectiscidea) 63, 66 subangulata (Plectiscidea) 65, 85 nigricoxus (Helictes) 100 subcurvatus (Plectiscus) 74 nigritus (Plectiscus) 86 subdola (Eusterinx) 90, 91, 92 nuptialis (Plectiscus) 70 subsimilis (Ephalmator) 66 obscurella (Eusterinx) 90, 91, 93 subsimilis (Plectiscus) 73 occupator (Lissonota) 54 substantiva (Plectiscidea) 65, 79 ochropus (Pantisarthrus) 61 subsulcatus (Proclitus) 59 oligomera (Eusterinx) 90, 91 subteres (Plectiscidea) 63, 67 orbitator (Megastylus) 105 subtilis (Plectiscus) 74 Oxytorus 52 suerinensis (Cylloceria) 55 paganus (Proclitus) 59 sylvestris (Cylloceria) 55, 57 Pantisarthrus 61 sylvestris (Tryphon) 57 parvula (Plectiscidea) 64, 69 tartarea (Eusterinx) 90, 94 petiolatus (Plectiscus) 73 tetraglyptus (Idioxenus) 100 Phosphoriana 102 tener (Plectiscidea) 63, 69 Phosphorus Thomson 102 tenuicincta (Eusterinx) 89, 90, 96 Phosphorus Van Rossem 102 tenuicornis (Plectiscidea) 64, 70 Phosphorus Voet 102 terebrata (Gnathochorisis) 88 pilicornis (Megastylus) (Helictes) 101 terebrator (Plectiscidea) 65, 83 Plectiscidea 62 townesi (Hemiphanes) 51 Plectiscus auctores 62 trichops (Catomicrus) 93 Plectiscus (Proclitus) 58 vagator (Plectiscidea) 64, 73 posticata (Plectiscidea) 65, 80 variator (Idioxenus) 101 praepositus (Plectiscus) 83 ventosa (Plectiscidea) 65, 84 praetor (Proclitus) 59 Voetia 102 Proclitus 58 xanthocephalus (Gnathochorisis) 88

Proeliator : . . . . 103 xanthoneuris (Plectiscus) 68 properator (Oxytorus) 52 zonatus (Proclitus) 59 DEEL 130 1987

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

^^SN. O^

Tijdschrift voor Entomologie, deel 130, 1987 INHOUD

R. E. Blackith and R. M. Blackith, Tridactylids and tetrigids (Orthoptera) from Sulawesi, Indonesia 1 R. V. Hensen and L. H. M. Blommers, Review of the Malagasy species of Belonogaster Saussure (Hymenoptera, Vespidae) 11

J. WiEDENSKA, Morphologie der Larven und Puppen einiger Phylidorea- Anen (Diptera, Limoniidae) 33 G. VAN Rossem, A revision of western Palaearctic oxytorine genera. Part VI. (Hymenoptera, Ichneumonidae) 49 B. K. RusSEV, Ecology, life history and distribution of Palingenia longicauda (Olivier) (Ephemeroptera) 109 S. fig A. J. Gardiner and G. Compton, New species of the wasp genus Diaziella (Hymen- optera, Chalcidoidea, Sycoecinae) 129 M. C. D. Speight, External morphology of adult Syrphidae (Diptera) 141 M. R. DE Jong, Taxonomy and biogeography of Oriental Prasiini. 3. The fatiloqua and parvula groups of the genus Lembeja Distant, 1892 (Homoptera, Tibicinidae) 177 W. NijVELDT, On four new Palaearctic species of the genus Cecidomyia (Diptera, Cecidomyii- dae) 211

Gepubliceerd 30 november 1987

ISSN 0040-7496 Tijdschrift voor Entomologie 130: 1 — 10 Gepubliceerd 30 november 1987

TRIDACTYLIDS AND TETRIGIDS (ORTHOPTERA) FROM SULAWESI, INDONESIA

R. E. BLACKITH and R. M. BLACKITH

Trinity College, Dublin, Ireland

Abstract

The tridactylids and tetrigids from the Project Wallace and other expeditions to Sulawesi are identified and where available notes on their biology are appended. Two widespread south-east Asian species constitute the surprisingly small tridactyhd fauna so far known. Four tetrigid species new to science are described, six new to Sulawesi are listed, and the

presence of 13 previously recorded there is confirmed. Two tetngid species described from Sulawesi are syaonymised. The tetrigid faunas of North, Central, and South Sulawesi are distinct; they may have arrived on different fragments of continental plate and failed to mix appreciably. The prospects for a radical revision of the Tetrigidae of the region are outlined.

Introduction low that once thought needed for genetic cohe-

Hancock (1915) lists 47 Indian tetrigid speci- sion between the populations. The large number mens which he reworked after Kirby (1914) had of endemic genera is one obvious consequence, determined them. They agreed fully on the but we also meet instances where a species identity of only two specimens; moreover, of found on one island is very similar, but not the nine specimens which Kirby labelled Hedo- identical, to a related species from another is- tettix gracilis De Haan, Hancock deemed there land. Whether it should be described as new be- to be six species in four genera (none of them comes an existentialist problem, where deci- Hedotettix). Again, within the 13 specimens sions are made with inadequate evidence to sup- which Kirby regarded as Euparatettix persona- port them. In view of the high degree of tus (Bolivar), Hancock found eleven species m endemism in south-east Asian tetrigids, the at- five genera. These discrepancies suggest that not tribution of a given specific name to material only are tetrigids a "difficult group" but that from widely separated continents is questiona- one of the difficulties is the wide disparity be- ble. tween one worker's appreciation of infra-specif- Cousin (1961) has drawn attention to the ic variation and another's. existence of "sibling" species (espèces sosies) in A successful revision of the group, now over- gryllids, whereby populations doubtfully sepa- due, may depend on establishing a numerical rable morphologically but long isolated geo- basis for deciding what constitutes variation graphically, e.g. in Africa and South America, within species, genera etc. We hope to use the may be capable of hybridising and giving rise to long series which were collected during Project fertile offspring. White (1978: 32) questions the

Wallace to this end. If no such basis is estab- role of cohesive forces (gene-flow) in prevent- lished, further revisions may simply pit another ing speciation, and notes that we have no means set of concepts against those implicit in previous of measuring them even approximately. He revisions, and add to the synonymy. There are considers it legitimate to regard the origin of ge- so few reliable characters that recourse to a netic isolating mechanisms rather than subdivi- multivariate ordination, using numerous charac- sion of the gene-pool as the prime cause of spec- ters in combination, is a possible solution. iation. Since tetrigids have, superficially at least, A particular difficulty with tetrigids from the uniform caryotypes the prospects for unequivo- islands of the Oriental Region is that many spe- cal resolution of their taxonomie problems by cies are weak flyers or apterous, so that gene cytogenetic analysis seem remote. However, the flow between the islands is likely to be far be- shapes of different species are characteristic, and Tijdschrift voor Entomologie, deel 130, 1987

susceptible to numerical analysis. Tridactyloidea: Tridactylidae A partial revision of south-east Asian tetrigids is under way, based on the multivariate Tridactylus riparius Saussure, 1877 analysis of 80 morphological characters assessed Material studied: lOcî, 10? (Blackith); 59, on each specimen. This analysis will be de- 7.VÌÌ.1985 (Butlin); 3(5, 5$, 15— 18.ix.1985 (Ashe). scribed elsewhere. Provisional identifications are offered here for tetrigids, as Kevan (1966) There is no discernable variation in the abun- recommends, although tridactyhds are defini- dance of this species over the nine months cov- tively identified from the male genitaUa (Black- ered by these collections, the first ever made, so ith & Blackith, 1979). To avoid repetition, all far as we can judge, from Sulawesi. collecting localities are Indonesia, Sulawesi and T. riparius occurs on silt and silt-filled gravel unless otherwise stated, in the Dumoga-Bone banks along the Dumoga, Bone, Toraut, Tum- National Park, Sulawesi Utara. All material col- pah and Pononontuna Rivers. A creek on the

lected by us is dated between 3.1.1985 and left bank of the Toraut upstream from the Maze 28.iii.1985; for long-lived insects in an almost was chosen for a capture - recapture experi- unvarying habitat on the equator, with, locally, ment. This was an area of 12 sq. m. covered no clear rainy season, further precision as to with long sparse grass and inhabited by frogs dates seems pointless. and water-snakes. On 2.ii.85, 25 males and 18 For biogeographical purposes we consider females were marked with a spot of yellow oil Sulawesi to be divided into three regions; north paint on the pronotum and released. Two days of the equator; central (between the equator and later 102 males and 55 females were captured of latitude 4°S); and south of latitude 4°S. These ar- which four and one respectively, were marked. bitrary divisions are based on major gaps be- A simple Lincoln Index calculation suggests tween areas where collecting has been done. that the population contained 637 males and 990 Anatomical nomenclature follows Albrecht females, roughly 128 individuals per sq. metre. (1953) except that the armament of the hind tib- We think this is as dense a population as occurs

iae is called teeth, rather than spinules, which in the region. imphes articulation. Several characters useful in Tridactylus opacus Walker, 1871 tetrigid taxonomy have received little or no attention in the literature; two that are used here Material studied: 2 (Î, 15-18.ix. 1985 (Ashe). are the organisation of the thoracic pleura (fig.

1) particularly the presence or absence of a visi- These few specimens were taken in a Malaise ble suture on the mesepisternum, and the ar- trap near "The Maze" on the Toraut River. rangement of prominent sensilla on the inner Both species of Tridactylus range from India face of the hind femora, proximal to the genicu- to Sulawesi, but neither occurs in Australia (K. lar region (fig. 2). These sensilla are often ar- K. Günther, 1978). ranged in two groups, a central cluster roughly half-way between the dorsal and ventral mar- Tetrigoidea: Tetrigidae gins of the inner face, and a row or cluster much As there is no substantially complete or nearer the dorsal margin. In some species, there agreed classification of this family into sub-fam- is a process or fold in the dorsal carena of the ilies it will be treated provisionally as an entity. hind femur here called the pregenicular fold, proximal to the ante-genicular tooth (fig. 2b). Diotarus pupus Bolivar, 1887 (fig. 2d)

Two characters that our long series show to be Material studied: 4cî, 49, Edwards' Camp; \i, sufficiently unreliable to be more of a hindrance '1440' summit; \S, 19, Gunung Muat, on crests of than a help are the colour pattern and the rela- ridges in forest, among leaf litter (Blackith). tive lengths of the pulviUi on the first segment of the hind tarsi. New to Sulawesi, but occurs in the Phil- ippines. Abbreviations: BMNH = British Museum (Natural History); LEM = Lyman Entomological Museum Hirrius montanus Günther, 1937 (Macdonald College of McGill Univ., Quebec); MB Material studied: \S, Sulawesi Tengah, Mount = Museum Bogorensis, Java, Indonesia; = MNHNP Tambusiasi, 1200 m, 3-13.iv.l980 (Brendell) Muséum National d'Histoire Paris; Naturelle, NMI (BMNH). = National Museum of Ireland, Dublin; RNHL = Rijksmuseum van Natuurlijke Historie, Leiden. Described from south Sulawesi. Blackith & Blackith: Sulawesi tridactylids and tetrigids

Eucriotettix aff. dammermanni 47 adults were marked on each occasion. Num- Günther, 1938 (fig. 2c) bers, rates of immigration, and death, were cal-

Material studied: lOcî, 189, widely distributed culated by Jolly's method as programmed by within the Dumoga-Bone National Park (Blackith); Davies(1971). \â, 28.vi.1985 (Butlin); IS, 18.ii.1985 (Hoiioway); The population was enclosed by areas inhos-

1 2 , Plot A (BMNH Fogging Team) (BMNH). pitable to the species except up-stream, and a few individuals including one marked specimen, This is a strong flyer and was often taken at were found 150 m away from the experimental lights. area, which occupied 12 sq. m. of the stream Kevan (1966) comments that the taxonomy of bed. By the end of the experiment, about half the large genus Eucriotettix Hebard is "in rather the individuals bore paint. The maximum popu- a chaotic state so that accurate determinations lation was estimated as 214, with standard error are virtually impossible". E. dammermanni was of 64. Probabilities of survival were never sig- described from Sevesi Island (between Java and nificantly less than unity, and estimates of re- Sumatra) and, if the identification is correct, is cruitment or loss never significantly greater new to Sulawesi where it seems to be associated than zero. with running water. Our impression is that the population is re- markedly stable, with long-lived adults, much Eucriotettix ridleyi Gttnther, 1938 more than three months, and few nymphs. It is

Material studied: 1 5. 1 ?, Hog's Back Camp, on hard to conceive of environmental factors likely forest floor (Blackith); 1 $, same data, on logs, to trigger substantial change, apart from non- 29.vi.1985 (Butlin). seasonal phenomena such as vulcanicity, wind- blows, or man's activity. Described from Singapore. If the identifica- However, Dr. R. Butlin (personal commu- tion is correct, it is new to Sulawesi. The species nication), who took part in Project "Wallace be- is unusual in having no sensilla immediately tween June and August 1985, examined the proximal to the genicular area inside the hind population several times during that period and legs. saw no marked individuals. Various currently un verifiable explanations suggest themselves; Scelimena celebica (Bolivar, 1887) there may have been a sharp onset of mortahty, or the marks may have worn off.

Material studied : 3 5 , 5 9 (Blackith). Tondanotettix modestus Günther, 1937

This is the only species of Günther's "Sceli- Material studied: 1<5, Edwards' Camp, on ridge in menae verae" found during the expedition. It leaf litter (Blackith); 2S, 29, Gunung Mogogonipa, inhabits rock-strewn river banks and gulhes on logs, 5. vii. 1985 (Butlin). with running water in rain-forest, often at low light intensities where no other riverine tetngid Described from north Sulawesi. The metepis- can survive. As Humbert, quoted by Bolivar ternum has a large (0.25 mm) rounded process

(1887) notes, it swims and takes off from water, with no obvious sensilla, apparently engaging

flashing its striking blue wings. We collected it with the ventral margin of the pronotum. wherever there was permanent running water in Tegotettix deep shade and boulders with a matrix suffi- corniculatus celebensis ciently porous to allow continuous algal growth Günther, 1937

on which it feeds (Blackith, in press). Described Material studied: 19, Sulawesi Tengah, Ramu Riv- previously from North Sulawesi, where it is ap- er Area, nr. Morowali (Brendell) (BMNH). parently endemic. Between 28.Ì.85 and 25.iii.85 an isolated pop- Described by Günther, as a subspecies of Te- ulation in a deep gully about 5 m below the for- gotettix corniculatus (Stal), from south Sulawesi. est floor, through which a stream runs into the Euparatettix left bank of the Toraut in the Maze, was sub- celebicus (Hancock, 1907) jected to five successive capture-recapture ex- Material studied: 29, Sulawesi Tengah, Ramu Riv- periments. A different coloured or positioned er Area, nr. Morowah, 27.i—20. iv. 1980 (Brendell) mark was placed on the pronotum on each occa- (BMNH). sion, using dots of oil-paints. Between 24 and Tijdschrift voor Entomologie, deel 130, 1987

Described from south Sulawesi in the genus range from Malaysia to the Philippines, and

Hedotettix Hancock, this species is said by Günther (1937) records it from north Sulawesi. Günther (1937) to be closely allied to E. personatus, E. tricarinatus, and Paratettix histricus, Loxilobus rugosus celebensis Günther, 1937

all three of which occur on the island, and may have speciated there. Material studied: iS , 39, Lombongo Village, along tributaries of the Bone River (Blackith). Euparatettix personatus (Bolivar, 1887)

Material studied: la, 29, Sulawesi Tengah, Ramu Günther (1937) described this form as a sub- River Area, nr. Morowali, 27.i—20.iv.l980 (Brendel!) species of L. rugosus Bohvar, which is a Bor- (BMNH). nean species. However, during the description

he calls it L. celebensis, probably indicating that

Recorded by Günther (1937) from south Su- it had virtually specific rank in his mind. The lawesi. type locality is south Sulawesi.

Paratettix tricarinatus Bolivar, 1887 Systolederus ophthalmicus Bolivar, 1887

Material studied: òS, 39, Sulawesi Tengah, Ramu S. carli celebensis Günther, 1937: 189. Syn. nov.

Camp, Kolonodale area, at black light, 29.Ì.1980 (P. S. fruhstorferi Günther, 1937: 189. Syn. nov. G. Kevan) (LEM); M, 39, Sulawesi Utara, Gua Ka- (But- pur, on Limestone with grass cover, 7.viii.l985 Material studied: 8d, 109 (Blackith); la, Im). 6.viii.l985 (Butlin); 19, 28.viii.1985 (Kirk-Spriggs) (BMNH); 1(5, X.1985 (Ashe).

The Ramu Camp specimens bear a label with Günther (1937) erected 5. carli celebensis and the same determination by D. K. McE. Kevan. S. fruhstorferi because of differences in the rela- Described from the Philippines, new to Sulawe- tive lengths of the pulvilli on the first segment si. of the hind tarsi and in colour patterns. He was

unable to see the type of S. ophthalmicus and

Paratettix aff. mimus had to rely on Bolivar's limited description Bolivar, 1887 based on a single female. Inspection of this type and of a long series of specimens from the To- Material studied: 5d, Sulawesi Tengah, Ramu raut region of Minahassa shows that colour pat- Camp, Kolonodale Area (P. G. Kevan) (LEM). tern and the relative lengths of the pulvilli fall into the category of characters which are collec- P. mimus was described from the Philippines, tively useful but individually unreliable. but is new to Sulawesi. There is an apparently continuous range of Hedotettix costatus Hancock, 1912 relative lengths (denoting the lengths of the pul- viUi seriatim by pi, p2 and p3) from pi = p2 = Material studied: Icî, 19, 1. vii. 1985 (Butlin); 59, (as Bolivar claims for the type of S. ophthal- Sulawesi Utara, nr. Dolodua, from egret's crop, p3 5.ÌV.1986 (C.Vermeulen). micus), through p3 = pi -I- p2 (as in the description of S. carli celebensis) to i^are specimens with

These specimens might well be H. gracilis De p3 > pi -I- p2 as in the description of S. fruhstor- Haan and H. costatus may itself be a synonym feri. In fact, careful measurement of Bolivar's

of H. gracilis; H. costatus is recorded by type shows that pi = p2 = 0.20, but p3 = 0.25 Günther (1937) from south Sulawesi. The spe- mm on both legs. cies seems to be adapting to life in paddy fields. No character has been found to distinguish forms within this range, although the sensory Loxilobus insidiosus Bolivar, 1887 pads on the tips of the maxillary palps (when

Materia! studied: Id, 29, Lombongo Village examined at X 160) do appear to be smaller and (Blackith); IS, 29, Lombongo Village, 6.viii.l985 more elliptical in the one specimen we have at-

(Budin) ; 2 9 , Manado (Blackith). tributable on Günther's description to S. fruhstorferi; this may be idiosyncratic. Kevan (1966) notes that the genus Loxilobus The locally common species inhabits boulders

Bolivar is almost certainly polyphyletic and its in fairly open rivers, not in deep shade, particu- taxonomy chaotic. The above species has a wide larly the Tumpah River and a waterfall some 4 BlACKITH & BlACKITH: Sulawesi tridactylids and tetrigids

km north-east of Lombongo. Aspects of its bi- eps epm epm. ology have been described by Blackith (in

press), and it appears to be endemic to north Su- lawesi.

Coptotettix alfurus Günther, 1937

Material studied: 3cî, 49, Huntuk trail and '1440' summit (Blackith); 6cJ, 79, Gunung Poniki, leaf-litter and sphagnum, 14— 15.viii.l985 (Budin); Icî, 39, Gunung Mogogonipa, 30. vii. 1985 (Butlin); 19, Gun- ung Poniki, 18.X.1985 (Monk) (BMNH).

An apterous, high altitude, form living in leaf-litter, with several colour patterns. De- scribed from south Sulawesi.

Coptotettix interruptus Bolivar, 1887 Fig. 1. Probolotettix kevani sp. n. Organisation of pleural segments of meso- and metathorax. e = ely- Material studied: 4(5, 39, streams feeding the Bone tron, sp = spiracle, eps, = mesepisternum, epm2 = River (Blackith); \i, 6.vii.l985 (Kirk-Spriggs) mesepimeron, eps'j = meta-anepisternum, = (BMNH); 19, Gunung Muat, Lakes Bungalow, epsj metakatepisternum, epm'j = meta-anepimeron, epmj 14.vii. 1985 (Butlin). = metakatepimeron, pr = (proprioceptive) process on metakatepimeron. Scala-line 0.25 mm. The possibility that this is the alate form of C. alfurus seems to be discounted by the fact that eye margins. Palps only slightly elliptical in the two forms live in quite different habitats, at cross section. different altitudes. Described from Java, pre- Vertical furrows of pronotum c and d deep, viously recorded from Sulawesi by Günther becoming obsolete towards median carena, (1937). hnked at base by deep transverse furrow k. Fur- Hyboella overbecki Günther, 1939 row e obsolete. Prozonum upturned against back of head. Infrascapular area (viewed X 160) Material studied: Id, Gua Kapur, nr Gorontalo, with smooth lower carena. Pronotai disc leaf litter in woods on limestone, 7.viii.l985 (Butlin); brown, maculate black. Elytra elliptical Id, Gunung Poniki, Ice Station Zebra, 14.viii.l985 (1.6 mm (Butlin). X 0.6 mm). Wings fully developed, exceeding pronotum by about 1.4 mm. Cells in cubital Previously recorded from Java and Sumatra area black, cross-veins white. Transverse suture (Günther, 1955) and new to Sulawesi. The Gua on mesepisternum weakly developed.

Kapur specimen is renerai. Ventral carenae of fore-and mid-femora without fringe of long golden hairs. Fore-femora Probolotettix corticolus sp. n. lobed. Hind femora with eight fragae on outer (figs. 2b, 3) face, but no callosities. Five conspicuous white sensilla on inner face of hind femora, proximal

Holotype, 9, Indonesia: Sulawesi Utara, Du- to genicular area. The largest sensillum mea- moga-Bone National Park, 13.iii.l985 (Black- sures 0.6 mm X 0.45 mm. Hind tibia with six ith) (RNHL). teeth on outer, five on inner, margins. Pulvilli on first segment of hind tarsi 0.6, 0.6 and 1.4 Paratypes, 2â , 29, same data (Blackith) (BMNH; LEM; NMI; MB). mm long. Ovipositor valves slender (upper valve 1.2 mm X 0.3 mm). Other material studied: 2(5, 3 9 standing over Head and anterior segments of pronotum as a label "Criotettix sp." in the National Museum in fig. 3. Vertex (4.7 mm), wider than eye (4.0 of Ireland have more robust ovipositor valves mm). Frons visible throughout, in side view, be- (0.9 X 0.5 mm) and are from Java. It seems un- tween eyes. Scape dorso-ventrally compressed, wise to designate these specimens as paratypes pedicel sub-spherical. Lateral carenae of vertex although they appear to belong to P. corticolus. joining median carena of fastigium, terminating This species was found on the bark of trees, caudad in small black horns adpressed to inner whether upright or fallen, in deep forest along Tijdschrift voor Entomologie, deel 130, 1987

face of hind femora as fig. 2a, pregenicular fold on dorsal carena. Pulvilli on first segment of hind tarsi produced into sharp points. The species is distinguished from P. corticolus by the arcuate frons, and the lateral carenae of the vertex which are distinctly higher than the median carena in profile (fig. 4). The species differs from all those assigned to the genus by Günther (1939) in having the vertex visible between the eyes in profile, a feature

which it shares with P. corticolus. It is, however, very similar to P. sundaicus Günther from

which it differs, apart from the projection of the vertex, in size, although P. sundaicus appears to be confined to western Indonesia.

Fig. 2. Arrangement of sensilla on internal face of The species is widespread in Sulawesi Utara hind tibia. (Diagrammatic), slpr = semilunar process, and inhabits river banks together with S. oph- = = agt antegenicular tooth, pgf pre-genicular fold; a, thalmicus. It flies strongly and comes to light Probolotetüx kevani sp. n.; b, Probolotettix corticolus readily. sp. n.; c, Eucriotettix aff. dammermanni Günther; d, P. kevani is also the predominant species re- Diotarus pupus Bolivar. Scala-line 0.5 mm. covered from the crops of egrets and herons feeding in and around paddy fields in areas of the Huntuk trail in the area between the Toraut north Sulawesi from which forest had been and Tumpah rivers. Records of partly cortico- cleared. It appears to be able to thrive in rice- lous tetrigid species are scattered throughout growing areas. These specimens, recovered by the literature, but in this case the species has Dr. Charlotte Vermeulen, were sent to me ac- been found only on bark. companied by specimens collected directly from the paddy fields, many of which proved to be P. Probolotettix kevani sp. n. kevani.

(figs. 1,2a, 4) Although there is some overlap, the number of teeth on the inner and outer margins of the

is (6 Holotype, 6-, Indonesia: Sulawesi Tengah, hind tibiae of P. kevani greater — 11, modal Ramu Camp, Kolonodale area, 5.ii.l980 (P. G. value 9) than in P. corticolus (5—7, modal value Kevan) (LEM). 6). This result is unexpected, because Descamps Paratypes, AS, 2 9, Sulawesi Utara, Dumoga- (1976) found that corticolous species generally Bone National Park (Blackith); (BMNH, NMI, had more hind tibial teeth than other acridids. MB, MNHNP); lc5, 29, same locality, 17.v— Mazarredia celebica Bolivar, 1887 16.vii.1985 (Butlin) (LEM, BMNH, NMI); 19, same data, at Hght, 19. vii. 1985 (Butlin) Material studied: \S, '1440' summit, 4.x. 1985 (K. (RNHL); Icî, Gunung Poniki, at light, Monk) (BMNH); Id, Gunung Ambang, 1200 m. Fog 15.viii.l985 (Butlin) (RNHL); 19, same locali- 7, 18.ii.1985 (BMNH fogging team) (BMNH). ty, 6.VÌÌ.1985 (Kirk-Spriggs) (BMNH). The genus Mazarredia Bolivar was erected for 13 south-east Asian species ranging from Sri Body colour brown, dorsal areas of epimera Lanka to the Philippines. One of these, M. ce- black, raised pronotal disc bordered russet, pig- lebica, was described from a single female from mented across pronotum, anterior part black, north Sulawesi. Bolivar notes that the depres- posterior yellow. Eyes globular, ocelli large sions behind the "shoulders" of the pronotum, (0.18 X 0.12 mm) frons arcuate in profile, pro- characteristic of the genus, are weakly devel- zonal carenae of pronotum divergent cephalad, oped in this small species, which makes it a pas- mesepisternal suture obsolete, elytra 1.1 X 0.5 sage form to several other genera of the region. mm, wings exceeding pronotum by 2 mm. Later, Bolivar (1909) segregated M. celebica Fringe of white setae on ventral carena of into the genus Xistrella Bolivar. However, mid-femora. Hind tibia with eight teeth on in- Günther (1955) expresses the view that Xistrella ner, seven on outer, margins. Sensilla on inner should be returned to a group of genera includ- Blackith & Blackith: Sulawesi tridactylids and tetrigids

Figs. 3 6. Upper part of in — head profile of: 3, Probolotettix corticolus sp. n., holotype 9 ; 4, P. kevani sp. n., paratype 9 ; 5, Mazarredia bolivan sp. n., paratype 9 ; 6, Thoradonta budini sp. n., head and anterior part of pronotum, holotype 9 . Scala-line 0.5 mm.

ing Mazarredia pending further revision, with Mazarredia bolivari sp. n. which we provisionally concur. Günther's ear- (fig- 5) her major revision of 1938— 1939 had been unable to include M. celebica as the type was un- Holotype, 9, Indonesia, Sulawesi Utara, Du- available to him. moga-Bone National Park, at u-v. hght, We have been able to examine Bolivar's type 30.Ì.1985 (Holloway) (BMNH). and compare it with two specimens from the Paratypes, AS, 59, same data (Blackith) Project Wallace expedition. The 80-character (BMNH; LEM; NMI; MB; MNHNP; morphometric analysis shows that the Project RNHL); 19, Sulawesi Tengah, nr. Morowali, Wallace specimens identical are virtually with Ramu River at light, 27.i—20.iv.l980 (Brendell) the type. It seems that this is a high-altitude spe- (BMHN). cies of some rarity, in view of the paucity of specimens discovered. Sulawesian species of Mazarredia can be dis- Leaden grey body colour flecked with dull tinguished from those of Probolotettix by the yellow. Head slightly exserted, eyes raised just absence, in Mazarredia, of sensilla proximal to above the pronotum. Frons ]ust visible before the genicular area of the internal face of the hind upper part of eyes. Vertex slightly narrower femora. However, the boundaries of these gen- than an eye, with minute horns on lateral care- era need clarification. M. celebica has patches of nae. Maxillary palps compressed. Anterior mar- scabrous cuticle on either side of the unpaired gin of pronotum tilted sharply upwards towards ocellus, possibly homologous with the fastigial head. Pronotum weakly depressed behind foveolae of gomphocerine acridids. shoulders. Panier (saddle-bag)-shaped processes Tijdschrift voor Entomologie, deel 130, 1987 on mesozonum. Lateral carena of pronotum in- in which the lateral carenae of the pronotum are terrupted by shallow protuberance (grey against percurrent, and in which the wings are subequal brown pronotal disc) cephalad of which the to the pronotum; the ocelli of M. bolivari are lateral carenae are obsolescent. Transverse care- distinctly larger (0.17 mm wide against 0.11 mm na on episternum of mesothorax. Metasternal for M. celebica). pits and sutures deeply incised. Elytra 1.45 X This species is a strong flyer and comes to 0.50 mm. Ocelli 0.20 X 0.17 mm. Wings ex- light. It IS abundant along the water-courses ceeding pronotum by 1.5 mm. Pronotal length draining the Dumoga and Bone valleys. The 10.5 mm, hind femoral length 4.9 mm. Long species is often found on vegetation some 20 cm (0.2 mm) golden setae spaced at a distance about above the ground, an unusual position for a te- equal to their length along ventral carena of trigid. fore- and mid-femora and proximal half of hind We name this species to celebrate the cente- femora. Hind tibiae with six teeth on inner, five nary (l.xi.1987) of Ignacio Bolivar's seminal on outer margin. "Essay" on the tetrigids.

This species is distinguished from M. celebica

Table 1. Distribution of tetrigids on Sulawesi (+ = material identified by us but not recorded by Günther, 1937; (-I-) = material seen by us, and recorded by Günther, = — = 1937; ( ) material not seen by us, but recorded by Günther, 1937; material not recorded from the area) SPECIES NORTH CENTRAL SOUTH

Diotarus pupHS Bolivar

Kraengia apicalis Bolivar ( )

Ophiotettix cygnicollis Walker ( ) HirriMS sarasinorum Günther Hirrius scrobiculatHS Günther Hirrius montanus Günther Criotettix bispinosus Dalman Eucriotettix dammermanni Günther Eucriotettix ridleyi Günther Scelemina celebica (Bolivar) Tegotettix armatus Hancock

Tegotettix c. celebensis Günther Bullaetettix sarasinorum Günther Tondanotettix meridionalis Günther Tondanotettix modestus Günter Pseudoparatettix luwuensis Günther Euparatettix celebicus Hancock Euparatettix personatus Bolivar Paratettix tricarinatus Bolivar Paratettix mimus Bolivar Paratettix femoralis Bolivar Paratettix histncus Stal Hedotettix costatus Hancock Indatettix sp. Loxilobus insidiosus Bolivar ( + ) Loxilobus r. celebensis Günther + Spadotettix heinrichi Günther ( ) Systolederus ophthalmicus Bolivar (+) Coptotettix alfurus Günther +

Coptotettix interruptus Bolivar -I-

Hyboella overbecki Günther -I-

Probolotettix corticolus sp. n. -I- Probolotettix kevani sp. n. + celebica Mazarredia Bolivar ( + ) Mazarredia bolivari sp. n. + Thoradonta butlini sp. n. BlACKITH & Blackith: Sulawesi tridactylids and tetrigids

Thoradonta budini sp. n. cies are common to centre and south, and only (fig- 6) six species to north and south, out of a total 36 tetrigid species recorded from the island. This strongly suggests that the tetrigid faunas Holotype, 9, Indonesia, Sulawesi Utara, Du- finding moga-Bone National Park, '1440' summit, of the three areas are distinct. Audley-Charles 24.vii.1985 (Butlin) (BMNH). (1981) reviews the evidence suggesting that the three regions of Sulawesi may be derived from distinct fragments of continental plate coming Pronotal disc flavous, sides and legs darker, together as part of the tectonic upheavals of the scabrous white, fragae on hmd femora suffused sea-floor in that region, and each fragment may with pink. Sooty black patches well behind have brought its own fauna with it. Mixing of shoulders of pronotum. Head and anterior part the faunas, in the dense forest cover of the is- of pronotum as fig. 6. Frons almost straight, land, may have been very slow. vertex, top of eyes and prozonum of pronotum However, there are probably biasses in these colinear. Antennae set well below eyes, upper records. Of the 25 species mentioned in margin of scape 0.3 mm lower than lowest mar- Giinther's 1937 paper only 13 have been found gin of eyes. Terminal antennal segments fuscous on Sulawesi again. Moreover, only six of and compressed, pedicel roughly square in pro- Giinther's 28 records have been confirmed both

file (0.13 X 0.13 mm). as to species and region, as table 1 shows. Var- Prozonum of pronotum projecting slightly ious reasons suggest themselves, including the over occiput. Grooves c, d and e all cut lateral possibility that some species are very local and carenae of pronotum anterior to the shoulders that their habitat(s) may have been destroyed by

where the carenae are obsolescent, marked by forest clearing. It is also possible that some spe- change of colour. Infra-scapular area of prono- cies may have been misidentified. Nevertheless, tum wide (0.25 mm). Pronotum 6 mm long. Ap- the fact that only 13 species out of 36 listed terous. Hind femora 4.3 X 1.7 mm, with 17 fra- from Sulawesi (table 1) were recorded both by gae on dorsal external segment and 14 on mid- Günther and subsequently suggests that the col- segment. No visible suture on episternum of lecting effort required to establish the tetrigid mesonotum, no sensilla on internal face of hind fauna with any precision is much greater than femora proximal to the genicular area. Hind tib- has been available so far. ia with seven teeth on inner margin, six on outer A curious bias is the apparent predominance margin. Hind femora just projecting beyond of small forms in the Sulawesian tetrigid faunas.

pronotum. Metaspinal pits exceptionally deep It is hard to know quite what size distribution and wide. to expect, but forms with a pronotum length of

This species is unique among known mem- more than 12 mm seem notably wanting. Only

bers of the genus, which is allied to Mazarredia, two of the usually large "scelimenae verae" with in having the lateral lobes of the pronotum pronotum lengths ranging from 15 to 25 mm are

smoothly rounded, but is otherwise representa- confirmed as present on the island. If it is true tive of the genus, with modifications appropri- that tridactylids have arrived on the island rela-

ate to an apterous species. tively recently It may also be true that part of the tetrigid fauna was also late in arriving geo- Discussion logically speaking and that there has not been

There is little biogeographical information to time for all available niches to be occupied. be gleaned from the tridactylids, both species of It seems that there are (at least) three distinct which are widespread from India to Indonesia. tetrigid faunas on the island which have not

The paucity of species, and the apparent ab- fully mixed. Since this conclusion is likely to

sence of records from expeditions earlier than hold, to some extent, for other orthopteroids, it Project Wallace in Minahassa, suggests that the may help to explain Ramme's (1940) conclusion tridactyloids may have reached Sulawesi only when writing of orthopteran biogeography

recently, as the distribution is grossly unbal- "Aber noch immer ist Celebes in vieler Bezei- anced. hung einer Sphinx" (Even today, the Celebes is The comparison of tetrigid records in the in many ways a Sphinx). three regions (table 1) into which we divide the

island is striking; only two species are recorded Acknowledgements from both the north and centre, only five spe- We are deeply grateful to the following for 10 Tijdschrift voor Entomologie, deel 130, 1987 specimens and information to supplement our quelques Gryllides du Continent Américain. — own collecting: Dr P. Ashe, Trinity College, Bulletin Biologique de la France et de la Belgique Dublin, for tridactylids and tetrigids; Dr R. K. 95:155—174. Davies, R. G., 1971. Computer Programming in Butlin, U. of East Anglia, for tridactylids and Quantitative Biology. — Academic Press, Lon- tetrigids; Dr HoUoway, Commonwealth J. don. Institute of Entomology, for tetrigids; Dr D. K. Descamps, M., 1976. Les Nicarchi, Ommatolampini Entomological Museum, McE. Kevan, Lyman dendrosclerophiles de la forêt néotropicale (Acri- for the loan of tetrigids collected by Dr P. G. domorpha, Ommatolampinae). — Annales de la

Kevan; Dr Vicenta Llorente del Moral, Instituto Société entomologique de France, n. sér. 12 (3): Espahol de Entomologia, for the loan of Boli- 509—526. var's types; Dr K. Monk, Reading University, Günther, K., 1937. Orthoptera Celebica Sarasiniana. Fam. Acrididae, Subfam. Acrydiinae. — Treubia for tetrigids; Dr D. R. Ragge, British Museum 16:165—195. (Natural History) for the loan of specimens in Günther, K., 1939. Revision der Acrydiinae (Orthop- that Museum; and Dr C. Vermeulen, Integrated tera). III. Sectio Amorphopi (Metrodorae Bol. Pest Control Research in the Dumoga-Bone 1887, Auct.). — Abhandlungen und Berichte der National Park for specimens from birds' crops. Museum für Tierkunde und Volkerkunde zu Our special thanks also go to all members of Dresden (A) 20: 1—335.

Phase 1 of Project Wallace who helped us so Günther, K., 1955. Über die Dornschrecken (Orth. unsparingly. Acrid. Tetrigidae) von Sumba und Flores mit fau- nenhistorischen zur Verbreitung This paper is partly based on material col- Anmerkungen einiger Gattungsgruppen der Tetrigidae im sud- lected while the authors were participants in ostasiatischen Inselbereich. — Wissenschaftliche Project Wallace, sponsored by the Royal Ento- Ergebnisse der Sumba-Expedition: Museums für mological Society of London and the Indone- Volkerkunde und des Naturhistorischen Museums (Results sian Institute of Sciences of Project in Basel 1949: 147— 175. Wallace no. 28) Günther, K. K., 1978. Die Tridactyliden Australiens (Tridactylidae, Caelifera, Orthopteroidea, Insec- References ta). — Mitteilungen aus dem Zoologischen Mu- Albrecht, F. O., 1953. The Anatomy of the Migratory seum in Berlin 54: 223—255.

Locust. — London, Albione Press. Hancock, J. L., 1915. Indian Tetrigidae (Acrydiinae). Audley — Charles, M. G., 1981. Geological History — Records of the Indian Museum, Calcutta 11: of the Region of Wallace's Line: 24—35 In: Wal- 13—54. lace's Line and Plate Tectonics. — Ed. T. C. Kevan, D. K. McE., 1966. Some Orthoptera-Caelifera Whitmore, Oxford, Clarendon Press. from the Philippine, Bismark, and Solomon Is- Blackith, R. E. in press. Primitive Orthoptera and lands, with a few interesting records from New Primitive Plants. — Bolletino della Società ento- Guinea and the Moluccas. — Entomologiske mologica italiano. Meddelelser 34: 375—420.

Blackith, R. E. & R. M. Blackith, 1979. Tridactyloids Kirby, F. W. I., 1914. The Fauna of Briush India, 1 of the Western Old World. — Acrida 8: 189— Orthoptera, Acrididae: 1 — 80. 217. Ramme, W., 1940. Beiträge zur Kenntnis der Acridi- Bolivar, L, 1887. Essai sur les Acridiens de la Tribu den-Fauna des Indomalayischen und benachbarter des Tettigidae. — Annales de la Société entomolo- Gebiete (Orth.), mit besonderer Berücksichtigung

gique de Belgique 31 : 175—313. der Tiergeographie von Celebes. — Mittedungen Bolivar, L, 1909. Nouvelles espèces d'Acridiens du aus dem Zoologischen Museum in Berlin 25 (1): Musée de Genève. — Boletin de la Real Sociedad 1—243.

Espanola de Historia Natural 9: 393—403. White, M. J. D., 1978. Modes of speciauon. — San Cousin, G., 1961. Essai d'analyse de la spéciation chez Francisco, W. H. Freeman. Tijdschrift voor Entomologie 130: 11 — 31 Gepubliceerd 30 november 1987

REVIEW OF THE MALAGASY SPECIES OF BEWNOGASTER SAUSSURE (HYMENOPTERA, VESPIDAE)

R. V. HENSEN

/. B. Bakkerlaan 69-111. 3582 VV Utrecht

and

L, H, M, BLOMMERS

Herenstraat 102, 3911 ]H Rhenen

Abstract The Malagasy species of the Afrotropical wasp genus Belonogaster are reviewed and keyed. Twenty-one species are recognized, of which twelve are newly described: B. ambtko, B. betsileo, B. dayi, B. discifera, B. erythrocephala, B. fanemitra, B. mandraka, B. scutifera,

B. tanosy, B. tipuliformis , B. trandraka, and B. vadoni. B. malagassa Saussure is newly synonymized with B. bicolor Saussure, B. keiseri Richards with B. madecassa (Saussure).

The male of B. maromandia is described for the first time. Lectotypes are designated for B. malagassa Saussure, B. ornata Saussure and B. pomicolor Saussure.

Introduction Afrika, Tervuren, Belgium (Dr E. de The Malagasy and the Afrotropical species of Cooninck) Belonogaster were recently revised by the late MHNG Muséum d'Histoire Naturelle, Geneva, O. W. Richards (1982). As Richards (1982: 101) Switzerland (Dr C. Besuchet) already suspected, the material available to him MNHN Muséum National d'Histoire Natu- in fact a was too limited to permit thorough relle, Paris, France, (Mme. J. Casevitz- treatment of the Malagasy species. The present Weulersse) study was based primarily on material collected RMNH Rijksmuseum van Natuurlijke Histo- by the authors (resp. in 1984 and 1971-1973), rie, Leiden, The Netherlands (Dr C. which appeared to contain several new and van Achterberg) imperfectly known species. At a later stage, the SEM Snow Entomological Museum, Law- collection of the Paris Museum has been exam- rence, Kansas, USA (Dr R. "W. Brooks) ined, including the material of the Seyrig col- Specimens collected by the second author will lection, and most of this material is treated here be deposited in the Zoologisch Museum, Am- as well. It seems that Richards misinterpreted sterdam, The Netherlands (ZMA), those of the some species, and overlooked some characters first author are part of his private collection of major diagnostic value. Therefore we feel (CH). obliged to give a new key, and to list the species again, with the most important references. Morphology One of the most important characters in We wish to thank the authorities of the Belonogaster is the length of the stalk of the following museums for enabling us to study the second metasomal tergite. The character, how- material under their care. ever, is not easily appreciated, if not explicitly BMNH British Museum (Natural History), defined, because there is no discontinuity be- London, England (Mr C. O. Vardy & tween the stalk and the remaining part of the Dr M. C. Day) tergite. For the length of the stalk we have taken KMMA Koninklijk Museum voor Midden the distance between the anterior margin of the

11 .

12 Tijdschrift voor Entomologie, deel 130, 1987 stalk and the point were the tergite has the same as long as flagellar width; mesosoma largely width as anteriorly; the width is measured were black, mesopleuron, metapleuron ventrally, the stalk is at its narrowest, usually just before scutellum and metanotum ferruginous; $ the middle. unknown ambiko sp. n. The males of Belonogaster have excellent — Mesonotum dull, finely reticulate between diagnostic characters in the shape of their ter- the punctures; setae of mesoscutum half as minal flagellomeres, and therefore we have long as flagellar width; mesosoma ferrug- selected males for holo- and lectotypes, as far inous; (5 unknown trandraka sp. n. as possible. The females are often less easily 7. Stalk of second metasomal tergite half as identifiable. The colour pattern of most species long as wide (fig. 8); meso- and metasoma is characteristic, but it may prove to be less entirely black; mesoscutum longer than wide; constant than the limited material available to length of forewing 14 mm; $ unknown .... us suggests. betsileo sp. n. — Stalk of second metasomal tergite at least Key to the Malagasy species of 1.4 times as long as wide; meso- and met- Belonogaster asoma usually at least partly green; mesoscutum as long as wide, or larger species

1. Hindwing: M + Cu divides after cu-a (fig. 8 1) 2 8. Mesoscutum 1.1 — 1.2 times as long as wide; — Hindwing: M + Cu divides before cu-a (fig. length of forewing 14.5 —29 mm 9 2) 3 — Mesoscutum as long as wide; length of 2. Body yellowish to brown; first flagellomere forewing 10— 14 mm 18 nearly as long as second + third + fourth 9. Fore coxae and mesopleuron ventrally with (fig. 3); no tubercle present between anten- numerous long black setae among the shorter nal insertions madecassum Saussure white pubescence or tomentum 10 — Body with vivid green, whitish-yellow and — Fore coxae with white pubescence, rarely brown markings; first flagellomere only with a few longer dark setae 15 slightly longer than second + third (fig. 4); 10. Length of forewing 25 —29 mm; body and frons with rounded tubercle between anten- legs entirely ferruginous guerini Saussure nal insertions; dayi sp. n. — Length of forewing 16— 21 mm; body and

3. Petiolus ventrally transversely striate ... 4 legs partly black and/or yellow 11

— Petiolus ventrally smooth 5 1 1 Propodeum with impressed median line over 4. Petiolus comparatively short (fig. 12); ce- its entire length; wings yellowish hyaline; phalic foveae small, as large as surrounding mesosoma ferruginous; stalk of second met- punctation; male: head black, clypeus, scapes asomal tergite 3 times as long as wide; ^ ventrally, fore and middle coxae and femora unknown bicolor Saussure ventrally white .... brevipetiolata Saussure — Propodeum without impressed median line; — Petiolus longer, more slender (fig. 15); ce- coloration of body and wings different 12 phalic foveae larger than surrounding punc- 12. Forewing greyish with the apical third light tation; head of $ ferruginous, with broad yellow; head and mesosoma bright yellow white bands along inner orbits; legs without and green; stalk of second metasomal seg- white markings, partly reddish ment 3.5 times as long as wide; $ unknown erythrocephala sp. n. tipuliforynis sp. n.

5. Mesoscutum strongly, rather densely punc- — Forewing uniformly yellow to brown; head tate, and with long black setosity; metapleu- and mesosoma ferruginous to fuscous or ral flange produced into a short tooth, im- nearly entirely yellow; stalk of second seg- mediately behind the spiracle (fig. 33); legs ment at most three times as long as dark green to black, without yellow pattern wide 13 6 13. Stalk of second metasomal tergite nearly — Mesoscutum at most sparsely, weakly punc- three times as long as wide (fig. 28); me- tate, and without long black setosity; mesoscutum distinctly punctate; legs and mes- tapleural flange without tooth; legs usually osoma dark ferruginous; wings brown partly yellow 7 $ scutifera sp. n. 6. Mesonotum nearly smooth and shiny be- — Stalk of second tergite at most twice as long tween the punctures; setae of mesoscutum as wide (fig. 23); punctures of mesoscutum Hensen & Blommers: Review of Malagasy Belonogaster 13

Figs. 1 — 10. 1, B. madecassa, $, right hindwing. 2, B. eumenoides, Ç, right hindwing. 3, B. madecassa, $, left antenna. 4, B. dayi, $, left antenna. 5 — 6, B. ambiko, $. 5, apical antennal segments; 6, metasomal basis. 7 — 8, B. betüleo, $. 7, apical antennal segments; 8, metasomal basis. 9, B. dayi, $, metasomal basis; 10, B. bicolor, 9, metasomal basis. 1, 2, 6—10: 9 X; 3, 4: 18 X; 5: 36 X. 14 Tijdschrift voor Entomologie, deel 130, 1987

hardly discernible; coloration different; Q 20. Mesosoma finely punctulate-reticulate, with- unknown 14 out coarser punctures, and without any black 14. Mesosoma yellow, pronotum laterally with setosity hildebrandti Saussure green spots; legs yellow; wings yellow — Mesosoma finely punctulate-reticulate, with vadoni sp. n. some distinct larger punctures, which bear — Mesosoma ferruginous, dorsally infuscated; black setae, at least in $ 21 legs black; wings brown mandraka sp. n. 21. Terminal flagellomere of $ dorsiventrally 15. Forewing dark grey, with the tip light red- flattened and strongly curved (fig. 30); malar brown; mesosoma black; $ unknown space of $ short, shorter than width of apicalis Saussure flagellum (fig. 29); gena of $ in lateral view — Forewing uniformly yellow to brown; mes- 0.4 times as wide as eye tanosy sp. n. osoma never entirely black 16 — Terminal flagellomere of (5 slightly flattened 16. Fifth segment of fore tarsi of $ not dilated; laterally, less strongly curved; malar space last antennal segment of $ weakly curved, of $ long, longer than flagellar width (fig. apically broad and hardly pointed; mesosoma 18); gena of $ in lateral view 0.5 — 0.7 times bright green and reddish yellow as wide eye 22 prasma Saussure 22. Terminal flagellomere of (5 rather strongly

— Fifth segment of foretarsi of (5 dilated and curved, widest subapically (fig. 25); colora- flattened (fig. 27); last antennal segment of tion: mesosoma green, with the mesopleuron yellow to yel- (5 strongly curved, apically sharply pointed and usually the metanotum (fig. 26); mesosoma ferruginous to fuscous lowish ferruginous; tibiae nearly entirely or very dark green, with at most metanotum yellow ornata Sausure and part of propodeum yellow 17 — Terminal flagellomere of $ weakly curved,

17. Fifth segment of fore tarsi of (5 longer than widest subbasally (fig. 17); coloration: mes- wide (fig. 41); mesosoma dorsally fuscous osoma ferruginous to fuscous; tibiae ferrug- to dark green, metanotum usually yellowish, inous, in (5 with white longitudinal streaks mid and hind tibiae yellow eumenoides Saussure inaromandta Richards — Fifth segment of fore tarsi of 3' wider than Annotated list of the Malagasy species of long (fig. 27); mesosoma entirely ferrugi- Belonogaster Saussure nous, mid and hind tibiae brown with white Belonogaster sp. n. streak (5 scutifera sp. n. ambiko 18. Forewing of $ greyish, apical third bright (figs. 5, 6) yellow; antenna of $ with dense pubescence Male. on inner side (fig. 38); apical segment of Body length 18.8 mm, length of forewing 13.5 foretarsus of S dilated, black (fig. 39); mes- mm. osoma, including propodeal valves nearly Coloration. — Head ferruginous, mandibles entirely green fanemitra sp. n. and clypeus apically pale yellow, face with pale — Forewing of $ uniformly yellow to brownish; yellow side stripes along inner orbits, and spot antenna of Q with insignificant pubescence between antennal insertions; vertex nearly black. on inner side; apical segment of foretarsus Antennae black, scape ventrally greenish yellow,

of (5 not dilated, not darker than preceding flagellum ventrally, apical three segments en- segments; mesosoma never entirely green, tirely yellowish. Mesosoma black, pronotum ven- propodeal valves always whitish 19 trally, mesopleuron largely, metapleuron ven- 19. Last flagellar segement of $ strongly dilated trally, scutellum and metanotum ferruginous, and flattened, dark brown in contrast to propodeal valves pale yellow. Legs dark green, preceding segments (fig. 13); female: dis- fore coxae anteriorly pale yellow, fore and middle tance between antennal insertions as long tibiae anteriorly with white streaks, middle and as distance between antennal insertion and hind tibiae and all tarsi black. Wings hyaline. inner orbit discifera sp. n. Metasoma black, petiolus anteriorly and ven- — Last flagellar segment of $ not dilated, trally dark green, posterior tergites and sternites yellowish like preceding segments; female; dark ferruginous. distance between antennal insertions longer Pubescence. — Clypeus with rather dense than distance between antennal insertion white tomentum, and rather long black setae; and inner orbit 20 frons, mesosoma and fore coxae with very long Hensen & Blommers: Review of Malagasy Belonogaster 15

black setae (as long as flagellum width); mes- Description. osoma with sparse white tomentum; mid and Body length 18.9 mm, length of forewing 14.3 hind femora ventrally only proximally with a mm. few pale setae; metasoma with short black setae Coloration. — Black, the following parts fer- on tergites, third to seventh sternite only with ruginous: mandibles, clypeus, frons including eye white tomentum. emargination, ventral side of scape, apical three Head. — Clypeus bluntly angled below, angle flagellomeres, fore legs except the tarsi. Face about 130°; gena 0.4 times as wide as eye in along inner orbits yellowish. Wings yellowish lateral view; cephalic foveae absent; clypeus hyaline, veins and stigma yellow. sparsely finely punctate, vertex sparsely rather Pubescence. — Clypeus with short white pu- strongly punctate, shiny; third antennal segment bescence, frons and vertex with long fine grey- 0.8 times as long as fourth + fifth, these both brown setae; mesosoma with sparse white to- 2.3 times as long as wide; eighth segment 1.7 mentum, and long fine grey-brown setae; fore times as long as wide, terminal segment long, coxae with short white pubescence, and some slender, strongly curved (fig. 5). longer grey setae; mid and hind femora with Mesosoma. — Mesoscutum as long as wide; short white pubescence ventrally; metasoma scutellum without median impressed line; me- with short white pubescence. tapleural flange produced into a flat tooth, behind Head. — Clypeus bluntly angled below, angle the mesopleural spiracular lobe; propodeum about 150°; gena 0.4 times as wide as eye in with anterior depression small, but deep, median lateral view; cephalic foveae absent; clypeus impressed line weak, fading near the small shiny, punctulate, with sparse fine punctures; posterior depression; integument shiny, rather vertex shiny, sparsely punctulate, and with dis- densely strongly punctate, punctulation dense, tinct punctation; third antennal segment 0.9 but shallow; propodeum with a few transverse times as long as fourth + fifth, these resp. 1.9 striae above the posterior depression. Hindwing: and 1.8 times as long as wide; eighth segment M + Cu divides before cu — a. 1.7 times as long as wide (fig. 7); third to ninth Metasoma. — Petiolus 1.3 times as long as ventrally keeled; last segment slender, strongly hind tibia, ventrally not transversely striate, curved (fig. 7). posteriorly not swollen, spiracles not prominent Mesosoma. — Mesoscutum 1.1 times as long (fig. 6); stalk of second tergite 0.5 times as long as wide; scutellum without median impressed as wide; seventh sternite truncate. line; metapleural flange without tooth; propodeum with distinct anterior pit, without median Holotype. — $. "Madagascar; (Tamat.); Pé- impressed line; posterior depression well-devel- rinet; 950 m; 48° 16 E, 18° 56 S; lO.v.1984; leg. oped; integument shiny, punctulate and rather R. Hensen & A. Aptroot" (RMHN). sparsely punctate, punctures fine but very distinct. Hindwing: M + Cu divides before cu— a.

Etymology. — "Ambiko" is one of the peculiar Metasoma. — Petiolus 1.2 times as long as Malagasy hedgehogs, of the family Tenrecidae. hind tibia, ventrally not transversely striate,

The name is chosen to illustrate the long pu- posteriorly hardly swollen, spiracles strongly bescence of B. aynbiko, and its more or less prominent (fig. 8); stalk of second tergite 0.5 similar colour. times as long as wide; seventh sternite truncate.

Belonogaster apicalis Saussure Holotype. — $, "Betsileo, Madagascar, 82—30" Belonogaster apicalis Saussure, 1900: 207, 208; Ri- (BMNH). The specimen bears Ri- chards, 1982: 47, 105, figs. 85, 86. chards' identification label, and his description and figure of B. brevipetiolata appear to have It seems very doubtful whether the two males $ been based on this specimen. in the RMNH, examined and mentioned by Richards (1982) indeed belong to this species. Etymology. — Named after the Betsileo, one Belonogaster betsileo sp. n. of the original Malagasy tribes, inhabiting (figs. 7, 8) roughly the highland region between Antsirabe

Belonogaster brevipetiolata, Richards, 1982: 101, and Fianarantsoa. 102, fig. 80 (partim). 16 Tijdschrift voor Entomologie, deel 130, 1987

Belonogaster bicolor Saussure Madagascar, Ambodivoangy, i.1962, both J. (fig. 10) Vadon (KMMA). Belonogaster bicolor Saussure, 1900: 207, 208; Ri- Belonogaster dayi sp. n. chards, 1982: 47, 104. (figs. 9, 36, 37,42) Belonogaster malagassus Saussure, 1900: 210. New synonymy. Male. Body length 17.5 length of forewing 12.5 Types. — The lectotype of B. malagassus, by mm, mm. present designation, is a female, labelled "Ma- Coloration. — Head yellow, vertex green, dagascar, F. Sikora". It evidently belongs to B. antennae ferruginous. Mesosoma green, meso- bicolor, and is the only one of the four specimens and metapleuron, posterior margin of scutellum, standing under B. malagassus in the MHNG agreeing completely with the description. The metanotum, propodeum along median line and two paralectotypes are labelled "Madagasc", apically yellow. Legs yellow, hind coxae and all femora green, tarsi ferruginous to fuscous, last resp. "Nossible". The fourth specimen, labelled segment of fore tarsi proximally white, last "Madagasc, Annanarive" agrees in no way with of mid tarsi light, contrasting the description, being smaller and nearly entirely segment and hind with darker preceding tarsomeres. Metasoma black, and cannot be considered a syntype. It green, second to sixth tergite and sternite with seems to belong to a still undescribed species, broad yellow band apically. proximally related to B. mandraka sp. n. Wings grey-brown, hyaline, veins brown, apical third yellow, veins yellow. Material examined. — 1 $, without locality Pubescence. — Clypeus with dense white (BMNH); 1$ , Ifanadiana, Ranomafana, 900 m, pubescence, vertex and mesosoma dorsally with 29.xii.I971, L. & R. Blommers (ZMA); $, Région d'Ambanja, A. Seyrig (MNHN). sparse brownish setae; fore coxae with dense white pubescence and a few longer dark setae; Belonogaster brevipetiolata Saussure mid and hind femora with dense white pubes- (figs. 11, 12) cence beneath; metasoma posteriorly without

Belonogaster brevipetiolata Saussure, 1891: 98, pi. dark setosity.

4, fig. 1 (partim); Richards, 1982: 101, figs. 79, Head. — Clypeus bluntly angled below, angle 80 (partim). about 140° gena about 0.4 times as wide as ; eye Most specimens examined by Richards belong in lateral view; cephalic foveae absent; clypeus to the related species B. erythrocephala sp. n., sparsely punctate; vertex densely punctulate, which was regarded as a variety by Saussure with a few indistinct punctures; third antennal (1891: 98). The two possible syntypes, seen by segment 0.85 times as long as fourth + fifth, Richards in the Paris Museum, belong indeed these resp. 2.9 and 2.6 times as long as wide; to the present B. brevipetiolata. The $ identified eighth segment twice as long as wide; third to and described by Richards (1982) as B. brevipe- eleventh ventrally keeled; last segment compar- tiolata belongs to a third species, B. betsileo sp. atively short and broad, weakly curved (fig. 36). n. Mesosoma. — Mesoscutum 1.1 times as long Diagnosis. — Like B. erythrocephala, except as wide; scutellum only posteriorly with median for the following. Coloration: head and meso- impressed line; metapleural flange without soma without red markings; $ with clypeus and tooth; propodeum with anterior depression part of frons, scapes ventrally, fore and middle small (propodeum damaged, other details not coxae and femora ventrally white. Cephalic fo- observable); integument punctulate-reticulate, veae small, as large as surrounding punctures mesopleuron with some weak punctures, prop- on vertex. Terminal flagellomere of $ more odeum with fine transverse striae. Foreleg: last slender (fig. 11), finely pubescent on inner side. tarsomere strongly dilated and flatened (fig. 37). Petiolus distinctly shorter and stouter (fig. 12), Hindwing; M + Cu divides after cu— a. as long as hind tibia. Metasoma. — Petiolus 1.15 times as long as hind tibia, ventrally not striate, posteriorly distinctly swollen, spiracles prominent; stalk of Material examined. — 1 $, without locality second tergite 2.5 times as long as wide; seventh (BMNH); 1 9, Périnet, 20.xii.l955, E. McC. sternite trunctate. Callan (BMNH); 1 $, Manjakandriana, Man- draka, 1200 m, 12.xii.1971, L & R. Blommers Female.

(ZMA); 1 $, Fampanambo, 1962, 1 $, N. E. Body length 19.5- .3 mm, length of fore- Hensen & Blommers: Review of Malagasy Belonogaster 17

Figs. 11 19. 11 B. — — 12, brevipetiolata. 11, $, apical antenna! segments; 12, 9, metasomal basis. 13—14, B. disafera. 13, apical antenna! segments; S, 14, $, metasomal basis. 15 — 16, B. erythrocephala. 15, $, metasomal' basis; 16, apical antenna! segments. $, 17—19, B. eumenotdes. 17, S, apical antenna! segments; 18 $ liead- 19, 9, metasomal basis. 12, 14, 15, 19: 9 X; 18: 18 X; 11, 13, 16, 17: 36 X. 18 Tijdschrift voor Entomologie, deel 130, 1987

wing 15.2 — 15.7 mm (holotype: largest speci- "John Noyes and I stayed at the Station Hotel men). at Andasibe for a week at the start of the winter Coloration. — Head green, mandibles, clypeus rains. We collected near the Lemur reserve, a apically, frons along inner orbits below antenna! kilometre south, and also on a small hill to the

insertions, large part of genae yellow. Antennae immediate south of the Hotel, where I sited three fuscous, scape ventrally green, flagellum ven- malaise traps. After erecting the second of these trally and apically bright ferruginous. Mesosoma in an area specially cleared adjacent to a path,

green, meso- and metapleuron, scutellum later- I noticed a green Belonogaster fly languidly past. ally, metanotum, propodeum along median line It flew rather like a tipulid, but inaccessible

and apically yellow; tegulae and post-spiracular amongst the vegetation, so I could not get at

plate ferruginous. Legs green, coxae, trochanters, it with a net. On each and every subsequent visit

tibiae and last segment of all tarsi yellowish; to the trap, sometimes several times a day, one remaining part of tarsi black, pulvilli white. or more of these green Belonogaster would Metasoma green, stalk of second tergite fuscous, materialise and fly away from me, in various

second to fifth tergites and sternites, with whit- directions. I became aware that they appeared

ish-yellow apical band, which is medially more only when I was on one particular side of the or less interrupted by a brownish hyaline part; trap and that their flight paths radiated from sixth segment whitish-yellow. Wings proximally that point! After considerable search, seven days grey-brown, hyaline, veins brown, apical third later I finally found the nest (photo, fig. 42), yellow, veins yellow, stigma yellow. which depended from a twig into open space Pubescence. — Clypeus ventrally with brown at head height. It consisted of a single filament setae, vertex and mesosoma dorsally with sparse, with twelve radially arranged cells, one above short curved setae, fore coxae with rather dense the other, much like some Oriental Stenogaster white pubescence, and many longer black setae; nests. A maximum of 13 wasps was seen clus- mid and hind femora with some white setae tered in a regular fashion about the filament; beneath; sixth tergite and sternite posteriorly the photograph shows eight. So far I could tell, with some black setae. all were female. Their position and behaviour Head. — Clypeus acute below, angle about were such that the nest was virtually indétectable, 100°; gena half as wide as eye in lateral view; giving the impression of leguminous seed heads cephalic foveae absent; clypeus sparsely punctate, or some such plant material. Even when coor- dorsal two thirds densely punctulate; vertex dinate sight lines were given to observers so punctulate- reticulate, with a few indistinct punc- that it was clear they must have been looking tures; third antennal segment as long as fourth at the nest, they nevertheless did not see it for + fifth, these resp. 1.3 and 1.2 times as long between several seconds and close to a minute. as wide; eighth segment 1.1 times as long as Coupled with the non-agressive "decoy" behav- wide. iour, this species exhibits very sophisticate ad- Mesosoma. — Scutellum without median im- aptations for the maintenance of open free- pressed line; propodeum with anterior depres- hanging colonies." sion small, posterior impressed line extending slightly more than half length of propodeum; Belonogaster discifera sp. n. last tarsomere of foreleg not dilated; otherwise (figs. 13, 14) as in the male. Male. Metasoma. — As in the male. Body length 16.1 mm, length of forewing 12.2

Holotype. — $, "Madagascar: Tamat.; Périnet; Coloration. — Ferruginous; antenna dorsally dark, 27.ÌV— 3.V.1983; J. S. Noyes, M. C. Day; BM ventrally pale green, second to third fla- 1983—201" (BMNH). gellomere dorsally infuscated, last flagellomere Paratypes. — 1 $, same data as holotype (CH); dark brown; mandibles white; lateral thirds of 1$, Perinet, 20.xii.l955, E. McC. Calllan clypeus and frons, and area between antennal

(BMNH); 1 $, Rogez, Forêt côte Est, i.1937, insertions white; legs dark green, fore and middle 3$, Périnet, Forêt côte Est, ii.l939, 1 $, Perinet, coxae anteriorly white, rest of fore coxae fer-

ii.1931, all A. Seyrig (MNHN, 1 $ CH). ruginous, tibiae and mid femur anteriorly with pale green streak, apical tarsomeres ferruginous; Note. — The following account on the nest propodeum greenish, propodeal valves white; (fig. 42) and behaviour was sent by Dr Day. petiolus and anterior half of second metasomal Hensen & Blommers: Review of Malagasy Belonogaster 19

segment dark green. Wings yellowish hyaline, sally punctulate, with sparse fine punctures; veins and stigma brownish yellow. vertex punctulate-reticulate, with a few indistinct Pubescence. — Clypeus with dense white punctures; third antennal segment 1.2 times as pubescence; vertex and mesosoma dorsally with long as fourth + fifth, these resp. 1.5 and 1.3 many rather long black setae; fore coxae with times as long as wide, eighth segment 0.9 times dense white pubescence and a few longer black as long as wide. setae; mid and hind femora ventrally without Meso- and metasoma like in the $. erect setosity; metasoma posteriorly without Holotype. — $, "Rep. Malgache, Manjakan- dark setae. driana, Mandraka, 1300 m, ll.iii.l973, L. & R. Head. — Clypeus acute below, angle about Blommers", "nest nr. 73.15" (ZMA).

120°; gena 0.4 times as wide as eye in lateral Paratypes. — 1 $, 5 (5, same data as holotype view; cephalic foveae absent; clypeus punctulate; (ZMA; 1 S RMNH); 1 $, Sambirano, Manon- vertex punctulate-reticulate, with a few indistinct garivo, 1150 m, xii.1960, P Griveaud (MNHN). punctures; third antennal segment 0.8 times as long as fourth + fifth, these resp. 2.6 and 2.4 Note. — Most specimens were collected from times as long as wide; eighth segment 2.2 times an old nest, with practically no cell walls left, as long as wide; seventh to eleventh with raised hanging in a tuft of grass on top of a road bank line on inner side, twelfth strongly dilated and facing south. flattened (fig. 13), about 1.4 times as long as wide. Mesosoma. — Mesoscutum as long as wide; Belonogaster erythrocephala sp. n. scutellum without median impressed line; me- (figs. 15, 16) tapleural flange without tooth; propodeum with Belonogaster brevipetiolata; Saussure, 1891'. 98, pi. anterior depression distinct, without impressed 4, fig. 1 (partim); Richards,, 1982; 101, 102, figs. median line, posterior depression small, shallow; 79, 80 (partim). integument punctulate-reticulate, and sparsely This species was regarded as a variety of B. shallowly punctate, punctures most distinct on brevipetiolata by the Saussure, and Richards did mesopleuron. Hindwing: M + Cu divides before not make any distinction at all. Both sexes can cu— a. easily be separated on basis of the characters Metasoma. — Petiolus 1.15 times as long as given in the key. hind tibia, ventrally not striate, posteriorly not swollen, spiracles prominent (cf. fig. 14); stalk Male. of second tergite 1.6 times as long as wide; Body length 21.0 mm, length of forewing 14.2 seventh sternite truncate. mm. Coloration. — Black; head ferruginous, face Female. with broad pale-yellow side stripes along inner Body length 16.4 mm, length of forewing 12.3 orbits and spot between antennal insertions; mm. ocelli surrounded by black rings; pronotum, sides Coloration. — Ferruginous; vertex, flagellum of scutellum and metanotum, and fore femur dorsally, propodeum and metasoma infuscated; and tibia ferruginous. Wings hyaline, veins and scape, legs except the fore coxae, trochanters stigma yellow. and tarsi, petiolus and stalk of second metasomal Pubescence. — Clypeus with distinct white tergite black with greenish shine; propodeal tomentum and longer black setae; frons and valves white. Wings yellowish hyaline, veins and vertex with rather long black setae; mesosoma stigma brownish yellow. with sparse white tomentum, mesoscutum and Pubescence. — Clypeus with yellow setae on mid and hind femora ventrally with short black ventral half; vertex and mesosoma dorsally with setae; fore coxae with long black setae; all tergites many rather long black setae; fore coxae with and sternites with sparse short brown setosity. long black setae; mid and hind femur with a Head. — Clypeus acute below, angle less than few outstanding white setae ventrally; metasoma 90°; gena 0.85 times as wide as an eye in lateral apically without dark setosity. view; cephalic foveae present, larger than sur- Head. — Clypeus acute below, angle about rounding punctures; clypeus sparsely rather 100°; gena 0.5 times as wide as eye in lateral strongly punctate, dorsal two thirds very finely view; cephalic foveae absent; clypeus on ventral punctulate; vertex punctulate-reticulate, more two fifth granulate, with coarse punctures, dor- densely punctate; third antennal segment 1.3 20 Tijdschrift voor Entomologie, deel 130, 1987

times as long as fourth + fifth, these resp. 1.1 Besuchet, Geneva, and only this allowed us to and 0.9 times as long as wide; eighth segment solve these taxonomie problems. 0.85 times as long as wide; terminal segments (fig. 16) with sparse dark setosity on inner side. Male.

Mesosoma. — Mesoscutum 1.2 times as long Body length 17.6 mm, length of forewing 1 1.2 as wide; scutellum with median impressed line; mm. metapleural flange without tooth; propodeum Coloration. — Ferruginous; the following with anterior depression small, median im- parts pale yellow. Clypeus and frons, except for pressed line extending hardly farther than the a median stripe below antennal insertions, small posterior depression; integument dull, very mandibles, scapes ventrally, narrow transverse finely punctulate-reticulate, and with sparse, line on pronotum, humeral plates, propodeal rather strong punctation. Hindwing; M + Cu valves, fore and mid coxae anteriorly. Pale yellow divides after cu— a. stripes along hind coxae, fore and mid femora, Metasoma. — Petiolus 1.1 times as long as all tibiae; last tarsal segments yellow. Black lines hind tibia, ventrally transversely striate, poste- present along edges of mesoscutum, scutellum, riorly hardly swollen, spiracles rather prominent metanotum, metapleuron, and along median line (cf. fig. 15); stalk of second tergite 0.7 times and parapsidal grooves of mesoscutum. Meta- as long as wide. soma except the petiolus fuscous, second tergite with yellow apical band in one specimen. Wings Female yellowish hyaline, veins yellow. Like the male, except for the following. Body Pubescence. — Clypeus with short white pu- length 21.5 mm, length of forewing 17.6 mm. bescence; vertex with long black setae; meso- Head without yellow pattern; mesosoma black; soma dorsally with sparse rather long black legs black, coxae, particularly the fore coxae, and setosity; fore coxae with white pubescence, and base of fore tibiae dark ferruginous. Clypeus with a few black setae; mid and hind femora without brown setosity. Petiolus 1.1 times as long as hind longer setae; metasoma with short black setosity tibia. on posterior tergites. Head. — Clypeus bluntly angled below, the Holotype. - S, "Rep. Malgache, Nosy Be, angle about 120°; clypeus as long as wide; gena Dzamandzar, 'lO.i. 1972, L. & R. Blommers" 0.4 times as wide as eye in lateral view; cephalic (ZMA). foveae absent; interocular width on vertex Paratypes. — 1 9, same data as holotype slightly longer than third antennal segment; (ZMA); 1 $, Nosy Be, 0—100 m, 5.vi.l984, R. clypeus shiny, finely punctulate; vertex more Hensen & A. Aptroot (CH); 1 $, Fampanambo, closely and strongly punctulate, and with sparse 19.xi.1957, F. Keiser (BMNH). shallow larger punctures; third antennal segment 0.77 times as long as fourth + fifth, these Belonogaster eumenoides Saussure resp. 2.8 and 2.5 times as long as wide; eighth (figs. 2, 17—19, 43) segment 2.0 times as long as wide; third to Belonogaster eumenoides Saussure, 1891: 94; Ri- eleventh with raised longitudinal line ventrally; chards, 1982: 47, 110, figs. 91, 92 (partim). last segment slightly flattened bilaterally, me-

This species is according to Richards (1982: dially and subapically equally wide, weakly curved 110) a very variable one. We think that in fact (fig. 17). a mixture of species was recognized under the Mesosoma. — Mesoscutum as long as wide; name B. eumenoides. For one of these species pronotal keel absent; scutellum without im- two names of Saussure (1900) appear to be pressed median line; metapleural flange without available: B. ornata and B. pomicolor. This tooth; propodeum with anterior depression species is redescribed here under the name B. small; posterior depression shallow, small, ornata. Furthermore, B. tanosy, B. fanemitra, and continued in short impressed median line dor- B. disci/era fall in this category. B. malagassa, sally; integument dull, closely punctulate-retic- placed by Richards (1982: 110) in the synonymy ulate, and with sparse shallow punctation. of B. eumenoides, is a synonym of B. bicolor Hindwing: M + Cu divides before cu-a. (q. v.). Metasoma. — Petiolus 1.1 — 1.2 times as long The type material of B. malagassa, B. ornata as hind tibia, ventrally not transversely striate, and B. pomicolor was kindly sent to us by Dr posteriorly hardly swollen, spiracles weakly 21 Hensen & Blommers; Review of Malagasy Belonogaster

madecassa. apical antennal segments; Figs 20—79 ^0 B fanemitru, 9, metasomal basis. 21—22, B. 21, 3, 24-25, B. ornata, $. 24, metasomal basis, 77 $ metasomal basis. 2^, B. mandraka, S, metasomal basis. antennal segments; 27, left fore tarsus, 25' abical antennal segments. 26—28, B. scutifera. 26, 5. apical $, 22-24, 28: X; 27, 29: 18 X; 21, 25, 26: dorsal view; 28, $, metasomal basis. 29, B. tanosy, Ç, head. 20, 9 36 X. 22 Tijdschrift voor Entomologie, deel 130, 1987

prominent (cf. fig. 19); stalk of second tergite type-series, including lectotype; MNHG); 2 9, 1.5 times as long as wide; seventh sternite Mandjakandriana, Angavokely, 1600 m, truncate. 17. vi. 1972, 1 9, same locality, 17.1.1973, L. & R. Blommers (ZMA); Female. 3 9, 17 km W. Amba- tolampy, 1650 m, 30.1.1985, Wenzel (SEM); Body length 16.8 — 17.5 mm, length of fore- J. 1 9, Tananarive, Parc de Tzimbazaza, 29.x. 1984, v^'ing 12.7 — 13.0 mm. R. Brooks (SEM). Coloration. — Ferruginous, the following parts more or less darkened. Antennae dorsally, Note. — The brown-coloured nests (nrs. 73.19 mesoscutum, propodeum, mid and hind legs, and and 73.20; fig. were found both attached metasoma. Propodeal valves pale yellow. Wings 43) to a loose tree root, on a steep bank along the brownish hyaline, veins yellow. The palest spec- highway near the hamlet Ambatoloana. Spec- imen has only the metasoma posteriorly dar- imens marked "Nest nr. ..." have been collected kened. The darkest specimens (two from original as adults with the nest, those marked "uit nest type-series) are entirely black, with greenish nr. ..." (= from nest nr. ...) emerged later. shine, and the petiolus green. Two females from Angavokely were caught very close Pubescence. — Clypeus ventrally with yellow to a similar, more greyish nest (nr. 72.30), fixed setae; vertex and mesosoma dorsally with rather to bare rock at 2.5 meters above the ground. short black setosity; fore coxae with rather dense white pubescence, and many longer black setae; Belonogaster fanemitra sp. n. mid and hind femora ventrally with short sparse (fig. 20, 38, 39) white pubescence; metasoma with short brown pubescence on posterior tergites. Male. Head. — Clypeus acute below, angle about Body length 16.0 mm, length of forewing 12.2 90°; clypeus 1.2 times as long as wide; malar mm. space as long as maximal flagellar width; gena Coloration. — Head and mesosoma green, sides 0.5-0.7 times as wide as eye in lateral view; mandibles and of face whitish, flagellum cephalic foveae absent; apical third of clypeus brown, apical flagellomeres ferruginous. Legs sparsely fovealate, proximal two thirds punctu- green, fore and mid tibiae suffused with white, tarsi to late and with sparse larger punctures; vertex dull, fore with second fourth segment whitish, punctulate-reticulate, with sparse shallow punc- fifth fuscous, mid tarsi more or less similar, hind tures; interocular distance on vertex as long third tarsi entirely fuscous. Metasoma fuscous, proximal half of petiolus green, stalk of second tergite + fourth + fifth antennal segment, or a little white. apically paler. more or less; third antennal segment 1.1 times greenish Wings brownish, Morphology. of B. as long as fourth + fifth, these resp. 1.3 and — Like male eumenoides, 1.1 times as long as wide; eighth segment 0.8 except for the following. Flagellum strongly times as long as wide. pubescent ventrally (fig. 38); metasoma without Mesosoma. — Pronotal keel distinct, often black setosity. Vertex without discernable punc- rather strong; otherwise like the male. tation; third antennal segment 0.9 times as long Metasoma. — Petiolus 1.1 — 1.2 times as long as fourth + fifth; these resp. 2.0 and 1.9 times as hind tibia, ventrally not striate, posteriorly as long as wide; eighth 1.5 times as long as wide; weakly or not swollen, spiracles weakly or hardly third to eighth ventrally keeled; last segment cylindrical, rather strongly curved (fig. 38). Prop- prominent (fig. 19); stalk of second tergite without impressed median line; mesos- 1.4 — 1.6 times as long as wide. odeum cutum without discernable punctation. Last seg-

Material examined. — 5 9 1 Ó'» Manjakan- ment of fore tarsi dilated, hardly longer than driana, Mandraka, 1350 m, ll.iii.l973, L. & R. wide (fig. 39). Petiolus 1.3 times as long as hind

Blommers, "Nest Nr. 73.19", 1(5 , same data, tibia, posteriorly swollen, spiracles prominent; "uit nest Nr. 73.19", 3 $, same data, "Nest Nr. stalk of second tergite about twice as long as 73.20", 2$, same data, "uit nest Nr. 73.19/20 wide. (ZMA); 1 9, Périnet, lO.v.1984, R. Hensen &

A. Aptroot (CH); 1$ , Betsileo, 82-30 (BMNH); Female. 2 9, Tananarive-E., 1350 m, I4.iv.l984, R. Body length 16.7 mm, length of forewing 14.0 Hensen & A. Aptroot (CH); 1 9, Périnet, 900 mm. m, I6.ii.1972, L. & R. Blommers (ZMA); 7 9 Coloration. — Head and mesosoma dark 1 S, Andragoloaka, 4 9, Annanarivo (original green, mandibles and apex of clypeus ferrug- Hensen & Blommers: Review of MaL Belonogaster 23

inous, flagellum ferruginous, proximally dark, which we have examined. It seems doubtful if apically lighter; mesoscutum infuscated. Legs these two, a 9 and a $, are conspecific indeed. green, tarsi largely fuscous. Metasoma fuscous, basal half of petiolus green, stalk whitish green. Belonogaster madecassa (Saussure) Wings yellowish grey, apex of forewing bright (figs. 1,3,21,22,45) yellow. Raphigaster madecassus Saussure, 1853: 16, pi. 2, Morphology. — Like female of B. eunienoides, f.g. 7. Belo7iogaster madecassus; Smith, 1857: 94. except for the following. Gena 0.6 times as wide Belonogaster longestylus Saussure, 1891: 97. as eye in lateral view; vertex without discernable Belonogaster madecassa\B^\cV\&vàs, 1982:46, 103, figs. punctation; interocular distance on vertex as long 82,83. as third + fourth antennal segment; third an- Belonogaster keiseri Richards, 1982: 46, 104, fig. 84. tennal segment 1.1 times as long as fourth + New synonymy. fifth, these resp. 1.3 and L2 times as long as Synonymy. — Richards saw only two males wide; eighth segment 0.9 times as long as wide. of this species, and described one of these as Pronotal keel absent; punctures of mesoscutum the new species B. keiseri. Examination of the not discernable; propodeum with posterior de- type, and several more males, including a nest- pression shallow, semicircular, not continued in series, convinced us that there are no specific impressed line dorsally.. Petiolus 1 .2 times as long differences between B. madecassa and B. keiseri. as hind tibia, posteriorly a little swollen, spiracles

weakly prominent (fig. 20); stalk of second Distribution. — This species appears to in- tergite twice as long as wide. habit the entire island of Madagascar, and occurs as well on the Comores (here recorded for the Holotype. — $, "Madagascar, Sambirano, Ma- first time). nongarivo, 1150 m, P. Griveaud, xii.1960" Material examined. — 2 9, Tamatave, sealevel,

(MNHN). 19.V.1984, 1 9, Ambanja, 50 m, 3.vi.l984, both Paratype. — $, Périnet, 21.iii.l931, A. Seyrig R. Hensen & A. Aptroot (CH); 1 9> Tamatave, is in bad condition. (MNHN). The specimen Ivoloina, ll.ii.1972, 10 9 3 (5, Tsaramandroso, Ampijoroa, 1. v. 1972, nest 72.23, all L. & R. Etymology. — "Fanemitra" is the Malagasy Blommers (ZMA); 1 9, Bereboka, 60 km NE word for "aculeate wasp '. Morondava, 18— 23.V.1983, J. S. Noyes, M. C Day (BMNH); 1 9, Grande Comore, Mitsoudje, Belonogaster guerini (Saussure) xii.1970, J. Brunhes (ZMA); 1 9, Mandjakan- (fig. 44) driana, Mandraka, 900 m, 10.ii.1973, 2 9, Nosy Raphigaster guerini Saussure, 1853: 17, pi. 2, fig. 8 Be, Dzamandzar, — 10. i. 1972, 1 9. Fénérive- 3. Est, Foulpointe, 11 — 12. ii. 1972, 1 9> Ifanadiana, Belonogaster guerini; Smith, 1857: 94; Richards, Ranomafana, 29.xii— 2.1.1971/72, all L. & R. 1982:46, 102, fig. 81. Blommers (ZMA); 1 9, Andriba, RN 4, kp 220, A photograph of the nest is given in fig. 44. 600 m, 25. i. 1973, 2 9> Tamatave, Fanadrana, Material examined. — 2 $, Ambohimanga, 5.ÌÌ.1972, all L. Blommers (ZMA); 8 9, Région 1600 m, 17. iv. 1984, 1 $, Tamatave, sealevel, d'Ambanja, 1 9, Fort Dauphin, via. 1940, 1 9, 19.V.1984, 1 9, Périnet, 950 m, lO.v.1984, all Ihosy, 1 9,Tampika,vi.l929, l9,Bekily,iii.l930, R. Hensen & A. Aptroot (CH); 1 $, Fort 1 S, Fort Dauphin, v. 1937, all A. Seyrig Dauphin, 24.iii.1966, J. Gutierrez (ZMA); 2 $, (MNHN); 2 9, Prov. d'Ananalava, Maromandia, Brickaville, Ambila-Lemaitso, 10.x. 1971, 2 $, R. Decary, 1923 (MNHN). Fénérive-Est, 15.x. 1971, 1 $, Ifanadiana, Ranom- Note. — Nest nr. 72.23 (fig. 45) was attached afana, 29.xii — 2.i. 197 1/72, 1 $, Tamatave, to the underside of a mango (Mangifera indica) Fanandrana, 5.ii.l972, all L. & R. Blommers leaf at about 1.75 m above the ground. (ZMA).

Belonogaster mandraka sp. n. Belonogaster hildebrandti Saussure (fig. 23) Belonogaster hildebrandti Saussure, 1891: 95, pi. 17, Female. fig. Richards, 1982: fig. 11; 47, 108, 90. Body length 24.3 mm, length of forewing 19.5 Known only from the type series, and two mm. specimens in the BMNH, identified by Richards, Coloration. — Ferruginous; the following 24 Tijdschrift voor Entomologie, deel 130, 1987 parts darkened: antennae except the apical seg- Belonogaster maromandia Richards ments, pronotum, mesonotum and metanotum. (figs. 40—41) Legs fuscous, fore coxae, trochanters and femora Belonogaster maro'rnandia Richards, 1982: 47, 108. greenish; metasoma fuscous, petiolus before the The species was described by Richards on basis spiracles green, stalk of second metasomal tergite of two females. The male is described here for anteriorly pale yellow, posteriorly green. Wings the first time, and some remarks on the types light brown, veins and stigma brown. and other females are given below. Pubescence. — Clypeus and vertex with rather long black setae; metasoma dorsally with many Male. rather short curved black setae; fore coxae with Body length 20.0—24.0 mm, length of fore- long black setosity; mid and hind femora ven- wing 14.5 — 15.5 mm. trally with rather long black setosity; metasoma Coloration. — Head ferruginous, mandibles, with black setae on all tergites and sternites. sides of face and antenna ventrally pale yellow. Head. — Clypeus ventrally acute, angle about Mesosoma ferruginous, dorsally infuscated, me- 95°; gena in lateral view 0.6 times as wide as sopleuron ventrally pale yellow. Legs ferrugi- eye; cephalic foveae absent; clypeus with ventral nous, all coxae and fore and mid femora ventrally third granulate, with sparse coarse punctures, white, mid and hind tibiae and tarsi yellow. dorsally punctulate, with sparse finer punctures; Metasoma ferruginous, partly infuscated. Wings vertex punctulate-reticulate, punctures not dis- reddish yellow. cernable. Third antennal segment 1.1 times as Pubescence. — Clypeus with dense white long as fourth + fifth, these resp. 1.6 and 1.4 tomentum; vertex and mesosomal dorsum with times as long as wide; eighth segment 1.2 times short black setae; fore coxae with dense white as long as wide. pubescence and a few brown setae; mid and hind 1.1 as Mesosoma. — Mesoscutum times long femora ventrally with dense white setosity. as wide; metapleural flange without tooth; prop- Head. — Clypeus bluntly angled below, angle odeum with anterior depression indistinct, with- 120° about ; gena 0.4 times as wide as eye in out line, posterior median impressed depression lateral view; cephalic foveae absent; clypeus shallow; punctulate-reticulate, integument punc- sparsely finely punctate, shiny, vertex densely indistinct, tures on mesoscutum on mesopleuron punctulate, dull, with a few coarse punctures; well discernable; propodeum with some trans- third antennal segment 0.78 times as long as verse striae above posterior depression. Hind- fourth + fifth; these resp. 3.4 and 3 times as wing: M + Cu divides before cu— a. long as wide; eighth segment 2.3 times as long Metasoma. — Petiolus 1.2 times as long as as wide; sixth to tenth keeled on inner side; hind tibia, ventrally not striate, posteriorly last segment apically pointed (fig. 40). slightly swollen, spiracles prominent (fig. 23); Mesosoma. — Mesoscutum 1.2 times as long stalk of second tergite 1.9 times as long as wide. as wide; scutellum with median impressed line; metapleural flange without tooth; propodeum Holotype. — 9> 'Rep. Malgache, Manjakan- with anterior depression small, median im- driana, Mandraka, 1200 m, 26.V.1972, L. & R. pressed line absent, posterior depression weak; Blommers" (ZMA). integument dull, finely punctulate-reticulate, with sparse punctation, punctures of mesoscu- The type and only specimen of this species tum distinct. Fifth segment of fore tarsi dilated is stylopized, and its morphology may therefore and flattened (though less than in B. scutifera) differ from the normal situation. Important (fig. 41). Hindwing: M + Cu divides before cu- characters, like the absence of an impressed line a. on the propodeum, the short stalk of the second Metasoma. — Petiolus 1.25 times as long as tergite, and the the aberrant colour pattern, hind tibia, ventrally not striate, posteriorly little strong setosity, which separate B. mandraka swollen, spiracles prominent; stalk of second from its closest relatives, viz., B. B. bicolor, tergite 2.4 times as long as wide; seventh sternite apicalis and B. maromandia, indicate that the truncate. type cannot be regarded as an anomalie specimen of one of the known species. Hensen & Blommers: Review of Malagasy Belonogaster 25

Figs. 30 — 35. 30— 31, B. tanosy. 30, $, apical antennal segments (posterior view; 30a, dorsal view); 31, 9. metasomal basis. 32, B. ttpuliformis, $, metasomal basis. 33 — 34, B. trandraka, $. 33, posterior part of metasoma, right side, lateral view, with tooth of metapleural flange indicated (wings omitted); 34, metasomal basis. 35, B. vadoni, $, metasomal basis. 31 — 35: 9 X; 30: 36 X.

Female Material examined. — 6 $ 2 (J, Region The coloration is rather variable; mesosoma d'Ambanja, A. Seyrig (MNHN; 1$ CH, 1 $ dark green with metanotum yellow to ferrug- ZMA); 1 $, Prov. d'Analalava, Maromandia, R. inous, and mesopleuron ventraiiy, metapleuron Deca.ry, 1923 (MNHN; paratype). and propodeum to a variable extent ferruginous; legs black to greenish, the tibiae and tarsi always Belonogaster ornata Saussure light ferruginous. The fore coxae and ventral (figs. 24,25) side of mesopleuron may bear some brown Belonogaster ornata Saussure, 1900: 209. setosity. Typically the propodeum bears some Belonogaster pomicolor S-àussuìie, 1900: 209. oblique striae apically; these are absent in some Belotzogaster eumenoides; Richards, 1982: 47, 110, specimens. figs. 91, 92 (partim).

It is possible that B. po?nicolor is a distinct 26 Tijdschrift voor Entomologie, deel 130, 1987

species, but as long as the male is not known ment; third antennal segment 1.1 times as long we prefer to leave it in the synonymy of B. ornata. as fourth + fifth, these resp. 1.6 and 1.3 times as long as wide; eighth segment as long as wide. Male. Pronotal carina weak. Petiolus 1.1 — L2 times Body length 14.0 mm, length of forewing 9.7 as long as hind tibia, spiracles weakly prominent; mm. stalk of second tergite 1.8 — 2.0 times as long Coloration. — Head pale yellow, vertex green; as wide. antennae ferruginous, scape ventrally yellow, dorsally green. Mesosoma green, meso- and Types. — The lectotype of B. ornata, by metapleuron yellowish ferruginous; posterior present designation, is a $, "Madagasc", margin of scutellum and large parts of metan- "ornatus Sauss., $" (MHNG). Paralectotypes are yellow; tegula ferruginous; propodeal all lectotype of B. otum 5 $ , 4 (5, MHNG. The valves white. Legs green, fore coxae and middle pomicolor, by present designation, is a $, labelled coxae anteriorly and lines along fore and middle "Madagasc" (MHNG); paralectotypes are 5 $, femora and tibiae white; posterior tibiae yellow, all MHNG. proximally and apically infuscated; tarsi fuscous. Other specimens. — 1 (5 1 $, Fampanambo, Petiolus and second metasomal tergite green, x.l962,J. Vadon (KMMA); 2$, Perinei, 950 m, rest of metasoma brown, second tergite with 10.V.1984; R. Hensen & A. Aptroot (CH); 5 $, white band apically, third to sixth tergite and Ivondro, i.1941, 1 $, Fort Dauphin, viii.1940, second to fifth sternite with white lateral spots all A. Seyrig (MNHN; 1 $ CH). apically. Wings light brownish hyaline, veins light brown. Belonogaster prasina Saussure Morphology. — Like the male of B. eunie- Belonogaster prasinus Saussure, 1891: 92, pi. 19, fig. Richards, figs. 87—89. noides, except for the following. Gena 0.3 times 5; 1982: 47, 107, as wide as eye in lateral view; interocular distance Some specimens have the mesosoma nearly on vertex slightly shorter than third antennal entirely yellow. The metasoma usually exhibits segment; third antennal segment 0.7 times as well defined yellow bands but sometimes it is long as fourth + fifth, these resp. 3.3 and 3.0 entirely light ferruginous. Variation in size is times as long as wide; eighth segment 3 times considerable; forewing of female 14.5 —21.0 mm. as long as wide; third to eleventh with raised longitudinal line ventrally; last segment strongly Material examined. — 1 $, Fampanambo, curved, widest subapically (fig. 25); pronotal keel i.1959, J. Vadon (KMMA); 2 $, Ifanadiana, absent; petiolus 1.2 times as long as hind tibia, Ranomafana, 900 m, 29.xii.1971, 2 $, Nosy Be, spiracles weakly prominent; stalk of second Dzamandzar, 8/U.Ì.1971, all L & R. Blommers tergite two times as long as wide (fig. 24). (ZMA); 2 (5 5 $, Fort Dauphin, v.1937, 1 $ 2 $, Région d'Ambanja, 7 $, Sambirano, 1 $, Female. Périnet, ii.1931, all A. Seyrig (MNHN, 1 S Body length 15.2 — 16.8 mm, length of fore- 3 9 CH). wing 10.6 — 12.9 mm. Coloration. — Head green, mandibles and Belonogaster scutifera sp. n. genae, and often clypeus and frons yellow; an- (figs. 26—28) tennae green, scapes ventrally often yellow, Male. flagellum ferruginous. Mesosoma dark green, Body length 20.2 mm, length of forewing 16.1 mesopleuron and often metapleuron, metano- mm. tum yellow to ferruginous, propodeum often Coloration. — Ferruginous; the following ferruginous; propodeal valves whitish. Legs parts pale yellow: mandibles, clypeus except for green, with at least fore and mid tibiae, often a median stripe, frons except medially below also hind tibiae yellow. Coloration of metasoma antennal insertions, ventral half of eye emar- variable: fuscous to green, with or without yellow ginations, propodeal valves, all coxae ventrally, spots. Wings light brownish hyaline to yellow, fore and middle trochanters and all femora veins and stigma light brown to yellow. anteriorly, small apical spots on fore tibiae, Morphology. — Like female B. eumenoides, streak anteriorly on middle and hind tibiae, fifth except for the following. Gena 0.55 times as wide segment of fore tarsi proximally; remaining part as eye in lateral view; interocular distance on of fore tarsi black; antennae dorsally darkened; vertex as long as third + fourth antennal seg- metasoma behind stalk darkened, except broad Hensen & Blommers: Review of Malagasy Belonogaster 27

apical margins of second to fifth segment. Wings as long as wide, eighth segment 1.1 times as yellowish hyaline, veins yellow. long as wide. Pubescence. — Clypeus with dense white Mesosoma, — Like the male but fifth tarsal tomentum, and sparse black setae; vertex with segment not dilated. rather long black setae; mesosoma with sparse Metasoma, — Petiolus 1.2 times as long as white tomentum and sparse very short black hind tibia, ventrally not striate, posteriorly setae; mid and hind femora ventrally with black weakly swollen, spiracles slightly prominent (fig. setae, particularly along the posterior margins. 28); stalk of second tergite 3 times as long as Head, — Clypeus bluntly angled below, angle wide. about 120°; gena 0.35 times as wide as eye in lateral view; cephalic foveae absent; clypeus sparsely finely punctulate, shiny, vertex densely Holotype. — $, "Mus. Roy. Afr. Centr., Ma- dull, with a coarse punctures; punctulate, few dagascar Est: Ambodivoangy, v. I960, J. Vadon" third antennal segment 0.8 times as long as (KMMA), fourth + fifth, these resp. 4 and 3.5 times as Paratype. — $, Fampanambo, ii.l96l,J. Vadon long as wide; eighth segment 1.6 times as long (KMMA), as wide; ninth and tenth segment keeled on inner side, last segment apically pointed (fig. 26). Mesosoma. — Mesoscutum 1.15 times as long Belonogaster tanosy sp. n. as wide; scutellum with median impressed line; (fies. 29—31) metapleural flange without tooth; propodeum Male, with anterior depression obsolete, median im- Body length 15,4 — 17.0 mm, length of fore- pressed line absent, posterior depression weak; wing 10.9 — 11.3 mm (holotype: largest speci- integument dull, finely punctulate-reticulate, men). with sparse punctation, punctures of mesoscu- Coloration. — Head pale yellow, vertex dark tum distinct, though shallow. Legs: fifth segment greenish brown; mandibles pale yellow; anten- of fore tarsi strongly dilated and flattened, shield- nae yellowish ferruginous, first three segments like (fig. 27). Hindwing: M + Cu divides before with greenish tinge. Mesosoma dark green, pron- cu— a. otum laterally, mesopleuron, metapleuron, pos- Metasoma. — Petiolus 1.15 times as long as terior margin of scutellum, metanotum laterally hind tibia, ventrally not transversely striate, and propodeal valves pale yellow; tegulae green- posteriorly swollen, spiracles prominent; stalk ish brown; fore legs and mid coxae pale yellow, of second tergite three times as long as wide; mid and hind legs green, with the last tarsomeres seventh sternite truncate. ferruginous. Metasoma fuscous, petiolus and stalk of second tergite dark green. Wings yel- Female. lowish hyaline, veins yellow. Body length 23.0 mm, length of forewing 17.8 Morphology. — Like the male of B. eume- mm. noides, except for the following. Clypeus 1.1 Coloration. — Ferruginous, femora and tibiae times as long as dorsally wide; gena 0.3 times and antennae dorsally darkened; propodeal as wide as eye in lateral view; interocular width valves reddish yellow; sixth sternite black. on vertex as long as third antennal segment; Pubescence. — Like the male, but clypeus only third antennal segment 0.79 times as long as with black setae and some tomentum, mid and fourth + fifth, these resp. 3,1 and 2.8 times as hind femora ventrally with many black setae, long as wide; eighth segment 2.1 times as long metasoma with brown to black setae on tergites as wide; third to tenth with raised longitudinal and sternites behind the petiolus, setosity of sixth line ventrally, the ones on the apical segments sternite rather long and dense. flattened and polished; last two segments dorsi- Head. — Clypeus acute ventrally, angle about ventrally flattened and concave on inner side; 100°; gena 0.5 times as wide as eye in lateral last segment strongly curved (fig. 30); pronotal view; cephalic foveae absent; clypeus finely punc- keel weak; petiolus 1.05 times as long as hind tulate, and sparsely strongly punctate; vertex tibia, posteriorly distinctly swollen, spiracles dull, punctulate-reticulate, and with sparse shal- strongly prominent (cf. fig. 31) weakly prom- low punctures; third antennal segment as long inent in some of the paratypes); stalk of second as fourth + fifth, these resp. 1.4 and 1.6 times tergite 1.5 times as long as wide. 28 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 36—41. 36— 37, B. dayi, $. 36, apical antennal segments; 37, fore tarsus. 38— 39, B. fanemitra, $. 38, apical antennal segments; 39, fore tarsus. 40—41, B. maromandia, $. 40, apical antennal segments; 41, fore tarsus. 37, 39, 41: 18 X; 36, 38, 40: 36 X.

Female. strongly prominent; stalk of second tergite 1.4 Body length 14.7 — 15.4 mm; length of fore- times as long as wide. wing 9.7 — 10.4 mm. Coloration. — Head ferruginous, clypeus me- Holotype. — Q, "Madagascar, Fort Dauphin, dially and ventrally, and sides of frons yellow; 500 m, 15.iv.1968, K. M. Guichard" (BMNH). vertex dark fuscous; scape greenish black, fla- Paratypes. — 1 9 2 (5", same data as holotype gellum dark ferruginous. Mesosoma ferruginous, (BMNH, 1 $ CH); 1 $, Fénérive, 22.xii.1955, pronotum, mesoscutum and scutellum anteriorly E. McC Callan (BMNH); 3 $, Fénérive-Est, fuscous; propodeum medially darkened, propo- 15.X.1971, L. & R. Blommers (ZMA); 8 $, deal valves yellow. Legs dark green, fore coxae Sambirano, 1 $, Fort Dauphin, vii. 1940, all A. ferruginous, mid coxae dark ferruginous. Wings Seyrig (MNHN, 1 $ CH, 1 $ RMNH). yellowish hyaline, veins yellow. Metasoma fus- tipuliformis sp. n. cous, anterior half of petiolus and stalk of second Belonogaster tergite green. (fig. 32) Specimens from other than the type locality Female. differ as follows: propodeum green; fore and Body length 19.2 mm, length of forewing 14.5 mid tibiae and fore tarsi yellow, mid tarsi and mm. hind tibiae and tarsi fuscous; second metasomal Coloration. — Head green, mandibles, genae tergite with a pair of yellow spots apically. and sides of clypeus and frons yellow; antennae Morphology. — Like female B. eu7nenoides, ferruginous, scape green. Mesosoma yellow, except for the following. Malar space shorter pronotum except ventral corners, mesoscutum than maximal flagellar width; gena 0.4 times and propodeum anterodorsally green. Legs yel- as wide as eye in lateral view; interocular distance low, mid and hind coxae pale green, femora on vertex nearly as long as third + fourth + green, tarsi largely black. Metasoma fuscous, fifth antennal segment; third antennal segment petiolus anterodorsally and ventrally, and stalk 1.1 times as long as fourth + fifth, these resp. of second tergite green, second to sixth tergite 1.2 and 1.1 times as long as wide; eighth segment with apical yellow band which is interrupted in 0.9 times as long as wide. Pronotal keel weak. the middle on second to fourth; second to sixth Petiolus (fig. 31) 1.3 times as long as hind tibia, sternite largely yellow. Wings proximally yel- posteriorly distinctly swollen, spiracles rather lowish grey, apical third bright yellow. Hensen & Blommers: Review of Malagasy Belonogaster 29

Figs. 42 —45. Nests of Belonogaster. Al, E. dayi, at Périnet (courtesy of Dr W. G. d'Arcy, Missouri); 43, B. eumenoides, at Manjakandriana, Mandraka, nest nr. 73.20; 44, B. guerini, at Fénérive, nest or. 71.58; 45, B. madecassa, at Tsaramandroso, nest nr. 72.23.

Pubescence. — Clypeus ventrally with brown Head. — Clypeus acute below, angle about setae, vertex and mesosoma dorsally with sparse 100°; gena 0.7 times as wide as eye in lateral short yellowish setae; fore coxa with dense white view; cephalic foveae absent; clypeus sparsely pubescence, and several longer black setae; mid punctate, dorsal two thirds densely punctulate; and hind femora with some white setae beneath; vertex densely punctulate, with a few indistinct metasoma without black setosity. punctures; third antennal segment 1.1 times as long as fourth + fifth, these resp. 1.5 and 1.4 30 Tijdschrift voor Entomologie, deel 130, 1987

times as long as wide; eighth segment 1.1 times behind the mesopleural spiracular lobe (fig. 33); as long as wide. propodeum with distinct anterior depression, Mesosoma. — Mesoscutum 1.1 times as long impressed median line only present just above as wide; metapleural flange without tooth; prop- the small, shallow posterior depression. Integ- odeum with anterior depression small, posterior ument punctulate-reticulate, and densely, rather depression small but deep, impressed median coarsely punctate, interspaces as large as the line very short; integument punctulate-reticu- punctures on the mesoscutum. Hindwing: M + late, sparsely shallowly punctate, fairly shiny. Cu divides before cu— a. Hindwing: M + Cu divides after cu-a. Metasoma. — Petiolus 1.5 times as long as Metasoma. — Petiolus 1.2 times as long as hind tibia, ventrally not striate, posteriorly hind tibia, ventrally not striate, posteriorly dis- hardly swollen, spiracles not prominent (fig. 34); tinctly swollen, spiracles prominent (fig. 32); stalk of second tergite as long as wide. stalk of second tergite 3.5 times as long as wide. Holotype. — $, "Mus. Roy. Afr. Centr., Ma-

Holotype. — $, "Madagascar; (Tamat.); Pé- dagascar Est: Ambodivoangy, v. I960, J. Vadon" rinet; 950 m; 48° 16 E, 18° 56 S; lO.v.1984; leg. (KMMA). R. Mensen & A. Aptroot" (RMNH). Paratypes. — 1 ?, "Madagascar, collection le Etymology. — "Trandraka" is the local name Moult", 1 $, Sambirano, Manongarivo, 1150 m, for one of the peculiar Malagasy hedgehogs, or xii.1960, R Griveaud (both MNHN). Tenrecs, insectivorous mammals of the family Tenrecidae. The name is chosen because of the Belonogaster trandraka sp. n. long erect pubescence and similar colour of the (figs. 33, 34) wasp. Female. Body length 20.6 mm, length of forewing 14.2 Belonogaster vadoni sp. n. mm. (fig. 35) Coloration. — Ferruginous; antennae infus- Female. cated, except the apical segments ventrally; legs Body length 26.3 mm, length of forewing 17.7 dark brown, but fore coxae ferruginous, mid mm. coxae, fore and mid femora and fore tibiae Coloration. — Yellow, antennae bright fer- greenish; propodeal valves pale yellow; meta- ruginous, pronotum dorsally with a pair of green soma fuscous, petiolus ventrally and dorsally on markings; petiolus suffused with green, second anterior half green. Wings light brown, veins and third tergite anteriorly green. Wings yellow, and stigma yellowish to reddish ferruginous. veins and stigma yellow. Pubescence. — Clypeus with rather long yel- Pubescence. — Clypeus with yellow, frons and low setae; vertex and mesosoma dorsally with vertex with conspicuous short black setae; mes- long rather dense black setosity, setae half as osoma with black setosity, on mesonotum rather long as flagellar width; fore coxae only with short dense and long; fore coxae with long dense black white pubescence; mid and hind femora ventrally setosity, and white tomentum; mid and hind with sparse white pubescence; setosity of pos- femora ventrally only with short pale pubes- terior tergites and sternites light brownish, com- cence; metasoma with yellowish tomentum and paratively short. yellow setae. Head. — Clypeus acute below, angle about Head. — Clypeus acute below, angle about 100°; gena in lateral view 0.6 times as wide as 90° gena 0.7 times as wide as eye in lateral ; eye; cephalic foveae absent; ventral third of view; cephalic foveae absent; clypeus ventrally clypeus granulate, sparsely foveolate, dorsal two coarsely punctate, dorsal three-fifth densely thirds punctulate, with sparse coarse punctures; punctulate and sparsely rather finely punctate; vertex punctulate-reticulate, with distinct punc- vertex dull, punctulate-reticulate, punctation in- tation. Third antennal segment 1.1 times as long distinct; third antennal segment 1.1 times as long as fourth + fifth, these resp. 1.1 and 1.0 times as fourth + fifth, these both 1.2 times as long as long as wide; eighth segment 0.9 times as as wide, eighth segment 0.9 times as long as long as wide. wide. Mesosoma. — Mesoscutum as long as wide; Mesosoma. — Mesoscutum 1.1 times as long scutellum without impressed median line; me- as wide; scutellum with median line weakly sopleural flange produced into a flat tooth, impressed on posterior half; metapleural flange Hensen & Blommers: Review of Malagasy Belonogaster 31 without tooth; propodeum without anterior de- References pression, median impressed line indicated on Richards, O. W., 1982. A revision of the genus posterior half, posterior depression well deve- Belonogaster de Saussure (Hymenoptera; Vespi- loped; integument dull, finely punctulate-retic- dae). — Bulletin of the British Museum (Natural History), Entomology 44 ulate, with sparse shallow, indistinct punctation; (2): 31 — 114. Saussure, H. L. F. de, 1853 54. Monographie des propodeum with strong oblique striae above the — guêpes sociales la — ou de tribu des Vespiens: i. xxi posterior depression. Hindwing: M + Cu divides + 1-590. — Paris cSc Genova (1—96, 1853, before cu— a. 97—256, 1854). Metasoma. — Petiolus 1.3 times as long as Saussure, H. L. F. de, 1891. In: Grandidier, A., Histoire hind tibia, ventrally not transversely striate, Physique naturelle et politique de Madagascar. 20. posteriorly swollen, spiracles prominent (fig. Histoire Naturelle des Hyménoptères. Première partie: i— xxi -|- 1^590, 27 pis. — 35); stalk of second tergite 1.7 times as long Paris. Saussure, H. L F. de, 1900. Wissenschaftliche Er- as wide. gebnisse der Reisen in Madagascar und Ostafrika in der Jahren 1889—1895 von Dr. A. Voeltzkow. Holotype. — 9> "Coll. Mus Tervuren, N. E. Hymenoptera Vespidae. — Abhandlungen der

Madagascar, Fampanambo, i.1959, J. Vadon" Senckenbergischen Naturforschenden Gesell- (KMMA). schaft 262: 203—210. Smith, F., 1857. Catalogue of the hymenopterous insects in the collection of the British Museum. Part 5. Vespidae: i— iv + 1—147. — London.

Tijdschrift voor Entomologie 130: 33 —47 Gepubliceerd 30 november 1987

MORPHOLOGIE DER LARVEN UND PUPPEN EINIGER PHYLIDOREA-ARTEN (DIPTERA, LIMONIIDAE)

JOLANTA WIEDENSKA

Instytut Biologi! Êrodowiskowej, Lòdi, Polen

Abstract Die Larven und Puppen von vier Arten der Gattung Phylidorea Bigot (Limoniidae, He-

xatominae) werden beschrieben, Phylidorea (s. Str.) nigricollis (Mg.) und Phylidorea {Euphylidorea) nigronotata (Siebke) zum ersten Mal. Die Beschreibungen und Illustrationen von Phylidorea (s. str.) squalens Zett. und Phylidorea {Euphylidorea) fulvonervosa (Schumm.) sind umfassender und genauer als frühere.

Als Material für die Beschreibungen wurden die Exuvien von Larven des 4. Stadiums und von Puppen benutzt.

Einleitung

In Europa wurde bisher das Vorkommen von (s. Str.) nigricollis (Mg.) und Phylidorea 15 Arten der Gattung Phylidorea Bigot (sensu {Euphylidorea) nigronotata (Siebke). Die Be- Alexander, 1972) festgestellt. Dagegen sind nur schreibungen von Phylidorea (s. str.) squalens sieben Arten in den Praeimaginalstadien be- Zett. und Phylidorea {Euphylidorea) fulvoner- kannt. vosa (Schumm.) dagegen enthalten neue, bisher Die Beschreibungen der Larven und Puppen in der Literatur unbekannte Einzelheiten. von diesen Arten sind meist unvollständig (Ba- Es wurde die Taxonomie von Phylidorea Bigot ling, 1878, 1886; De Meijere, 1916; Levy, 1918; nach Alexander (1972) und Mendl (1978) an- Brindle, 1958, 1967; Hennig, 1968). Die Autoren gewandt, obwohl Stary (1981) und Savtshenko dieser Arbeiten haben oft solche Merkmale be- ( 1986a, b) innerhalb dieser Gattung wesentliche rücksichtigt, die für die ganze Gattung oder viele Veränderungen vorschlagen. Phylidorea- Arten zutreffen, deshalb sind diese Herrn Doz. Dr. Krzysztof Jazdzewski danke Charakteristiken keine guten diagnostischen Be- ich sehr herzlich für wertvolle Anregungen zu schreibungen. Von grösserer Bedeutung sind die meiner Arbeit. von Brindle und Bryce (I960) zusammengestell- ten Bestimmungstabellen, weil sie nicht nur den Analsegmentbau sondern auch den Kopfkapsel- Material und Methoden bau berücksichtigen. Diese Bestimmungstabellen Die Untersuchungen wurden von Herbst 1980 werden jedoch von den Autoren selbst für ziem- bis Herbst 1983 durchgeführt. Das Material lich provisorisch gehalten, weil sie nicht alle Ar- wurde im Lubrzanka-Fluss (rechter Nebenfluss ten enthalten und die angegebenen Merkmale des Czarna Nida-Fluss, àwiçtokrzyskie Gebirge) meistens keine diagnostischen Merkmale einzel- und im Grabia-Fluss (rechter Nebenfluss des Wi- ner Arten sind. Alle bisher bekannten Jugend- dawka-Fluss, Central Polen) gesammelt. stadien der Limonaden sind in den Bestim- Die Larven des 4. Stadiums leben im Ufer- mungstabellen von Rozkosny und Pokorny schlamm an der Wasserlinie. Sie wurden mit ei- (1980) enthalten. nem hydrobiologischen Netz (Maschenweite: 0,5 Meine Arbeit betrifft die Larvenmorphologie mm) entnommen, vorsichtig ausgespült und in des 4. Stadiums und die Morphologie der männ- Thermosflaschen in das Laboratorium transpor- lichen Puppen von vier Arten der Gattung tiert. Dort wurde jede Larve einzeln in einer Phylidorea Bigot. Zwei Arten werden zum ersten Kunstoffschale mit sauberem Sand und Fluss- Mal von mir beschrieben, nämlich Phylidorea wasser bis zur Imago zogen. Beobachtungen an

33 34 Tijdschrift voor Entomologie, deel 130, 1987 lebenden Tieren erfolgten durch den Schalen- ich das Vorkommen von 7 Arten der Gattung deckel. Phylidorea Bigot festgestellt, aber nur vier Arten Imago, zugehörige Exuvien der Puppe und des habe ich bis zur Imago oder Puppe züchten kön- letzten Larvenstadiums wurden nach dem nen. Schlüpfen in 75% Alkohol konserviert. Die Phylidorea (Ph.) nigricoUis (Mg.) Kopfkapseln der Larvenexuvien wurden in mi- kroskopischen Dauernpräparaten in Canadabal- Limnobia nigricoliis Meigen, 1830: 276. Limnophila nigricoliis (Mg.); de Meijere, 1921: 85. sam verarbeitet. Limnophila {Phylidorea) nigricoliis (Mg.); Stary, Von etwa 90 züchteten Larven haben sich acht 1970: 146. zur Imago entwickelt, weitere acht haben sich Verbreitung: Mittel- und Nordeuropa (Stary verpuppt, die übrigen erreichten das 4. Stadium. 1970; Savtshenko 1986 b). Für meine Beschreibungen benutzte ich nur Ex- Material: 3 (5, 5 9- Lubrzanka-Fluss: Zagnarisk — emplare, die sich zumindest bis zur Puppe ent- Gruszka, 1 Larve: 22. xi. 1980 gesammelt, 29.xii.1980, wickelt hatten. S geschlüpft; Marzysz, 1 Larve: 20. v. 1981 gesammelt, Dieser Material bildet nur ein Teil meiner von 30.V.1981, $ geschlüpft; Marzysz, 1 Larve: 20.V.1981 Swiçtokrzyskie Gebirge bearbeitenen Samm- gesammelt, 21. v. 1981, $ verpuppt; Marzysz, 1 Larve: lung/ Wiederiska 1986/. Auf diesem Gebiet habe 14. V. 1982 gesammelt, 21.V.1982, $ verpuppt. Grabia-

nigricoliis. Fig. 1. Analsegment von Phylidorea (Ph.) nigricoliis. Dorsalansicht. Fig. 2. Kopfkapsei Phylidorea {Ph.) Dorsalansicht. WiEDENSKA: Larven und Puppen Phylidorea- Arten 35

Fluss: Grabica, 1 Larve: 18. v. 1982 gesammelt, Länge: 1,5 — 1,9 mm. Die Lateralplatten mit den 06.vi.1982, $ geschlüpft; Grabica, 1 Larve: 06.V.1983 tiefen Dorsalspalten in Externo- und Interno- gesammelt, 19.v. 1983, geschlüpft; Zamosc, 1 Larve: $ lateralia geteilt. Internolateralia hinter der Fron- 12.iv.1983 gesammelt, 22.iv.1983, $ geschlüpft; Za- talplatte verbunden, am Hinterrand der Kopf- mosc, 1 Larve: 12. v. 1983 gesammelt, 23. v. 1983, $ geschlüpft. kapsel durch die schmale, nicht sehr tiefe Coronalspalte geteilt. Alle Platten fein und fast Länge der Larven des 4. Stadiums 17 — 20 mm. durchsichtig; stark skierotisiert sind nur die Lei- Körper bedeckt mit sehr kurzer und zarter, aber sten, die die Ränder dieser Platten bilden. Die dichter brauner und schillernder Behaarung. Externolateralia rinnenförmig, auf der Ventral- Verletztes Segment charakteristisch ange- seite durch starke Leisten mit dem Hypopharynx schwollen. verbunden. Die Verbindungsstelle der länglichen Auf der Ventralseite des Analsegments (Fig. Leisten, die die Antennen und Mundgliedmassen 1) vier ziemlich grosse, ovale Analpapillen. Stig- tragen, ist am kräftigsten skierotisiert. menfeld auf der Dorsalseite gelegen, Stigmen Antennen (Fig. 3a) zweigliederig. Basalglied deutlich pigmentiert, oval. Vier Randlappen: die 75,7—100,7 |im lang, 22,2—30,7 \im breit. Api- zwei Ventrallappen lang und breit, die beiden kalglied 41,5 — 56,9 |im lang, 6,3 — 7,9 l^m breit, Laterallappen kurz und stämmig, nur etwa halb fein gestreift. Auf dem Basalglied zwei schlanke so lang wie die Ventrallappen. Randlappen mit Sensillen und eine Borste, die etwa halb so lang hellbraunen Streifen (schmale auf den Lateral- wie das Apikaiglied ist. Sockel breit, stark skle- lappen, breite an der Basis auf den Ventrallap- rotisiert, unregelmässig geformt. pen), die sich auf dem Stigmenfeld nicht mit Clypeolabrum (Fig. 4a): Länge 149,6— 174,1 einander verbinden. Distalränder der Ventral- \xm. Breite 290,2 — 340,3 |im, ziemlich stark skle- lappen fast schwarz, stark skierotisiert, mit einer rotisiert. Sutur zwischen Clypeus und Labrum dichten Reihe langer und weicher Haare ge- undeutlich. Lateralränder mit stark sklerotisier- säumt. Das Stigmenfeld nackt. ten "Flügelchen". Am Vorderrand des Clypeo- Kopfkapsel (Fig. 2), wie bei anderen labrums (Fig. 5a) verschiedenartige Strukturen, Phylidorea- Anen schwach und zart skierotisiert.

Fig. 3. Antennen Phylidorea-Larven: a, Ph. (Ph.) nigricollis; b, Ph. (Ph.) squalens; c, Ph. (E.) ntgronotata; d, Ph. (E). fulvonervosa. 36 Tijdschrift voor Entomologie, deel 130, 1987 die symmetrisch angeordnet sind: links und Mandibeln (Fig. 6a, e) 352,8—405,1 |im lang, rechts von der Medianen zwei kräftige, etwas 129,2 — 156,5 (im breit. Ziemlich stark skiero- abgestumpfte Dorne; seitlich davon, auf vorste- tisiert, besonders im unregelmässigen und ziem- henden Erhöhungen, je drei fingerförmige, ne- lich bauchigen Proximalteil. Der Apikaiteil beneinander liegende Fortsätze; daneben je zwei dorso-ventral abgeflacht, mit langem, gleichmäs- feine Sensillen (finger- und lanzettförmig) und sig verjüngtem und zur Mitte umgebogenem auf den am weitesten lateral gelegenen, deut- Zahn. Am Innerrand dieses Zahns nahe seiner lichen Erhöhungen je drei Fortsätze mit schar- Basis eine schwach skierotisierte, fast durchsich- tigen Rändern. Auf der Fläche etwas vom Vor- tige, schmale Klinge, die fast die Hälfte der derrand entfernt, lateral je ein selbst ziemlich Zahnlänge erreicht. Unter dem ersten, dem Api- kräftig skierotisierter Fortsatz mit unregelmäs- kalzahn, noch vier Zähne: der zweite und dritte sigen Rändern auf stark skierotisierten Erhebun- Zahn blättrig, fein, in der Höhe ausgebreitet; gen. Daneben, zur Medianen hin, zwei weiche, der vierte Zahn dreieckig, spitz endend und stark streitkolbenförmige, niedrige Warzen. skierotisiert; der fünfte Zahn klein, schmal, ab- Epipharynx (Fig. 5b) auf der Ventralseite des gestutzt, mit unregelmässigem Rand. Die Ka- Clypeolabrums gelegen. Symmetrisch zur Kör- nalmündungen befinden sich an der Basis des perachse befinden sich von vorn nach hinten: zweiten Zahns und auf dem Basalvorsprung der ein Paar zweigliederige, fingerförmige Fortsätze Mandibel weit unterhalb des fünften Zahns. Auf und drei Paar Sensillen. Ausserhalb dieser Rei- der Aussenfläche der Mandibel, ein Drittel von hen auf sehr kräftig skierotisierten, ausgedehn- der Basis entfernt, zwei lange Borsten. ten Erhöhungen zwei lange, steife Borsten, die MaxiUen (Fig. 7a): etwa 500 |im lang, an der an der Rändern dieser Erhöhungen liegen. Nä- Basis 106,4— 169,9 |im breit. Im Basalteil ziem- her zum Aussenrand des Epipharynx ein kleines lich stark skierotisiert, zur Spitze hin schwächer. Büschel niedriger, dicker Haare. Der membranartige Apikaiteil an den stark skle-

100 xjm

Fig. 4. Clypeolabrum der Phylidorea-La.rven: a, Ph. (Ph.) nigricollis; b, Ph. (Ph.) squalens; c, P/j. (E.) nigronotata; d, Ph. (£.) fulvonervosa. —

WiEDENSKA: Larven und Puppen Phylidorea- Arten 37

rotisierten, spitze zulaufenden Keil angelehnt. Die unbewegliche Puppe ist dunkelbraun, hell Die Innenflächen der Maxillen im Apikaiteil nur an den Segmentgrenzen. Die Länge der dicht behaart. männlichen Puppe beträgt etwa 15 mm, der Hypopharynx (Fig. 8a, b) membranös, von weiblichen etwa 17 mm. der Querbrücke und zwei Lateralarmen begrenzt. Die Kopfscheide (Fig. 9a) ist flach, dunkler Am Vorderrand lateral zwei säulenartige Labial- als der Puppenkörper. Die Labralscheide (Breite palpen, von 34,7 —35,9 ^m Höhe; die Entfer- 344 — 358 |im) ist gläschenförmig, etwas vor die nung zwischen beiden beträgt 58,3 —63,7 )im. Labialscheide vorgeschoben. Die Maxillentaster- Auf der platten Apikaifläche des Palpus eine scheiden (Länge etwa 630 ^m) sind allmählich schlanke Papille (Länge 10,8 — 12,5 jam). Die zur Spitze hin verschmälert, etwas nach oben Querbrücke (Länge 44,4—72,8 |im. Breite gekrümmt. Die Labialtasterscheiden (Länge 195,2 — 228,7 )a.m) ist stark gleichmässig skle- etwa 230 |im) sind schlank, von charakteristi- rotisiert, mit fast parallelen Rändern. Die La- scher Gestalt: im Mittelteil stark gewölbt, im teralarme (Länge 177,0— 215,7 |im. Breite Distalteil dagegen abrupt verengt. 130,9— 172,4 |im) sind unregelmässig und platt. Pronotalhörner (Fig. 10a) 900—930 )im lang, Ihre Aussen- und Vorderränder sind umgebogen röhrchenförmig, an der Basis rund, im Apikaiteil und stärker skierotisiert als die übrigen Teile. abgeplattet. Oberfläche mit skierotisierten La- Hypostomium nicht entwickelt. mellen bedeckt, sehr fein gekerbt. Die Ränder

Fig. 5. Vorderrand des Clypeolabrums (a, c, e, — g) und Vorderteil des Epipharynx (b, d, f, h) der Phylidorea-Larwen: a b, Ph. (Ph.) tiigricollis; c d, Ph.(Ph.) squalens; e— f, Ph. (E.) nigronotata; g— h, Ph. (E.) fulvonervosa. 38 Tijdschrift voor Entomologie, deel 130, 1987 WiEDENSKA: Larven und Puppen Phylidorea-Arten 39

100 Aim

Fig. 7. Maxillen der Pbylidorea-Larven: a, Ph. (Ph.) nigrkollis; b, Ph. {Ph.) squalens; c, Ph. (E.) iro7iotata; d, Ph. (E.) fulvonervosa. der Atmungsspalte membranartig, mit spitz ge- (AN) an dem gemeinsamen Stamm sind lang franstem Chitin verstärkt. Da die Sklerotisierung und schlank. der Pronotalhörner stark ist, sind Tracheen- stamm und Atmungskammer unsichtbar. Phylidorea (Ph.) squalens (Zett.) Die Flügelscheiden reichen bis zum Ende des Limnobia squalens Zetterstedt, 1838: 834. 2. Abdominalsegments. Die gleichartigen Bein- Limnophila bicolor (Zett.); Lundström, 1912: 63. scheiden reichen fast bis zum Ende des 3. Ab- Limnophila squalens (Zett.); de Meijere, 1921: 73. dominalsegments. Litnnophila {Phylidorea) squalens (Zett.); Stary, 1970: 147. Die Abdominaltergite und -sternite sind mit Praeimaginalstadien: unregelmässigen Plättchen bedeckt (Fig. IIa), Limnophila (^Phylidorea) squalens (Zett.); Brindle & die ein ziemlich symmetrisches Muster bilden: Bryce, I960: 217, Fig. 13 (Analsegment); Brindle zwei horizontale Reihen mit dicht nebeneinan- 1967: 199, Fig. 48, 52 (Analsegment), Fig. 123 der angeordneten Plättchen und zwei vertikale (Pronotalhorn). Reihen an den Segmentseiten bilden ein Recht- Verbreitung: Mittel- und Nordeuropa, Nord- eck, dessen Fläche unregelmässig mit Plättchen und Ostsibirien (Stary, 1970; Savtshenko, bedeckt ist. Die Sternite sind dichter als die Ter- 1986b). gite mit Plättchen bedeckt. In der Mitte von Ter- git und Sternit befindet sich das unpaare Stigma. Material: 2 Q. Lubrzanka-Fluss: Ameliówka, 1 Am Hinterrand des vorletzten Abdominalseg- Larve: 20. v. 1981 gesammelt, 25. v. 1981, $ verpuppt; ments steht eine Reihe dicht angeordneter, spit- Zagnarisk — Jaworze, 1 Larve: 23. iv. 1983 gesammelt, zer Dorne. 04.V.1983, (5 geschlüpft. Analsegment des $ (Fig. 12a) schlank mit ei- Die Länge der Larven des 4. Stadiums beträgt nem Paar unregelmässiger und ziemlich weicher etwa 10— 12 mm. Bei makroskopischer Betrach- Lateralvorsprünge (L). Zwei Paar Dorsalvor- tung ist die Larve anderen Phylidorea-Arten ähn- sprünge: die Anterodorsalvorsprünge (AD) sind lich, aber bedeutend kleiner. Ihr Körper ist mit klein, die Posterodorsalvorsprünge (PD) dage- einer dunkelbraunen und intensiv schillernden gen grösser und stämmig. Die Analvorsprünge Behaarung bedeckt. Analpapillen klein und ku- 40 Tijdschrift voor Entomologie, deel 130, 1987

gelförmig. Ventrallappen lang und breit an der glieds eine steifen Borste, die etwas länger als das Basis. Laterallappen fast so lang wie die Ventral- Apikalglied ist. Sensillen fehlen. lappen. Randbehaarung sehr lang, pigmentierte Clypeolabrum (Fig. 4b) 111,0— 145,1 ^m lang, Streifen auf den Randlappen deutlich und breit. 146,8—207,8 |im breit, trapezförmig mit sanft Die Länge der Kopfkapsel beträgt 0.9 — 1,3 abgegrundeten Ecken. Lateralränder ohne skle- mm. rotisierte "Flügelchen". Am Vorderrand (Fig. 5c) Antennen (Fig. 3b): Basalglied 24,6 \im lang, zwei Paar stärkerer Borsten, seitlich davon 17,7 (am breit; Apikaiglied 35,3 |im lang, 6,3 ^m scharfe, in einer Vertiefung angelegte Dorne, breit. Das Apikalglied ist fein gestreift und sitzt flankiert von einer Papille. auf dem Basalglied versenkt in einer kragenför- Epipharynx (Fig. 5d) am Vorderrand nahe des migen Krause. Auf der Distalfläche des Basal- Medianen mit einem Paar lanzettförmiger Papil-

100 /Um

Fig. 8. Hypopharynx der Phylidorea-La.T\ren und seine Grundbestandteile: a, Hypopharynx von Ph. (Ph.)

nigricollis; b, Querbrücke und Lateralarm von Ph. {Ph.) nigricollts; c, Ph. {Ph.) squalens; d, Ph. (£.) nigronotata;

e, Ph. (E.) fulvonervosa; i. Labium von Ph. {Ph.) squalens; g, Ph. {E.) nigronotata; h, Ph. {E.) fulvonervosa. WiEDENSKA: Larven und Puppen Phylidorea-Arten 41

len, dahinter ein Paar zweigliederige Fortsätze Analsegment $ (Fig. 12b) kurz und untersetzt mit kleines Papille an der Basis, seitlich davon mit zwei Paar Lateralversprüngen und zwei Paar zwei stark skierotisierte Erhöhungen mit dichter verschiedenartigen kegelförmigen Dorsalvor- Behaarung. Einzeln stehende Borsten fehlen. sprüngen. Vor jedem Anterodorsalvorsprung Mandibeln (Fig. 6b, f) nur 196,6—273,1 |im eine kräftige, kurze, spitze Borste. Die beiden lang und 84,2 — 114,9 |Jm breit. Die schlanke und kurzen Analvorsprünge haben separate Stämme. durchsichtige Klinge am Innenrand des Apikai- zahns erreicht fast zwei Drittel seiner Länge. Der Phylidorea (Euphylidorea) nigronotata zweite Zahn ist platt mit einer ziemlich breiten (Siebke) Klinge, der dritte bedeutend kleiner und schlank. Limnobia nigronotata Siebke, 1870: 305. Anstelle des fünften Zahns steht eine geringe Phylidorea (Macrolabina) nigronotata (Siebke); Savtshenko, Erhöhung mit der Kanalmündung. Auf der Aus- 1986 a: 20. senfläche der Mandibel befindet sich zwei unter- Verbreitung: Nord- und Mitteleuropa, Sibi- schiedlich lange, ziemlich dicke aber weiche Bor- rien, Zentralasien (Savtshenko 1986 a). sten.

Maxillen (Fig. 7b) etwa 250 fam lang und etwa Material: 3 $ Lubrzanka-Fluss: Marzysz, 1 Larve: Ol. iv. 1982 gesammelt, iv. 100 |im breit, fast auf ganzer Länge gleich breit. 13. 1982 $ verpuppt. Gra- bia-Fluss: Zamosc, 1 Larve: 07. iv. 1982 gesammelt, Hypopharynx (Fig. 8g, f): die Querbrücke 22,2 13.iv.1982 (5 verpuppt: Zimne Wody, 1 Larve: |im lang in der Mitte, 84,8 |im breit; die Latera- 06.V.1983 gesammelt, 08.V.1983 S verpuppt. larme 155,3—191,2 um lang und 69,4—101,3 ^m Die Larven des 4. Stadiums 15 —20 mm lang, breit. Die Querbrücke ist unterschiedlich stark Ihr Körper ist sehr zart und schillernd behaart. skierotisiert, am schwächsten in der Vertiefung Das vorletzte Segment ist — wie bei allen in der Mitte. Die Labialpalpen 29,6 )im, die Pa- Phylidorea- Anen — angeschwollen. pillen 10,8 |im höh. Die Analpapillen sind lang und sehr faltig. Hypostomium nicht entwickelt. Alle vier Randlappen sind lang und schlank, die Die Puppe ist hellbraun, ihre Länge beträgt Laterallappen erreichen zwei Drittel der Ven- etwa 9 mm. trallappenlänge. Randlappen mit sehr blass pig- Die Kopfscheide (Fig. 9b) hat dieselbe Farbe mentierten Streifen und ziemlich langer Randbe- wie der Körper. Die Labralscheide (Breite 200 haarung. |j,m) ist oval, lang und deutlich sichtbar ausser- Die Kopfkapsel ist 1,7 — 1,8 mm lang und halb vor die Basis der Labialtasterscheiden vor- schwach skierotisiert. geschoben. Die Maxillentasterscheiden (Länge: Antennen (Fig. 3c): Basalglied 51,2 —73,4 )am etwa 340 |am) wirken untersetzt, sie sind allmäh- lang, 23,9—26,7 |im breit; Apikaiglied fein ge- lich zur Spitze hinverschmälert. Die Labialtaster- streift, 37,0—47,8 \\.m lang, 8,0 —9,7 i^m breit. scheiden (Länge: etwa 140 ^m) sind kurz, unter- Sockel besonders gross, breit und skierotisiert. setzt, schaufeiförmig. Im Proximalteil des Basalglieds ein Porus; auf Die Pronotalhörner (Fig. 10b) (Länge der Distalfläche zwei geringe, untersetzte Sensil- 230— 270 i^m) sind klein, untersetzt, len und eine starke steife Borste, die etwas kürzer kegeistumpfförmig, mit gerundeten Ecken. Im als das Apikaiglied ist. Durchmesser sind sie an der Basis rund, bauchig, Clypeolabrum (Fig. 4c): 133,7 — 140,5 |im dagegen im Apikaiteil abgeflacht. Ihre Fläche ist lang, 263,1 — 260,6 |im breit, ziemlich stark skle- eben. Die Atmungskammer und der Tracheen- rotisiert. Lateralränder mit stark skierotisierten stamm scheinen durch. "Flügelchen". Am Vorderrand (Fig. 5e) median Die Flügelscheiden reichen bis zum Ende des zwei stark skierotisierte Plättchen von unregel- 2. Abdominalsegments, die Beinscheiden fast bis mässiger Form zum Vorderrand hin hochgebo- zum Ende des 3- Abdominalsegments. gen, mit ausgefransten Oberkanten. Auf diesen Die kutikulare Oberflächenstruktur auf Tergi- Plättchen je zwei Borsten: eine kurze steife und ten und Sterniten (Fig. IIb) besteht aus spitzen daneben eine lange schwanke. Symmetrisch zu Dornen, die ein Rechteck bilden. Seine Fläche ist den Seiten hin je zwei verschiedengebaute Sen- unregelmässig mit Dornen bedeckt. An den La- sillen: eine zylindrisch abgestumpft, mit breiter teralrändern der Tergite und Sternite befinden Basis, die andere zweigliedrig mit schmälern Ba- sich unregelmässige, dunkelbraune Pigment- salglied. Und etwas weiter auf die Fläche gerückt streifen. Im Distaiteli und in einem Zweitel der stehen je zwei Sinnesgruben. Länge jedes Abdominalpleurits steht ein grosser Epipharynx (Fig. 5f) auf der Dorn. Ventralseite des deel 1987 42 Tijdschrift voor Entomologie, 130,

o, o WiEDENSKA: Larven und Puppen Phylidorea- Arten 43

Clypeolabrums gelegen. Ausgerüstet mit zwei lang wie der Apikaizahn. Der zweite und dritte Paaren zweigliedriger, verschiedenlanger, fin- Zahn sind schmal, mit fast parallelen Rändern. gerförmiger Fortsätze. An deren Basis ein zartes Der vierte Zahn ist kegelförmig, an der Spitze Haarfeld. Auf den seitlichen Erhebungen steht je abgestutzt, ziemlich kräftig entwickelt. Die Ka- eine lange steife Borste. nalmündungen liegen an der Basis des zweites Mandibeln (Fig. 6c, g): 302,7—315,7 \am lang, Zahns und etwas unterhalb des kleinen fünftes 115,5—135,4 um breit. Die schmale Klinge am Zahns. Innenrand des Apikaizahns ist mehr als halb so MaxiUen (Fig. 7c): etwa 450—500 \i lang und

150 AJm

I 1

Fig. 10. Pronotalhörner der Phyltdorea-Vnç^Qn: a, Ph. (Ph.) fiigricollts; b. Ph. (Ph.) squalens; c, Ph (E) nigronotata;à,Ph.(E.)fulvonervosa. 44 Tijdschrift voor Entomologie, deel 130, 1987

etwa 130 |i breit, lang und schlank. Der dunkel etwas verengt; ihre leicht konkav gebogener skierotisierte Keil, der den häutigen Apikaiteil Rand reicht nicht an die Basis den Labialtaster stüzt, ist sehr lang. heran. Die Maxillentasterscheiden (Länge etwa Hypopharynx (Fig. 8d, g): Querbrücke 470 (im) sind an der Basis breit, bauchig und im 26,7—50,6 |am lang, 156,5—162,7 ^m breit; La- Distaiteli stark verschmälert. Die Labialtas- teralarm 176,4—190,6 um lang, 129,7—138,8 terscheiden (Länge etwa 300 (am) sind unter- |am breit. Die Querbrücke ist oval mit bis- setzt, keulenförmig. kuitförmig verbreiterten Enden. Der Lateralarm Die Pronotalhörner (Fig. 10c) sind schwach ist sehr unregelmässig geformt. Die Höhe der skierotisiert, so dass Tracheenstamm und At- Labialpalpen misst 13,1 |^m und die der Papillen mungskammer durchscheinen. Ihre Form ist ke- 12,5 )im; die Entfernung zwischen den Labialpal- gelstumpfartig, im Durchmesser sind sie an der pen beträgt 43,8 |im. Basis rund, zum Apikaiteil hinabgeflacht oval, Hypostomium nicht entwickelt. die Spitze ist abgestumpft. Ihre Länge beträgt Die Puppe ist dunkelbraun und etwa 17 mm etwa 450 (im. lang. Der Kopf wirkt ziemlich schmal im Ver- Die Flügelsscheiden reichen bis zum Ende des hältnis zum Körper. Die Mundgliedmassen- 2. Abdominalsegments, die Beinscheiden fast bis scheiden sind in Fig. 9c abgebildet. Die Labral- zum Ende des 3. Segments. scheide (Breite etwa 250 (am) ist im vorderen Teil Die Musterung auf den Abdominalsegmenten

I 1

Fig. 11. Kutikulare Oberflaächenstruktur von Phylidorea-Puppen: a, Ph.GPh.) nigricolUs; b, Ph. (Ph.) squalens; c. Ph. (£.) nigronotata; d, Ph. (E.) fulvonervosa. WiEDENSKA: Larven und Puppen PhyUdorea- Arten 45

ist unterschiedlich. Auf den Tergiten ist nur je- sprünge sind kegelförmig: die Anterodorsalvor- weils eine Reihe Plättchen am Distelrand deut- sprünge kleiner, die Posterodorsalvorsprünge lich zu erkennen. Auf den Sterniten ist die übli- grösser. Zwischen den Anterodorsalvorsprün- che Musterung in Form von Rechtecken gen liegt eine nicht allzugrosse skierotisierte Er- ausgebildet. Die Plättchen sind breit, ziemlich höhung mit zwei Larvalstigmenresten. Die flach und unterschiedlicher Grösse (Fig. 11c). Analvorsprünge sind lang, schlank, scharf zuge- Nahe der Distalränder sind sie gross und stehen spitzt und gehen von einem gemeinsamen recht dicht in Reihen, auf der übrigen Fläche sind Stamm aus. Die Spitzen aller Vorsprünge sind nur einzelne Plättchen anzutreffen. Am Distal- schwarz pigmentiert. rand des vorletzten Abdominalsegments stehen grosse, starke, dicht nebeneinander ansetzende Phylidorea (Euphylidorea) fulvonervosa Dornen; ein kleineren Dorn auch imm Distaiteli (Schumm.) jedes Pleurits. Limnobia fulvonervosa Schummel, 1829: 164. Liriinophila fulvonervosa (Schumm.): de Meijere, Analsegment (5 (Fig. 12c) ist lang und schlank. 1921:82. Lateralvorsprünge fehlen. Die Dorsalvor-

<^C^#^l^^^^^V

Fig. 12. Analsegment $ der Phylidorea-Puppen. Dorsalansicht, a, Ph. (Ph.) nigricollis; b, PH. (Ph.) squalens; c, Ph. (E.) nigronotata. 46 Tijdschrift voor Entomologie, deel 130, 1987

Limnophila {Phylidorea) fulvonervosa (Schumm.); Basis des drittes Zahns und dicht unter dem fünf- Stary, 1970: 145. ten Zahn. Auf der Aussenfläche der Mandibel, Phylidorea {Paraphylidorea) fulvonervosa nahe an ihrer Basis, inseriren zwei schwanke, (Schumm.); Savtshenko 1986b: 290. unterschiedlich lange Borsten. Praeimaginalstadien: Maxillen (Fig. 7d): etwa 480 |im lang und 150 Limnophila (Phylidorea) fulvonervosa (Schumm.); Brindie & Bryce, I960: 217, Fig. 13 (Analsegment |am breit. Im Apikaiteil ein, zwei oder drei keil- und Antenna); Brindie, 1967: 199, Fig. 46 förmige Sklerotisierungen, die den unregelmäs- (Analsegment). sig geformten, membranösen Teil verstärken. Verbreitung: Europa (Stary 1970; Savtshenko Hypopharynx (Fig. 8e, h): Querbrücke 1986 b), 35,3 —63,7 ^m lang, 130,9— 148,5 |im breit; La- teralarm 245,8—270,8 |im lang, 134,3—150,2 |im breit. Die Querbrücke ist unregelmässig Material: 2 (5, 1 $. Lubrzanka-Fiuss, Zagnahsk- ge- Jaworze: 1 Larve: 12. xi. 1981 gesammelt, 08. xii. 1981, formt, in der Mitte mit einer tiefen, schwach (5 verpuppt; 1 Larve: 23. iv. 1983 gesammelt, skierotisierten Vertiefung. Der Lateralarm ist 04.V.1983, 9 geschlüpft; 1 Larve: 23. iv. 1983 gesam- platt, im Mittelteil stark skierotisiert, verstärkt melt, 09.V.1983, verpuppt. S durch parallele Leisten. Die Höhe der Labialpal- 15 Lärvenlänge des 4. Stadiums — 20 mm. Kör- pen beträgt 41,5 —49,5 l-im, ihre Entfernung von- perbehaarung sehr zart, tast unsichtbar. Ventral- einander 56,9—64,3 l^m. Die Höhe der Papillen lappen lang, mit langer Randbehaarung und beträgt 10,2 — 11,2 jjm. breit gestreuten Pigmentstreifen. Die Laterallap- Hypostomium nicht entwickelt. pen sind kurz, etwa halb so lang wie die Ventral-

lappen. Die Puppe ist hellbraun. Die Länge einer weib- Die Kopfkapsellänge beträgt etwa 1,6 mm. lichen Puppe beträgt etwa 17 mm. Die Mund- Antennen (Fig. 3d): Basalglied 52,9— 55,2 |im gliedmassenscheiden sind in Fig. 9d dargestellt. lang und 25,6—29,6 |im breit; Apikaiglied Die Labralscheide ist breit 344 |im, fast quadra- 33,6—37,6 |_im lang und 5,1 —9,1 l^m breit, deut- tisch mit sanft gerundeten Ecken und etwas unter lich gestreift. Auf der distalen Fläche des Basal- die Labialtasterbasis gezogen. Die Maxillentas- glieds, nahe am Anzatz des Apikaiglieds, stehen terscheiden (Länge: etwa 630 |im) sind an der eine Papille und eine lange, steife Borste (Länge Basis eng, im Mittelteil deutlich verbreitert und 65,4 —68,3 |im). An der Basalgliedbasis ist ein zum Distaiteli hin sanft zugespitzt. Die Porenring auf der Innenfläche der Antenne zu Labialtasterscheiden (Länge: etwa 190 (im) wir- erkennen. ken untersetzt; ihr Ansatz am Labium ist sehr Clypeolabrum (Fig. 4d): 158,2 — 194,6 \im breit, ihr Apikaiteil hat eine charakteristische lang, 245,2 — 261,2 jim breit, trapezeförmig. Gestalt — er gleicht einem Fuss in Seitenansicht. "Flügelchen" fehlen. Am Vorderrand (Fig. 5g) in Pronotalhörner (Fig. lOd) etwa 600 \xn\ lang, der Mitte eine stark skierotisierte Erhöhung, kelchförmig, an der Basis rund, im Apikaiteil symmetrisch zu den Seiten hin je eine lange, abgeflacht. Die Atmungsspalte ist sehr breit, schwanke Borste, je eine spitze Dorne und je halbkreisförmig, Atmungskammer und Tra- zwei flache Papillen: eine spitzlanzettförmige cheenstamm scheinen durch. und eine fingerförmige. Etwas weiter hinten la- Die Flügelscheiden reichen bis zum Ende des

teral je zwei Sinnesgruben. 2. Abdominalsegments, die Beinscheiden fast

Epipharynx (Fig. 5h) mit zwei Paar zweiglie- bist zum Ende des 3. Segments. deriger Papillen: die Papillen des ersten Paares Die abdominalen Tergite und Sternite sind mit sind grösser als die des zweiten. Lateral stark langen, spitzen, markant breit ansetzenden Dor- skierotisierte Erhöhungen mit kurzer Behaa- nen bedeckt (Fig. lld), die eine regelmässige rung. Am inneren Rand beider Erhöhungen eine Rechteck-Musterung bilden. Am dichtesten ste- kurze, steife Borste. hen die Dorne im capitad weisenden Teil jedes Mandibeln (Fig. 6d, h): 290,2—352,8 jim lang, Rechtecks. Lateral, an Tergiten und Sterniten 128,0— 143,4 )im breit. Die helle Klinge am In- entlang, ziehen sich unregelmässige, längliche nenrand des Apikaizahns ist ziemlich kurz; sie dunkelbraune Flecke hin erreicht nicht die Hälfte der Apikaizahnlänge. Die Abdominalspitzen der männlichen Pup- Zweiter und dritter Zahn sind blattförmig (bei penexuvien sind bei der Aufzucht leider verlo- den einzelnen Exemplaren unterschiedlich aus- rengegangen. geprägt). Die Kanalmündungen liegen an der WiEDENSKA: Larven und Puppen Phylidorea-Arten 47

Literaturverzeichnis sche, vorwiegend holländische, Limnobiiden, ins-

Alexander, C. P., 1972. New subgenera of North besondere über ihre Kopulationsorgane. — Tijd- American crane flies (Tipulidae: Diptera). — schrift voor Entomologie 64; 54— 118. Entomological News 83: 29— 37. Mendl, H., 1978. Limoniidae. In: Limnofauna Euro- (lilies Beling, T., 1878. Zweiter Beitrag zur Naturgeschichte paea J. Ed.): 367 — 377. — Stuttgart. (Metamorphose) verschiedener Arten aus der Fa- Rozkosny, R., & P. Pokorny., 1980. 3. celed Bahnomil- milie der Tipuliden. — Verhandlungen der Zoolo- koviti — Limoniidae. In: Klic vodni'ch larev hmyzu gisch-Botanischen Gesellschaft in Wien 28: (Rozkosny R. Ed.): 245—257. — Praha. 21—56. Savtshenko, E. N., 1986 a. Palearctic Limoniid Flies Beling, T., 1886. Dritter Beitrag zur Naturgeschichte of "nigronotata" group of the genus Phylidorea (Metamorphose) verschiedener Arten aus der Fa- (Diptera, Limoniidae). — Vestnik Zoologii 5: milie der Tipuliden. — Verhandlungen der Zoolo- 20—26, gisch-Botanischen Gesellschaft in Wien 36: Savtshenko, 1986 b. Komary-limoniidy (obscaja Cha-

171—214. rakteristika, podsemejstva pediciiny i geksato- Brindle, A., 1958. Notes on the identification of miny). — Fauna Ukrainy 14: 1 — 380. — Kiev. Limnophila larvae (Diptera — Tipulidae). — Schummel, T. E., 1829. Beschreibung der, in Schlesien Transactions of the Society for British Entomology einheimischen, Arten einiger Dipteren Gattun- 13: 58—68. gen. I. Limnobia. Meigen. — Beiträge zur Entomo- Brindle, A., 1967. The larvae and pupae of the British logie 11: 97—201. Siebke, H., Cylindrotominae and Limoniinae (Diptera, Tipu- 1870. Beretning om en i Sommeren 1869 lidae). — Transactions of the Society for British foretagen entomologisk Reise gjennem Ringerike, Entomology 17: 151 — 216. Hallingdal og Valders. — Nyt Magazin for Natur- Brindle, A., & D. Bryce, I960. The larvae of the British videnskaberne 17: 246— 314. Stary, Hexatomini (Dipt., Tipulidae). — Entomologist's J., 1970. Revision der Arten der Unterfamilie Gazette 11: 207—224. Limoniinae (Tipulidae, Diptera) aus den Samm- Hennig, W., 1968. Die Larvenformen der Dipteren. lungen des Mährischen Museums in Brno mit be- Teil 2: 1—458. — Berlin. sonderer Berücksichtigung der Fauna Mährens. IL Levy, L., 1918. Contributions à l'étude des métamor- Tribus Hexatomini und Eriopterini. — Acta Musei phoses aquatiques des Diptères. — Annales de moraviensis 55: 133 — 194. Stary, 1981. biologie lacustre 9: 201 — 248. J., Nachträge und Berichtigungen zur Lundström, C, 1912. Beiträge zur Kenntnis der Dip- Limoniiden-Fauna der Tsechchoslowakei (Dip- teren Finnlands. VIIL Suppl. 2. Mycetophilidae, tera). IL — Acta Rerum naturalium Musei Slove- Tipulidae, Cylindrotomidae und Limnobiidae. — nia 27: 99—122. Wiedeiiska,!., Acta Societatis pro fauna et flora fennica 36: 1 — 70. 1986. Sygaczowate (Diptera, Limonii- Meigen,J. W., 1830. Systematische Beschreibung der dae) Gor Swiçtokrzyskich. Cz. I. Limoniidae doliny bekannten europäischen zweiflügeligen Insecten Lubrzanki. — Fragmenta Faunistica 30: 99— 120. 6: 1 —405. — Schulzische Buchhandlung, Hamm. Zetterstedt.J. W., 1838. Insecta Lapponica: 1 — 1139. — Lipsiae. De Meijere, J. C H., 1916. Beiträge zur Kenntnis der Dipterenlarven und -puppen. — Zoologische Jahrbücher, Abteilung für Systematik, Geographie Anschrift des Verfassers: Dr Jolanta Wiedenska, und Biologie der Tiere 40: 177 — 322. Instytut Biologii Srodowiskowej, Uniwersytet Lódzki, 90—237 Lodz, ul. Banacha 12/16, Polen. De Meijere, J. C. H., 1921. Studien über palaearkti-

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Tijdschrift voor Entomologie 130; 49— 108 Gepubliceerd 30 november 1987

A REVISION OF WESTERN PALAEARCTIC OXYTORINE GENERA. PART VI. (HYMENOPTERA, ICHNEUMONIDAE)

G. VAN ROSSEM

Berkenlaan 25, Ede, The Netherlands

Introduction sem, 1980, is preoccupied. It is proposed to re-

A taxonomie guide and a key to the genera of place it by Phosphoriana nomen novum. the Oxytorinae was given by Townes (1971). A A new species is introduced in the genus generic key to the Palaearctic Oxytorinae was ProeliatorVzn Rossem: P. invictus. pubHshed by Van Rossem (1982). Almost all the Of the genera Oxytorus, Cyllocena and Meg- type-species of the genera are specified by astylus type material has become available, Townes (1971). This part of the revision of which allowed me to make some remarks. western European Oxytorinae includes a survey of the genus Hemiphanes Förster together with Materials and methods the record of three new species, viz., H. hor- All observations were made with a Zeiss bin- tense, H. inusitatum and H. montanum. ocular compound microscope. The length of the

The genus Apoclima Förster is re-introduced front wing was measured with the ocular micro- with the description of one new species, A. hae- meter at 10 X enlargement. The length of the selbarthi. ovipositor was taken from the apex of the gas-

One new species of Pantisarthrus Förster is ter. described, P. gracilis. For the terms used, see Townes (1969: 36 The Förster (1871) and Thomson (1888) type material of the genus Plectisadea Viereck was studied. Two new subgenera, Plectisadea and Acknowledgements

Fugatrix, are introduced and five new species, The author is indebted to Dr E. J. Fittkau, P. indomita, P. foersteri, P. substantiva, P. blan- Director of the Zoologische Staatssammlung at dita and P. ventosa are proposed. Plectisadea München, for permission to study the Förster nemorensis is a nomen novum for Ephalmator collection over a period of seven years. I thank subsimilis Van Rossem. The Plectisadea species Mr Erich Diller for his help and hospitality dur- described from males by Förster i.e. are re- ing my visits to the München Museum.

garded as species inquirendae, for at present it is I am grateful to Dr Henry Townes (Gaines- impossible to find the matching females. ville, Florida) for information, the loan of speci- The name Gnathochorisis Förster, 1869, takes mens and continuous concern with respect to priority over Laepserus Förster, 1869. Gnatho- my work. chorisis crassulus (Thomson), that was presumed For the loan of type material it is a pleasure to (Van Rossem, 1980) to be a colour from of thank: Dr R. Danielsson (Universitetets Zoo-

Gnathochorisis dentifer (Thomson), is now con- logiska Institutionen, Lund); Dr M. Kak (Mu- sidered to be a separate species. zeum Przyrodnicze, Wrociaw); Dr B. Petersen

A new approach to the genus Eustennx (Zoologisk Museum, Kabenhavn); Dr J. P. Förster is presented with the recognition of six O'Connor (National Museum of Ireland, Dub-

new subgenera. lin); Dr K. J. Hedqvist (Naturhistoriska Of the genus Helictes Haliday a revision of Riksmuseet, Stockholm); Dr habil. G. Morge the type material, males only, is published, in- (t) (Curator of Strobl Collection at Admont, cluding two newly described species, H. incon- Eberswalde-Finow, DDR); Dr M. G. Fitton gruens and H. fabularis. At present the recogni- (British Museum (Natural History), London;

tion of the females is not possible. Dr E. Königsmann (t) (Zoologisches Museum The name of the genus Phosphorus Van Ros- Humboldt Universität, Berlin, DDR); Dr J.

49 — —

50 Tijdschrift voor Entomologie, deel 130, 1987

Oehlke (Institut für Pflanzenschutzforschung Key to Hemiphanes females Kleinmachow, Eberswalde-Finow, DDR); Mr (The females of //. gravator, H. hortense and K. W. R. Zwart (Laboratorium voor Entomolo- //. inusitatum are unknown) gie, Wageningen); Dr D. R. Kasparyan (Zoo- of Sciences, Lenin- logical Institute, Academy 1. Apical margin of clypeus with a deep medi- grad); Dr K. Horstmann (Zoologisches Institut an semicircular notch. Upper and lower III der Universität, Würzburg). teeth of mandible equal in length Lastly I am very much indebted to Dr E. //. montanum spec. nov. Haeselbarth, München, and Mr C. J. Zwakhals, — Apical margin of clypeus only weakly in- Arkel, for their contribution of numerous speci- dented. Lower tooth of mandible shorter mens of Oxytorinae, frequently undescribed than upper 2 locali- species, or specimens from remarkable 2. Lateral lobes of mesoscutum with close ad- ties. pressed hairs. Gaster more fuscous, with I wish to thank my friend Dr C. G. Johnson, yellov/ish spots medially on tergites two Harpenden (U.K.) for reading the manuscript and three //. townesi Van Rossem and improving the English. — Lateral lobes of mesoscutum hairless except The author owes thanks to the Uyttenboo- for some hairs proximally and laterally. The gaart-Eliasen Stichting at Amsterdam for grants gaster from tergite three yellow to travel to München. //. flavipes Förster

Hemiphanes flavipes Förster

Genus Hemiphanes Förster Hemiphanes flavipes Förster, 1871 : 101. Hemiphanes Van Rossem, 1980: 86. Hemiphanes Vörster, 1871: 101 — 102. flavipes;

Hemiphanes; Townes, 1971 : 184— 185. Hemiphanes; Van Rossem, 1980: 85—88. Characteristics of the male: Front wing 4.5 5.3 mm. Mandible yellow, lower tooth shorter The species described below include three than the upper. Clypeus yellow, margin trun- hitherto unknown. I regret that I have only one cate. Anterior tentorial pits open. Antenna long, specimen of each, two of them in rather poor reaching length of body, scape rather swollen, condition. I consider the gender of Hemiphanes pedicel large, yellow. Tyloids on flagellar seg- to be neuter. ments 9— 11. Postannellus 5.2—6.0 times as long as apical width. Apical corners of prothorax and tegulae yellow. Front parts of notauli Key to Hemiphanes males deep. Median lobe of mesoscutum with ad- (The male of//, montanum is unknown) pressed hairs, lateral lobes polished. Epomia present. Apex of scutellum with some rough

\. Hind aspect of head deeply concave 2 sculpture. Propodeum with apical transverse ca- — Hind aspect of head not concave 3 rina strong. Basal transverse carina absent. Me- 2. Tyloids on flagellar segments 9— 12. Occip- dian longitudinal carinae present. Front wing

ital carina interrupted. Postpetiole striate . . . without areolet. Nervelus placed basally of bas- H. gravator Förster al vein. Nervellus about vertical, intercepted — No tyloids present. Occipital carina not low, discoidella present. Legs, including coxae, present. First tergite with different longi- yellow, hind coxae comparatively large. Hind tudinal sculpture ...//. hortense spec. nov. femur 5.0—5.7 times as long as wide in the mid- 3. Postanellus conspicuously stout, 3.0 times dle, rather short and slender. First gastral seg- as long as apical width. No tyloids present ment wide towards apex, end of first sternite //. musitatum spec. nov. and spiracles situated in the middle. — Postanellus not stout, 5.0— 7.6 times as Characteristics of the female: Front wing long as apical width 4 5.4—6.0 mm. Clypeus 2.0 times as wide as long. 4. No tyloids present. Nervulus opposite Postanellus 8.0 times as long as apical width. basal vein //. townesi Van Rossem Gena, temple and vertex polished. Occipital ca- — Tyloids on flagellar segments 9— 10 (or 9 rina closed. Pronotum for the greater part pol- 11), running over the whole length of the ished, epomia weak, with some parallel running segment. Nervulus slightly basally of basal ridges. Notauli running beyond centre of meso- vein //. flavipes Förster scutum. Mesopleurum poHshed, prepectal cari- —

Van Rossem: Western Palaearctic Oxytorinae 51

na reaching subtegular ridge. Lateral parts of ished. Frons not concave. Pronotum polished, first tergile with longitudinal striation and with epomia present. Mesoscutum with close ad- some continuation on tergite two. Following pressed hairs. Scutellum punctured, apex with tergites polished. Ovipositor slightly protruding rough sculpture. Propodeum with irregular beyond tip of gaster. Gaster rather depressed sculpture, the median longitudinal and lateral towards apical part. Otherwise as the male. longitudinal carinae proximally obliterated. Ap- ical transverse carina strong, medially devel- Distribution. — Germany. Sweden up to Lapland oped into a keel, lying near to apex of propo- (VanRossem, 1980). deum. The minute propodeal spiracles circular. Dorsal rim of metanotum with a triangular pro- Hemiphanes gravator Forster jection, lying opposite the front end of lateral Hemiphanes gravator Förster, 1871: 102. longitudinal carina. Mesopleurum polished, Hemiphanes gravator; Van Rossem, 1980: 86. prepectal carina present. Front wing without areolet. Nervulus lying shghtly distally of basal The female is unknown. vein. Nervellus intercepted low, the discoidella Characteristics of the male: Front wing 4.0 running almost to wing margin. Coxae and legs 5.0 mm. Mandible yellow, the lower tooth whitish to light brown. First tergite with rough shorter and narrower than upper. Apical margin sculpture, spiracles at 0.40 of the length, end of of clypeus yellow, truncate. Anterior tentorial first sternite at 0.30 of the length of the seg- pits open. Face rather protuberant below the ment. Front half of second tergite with longi- antennal base (more than in H. flavipes). Hind tudinal striation and coriaceous sculpture. Api- aspect of head deeply concave. Occipital carina cal half with weak striation and a median yellow interrupted. Scape ventrally yellow. Postannel- brown spot. Following tergites more polished. lus 5.0—6.0 times as long as apical width. Ty- Third and fourth tergite with yellow brown loids situated on flagellar segments 9— 12. No- area. Apical part of gaster depressed. tauli present. Scutellum flat, closely punctured. Characteristics of the female: Front wing Wing without areolet. Nervulus somewhat dis- about 4.5 mm. Otherwise as the male. First ter- tally of basal vein. Legs, including coxae, yel- gite with close longgitudinal striation. Oviposi- low. Hind femur 4.7—6.0 times as long as wide. tor not protruding beyond tip of gaster. Propodeum with only apical transverse carina present, more smoothly sculptured than in H. Distribution. — China, Shaowu Hsien, 1200— 1500 flavipes. First gastral segment wide towards m altitude (coll. Townes). apex, postpetiole striate, with yellow spot. Ster- nite in front of the middle. Second tergite Hemiphanes hortense species nova striate, with brown spots laterally. Characteristics of the holotype {â , Nether- lands, Asperen (Prov. Zuid-Holland), 8. vi. 1973, Distribution. — Austria. Germany. Sweden (Van leg. and coll. C. J. Zwakhals): Front wing 2.8 Rossem, 1980). mm. Head fuscous. Mandible yellowish brown, the lower tooth slightly the longer. Clypeus flat, Hemiphanes townesi Van Rossem rather protruding, the margin truncate. Clypeus Hemiphanes townesi Van Rossem, 1980: 86— 87. and face, frons, vertex and gena coriaceous. Hind aspect of head deeply concave (seen from Characteristics of the male: Front wing about dorsal side V-shaped), occipital carina absent, 5.8 mm. Mandible whitish, teeth brown. The lower part of genal carina present. Postannellus lower tooth about 0.5 times shorter than upper. as long as second flagellar segment, slender. No Clypeus about 2.4 times as wide as long, whit- tyloids present. Epomia absent. Notauli strong ish to brownish, convex, margin truncate. No to about middle of mesoscutum. Mesoscutum, groove between clypeus and face. Face below scutellum, and propodeum coriaceous. Propo- antennal sockets closely punctured, with ad- deum with only apical transverse carina and pressed long, silvery hairs. Malar space wide, pleural carina present. Mesopleurum coria- about 1.5 times the width of mandibular base, ceous, prepectal carina weak but reaching the polished. Scape globular, pedicel large, whitish margin. Nervellus intercepted about in the mid- to brownish. Postannellus slender, about 8.0 dle, discoidella very weak. Legs including mid- times as long as apical width. Antenna without dle and hind coxae, yellowish brown, long and tyloids. Frons, vertex, temple and gena pol- very slender, especially the hind tibia. First ter- 52 Tijdschrift voor Entomologie, deel 130, 1987

gite with conspicuous longitudinal sculpture. stout, middle and hind femur more slender. Left Second tergite with weak longitudinal sculp- middle leg missing beyond femur. In the speci- ture. Gaster fuscous. men the right hind leg missing beyond coxa; also tarsi of left hind leg missing beyond basitar- Material examined. — The holotype only. sus. First tergite coriaceous, rather wide apically, about L6 times as long as apical width. A Hemiphanes inusitatum species nova conspicuous pit at base of petiole between the Characteristics of the holotype (a, Italy, median dorsal carinae. Following tergites coria-

Prov. Bolzano, Sarntal, 1250 m, 1. vi. 1977, leg. ceous, with a yellow brown band medially from and coll. C. J. Zwakhals): Front wing 3.7 mm. apical margin of tergite two. Ovipositor not Mandible (teeth reddish), clypeus and face yel- projecting beyond tip of gaster. low. Clypeus not protruding, short, about 3.6 times as wide as long. Hind aspect of head not Material examined. — The holotype only. concave. Occipital carina present and closed. Postannellus stout, 3 times as long as apical Genus Oxytorus Förster width. No tyloids present. Antenna light Oxytorns Förster, 1868: 199. brown. Epomia absent. Mesoscutum coria- Oxytorus; Townes, 1971 : 1 85. ceous, notauli absent. Propodeum coriaceous, Oxytorus; Van Rossem, 1980: 88. no carinae present except for stubs of median longitudinal carinae. Lower part of mesopleu- Type-species: Oxytorus armatus Thomson, rum coriaceous, upper part polished. Prepectal 1883. carina almost obsolete. Front and middle coxae white, hind coxae fuscous. Legs yellowish Oxytorus luridator (Gravenhorst) brown. First tergite L8 times as long as apical Ichneumon luridator Gravenhorst, 1820: 379. width, coriaceous. Gaster fuscous, second ter- Oxytorus luridator; Van Rossem, 1980: 90. gite with brownish apical band. Atractodes properator HaMij, 1838: 120. Oxytorus luridator; Fitton, 1976: 332. Material examined. — The holotype only. Characteristics of the lectotype of Atractodes

The specific name Is from the Latin "inusita- properator Hahday. Labels: a printed label tus", meaning "uncommon" or "unusual". "England?"; a label "Haliday, 20.2.82"; a label "named by Claude Morley Atractodes propera- Hemiphanes montanum species nova tor Hal, Type, vi. 193"; a circular label with red Characteristics of the holotype (9, Austria, margin "Type CM"; a small label illegible; lec- T., Niederthai Gubener Hütte, 2000 m, totype label Fitton, 1975. National Museum of

1. ix. 1979, leg. and coli. Haeselbarth): Front Ireland, Dublin. Female. The specimen repre- wing 4.7 mm. Mandible yellowish, teeth brown, sents Oxytorus luridator (Gravenhorst). of the same length. Clypeus yellow, the apical part depressed and with a deep median semicir- Oxytorus luridator (Grav.) cular indentation, the side parts of which stand forma nigricoxa Kiss von Zilah out flap-like. Malar space as wide as base of Callidiotes luridator (Grav.) forma nigricoxa Kiss von mandible. Head entirely coriaceous, excepting Zilah, 1924: 118. the clypeus. Flagellum yellowish, postannellus slender, 7.0 times as long as apical width. Epo- The two syntypes (Transylvania, Borosjenö, mia absent. Mesoscutum coriaceous, notauli leg. Diöszeghy) are labelled as males, but both present towards margin, shallow. Scutellum and are females. They show fuscous coxae which is propodeum coriaceous. Pleural carina present. in fact a male character. The specimens are kept Median longitudinal carina present only to- in the Természettudomânyi Mûzeum, Buda- wards base of propodeum. Mesopleurum coria- pest. I thank the curator Dr J. Papp for sending ceous and weakly striated. Prepectal carina them to me. strong, reaching the margin. Nervulus distad of basal vein. Nervellus not intercepted. Discoidel- Note. — In the description of O. luridator la absent. All coxae fuscous. Second trochanter (Van Rossem I.e.) there is a wrong description of all legs pale in colour. Front and middle legs of the mesopleural sculpture: "transverse" brownish, hind femur fuscous. Front femur ridges should be "longitudinal" ridges. Van Rossem; Western Palaearctic Oxytorinae 53

bristles (fig. 1) A. signaticorne Förster — Third flagellar segment notched on apical half (the tyloid), the base of fourth flagellar segment scarcely notched. The third flagellar segment with a strong tooth below the tyloid at the apex of which two minute hairs (fig. 2) A. haeselbarthi spec. nov.

Apoclima haeselbarthi species nova Characteristics of the holotype of Apoclima

haeselbarthi {S , Germany, Bayern, Oberam- mergau, Laber, 1400— 1600 m, 5. ix. 1980, leg. and coll. Haeselbarth (München)): Front wing 3.5 mm. Palpi brown. Lower margin of mandible turned inwards. Clypeus convex, coriaceous as malar space, face and vertex. Malar space and gena wide. Occipital carina widely interrupted medially. Occiput not so strongly curved inwards as in A. signaticorne. Antenna brown, including scape and pedicel tips missing in specimen. Pedicel large. Third flagellar segment notched on apical half (the tyloid). Base of ^W' fourth flagellar segment very weakly notched. The third flagellar segment with a strong tooth below the tyloid, at the apex with two minute hairs. Epomia present. Mesoscutum coriaceous, with notauli vaguely meeting in centre. Scutel-

Figs. 1, 2. Base of male entenna. 1, Apoclima signati- lum with weak carina running towards apex. corne Förster, left antenna (right one broken); 2, A. Propodeum with irregular sculpture, apical haeselbartht spec, nov., right antenna. Enlargement, transverse present, other carinae not strongly caSOX. developed. Mesopleurum polished. Prepectal carina rather underdeveloped, not reaching margin. Front wing without areolet, with two bullae. Portion of cubitus between intercubitus Genus Apoclima Förster and second recurrent vein 0.3 of the length of recurrent vein. Nervellus inclivous. Discoidella Apoclima Förster, 1871 : 97—98.

Apoclima; Townes, 1971 : 191—192. present. Legs brownish. Claws of front legs and Apoclima; Van Rossem, 1980: 97—98. left hind leg missing beyond femur in the specimen. Femora stout. All coxae fuscous and with

In Dr E. Haeselbarth's material I found an long erect hairs. First tergite coriaceous, about

Apoclima male which I compared with the holo- 1.1 times as long as apical width, glymma pre- type of Förster's Apoclima signaticorne . The lat- sent, median dorsal carina strong to about 0.75 ter has a weak projection on segment three of of length. Dorsolateral carina strong, the spira- the flagellum that merely delineates the notch of cles protruding. The dorsal profile of first ter- the segment (fig. 1). In Haeselbarth's specimen, gile strongly convex in the middle. Apex of first which undoubtedly is another species, the pro- sternite in front of the middle. Second tergite jection is strong and toothlike (fig. 2). for the greater part coriaceous. The entire gaster fuscous, except for vague brownish apical mar- Key to Apoclima males gin of tergite two. All tergites, except the first, 1. Third flagellar segment notched on apical with conspicuous suberect hairs. half, the base of fourth flagellar segment Female unknown. somewhat notched (the tyloids). The third flagellar segment with a weak projection below the tyloid at the apex of which two Material examined. — The holotype only. —

54 Tijdschrift voor Entomologie, deel 130, 1987

I name this species after Erasmus Haeselbarth Key to females

(München) who contributed his extensive Oxy- (The female of C. suerinensis (Brauns) is un- torine material. known)

Apoclima signaticorne Förster 1. Frons with rough sculpture. Nervellus in- 97 Apoclima signaticorne Förster, 1871 : —98. tercepted below the middle, upper part in- 97 Apoclima signaticorne ; Van Rossem, 1980: —98. clivous, lower part reclivous. Dorsolateral carina of the first tergite running to the spi- Characteristics of the male. Length front racles. Second to fourth tergite orange, pol- wing 3.2 mm. The third flagellar segment ished. Basal half of second tergite with notched on apical half, the base of fourth flagel- some rough longitudinal wrinkling lateral- lar segment somewhat notched (the tyloids). ly. Ovipositor about the length of hind tibia The third flagellar segment with a weak projec- C. langei (Brauns) tion below the tyloid, at the apex of which there — Frons polished or with fine sculpture or are two bristles (fig. 1). Pedicel large. Postannel- with regular longitudinal striation. Combi- lus 3.7 times as long as wide. Postocciput con- nation of characters not as above 2 cave, occipital carina interrupted. First tergite 2. The last joint of flagellum conspicuously wide and short. Apex of scernite and spiracles m swollen. Second tergite with fine longitudi- front of the middle. Glymma present. Median nal striation. Ovipositor 0.34 of length of dorsal carina not reaching the spiracles. front wing C. borealis (Roman) Characteristics of the female. Length of front — The last joint of flagellum not swollen ... 3 wing 3.4 mm. Pedicel large. Flagellum slender, 3. Mandible and clypeus yellow. Head and postanellus 4.0 times as long as wide. Postocci- mesoscutum pohshed, without punctures. put rather concave. Occipital carina widely in- Hind femur fuscous. Nervellus intercepted terrupted. Portion of cubitus between intercubi- deeply below the middle. Ovipositor about tus and second recurrent vein 0.36 of the length 0.46 of length of front wing of recurrent vein. Nervellus vertical, intercepted C. alpigena (Strobl) below the middle, front part of discoidella pre- — Clypeus not yellow. Hind femur orange or sent. First tergite L4 times as long as apical yellow 4 width, coriaceous. Glymma weak. Median dor- 4. All coxae and femora orange 5 sal carina not reaching the spiracles, the latter at — Alle coxae fuscous 6 0.32 of the length. Basal half of second tergite 5. Mesopleurum polished, widely and finely coriaceous. Ovipositor about the length of hind punctured. Some striation may be present tibia beyond tip of gaster, apex somewhat up- in the lower hind corner below the specu- curved, no dorsal notch present, the tip acumi- lum and some below the subtegular ridge. nate. Frons polished or with indistinct coriaceous sculpture. Ovipositor 0.50—0.67 of length Distribution. — A very rare species. In total four of front wing specimens are registered; the male holotype is from C. caligata (Gravenhorst) Aachen (coll. Forster), a female from Blankenburg — Mesopleurum with close longitudinal stria- (Thüringen) (coll. Schmiedeknecht) and a male and fe- tion. Frons with regular longitudinal stria- male from Wiesen (Spessart) (Germany) (both coll. tion. Ovipositor 0.85 of length of front Flaeselbarth). wing C. invicta spec. nov.

6. Apical half of second tergite and all follow-

Genus Cylloceris Schiodte ing tergites orange . C. fusciventns{¥\.Q\\én) Cylloceria Schiodte, 1838: 140. — Gaster fuscous 7 Cylloceria; Townes, 1971: 192. 7. Postannellus long, 9.6 times as long as api- Cylloceria; Van Rossem, 1980: 98—107. cally wide. Ovipositor 0.9 LO as long as the front wing 8 One undescribed species (Cylloceria invicta) — Postannellus shorter, 7.0—8.8 times as long is inserted. The lectotype of Gravenhorst's Try- as apically wide. Ovipositor 0.6— 1.1 of the phon sylvestris was brought to light. It proved length of front wing 9

to be the same as Cylloceria stnolata (Hellen). 8. First and second tergite with very rough The type specimen of Lissonota occupator sculpture. Basal margin of third and fourth Gravenhorst, 1829, is lost. tergite with a rough band of sculpture. Van Rossem: Western Palaearctic Oxytorinae 55

Mesopleurum polished. Apical margins of phasized tooth-like. The basal part of seg- tergites black. Postanellus long, 9.0 times as ment four with a weak notch 6 long as apically wide. Ovipositor 0.9 of 6. Nervellus intercepted süghtly below the length of front wing middle. Second and third tergite with some C. imperspicua spec. nov. longitudinal sculpture. Gena polished, — First tergite with regular and rather fine widely punctured sculpture. Second tergite finely striated. C. sylvestris (Gravenhorst) Third tergite also showing striation. Meso- — Nervellus intercepted over the middle. Sec- pleurum polished. Postanellus 9.6 times as ond, third and fourth tergites without lon- long as apically wide. Ovipositor 0.9— 1.0 gitudinal wrinkling. The base of the second of the length of front wing and third tergites with weak coriaceous C. sylvestris (Grav.) sculpture. The apical margins of tergite two 9. Postannellus 8.0—8.8 times as long as api- to five with brownish to orange colour. cally wide. Second tergite with fine sculp- Gena polished, punctures absent ture, the base finely striated. Nervellus in- C. melancholica (Gravenhorst) tercepted slightly below, or in the middle. Ovipositor 0.9— 1.1 of the length of front Cylloceria borealis (Roman) wing C. sylvestris (Grav.) Lampronota borealis Roman, 1925, Arkiv for Zoologi — Postannellus 7.0 times as long as apically 17A(4):20—21. wide. Second tergite coriaceous. Nervellus intercepted slightly over the middle. Ovi- Characteristics of the lectotype of C. borealis. positor about 0.6 of the length of front Female. Front wing 6.4 mm long. Palpi brown. wing C. melancholica (Gravenhorst) Mandible and clypeus brown, the latter polished and 2.2 times as wide as long. Malar space wide, Key to males 0.33 of width face. Face, frons and vertex finely (Males of C. invicta spec, nov., C. alpigena sculptured. The last joint of flagellum conspicu- (Strobl) and C. imperspicua spec. nov. are un- ously swollen, a character not shown in other known). Cylloceria species. Postannellus 7.1 times as long as apically wide. Pronotum finely sculp- 1. Frons with rough sculpture. Second to tured with some wrinkling. Mesoscutum with fourth tergite orange, polished indistinct sculpture. Notaulus strong. Upper C. langei (Brauns) part of scutellum polished. Propodeum with — Frons polished or with fine sculpture .... 2 rough sculpture, the longitudinal carinae strong. 2. All coxaeand all femora including front and Mesopleurum for the greater part polished. Pre- middle tibiae orange pectus with sculpture. All coxae brown, other C. caligata (Gravenhorst) parts of legs orange-yellow to brown in colour. — All coxae brown or black 3 Claws strong. First tergite with rough, somew- 3. Flagellar segments four and five weakly hat striated sculpture. The first abdominal seg- notched. Third and following tergites pol- ment robust, with a broad apical edge. Second ished C. suerinensis (Brauns) tergite with fine longitudinal striation. Base of — Flagellar segments three and four notched third tergite with longitudinal wrinkling, the 4 further part polished. The apical tergites pol- 4. Gaster from tergite two reddish to orange ished. Ovipositor 0.34 of length of front wing. in colour. Second tergite coriaceous and Characteristics of the male. Front wing 6.4 with longitudinal wrinkling. The following mm long. Palpi yellow. The sculpture of the tergites for the greater part polished head corresponding to that of the female. Third C. fusciventris (Hellen) segment of the flagellum apically with a deep — Gaster fuscous 5 semicircular notch, emphasized tooth-like at 5. Third segment of flagellum apically with a base of notch. Basal part of segment four also deep semicircular notch. The base of the with a weak notch. Pronotum with irregular notch is emphasized tooth-like. The basal sculpture. Mesoscutum almost polished, notauli half of segment four with a weak notch. strong, meeting. Propodeum roughly sculp- Notauh meeting C. borealis (Roman) tured, the longitudinal carinae strong. Meso- — Third segment of flagellum apically with a pleurum polished. Prepectal carina bending to- semicircular notch, the base not em- wards the margin. Coxae brown, other parts of —

56 Tijdschrift voor Entomologie, deel 130, 1987 legs yellow in colour. Claws strong. First tergite regular, longitudinal striation. Occiput punc- with rough sculpture. Second and third tergite tured. Gena more polished, but punctured to- with longitudinal wrinkling. The broad apical wards genal carina. Antennal scape closely margin of the third tergite pohshed. The apical punctured. Postannellus 7.8 times as long as api- tergites polished. cally wide. Pronotum closely striated. Mesoscu- tum finely punctured, backwards and laterally Material examined. — Sweden: female holotype indistinct. Notaulus strong, particularly to- and male paratype, Ângermanland, leg. C. Stil (coll. wards base. Propodeum with regular and rough Stockholm). Roman, Naturhistoriska Riksmuseet, sculpture, the longitudinal carinae strong. Lectotype label of Townes. Mesopleurum with close, longitudinal striation.

(The mesopleurum in C. caligata is polished and Distribution. — According to Roman (I.e.), the punctured, which is a distinguishing character). species is found in northern Sweden, Finland and the U.S.S.R., Siberia (Jenissei region) and Kamchatskaya. Prepectal carina indistinct. Front coxae brown.

Three specimens, all females, were received on loan Middle and hind coxae and other parts of legs from the Leningrad Museum: 5 km N. Pusjkina, Le- orange, except for trochanters, hind tibia and ningrad, lesopoloça (forest area near Leningrad), leg. tarsus. Nervellus intercepted over the middle. D. R. Kasparyan, 14.viii.l980; Romanovka (east of First tergite with regular and rather subtle Baykal Lake), bliz Jamburga, 30.V.1905, leg. Bar- sculpture. Spiracles at 0.38 of the length of the ovskij; R. Nelgeche, berch. Verchojan, okr. 11 segment. Median dorsal carina developed to the 12.viii.l927, leg. Tkatsjenko. spiracle. Second and third tergite finely coriaceous. Ovipositor 0.85 of length of front wing. Cylloceria caligata (Gravenhorst, 1829)

Phytodietus caligatus Gravenhorst, 1829: 936. Material examined. — Female holotype from the Lampronota crenicornis Curtis, 1832: 407. Zoological Institute at Leningrad: U.S.S.R., Olenek, Cylloceria crenicornis; Fitton, 1976: 323. Yakutia, 67—689, viii.1874, leg. A. Czekanowski. It Characteristics of the female lectotype of Lampronota also bears a number: 74478. crenicornis. Labels; a separate label: Haliday 20-2- "Invictus" is the Latin for "irrefutable", "im- 82; lectotype label of Fitton (National Museum of movable". Ireland, Dublin). The specimen has the character-

istic orange coloured coxae. Chalinoceras mancus Ruthe, 1855: 82. Cylloceria imperspicua species nova Cylloceria manca; Fitton, 1978: 76. Characteristics of the holotype of C. imperspicua. Female. Front wing 8.22 mm long. Labels: Germany: Spandau (BMNH). The lec- Palpi brown. Basal half of mandible coriaceous totype is a specimen of Cylloceria caligata. and with strong punctures, apical part polished, This was also suggested by Fitton. teeth robust. Clypeus with vague coriaceous sculpture, 2.0 times as wide as long. Malar space Material examined. — Five specimens, all females, wide, 0.37 of width of face. Face towards malar were received on loan from the Leningrad Museum: space polished. Medially, below antennal sock- females, ? Sumucha), 28.vii.1896, leg. K. Ko- Sumuch ets a triangular protuberance with vertically kujeva (Western Siberia); Kurjat, 18.vi.l886, merig, wrinkled sculpture. Postannellus extremely leg. K. Jarosjevskago; Kurjat, 18.viii.l888, Na Zet long, 9.0 times as long as apically wide. Frons poyl, leg. K. Jarosjevskago; Kurjat, 18.vi.l889, merig strongly concave. Vertex with fine coriaceous A. offic, leg. K. Jarosjevskago. 1 i, Bologoye, Val- dajsk, v.vii.1907, leg. Zajtseez (400 km south of Le- sculpture. Gena wide, polished. Pronotum me- nmgrad). dially wrinkled. Mesoscutum with strong notauli, these not meeting. The median lobe in the Distribution. — The species occurs through-out the backward part with rough wrinkled sculpture. Palaearctic Region. The basal part of the median lobe punctured and the lateral lobes indistinctly punctured. Propo- Cylloceria invicta species nova deum with rough sculpture, the longitudinal ca- Characteristics of the holotype. Female. rinae present. Mesopleurum widely punctured Front wing 8.8 mm long. Palpi brown. Mandi- and for the greater part polished. Prepectal cari- ble brown, medially with a light brown spot. na strong, meeting the margin. A robust subte- Clypeus brown. Other parts of head black. Face gular ridge present. Nervellus intercepted over longitudinally striated. Malar space with irregu- the middle. Coxae fuscous and polished. Front lar sculpture, 0.34 of width face. Frons with and middle tibia and femur and the hind femur Van Rossem: Western Palaearctic Oxytorinae 57

orange in colour. The front and middle tarsus Characteristics of the lectotype of Tryphon brown, the hind tibia and tarsus black. The first sylvestris {6, a white tag, 69; a green tag; a abdominal segment robust, the apical edge 0.85 white tag handwritten conf (confer?) melancho- of the length of the segment. The first and sec- lica): Front wing 7.1 mm long. Mandible ond tergite showmg rough and irregular sculp- brown. Clypeus: the apical part polished, near ture. Third and fourth tergite basally with a basal margin somewhat sculptured. The apical rough band of sculpture, backward more coria- margin truncate. Face closely punctured, me- ceous. Gaster entirely black. Ovipositor 0.91 of dially somewhat convex. Frons polished. Vertex the length of front wing. and gena finely punctured (implantations of hairs). Apex of third flagellar segment with a Material examined. — Sweden: 9, holotype, Dalar- semicircular notch, base of fourth segment also na, Idre (Fjätervilen), 24. vii— l.viii.l982, Malaise with a notch. Pronotum with rather coarse trap, leg. & coll. Van Rossem. semicircular striation, epomia present. The median lobe of mesoscutum convex, closely punc- "Imperspicuus" is the Latin for "inscrutable". tured, the lateral lobes with fine punctures. Pro- podeum roughly sculptured, with all longitudi- Cylloceria sylvestris (Gravenhorst) nal carinae present. Mesopleurum with fine Tryphon sylvestris Gravenhorst, 1829: 138. punctures, the prepectal carina to the margin. Cylloceria sylvestris; Pfankuch, 1906: 87. Below the subtegular ridge some longitudinal Lampronota melancholica (Grav.) var. striolata striation. Nervellus intercepted below the mid- Hellen, 1915:48. dle. All trochanters fuscous. Lampronota striolata Hellen, 1937: 12. coxae and All fem- Cylloceria striolata; ]ussi\3i, 1965: 101. ora, front and middle tibiae yellow to orange. Cylloceria striolata; Van Rossem, 1980: 130—104. Hind tibia and tarsus conspicuously fuscous. First tergite roughly sculptured and somewhat Through the kindness of Dr M. Kak (Mu- longitudinally wrinkled. Second tergite coria- zeum Przyrodnicze, Wrociaw) I got the oppor- ceous and with some longitudinal striation. The tunity to study the type material of Tryphon syl- third tergite with indistinct coriaceous sculp- vestris Gravenhorst. It appears that the type ture. material in question consists of two male specimens. The one I labelled as the lectotype is From Dr R. Jussila (Paattinen, Finland) I re- identical with Cylloceria striolata (Hellen). ceived a female and male of Cylloceria striolata Dr Kak wrote that he was not quite sure that (Hellen) from Finnish Lapland. In a collection the specimen which I labelled as the type of Oxytorinae sent by Dr D. P. Kasparyan from

Tryphon sylvestris is a Gravenhorst specimen. the Zoological Institute at Leningrad there were However, the other specimen represents Cyllo- four dubious specimens of the same species. Al- ceria caligata, which species was also described though these specimens show rather striking by Gravenhorst (I.e., p. 138). variability, I decided to place the four Russian

Table 1 . Cylloceria sylvestris (Gravenhorst). For explanation, see text. specimen length length ratio length ratio width nervellus front ovipositor/ postannellus/ malar space/ wing length apical width face front wing width

Reservoir Vudjavr 20.viii.l930 — —

58 Tijdschrift voor Entomologie, deel 130, 1987

specimens tentatively with C. sylvestris. For Chalinoceras longicornis Ratzeburg, 1852: 130. Sensu

comparison table 1 gives: the length of the front Viereck, 1914. The type is lost. wing in mm; ratio of the length of the ovipositor to the length of the front wing; ratio of the Cylloceria melancholica forma length of postannellus to its apical width; ratio denticornis (Haliday, 1838) of the width of malar space to the width of face; Lampronota denticornis WzYiAzy , 1838: 121. interception of the nervellus. ? Cylloceria accusator (Fabricius) sensu Fitton, 1976: Characteristics of the female. Front wing 32 (8): 334. The Fabrician type material does not belong to the 7.1 —8.9 mm long. Palpi brown. Malar space Oxytorinae (Van Rossem, 1980). wide, 0.34—0.40 of width face, somewhat co- Cylloceria melancholica f. marginaler Schiodte, 1839: riaceous. Face laterally widely punctured, me- 24 (sensu Van Rossem, 1980). dially somewhat convex. Postanellus long, 8.0 9.6 as long as apically wide. Second and third Characteristics of the female lectotype of flagellar segments long. Frons, vertex and gena Lampronota denticornis. Labels: "British"; a polished and with fine punctures (implantations separate label: 20-2-82; lectotype label of Fit- of hairs). Pronotum with coarse sculpture, ir- ton, 1975 (Nat. Mus. Ireland, Dublin). The sec- regularly striated. The basal part of median lobe ond to sixth tergite have an orange to yellowish of the mesoscutum closely punctured. The hind margin and a brownish to light brown col- lateral lobes indistinctly punctured. Propodeum our. roughly sculptured, the carination indistinct or absent. Mesopleurum polished, but in the last Material examined of Cylloceria melancholica specimens of the table, finely striated. The two (Gravenhorst). — U.S.S.R. : several 9, Guzeril, Kav- prepectal carina reaching to the margin. Coxae kazck (Caucasus), Zapov (National Park), ysjtsj, Zje- and trochanters fuscous. Other parts of legs yel- lobnoi, leg. D. R. Kasparyan, 22. vi. 1976; 2 sp., Te- lowish to orange to brown (hind tibia). Hind bertsinsky, Zapov (Nat. Park), g. M. Chatipara, chv les (forest), leg. D. R. Kasparyan, H.vii. Berbi- tarsus fuscous. Nervellus intercepted below the 1976; guno, 26. V. 1891, Lisuv. ber., leg. K. Kokujeva. Identi- middle or in the middle. First tergite with regu- fication dubious, ovipositor too short, Sejdozero, 20 lar and rather fine sculpture, slightly striated. km south east of Revdy, Murmansk district, leg. D. R. Second tergite with regular and fine striation, Kasparyan, 25. vii. 1974 (label Aubert; Cylloceria). All the apical margin polished. Third tergite with specimens from the Zoological Institute at Leningrad. some fine striation and coriaceous sculpture. The fourth tergite sometimes also showing in- Distribution. — Cylloceria melancholica (Graven- distinct striation and some coriaceous sculpture. horst) is widely spread in the Western Palaearctic Re- gion. Ovipositor 0.9— 1.1 of the length of the front wing. Genus Proclitus Forster

Cryptus (Clepticus) Haliday in Curtis, 1838: 112 Material examined. — Finland: (J9, Suomi Ini. 121. Utsjoki, 10 & H.vii. 1961 (coll. R. Jussila). U.S.S.R.: Proclitus Forster, 1868: 172. 4 9, Reservoir Vudjavr, Chibin. g. Kolsk (mountain),

Proclitus; Forster, 1871 : 113. leg. Tsevurova, 30. vili. 1930; Jujasnor chibinsja, gory Plectiscus (Proclitus); Thomson, 1888: 1306—1307. Kolsk, leg. Fridolin, 8.viii.l937; Burunduk, kos. r.

Proclitus; Townes, 1971 : 194. Adzva, Arkh. g. leg. Kuluk, 29.vii.1909; Kozlovo, 50 = Proclitus; Aubert, 1977: 142. km s. Kaigi (? Kalga), Tsit. (=? Chitinskaya), obi. ( Proc/zr^s; Van Rossem, 1983^: 153—165. oblast = province), leg. Kasparyan, 16. vii. 1975 (all Zoological Institute, Leningrad, curator D. P. Kaspa- (coll. Key to the males ryan). Poland: S -, lectotype, "Warmbrunn" Gravenhorst, Wroclaw). The males of P. comes (Haliday in Curtis), P. fulvicorms Forster and P. rudis Forster are un- Cylloceria melancholica (Gravenhorst, 1820) known. I have not seen the male of P. fulvipec- Ichneumon melancholicus Gravenhorst, 1820: 372. tus Forster. These four species are not included Lampronota Haliday, 1838: 121. fracticornis in the key. Cylloceria fracticornis; Fitton, 1976: 334. The distinction of the males of P. praetor Characteristics of the female lectotype of Lampronota (Haliday in Curtis), P. ardentis species nova, P. fracticornis. Labels: "British"; a separate label: attentus Forster and P. alhidipes Forster is al- Haliday 20-2-82; lectotype label of Fitton (Nat. Mus. Ireland, Dublin). In my key the specimen most impossible; nevertheless I have included runs to C. melancholica. them in the key. I based the distinction on the —

Van Rossem; Western Palaearctic Oxytorinae 59

ratio of the length of the first abdominal seg- 2. Ovipositor exceptionally long, 0.80— 1.05 ment to the apical width. I have used only those of length front wing. First abdominal seg- units which are clearly separated. This implies ment 2.5— 3.0 times as long as the apical that other data can overlap. width P. comes (Haliday in Curtis) — Ovipositor not exceptionally long, 0.60

1. Anterior tentorial pits impressed and with a 0.70 of length front wing. First abdominal carina between the eye margin and the segment 2.7—4.5 times as long as the apical lateral corner of the clypeus. Clypeus flat, width 3 the apical margin arcuate, 2.5 times as wide 3. First abdominal segment 3.5—4.5 times as as long P. paganus (Haliday m Curtis) long as the apical width. Ovipositor 0.59 — Anterior tentorial pits not impressed and 0.69 of length front wing. Clypeus for the

without carina between eye and clypeus . . 2 greater part yellow 2. Face wide, about 0.45 of frontal width (in- P. praetor (Haliday in Curtis) cluding the eyes) and with a row of con- — First abdominal segment 2.7—3.0 times as spicuous long, erect setae along the inner long as apical width. Ovipositor 0.63—0.65 margin of eye. Clypeus exceptionally wide, of length front wing. Clypeus fuscous almost 3 times as wide as long, the apical P. ardentis spec. nov. margin arcuate. Lateral corner of clypeus 4. Anterior tentorial pits impressed and with a reaching beyond line of inner margin of carina between eye margin and lateral cor- eye. Clypeus with widely placed long setae ner of clypeus 5 P. suhsulcatus Förster — Anterior tentorial pits not impressed and — Not this combination of characters 3 with a weak or no carina between eye mar-

3. First tergite coriaceous. Lateral carinae of gin and clypeus 7 scutellum running to apex of scutellum, not 5. Ovipositor 0.27—0.31 of length of front meeting P. zonatus (Gravenhorst) wing. Clypeus flat, 2.4 times as wide as — First tergite not coriaceous. Lateral carinae long, the apical margin weakly arcuate. of scutellum not running to apex, only pre- Margin of tergite two yellowish brown, ter- sent at proximal corners of scutellum .... 4 gite three yellowish brown or entirely fus- 4. First abdominal segment 4.0—4.7 times as cous P. paganus (Hahday in Curtis) long as the apical width. Clypeus for the — Ovipositor 0.37—0.48 of length of front greater part yellow wing 6 P. praetor (Hahday in Curtis) 6. Ovipositor 0.37—0.47 of length of front — First abdominal segment less than 4.0 times wing. Median dorsal carina of first tergite as long as the apical width. Clypeus fuscous present. Margin of tergite two and tergite 5 three entirely more yellow in colour

5. First abdominal segment 3.5—3.7 times as P. fulvicornis Förster long as the apical width — Ovipositor 0.48 of length of front wing. P. ardentis spec. nov. Clypeus medially strongly convex, the api- — First abdominal segment 3.1 —3.3 times as cal margin arcuate. Median dorsal carina long as the apical width P. attentus Förster absent. Abdominal tergites from margin of tergite two to apex orange in colour

Remark. — There is a male paralectotype of P. edwardsi Roman Proclitus edwardsi with the same labels, but I 7. Ovipositor very short, 0.26 of length front doubt whether this male agrees with the female. wing. Malar space with a groove. Lateral The specimen does not show the impressed an- carinae of scutellum running to apex of scu- terior tentorial pits and neither the shape of the tellum, not meeting. Apex of scutellum clypeus agrees. striated or with somewhat rough sculpture. First, and in some specimens also the sec- Key to the females ond tergite coriaceous (Females of P. suhsulcatus Förster and P. albt- P. zonatus (Gravenhorst) dipes Förster are unknown) — Ovipositor longer, 0.36—0.48 of length L Ovipositor long, 0.60—L05 of length front front wing. Lateral carinae of scutellum on- wing 2 ly at proximal corners of scutellum 8 — Ovipositor shorter, less than 0.60 of length 8. Anterior tentorial pits conspicuous, open. front wing 4 A carina and laterally of the carina a groove 60 Tijdschrift voor Entomologie, deel 130, 1987

between eye margin and lateral corner of narrow. All parts of head fuscous. Frons be- clypeus P. fulvipectHS Förster tween antennal sockets impressed. Pronotum — Anterior tentorial pits not conspicuous. fuscous, pohshed. Epomia absent. Other parts There is a groove between eye margin and of thorax pohshed and fuscous. Prepectal carina clypeus, but no carina present 9 present, but weak. Front and middle legs, in- 9. Ovipositor 0.42—0.48 of length front wing. cluding coxae yellow. Hind coxae yellow. Apex First abdominal segment 2.7—3.0 times as of hind femur, tibia and hind tarsus brown. First long as the apical width. Radius originating abdominal segment 3.5 times as long as the api- at 0.54 of lower margin of stigma cal width. Median dorsal carina present on post- P. attentus Förster petiole. All other tergites polished. Apical mar- — Ovipositor 0.39 of length of front wing. gin of tergite two yellow. Tergites three, four First abdominal segment short, 2.0 times as and five for the greater part yellowish brown. long as the apical width'). Radius originat- All tergites, except the first, with adpressed long ing at 0.50 of lower margin of stigma setae.

P. rudis Förster The trivial name "ardentis" is from the Latin for "sparkling". Proclitus ardentis species nova Characteristics of the holotype of Proclitus Material examined. — Austria: 9, holotype, ardentis. Female. Front wing 4.97 mm long. Oberösterreich, Riedl im Haselgraben, 12. ix. 1985; 2 (?, paratypes, same locality and date as holotype; 1 9, Clypeus somewhat convex, fuscous, its apical paratype, Oberösterreich, Felsleiten bei Eidenberg, margin protruding, with rather long setae. Ma- 10. ix. 1985; 9, paratype, Oberösterreich, Brunnwald lar space narrow, about as wide as the mandibu- bei Bad Leonfelden, 21.viii.l985; â, paratype, lar base, with a groove eye margin and between Oberösterreich, Schauerschlag, bei Zwettl R., clypeus. Clypeal fovea somewhat impressed, 15.ix.1985 (the entire series leg. & coll. Martin but not forming a distinct carina between eye Schwarz, Zwettl, Osterreich). margin and clypeus. Face and other parts of head fuscous. Antennal sockets slightly el- Distribution. — Only four localities in Austria are evated. Upper part of face between antennal kwown. sockets with a circular impression. Flagellum slender. Postannellus 5.6 times as long as the ap- Proclitus edwardsi Roman ical width. Pronotum polished, epomia almost Proclitus edwardsi Roman, 1923: 73—74. obhterated. Mesoscutum polished, notaulus only present as a dent on the mesoscutal margin. Characteristics of the lectotype of Proclitus Propodeum almost entirely polished, carinae edwardsi. Female. Front wing 4.71 mm long. present but rather weak. Mesopleurum pol- Palpi whitish. Mandible yellow. Malar space ished, prepectal carina almost obliterated. Front 0.35 of width face. Clypeus medially strongly and middle legs, mluding the coxae, yellow. convex, the apical margin arcuate. Anterior ten- Hind coxae yellow, hind femur, tibia and tarsus torial pits strongly impressed and with a carina brownish. First abdominal segment 2.9 times as between eye margin and lateral corner of cly- long as the apical width. Postpetiole with some peus. Face, frons, vertex and gena polished, fus- longitudinal striation, median dorsal carina cous. Pronotum fuscous, polished, epomia pre- weakly present. Second tergite polished, thyri- sent, short. Mesoscutum polished, notaulus dia present and rather large, the apical tergal short. Scutellum polished, the margin only pre- margin yellow. Third tergite polished and al- sent at the front corner. Scutellar fossa deep. most wholly yellow. Remaining tergites fus- Propodeum polished, apical transverse carina cous, polished and with adpressed long setae. strong. Median longitudinal carina developed as Ovipositor 0.65 of the length of front wing. stubs on apical transverse carina. Mesopleurum polished, prepectal carina not reaching to the Characteristics of a male paratype of P. ar- margin. Legs, including coxae yellow. Coxae dentis. Front wing 4.62 mm long. Clypeus polished. Front femur stout, about 3.0 times as somewhat convex, fuscous, its apical margin long as wide. Postpectal carina absent. First ter- protruding, setae not conspicuous. Malar space gite fuscous, polished, median dorsal carina absent. The spiracle in the middle. The first seg-

ment is 2.2 times as long as apically wide. All

') Based on the holotype only. other tergites polished, from the apical margin —

Van Rossem: Western Palaearctic Oxytorinae 61

of tergite two orange in colour. The ovipositor as apical width P. dispar Van Rossem 0.48 of the length of the front wing. — Section gh of radiella present. First tergite The species is a parasite of Brachypeza radia- 1.4—2.3 times as long as apical width .... 4 ta Jenkins. 4. Section gh of radiella equal to or shghtly shorter than section dg. First tergite 1.8 Material examined. — England: lectotype, labels: 2.3 times as long as apical width B.M. Type Hym. 3.b.l623; Shefford Beds, viii.1918, P. inaequalis Förster leg. F. W. Edwards, ex Brachypeza radiata Jenk. in — Section gh of radiella longer than section Pleurotus; Proditus edwardsi Roman, 9, label: Prodi- dg. First tergite 1.4—2.5 times as long as tus Edwardsi Rn n.sp. type. Lectotype label Fitton, apical width P. lundus Förster 1977.

Pantisarthrus gracilis species nova Genus Pantisarthrus Forster Pantisarthrus Förster, 1871: 109—110. Characteristics of the holotype. Female. Pantisarthrus; Townes, 1971: 193— 194. Front wing 3.5 mm. Palpi yellow. The mandible Pantisarthrus; Van Rossem, 1980: 11C^113. not turned inwards, the lower tooth much smaller than upper. Clypeus convex, yellowish I found an undescribed species in Haesel- brown in colour, the margin truncate. Clypeus barth's collection. It is described below. with erect, long hairs. Occiput slightly concave and sloping beyond ocelli. Pronotum polished, Key to the species with epomia. Mesoscutum polished, notaulus present towards centre of mesoscutum, but 1. First tergite exceptionally long, 2.7—3.0 shallow. Prescutellar fovea wide and deep. Pro- times as long as apical width, with rough podeum with strong apical transverse carina and irregular longitudinal sculpture. Profile with an obtuse tooth between median longitudi- of tergite trapezium shaped. Claws of front nal carinae (also a character of other Pantisarth- leg conspicuously stronger than of middle rus species). Mesopleurum polished, prepectal and hind leg. Ovipositor 0.16 of length hind carina present, not reaching to the margin. Por- tibia P. gracilis spec. nov. tion of cubitus between intercubitus and second j— First tergite not more than 2.5 times as long recurrent vein 0.5 times as long as recurrent as apical width. Sculpture of first tergite co- vein. In the hind wing gh shorter than dg. Dis- riaceous. In some males surpassing 2.5 coidella absent. Legs, including the coxae, yel- times but in these the first tergite coria- lowish. Dorsal part of hind tibia brown. Claws ceous 2 of front leg stronger than those of middle and 2. Second tergite and following polished but hind legs. Hind femur conspicuously club- with some, not very obvious robust, irregu- shaped. First tergite exceptionally long, 2.7 larly placed punctures. Mesoscutum strong- times as long as the apical width. The tergite has ly convex, prescutellar groove conspicuous- rough longitudinal sculpture and a trapezium ly deep. A not very distinct species, of shaped profile. All other tergites fuscous and

which only the holotype is extant highly polished. Second tergite with a yellow P. rudepunctatus Strobl apical band. Ovipositor 0.16 of the length of — Second tergite and following polished, hind tibia. without punctures 3 Male unknown. 3. Section gh of radiella (fig. 3) in hind wing absent. First tergite 2.0—2.6 times as long Material examined. — Italy: 9, holotype, Prov. Bolzano, Kaltern Leuchtenburg, 500 m, 22. ix. 1978;

9 , paratype, Prov. Bolzano, Kaltern Leuchtenburg, 500 m, 22. ix. 1978 (both leg. and coll. Haeselbarth, München).

Pantisarthrus inaequalis Förster

Pantisarthrus inaequalis Förster, 1871 : 1 10. Pantisarthrus ochropus Förster, 1871: 110. Pantisarthrus pseudochropus Strohl, 1903: 137. Fig. 3. Hind wing of an ichneumonid. dgh = radiella; Aniseres subalpinus Strobl, 1903: 138. jmp = nervellus; mn = discoidella. After Townes Pantisarthrus inaequalis; Van Rossem, 1980: 110 (1969). 111. 62 Tijdschrift voor Entomologie, deel 130, 1987

Characteristics of male and female. Length Förster (1871) published keys to his Plectiscus front wing 3.0—3.6 mm. Tyloids absent. Malar females and males and at the same time intro- space wide, with a groove. Face, frons and ver- duced 51 new species in his key to the females.

tex polished. Occipital carina present. Mesoscu- The description of these is extraordinarily con- tum convex, poHshed, notaulus absent. Scutellar cise and based upon characters which do not carina almost reaching to apex. Section gh of ra- lead to identification. Apparently Förster con- diella equal to or shorter than section dg. First fused the conception of a species with mere in- tergite 1.8—2.3 times as long as wide, coria- dividual differences, describing specimens rath- ceous. Median dorsal carina present. er than species.

The Förster type material is in the Zoolog- Common and widely distributed. ische Staatssammlung at München and in excellent condition. Aubert (1975) first revised this Pantisarthrus luridus Forster collection and recognized 14 species.

Pantisarthrus luridus Forster, 1 871 : 110. A subgenus, Fugatrix, is introduced to incor- Pantisarthrus luridus; Van Rossem, 1980: 111 — 112. porate one species, Plectiscus communis Förster in the genus Plectiscidea. This species was Characteristics of male and female. Length placed in Dtalipsis by Townes (1971). I think front wing 3.0—3.6 mm. Section dg of radiella that the character used by Townes to separate about 0.6 of section gh. First tergite L'I—2.0 Dialipsis and Plectiscidea, that is the ratio be- times as long as wide. tween the length of the petiolar area and the length of combined areola and basal area, does Common and widely distributed. not exclude other Plectiscidea species from Di- alipsis. Neither the shape of the postannellus Pantisarthrus rudepunctatus Strobl separates these two genera. It seems to me that Pantisarthrus rudepunctatus Strohl, 1903: 137— 138. only the exceptionally large clypeal fovea is a Pantisarthrus rudepunctatus; Van Rossem, 1980: 112. true character of Dialipsis. Townes considers this character to be only an aberrant one. Con- Characteristics of the female. Length front sequently the other species belong to the subge- wing 3.0 mm. Section dg of radiella 0.5 of sec- nus Plectiscidea. tion gh. First tergite 2.4 times as long as wide, On the whole 36 species are dealt with; of coriaceous. Other tergites, according to Strobl, these five are new: Plectiscidea indomita, P. with robust, irregularly placed punctures. I did foersten, P. substantiva, P. blandita and P. ven- not find these to be very conspicuous. tosa. Plectiscidea nemorensis is a nomen novum for Ephalmator subsimilis Van Rossem. Distribution. — Only the holotype of Styrian Alp

(Austria) is extant. Morphology

It is difficult to find proper morphological Pantisarthrus dispar Van Rossem characters for comparison and for the separa- Pantisarthrus dispar Van Rossem, 1980: 112— 113. tion of Förster's types. I used the following eight criteria:

Characteristics of male and female. Length of (1) ratio of width malar space to the width of front wing 2.8—3.3 mm. Section gh of radiella face (enlargement 100 x); absent. First tergite 2.0—2.6 times as long as (2) ratio of length of postannellus to its apical wide. width (enlargement 100 x); (3) presence or absence of notaulus; and widely Common distributed. (4) development of the lateral scutellar carina; (5) ratio of length of first abdominal segment to Genus Plectiscidea Viereck its apical width (enlargement lOOx);

Plectiscus auctores, before 1914. (6) ratio of length of first abdominal segment to Plectiscidea Viereck, 1914: 118. the length of the front wing (enlargement

8-1 Plectiscus; Förster, 1871 : —90. 40X); Plectiscus; Strobl, 1903: 125—130. (7) position of the spiracles of the first abdomi-

Plectiscidea; Townes, 1971 : 196 197. — nal segment in proportion to the length of Plectiscidea; Aubert, 1975: 3—5; 7—8. the segment (enlargement lOOx); Van Rossem: Western Palaearctic Oxytorinae 63

(8) length of ovipositor in proportion to the — Ovipositor longer than 0.30 of length of

length of the front wing (enlargement 40 X ). front wmg 15

3. Ovipositor 0.09 of length of front wing. These criteria offer a rather meagre founda- Postannellus long, 6.0 times the apical tion. Perhaps scanning electron microphotogra- width and with a conspicuous character, phy would reveal better characters, but I had no viz., a medial notch, thus giving the impres- access to such an mstrument. sion of two short inflated segments. Anten- A word must be said about the males. Förster na, legs, propodeum and gaster with long described 26 species, founded solely on the hairs. Abdominal segment 2.3 times as long males. I regard these as species inquirendae as it as the apical width. The first tergite coria- is impossible at present to find the matchmg fe- ceous P. nemorensis nom. nov. males. — Ovipositor more than 0.10 of length of front wing 4 Note to the key 4. Ovipositor 0.14—-0.18 of length of front An extensive series of Plecttsadea collaris col- wing. Postannellus 4.0—4.8 times as long as lected in Austria by Martin Schwarz gave me a the apical width. The first abdominal seg- better understanding of the variability of this ment 1.4— 1.8 times as long as the apical species. I, therefore, doubt if my key between width P. bistnata (Thomson) 17 and 18 is reliable. — Ovipositor longer than 0.18 of length of The following observations should also be front wing 5 noted: 5. Postannellus 4.0—4.8 times as long as the Plectiscidea monticola is distmguished by the apical width 6 short first abdominal segment. — Postannellus 5.0—6.5 times as long as the Plectiscidea agitator, of which only the holo- apical width 8 type is available, could be identical with P. col- 6. The length of the first abdominal segment laris. 1.3 times as long as the apical width due to Plectiscidea conjuncta can be distinguished the conspicuous breadth of the tergite. Ovi- from P. collaris by the ratio of the length of the positor 0.27 of the length of the front wing. first abdominal segment to the length of the Postannellus 4.2 times as long as the apical front wing (0.14—0.16 in P. conjuncta and width P. subteres (Thomson) 0.17—0.19 in P. collaris). In P. conjuncta the — The length of the first abdominal segment spiracles of the first abdominal segment lie at 2.0—2.4 times as long as apical width .... 7 0.31 —0.37 of the length of the segment. In P. 7. Ovipositor 0.20—0.22 of the length of front collaris at 0.37—0.42 of the length of the seg- wing. Postannellus 4.0—4.6 (4.8) times as ment. long as apical width. The length of the first Of P. conjuncta only the holotype and three abdominal segment 2.0—2.3 times as long other specimens were available. Two of these as apical width P. indomita spec. nov. are type specimens of P. ßavicoxis of which the — Ovipositor 0.25—0.27 of the length of front status is doubtful. wing. Postannellus 4.3—4.6 times as long as the apical width. The length of the first ab- Key to Plectiscidea females dominal segment 2.2—2.4 times as long as the apical width P. moerens (Forster) 1. Length of ovipositor 0.72—0.85 of length 8. Postannellus 5.0—5.6 times as long as the of front wing. Petiolar area of propodeum apical width 9 1.5—2.0 times as long as combined areola — Postannellus 6.0—6.5 times as long as the and basal area. Postanellus 3.6—4.5 times as apical width 12 long as the apical width. First abdominal 9. First tergite with longitudinal striation and segment 1.8—2.5 times as long as the apical some not very conspicuous coriaceous width. First tergite coriaceous. Front wing sculpture between. The spiracles rather 2.3 —3.6 mm long . . Fugatnx subgen. nov. protruding. The first abdominal segment — Length of ovipositor less than 0.70 of 2.1 times the apical width. Postannellus 5.0 length of front wing. (Subgenus Plectisci- times as long as the apical width. Malar dea) 2 space 0.41 of width face. Ovipositor 0.21 of 0.09 2. Ovipositor relatively short, —0.30 of the length of front wing . P. tener (Forster) length of front wing. 3 — First tergite with coriaceous sculpture .. 10 64 Tijdschrift voor Entomologie, deel 130, 1987

10. Ovipositor 0.19—0.22 of length of front the apical width 22 wing. Postannellus 5.0—5.5 times as long as 18. Ovipositor 0.55 of the length of front wing. the apical width. First abdominal segment Postannellus long, 6.0 times the apical

1.8—2.0 times as long as the apical width . . . width. Length of first abdominal segment P. parvula (Forster) 2.7 times the apical width. Front wing 4.8 — Ovipositor 0.24—0.30 of length of front mm long P. erythropyga (Forster) wing. Postannellus 5.0—5.6 times as long as — Ovipositor shorter, 0.35—0.52 of the the apical width 11 length of front wing 19 11. Ovipositor 0.24—0.27 of length of front 19. First abdominal segment short, 2.0 times wing. Postannellus 5.5—5.6 times as long as the apical width. Ovipositor 0.44 of length the apical width. First abdominal segment of front wing. Postannellus 5.0 times as 2.0—2.3 times as long as apical width long as the apical width. Front wing 3.7 mm P. tenuicornis (Forster) long P. monticola (Forster) — Ovipositor 0.28—0.30 of length of front — Length of first abdominal segment 2.3—2.9 wing. Postannellus 5.0—5.5 times long as times the apical width 20 the apical width. First abdominal segment 20. Postannellus 6.0 times as long as the apical 1.9—2.3 times as long as apical width width. Ovipositor 0.40 of the length of P. cinctula (Forster) front wing. First abdominal segment 2.3 12. Ovipositor 0.22—0.23 of length of front times as long as the apical width. Front wing. Postannellus 6.0 times as long as the wing 4.5 mm long .... P. agitator (Forster) apical width. The first abdominal segment — Postannellus 5.0—6.3 times as long as the 2.3—2.6 times as long as apical width apical width. Ovipositor 0.35—^0.52 of P. amicalis (Forster) length of front wing 21 — Ovipositor 0.25—0.30 of length of front 21. The first abdominal segment measures wing 13 0.14—0.16 of the length of the front wing. 13. First abdominal segment 2.4—2.7 times as Ovipositor 0.36—0.38 of length of front long as apical width. Ovipositor 0.25—0.31 wing. Postannellus 5.2—5.3 times as long as of length of front wing. Postannellus 6.5 the apical width. First abdominal segment 2.2 times as long as apical width —2.4 times as long as the apical width . . . P. helvola (Forster) P. conjuncta (Forster) — First abdominal segment 2.0—2.3 times as — The first abdominal segment measures long as apical width 14 0.17—0.19 of the length of the front wing. 14. Ovipositor 0.25—0.30 of length of front Ovipositor 0.35—0.52 of length of front wing. Postannellus 6.0 times as long as the wing. Postannellus 5.0—6.3 times as long as apical width. Length of first abdominal seg- the apical width. First abdominal segment 2.1 2.3 ment —2.3 times the apical width —2.9 times as long as the apical width . . . P. vagator (Forster) P. collans (Gravenhorst) — Ovipositor 0.27—0.30 of length of front 22. Postannellus 3.7 times as long as the apical wing. Postannellus 6.0—6.5 times as long as width. Ovipositor 0.44 of length of front the apical width. Length of first abdominal wing. First abdominal segment 2.4 times as segment 2.0 times the apical width long as the apical width. Front wing 4.7 mm P. melanocera (Forster) long P. foersteri spec. nov. 15. Notaulus indicated by a groove on the — Postannellus 4.2—4.6 times as long as the mesoscutal margin 16 apical width 23 — Notaulus not present or evanescent 25 23. Ovipositor 0.38 of length of front wing. 16. First abdominal segment conspicuously Postannellus 4.3 times as long as apically long, 3.1 —3.5 times the apical width. Post- wide. First abdominal segment 2.3 times as annellus (4.8)—5.2 times as long as the api- long as the apical width. Front wing 3.7 mm cal width. Ovipositor 0.33—0.36 of length P. nava (Forster) of front wing .... P. canaliculata (Forster) — Ovipositor 0.41 —0.48 of length of front — Length of first abdominal segment less than wing. Postannellus 4.0—4.6 times as long as 3.0 times the apical width 17 the apical width 24 17. Postannellus 5.0—6.3 times as long as the 24. Malar space 0.29—0.35 of width face. Ovi- apical width 18 positor 0.43—0.48 of length of front wing. — Postannellus less than 5.0 times as long as Postannellus 4.0—4.5 times as long as the Van Rossem: Western Palaearctic Oxytorinae 65

apical width. First abdominal segment space 0.37 of width face 2.4—2.7 times as long as the apical width. P. mendica (Förster) Front wing 4.2—5.2 mm long — The spiracles of the first abdominal segment P. substantiva spec. nov. situated at 0.40—0.43 of the length of the — Malar space wide, 0.41 —0.42 of width face. segment 31 Ovipositor 0.41 —0.43 of length of front 31. The spiracles of the first abdominal segment wing. Postannellus 4.2—4.6 times as long as situated at 0.40 of the length of the seg- apical width. First abdominal segment ment. The length of the first abdominal seg- 2.3—2.6 times as long as the apical width. ment 2.3 times the apical width. Postannel- Front wing 3.7—4.5 mm long lus 5.5 times as long as the apical width. P. crassicornis (Förster) Malar space 0.38 of width face 25. Postannellus extremely long, 7.0 times the P. fraterna (Forster) apical width. Ovipositor 0.33—0.35 of — The spiracles of the first abdominal segment length of front wing. First abdominal seg- situated at 0.43 of the length of the segment 2.7 times as long as apical width ment. The length of the first abdominal seg- P. posticata (Förster) ment 2.1 times the apical width. Postannel- — Postannellus shorter, less than 5.8 X the ap- lus 5.0 times as long as the apical width. ical width^) 26 Malar space 0.35 of width face 26. Ovipositor 0.40—0.47 of length of front P. deterior (Förster) wing. Postannellus 5.0—;5.7 times as long as 32. Malar space 0.42 of width face. The first ab- the apical width. Length of first abdominal dominal segment 2.5 times as long as the segment 1.8—2.8 times the apical width 27 apical width, the spiracles situated at 0.34 of — Ovipositor shorter, 0.33—0.38 of length of the length of the segment front wing. Postannellus 4.3—5.4 times as P. blandita spec. nov. long as the apical width. Length of first ab- — Malar space 0.33—0.35 of width face. The dominal segment 1.6—2.6 times the apical first abdominal segment 2.1 —2.3 times as width 33 long as the apical width, the spiracles situ- 27. Ovipositor 0.40 of length of front wing. ated at 0.34—0.37 of the length of the seg- Postannellus 5.3 times as long as the apical ment. Postannellus 5.0—5.3 times as long as width. Malar space 0.40 of width face. the apical width.') Ovipositor 0.42—0.47 of Length of first abdominal segment 2.7 times the length of the front wing. Front wing the apical width, the spiracles situated at 3.5—4.5 mm long .. P. terebrator {Vörsitr) 0.37 of the length of the segment 33. Postannellus 4.3 —4.6 times as long as the P. connexa (Förster) apical width. Ovipositor 0.32—0.37 of — Ovipositor more than 0.40 of length of length of front wing. Length of first ab- front wing 28 dominal segment 1.6— 1.8 times the apical 28. Length of first abdominal segment 1.8 times width P- ventosa spec. nov. the apical width, the spiracles situated at — Postannellus 5.0—5.4 times as long as the 0.32 of the length of the segment. Postan- apical width 34 nellus 5.7 times as long as the apical width. 34. Postannellus 5.4 times as long as the apical Ovipositor 0.42 of length of front wing .... width. Ovipositor 0.36 of length of front P. mesoxantha (Förster) wing. Malar space 0.31 of width face. Scu- — Length of first abdominal segment more tellar carina shghtly beyond the scutellar than 1.8 times the apical width 29 corner and turning inwards, but not meet- 29. The spiracles of the first abdominal segment ing. Length of first abdominal segment 2.2 situated between 0.40—0.50 of the length times the apical width. Front wing long, 5.2 of the segment 30 mm P. subangulata (Förster) — The spiracles of the first abdominal segment — Postannellus 5.0—5.2 times as long as the situated between 0.34—0.37 of the length apical width. Malar space 0.31 —0.38 of of the segment 32 width face. Front wing 3.5—4.3 mm long 30. The spiracles of the first abdominal segment 35 situated in the middle of the segment (0.50). The length of the first abdominal segment

2.0 times the apical width. Postannellus 5.0 ') In P. terebrator the postannellus can reach to 6.3- times as long as the apical width. Malar 6.6 times the apical width. —

66 Tijdschrift voor Entomologie, deel 130, 1987

35. Postannellus 5.0 times as long as the apical The male was originally described with

width. Malar space 0.37 of width face. Ovi- Ephalmator subsimilis. It is not taken into con- positor 0.36 of length of front wing. Scutel- sideration here. lar carina slightly beyond scutellar corner. Characteristics of the female. Front wing The spiracles of the first abdominal segment 3.0—4.1 mm long. Malar space 0.38—0.40 of situated at 0.43 of the length of the seg- width face. Anterior tentorial pits in the Dutch ment. First abdominal segment 2.5 times as specimen rather impressed. Postannellus 5.6 long as the apical width. Front wing 4.0 mm 6.0 times as long as apical width. The postannel- long P. eurystigma (Thomson) lus with a weak medial notch giving the appear- — Postannellus 5.0 times as long as the apical ance that the postannellus consists of two seg- width. Malar space 0.33 of width of face. ments. Notauh not present or fading. Propo- Ovipositor 0.37—0.38 of length of front deum coriaceous, median longitudinal carina wing. Scutellar carina slightly beyond cor- weak, present as stubs from the apical trans- ner and curving inwards, not meeting. The verse carina. The latter strong. Legs, including spiracles of the first abdominal segment sit- coxae, with long hairs. First abdominal segment uated at 0.32—0.34 of the length of the seg- 2.0—2.3 times as long as apical width. The first ment. First abdominal segment 2.0—2.6 tergite coriaceous, median dorsal carina absent, times as long as the apical width. Front the spiracles at 0.37—0.43 of the length. Second wing 3.6—4.3 mm long tergite with an apical yellow band. Third tergite P. humeralis (Forster) for the greater part yellow. The ovipositor exceptionally short, 0.09—0.10 of the length of Subgenus Plectiscidea Viereck the front wing. The main characters are shown

Plectiscidea Viereck, 1914: 118. Type: Plectiscus colla- on table 2. ris Gravenhorst. Original designation. Material examined. — Austria: 9, T. Pertisau, 1550 m, 12. vii. Front wing 2.7—6.0 mm long. Postannellus 1977 (paratype of Ephalmator subsimilis) (leg. & coll. Haeselbarth). Netherlands: 2, Naarder- 4.0—7.0 times as long as apically wide. Petiolar meer, Malaise trap, (Loc. ix), 6.viii.l974, leg. Bunnik area of propodeum around 1.2 times as long as & Van Wijngaarden (coll. K. W. R. Zwart, Wagening- combined areola and basal area. First abdominal en). segment 1.3—3.5 times as long as apical width. Ovipositor 0.09—0.55 of length of front wing. Remark. — The name "subsimilis" is preoccupied (Förster, 1871, Verb, naturh. Ver. pre- Plectiscidea nemorensis nomen novum uss. Rheinl. 28: 86). The new name is from the Ephalmator subsimilis Van Rossem, 1980: 122 (nee Latin for "from the holy wood"; the locahty Förster, 1871). Paratype examined in 1985. See al- "Naardermeer" is a nature reserve. so Van Rossem (1982: 169).

Table 2. Plectiscidea nemorensis nom. nov.: frw — the length of the front wing in mm; ovip/frw — ratio of ovipositor length to length of the front wing; psta l/w —ratio of length of postannellus to its apical width; malsp /ƒ — ratio of width of malar space to the width of face; abds l/w — ratio of length of first abdominal segment to its apical width; spir/abds — the position of the spiracles of the first abdominal segment in relation to the lenght of the segment; not — notaulus (if not filled in, no attention is given to the notaulus); pr — notaulus present; weak or prw — notaulus weakly present; not pr — notaulus absent; pet. area/area + bas — ratio of length of petiolar area to length of combined areola and basal area (used only for the subgenus Fugatrix)\ abds/frw — ratio of length of first abdominal segment to length of front wing (used only for P. conjuncta and P. collaris); not filled in — not determined. Van Rossem: Western Palaearctic Oxytorinae 67

Plectiscidea bistriata (Thomson) 17.V.1976; Hedemünden C, 12.V.1966; Dransfeld, 12.vi.1966 (all leg. & coll. Haeselbarth). Netherlands: PlectiscMS bistriatus Thomson, 1888; 1288. Lectotype Naardermeer (Malaise trap), 11. vi. 1974, leg. Bunnik designation by Van Rossem. There is a male in the & Van Wijngaarden (coll. Zwart, Wageningen). Swe- type series (Fitton, 1982). den: lectotype, Herrevadskloster (Skine), vi. 1882 (coll. Thomson, Entomology Museum, Lund). Characteristics of the lectotype. Female. Front wing 3.5 mm. Clypeus small, square, convex, somewhat protruding, the apical margin Plectiscidea subteres (Thomson) truncate. Face polished, slightly produced for- Plectiscus subteres Thomson, 1888: 1300. Holotype ward below the antennal sockets. Postannellus label of R.Hinz (1962). 4.0 times as long as the apical width. Occipital carina closed. Mesoscutum polished, notauli Characteristics of the holotype. Female. weakly present. Scutellum with carina to apex. Front wing 4.2 mm. Clypeus polished, outer Propodeum almost completely polished, with margin truncate, medially protrudmg, with erect rather long hairs. Mesopleurum polished, erect hairs, width 0.61 of width face. Face, prepectal carina present but widely away from frons, vertex and temple polished. Occipital ca- the margin. Legs and coxae brownish. First ab- rina complete. Postannellus 4.2 times as long as dominal segment 1.5 times as long as apical the apical width. Pronotum polished, epomia width. Postpetiole finely coriaceous. Apical half strong. Mesoscutum polished, notauli present of abdomen compressed. Ovipositor 0.18 of the but obHterated posteriorly. Scutellum with the length of the front wing. Compare table 3. carina partly present. Most of the propodeum The two main characters of this species are with irregular sculpture, carinae rather strongly the short ovipositor: 0.14—0.18 of the length of developed. Mesopleurum polished, prepectal the front wing and the rather short and wide carina present but well away from the margin. first abdominal segment (1.4— 1.8 times the api- Coxae and legs yellowish brown, legs slender. cal width). Length of the abdominal first segment 1.3 times The male was described by Thomson (1888). the apical width. The apical margin wide. First tergite coriaceous, median dorsal carina to 0.42 Distribution. — The species is widely spread in the of the length of the tergite. The spiracle situated Palaearctic region: Austria, Germany, the western at 0.37 of the length of the segment. The end of Netherlands and Sweden. the first sternite is at 0.28 of the length of the

segment. The second tergite is coriaceous, the Material examined. — All 9. Austria: Kärnten, apical margin is polished and light brown in col- Bodental 1100 m, 30.vi.l981 (leg. & coll. Zwakhals); Sonnenwendgebirge, 1300— 1500 m. Hint. Swjoch our. The ovipositor is 0.27 of the length of the 21.vi.1959; St. Haus Gföhlalm, 1300 m, 27.vi.1972. front wing. Compare table 4. Germany: Oberbayern, Gauting, 5.vi.l972; ibidem, Male unknown.

Table 3. Plectiscidea bistnata (Thomson). For explanation of abbreviations, see table 2. —

68 Tijdschrift voor Entomologie, deel 130, 1987

Table 4. Plectiscidea suhteres (Thomson). For abbreviations, see table 2.

frw ovip/ psta abds frw 1/w 1/w

holotype P. suhteres 4.0 0.27 4.2 1.3

Material examined, - Germany: (?), 2, holotype, Distribution. — Widely spread in the western Pal-

f. 22.V.1886, leg. and dII. Thomson (Entomological aearctic Region. Museum, Lund). "Indomitus" is the Latin for "invincible", "indomitable". Plectiscidea indomita species nova Plectiscidea moerens (Forster) Characteristics of the holotype of P. indomi- Plectiscus moerens Förster, 1871: 87. Holotype la- ta. Female. Front wing 3.8 mm long. Mandible belled by Van Rossem. yellow, teeth about the same length. Clypeus Plectiscus xanthoneuris Förster, 1871: 87. Holotype rather convex, somewhat protruding. Malar labelled by Van Rossem. New synonym. space rather wide, 0.43 of width of face. Postan- Plectiscus flavizonus Förster, 1871: 88. Holotype la- nellus 4.0 times as long as the apical width. Epo- belled by Van Rossem. New synonym. mia present. Notaulus present, short. Lateral Plectiscus eversorius Förster, 1871: 88. Holotype labelled by Van Rossem. New synonym carina of scutellum running somewhat behind the corner and slightly curving inwards. Propo- Characteristics of the holotype of P. moerens. deum dorsally with coriaceous sculpture, Female. Front wing 3.4 mm long. Postannellus laterally with irregular fine sculpture. Prepectal 4.3 times as long as apical width. Epomia pre- carina present. First abdominal segment 2.0 sent, short. Notaulus short. Lateral carina of the times as long as the apical width. The spiracles scutellum somewhat behind the corner. Propo- situated at 0.40 of the length of the segment. deum polished, with rather long hairs. Prepectal The first tergite medially rather convex, with carina present. First abdominal segment 2.4 coriaceous sculpture. The second tergite with times as long as apical width. Spiracles situated vague coriaceous sculpture. The other tergites at 0.33 of the length of the segment. First tergite polished. Ovipositor 0.22 of the length of the coriaceous, with rather long lateral hairs. Sec- front wing. Compare table 5. ond tergite polished. Ovipositor 0.27 of the Male unknown. length of the front wing. Compare table 6.

The species is characterized by the ratio of Material examined. — Austria: paratype, T. Lech- length and width of the postannellus, 4.3—4.6 taler A., Bichlbachle, 1350 m, 14.viii.l974. Germany: times as long as the apical width, the length of Hohenschwangau, Säulingweg, 1120 9, holotype, the ovipositor, 0.25—0.27 of the length of the 1300 m 16.vii.1974; 2 9, paratypes, Weszling, Hoch- front wing and the ratio of length and width of stadt, 19.V.1974 (all leg. & coll. Haeselbarth); 9, para- the first abdominal segment, 2.2 2.4 times as lectotype of Plectiscus sodalis Förster, Lousberg, 19.v — long as the apical width. The second tergite al- (coll. Förster, München). Italy: BS, Valvestino Malga Tombea, 1800 m, 14.vi.l976 (leg. & coll. Haesel- most polished in most specimens. barth). Male unknown.

Table 5. Plectiscidea indom Van Rossem: Western Palaearctic Oxytorinae 69

Table 6. Plectiscidea 70 Tijdschrift voor Entomologie, deel 130, 1987

Table 8. Plectiscidea parvula (Forster). For abbreviations, see table 2. Van Rossem: Westerti Palaearctic Oxytorinae 71

urus. Female. Front wing 2.8 mm long. Postan- width. First tergite coriaceous. Ovipositor 0.29 nellus 5.5 times as long as apical width. Notauli of length of front wing. weak, with a short carina on the inner margin. Characteristics of the lectotype of P. curti- First tergite coriaceous. First abdominal seg- cauda (Thomson). Female. Front wing 4.3 mm ment 2.0 times as long as apical width. Oviposi- long. Postannellus 5.1 times as long as apical tor 0.26 of length of front wing. width. Epomia short. Notaulus rather strong,

The species is characterized by the length of but short. Lateral carina of scutellum not run- the ovipositor (0.24—0.27 of the length of the ning behind the corner. Propodeum with irreg- front wing). The postannellus is 5.5—5.6 times ular sculpture, the apical transverse carina with as long as the apical width. weak apophyses. The first abdominal segment The male was described by Thomson (1888). 2.5 times as long as apical width. The spiracles lying at 0.35 of the length of the segment. The Material examined. — Austria: 9, Kärnten, Zell first tergite coriaceous. About 0.60 of the sec- Pfarre, 1200 m, 14.vii.l979; 9, Kärnten, Himmel- ond tergite with coriaceous sculpture. Oviposi- Zwakhals); berg, 1000 m, 15. vili. 1980 (both coll. 9, tor 0.27 of the length of the front wing. Fliess (Tirol), 1550 m, ll.viii.l971 (coll. Haeselbarth). Male unknown. Germany; 9, holotype of P. tenuicorms, Lousberg, 25.x.; 9, holotype of P. brachyurus, Aachen (both Remark. — The lectotype and paralectotype leg. & coll. Forster, Münden); 2 9, Hann. München, of P. determinata are placed provisionally un- 13.viii.l965; 9, Hedemühden, 12.V.1966; 9, Lip- poldshausen, B, 20.V.1967; 9, Hessen, Witzenhausen, der P. cinctula. In both specimens the ratio of 14. V. 1966; 9, Niedersachsen, Wiershausen, the length and width of the postannellus does

19. vili. 1966; 2 9, Bayern, Herrsching, Kerschlacher not agree with the holotype of P. cinctula. Forst, 19. V. 1974; 9, Oberbayern, Weszling, Compare table 10. 2.viii.l973. Italy: 9, Campenjoch, Südtirol, 1350 m, A, 3. ix. 1967 (all leg. & coli. Haeselbarth). Nether- Material examined. — Austria: 9, T. Lechtaler Al- Malaise trap, 4. vi. 1974, leg. lands: 9, Naardermeer, pen, Bleispitze, 1900—2200 m, 14.viii.l974 (leg. & Malaise trap, Bunnik & Van Wijngaarden; 9, coll. Haeselbarth). Germany: 9, holotype of P. cinc- Wijngaarden (both speci- 8.x. 1974, leg. Bunnik & Van tula, Aachen (coll. Förster, München); 9, lectotype of coll. Zwart, Wageningen). Norway: 9, Opp- mens P. curticauda (Thomson), ? Kaltenkirchen, 20.vii.l978 (leg. coli. Van land, Lom-Lia, 26.vi.— & 23.viii.1886 (coll. Thomson, Lund); 3 9, Hedemün- Klysna "Norr- Rossem). Sweden: 9, Örebro Län, den, C, 12. v. 1966; 9, Oberbayern, Deisenhofen, vii. —2.viii.l979; Värmland, Transtrand, berga", 8. 9, 26. vii. 1958; 2 9, Bayern, Hohenschwangau, Säuling- 1" 21 Storbacken, Station, —31. vii. 1981, margin of weg, XnO—UQQ m, 16.vii.1974 and Wildsulz, 1420— forest, close vegetation (leg. coll. Van Rossem). & 1560 m, 16.vii.1974 (all leg. & coll. Haeselbarth). Sweden: 2 9, Fjätervilen — Idre, Dalarna, 2.viii— species is widely spread in the Distribution. — The 12.viii; 24. vii— l.viii.l982 (leg. & coli. Van Rossem); Western Palaearctic Region. 9, Dalarna, Transtrand, Hemfjällstangen, vii. 1976 (leg. & coli. Van Rossem). Plectiscidea cinctula (Förster) Distribution. — A widely spread species which Plectiscus cinctulus Förster, 1871: 89. Holotype la- may tend to mountainous regions. belled by Van Rossem. Plectiscus determinatus Förster, 1871: 88. Lectotype designation by Van Rossem. New synonym. Plectiscidea amicalis (Forster) Plectiscus curticauda Thomson, 1888: 1302. Lectotype Plectiscus amicalis Förster, 1871: 87. Lectotype desig- designation by Aubert (1977). The specimen has nation by Van Rossem. an Aubert label "gleiche Art wie hier unten als Plectiscus sodalis Förster, 1871: 88. Lectotype desig- tenuicorms Frst." New synonym. nation by Van Rossem. New synonym.

Characteristics of the holotype of P. cinctu- Characteristics of the lectotype of P. amicalis. lus. Female. Front wing 3.8 mm long. Apical Female. Front wing 4.0 mm long. Postannellus part of clypeus yellow. Postannellus 5.0 times as 6.0 times as long as apical width. Notaulus long as apical width. Notaulus weak, only visi- weakly indicated on the margin. Mesopleurum ble on the margin. Scutellar carina running be- polished, prepectal carina not reaching the mar- yond the corner, not meeting at apex. Propo- gin. The first abdominal segment 2.6 times as deum with vague coriaceous sculpture. Prepec- long as the apical width. First tergite in part co- tal carina not reaching the margin. First riaceous. Second tergite with vague coriaceous abdominal segment 2.3 times as long as apical sculpture. Caster with rather conspicuous hairs. 72 Tijdschrift voor Entomologie, deel 130, 1987

Table 10. Plectiscidea cinctula Van Rossem: Western Palaearctic Oxytorinae 73

Table 12. Plectiscidea helvola (Förster. For abbreviations, see table 2. 74 Tijdschrift voor Entomologie, deel 130, 1987

ished, but the second tergite with vague coria- with P. canaliculata. The postannellus is rather ceous sculpture. Ovipositor 0.27 of the length short. The ovipositor reaches 0.32 of the length of the front wing. Compare table 14. of the front wing and the first abdominal seg-

The species is characterized by the long post- ment measures 3.5 times the apical width. annellus, 6.5 times the apical width; the length of the ovipositor, 0.27—0.30 of the length of the Material examined. — Germany: 9, lectotype of front wing and the short first abdominal seg- P. canaliculata Lousberg, 15. ix; 9, lectotype of P. subtilus, Aachen; locality (pencil label "sub- ment, 2.0 times the apical width. 9, no tilis"); lectotype of P. distincta, Aachen, 5. vii; Male unknown. 9, 9, lectotype of P. subcurvata, Lousberg, 5.v (all coll. Förster, München). Austria: 9, Kärnten, Bodental, Material examined. — Germany: 9, holotype of 1100 m, 13.vii.1981 (leg. & coll. Zwakhals). Ger- P. melanocera, Lousberg, 13. ix; $, lectotype of many: 9, Bayern, Hohenschwangau, 830— 1050 m, P. proxima, Lousberg, 11. ix (coll. Forster, München); 16.vii.1974 (leg. & coll. Haeselbarth). 2 9, Hedemünden, C, 12.V.1966 (leg. & coll. Haesel- barth). Netherlands: 9, Naardermeer, Malaise trap, Distribution. — Widely spread in the western Pal- 20.viii.l974; 9, Naardermeer, Malaise trap, 3.ix.l974; aearctic Region. 9, Naardermeer, Malaise trap, 8.X.1974 (all leg. Bun- nik & Van Wijngaarden) (coll. Zwart, Wageningen). Plectiscidea erythropyga (Förster)

Distribution. — Widely spread in the western Pal- Plectiscus erythropygus Förster, 1871: 88. Lectotype aearctic Region. designation by Aubert (1975).

Plectiscidea canaliculata (Förster) Characteristics of the lectotype of P. erythro-

Plectiscus canaliculatus Förster, 1871: 86. Lectotype pyga. Female. Front wing 4.8 mm long. Mandi- designation by Aubert. Report on type by Aubert bular teeth of the same length. Clypeus for the (1975). greater part yellow. Malar space impressed. Plectiscus subtilis Förster, 1871: 86. Lectotype desig- Postannellus 6.0 times as long as apical width. with P. ca- nation by Townes. Placed as synonym Epomia strong, almost reaching the mesoscutal naliculata by Aubert (1975). margin. Notaulus strong, reaching to about 0.3 Plectiscus distinctus Förster, 1871: 88. Lectotype des- of the distance to the center of mesoscutum. ignation by Aubert. Placed as synonym with Upper part of propodeum more or less pol- P. canaliculata by Aubert (1975). Plectiscus suhcurvatus Förster, 1871: 89. Lectotype ished, apical transverse carina strong. Meso- designation by Van Rossem. New synonym. pleurum polished, prepectal carina not reaching the margin. Hind coxa punctured by implanta- Characteristics of the lectotype of P. canali- tions of hairs. First abdominal segment 2.7 culata. Female. Front wing 3.1 mm long. Malar times as long as apical width. The first tergite space 0.33 of width face. Postannellus 5.2 times with longitudinal striated sculpture, also as long as apical width. Epomia present but somewhat coriaceous. In the basal half of the short. Notaulus strong on the mesoscutal mar- second tergite vague coriaceous sculpture, the gin. Lateral carina of scutellum at the corner. apical margin yellowish brown. Third tergite Propodeum with some vague coriaceous sculp- with a narrower yellowish margin. The apical ture and widely placed rather long hairs. Pre- part of the gaster dirty yellowish. Length of pectal carina present, short, not beyond ventral ovipositor 0.55 of the length of front wing. corner of pronotum. First abdominal segment Compare table 16. long, 3.1 times as long as apical width. First ter- Male unknown. gite coriaceous and with a vague median dorsal carina. Ovipositor 0.36 of the length of the Material examined. — Germany: 9, lectotype of front wing. Compare table 15. P. erythropyga, Lousberg, 15.x. (coll. Förster, Male unknown. München).

Remark. — The species is characterized by Plectiscidea monticela (Forster) the long first segment, 3.1 3.5 times abdominal — Plectiscus monticala Förster, 1871: 89. Lectotype des- as long as apical width. I am well aware that the ignation by Aubert. Report on type by Aubert lectotype of P. subcurvata does not quite agree (1975). Van Rossem: Wester7i Palaearctic Oxytorinae 75

Table 13. Plectisc 76 Tijdschrift voor Entomologie, deel 130, 1987

Table 15. Plectiscidea canaliculata (Forster). For abbreviations, see table 2. Van Rossem: Western Palaearctic Oxytorinae 77

Table 17. Plectiscidea monticala (Forster). For abbreviations, see table 2. 78 Tijdschrift voor Entomologie, deel 130, 1987

Table 19. Plectiscidea collaris {Grzvenhorst) Van Rossem: Western Palaearctk Oxytorinae 79

Table 2Q. Plectiscidea nava (Förster). For abbreviations, see table 2. 80 Tijdschrift voor Entomologie, deel 130, 1987

Table 11. Plectiscidea crassicornis (Forster). For abbreviations, see table 2. Van Rossem: Western Palaearctic Oxytorinae 81

Table 24. Plectiscidea connexa (Förster). For abbreviations, see table 2. 82 Tijdschrift voor Entomologie, deel 130, 1987

Table 26. Plectiscidea mendica (Forster). For abbreviations, see table 2. Van Rossem: Western Palaearctic Oxytorinae 83

Table 28. Plectiscidea deterior (Förster). For abbreviations, see table 2. Tijdschrift voor Entomologie, deel 130, 1987

Table 30. Plectiscidea terehrator Van Rossem: Western Palaearctic Oxytorinae 85

Table 32. Plectiscidea subangulata (Förster). For abbreviations, see table 2. 86 Tijdschrift voor Entomologie, deel 130, 1987

Table 34. Plectiscidea humeralis (Forster). For abbreviations, see table 2. Van Rossem: Wester7i Palaearctic Oxytorinae 87

Table 35. Plectiscidea (Fugatnx) communis (Förster). For abbreviations, see table 2. Tijdschrift voor Entomologie, deel 130, 1987

ra, 1400—2000 m (coll. Van Rossem). Netherlands: 6 Since my revision of the type material of Eu-

9, Asperen (Prov. Zd. Holl.), 3.ix.l972; l.vi, 30.vii, sterinx in 1982 I have come to the conclusion 9.vm, 7. XX, 10.ix.1973; 1 9, Nunspeet, 10.vii.l975; 4 that the diverging characters of the species al- 9, Ede, 22. V, 26. v, 6. vi, 12. vi. 1971 (Dutch specimens low the introduction of six subgenera, viz., Eu- coll. Zwakhals). Sweden; 2 9, Järnavik (Blekinge) sterinx Förster, 1868; Catomicrus Thomson, 12—20. vii. 1972; Orebro Län, Klysna-Norrberga, 1888; Ischyracis Förster, 1868; Divinatrix sub- S.vii—2.viii.l979 (coli. Van Rossem). genus novum; Dallatorrea Ashmead, 1902; and Distribution. — Widely spread in the Western Pal- Holomeristus Förster, 1868. A tentative key to aearctic Region. the subgenera follows here.

Gnathochorisis dentifer (Thomson). Austria: 2 6 1. Tyloids absent 2 Tirol, Aschbach, 1400 m, 6.vii.l976 and 1 9 — Tyloids present 3 18.viii.l975; 3 9, Kärnten, Himmelberg, 1100 m 2. Apophyses absent. Front wing not longer 14.vii.1979, ll.viii and 15.viii.l980; 1 d, Kärnten than 3.0 mm. Eyes not hairy Bodental, 1100 m, 13. vii. 1981 (all coli. Zwakhals) Germany: 4 9, Bechtaler Wald (Stegen-Wittental), Subgenus Eusterinx Förster 5.x & 2—9.XÌ.1984; 29.V.1985; no locality: 7.ix.l984 — Apophyses weakly developed or well de-

(leg. & coli. H. Hilpert). Netherlands: 1 â 3 9, Aspe- veloped. Front wing in most specimens lon- ren (Prov. Zd Holl.), 3.VÌ, 15.vi, l.ix.l972; 4 â, Ede, ger than 3.0 mm. Eyes often hairy 22.V, 26.V, 3.VÌ, 6.V1.1971 (coll. Zwakhals). Subgenus Catomicrus Thomson 3. Front wing without areolet 4 Distribution. — Widely spread in the Western Pal- — Front wing with areolet 5 aearctic Region. 4. Strong apophyses present. Front wing without areolet and in most specimens lon- Checklist of Gnathochorisis species ger than 3.0 mm. Sixth flagellar segment 1. Gnathochorisis flavipes Förster, 1871 : 113. with a tyloid, a concave polished area Gnathochorisis terebrata Strand, 1918: 159. Subgenus Ischyracis Förster The holotype has the following labels: Igna- — Apophyses absent. Front wing without lina (Litauen), viii.1916, leg. W. Horn; G. te- areolet and in most specimens not longer rebrata m. 9 Strand det. Typus (Institut für than 3.0 mm. There is a tyloid on sixth fla- Pflanzenschutsforschung Kleinmachow, gellar segment or the sixth and seventh fla- Eberswalde). The type agrees with G. flagellar segments are flattened vipes. Subgenus Eusterinx Förster Gnathochorisis terebrata; Oehlke, 1963: 409. 5. Second and third tergites in the female and Laepserus flavipes; Van Rossem, 1980: 115. tergites two, three, four and five in the male 2. Gnathochorisis dentifer (Thomson, 1888: divided by a conspicuous transverse suture. 1288)(comb.n.). The two parts of the tergites have a differ- Laepserus dentifer; Van Rossem, 1980: 118. ent sculpture . . Subgenus Divinatrix novum 3. Gnathochorisis crassulus (Thomson, 1888: — Tergites of female and male without suture 1289) (comb.n.). 6 Laepserus dentifer f. crassulus; Van Rossem, 6. Eyes crassate. Apophyses exceptionally 1980:119. strong. Ovipositor straight, 0.12—0.14 of 4. Gnathochorisis xanthocephalus (Strobl, 1903: length of front wing. (I have seen only two 113) (comb.n.). females of this subgenus) Laepserus xanthocephalus; Van Rossem, Subgenus Dallatorrea Ashmead 1980: 120. — Eyes normal. Apophyses not developed. 5. Gnathochorisis restrictus (Van Rossem, 1980: Ovipositor somewhat upcurved, 0.19—0.25 121) (comb.n.). of length of front wing (straight in E. aqui- Laepserus restrictus; Van Rossem, 1980: 121. lonigena). Tyloids on flagellar segments six and seven, a longitudinal carina; or on seg- Revision of the genus Eusterinx Förster ments six to eleven, laterally somewhat

Eustermx Förster, 1868: 172. concave and polished and with a carina Subgenus Holomeristus Förster Eusterinx; Förster, 1871 : 107— 109. Eusterinx; Aubert, 1968: 39. Key to the females Eustermx; Townes, 1971 : 202—204. £«iterz«x; Van Rossem, 1980: 131—132. (The females of E. minima and E. tartarea are Eusterinx; Van Rossem, 1982: 154— 169. unknown) Van Rossem: W^estem Palaearctic Oxytorinae 89

1. Eyes convergent towards clypeus 2 First to fourth tergite coriaceous. Oviposi- — Eyes not convergent towards clypeus .... 7 tor 0.10—0.14 of length of front wing 2. Apophyses of propodeum absent 3 E. (Dallatorrea) circaea Van Rossem — Apophyses of propodeum present 4 7. Front wing without areolet. Eyes without 3. Second and third tergite with different hairs. Head square, vertex deep. Tip of sculpture of proximal and distal half. These mandible twisted and with a sharp upper areas separated by a groove. Ovipositor and lower tooth. Second tergite slightly

0. 1 5 of length of front wing striated or with some rough longitudinal .... £. (Divinatrix) inaequalis Van Rossem sculpture or second tergite coriaceous or — Second and third tergite without groove. polished. Some specimens have conspicu- Ovipositor 0.22 of length of front wing .... ous thyridia. Length of front wing \.7—3.0 E. {CatomicTHs) pusilla (Zetterstedt) mm Subgenus Eusterinx 4. Eyes conspicuously hairy, converging to- — Front wing with areolet 8 wards clypeus. Notauli meetmg, with a 8. Malar space very narrow, 0.18 of width weak carina from pronotal margin. Scutel- face. Postannellus slender 4.5 times as long lum striated. Apophyses somewhat devel- as the apical width. Notauli not meeting on oped. Mesopleurum polished. Front wing mesoscutum. Propodeum without apo- with areolet not closed. Hind femur 6.3 physes. Second tergite proximally with times as long as wide. First to fourth tergite some longitudinal striation, apical half pol- coriaceous. Ovipositor 0.28 of length of ished. Ovipositor 0.14—0.19 of length of front wing front wing E. (Holomeristus) .... E. (Catomicrus) disparilis Van Rossem aquilonigena Van Rossem — Eyes without hairs or with inconspicuous — Malar space wide, 0.25—0.40 of width of setae. Strong apophyses present 5 face 9 5. First and second tergite with striation. 9. Malar space 0.23—0.25 of width of face. Third tergite with weaker striation. Strong Postannellus 3.0—5.0 times as long as the apophyses present. Ovipositor 0.20 of apical width. Notauli meeting on center of length of front wing mesoscutum. Propodeum without apo- E. {Ischyraas) bispinosa (Strobl) physes. Second tergite coriaceous and with — First to fourth tergite coriaceous 6 longitudinal striation. Ovipositor upcurved,

6. Eyes exceptionally large and convex, inner about 0.20—0.26 of length of front wing . . . margins strongly converging towards cly- .... E. {Holomeristus) tenuicmcta (Forster) peus, leaving a very narrow face. Notauli — Malar space wide, 0.40 of width of face. strong, meeting, restricting the median Postannellus 3.3 times as long as the apical lobe. The apical region of the median lobe width. The apical transverse carina some- with strong longitudinal sculpture. Propo- what lamelliform and with weak apophyses. deum with all carinae and robust, flattened Second tergite coriaceous. Ovipositor apophyses. Mesopleurum polished, with somewhat upcurved, 0.23—0.25 of length some longitudinal sculpture medially. Legs of front wing long and slender, including coxae orange in E. {Holomeristus) refractaria Van Rossem colour. Hind coxae for the greater part with rough sculpture. First abdominal segment Key to the males very slender, with long petiole. Postpetiole (Males of the Subgenus Eusterinx are tentatively apically with some striation. First to fouth included; males of E. {Catomicrus) disparilis and tergite coriaceous. Ovipositor 0.12 of E. {Dallatorrea) circaea are unknown) length front wing

E. (Dallatorrea) armata Ashmead 1. Front wing without areolet 2 — Eyes not excessively large and convex, con- — Front wing with areolet 9 verging to clypeus. Mesoscutum without 2. Propodeum with strong apophyses. First, special characters. Propodeum with all cari- second and third tergites striated. Sixth fla-

nae and robust, flattened apophyses. Meso- gellar segment with a tyloid, which is a con- pleurum polished, with some longitudinal cave polished area sculpture medially. Legs slender, brownish. E. {Ischyracis) bispinosa (Strobl) Hind coxa with rough sculpture. First ab- — Propodeum without apophyses 3 dominal segment slender, with long petiole. 3. Second tergite with conspicuous thyridia. .

90 Tijdschrift voor Entomologie, deel 130, 1987

Tyloids absent 4 E. {Eusterinx) argutula Forster — Second tergite without, or with indistinct 9. Apophyses of propodeum present. In £. thyridia 6 refractaria the apical transverse carina 4. Medial part of ciypeal margin somewhat somewhat lamelliform, thus developing risen, with a pair of very weak tubercles. weak apophyses 10 Postannellus 2.8 times as long as apically — Apophyses of propodeum absent 11 wide. Three flagellar segments after the 10. Apophyses rather strong. Antenna without postannellus about 0.71 of length of postan- tyloids. Hind femur stout, 3.8 times as long nellus. Second tergite finely striated. Thyri- as wide. First and second tergites entirely dia conspicuous coriaceous. Third tergite proximally coria- E. {Eusterinx) jugorum (Strobl) ceous — Medial part of clypeus not risen and with- E. {Catomicrus) tartarea Van Rossem out tubercles. Flagellar segments after the — Apophyses weak, caused by the somewhat postannellus short 5 lamelliform shape of the apical transverse 5. Postannellus 2.0 times as long as apically carina. Flagellar segments six and seven Wide. Flagellar segments after the postan- with a tyloid, a longitudinal carina. Hind nellus short less than 0.71 of length postan- femur stout. First, second and proximad nellus. Setae of flagellar segments erect. half of third tergite coriaceous. Tergites Thyridia in basal corners of second tergite two, three and four with narrow apical conspicuous, yellow in colour. Second ter- margin yellow gite basally with some coriaceous sculpture. E. {Holomeristus) refractaria

Following tergites polished. (There is no Van Rossem

male in Förster's material, it is inserted ten- 1 1 Second and third tergites with a transverse

tatively) . . E. {Eusterinx) obscurella Forster groove (groove weak on fourth tergite). — Postannellus 3.0 times as long as apically The proximal and distal regions of these wide. Flagellar segments after the postan- tergites with a difference in sculpture nellus less than 0.71 of length postannellus. .... E. (Divinatrix) inaequalis Van Rossem Setae of flagellar segments erect. Thyridia — Second and third tergites not so 12 of second tergite less conspicuous. Second 12. Tyloids on flagellar segments six to eleven. tergite with longitudinal striation, the distal The flagellar segments flattened, without margin and the following tergites polished setae. Second tergite for the greater part E. {Eusterinx) subdola Forster with longitudinal striation. The thyridia 6. Flagellum without tyloids 7 visible. The third tergite with some striation — Flagellum with tyloids 8 E. {Holomeristus) aquilonigena 7. Eyes hairy. Hind femur notably slender, 7.4 Van Rossem') times as long as wide. Second tergite — Tyloids on flagellar segments six to eight or striated and with coriaceous sculpture. Pro- six to nine. The flagellar segments flattened

podeum with indistinct apophyses. Length and with a carina. Thyridia not visible . . 13 of front wing about 3.8 mm 13. Second tergite and front part of third tergite E. (Catomicrus) pusilla (Zetterstedt) with longitudinal sculpture. Thyridia not — Eyes not hairy. Second tergite polished, or visible. Tyloids on flagellar segments six to with indistinct coriaceous sculpture. Propo- eight. Clypeus impressed deum without apophyses. Length of front E. {Holomeristus) tenuicincta Forster

wing about 1 .9 mm — Second and third tergites coriaceous. Ty- .... E. (Eusterinx) pseudoligomera Gregor loids on flagellar segments six to nine. Cly- 8. Antenna with tyloid on sixth flagellar seg- peus not impressed ment. Second tergite polished or proximally E. {Holomeristus) minima Strobl somewhat coriaceous. In some specimens the second tergite weakly striated Subgenus Eusterinx Forster

E. [Eusterinx) oligomera Forster Eusterinx¥örsitT, 1868: 172. — Antenna with tyloids on flagellar segments Type-species: Eusterinx oligomera Förster. six and seven (the second tyloid difficult to see). An indication of a tyloid on segment

eight. The flagellum more robust than in E. ') The separation of £. aquilonigena and E. tenuicinc-

{Eusterinx) oligomera ta is difficult without females from the same locality. —

Van Rossem: Western Palaearctic Oxytorinae 91

Front wing 1.7—3.0 mm long. Males with ridia absent in E. argutula one tyloid on sixth flagellar segment, and either E. {Eusterinx) oligomera Förster and E. two or none on flagellar segments six and seven. {Eusterinx) argutula Förster

Head square, vertex deep. Ratio gena-width :

eye-width = 7 : 4 (7 : 6) or 1 : 1. Mesoscutum Eusterinx (Eusterinx) jugorum (Strobl)

strongly convex, notauli varying from very Hemiteles pseudominutus var. jugorum Strobl, 1900: weak to strong. Propodeum with all carinae. In 243—244.

some specimens the costula is absent. Front wing without areolet. Nervellus reclivous. First Characteristics of the holotype (label partly gastral segment slender, spiracles at about 0.5 of illegible, 12/8, Styriae Alp Strobl, holotype label length. In E. obscurella and E. subdola the thy- of Horstmann, 1971): Male. Front wing 2.44 ndia are conspicuous, in other species weak or mm long. Medial part of clypeal margin some- absent. Second tergite in most species polished, what risen, with a pair of very weak tubercles (a occasionally somewhat coriaceous, seldom character not found in other species of the sub- striated. The ovipositor relatively long, 0.15 genus). Face finely coriaceous. Malar space 0.22 of length front wing, rather wide or wide. Postannellus 2.8 times as long as apically somewhat club-shaped and with long erect hairs wide. Tyloids absent. Three segments coming on sheath. after the postannellus about 0.71 of the length The subgenus includes five closely related of postannellus. Frons, vertex and gena pol- species. Males of different species are distin- ished. Gena wide. Pronotum indistinctly coria- guishable. The separation of the females of E. ceous. Mesoscutum polished, notauli strong, oligomera, E. argutula and E. pseudoligomera meeting and medially extended by a short fur- remains impossible. The followmg characters row. Propodeum laterally coriaceous. Meso- are not reliable: length of postannellus; the ratio pleurum polished, prepectal carina not reaching

of gena-width to eye- width; and the ratio of to the margin. Coxae and all other parts of the ovipositor length to length of hind tibia. legs conspicuously brown. Hind coxae coriaceous. Caster fuscous. First tergite coriaceous, The males are inserted into the general key to spiracles protruding. Second tergite finely Eusterinx males. The following key to the fe- striated. Thyridia conspicuous. The apical mar-

males is tentative. gin of the second tergite yellowish. Third tergite indistinctly coriaceous. The apical tergites pol- Key to the females ished. (The female of E. jugorum (Strobl) is unknown) The holotype from the "Kalbling" (2000 m) (Austria) is the only extant specimen. 1. Second tergite with conspicuous thyridia in proximal corners 2 Remark. — A female syntype (Horstmann, — Second tergite without or with weak thyri- 1971) oî Hemiteles pseudominutus Strobl, 1900, dia 3 has no original label. The specimen belongs to

2. Ratio gena-width : eye-width = 7:5 or Eusterinx {Eusterinx).

7 : 6. Postannellus 3.0 times as long as As the identification of the females of this

wide. Second tergite polished or with some subgenus is still uncertain, the specimen was la- coriaceous sculpture belled: Eusterinx {Eusterinx) species. E. {Eusterinx) obscurella Forster Postannellus 3.0 times as long as apically

— Ratio gena-width : eye-width =1:1. Post- wide. Spiracles of first gastral segment at 0.41 of annellus 4.5 times as long as wide. Second the length of the segment. Second and following tergite with longitudinal striation or weakly tergites polished. Ovipositor 0.18 of the length striated and somewhat coriaceous of front wing.

E. {Eusterinx) subdola Forster According to Horstmann "jugorum" is de- 3. Notauli weakly indicated directly behind rived from the Latin word "jugum" which mesoscutal margin. Second tergite polished, means "pass, narrow passage in mountains". thyridia weak .... E. {Eusterinx) pseudoligomera Gregor Eusterinx (Eusterinx) oligomera Förster Eusterinx — Notauli stronger, running towards centre of oligomera Förster, 1871 : 109. Eusterinx mesoscutum. Second tergite polished. Thy- oligomera; Townes, 1971 : 202. —

92 Tijdschrift voor Entomologie, deel 130, 1987

Eusterinx oligomera; Van Rossem, 1982: 163— 164. segment eight. The flagellum more robust than in E. oligomera. Mesoscutum convex, polished, Characteristics of the female: Front wing 1.7 notauli present but weak. Second tergite mm long. Clypeus flat, 2.0 times as wide as vaguely coriaceous, other tergites polished. long. Malar space wide, 0.3 of width of face, somewhat coriaceous. Face coriaceous, shghtly Distribution. — The Aachen Region (Forster col- convex with widely placed long hairs. Eyes lection). Italy: Dolomites (1300 m); Judikari Alpen (1720 m) (Haeselbarth collection). small, broadly elliptic. Ratio gena-width : eye- width = 7:4. Head square, frons and vertex Eusterinx (Eusterinx) pseudoligomera Gregor about 0.5 deeper than wide. Frons vaguely co- Eusterinx pseudoligomera Gregor, 1941: 8. riaceous. Vertex polished. Postannellus 3.3 Eusterinx pseudoligomera; Van Rossem, 1982: 168 times as long as wide. Mesoscutum polished, 169. notauli present. Propodeum coriaceous, with all carinae. Nervulus distad of basal vein. Areolet The males of this species can be tentatively absent. Nervellus reclivous. Lower part of distinguished from males of other species by mesopleurum coriaceous. Middle and hind legs, lack of antennal tyloids and by the vagueness of including tarsi, with long, subadpressed hairs. the thyridia; but females cannot be distin- First tergite coriaceous, spiracles at 0.4 of guished from those of other species, except by length. Apex of first sternite at 0.56 of length. their association in the field with males. Ovipositor club shaped, 0.12 of length front wing. Characteristics of the female: Front wing 1.9 Characteristics of the male: Front wing 1.7 mm long. Ratio gena-width : eye-width =1:1. mm long. A lateral tyloid on sixth flagellar seg- Malar space wide. Pronotum coriaceous. Meso- ment, somewhat flattened and pohshed. Prono- scutum pohshed, notauli obsolete. Propodeum tum polished, epomia weak. Mesoscutum pol- with all carinae. Mesopleurum coriaceous. First ished, notauli present. Propodeum weakly co- tergite coriaceous, median dorsal carinae pre- riaceous, with all carinae, except costula. Legs sent. End of first sternite at 0.69 of length of with close, long hairs. Second gastral segment segment. Second tergite pohshed, with weak variable in sculpture, from pohshed to some- thyridia. Ovipositor 0.21 of length of front what coriaceous. Striated sculpture may occur. wing. Characteristics of the male: Front wing 1.9 Distribution. — Apart from the Forster specimens mm long. Ratio gena-width : eye-width =1:1. from the Aachen region, the species occurs at rather Antenna without tyloids. Notaulus short. Up- striking altitudes (1200— 1300 m) in Austria and Italy (Dolomites). per half of mesopleurum polished, lower half somewhat coriaceous. Spiracles of first gastral segment at 0.5 of length of segment. End of first Eusterinx (Eusterinx) argutula Forster sternite at 0.7 of length of segment. First tergite

Eusterinx argutula : Forster, 1871 108 coriaceous, the others polished. Gaster rather Eusterinx argutula; Van Rossem, 1982: 164 165. — depressed.

Characteristics of the female: Front wing Distribution. — Czechoslovakia, the Gregor type 2.5—3.0 mm long. Ratio gena-width : eye- material is from Moravia, Ubusîn. Austria, Reiter width = 1:1 (or 8 : 7). Mesoscutum rather Alp, 1600 m; St. Haus, 1200 m. Germany, strongly convex, polished, notauli present but Lippoldshausen. faint. All tergites polished. Thyridia obsolete. There is no female in the Forster type materi- Eusterinx (Eusterinx) subdola Forster al. Eusterinx subdola Förster, 1871 : 108. Characteristics 2.6 of the male: Front wing Eusterinx subdola; Aubert, 1968: 39. mm long. Anterior tentorial pits large. Malar Eusterinx subdola; Van Rossem, 1982: 165. space wide. Face, frons, vertex and gena pol- Hemiteles pseudominutus Strobl, 1900: 243. ished, vertex deep. Postannellus 3.5 times as

long as wide. Sixth flagellar segment flattened There is no female of this species in the on one side, seventh segment slightly flattened Forster collection. Recognition of the female re- and here the microscopical longitudinal ridges mains uncertain. (glumes) absent. An indication of a tyloid on Characteristics of the supposed female by Van Rossem: Western Palaearctic Oxytorinae 93

comparison with the male: Front wmg 2.6 mm Distribution. — The type locality is Aachen. Italy: long. Postannellus 4.5 times as long as wide. Ra- Trento, M. Bondone Cornetto, 1900—2100 m (coll. Haeselbarth). tio gena-width : eye-width =1:1. Second tergite with longitudinal striation. Ovipositor 0.2 of length of front wing. Subgenus Catomicrus Thomson Characteristics of the male: Front wmg 3.0 Catomicrus Thomson, 1888: 1291. mm long. No tyloids present. Postannellus 3.0 times as long as wide. Femora rather stout, hind Type-species: Tryphon pusillus Zetterstedt femur 4.3 times as long as wide. Second tergite {Catomicrus tnchops Thomson). with large thyridia and longitudinal sculpture. Other tergites polished, with apical margins yel- Characteristics: Front wing 2.8—3.5 mm low and rather long adpressed hairs. long. Maies without tyloids. Eyes converging to Characteristics of the lectotype of Hemiteles clypeus in females, in males not converging. pseudominutus Strobl. Lectotype label of Eyes hairy in some species. Notauli strong and Horstmann, 1971. Male. Front wing 2.6 mm meeting in some species. Apophyses of propo- long. Postannellus 2.8 times as long as wide. deum weakly or strongly developed. Front Second tergite with longitudinal sculpture. wing with or without closed areolet. Ovipositor Hind femur 4.0 times as long as wide. 0.22—0.28 of length of front wing. I have placed two species in this subgenus.

Distribution. — The type locality is Lousberg (Aa- The single female of E. disparais is inserted ten- chen). I saw specimens from Austria, Tirol, 1400 m. tatively. Czechoslovakia, near Prague. Germany, Oberbayern. Italy, Bolzano Region. Netherlands, Asperen. Swe- den, Skâne; Lappland. Eusterinx (Catomicrus) pusilla (Zetterstedt) Tryphon pusillus Zetterstedt, 1838: 385. Male holotype.

Eusterinx (Eusterinx) obscurella Förster Catomicrus trich ops Thomson, 1888: 1291.

Eusterinx trichops; : Eusterinx obscurella Förster, 1871 : 108. Townes, 1971 203. Eusterinx obscurella; Aubert, 1968: 39. Eusterinx pusilla; Van Rossem, 1982: 159— 160. Eusterinx obscurella; Van Rossem, 1982: 167— 168. Characteristics of the female: Front wing 2.8 Characteristics of the type female from the mm. Malar space absent, eye margin almost type locality Aachen: Front wing 2.5 mm long. touching clypeal margin. Face narrow, polished, Postannellus 3.0 times as long as apically wide. eyes strongly converging to clypeus. Frons, ver-

Ratio gena-width : eye-width = 7:6. Thyridia tex and gena polished. Antenna rather short, to- on second tergite conspicuous, more outlined wards distal end gradually somewhat widening. by their colour. Ovipositor 0.15—0.22 of the Epomia distinct. Mesoscutum polished, notauh length of front wing. present, with short carina on pronotal margin. Characteristics of the male. The male was not Propodeum coriaceous, with all carinae. No described by Förster. The following description apophyses present. Mesopleurum somewhat co- is based on males tentatively considered to be of riaceous, prepectal carina not reaching the mar- this species. They are from Monte Bondone gin. Front wing with areolet not closed. Legs Cornetto, 1900—2100 m. Dolomites, Italy. slender. Hind femur notably slender, 6.6 times Front wing about 1.9 mm long. No tyloids pre- as long as wide, with close subadpressed hairs. First, sent. Ratio gena-width : eye-width = 6:4. second and third tergite coriaceous. Api- Eyes comparatively small, roundish. Pedicel cal margins of tergites two, three and four yel- proportionally large, sHghtly shorter than post- low. Ovipositor 0.22 of length of front wing. annellus. Apex of scutellum with rough sculp- Characteristics of the male: Front wing 3.8 ture. First tergite with longitudinal striation. mm. Malar space 0.27 of width of face. Eyes Large thyridia in proximal corners of second hairy and with inner margins parallel. Face, tergite, yellow in colour, some coriaceous frons, vertex and gena polished. Face with erect sculpture between them. Rest of tergite and hairs. Epomia distinct. Notauli strong, with a other tergites polished. carina along inner edge. Propodeum without —

94 Tijdschrift voor Entomologie, deel 130, 1987

distinct dorsal face. Apical transverse carina ly- fourth tergite yellowish. Ovipositor 0.28 of ing towards distal edge of propodeum, with length of front wing. weak apophyses. Prepectal carina present. Hind Male unknown. femur slender, 7.4 times as long as wide. First Distribution. — Only the holotype from Sweden, tergite coriaceous, medially convex, median Messaure (Lapland) is extant. dorsal carinae weak. Second tergite coriaceous tergite proximal- and with weak striation. Third Subgenus Ischyracis Forster ly coriaceous. IschyracisYörsier, 1868: 175 Type-species: Catomicrus alpigenus Strobl. Distribution. - Sweden. Ischyracis; Perkins, 1962: 431 —432.

Ischyracis; Townes, 1971 : 203. Eusterinx (Catomicrus) tartarea Van Rossem Eusterinx tartarea Van Rossem, 1982: 159. Front wing 3.1 —3.6 mm long, without areolet. In the female the eyes converging to the cly- Characteristics of the male: Front wing 3.8 peus, in the males not so. Sixth flagellar segment mm. Clypeus 1.3 times as wide as long, pol- of the male with a tyloid, a concave pohshed ished, somewhat convex. Face polished. Malar area. Strong apophyses present. space wide, 0.3 of width of face, with a groove. I have placed only one species in this subge- Frons, vertex and gena polished, with rather nus, viz. E. alpigena (Strobl) = Eusterinx bispt- long subadpressed hairs. Antenna without ty- nosa (Strobl), which species was designated by loids. Epomia present. Notauli rather strong, Perkins as the type species of Ischyracis. with a short carina. Apex of scutellum striated. Propodeum with all carinae and conspicuous Eusterinx (Ischyracis) bispinosa (Strobl) apophyses. Prepectal carina to the middle of Hemiteles bispinosus Strobl, 1900: 234—235. pronotal margin. Front wing with areolet. Hind Ischyracis bispinosus; Aubert, 1970. 279. coxae with rough coriaceous sculpture. Hind fe- Eusterinx bispinosa; Horstman, 1974: 53. mur stout. First tergite coriaceous, median dor- Catomicrus alpigenus;SxToh\, 1903: 116— 117. sal carinae absent. Lateral dorsal carina indi- Eusterinx alpigena; Van Rossem, 1980: 131 — 132. cated. Second tergite for the greater part coriaceous, third tergite proximally coriaceous. Characteristics of the female: Front wing Second tergite with a carina from spiracle to 3.1 —3.6 mm long. Eyes convergent to clypeus, proximal margin. All tergites with rather long not touching clypeal margin. Eyes not hairy. adpressed hairs. Malar space present. The clypeal fovea situated Female unknown. in a conspicuous depression between eye and clypeus. Face and frons polished. Strong notauli Distribution. — Only the male holotype from Italy, meeting in the middle. Front wing without St. Peter, Ahrntal, 1350 m, Südtirol, is extant (coll. areolet. Propodeum with strong apophyses. Haeselbarth). Hind coxa black, coriaceous. Hind femur 5.0 times as long as wide. Gaster fuscous. First tergite with strong striation, second and third ter- Eusterinx (Catomicrus) disparilis Van Rossem gite with weaker striation. Ovipositor 0.2 of Eusterinx disparilis Van Rossem, 1982: 159. length of front wing. Characteristics of the male: Front wing 3.2 Characteristics of the female: Front wing 3.5 3.6 mm long. Eyes not convergent. Sixth flagel- mm. Malar space 0.3 of width of face. Face pol- lar segment with a tyloid, a concave poHshed ished, with widely placed setae. Eyes hairy, area. Notauli meeting in the middle. Propo- converging to clypeus. Frons, vertex and gena deum with strong apophyses. Front wing with- polished. Mesoscutum with adpressed hairs, no- out areolet. Hind femur robust, 4.4 times as tauli meeting, with weak carina from margin. long as wide, with conspicuous adpressed hairs. Scutellum striated. Propodeum coriaceous. Median sternal groove (mesolcus) deep. Gaster Apophyses somewhat developed. Front wing fuscous. Spiracles of first segment at 0.5 of with areolet not closed. Legs slender, hind fe- length. First tergite with rong striation, weaker mur 6.3 times as long as wide. First to fourth on second and third tergite. Tergites two to four tergite coriaceous. Apical margin of second to coriaceous. Van Rossem: Western Palaearctic Oxytorinae 95

Distribution. — Austria, Admont; Natterriegel. Distribution. — Italy, Riva S. Garda (Haeselbarth). Germany, Thüringen, Blankenburg (Schmiede- Netherlands: Asperen (Prov. Zuid-Holland) (Zwak- knecht); Lippoldshausen (Haeselbarth). Netherlands, hals). U.S.A.: Spring Br. Pa., 25.viii.1945 (leg. H. K. Asperen (Prov. Zuid-Holland) (Zwakhals). Townes).

Divinatrix subgenus novum Subgenus Dallatorrea Ashmead Front wing 3.0—3.20 mm long, with areolet. Dallatorrea Ashmead, 1902: 205.

Eyes strongly convergent to clypeus in the fe- Da!latorrea;lownes, 1971 : 203. male; slightly converging in the male. Mesoscu- Type-species: Dallatorrea armata Ashmead. tum with deeply impressed notauHces. Apo- physes absent. Second and third tergites in the Front wing 5.2—6.0 mm long, with areolet. female and tergites two to five in the male divid- Eyes crassate, converging, or strongly converg- ed by a conspicuous transverse suture. The two ing to clypeus. Antenna short. Mesoscutum parts of the tergites with a different sculpture. with close adpressed hairs. Notauli strong.

Ovipositor 0. 1 5 of length of front wing. Apophyses very robust, flattened. Ovipositor

There is only one species in this subgenus, E. 0.11 —0.14 of length of front wing. inaequalis, which is the type-species. "Divina- There are two species in this subgenus. I have trix" is the Latin for "prophetess". examined only females.

Eusterinx (Divinatrix) inaequalis Van Rossem Eusterinx (Dallatorrea) armata (Ashmead)

Eustermx inaequalis Van Rossem, 1980: 132. Dallatorrea armata Ashmead, 1902: 205.

Eusterinx armata; Townes, 1971 : 203. Characteristics of the female: Front wing 3.2 mm long. Clypeus somewhat protruding, about Characteristics of the female: Front wing 5.2 as wide as long. Large eyes strongly convergent mm. Mandible turned in, upper tooth long and to clypeus, touching clypeal margin. Face and sharp, lower tooth minute. Clypeus protruding, frons polished. Scape and pedicel yellow. Pro- its width about 2.0 times the lower width of notum coriaceous. Mesoscutum with strong, face. Malar space absent. Eyes very large and deeply impressed notauli, meeting in the mid- strongly convex, with inconspicuous setae. In- dle, with conspicuous transverse ridges. Prono- ner margins of eyes strongly converging to cly- tal margin turnmg inwards towards notauli. peus. Width frons at ocelli more than 3.5 times Mesoscutum with adpressed hairs. Prepectal ca- the width of face at clypeus. Face, frons and rina reaching pronotal margin somewhat below gena polished. Antenna short. Occipital carina wing base. Propodeum completely areolated. closed. Epomia strong. Pronotum polished. Front wing with areolet. Front and middle Mesoscutum with close adpressed hairs. Notau- coxae yellow, hind coxa coriaceous, brown; li robust, meeting in centre of mesoscutum, hind femur 6.4 times as long as wide. The most bounding a convex median lobe. The distal zone conspicuous character is the sculpture of the of median lobe with strong, longitudinal second and third tergile. Proximal 0.75 part of wrinkles. Propodeum completely areolated with second tergite coriaceous and striate, distal part strong carinae. Apophyses very conspicuous, polished. About 0.5 part of third tergite striated, robust and flattened. Mesopleurum polished, distal part polished. The two halves separated with some longitudinal wrinkles medially. by a conspicuous suture, almost giving the im- Front wing with areolet. Nervulus proximal of pression of two tergites. A weak indication of basal vein. Nervellus vertical, discoidella absent. the suture on tergite four. Ovipositor 0.15 of Legs long and slender, including the coxae yel- length of front wing. lowish to orange in colour. Hind coxae with Characteristics of the male: Front wing 3.0 rough sculpture, about 3.0 times the size of mm long. Eyes slightly converging, not touch- middle coxae. Hind femur strongly developed. ing clypeal margin. Malar space present. The Fringe at apex of hind tibia minute. Petiole long. tergites two, three, four and five with the same First tergite coriaceous, with some longitudinal conspicuous character as the female's second striation on post-petiole. Spiracles about in the and third tergites. middle. Second, third and fourth tergite proxi- — —

96 Tijdschrift voor Entomologie, deel 130, 1987 mally coriaceous. Ovipositor 0.12 of length of Front wing 2.6—4.0 mm long, with areolet. front wing. The ovipositor with a long and slen- Malar space wide. Males with tyloids on sixth to der tip. eleventh flagellar segments, flattened and slightly concave areas with a longitudinal carina on Material examined. — 19, Corvallis (Oregon, one side or only a longitudinal carina. Eyes not U.S.A.), 1 1 July 1978, from collecdon Townes. convergent to clypeus. Notauli present. Propo- deum completely areolated. (£. {Holomeristus) Eusterinx (Dallatorrea) circaea Van Rossem refractaria has weakly developed apophyses). Eusterinx circaea Van Rossem, 1982: 157— 158. Front wing with areolet. Thyridia weak. Ovipo- sitor somewhat upcurved, 0.14—0.26 of length Characteristics of the female: Front wing of front wing. 5.2—6.0 mm long. Mandible yellowish, the tip The subgenus includes four species. twisted. Upper tooth with sharp point, the lower tooth inside and less than 0.3 of length of up- Eusterinx (Holomeristus) per tooth. Clypeus convex, strongly protruding, tenuicincta (Förster) about as wide as long. Clypeus not distinctly Holomeristus tenuicinctus Förster, 1871 : 80—81. separated from face by a groove. Malar space Holomeristus tenuicinctus; Aubert, 1970: 274. wide, about 0.5 of width clypeus, with a groove Eusterinx tenuicincta; Townes, 1971 : 202. between eye and clypeus. Eye margins converg- Eusterinx tenuicincta; Van Rossem, 1982: 160— 161. ing to clypeus. Face and frons polished. Ocelli robust. OOL : POL = 6:4. Vertex narrow, Characteristics of the female: Front wing 3.5 occiput steeply sloping behind ocelli. Occipital mm long. Eyes not convergent to clypeus. Cly- carina closed. Antenna slender, but short. Epo- peus yellow, impressed, 1.8—2.1 times as wide mia present. Mesoscutum polished, with close as long.Mandible twisted, with a single tooth. adpressed hairs. Notauli strong, with a sharp Malar space 0.23—0.25 of width face. Gena notch on margin. Scutellum and postscutellum wide. Scape and other parts of antenna yellow. rugulose. Propodeum laterally with conspicu- Postannellus 3.0—5.0 as long as the apical ous hairs. Median longitudinal carinae around width. Last joint of antenna large and inflated. area superomedia strongly developed. Apo- Pronotum polished, epomia long. Mesoscutum physes very robust, flattened. Mesopleurum fuscous, polished, with conspicuous notauh polished, ventrolateral margin rugulose and coming together in centre of mesoscutum and with longitudinal striation. Front coxae yellow, with a short carina on their front side, meeting polished. Middle coxae with long hairs, ventral- the epomia. Propodeum with all carinae, apical ly rugulose. Hind coxae fuscous, with long transverse carina somewhat lamelliform, but not hairs, rugulose, dorsally with a polished concav- developing apophyses. Mesopleurum polished, ity towards trochanter. Front and middle legs prepectal carina extending to subtegular ridge. slender, hind femur robust, with rugulose sculp- Sternaulus not developed but indicated by weak ture. Hind tibia and tarsus slender. Claws of all ridges to edge of mesopleurum. Legs, including legs small. Front wing with areolet. First gastral coxae yellowish with brownish hind femur, tip segment with slender petiole, spiracles at 0.5 of of hind tibia and hind tarsus. Areolet present. length. End of first sternite at 0.7 of length. First tergite fuscous and with longitudinal stria- First tergite coriaceous, with longitudinal striation. Second tergite fuscous and striated, with tion. Second to fourth tergite coriaceous, apical pohshed yellow apical margin. Thyridia vague. margins polished, yellow. Ovipositor 0.11 Third tergite in some specimens with narrow 0.14 of length of front wing. front margin striated, the greater part brownish Male unknown. and pohshed, with yellow apical margin. Other tergites fuscous and polished. Ovipositor 0.20 Distribution. — The female holotype is from 0.26 of length of front wing, upcurved. Its Hochstadt, Oberbayern (Germany) (coll. Haesel- sheaths slender. barth). One female was collected in Italy, Prov. Bol- Characteristics of the male: Front wing 3.1 zano, Sarntal, 1250 m, 28. vi. 1976, leg. and coll. C. J. Zwakhals. 3.6 mm long. Mandible twisted, narrow, with a sharp point. The clypeus impressed, yellowish Subgenus Holomeristus Förster brown, with a marginal fringe of setae. Malar

Holomeristus Förster, 1868: 171. space 0.22 of width of face. All parts of head Type-species: Holomeristus tenuicinctus Förster. fuscous and polished. Gena wide. Postannellus Van Rossem: Western Palaearctic Oxytorinae 97

3.7 times as long as the apical width. Antenna hat upcurved, 0.23—0.25 of length of front yellow, with vague tyloids on flagellar segments wing.

6, 7 and 8. Pronotum fuscous, epomia long. Characteristics of the male: Tyloids, longi- Mesoscutum fuscous, polished. Notauli con- tudinal carinae, on flagellar segments six and spicuous, meeting in center with a short carma seven. Hind femur stout. First, second and on front side. Propodeum with all carinae. proximal part of third tergite coriaceous. Ter- Mesopleurum polished, prepectal carma to sub- gites two, three and four with narrow apical tegular ridge. Sternaulus absent, mdicated by margin yellow. weak ridges. Legs yellowish. Front and middle legs very slender. Areolet present, m some spec- Distribution. — The holotype is from Kytin (Bohe- imens not closed. First abdominal segment fus- mia), Czechoslovakia. Italy, Bolzano, Feldthurns, 1200 m. Sweden, Skane, Rostanga. cous, slender, about 3.25 times as long as the apical width. First tergite with longitudinal striation. Second tergite with longitudinal striation, Eusterinx (Holomeristus) minima (Strobl) fuscous, with polished apical margin brown. Holomeristus minimus Strobi, 1903: 119. Third tergite with longitudinal striation proxi- Eusterinx minima; Van Rossem, 1982: 161 — 162. mally, the remaining part polished. Of this species only the Strobl holotype male

Distribution. — The Palaearctic Region, including is extant. It has the tyloids on the flagellar seg- Japan. The Nearctic Region, including Alaska. ments six to nine. The sculpture of the second and third tergites is coriaceous. In E. tenuicincta Remark. — Mr. Andreas Zumdick (Kiel) these tergites have longitudinal striation. The

bred the species from Polyporus squamosus Fr. sculpture of the tergites is rather variable in Eu- collected in the Stifter Wald (near Kiel, Ger- sterinx and does not offer definite characters to many) between July and September 1983. The separate the species. mushrooms were infested with Diptera larvae: Mycetophilidae, Limoniidae and Muscidae. Characteristics of the holotype: Front wing 2.6 mm long. Clypeus slightly protruding and Eusterinx (Holomeristus) refractaria somewhat convex, 1.5 times as wide as long. Van Rossem Face, frons and vertex polished. Antenna with Eusterinx refractaria Van Rossem, 1982: 158. tyloids on flagellar segments 6—9. Notauli strong, meeting in middle of mesoscutum. Legs Characteristics of the female: Front wing 3.9 very slender, with long hairs. Prepectal carina to mm long. Eyes not convergent to clypeus. Cly- about 0.5 of pronotal margin. First, second and peus rather convex, polished, 2.0 times as wide third tergite coriaceous. as long. Face polished, with widely-placed fine Female unknown. punctures. Malar space wide, 0.4 of width face, with a distinct furrow. Frons, vertex and gena Distribution. — The holotype was collected in Austria, Styrian (Alpenwiesen Natterriegel). pohshed. Postannellus 3.3 times as long as wide. Alp des It is kept in the Strobl collection at Admont (Austria). Pronotum with distinct epomia. Mesoscutum with close adpressed hairs. Notauli strong, with a short carina from margin. Propodeum with all Eusterinx (Holomeristus) aquilonigena carinae, the apical transverse carina somewhat Van Rossem lamelliform and thus developing short apo- Eusterinx aquilonigena Van Rossem, 1982: 156— 157. physes. Mesopleurum polished, prepectal carina to about the middle of pronotal margin. Front Characteristics of the female: Front wing 3.0 wing with areolet. Nervellus vertical, not inter- mm long. Mandible twisted, with a sharp upper cepted. Front and middle coxae yellow; hind tooth. Lower tooth 0.5 of length of upper tooth. coxae brown, coriaceous; hind femur stout; Clypeus 1.5 times as wide as long, the apical claws of all legs strong. End of first sternite at margin with a close row of bristles. Malar space 0.6 of length of segment. First tergite without very narrow, 0.18 of width of face. Face pol- dorsolateral and median dorsal carinae. Second ished, with a row of erect hairs along inner mar- tergite and proximal part of third tergite coria- gins of eyes and with two rows medially. Frons, ceous. Following tergites polished. All tergites vertex and gena polished. Gena with two rows with apical margin yellow. Ovipositor somew- of subadpressed hairs. Scape subcyhndrical. 98 Tijdschrift voor Entomologie, deel 130, 1987

Epomia weak. Notauli with a fine carina on low, with a key to the males. Two new species their front side. The notauH not meeting in cen- are proposed: Helictes fabularis and H. incon-

ter. Propodeum with all carinae. Prepectal cari- gruens. The identity of the males could only be na not reaching the margin. Front wing with based on a single character namely the position areolet. Legs slender, including coxae, yellow. of the tyloids on the flagellar segments. I have First tergite coriaceous, medially convex. Sec- been unable to recognize the females. Only with ond tergite coriaceous and with longitudinal Helictes erythrostoma I have females and a male striation on front part, apical half polished. from the same locality and date. Thyndia lying near basal margin behind end of ventrolateral carina of first tergite. Other ter- Key to Helictes males gites polished. Ovipositor 0.14—0.19 of length (Postannellus counted as first segment) of front wing. Characteristics of the male: Front wing 3.1 1. Segment five of flagellum with a tyloid ... 2 mm long. Tyloids on flagellar segments 6— 11, a — Segment five of flagellum without a tyloid 3 6 flattened zone without setae. Second tergite 2. Tyloids on flagellar segments 5— — 7. Ter- more rough than in female, for the greater part gite two With vague microsculpture with longitudinal striation. Third tergite in front H. erythrostoma (Gmelin) 6 7 part with some longitudinal striation. — Tyloids on flagellar segments 5— — —8. Tergite two with microsculpture Distribution. — Sweden. Skane. H. conspicuus (Förster) 3. Tyloid on flagellar segment six Genus Helictes Haliday H. fabularis spec. nov. Helictes Haliday, 1838: 115. — Tyloids on flagellar segments 6—7— 8 or Myriarthrus¥öTsx.e.r, 1869: 172. 6—7—8—9 4

Myriarthrus; Förster, 1871 : 102. 7 4. Tyloids on flagellar segments 6— —8. Ter- Idioxenus Förster, 1868: 171. gite two polished or with indistinct micros- Idioxenus; Förster, 1871: 94. culpture H. borealis (Holmgren) MegaStylus; Holmgren, 1855: 129. — Tyloids on flagellar segments 6 7 8 9. MegaStylus {Helictes)\i:homson, 1888: 1312. — — — //e/zcre5;Strobl, 1903: 139. Tergite two polished H. incongruens spec. nov. Helictes; Townes, 1971 : 204. Helictes; Aubert, 1977: 148. Helictes erythrostoma (Gmelin) Ichneumon erythrostoma Gmelin, 1790: 2721. Type Ichneumon erythrostoma Gmelin, 1790, was destroyed. designated as the type-species of Helictes Hal- Plectiscus erythrostoma; Gravenhorst, 1829: 718. iday by Westwood in 1840. Gmelin's material is Cryptus {Helictes) fulvicornis Haliday, 1838: 115. lost. It is therefore to best follow Gravenhorst, Lectotype Fitton, 1976: 333. 1829, with respect to this species and to desig- Idioxenus mediator (Schiodte, 1838) sensu Förster, nate Gravenhorst's male as the neotype. 1871:95.

For no good reason Förster rejected the name Idioxenus inaequalis Förster, 1871 : 95. Helictes and introduced the genus Myriarthrus with Plectiscus erythrostoma Gravenhorst as the The type material of Ichneumon erythrosto- type. Förster did not see the Gravenhorst type ma Gmelin has been destroyed. This species is material. His other species in Myriarthrus be- named according to the revision by Graven- long to Megastylus Schiodte. Moreover Förster horst, 1829, whose specimen is regarded as the described the genus Idioxenus with Megastylus neotype. No original Gravenhorst label is pre- mediator Schiodte, 1838, as the type-species. sent. There are two existing labels; that of Au-

The Schiodte type specimen is a true Megastylus bert which says Helictes erythrostoma Gmel. (Townes, 1971; van Rossem, 1974). The lectotype of Förster's Idioxenus mediator is a male ') If Förster, 1871, designates as type-species for Idi- specimen of Helictes erythrostoma (Gmelin) oxenus gen. nov. a species that he cites in some such (sensu Gravenhorst). The other species placed 1) manner as Megastylus mediator Schiadte, 1838, the by Förster in Indioxenus also belong to Heli- type-species of Idioxenus is that which was before ctes. Förster, and not that named by Schiodte, and its name

Redescriptions of the Western Palaearctic is to be cited as Idioxenus mediator Förster, 1871. species of Helictes Haliday, 1838, are given be- (Int. Code, 1985, Art. 70c). Van Rossem: Western Palaearctic Oxytorinae 99 male (= conspicuus = inaequalis) and the neo- Characteristics of the female: Description type label of the present author. based on Haeselbarth's specimen from Lippoldshausen, 20. v. 1967, from the same lo- Characteristics of the neotype of Helictes caHty (A) as a male. Front wing 3.6 mm. Palpi erythrostoma: Male. Front wing 3.5 mm long. whitish. Face with microsculpture and ad- 6 Tyloids of flagellar segments 5— — 7. Tergite pressed hairs. Frons polished. Vertex and occi- two almost polished. The specimen is m bad put with very fine microsculpture and widely condition and glued on a square of mica. Right placed adpressed hairs. Temple polished, with- antenna missing beyond postannellus; left an- out hairs. Antenna yellowish brown, slender. tenna missing beyond segment eight. Right Postannellus long and slender, 8 times as long as middle leg and left hind leg missing. apical width. Pronotum with fine microsculp- Characteristics of the lectotype of Cryptus ture, epomia present. Mesoscutum with micros- (Helictes) fuhicornis Haliday. Labels: lectotype culpture, notauh almost obliterated. Scutellum label of Fitton, 1975. Ireland. Female (National with microsculpture, without margin. Propo- Museum of Ireland, Dublin). I have seen the deum with strong microsculpture. Median lon- lectotype. It is a reddish brown specimen. gitudinal carina strong down to apical trans- There is a male specimen of Idioxenus media- verse carina. Pleural carina present. Mesopleu- tor sensu Förster labelled as follows: a Förster rum with microsculpture and adpressed hairs on label "Aachen, male, 27 gl."; a box label front half up to apex of prepectal carina. Pre- ''mediator" Schiodte"; a label of Aubert "He- pectal carina not reaching margin. Tegulae lictes erythrostoma Grav. (= mediator auct. nee. whitish. Legs, including coxae, light brown. Schiodte)". Middle and hind coxae with close sculpture. Characteristics of Förster's specimen of Tergites one, two and three with microsculp- Helictes mediator: Male. Front wing 3.6 mm ture. Ovipositor not extending beyond subgeni- 6 long. Tyloids on flagellar segments 5— — 7. tal plate. Tergite two with fine microsculpture.

The lectotype of Idioxenus inaequalis is la- Material examined. —

100 Tijdschrift voor Entomologie, deel 130, 1987

Helictes conspicuus (Forster) culpture. First and second tergite with vague microsculpture. Idioxenus conspicuus Forster, 1 871 : 95.

Idioxenus inquilinus Forster, 1871 : 95. There are three paralectotypes {2 6 1 9), all

Idioxenus intricator Forster, 1871 : 95. from Lousberg. There are also specimens (8 â Idioxenus tetraglyptus Forster, 1871 : 95. 14 9), labelled by Förster Idioxenus tetraglyp- eltctes nigricoxus iirohl, 1903: 139. H tus, but these are not type specimens. The lectotype of Helictes nigricoxus Strobl, The holotype of Idioxenus conspicuus is la- 1903, is labelled: "Admont 18 August" and balled as follows: a Förster label "Aachen ê";a. bears the lectotype label of Aubert, 1977, who label "conspicuus Frst." and the holotype la- box identified this specimen as Helictes erythrosto- bel of the present author. ma Grav., male. I have not seen this specimen, but following the original description of Strobl, Characteristics of the holotype of Helictes viz. "das fünfte bis achte Geiszeiglied start aus- conspicuus: Male. Front wing 3.4 mm long. Face gerandet" I am inclined to place it under Heli- with microsculpture and adpressed hairs. Ver- ctes conspicuus. Strobl also noted: "nach Frst. tex, occiput and temple with adpressed hairs. Tab. gelangt man auf inquilinus" 6 7 Tyloids on flagellar segments 5— — — 8. Pro- notum with epomia and microsculpture. Mesos- Material examined. — The localities mentioned above are recapitulated. cutum with microsculpture and notauli present. not Germany: S , holotype of Idioxenus conspicuus, Aachen (coll. Förster, Propodeum with microsculpture down to posi- München); 2 S, Wiershausen (Niedersachsen), A, tion of apical transverse carina, the latter being 22.V.1966; 4 â, Lippoldshausen, A & B, 20.V.1967; 2 obliterated. A part of mesopleurum almost pol- S, Ziegenhagen (Hessen), Ac, 13.viii.l966; Aa, ished. Prepectal carina reaching not margin. 15.viii.l966; S, Münden (Hann.), 13.viii.l965; i, Upper part of hind coxae with somewhat more Glonn (Ober Bayern), 14.vii.l968; S, Weszling rough sculpture. Second tergite with fine micro- (Ober Bayern), 12.viii.l972; 2 d, Weszling, Hochs- sculpture. tadt (Bayern), 22. vi. 1974; S, Starnberg, Kerschlach (Bayern), 27.vii.1974; Sachsenkam, Kirchseefilz The lectotype of Idioxenus inquilinus is la- S, (Ob. Bayern), 19.vii.l972 (all leg. & coli. Haesel- belled with a Förster label "S, 30 gl. 5—8 aus- barth). Italy: i, TN, M. Baldo Bocca Navene, 1400 gebuchtet". Lectotype label of Aubert, 1977. m, 9.VÌÌ.1972; 6, Algund (Südtirol), 1800 m, A, Characteristics of the lectotype of Helictes in- 24. vii. 1966; male, Idrosee (Brescia), Vesta, 500 m, quilinus: Male. Front wing 4.0 long. Face mm 15.vi.1958; S, Campi, Riva s. Garda, 1400 m. E, with microsculpture. Tyloids on flagellar seg- 7.ÌX.1967 (all leg. & coll. Haeselbarth); 2 S, Sarntal 6 7 ments 5— — — 8. Mesoscutum, first and sec- (Bolzano), 1250 m, 26.vi.1976 & 30.vl (both leg. & ond tergite with microsculpture. coll. Zwakhals). Netherlands: 10 S, Ede (Prov. There are two paralectotypes from Lousberg Geldl), 28.ix.1970, 14.x, 17.x, 24.x, l.xi, 15.xi; (the type locality), probably a specimen of H. 12.vi.1971; i, Overveen (Prov. Noord-Holland), 9.VÌ.1974; 4 â, Asperen (Prov. Zuid-Holland) erythrostoma and also one female from Lous- i.v.1967; 18.V.1970; 24.V.1972; 27.V.1972; ê, Arkel berg. (Prov. Zuid-Holland), 10.viii.l967; S, Schelluinen, The holotype of Idioxenus intricator is la- 23.viii.1967; i, Hoornaar, ll.vii.l967. All material belled with a Förster label, 28 gl., "d, Lousberg leg. and coli. Zwakhals. Poland: S, Polanowice (10 12.10." and a holotype label of the present au- km N. of Wroclaw), 15.vii.l967, leg. W. J. Pulawski thor. (coll. van Rossem). Switzerland: S, Tarasp Lai Nair Characteristics of the holotype of Helictes in- (Gr.), Il.viii.l973 (coll. Haeselbarth); S, Bern, tricator: Male. Front wing 4.0 mm long. Face Delémont, 21. v. 1975, leg. R. T. Simon Thomas (coll. van Rossem). Sweden: S, Lapland m., 30.viii. Bhn with microsculpture. Tyloids on flagellar seg- 6 7 (Boheman) (in type series of Megastylus borealis ments 5— — —8. Mesoscutum, first and sec- Holmgr., Riksmuseet, Stockholm). ond tergite with microsculpture.

The lectotype of Idioxenus tetraglyptus is la- Distribution. — The species is widely spread in the belled with a Förster label, "(5, 25 gl., Lous- western Palaearctic Region. berg, 9.6." and a lectotype label of the present author. Helictes borealis (Holmgren)

Characteristics of the lectotype of Helictes te- Megastylus borealis Holmgren, 1855: 129. traglyptus: Male. wing 3.2 Front mm long. Face Idioxenus coxalis Forster, 1871 : 95. with microsculpture. Tyloids on flagellar seg- Idioxenus propinquus Forster, 1871 : 95. 6 7 ments 5— — —8. Mesoscutum with micros- Idioxenus invalidus Forster, 1871 : 95. —

Van Rossem: Western Palaearctk Oxytorinae 101

weak, not reaching the margin. Idioxenus variator Förster, 1871 : 95. Tegulae white. Megastylns (Heltctes) ptlicornis Thomson, 1888: 1312. Legs including coxae yellow. All tergites polished.

The lectotype of Megastylus boreahs Holm- The holotype of Idioxenus propinquus is la- gren (Riksmuseet, Stockholm) is labelled: "Lp belled with a Forster label "Aachen, S ". (= Lapland) m, S.viii. Bhn (= Boheman)". Lec- Characteristics of the holotype of Helictes totype label of the present author. propinquus: Male. Front wing 4.3 mm long. Ty- Characteristics of the lectotype of Heltctes loids on flagellar segments 6—7—8. Second ter- borealis: Male. Front wing 3.7 mm long. Palpi gite almost polished, some vague microsculp- and mandible yellow. Clypeus convex, yellow. ture present. Face coriaceous, with long, subadpressed hairs. The lectotype of Idioxenus invalidus has two Malar space wide, 0.26 of width of face. Eyes Forster labels: "Aachen, â, 30 gl"; "6— 8..." with setae. Tyloids on flagellar segments 6—7 (illegible) and the lectotype label of the present 8. Frons and vertex polished. Gena with long author. hairs. Occipital carina closed. Pronotum with vague sculpture, epomia present. Mesoscutum Characteristics of the lectotype of Helictes in- with adpressed setae. Notauh present. Propo- validus: Male. Front wing 3.4. mm long. Ty- 7 deum poHshed, carinae absent except for stubs loids on flagellar segments 6— — 8. second ter- of longitudinal carinea. Mesopleurum polished, gite polished. prepectal carina not reaching te margin. Coxae There are two males mounted on the pin. I and legs yellow. Middle and hind coxae with consider the right-hand specimen to be the type. long hairs. Hind femur slender, hind tibia slen- The other specimen also represents H. coxalis. der and very long, Nervellus vertical, discoidel- There are also three female paralectotypes; one la absent. First gastral segment rather slender, of these is the holotype of Megastylus {Helictes) tergite almost polished. Following tergites pol- pilicornis Thomson, holotype label of M. G. Fit- ished. Second tergite proximally and distally ton, 1980. yellow. Third tergite proximally yellow. The lectotype of Idioxenus variator has a The two males labelled: "Lapland m., 4.viii. Förster label "Lousberg, 17.10. 6, 29 gl" and Bhn" and "Lapland in. Bhn", respectively, I la- the lectotype-label of the present author. belled as paralectotypes. Characteristics of the lectotype of Helictes The lectotype of Idioxenns coxalis was la- variator: Male. Front wing 4.2 mm long. Ty- belled by Aubert and has a Forster label "Aa- loids of flagellar segments 6— 7—8. Second ter- chen". gite almost polished, some vague microsculp-

The neotype of Idioxenus coxalis has a ture is present.

Forster label "Aachen, â und 9 , 29 gl" a cab- Ther are two other Förster males, one la- inet label coxalis Frst. and a lectotype label of belled "Aachen". The other without a label. Aubert, 1977. There are two specimens mounted on the pin. I consider the right-hand Material examined. — I studied 147 S from the fol- specimen, a male, to be the neotype. lowing localities. Austria: St. Schladming, 1250 m, Characteristics of the neotype of Helictes Heidelbeere; Flintsbach, Inn.B, 550 m; Walchsee, Ti- rol, 800 m; T. Pertisau, 1550 m; Fl. 3, Schwestern coxalis: Male. Front wing 3.4 mm long. Palpi Grat, 2000 m; Reiter Alm, 1600 m, Heidelbeere (all yellow. Clypeus convex, yellowish brown. Ma- coll. Haeselbarth); Kärnten, Himmelberg, 1000 m lar space wide, 0.33 of width of face. Clypeus (coll. Zwakhals). Germany: Bayern. Wiershausen; face with rather suberect hairs. and long, close, Jettenhausen; Ammergeb. Jausen, 1400— 1600 m, Face with microsculpture. Scape ventrally yel- Nickelswald 1100— 1300 m; Starnberg, Kerschlach; low, with long hairs. Tyloids on flagellar seg- Leutstetten; Glonn; Niederaudorf, 1000 m; Herrsch- 7 ments 6— — 8. Frons vertex and gena polished. ing Widdersberg Mischwald; Gauting (all coll. Hae- Lower gena with long hairs. Pronotum pol- selbarth); EUmau 1050 m; Garmisch 700—1400 m; il- (coll. Bauer, Zool.Staatss. München). ished, epomia present. Mesoscutum strongly legible 750 m Hann. Münden; Geierlambach, Heidelbeere; convex, with fine microsculpture. Notauh pre- Lippoldshausen; Nd. Sachsen, Bramwald; Holledau, sent, but not reaching the margin. Scutellum Heidelbeere; Fl.Triesenberg, 1450 m; Hedemünden; with lateral carina only at proximal corners and Meensen; Dransfeld (ail coll. Haeselbarth); illegible laterally with long hairs. Propodeum polished, leg. Pfankuch (Zool.Staatss. München); Augsburg with hairs, pleural carina present. Mesopleurum (Zool.Staatss. München); Goslar a. H. Haldenstieg, polished, with prepectal carina present but Grauhöferholz; Harz, Harzburg, Radautal; Allgäu, —

102 Tijdschrift voor Entomologie, deel 130, 1987

Riezlern, 1150 m (coll. Bauer, Zool. Staatss. Distribution. — The species gives the impression of München). Italy: St. Peter, Ahrntal, Südtirol, 1350 being a boreal and alpine element. 1600 m; Martelltal, Südtirol, 2100 m; Tremalzo, Judi- kar Voralpen, 1730-1900 m; Karthaus Südtirol, 1200 The name "incongruens" is the Latin for m; Merano, 700 m; Feldthurns (Bolzano) 1200 m; "disagreeing". Partschins, Südtirol, 850 m; Tirol, Südtirol, 2250 m; Malcesine (VR) 500 1300 m. Bosco ceduo; Campi, — Helictes fabularis species nova Riva s. Garda, 240— 1200 m (22 specimens) (all coll. Haeselbarth); Bolzano, 800 m; Sarntal (Bolzano) 1250 In the type series of Megastylus borealis m (coll. Zwakhals). Netherlands: Ede (Prov. Gelderl.) Holmgren there is a male which differs conspic- (coll. Zwakhals and coll. van Rossem); Heilo and Ber- uously from the other syntypes by having a sin- gen (Prov. Nd Hoi.); Asperen and Arkel Prov. Zd gle tyloid. I consider this specimen to represent Hoi.) (14 specimens); Halsteren (Prov. Nd Brab.) an undescnbed species. (coll. Zwakhals). Norge: Opland, Lom-Lia (4 speci- The holotype of Helictes fabularis has the fol- mens) (coll. van Rossem). Sweden: Dalarna, Boda = lowing labels: "Lp (= Lapland) in., Bhn" ( Kyrkby, Silverberg; Fjätervalen Idre; Transtrand, Boheman) (Riksmuseum, Stockholm). Hemfjäll Stangen (coll. van Rossem); Höör (Skâne) (coll. Zwakhals). Switzerland: Wallis, Fiesch, 1200 m (coll. Zwakhals). Characteristics of the holotype of Helictes Localities of type material are not repeated. fabularis: Male. Front wing 4.7 mm long. Palpi Collecting dates between May and November. and mandible yellow. Clypeus convex, polished, front margin protruding. Face wide, pol- is spread in the Distribution. — The species widely ished to slightly coriaceous, with widely placed western Palaearctic Region. setae. Malar space wide, 0.45 of width of face. Frons and gena slightly coriaceous. Gena and Helictes incongruens species nova occiput with long, subadpressed hairs. Occipital The holotype of Helictes incongruens species carina closed. The single tyloid on the sixth fla- nova has the following labels: "Judikar Voral- gellar segment. Postannellus long. Scape large, pen, C. Tombea 1800 m, 18.vi.l958" (leg. & broadly ovate. Pronotum polished, with strong coll. Haeselbarth). "Helictes sp." det. Townes, epomia and a rather characteristic downward 1964, and the holotype label of the present au- slope of the hind margin dorsally. Mesoscutum thor. with indistinct microsculpture and vaguely out- Characteristics of the holotype of Helictes in- lined notauli (damaged by pin). Propodeum pol- congruens: Male. Front wing 3.6 mm long. Palpi ished, with erect setae. Only the pleural carina whitish. Clypeus convex, polished, fuscous, present. Mesopleurum pohshed, prepectal cari- front margin light brown, width 0.59 of width na not reaching the margin. Legs, including the face. Lower part of face polished, towards an- coxae yellowish. Hind coxa relatively slender. tennal sockets somewhat rough, with suberect Hind femur and tibia exceptionally slender. hairs. Frons, vertex an occiput pohshed. Vertex Nervellus indistinctly intercepted, discoidella and occiput with widely placed hairs. Tyloids absent. First gastral segment slender, polished 7 8 and with the spiracles at 0.5 of the length. on flagellar segments 6— — —9. Pronotum The polished, epomia present. Mesoscutum almost first sternite ending in the apical half. The other polished. Lateral carina of scutellum only pre- tergites fuscous, polished, with short suberect sent beyond the corner and not meeting at apex. hairs. Mesopleurum polished, prepectal carina not Material examined. — The holotype only (Sweden, reaching the margin. Legs long and slender. Te- Lapland). In a collection of the Museo de Ciencias gulae white. Propodeum polished, only pleural Naturales (Santa Cruz de Tenerife) (Dr G. Ortega) I carina present. All tergites polished. found specimens from Gran Canaria, Gomera and Palma (Islas Canarias).

Material examined. — Italy: â , Brescia, Judikar Voralpen, C. Tombea, 1800 m, 18.vi.l958, holotype. The name "fabularis" is the Latin for "mythi-

Paratypes: i , Judikar Voralpen, C. Tombea, 1800 m, cal' 18. vi. 1958. Austria: d, Steiermark, Schladming, 1250 m, 11. vi. 1972, Heidelbeere. Germany: 6, Reither Phosphoriana nomen novum Alm, 1600 m, Heidelbeere. Preceding specimens leg. Phosphorus Voet, ? 1769: 84. and coll. E. Haeselbarth, München. Sweden: â , Lap- land in., S.viii., Bhn (= Boheman) (specimen in type Phosphorus Thomson, 1857: 27 (= Voetia Strand, series of Megastylus borealis Holmgren; Riksmuseum 1943). Stockholm). PhosphorusVunKossem, 1980: 129—131. —

Van Rossem; Western Palaearctic Oxytorinae 103

front The name Phosporus, which I re-introduced Characteristics of the female: Length slender, 7.0 9.0 in 1980, is preoccupied. I propose to use Phos- wing 5.0 mm. Postannellus — phoriana as the replacement name, gender femi- times as long as wide. The pronotum with the nine. The type-species of Phosphoriana is Enty- same characteristics as in the male. Mesoscutum poma rugosissimum Strobl, 1903, the type-spe- steeply rising, polished, with conspicuous no- cies of Phosphorus Van Rossem by monotypy. tauli. Propodeum with pleural, lateral longitudinal and apical transverse carina. Prepectal carina Phosporiana rugosissima (Strobl) strong. Colour of the legs the same as in the Entypoma rugosissimum Strobl, 1903: 114. male. Hind femur robust, 4.3 times as long as Phosphorus rugosissimus; Van Rossem, 1980: 129 wide. Front wing with areolet. Nervellus inter- 131. cepted below the middle, discoidella present. First tergite long and slender, about 3.0 times as Hitherto the male of this species was un- long as wide apically, with rough sculpture, spi- all known. I found two males in the collection of racles at 0.76 of length. Apical margins of Haeselbarth. A description follows here. tergites ivory-yellow. Ovipositor 0.25 of length front wing. Characteristics of the male: Length front Distribution. rare species. The holotype of wing 4.6 mm. Palpi white. Mandible yellow, — A Strobl is from Johnsbachgraben (Austria). There are lower tooth slightly shorter. Clypeus with api- three specimens from Germany in the collection of E. cal half flattened and yellow, about 1.7 times as Haeselbarth (München). wide as long. Face below the antennae protuberant, with a conspicuous groove between the antennal sockets. Below each antennal socket a Genus Proeliator Van Rossem triangulate ivory spot, the base proximal to the /^roe/!Äfor Van Rossem, 1982: 152—154. socket. Malar space as wide as apex of postan- brought to notice an un- nellus. OOL : POL =2 : l^). Frons, vertex and Dr. H. Townes my gena polished. Face with some vague and shal- described species in the type series of P. propri- low punctures. Gena narrow, about 0.35 of HS Van Rossem. A description follows here. width eye. Antenna long and slender, postannellus 7.0 times as long as the apical width. No Key to the Proeliator females tyloids present. The pronotum striking, having (The males of P. invictus and P. captiosus are two elevations, ivory in colour, with a sharp unknown) groove between, directly behind the postocciput. The mesoscutum strongly inchned upwards 1. Length of ovipositor 0.14—0.17 of length from the pronotum, the median lobe conspicu- of front wing ously separated from the lateral lobes by wide Proeliator invictus spec. nov. but shallow notauli. Towards the centre of the — Length of ovipositor 0.23—0.30 of length mesoscutum the median lobe with a V-shaped of front wing 2 depression. The propodeum with some trans- 2. Lower tooth of mandible very small, giving verse, irregular sculpture. Pleural, lateral longi- the impression of a single-toothed mandi- tudinal and median longitudinal carinae present. ble. Last tarsal joint of hind leg robust, Mesopleurum polished prepectal carina strong. claws strong. Alaska All coxae and most of the front and middle legs Proeliator captiosus Van Rossem whithish yellow. Hind femur and tibia more — Lower tooth of mandible shorter than up- yellow. All femora rather stout. Hind tibia long per tooth, but visible. Last tarsal joint of and slender. First tergite long and slender, 3.5 hind leg not particularly robust. Europe .... times as long as the apical width, with longitudi- Proeliator proprius Van Rossem

nal sculpture, which is continued on tergites two and three. All tergites with a broad apical ivory band.

Material examined. — Germany: 2 â, Bayern, = Neuburg Donau, Finkenstein, 6. vii. 1982, leg. and OOL ocular-ocellar line. = lateral ocelli coli. Haeselbarth. POL distance between 104 Tijdschrift voor Entomologie, deel 130, 1987

Proeliator invictus species nova shape, 1.8—2.5 times als long as the apical Characteristics of the holotype of P. invictus. width. Median dorsal carina in most specimens Female. Front wing 3.4 mm long. Lower tooth short. The first tergite coriaceous. Second ter- of mandible about half the length of upper gite in part coriaceous. Following tergites pol- tooth. Clypeus elliptical, upper margin convex, ished. Ovipositor 0.23—0.30 of length of front for the rest clypeus impressed. Entire head pol- wing. Sheath with widely placed hairs. ished. Pedicel large. Antenna yellowish brown. Characteristics of the male: Tyloids of flagel- Pronotum polished, with epomia. Mesoscutum lar segments 6— 8. polished, with adpressed rather close hairs. No- tauH weak, only indicated on the margin. Pro- Distribution. — Germany: Spessart. Sweden: Mes- podeum with rather close erect hairs. Apical saure (Lapland). transverse, median longitudinal, and pleural carinae present. Mesopleurum polished, prepectal Proeliator captiosus Van Rossem carina to the margin. Front wing with areolet. Proeliator captiosusVznKosstm, 1982: 153— 154. Discoidella absent. Front and middle legs, including coxae yellow. Hind legs more brown- Characteristics of the female: Front wing 3.3 ish, hind femur with conspicuous long hairs. mm. Lower tooth of mandible very small, giv- First tergite coriaceous, 2.0 times as long as api- ing the impression of a single toothed mandible. cal width. Dorsolateral and median dorsal cari- Occipital carina closed. Pedicel large. Postan- na strong. Median dorsal carina to apical mar- nellus 4.0 times as long as wide. Epomia pre- gin. Following tergites polished. The fourth ter- sent. Front wing with areolet. Nervulus somew- gite and following with transverse rows of hat inclivous. Nervellus somewhat reclivous. widely placed suberect setae. Ovipositor 0.14 of Last tarsal joint of hind leg robust, claws strong. length front wing. Sheath with widely placed First tergite coriaceous, median dorsal carinae long hairs. not present. Following tergites brown, polished.

Ovipositor of paratype is 0.17 of length of Ovipositor 0.26 of length front wing. Sheath front wing. with widely placed long hairs. Male unknown. Male unknown.

Material examined. — Sweden, $ , holotype, Mes- Distribution, — U.S.A.: Mt McKinley, Alaska saure, 7.ix.l972, leg. Karl Muller; 9, paratype, Mes- (coll. Townes) saure, 12. ix. 1971, leg. Karl Muller (both coll. Townes, Gainesville (Florida). Genus Megastylus Schiodte The name "invictus" is the Latin for "indis- MegaStylus Schiodte, 1838: 139. putable, irrefutable". Megastylus; Townes, 1971 : 205. Megastylus; Van Rossem, 1974: 273—285. Megastylus; Van Rossem, 1983b: 121—132. Proeliator proprius Van Rossem Proeliator propriusYa.nKossem, 1982: 152 153. — Type-species: Megastylus cruentator Schiodte, 1838. Characteristics of the female: Front wing 3.5 mm. The lower tooth of mandible visible, short- Megastylus cruentator Schiodte er than upper tooth. Head polished, square. Oc- Megastylus cruentator ScWioàie, 1838: 139. cipital carina closed. Scapus subcylindrical, ped- Megastylus cruentator; Van Rossem, 1974: 276—278. icel large. Pronotum polished, epomia present. Megastylus cruentator; Van Rossem, 1983b: 123 & Mesoscutum convex, polished, with widely 126. placed subadpressed hairs. Notauli weak. Pro- Cryptus (Helictes) cruentatus Haliday, 1838: 115. podeum with a strong apical transverse carina Megastylus cruentator; Fitton, 1976: 333. and pleural carina, other carinae weak to obsolete. Propodeum with long, erect hairs. Front Characteristics of the lectotype of Cryptus wing with areolet. Nervellus vertical, discoidel- cruentatus Haliday. Labels: a label "named by

la absent. Front and middle coxae whitish, hind Claude Morley Helictes cruentatus Hal. Type coxae brown. Legs yellow, with rather long (unlabelled) vi. 1913; a circular label with red hairs, especially the hind tibia, the hind femur margin Type CM; lectotype label of Fitton, and the tarsi. The first tergite rather variable in 1975. Nat. Mus. Ireland, Dublin). Female. The — —

Van Rossem: Western Palaearctic Oxytorinae 105

specimen represents Megastylus cruentator History), Entomology 45: 1 — 1 19. SchÌ0dte. Förster, A., 1868. Synopsis der Familien und Gat- tungen der Ichneumonen. — Verhandlungen des Naturhistorischen Vereins der Preusischen Rhein- MegaStylus orbitator Schiedte lande und Westfalens 25: 135—221. Megastylus orbitator Schiodte, 1839: 139 (type lost). Förster, A., 1871. Uebersicht der Gattungen und Ar- Megastylus orbitator; Van Rossem, 1983b: 127 129 — ten der Familie der Plectiscoiden. — Verhand- (neotype) lungen des Naturhistorischen Vereins der Preus- Misoleptus maderensis Wollaston, 1859: 21 {Misolep- sischen Rheinlande und Westfalens 28: 71 — 123. tus is a lapsus for Mesoleptus) Gmelin, J. F., 1790. Caroli a Linné, Systema Naturae "^Megastylus maderensis; Fitton, 1976: 356. per regna tria naturae. . .etc., editio 13 2(5): 2125—3020. — Lipsiae. Ctiaracteristics of the holotype. Labels: Ma- Gravenhorst, J. L. C, 1820. Monographia Ichneumo- deira Wollaston (printed); a blue label: Misolep- num Pedemontanae regionis. — Memorie della tus maderensis W.; B. M. Type Hym. 36. 1999; Academia delle Scienze di Torino 24: 275—388. holotype label Fitton 1974. Male. Front wing Gravenhorst, J. L. C, 1829. Ichneumonologia Euro-

paea 1 : 1—827; 2: 1—989. — Vratislaviae. 2.4 mm long. The specimen is quite small and Gregor, F., 1941. Stredoevropské druhy rodif Euste- stuck to the mounting slip in such a way that nnx (Forst.) Thoms. (Hym. lehn.). — Entomolo- ventral examination is impossible. Nevertheless gické Listy, Brno 4: 5— 8. I hold it to be close to M. orbitator. Haliday, A. H., 1838. Descriptions of new British insects, indicated in Mr. Curtis's Guide. — Annals References ofNatural History 2: 112-121,183-190. Ashmead, W. H., 1902. Papers from the Harriman Hellen, W., 1915. Beiträge zur Kenntnis der Ichneu- Alaska Expedition. 28. Hymenoptera. — Pro- moniden Finlands. I. Subfamilie Pimplinae. — Ac- ceedings of the Washington Academy of Sciences ta Societatis pro Fauna et Flora Fennica 40: 5— 89. 4: 117—274. Hellen, W., 1937. Für die Fauna Finnlands neue Ich- 1968. Révision du genre Eustennx neumoniden (Hym.). — Notulae entomologicae Aubert, J. -F., Forst, et descriptions d'autres Ichneumonides Mi- 17:5—13.

croleptinae inédites. — Bulletin de la Société ento- Holmgren, A. E., 1855. Försök till uppstälning och

mologique de Mulhouse 1968: 37—41. beskrifning af de i Sverige funna Tryphonides. — Aubert, J.-F., 1969. Deuxième travail sur les Ichneu- Kungliga Svenska vetenskapsakademiens Hand-

monides de Corse (Hymenoptera). — Veröffentli- Imgar : 93—246. chungen der zoologischen Staatssammlung, Horstmann, K., 1974. Typenrevision der von Strobl

München 13: 27—70. m der Gattung Hemiteles Gravenhorst s.l. be-

Aubert, J.-F., 1970. Révision des travaux concernant schriebenen Arten und Formen (Hymenoptera, les Ichneumonides de France et 7e supplement au Ichneumonidae). — Zeitschrift der Arbeitsge- catalogue de Gaulle. (100 espèces nouvelles pour meinschaft österreichischer Entomologen 25: 52-

la faune française). — Bulletin mensuel de la So- 56. ciété Linnéenne de Lyon 39: 269—280. Jussila, R., 1965. The Ichneumonidae of the Kevojoki Aubert, J.-F., 1975. Révision des Apertleptus Forst, et area in Inari Lapland (Finland). — Annales Uni- Plectiscidea Vier. {Plectiscus auct.) du Förster et de versitatis Turkuensis (A II) 34: 1 — 186. Strobl. — Opuscula Zoologica, München 138: 1 Kiss von Zilah, A., 1924. Zweiter Beitrag zur Kennt- 8. nis der ungarischen und siebenbürgischen Ichneu- Aubert, J.-F., 1977. Révision des Ichneumonides Pro- moniden-(Schlupfwespen-)Fauna. — Verhand- lictus Foerst., Pantisarthrus Foerst., Aniseres lungen und Mitteilungen des siebenbürgischen Foerst. et Helictes Hal. — Spixiana 1(2): 141 Vereins für Naturwissenschaften in Hermannstadt 149. 72—74:32—146. Curtis, 1832. British Entomology 9: pis. 384—433. Oehlke, 1963. Revision der im Deutschen Entomo- J., J., — London. logischen Institut aufbewahrten Typen paläark- Fhton, M. G., 1976. The Western Palaearctic Ichneu- tischer Ichneumoniden (Hymenoptera: Ichneu- monidae (Hymenoptera) of British authors. — monidae). — Beiträge zur Entomologie 13: 403 Bulletin of the British Museum (Natural History), 410. 303 373. Entomology 32: — Perkins, J. F., 1962. On the type species of Foerster's Fitton, M. G., 1978. The Ichneumonidae (Hymeno- genera (Hymenoptera: Ichneumonidae). — Bul- der ptera) described by J. F. Ruthe. — Zeitschrift letin of the British Museum (Natural History),

Arbeitsgemeinschaft österreichischer Entomolo- Entomology 1 1 : 383—483. gen 30: 75—79. Pfankuch, K., 1906. Die Typen der Gravenhorstschen Fitton, M. G., 1982. A catalogue and reclassification Gattungen Mesoleptus und Tryphon. — Zeit- of the Ichneumonidae described by C. G. Thom- schrift für Systematischen Hymenopterologie und son. — Bulletin of the British Museum (Natural Dipterologie 6: 81—96, 217—224, 289—296. —

106 Tijdschrift voor Entomologie, deel 130, 1987

der Ratzeburg, J. T. C, 1852. Die Ichneumonen Schmiedeknecht, O., 1911. Opuscula ichneumonolo- Forstinsecten in forstlicher und entomologischer gica 28 & 29: 2161—2322. — Blankenburg. Beziehung 3: 1 —272. — BerHn, Nicolai. Strand, E., 1918. Ueber W. Horn's litauische entomo- Roman, A., 1923. Ichneumonids reared from Diptera logische Kriegsausbeute 1916. Hymenoptera. — Nematocera. — Entomologist's Monthly Maga- Entomologische Mitteilungen, Berlin 7: 30—32, zine 59: 71—76. 149—160. Roman, A., 1925. Schwedische Schlupfwespen, alte Strobl, P. G., 1900. Ichneumoniden Steiermarks (und und neue. — Arkiv for Zoologie 17A(4): 1 —34. der Nachbarländer). — Mitteilungen des natur- Rossem, G. van, 1974. The Gravenhorst, Schiedte and wissenschaftlichen Vereins für Steiermark 37: Foerster types, belonging to the genus Megastylus 132—257. Schiodte, 1838, with keys to the species. — Strobl, P. G., 1903. Ichneumoniden Steiermarks (und Tijdschrift voor Entomologie 117: 273—285. der Nachbarländer) (Schluss). — Mitteilungen des Rossem, G. van, 1980. A revision of some Western naturwissenschaftlichen Vereins für Steiermark Palaearctic Oxytorine genera. — Spixiana, Supple- 40: 111—142.

ment 4: 79— 135. Thomson, J., 1857. Monographie du groupe des Tra- Rossem, G. van, 1982. A revision of some Western gocéphalites, de la famille des Cérambycides. —

Palaearctic Oxytorine genera. Part II. Genus Eu- Archives entomologiques 1 : 25—39. sterinx, Proeliator new genus. — Spixiana 5: Thomson, C. G., 1888. xxviii, Försök tili grupering af 149—170. slägtet Plectiscus, Grav. — Opuscula Entomologi- Rossem, G van, 1983a. A revision of Western Pal- ca 12: 1266—1318. aearctic Oxytorine genera. Part III. Genus Procli- Townes, H., 1969. The genera of Ichneumonidae, part tus. — Contributions of the American Entomo- 1. — Memoirs American Entomological Institute logical Institute 20: 153— 165. 11: 1—300. Rossem, G. van, 1983b. A revision of Western Pal- Townes, H., 1971. The genera of Ichneumonidae, part aearctic Oxytorine genera. Part IV. Genus Megas- 4. — Memoirs American Entomological Institute tylus. — Entomofauna 4: 121 — 132. 17: 179—206. Rossem, G. van, 1985. A revision of Western Pal- Viereck, H. L., 1914. Type species of the genera of aearctic Oxytorine genera. Part V. Genus Aperi- ichneumon flies. — Bulletin of the United States leptus. — Spixiana 8: 145— 152. National Museum 83: 1 — 186. der Ich- systematicus Coleoptero- Ruthe, J. F., 1855. Beiträge zur Geschichte Voet, J. E., 1769. Catalogus neumoniden. — Stettiner Entomologische Zeitung rum 1: 1—104. 16: 51—58, 79—89. Woilaston, T. V., 1859. Brief diagnostic characters of faunam insects. — Annals and Schiodte, J. G., 1838. Ichneumonidarum ad undescribed Madeiran Daniae pertinentium genera et species novae. — Magazine of Natural History (3)1: 18—28, 113 Revue Zoologique 1: 139— 141. 124. faunam 1838—1840. Insecta Lapponica de- Schiedte, J. G., 1839. Ichneumonidarum ad Zetterstedt, J. W., Daniae pertinentium genera et species novae. — scripta: 1 — 1 140. — Lipsiae. Magasin Zoologique 9: 1 —27. )

Van Rossem: Western Palaearctic Oxytorinae 107

INDEX

{Synonyms in italics)

accusator (? Cylloceria) 20 erythropyga (Plectiscidea) 36

Acroblapticus 48 erythrostoma Gmelin (Ichneumon) . . . 60 agitator (Plectiscidea) 37 erythrostoma Gravenhorst (Plectiscus) 60 alpigena (Cylloceria) 16 erythrostoma (Helictes) 60 alpigena (Eusterinx) 56 eurystigma (Plectiscidea) 47 alpigenus (Catomicrus) 56 Eusterinx 50, 52 ambulator (Plectiscus) 35 eversorius (Plectiscus) 30 amicalis (Plectiscidea) 33 fabularis (Helictes) 64 Apoclima 15 flavicoxis (Plectiscus) 38 aquilonigena (Eusterinx) 59 flavipes (Gnathochorisis) 50 ardentis (Proclitus) 22 flavipes (Hemiphanes) 12 argutula (Eusterinx) 54 flavizonus (Plectiscus) 30 armata (Eusterinx) 57 foersteri (Plectiscidea) 40 armatus (Oxytorus) 14 fracticornis (Lampronota) 20 attentus (Proclitus) 21, 22 fraterna (Plectiscidea) 44 binodulus (Plectiscus) ....'. 39 Fugatrix 48 bispinosa (Eusterinx) 56 fulvicornis (Cryptus) (Helictes) 60 bistriata (Plectiscidea) 29 fulvicornis (Proclitus) 21 blandita (Plectiscidea) 44 fulvipectus (Proclitus) 22 Blapticus Forster 48 fulvus (Plectiscus) 47 Blapticus Thomson 48 fusciventris (Cylloceria) 16 borealis (Cylloceria) 17 gilvus (Plectiscus) 48 borealis (Helictes) 62, 64 Gnathochorisis 48 brachyums {Plectiscus) 32 gracilis (Pantisarthrus) 23 caligata (Cylloceria) 18 gravator (Hemiphanes) 13 canaliculata (Plectiscidea) 36 habilis (Plectiscus) 45 captiosus (Proeliator) 6G haeselbarthi (Apoclima) 15 Catomicrus 55 Helictes 60 cinctula (Plectiscidea) 33 helvola (Plectiscidea) 34 circaea (Eusterinx) 58 Hemiphanes 12 collaris (Plectiscidea) 39 Holomeristus 58 collaris (Plectiscus) 39 hortense (Hemiphanes) 13 comes (Proclitus) 21 hostilis (Plectiscus) 47 communis (Plectiscidea) 48 humeralis (Plectiscidea) 47 conjuncta (Plectiscidea) 38 Idioxenus 60 connexa (Plectiscidea) 43 imperspicua (Cylloceria) 18 conspicuus (Helictes) 62 inaequalis (Eusterinx) 57 coxalis (Idioxenus) 62 inaequalis (Idioxenus) 60 coxator (Plectiscus) 32 inaequalis (Pantisarthrus) 23 crassicornis (Plectiscidea) 42 incongruens (Helictes) 6A crassulus (Gnathochorisis) 49, 50 indomita (Plectiscidea) 30 crenicornis (Lanipronota) 18 infirmus (Plectiscus) 48 cruentator (Megastylus) 66 inquilinus (Idioxenus) 62 cruentatus (Cryptus) (Helictes) 66 intricator (Idioxenus) 62 curticauda (Plectiscus) 33, 34, 38, 46 inusitatum (Hemiphanes) 14 Cylloceria 16 invalidus (Idioxenus 62 Dallatorrea 57 in vieta (Cylloceria) 18 dentifer (Gnathochorisis) 49, 50 invictus (Proeliator) 66 deterior (Plectiscidea) 44 Ischyracis 56 determinatus (Plectiscus) 33 jugorum (Eusterinx) 53 Dialipsis 24 Laepserus Forster dispar (Pantisarthus) 24 Laepserus Van Rossem disparilis (Eusterinx) 56 langei (Cylloceria) 17 distinctus (Plectiscus) 36 longicornis (Chalinoceras) 20 Divinatrix 57 luridator (Oxytorus) 14

edwardsi (Proclitus) : 22 luridator (Oxytorus) f. nigricoxa 14 108 Tijdschrift voor Entomologie, deel 130, 1987

luridus (Pantisarthrus) 24 propinquus (Idioxenus) 62 maderensis (Misoleptus) 67 proprius (Proeliator) 66 mancus (Chalinoceras) 18 proxiynus (Plectiscus) 35 mediator Förster (Idioxenus) 60 pseudocbropus (Pantisarthrus) 23 mediator Schiodte (Megastylus) 60 pseudoligomera (Eusterinx) 54 Megastylus 66 pseudominutus 54 Megastylus (Helictes) Thomson 60 pseudominutus (Hemiteles) var. jugorum 53 Megastylus Holmgren 60 pungens (Plectiscus) 42 melancholica (Cylloceria) 20 pusilla (Eusterinx) 55 melancholica f. denticornis (Cylloceria) 20 refractaria (Eusterinx) 59 melancholica f. marginator (Cylloceria) 20 restrictus (Gnathochorisis) 50 melanocera (Plectiscidea) 35 rudepunctatus (Pantisarthrus) 24 mendica (Plectiscidea) 44 rudis (Proclitus) 22 mesoxantha (Plectiscidea) 43 rugosissima (Phosphoriana) 65 minima (Eusterinx) 59 rugosissimum (Entypoma) 65 moerens (Plectiscidea) 30 rugosissimus (Phosphorus) 65 montanum (Hemiphanes) 14 signaticorne (Apoclima) 16 monticola (Plectiscidea) 36 sodalis (Plectiscus) 33 Myriarthrus 60 striolata (Cylloceria) 19 nava (Plectiscidea) 40 subalpinus (Aniseres) 23 nemorensis (Plectiscidea) 28 subangulata (Plectiscidea) 47 nigricoxus (Helictes) 62 subcurvatus (Plectiscus) 36 nigritus (Plectiscus) 48 subdola (Eusterinx) 54 nuptialis (Plectiscus) 32 subsimilis (Ephalmator) 28 obscurella (Eusterinx) 55 subsimilis (Plectiscus) 35 occupator (Lissonota) 16 substantiva (Plectiscidea) 41 ochropus (Pantisarthrus) 23 subsulcatus (Proclitus) 21 oligomera (Eusterinx) 53 subteres (Plectiscidea) 29 orbitator (Megastylus) 67 subtilis (Plectiscus) 36 Oxytorus 14 suerinensis (Cylloceria) 17 paganus (Proclitus) 21 sylvestris (Cylloceria) 19 Pantisarthrus 23 sylvestris (Tryphon) 19 parvula (Plectiscidea) 31 tartarea (Eusterinx) 56 petiolatus (Plectiscus) 35 tetraglyptus (Idioxenus) 62 Phosphoriana 64 tener (Plectiscidea) 31 Phosphorus Thomson 64 tenuicincta (Eusterinx) 58 Phosphorus Van Rossem 64 tenuicornis (Plectiscidea) 32 Phosphorus Voet 6A terebrata (Gnathochorisis) 50 pilicornis (Megastylus) (Helictes) 63 terebrator (Plectiscidea) 45 Plectiscidea 24 townesi (Hemiphanes) 13 Plectiscus auctores 24 trichops (Catom-icrus) 55 Plectiscus (Proclitus) 20 vagator (Plectiscidea) 35 posticata (Plectiscidea) 42 variator (Idioxenus) 63 praepositus (Plectiscus) 45 ventosa (Plectiscidea) 46 praetor (Proclitus) 21 Voetia 64 Proclitus 20 xanthocephalus (Gnathochorisis) 50 Proeliator 65 xanthoneuris (Plectiscus) 30 properator (Oxytorus) 14 zonatus (Proclitus) 21 Tijdschrift voor Entomologie 130; 109— 127 Gepubliceerd 30 november 1987

ECOLOGY, LIFE HISTORY AND DISTRIBUTION OF PALINGENIA WNGICAUDA (OLIVIER) (EPHEMEROPTERA)

BORIS K. RUSSEV

Institute of Zoology, Bulgarian Academy of Sciences, Sofia

Dedicated to the memory of the Dutch naturalist fan Swammerdam, who initiated the studies on the life history of Palingenia longicauda with great love and dedication in 1667

Abstract

The ecology and life history of Palingenia longicauda (Olivier) have been studied in the Bulgarian section of the river Danube, both for the aquatic larval stages and the flying

adult stage. The significance of this species for the fisheries is outlined. The distribution in Europe, the retreat from Western Europe and finally the complete disappearance from the Danube river system are described.

Introduction At present the nomenclature of this species

The first record of mayflies, as well as an ex- is as follows: planation of their name was given by Aristoteles Ephemera longicauda Olivier, 1791 (384 — 322 B.C.) in his Historia Animalium (see Ephemera flos-aquae Illiger, 1802 lilies, 1968; Francissen & Mol, 1984). Swam- Semhlis m.arginata Panzer, 1804 merdam (1752: 100) was correct in assuming Ephem.era swammerdiana Latreille, 1805 that Aristoteles, later cited by Plinius and Eli- Ephemera swammerdam-iana Shaw, 1806 anus, had studied the same insect, calling it Palingenia longicauda (Burmeister, 1839) Hemerobius, Ephemerus and Diaria, respectively. Clutius (1634) wrote about the abundance of Material and methods

Hemerobius in Dutch rivers some 350 years ago. I started my study on the life history and dis- He drew somewhat distorted pictures of the larva tribution of larvae of Palingenia longicauda along (in dorsal and ventral view), of the exuviae and the Bulgarian Danubian stretch in September, of the adult. 1952. For my qualitative and quantitative surveys

Swammerdam (1675) was the first, who re- I used the so-called fisherman's probe or gunter, ported on the life history of the larva and of utilized by sportsmen for collecting mayfly larvae the adult, which he called "Haft" of "Oeveraas", as bait. This is a metal cylinder, measuring and presented pictures and descriptions of both. 16— 18 by 32 — 34 cm, fixed on a wooden handle They were later included in the Dutch edition of 6—8 m long. In total 1033 larvae and nymphs of the "Bible of Nature" (Bijbel der Nature) by were collected from 22 localities along the Bul- Swammerdam (1737) — the first book of this garian bank of the Danube (table 1). kind and of primary importance for that time. The transversal distribution of larvae of

In 1752 it was translated into German, and in Palingenia longicauda was established during 1758 into English. Marsili (1726: 25) in his six several years of zoobenthal studies along and volumes with geographical, historical, astronom- across the river between km 845 and 375. These ical and hydrographical data on Hungary and studies were carried out aboard of the hydro- the Balkans, reported on the mass flight of this graphical ship "Ossum", property of the Bul- mayfly species over the Tissa river. garian Danube Shipping and Monitoring Au-

109 no Tijdschrift voor Entomologie, deel 130, 1987

Table 1. Larvae and nymphs of Palingenia longicauda on the clay bottom immediately by the right Danubian bank.

Date RUSSEV: Palmgenia longicauda 111 banks the distance from the banks can be considered a key, although indirect, ecological factor, affecting the distribution (Russev, 1977). The larvae were predominantly found up to 100 m from the Bulgarian bank (70% of the localities), seven localities (23%) were between 100 and 252 m and two (7%) between 789 and 880 m off the Bulgarian bank. This shows the inability of the larvae to maintain their positions at distances of more than 250 m from the banks, which should be attributed to the higher current Fig. 1. Larva of Palingenia ncauda (Photo A. Val- velocity and the inadequate sand substrate in the kanov). middle of the stream. No larvae were found at flow rates over 0.76 m/s (measured 0.5 m above Palingenia longicauda feed on the organic mat- the bottom) and on sandy substrate. ter, which is taken up by the stomach from the

Depth is hardly of influence on larval dis- clay that is consumed. Strenger (1973) noticed tribution. Larvae were found at depths of up that the mouth organs of Palingenia are well to 10.6 m. During periods of rapid fall of the adapted for scratching the detritus. water level (e.g. on 29 September, 1954) they Only eggs that have reached adequate clay sub- left their holes in the clay, while many of them strate, develop normally. Under laboratory con- died while they tried to follow the retreating ditions at water temperatures of 20—25 °C water. Unger (1927) found out that the embryonal de- The studies on the horizontal and vertical dis- velopment up to the hatching of the larvae lasted tribution of larvae of Palingenia longicauda at for 4—6 weeks. He assumed that larval devel- km 166.5 of the Tissa river led to the con- opment took three years, and because of conclusion (Csoknya & Halasy, 1974), that tinuous growth, the larvae passed through some "... the most uniform distribution of the zoo- twenty moultings. Already Swammerdam (1752) benthos is found 5 m from the bank towards made the observation of three different size the river bed, in the entire depth of the mud classes of larvae shortly before the metamor- samples (60 cm). In the region lying closer to phosis, from which he concluded that their lon- the bank (3 m) the young larvae (0.5 —20.5 mm) gevity was three years. We tried to distinguish are distributed fairly uniformly in the mud sam- the respective stages by biometrics and analysis ples. Between 3 and 7 m from the bank, however, of variance between larvae of various ages, but they occurred in the uppermost 20 cm layer. The we obtained no positive results. This was ob- largest larvae (40.5 —60.5 mm) are more fre- viously due to individual and nutritional pecul- quent 4— 5 m from the bank, and predominantly iarities of larvae of various ages leading to merg- in the mud layers (30— 50 cm). Intermediate ing sizes and weights within the various age larvae (20.5 —40.5 mm) exhibit a uniform dis- groups. Only in July, shortly after the emergence tribution in the region examined". of the adults but before the hatching of the young During our studies on Palingenia longicauda larvae, two age groups could be distinguished, we measured the following hydrological and hy- which confirmed the three-year life-cycle. In drochemical parameters: average current veloc- other cases we have observed just one age group ity ranging from 0.56 to 2.10 m/s; turbidity after emergence. The probability of absence of

13—1046 g/m'; floating deposits 37—8391 kg/ a certain age group at a particular site is very

s; transparency 0.9 —23 cm; temperature up to high, since new eggs do not reach all localities 28.2 °C; dissolved oxygen 5.55 —9.65 ml/1; ox- each year. ygen saturation 68— 123%; oxydability The larvae live in U-shaped holes dug in the 2.41 —9-40 mg/1 O2; biological oxygen demand clay. They make these holes using their well- for five days (BOD5) 5.74—0.35 ml 0^/1; total adapted legs. Particularly the forelegs provided hardness 7.67—13.7 dH"; pH 7.5—8.2; alkalinity with denticles on the lateral sides, as well as 2.00—3.06 mg equiv./l; HCO, 124.0—186.9 the mandibulae with their strong lateral chitine mg/1; CI' 11.8 — 17.5 mg/1; general mineraliza- denticles, are very suitable for digging (fig. 1). tion 239—439 mg/1 (Russev, 1968). According to Swammerdam (1752) these holes According to Swammerdam (1752), Unger are "... lange, und rechte, zuweilen aber auch (1927) and Schoenemund (1929) the larvae of krumme und schiefe hohle Röhren in Thone, 112 Tijdschrift voor Entomologie, deel 130, 1987

Table 2. Distribution of Palingenia longicauda larvae across the Danube off the Bulgarian river bank (samples collected from a ship by the Petersen Bottom Sampler. 1/10 m').

Date RusSEV: Palingenia longicauda 113

(Table 2, continued)

Date km Distance 114 Tijdschrift voor Entomologie, deel 130, 1987

Table 3. Observation on the flight of Palingenia longicauda over the Danube between the 845th and 375th km.

Date RusSEV; Pdlmgenia longicauda 115

(Table 3, continued)

Date 116 Tijdschrift voor Entomologie, deel 130, 1987

» ,1

*«4^' "* * 4» É ^

3* A

li # *

n» •>

Jk. j^

Fig. 2. Holes of Palingenia longicauda larvae in a clay bank of the Danube river at km 715 (Photo B. Russev).

it is removed from the base to the apex of the two caudal filaments are stretched sidewards at wing. During the moulting of the legs, the insect an angle of 45° towards the body, thus forming loses its equilibrium and falls sidewards. After an angle of 90° between themselves. The fore- releasing wings and legs, the imago makes in- legs are stretched forewards resembling horns. tensive movements for the complete release of The caudal filaments often touch the water sur- the caudal filaments and the entire body from face, when the male flies low over the water the exuviae (figs. 4— 10). Unsuccessful moult- surface. When a female emerges, a cluster of up ings were seen very frequently. to ten competing males may approach it. Males seem to emerge one hour before the females, Ethology of imagos. — The newly emerged while they also frequent the banks. This may lead male imago is more active and flies much faster to the wrong conclusion that males are more than the subimago. Its caudal filaments are about numerous than females. As was already observed three times as long (ca. 65 mm), the forelegs are by Swammerdam (1752), Cornelius (1848) and longer and its yellowish-brown coloration is Csongor & Moczar (1954) females do not have a much brighter. Males usually fly towards the subimago stage. This may be the reason why they middle of the stream searching for females, moult somewhat later. The females may be easily whereafter they either return to the bank or fly distinguished by their larger body, large white further upstream. In the latter case they make the wings, smaller eyes, and three times shorter cau- 45° turn until they take the downstream direc- dal filaments. They fly higher and faster than tion again. The movements and body position of males. Although we have observed matings in males searching for females is most typical. The flight, gradually losing height, most matings RusSEV: Palingenia longicauda lil were seen on the water surface. Drenkelfort (1910, cited after Brinck, 1957), however, considered mating at the surface exceptional. During copulation the position of the male is under the female, while he holds her head with the forelegs. Couples may fly in any direction. According to Brodskii (1973), in his work on the swarming behaviour of mayflies, species of the family Pa- lingeniidae show the very abundant third swarming type; the male's mating flight includes rapid horizontal flight parallel to the water surface.

Mass flights. — During mass flights the abundance of males and females gradually increases, Fig. 3. Mass metamorphosis of nymphs of Palingenia and may reach more than 120 individuals/m\ longicauda on the water surface of the Danube river The river turns brown and even darker strips and (Photo B. Russev). spots may be detected. A typical buzz of hundreds of thousands of wings can be heard and a fish Ecological factors. — The ecological factors odour be smelled. Most specimens fly between that influence emergence and development of one and three m above- the surface. After the the adult stages are not fully clear. The compen- mating the males fall at the water and are carried sation flight is strongly influenced by strong away by the stream. The fertilized females con- winds. This was clearly observed in June I960 at tinue their short life until they have layed their km 715. After mass flights on the 9th and 10th, eyes. Mass flights seldom last more than half an there was not a single mayfly on the 11th with hour. poor weather and strong western wind. The After mass flights the banks are covered with flights resumed after June the 20th under more subimaginai skins and look white (figs. 11 — 13). favourable weather conditions (table 3). Another

The water surface is still covered with skins, and example dates from June 1958 at km 536. exhausted males still moving their wings. Palingenia longicauda was most abundant on the 8th, 9th, 10th and 12th during calm, sunny and Compensation flights. — After the mating the warm weather, whereas the flight was of medium females fly upstream with c. 18 km/h (Russev, intensity on 11th, with a wind of 3 Beaufort. 1959). Males that were unable to mate also do so, During a gentle breeze (3 Beaufort or less), but usually at a lower speed of 14 km/h (figs. 14, fertilized females can still perform their compen- 15). Females may sometimes touch the water, sation flight, as already described by Russev and fly off again without getting drowned. We (1973, figs. 3 — 5). It seems that wind direction have observed, under experimental conditions, does not influence these flights, as could be con- that females oviposit immediately after a drop- cluded from observations where the stream fol- ping by wave-like movements. The total number lowed a west-east ( fig. 5 ) , or northwest-southeast of eggs laid numbers appr. 8—9000, which all get direction (at km 716 and 556), or any other di- dispersed in the water. rection. Orientation seems to be guided by the We have called the upstream flight of the fer- oculi, or, as was noted earlier by Russev (1959), tilized females the "egg-laying-preceding com- by brief touches of the water surface, where the pensation flight" (Russev, 1959). This adaptive insects may be able to detect the current direc- behaviour may compensate the downstream tion. Further studies are, however, needed to find movements of the nymphs during metamorpho- out the exact orientation mechanism. The larger sis, as well as the carriage of the eggs by the water weight and wings, as well as the shorter caudal current before they reach the bottom. Eggs are filaments of the females may be considered as very small (360/300 to 380/330 \im, see figs. morphological adaptations for carrying eggs, as 16—20) and may be carried ca. three km (Russev, well as for the compensation flight. 1973). This is corresponding to the "colonization Influence of light intensity on swarming, as cycle" concept of Müller (1954), and confirmed was proposed by Pongracz (1933, cited after by other studies (Müller, 1973, 1982; Keller, Csongor & Moczàr, 1954), should be investigated 1975, and others). further, and the same is true for Csongor & Moc- 118 Tijdschrift voor Entomologie, deel 130, 1987

longkauda (Photos B. Russev). Figs. 4—10. Stages of moulting of the subimago of Paltngenia RUSSEV; Palingenia longicauda 119

^'5S

river (Photos B. Russev). Figs. 11 — 13. Subimaginai exuviae of Paltngenia longicauda on the bank of the Danube 120 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 14 — 15. Upstream compensation flight of Palingenia longicauda. Danube river. (Photo B. Russev). sàr's statement that "the coincidence of high at- flight period lasted from 5 to 20 June. Only very mospheric pressure, high water and air temper- few observations are available outside this pe- ature, with changing moon phase furthers mass riod. A medium to mass flight was observed on emergence". According to our observations in 28 and 29 May, 1968 at km 576 and km 570 by the period 1955 — 1958 mass fhghts in the Da- eng. N. Mladenov (personal communication) nube coincided with the last quarter of the moon (table 3). These observations confirm the period (13 June, 20 June and 9 June, respectively), but we of records of mass flights by Swammerdam (13 do not consider this clear proof. June 1671), Triebke( 1840) (middle ofJune), Cor- We also tried whether summarized water tem- nelius (1848) (12 — 20 June), Dziçdzielewicz peratures affected the timing of swarming. The (1867) (10—25June),SelysLongchamps (1888) average water temperature at the town of Svish- (10—25 June), Moczary (1900) (10—20 June) tov was calculated for the period 15 June 1955 to and Unger (1927) (5 June—early July), but is 9 June I960, but no correlation was found be- somewhat different from Beretzk et al. (1957) tween annual averages and the timing of the (end of June— early July, and sometimes even flights. somewhat later). Swammerdam (1752) and Cor- nelius (1848) considered warm winters, hot Flight period. — Mass flights of Palingenia springs (particularly May), and limited precipita- longicauda off the Bulgarian bank in the Danube tion favourable for early timing of flights. The were observed between 8 and 15 June; the entire abundance of the flights varies from year to year. RusSEV: Palingenia longicauda 121

Table 4. Literature data on the use of Palingenia longicauda as food of various fish.

?) Û C\ ^ Q S^Q o e û< 5 o S-2 '> o -9 rt cd S p 3 Çc —•i/i'ijJ (u.::cS e jji^c^ e (U :- D X Q Pi Q XtótB OJoK; ÜCcïi

Acipenser gueldenstaedti Brand Acipenser ruthenus (L.) Acipenser stellatus Pali. Ameirus nebulosus Le Sueur Barbus barbus L. Cyprinus carpio L. Esox lucius L. Gymnocephalus cernuus (L.) Gymnocephalus schraetzer (L.) Huso huso (L.) Silurus glanis L. Stizostedion lucioperca (L.) Zingel streber (Siebold) Zingel zingel (L.)

In some years, e.g. 1961, no mass flight was seen Several authors have reported on the use of at all (table 3)- larvae of Palingenia as baits for fishing sterlet Flights in Bulgaria were observed between (Acipenser ruthenus (L.)), and barbel {Barbus

15.30—19.30 h (sunset on 12 June is at 19.45 h), barbus L.), Lota lota L., Silurus glanis L., Aspro usually with the following sequence. Metamor- cingel L., Chalcalburnus chalcoides danubicus phosis of nymphs and male subadults occurred Antipa and Cyprinus carpio L. (Swammerdam, between 15.30— 17.00 h; female adults emerged 1752; Antipa, 1909: 248; Bacescu, 1943; Csongor between 16.00— 17.00 h; mass flights were be- & Moczàr, 1954; Russev, 1956). Bulgarian fish- tween 17.30 and 18.00 h, and compensation ermen were using larvae of Palingenia as baits flights started after 18.00— 18.30 h. Flights start- from May to September, and sometimens during ed earlier or later only very seldom. winter at low water level, for fishing Lota lota Imago and subimago usually live not longer and also sterlet. Larvae were collected using the than two hours, provided that they have mated. fisherman's probe (gunter). Larvae can be kept Cornelius (1848) noted an adult longevity of 1.5 alive for several days in cool, moisty sand. hours, or up to 13 hours if males had not mated. The significance of Palingenia longicauda for fish nutrion in the Balkans has extensively been SiGNinCANCE OF PALINGENIA LONGICAUDA studied (table 4). FOR FISH NUTRITION Several studies revealed that Palingenia domi- Palingenia has extensively been used as fishing nates in the diet of fishes. In a study of Gheraco- bait. It has various local names, e.g. "oeveraas" pol & Selin (1968) and Gheracopol et al. (1969) and "haft" in The Netherlands, "Spork-Oese", in the Danube river 69.4% of the diet of the "Sprock", "Spaargoos", "Spaargaänse" in Ger- starlet consisted of this mayfly species. An aver- many (Westfalen), "Tiszavirag", "Theissbliite" age of 30—40% of the food weight of fishes was in Hungary, "Vetritze", "Rusalii" in Romania, found in many other studies (e.g. Russev, 1963). and "gandatsi" (for the larvae), "rusalki" and Also see fig. 21. "karchani" (for the adults) in Bulgaria. Szent- limiting the distribution Ivany & Ujhazy (1973) reported on New Gui- Factors nean and Hungarian folk songs devoted to may- Swammerdam (1752) already described the flies. Two Hungarian folksongs are about the negative influence of climatological factors on "Flower of the Tissa" {Palingenia longicauda). the distribution of Palingenia: "Diejenigen 122 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 16 — 20. Eggs of Palingefiia longicauda at various magnifications. 16, 1450 X; 17, 285 X; 18, 730 X; 19, 285 X; 20, 140 X (Photos N. Hinton).

Dinge, die das Aas an seiner Veränderung hin- bedecken; dessgleichen auch die grossen Dürre, dern, es tödten, seinen Anwachs aufj-ialten und als die sich nöthiget ihre Häusgen zu verlassen, verursachen, dass es das eine Jahr in geringerer und andere anzubauen und auszubohren". Nowa- Anzahl und später als im andern hervorkommt days we would add the bank slides, the harmful sind folgende: Ein harter langer Winter, viel effects of the deposits on the holes during periods Schnee und Regen, als welche die Röhrgen, dar- of high water level, the high wind and rain during innen sie leben, zu und wegspülen, und mit Sand the time of mass flights, when compensation RUSSEV: Palingenia longicatida 123

flights are impossible and the chance of being carried downstream are increased. Negative effects by fishes were already discussed, but also birds and spiders are heavy predators of this species. Mass flights of Palingenia were seen to be attacked by birds as spoonbill (Platalea leuco- rodia L.), sparrowhawk (Accipiter nisus L.), kit- tiwake (Rissa tridactyla), as well as wild ducks, swallows, sparrows, larks, wagtails, crows, kites and terns (Gorove, 1819; Csongor & Mocsàr, 1954, personal observations). On 1 1 June, 1958, on the pier of Krivina Port, as well as in many other cases, I have observed male subadults being trapped in spider webs, and immediately attacked by the spider. However, anthropogenetic factors have ex- erted the strongest negative influence on this species. These factors include the construction of canals, dams, reservoirs, hydrodynamic power plants, commercial irrigation pumps etc. Also increasing pollution was detrimentous to the species. Some indirect evidence of this will be discussed in the following chapter.

Distribution in Europe In several parts of western Europe Palingenia was abundant up to the end of the 19th century. Lestage (1937) and Tshernova (1949) already expressed their concern about the process of its extinction in the rivers of western Europe. They also listed the publications with distributional records. In southern and southeastern Europe it occurred in large numbers as late as the 1960's, but its seems that it is now extinct from most localities in Europe. The following is a survey of all records available, arranged per country. It should be under- stood that not all records are fully reliable. Re- cords given by Schäffern (1757) and Cremer (1938) probably all belong to Ephoron virgo (Oliv.), what may be concluded from the flight period reported. Also records for France and Bel- gium are doubtful. The Netherlands: Clutius (1634), Swammer- dam (1675, 1737, 1752), Selys Longchamps (1888), Albarda (1888) and Lauterborn (1918, p. 40) — the rivers of Rhine, Meuse, Waal, Lek, IJssel and some of their tributaries. Hungary; Marsili (1726, p. 125), Gorove (1819), Mocsary, S. (1875), Mocsary, A. (1900), Unger (1927), Pongracz (1933 after Csongor- Mòcsàr, 1954), Csongor and Mòcsàr (1954), Be- retzk et al. (1957), Csoknya and Ferencz (1972) — the river of Tissa, Mocsàry, S. (1875) — the following tributaries of the Tissa — Maros, 124 Tijdschrift voor Entomologie, deel 130, 1987

Germany: Triepke (1840) — the Oder and 1. 1634 — 1900. Palingenia longicauda occurs some of its tributaries; Cornelius (1848) — the widely in the lower and middle courses of large Lippe — a right tributary of the Rhein; Hagen and medium-sized rivers, (1859) — Prussia. 2. 1901 — 1927. Its becomes extinct in West- The USSR: Dziedzielewicz (1867, p. 161) — ern Europe, and is strongly diminishing is Cen- the Dnester (upstream from Strvionzh at Sam- tral Europe, bor town); Kinel, Krasucki, Noskiewicz (1927) 3. 1928 — 1978. It is still present in the lower — the Dnester; Mikulskii (1936, p. 64) — the course of the Danube river, as well as in the rivers Dnester (Strvionzh, Seret and from Lvov down- Tissa, Bodrog, Maros-Muresul, Uh, Laboree and stream); Moras and Bacescu (1973, p. 29) — 18 Lazorica (all within the water catchment of the km to the east from Kishinev; Tchernova (1949), Tissa, a left tributary of the Danube), and it was Markovskii (1955) and Olivati (1961) — the also present in the river Vardar and in a canal Kilian branch of the Danube. near the Maritsa river. Romania: Mocsàry, S. (1875) — the Danube at Palingenia used to find favourable conditions Orsova (The Iron Gates); Motas (1936) — the in the oligo-^-mesosaprobic water of the Bulgar- Danubian island of Ada-Kale (on the bottom of ian Danubian stretch. The gradual deterioration, the present day Iron Gates reservoir); Bacescu caused by pollution, led to the rapid extinction of (1943) — the Danube, the Prut, the Mures and this species. The most abundant flight ever seen the Olt; Bogoescu (1958, p. 58) — the Danube was observed in 1958, and before 1968 mass delta. Calatasi and Olteanu at the Danube; flights were still recorded. Later on the flights Busnita, Enaceanu and Brezeanu (1961, p. 207) became more and more scarce, and over more — the Ardjes (a left tributary of the Danube); limited parts of the river. After 1974 no fisher-

Busnita, Brezeanu and Prunescu-Arion (1961, p. man, crewman of anybody else ever noticed the 320) — the Olt and the Danube at the Jiul river- flight of this popular and large mayfly along the mouth; Brezeanu and Prunescu-Arion (1962, p. Bulgarian Danubian stretch. Not one single male 167) — the Snt. George branch of the Danube; was seen, nor one single nymph found in clay Enaceanu and Brezeanu (1966, p. 182) — be- bottoms of formerly typical localities during a tween the 488th and the 235th km of the Danube special investigation carried out along the entire river; Prunescu-Arion, Elian and Baltac (1965, p. Bulgarian bank of the Danube. For this reason, 164) — Macin branch of the Danube near Braila we assume that it has become extinct from this town; Enaceanu (1967, p. 298 and 416) — the part of the river, as happened in the upper parts Danube and the flooded lowlands; Bogoescu and of the same river earlier. Tabacaru (1969) — the Danube delta. Belgium: Lestage (1923) — "The only known specimen from Belgium was collected some 50 Acknowledgements years ago in the vicinity of Diest near Demer". It is my pleasure to express my gratitude to my Poland: Ulmer (1927, p. 240) — the Visla highly respected teacher the late Prof. A. Val- river and some of its tributaries. konov for his valuable guidance, particularly at Bulgaria: Buresh (1936) — the Iskar river at the beginning of my studies, as well as to N. Svoge; Russev (1956 and 1966) — the Bulgarian Mladenov, Dr V. Naidenov, Dr V. Beshkov and stretch of the Danube and channel of the Maritza E. Undjian for various kinds of observations, and river. to colleagues from other countries, viz.. Prof. Dr Yugoslavia: Ikonomov (1958) — the Vardar O. A. Tschernova (Moscow, USSR) for larvae river, 15 km to the south of Skopje City; Russev from the USSR, to Dr M. Erti (Bratislava, Cze- (1968) — the Danube. choslovakia) and to Dr M. Keffermüller (Poz- The data on the distribution of Palingenia lon- nan, Poland) for valuable advise about the distri- gicauda in Europe over the last 350 years, reveal bution of Palingenia longicauda in their country. that three, not precisely defined, periods can be I am grateful to the late Prof. Dr G. Pleskot distinguished. One has to bear in mind, that for (Vienna, Austria), Dr I. Miiller-Liebenau (Plön, many parts of the range insufficient studies are ERG), Dr Kr. Kumanski (Sofia, Bulgaria) for available. The periods are: their support and critical reviews of my work. RUSSEV; Palingenia longicauda 125

opterenfauna Westdeutschlands. — Decheniana References 97 (B): 147—167. Albarda, H., 1889. Catalogue raisonné et synony- Csoknya, M. & M. Ferencz, 1972. A study of mique des Névroptères observés dans les Pays Bas Palingenia longicauda Oliv, in the zoobenthos of et les pays limitrophes. — Tijdschrift voor Ento- the Tisza and Maros (Ephemeroptera). — Tiscia mologie 32: 211—376. 7: 47—57. Antipa, G., 1909. Fauna ichtiologicä a României. — Csoknya, M. & K. Halasy, 1974. Data on the distribu- Publicafiunile fondului "Vasilie Adamachi" 16: tion of mayfly larvae (Ephemeroptera). — Tiscia 248. — Academia Romàna, Bucurefti. 9:71—75. Arion, E. Prunescu-, L. Elian & M. Baltac, 1965. In- Csongar, G. & L. Moczar, 1954. A. Tiszaviràg. — fluenta viiturilor Dunarii asupra chimismului §i Müzeumi Fiizetek 6: 1 — 32. biologici girlei Saltava §i canalului Filipoiu din Demoulin, G., 1965. Contribution a l'étude des Palin- lanca inundabilä. — Hidrobiologia, Bucure§ti 6: geniidae (Insecta, Ephemeroptera). — Nova Gui- 151 — 167. nea, Zoology 33: 305—344.

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Brtek, J. & J. Rothschein, 1964. Ein Beitrag zur ura Academia Republicii Socialiste Romania. Kenntnis der Hydrofauna und des Reinheitszu- Ferencz, Sz., 1956. Untersuchungen des Fisch-Darm- standes des Tschechoslovakischen Abschnittes der inhaltes in den Gewässern von Szeged. — Acta Donau. — Biologické Prâce 10 (5): 5 —60. Universitatis Szegediensis (Acta Biologica) 2 Bures, I., Prinos kam izucavaneto na mrezokrilnata (1—4): 167—182. fauna na Balgarija (Insecta, Neuroptera). — Izves- Gheracopol, O. &^M. Selin, 1968. Contributii la Stu- tija na balgarskoto entomologicno druzestvo 9: diul Biologiei Cegil din Cursul Inferior al Dunärii 135 — 150. (Acipenser ruthenus L.). IL Regimul Alimentar al Burmeister, H., 1839. Handbuch der Entomologie II Cegil din Zona Galati. — Buletinul Institutului (2): 788—804. Cercetàri Proiectâri Piscicole 27 (4): 16 —22. Burnita, Th., Gh. Brezeanu & E. Prunescu-Arion, Gheracopol, O., M. Selin & G. Munteanu, 1969. Con-

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Vlijanie vod Dîmbovitî i Ardzesa na vodi Dunaja. Cercetàri Proiectâri Piscicole 28 (1): 67 — 80. — Revue de Biologie 6 (2): 199— 212. Gheracopol, O., M. Selin & G. Munteanu, 1970. Con- Clutius, A., 1634. De Hemerobio sive Ephemero in- tributii la Studiul Biologiei Pietrarului (Aspro zin- secto et majali verme: 96, 100. — Amsterdam. gel Linné, 1758) din Cursul Inferior al Dunarii. — Cornelius, C, 1848. Beiträge zur näheren Kenntnis Hidrobiologia, Bucureçti 11: 143 — 153. der Palingenia longicauda Oliv.: 1 — 37. — Büsch- Gorove, L., 1819. Eggy különös tüneménynek, az udy ler'sche Verlagsbuchhandlung, Elberfeld. nevezett Tiszavirgza-sânak leitràsa. — Tudomâ- Cremer, E., 1938. Beitrag zur Kenntnis der Ephemer- nyos Gyüytemény 8: 3 — 22. — —

126 Tijdschrift voor Entomologie, deel 130, 1987

Hagen, H., 1859. Über das Vorkommen von servationibus geographicis, astronomicis, hydro- Palingenia longicauda in Preußen. — Stettiner En- graphicis, historicis, physicis. VI, De insectis: 125, tomologische Zeitung 20: 431. — Haga, Amsterdam, Hagen, H., 1888. Unsere gegenwärtige Kenntnisse Mikulski, 1936, (Ephemeroptera), — Fauna J,, Jefki der Ephemeren. — Stettiner Entomoiogische Zei- slodcowodna Polska: Wydawnictwo Kasy Imenia tung 49: 221. Mionowskiego Instytutu Popierania Nauki, 'Wars- Ikonomov, P., 1958. Preliminary notes on the nymphs zawa, of Ephemeroptera found in Macedonian waters. — Mocsâry, A,, 1900, Ungarns Neuropteren (Magyaror- Verhandlungen der Internationalen Vereinigung sag Neuropteràe), — Természetrajzi Füzetek 13: für theoretische und angewandte Limnologie, 108—116, Stuttgart 13: 858—859. Mocsâry, S., 1875, Harcsaféreg Györ meilett, — Ter- (Eintagsfliegen). — lilies, J., 1968. Ephemeroptera mészettu-domânyi Közlöny 7: 409, In: Handbuch der Zoologie 4 (2) 2 (5): 1—63. — Motas, G,, 1936, Zborul nuptial al "Vetritzelor" sau Berlin, "Rusalülor" la Ada-Kaleh, — Revista Stiintifica Illiger, 1802. Magazin für Insektenkunde 1 (17): 187. "V, Adamachi" 22: 2, Jacob, U., 1977. Palingenia anatolica n.sp. (Ephemer- Motas, C, & M, Bâcesco, 1937, La découverte en Rou- optera, Palingeniidae) aus der Türkei. — Entomo- manie d'une nymphe d'éphémère appartenant au

logische Nachrichten 21 (12): 177—182. genre Behningia J, A. Lestage 1929. — Annales Jankovic, D., 1958. Ekologija Dunavské kecige Scientifiques de l'Université de Jassy 24 (2): (Acipenser ruthenus L.). — Bioloski Institut 25—29. N.R.Srbye 2: 145. Müller, K., 1954, Investigations on the organic drift Keller, A., 1975. Die Drift und ihre ökologische Be- in North-Swedish streams, — Report of the Insti- deutung. Experimentelle Untersuchung an tute of Freshwater Research, Drottningholm 35: Ecdyonurus venosus (Fabr.) in einem 133—149, Fließwassermodel. — Schweizerische Zeitschrift Müller, K,, 1973. Life cycles of stream insects. — für Hydrologie 37 (2): 294—331. Aquilo, ser. Zoologica 14: 105 — 112.

Kinel, J., A. Krasucki & J. Noskiewicz, 1927. Owady Müller, K., 1982. The colonization cycle of freshwater krajowe. — Przewodnik okréslania vzedów, rodzin insects. — Oecologia, Berlin 52: 202 —207.

i rodzajow, Lwów, Warszawa, Krakow, Olivari, G,, 1961, Bentos sovetskogo ucastka Dunaja,

Kolarov, P., 1959. Materiali vârhu hranata na cigata In: Dunai i pridunaiskie vodoemi v predelah SSSR. ot bälgarskoto krajbrezie na r. Dunav, — Priroda, — Trudi Instituta Gidrobiologii 36: 145 — 165, — Sofia 4: 62 — 65. Izdatelstvo Akademia Nauk USSR, Kiev, Landa, VI., 1969. Jepice — Ephemeroptera, In: Fauna Olivier, A, G,, 1791. Encyclopédie méthodique. In- CSSR 18: 1 — 352. — Ceskoslovenska Akademie sectes 6: 18. Praha. Ved, Panzer, 1804. J. D. Schaefferi Icon. Inst. Ratisb. Enum. Latreille, P., 1805. Histoire naturelle, generale et par- Syst. 177. ticulière, des Crustacées et des Insectes: 96. 13, Rothschein, J., 1959. Palingenia longicauda Oliv. Lauterborn, R., 1918. Die geographische und biolo- (Ephemeroptera) na vychodnom Slovensku. — Bi- gische Gliederung des Rheinstroms, III Teil. — ologia, Casopis Slovenska Academia vied 14 (2): Sitzungsberichte der Heidelberger Akademie der 139—141. Wissenschaften 1: 40. Russev, B., 1957. Larvite na vodnite nasekomi - os- Lestage, J.-A., 1922. Catalogue des Ephémères de novna hrana na cigata po bâigarskija brjag na Dun- France. — Annales de la Société Entomologique de ava, — Ribno stopanstvo, Sofia 1: 37 —40, France 91: 273—276. Russev, B,, 1959, "Vol de compensation pour la Lestage, J.-A., 1923. Étude sur les Palingeniidae ponte" de Palingenia longicauda (Oliv,) (Ephem,) (Ephémères) et description de deux genres nou- contre Ie courant du Danube, — Comptes rendus veaux et d'une espèce nouvelle de la Nouvelle Gui- de l'Académie bulgare des Sciences 12 (2): née. — Annales de la Société Entomologique de 165 — 168, Belgique 63: 95—112. Russev, B,, 1963, Hrana na cigata {Acipenser ruthe- Lestage, J.-A,, 1924. Notes sur les Ephémères de la nus L.) V reka Dunav pred bâigarskija brjag. — monographical revision de Eaton. — Annales de la Izvestija Opitnata stancia po sladkovodno ri- Société Entomologique de Belgique 64: 33 —60. bastvo, Plovdiv 2: 49— 72. Lestage, J.-A., 1937. Les Ephéméroptères de Belgique Russev, B., 1966. Hidrobiologicni izsledvanija na reka — IX, Palingenia longicauda Oliv. type disparu ou Marica. I. In: Fauna na Trakija 3: 231 —291. — n'ayant jamais existé. — Bulletin & Annales de la Izdatelstvo na Bâlgarskata Akademija na naukite, Société Entomologique de Belgique 77: 170— 175. Sofia. Markovskii,J,, 1955, Fauna bespozvonocnih nizovjev Russev, B., 1967. Zoobentosât na reka Dunav mezdu

rek ukraini, usiovija ejo sustestvovanija i puti is- 845 ija i 375 ija recen kiiometâr. IL Biocenolog-

polzovanija, III. Vodoemi kiliiskoi delti Dunaja: ija i dinamika. — Izvestija Zoologicski Institut s 157—159. — Izdatelstvo Akademia Nauk USSR, muzei, Sofia 23: 33 — 78. Kiev. Russev, B., 1968. Ökologische Untersuchungen über Marsili, L., 1726. Danubius Pannonico-Mysicus, ob- die Ephemeropterenlarven der Donau vor dem — —

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bulgarischen Ufer. — Limnologische Berichte der Entomologique de Belgique 32: 147. X. Jubiläumstagung Donauforschung, Sofia: Soldan, T., 1978. Revision of the genus Palingenia in 295—303. Europe (Ephemeroptera, Palingeniidae). — Acta Russev, B., 1973. Kompensationsflug bei der Ord- entomologica bohemoslovaca 75: 272 — 284. nung Ephemeroptera. — Proceedings of the First Strenger, A., 1973. Die Mandibelgestalt der Epheme- International Conference on Ephemeroptera, ridenlarven als funktionsmorphologisches Pro- 1970: 132—142. Leiden, Brill. blem. — Verhandlungsberichte der Deutschen Russev, B., 1977. Einfluß der Corioliskraft auf die Zoologischen Geselschaft 66: 75 — 79. — Gustav Breitenverteilung der Bodenablagerungen und der Fischer Verlag.

zugehörigen Biozönosen im bulgarischen Do- Swammerdam, J., 1752. Bibel der Natur (Haft, Ufer- nauabschnitt. — Archiv für Hydrobiologie, Suppl. aas, 100—114, Tab. 13—15). —Johann Friedrich 52 (Donauforschung 6) 1: 23 — 31. Gleditschens Buchhandlung, Leipzig.

Russev, B., 1978. Osobenosti i znacenie na zoobentosa Szent-Ivany J. & E. Ujhazy, 1973. Ephemeroptera in

na reka Dunav mezdu 845 ija i 375 ija recen the regiment of some New Guinean People and in kilometär. — In; Limnologija na bälgarskija sektor Hungarian folksongs. — Eatonia 17: 1 — 6. na reka Dunav: 145 — 306. — Izdatelstvo na Triepke, 1840. Einige Bemerkungen über Ephemera Bälgarskata Akademija na naukite, Sofija. flos-aque. III. — Entomologische Zeitung 1: jepice. Samal, J., 1935. Dvé zajîmavé ceskoslovenské 54—58. — Veda pfirodni, Praha 16: 184—186. Tshernova, O., 1949. K. poznaniju roda Palingenia oder der Schäffern, J., 1757. Das fliegende Uferaas Burm. (Ephemeroptera, Palingeniidae). — Ento- Haft (Wegen desselben an Il-ten Augustmon. an mologiceskoe obozrenie 30 (3 —4): 304 — 307. der Donau und sonderlich auf der steinernen Ulmer, G., 1927. Verzeichnis der deutschen Ephe- Brücke zu Regensburg ausserordentlich häufigen meropteren und ihrer Fundorte. — Konowia 6(4):

Erscheinung und Fluges )1 34. — Gebrüdern Zun- 234—262. kel, Regensburg. Unger, F., 1927. Magyar tavak es folyók természetes

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Tijdschrift voor Entomologie 130: 129 — 140 Gepubliceerd 30 november 1987

NEW SPECIES OF THE FIG WASP GENUS DIAZIELLA (HYMENOPTERA, CHALCIDOIDEA, SYCOECINAE)

A. J. GARDINER and S. G. COMPTON

Department of Zoology and Entomology, Rhodes University, Grahamstown, South Africa

Abstract

Eight new species of Diaziella are described from Borneo: alleni, laticeps, latipennis, longiceps, pallidiceps, retakensis, tumidigena, and wiehesi. A key to the known species

of the genus is added.

Introduction Methods The Sycoecinae are a well defined but enig- Specimens were stored in 70% ethanol. For matic group of fig wasps. Although generally detailed examination, examples of each species placed in the Torymidae, Boucek (in Boucek et. were slide mounted using a modified version al., 198 1 ) has suggested that they may have closer of Prinsloo's (1980) technique for the prepa- affinities with the Agaonidae. Diaziella Grandi ration of permanent slide mounts. is the only genus of the subfamily described from outside Africa, although representatives of a Key to the known species of Diaziella further, undescribed, genus have been collected (females) in Australia (Wiebes, pers. comm.). Grandi 1. Head yellow in colour; epistomal margin (1928) based Diaziella on two new species from with three slight protuberances, one in the Sumatra (D. bicolor and D. macroptera). Wiebes centre and one on either side of the central (1974) subsequently added a further two species protuberance (the protuberances being (D. philippinensis and D. falcata), both from broadly separated) (fig. 12) the Philippines. With the exception of D. falcata, pallidiceps spec. nov. the species are known only from the female sex. — Head brown or black in colour, epistomal

Material collected at light by B. Allen on the margin if having incisions or protuberances island of Borneo has been found to contain fe- then not as above (for example figs. 1, 2, males of a further eight speecies. This paper 19) 2 describes these new species of Diaziella and pro- 2. Epistomal margin with the central region vides a key to the known representatives of the forming an inverted w, the top of the w genus. leveling off slightly and then rising to the

Certain features are of particular value for dis- cheek margin (fig. 1). First funicle segment tinguishing between the species, these are: of the antenna with irregular sensilla; re-

1. The epistomal margin of the clypeus, which maining funicle segments with two slightly

is variable in shape and form. irregular transverse rows of sensilla. Sub-

2. The fore wings, which vary considerably in marginal to marginal ratio 1.8 : 1 or greater shape. (fig. 7) philippinensis 3. The mandibles. These have one or more cusps — Epistomal margin not as described above (for

on the subapical tooth. example figs. 2, 43, 45). Funicle segments 4. The hypopygium, which protrudes beyond the of antenna with one row of transverse sen- gaster to a varying extent. silla, or at most with the first and second

129 130 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 1, 7. Diaziella philippinensis Wiebes, female. 1, head; 7, fore wing. Figs. 2, 8. Diaziella macroptera Grandi, female (redrawn from Wiebes, 1974). 2, head; 8, fore wing. Figs. 3, 5. Diaziella latipennis spec, nov., female. 3, first five antennal segments; 5, fore wing. Fig. 6. Diaziella alleni spec, nov., fore wing. Figs. 9, 1 1. Diaziella bicolor Grandi, female (redrawn from Grandi, 1928). 9, fore wing; 11, mandible. Fig. 10. Diaziella falcata wiebes, female, hypopygium (redrawn from Wiebes, 1974). Gardiner & Compton: The fig wasp genus Diaziella 131

funicle segments having two or three rows. (fig. 8). Epistomal margin indented and with Submarginal to marginal ratio less than 1.8 two slight projections on either side of a

: 1 (figs. 5, 6, 9) 3 small central incision (fig. 2) macroptera

3. Antenna; Pedicel longer than the first funicle — Fore wing less than three and a half times segment (fig. 3); first funicle segment with as long as wide (fig. 67). Epistomal margin one irregular transverse row of large sensilla either slightly protruding with a central in- (fig- 3) .4 cision (fig. 53) or indented but without a — Antenna: Pedicel shorter than first funicle central incision (fig. 61) 9 segment (fig. 4); first funicle segment with 9. Epistomal margin slightly protruding and more than one row of sensilla, which may with a central u-shaped incision (fig. 53) form definite or integrated rows, but clearly head more rounded in shape). Mandible bi- not one row (fig. 4) 10 dentate tricuspidate; length approximately 4. Basal portion of stigmal vein almost per- two and a half times the width (fig. 54) .... pendicular to the wing margin (fig. 5). Wing alleni spec. nov. club shaped, its length less than two and — Epistomal margin indented and with a very a half times the width (fig. 5) 5 slight central protuberance (fig. 61) (head — Basal portion of stigmal vein at an acute more square in shape). Mandible bidentate angle to the wing margin (fig. 6). Wing more bicuspidate (the subapical tooth may have elongate, its length more than two and a a small protuberance on the inner margin); half times the width (fig. 6) 8 length almost four times the width (fig. 62)

5. Fore wing with few microtrichia, almost gla- wiebesi spec. nov. brous. Hypopygium just projecting beyond 10. Antennal insertion clearly closer to the ep- the end of the abdomen (fig. 10) ... falcata istomal margin than to the vertex (fig. 35).

— Fore wing with many microtrichia (fig. 17). Head clearly longer than broad (1.13 : 1) Hypopygium clearly protruding beyond the (fig. 35). Epistomal margin with a central end of the abdomen (fig. 66) 6 incision (fig. 35). Mandible bidentate tricus-

6. Two dark patches on the fore wing, one pidate, length approximately four times the below the stigmal vein and adjacent to the width (fig. 36) longiceps spec. nov. marginal vein the other above the stigmal — Antennal insertion slightly or distinctly vein (fig. 9). Mandible with subapical tooth closer to the vertex than to the epistomal tricuspid (fig. 11) bicolor margin (figs. 19, 27). Head as long as broad

— One dark patch on the fore wing, below the or slightly longer than broad (max 1.04 : stigmal vein and adjacent to the marginal 1) (Figs. 19, 27). Epistomal margin with a vein (fig. 5). Mandible with subapical tooth central truncate protuberance (figs. 19, 27). monocuspid or bicuspid (figs. 44, 46) .... 7 Mandible bidentate bicuspidate, length more 7. Head with the vertex raised towards the than six times the width (figs. 20, 28) 11 ocelli (fig. 45); epistomal margin with small 11. Mandible extremely long and narrow, the cone-like central protuberance (fig. 45). length more than nine times the width (fig. Mandible bidentate tricuspidate and three 28). Antennal insertion clearly closer to the times as long as wide (fig. 46). Antennal vertex than to the epistomal margin (fig. insertion clearly above ventral margin of the 27). Epistomal margin with a broad truncate eye (fig. 45) latipennis spec. nov. protuberance (width approximately one — Head with vertex truncate (fig. 43), epis- third the width of the head) (fig. 27); cheeks tomal margin more indented, with a central diverging towards the ventral margin (fig. (but not cone like) protuberance. The sides 27) tuniidigena spec. nov. of the protuberance smooth and leading up — Mandible length six to seven times longer to a fairly sharp central point (fig. 43). Mand- than wide (fig. 20). Antennal insertion ible bidentate bicuspidate, with a slight pro- slightly closer to the vertex than to the ep- tuberance on the inner margin of the sub- istomal margin (fig. 19). Epistomal margin apical tooth; four times as long as wide (fig. with a smaller truncate protuberance (width 44). Antennal insertion just above ventral approximately one seventh the width of the margin of the eye (fig. 43) head) (fig. 19); cheeks parallel or converging laticeps spec. nov. towards the ventral margin (fig. 19) 8. Fore wing almost four times as long as wide retakensis spec. nov. 132 Tijdschrift voor Entomologie, deel 130, 1987 Gardiner & Compton: The fig wasp genus Diaziella 133

Diaziella pallidiceps Gardiner spec. nov. (NH) London, some paratypes in RMNH (figs. 12-18) Leiden.

Female. — Head (fig. 12) yellow in colour; almost quadrate, slightly wider than long (1.15 Diaziella retakensis Gardiner spec. nov. (figs. 19—26) : 1); length of the compound eye approximately (fig. dark to black 2.5 times the length of the cheek (2.6 : 1); eyes Female. — Head 19) brown protruding laterally; epistomal margin with in colour; almost quadrate, slightly longer than

three slight protuberances one in the centre and wide (1.05 : 1) or as long as wide; length of the one on either side of the central protuberance. compound eye almost twice that of the cheek laterally; Antenna (fig. 15) eleven segmented: antennal (1.73 : 1); eyes slightly protruding ep- insertion closer to the epistomal margin than istomal margin with a truncate protuberance the to the vertex; scape almost four times as long width of which is an approximately one seventh

as wide (3.75 : 1) and three times the length the width of the head (epistomal margin below of cheek). (fig. of the pedicel (3 : 1); pedicel shorter than the the ventral margin the Antenna first funicle segment; funicle, first segment 23) eleven segmented; antennal insertion to : to than the epistomal longer than the others (29 : 24 ; 23 : 23 24), slightly closer the vertex first funicle segment with two transverse rows margin (distance ratio 4:5); scape four times as

of long sensilla; remaining segments including long as wide (3.9 : 1) and about three times the

the three segmented club, with one row of sen- length of the pedicel (2.81 : 1); pedicel shorter silla each. Mandible (fig. L3) bidentate, subapical than the first funicle segment; funicle, first seg-

tooth longer than the apical tooth; mandible ment longer than the remaining four (30 : 22 :

about four times longer than wide (3.9:1). Max- 23 : 22 : 23), first funicle segment with two rows

illary palp (fig. 14) three segmented (6 : 8 : of long sensilla; remaining segments including 9); labial palp (fig. 14) two segmented (1:1). the three segmented club with one row of sen- Thorax. Fore wing (fig. 17) 1.33 mm long; silla. Mandible (fig. 20) bidentate, bicuspidate, length approximately 2.5 times the width (2.55 subapical tooth more robust than the apical but of the same length; mandi- : the approximately 1 ) ; marginal vein twice the length of stigmal both

90° as long as wide (.6.77 : (2.04 : 1); stigmal vein arising at almost ble more than 6.5 times

to the wing margin; submarginal 1.5 times the 1). Maxillary palp (fig. 21) three segmented (5 :

length of the marginal (1.58 : 1); membrane 9 : 9); labial palp (fig. 22) two segmented (1 : 1). with dense microtrichia. Hind wing (fig. 18) 0.83 Thorax. Fore wing (fig. 25) 1.33 mm long; mm long; length about 4.5 times the width (4.6 length approximately 2.5 times the width (2.48

: 1 marginal vein twice the length of the stigmal : 1). Fore leg: tibia with one dorsal-apical tooth, ) ; 90° to one ventro-apical tooth and one spur; tibia to (2 : 1); stigmal vein arising at almost the and a half times tarsal ratio 0.83 : 1; tarsal segments in ratio wing margin; submarginal one

: membrane 23 : 12 : 8 : 24. Mid leg: tibia with one ventro- the length of the marginal (1.5 1); apical spur. Hind leg: tibia with two spurs (one with dense microtrichia. Hind wing (fig. 26) 0.82

much smaller than the other); tibia with a row mm long; length 4.5 times the width (4.56 : 1). of dorsal cones (usually three), tarsal segments Fore leg: tibia with two dorso-apical teeth, one ventro-apical spur; ; and one in ratio 36 : 35 : 30 : 12 26. ventro-apical tooth

Gaster. Hypopygium (fig. 16) length 0.5 1 mm, tibia to tarsal ratio 1:1; tarsal segments in ratio

with many spines and extending well beyond 14 : 7 : 4 : 14. Hind leg: tibia with two ventro- end of abdomen. Pygostyle with four long setae. apical spurs (one much smaller than the other); Total length 2.21 mm. tibia with a row of dorsal cones (usually six)

tarsal segments in ratio 32 : 37 : 27 : 20 : 27. Type material. — Holotype $ (slide mounted), Gaster. Hypopygium (fig. 24) 0.51 mm long, Brunei, Mt. Retak, iv.l981, leg. B. Allen, at light. with many spines and clearly extending beyond Paratypes: 6 $, same data as holotype (coll. dates end of abdomen. Pygostyle with four long setae. 23 — 28. iv. 1981). Holotype to be deposited in BM Total length 1.98 mm.

Figs. 12 — 18. Diaziella pallidiceps spec, nov., female. 12, head; 13, mandible, ventral aspect; 14, labiomaxillary complex; 15, antenna; 16, hypopygium; 17, fore wing; 18, hind wing. Figs. 19— 26. Diaziella retakensis spec, nov., female. 19, head; 20, mandible, ventral aspect; 21, maxillary palp; 22, labial palp; 23, first five antennal segments; 24, hypopygium; 25, fore wing; 26, hind wing. deel 130, 1987 134 Tijdschrift voor Entomologie, Gardiner & Compton: The fig wasp genus Diaziella 135

Type material. — Holotype $ (slide mounted), : 37. Hind leg: tibia with two ventro-apical spurs Brunei, Mt. Retak, iv.l981, leg. B. Allen, at light. (one much smaller than the other); tibia with a Paratypes: 9 9i same data as holotype (coll. dates row of dorsal cones (usually six) tarsal segments

23 — 28. iv. 1981). Holotype to be deposited in BM in ratio 50 : 56 : 36 : 25 : 36. (NH) London, some paratypes in RMNH Gaster. Hypopygium (fig. 32) 0.62 mm long, Leiden. with many spines, its projecting portion extremely long. Pygostyle with four long setae. Diaziella tumidigena Gardiner spec. nov. Total length 2.64 mm. (figs. 27—34)

Female. — Head (fig. 27) dark brown to black Type material. — Holotype $ (slide mounted), in colour; bell shaped, slightly longer than wide Sarawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B.

(1.04 ; 1); length of the eye slightly more than 1.5 Allen, at light. Paratypes: 2 $, same data as hol- times the length of the cheek (1.63 : 1); eyes not otype. Additional material: 1 $, Brunei, Mt. protruding beyond lateral margin of the cheek; Retak, iv.l981, leg. B. Allen, at light. Holotype to cheek diverging towards the epistomal margin; be deposited in BM (NH) London, a paratype in epistomal margin with an almost truncate pro- RMNH Leiden. tuberance the width of which is approximately one third the width of the head (the epistomal Diaziella longiceps Gardiner spec. nov. margin protuberance may protrude beyond the (figs. 35—42) ventral margin of the cheek). Antenna (fig. 31) Female. — Head (fig. 35) dark brown in co-

eleven segmented; antennal insertion clearly lour; distinctly longer than wide (1.15 : 1); length closer to the vertex than to the epistomal margin of eye between 1.5 — 2 times the length of the

(distance ratio 0.58 : 1); scape over four times as cheek (1.74 : 1); eyes protruding laterally; epis- long as wide (4.37 : 1) and about three times as tomal margin having a central incision with long as the pedicel (3.07 : 1); pedicel shorter than stepped sides. Antenna (fig. 39) eleven seg- the first funicle segment; funicle, first segment mented; antennal insertion closer to the epis- longer than the remaining four and with two tomal margin than to the vertex; antennal inser- rows of long sensilla, second segment longer tion well above the ventral margin of the eye;

than the remaining three and with two irregular scape 3.5 times as long as wide (3.5 : 1) and rows of long sensilla (37 : 31 : 28 : 28 : 28); almost three times the length of the pedicel (2.8 remaining segments including the three seg- : 1); pedicel shorter than the first funicle segmented club with one row of sensilla. Mandible ment; funicle, first segment longer than the re- (fig. 28) bidentate, bicuspidate, subapical tooth maining four and with three irregular rows of more robust than the apical but both of approx- long sensilla, second segment longer than any of imately the same length; extremely long and nar- the remaining three and with two irregular rows

row almost eleven times as long as wide (10.87 of long sensilla (40 : 34 : 31 : 30 : 31); remaining 1). Maxillary palp (fig. 29) three segmented (9 segments including the three segmented club

12 : 11); labial palp (fig. 30) two segmented (9 with one row of sensilla each. Mandible (fig. 36) 7). bidentate, tricuspidate, all cusps large with upper Thorax. Fore wing (fig. 33) 1.9 mm long; cusp of subapical tooth slightly larger than apical length almost 2.5 times the width (2.44 : 1); tooth; almost 3.5 times longer than wide (3.3 : 1).

marginal vein twice the length of the stigmal Maxillary palp (fig. 37) three segmented (5 : 7 :

(2.19 : 1); submarginal 1.5 times the length of the 7); labial palp (fig. 38) two segmented (9 : 7). marginal (1.49: 1 ) ; membrane with dense micro- Thorax. Fore wing (fig. 41) 1.71 mm long; trichia. Hind wing (fig. 34) 1.17 mm long; length length between 2 — 2.5 times as long as wide slightly more than four times the width (4.33 : 1 ). (2.31 : 1); marginal vein almost twice the length

Fore leg: tibia with two dorso-apical teeth, one of the stigmal (1.88 : 1); stigmal vein arising at ventro apical tooth and one spur; tibia to tarsal almost 90° to the wing margin; submarginal 1.5 ratio 0.96 : 1; tarsal segments in ratio 34 : 23 : 13 times the length of the marginal (1.5 : 1); mem-

Figs. 27 — 34. Diaziella tumidigena spec, nov., female. 27, head; 28, mandible, ventral aspect; 29, maxillary palp; 30, labial palp; 31, first five antenna! segments; 32, hypopygium; 33, fore wing; 34, hind wing. Figs. 35 —42. Diaziella longiceps spec, nov., female. 35, head; 36, mandible, ventral aspect; 37, maxillary palp; 38, labial palp; 39, first five antennal segments; 40, hypopygium; 41, fore wing; 42, hind wing. 136 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 43, 44. Diaziella laticeps spec, nov., female. 43, head; 44, mandible, ventral aspect. Figs. 45 — 52. Diaziella latipennis spec, nov., female. 45, head; 46, mandible, ventral aspect; 47, maxillary palp; 48, labial palp; 49, first five antennal segments; 50, fore wing; 51, hind wing; 52, hypopygium.

brane with dense microtrichia.. Hind wing (fig. other); tibia with a row of dorsal cones (usually

42) 1.07 mm long; length almost 4.5 times the six) tarsal segments in ratio 44 : 47 : 36 : 22 : 30. width (4.4 : 1). Fore leg: tibia with two dorso- Gaster. Hypopygium (fig. 40) 0.66 mm long, apical teeth, one ventro-apical tooth and one with many spines and extending well beyond the

spur; tibia to tarsal ratio 0.89 : 1", tarsal segments end of the abdomen. Pygostyle with four long

in ratio 33 : 14 : 11 : 31. Hind leg: tibia with two setae. ventro-apical spurs (one much smaller than the Total length 2.44 mm. Gardiner & Compton: The fig wasp genus Diaziella 137

Type material. — Holotype $ (slide mounted), nov., fig. 52) 0.63 mm long with many spines and Brunei, Mt, Retak, iv.l981, leg. B. Allen, at light. extending well beyond the end of the abdomen. Paratypes: 5 $, same data as holotype (coll. dates Pygostyle with four long setae. 23— 28.ÌV. 1981). Additional material: 3 $, Sa- Total length 2.93 mm. Apart from the charac- rawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B. teristic head and mandibles D. laticeps spec. nov. Allen, at light. Holotype to be deposited in BM is very similar to D. latipennis spec. nov. (NH) London, some paratypes in RMNH Leiden. Type material. — Holotype $ (slide mounted), Sarawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B. Diaziella laticeps Gardiner spec. nov. Allen, at light. Paratypes: 12 $, same data as (figs. 43, 44) holotype. Additional material: 2 $, Brunei, Mt. Female. — Head (fig. 43) brown in colour; Retak, 23—28.iv.1981, leg. B. Allen, at light and 2 Brunei, Temburong River, 300 m, almost quadrate, slightly wider than long (1.18 : $,

1); length of the eye slightly more than 1.5 times 26.iv.1981, leg. B. Allen, at light. Holotype to be deposited in London, some paratypes : BM(NH) the length of the cheek (1.7 1 ) ; eyes protruding slightly beyond lateral margin of the head; epis- in RMNH Leiden. tomal margin indented with a central protuberance and with the clypeus sloping back towards Diaziella latipennis Gardiner spec. nov. the posterior of the head. Antenna (as in D. (figs. 45—52) latipennis, fig. 49) eleven segmented; antennal Female. — Head (fig. 45) dark brown in co- insertion closer to the epistomal margin than to lour; vertex raised towards the ocelli; wider than the vertex and slightly above the ventral margin long (1.27 : 1); longitudinal diameter of the eye of the eye; scape four times as long as wide (4.17 not quite 2.5 times the length of the cheek (2.37

: 1) and about 2.5 times the length of the pedicel : 1); eyes protruding laterally; epistomal margin

(2.63 : 1); pedicel longer than the first funicle slightly indented and with a small central cone segment; funicle, first segment longer than the like protuberance. Antenna (fig. 49) eleven seg- remaining four and with one irregular row of mented; antennal insertion closer to the epis- long sensilla (30 : 25 : 22 : 22 : 24); remaining tomal margin than to the vertex and well above segments including the three segmented club the ventral margin of the eye; scape almost 4.5 with one row of sensilla. Mandible (fig. 44) bi- times as long as wide (4.41 : 1) and almost three dentate, bicuspidate, subapical tooth longer than times longer than the length of the pedicel (2.89

the apical and with a slight protuberance on its : 1 pedicel longer than the first funicle segment; ) ; inner margin; about 4.5 times as long as wide (4.6 funicle, first segment longer than any of the re-

: 1). Maxillary palp three segmented (4 : 5 : 2); maining four and with one slightly irregular row labial palp two segmented (1:1) (Maxillary and of long sensilla (30 : 24 : 24 : 25 : 25); remaining labial palps similar in appearance to D. latipen- segments including the three segmented club nis spec, nov., figs. 47 and 48 respectively). with one row of sensilla. Mandible (fig. 46) bi- Thorax. Fore wing (as in D. latipennis spec, dentate, tricuspidate (subapical tooth with two nov. fig. 50) 2.14 mm long; length twice the large cusps), three times as long as wide (3 : 1). width (2 : 1); marginal vein 1.5 times the length Maxillary palp (fig. 47) three segmented (5 : 6 :

of the stigmal (1.5 : 1); stigmal vein arising at 4); labial palp (fig. 48) two segmented (1 : 1). almost 90° to the wing margin; submarginal 1.5 Thorax. Fore wing (fig. 50) 2.3 mm long; times the length of the marginal (1.5 : 1); mem- length twice the width (2 : 1); marginal vein brane with dense microtrichia and a dark mark- almost 1.5 times the length of the stigmal (1.43 90° ing below the stigmal vein (adjacent to the mar- : 1); stigmal vein arising at almost to the ginal vein). Hind wing (as in D. latipennis spec, wing margin submarginal vein 1.5 the length of nov., fig. 51) 1.38 mm long; length almost five the marginal (1.58 : 1); membrane with dense times the width (4.78 : 1). Fore leg: tibia with two microtrichia and a dark marking below the stig- dorso-apical teeth, one ventro-apical tooth and mal vein (adjacent to the marginal vein). Hind one spur; tibia to tarsal ratio 1:1; tarsal segments wing (fig. 51) 1.3 mm long; length between 4.5 in ratio 33 : 21 : 13 : 40. Hind leg: tibia with two to five times longer than wide (4.64 : 1). Fore leg: ventro-apical spurs (one much smaller than the tibia with two dorso-apical teeth, one ventro- other); tibia with a row of dorsal cones (usually apical tooth and one spur; tibia to tarsal ratio 1

six) tarsal segments in ratio 68 : 62 : 43 : 25 : 41. : 1; tarsal segments in ratio 35 : 20 : 12 : 42. Hind Gaster. Hypopygium (as in D. latipennis spec. leg: tibia with two ventro-apical spurs (one much 138 Tijdschrift voor Entomologie, deel 130, 1987

56 53 Gardiner & Compton: The fig wasp genus Diaziella 139

smaller than the other); tibia with a row of dorsal longer than wide (6.43 : 1). Fore leg: tibia with cones (usually four) tarsal segments in ratio 6 : two dorso-apical teeth, one ventro-apical tooth 5:4:2:3. and one spur; tibia to tarsal ratio 0.95:1; tarsal

Gaster. Hypopygium (fig. 52) 0.59 mm long, segments in ratio 27 : 16 : 1 1 : 28. Hind leg: tibia with many spines and clearly projecting beyond with two ventro-apical spurs (one much smaller the end of the abdomen. Pygostyle with four long than the other); tibia with a row of dorsal cones setae. (usually six) tarsal segments in ratio 55 : 47 : 32

Total length 2.73 mm. : 22 : 33. Gaster. Hypopygium (fig. 58) 0.49 mm long, Type material. — Holotype $ slide mounted), with many spines and clearly extending beyond Brunei, Mt. Retak, iv.l981, leg. B. Allen, at light. end of abdomen. Pygostyle with four long setae.

Paratype: 1 $, same data as holotype (coll. dates Total length 2.15 mm. 23— 28.iv.1981). Additional material: 2 $, Sa- rawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B. Type material. — Holotype $ (slide mounted), Allen, at light. Holotype to be deposited in BM Brunei, Mt. Retak, iv.l981, leg. B. Allen, at light. (NH) London. Paratypes: 7 $, same data as holotype (coll. dates 23—28.iv.1981). Additional material: 2 $, Sa- Diaziella alleni Gardiner spec. nov. rawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B. (figs. 53—60) Allen, at light. Holotype to be deposited in BM Female. — Head (fig, 53) brown in colour; (NH) London, some paratypes in RMNH almost quadrate; slightly wider than long (1.08 : Leiden.

1); length of the eye twice the length of the cheek Diaziella wiebesi Gardiner spec. nov. (2.12 : 1); eyes protruding beyond the lateral (figs. margins of the head; epistomal margin initially 61—68) protruding and then forming a large central u Female. — Head (fig. 61) dark brown to black shaped incision. Antenna (fig. 57) eleven seg- in colour; almost quadrate, wider than long ( 1.25 mented; antennal insertion closer to the epis- : 1 ) ; length of the eye slightly more than twice the tomal margin than to the vertex; scape 4.5 times length of the cheek (2.26 : 1); eyes protruding

as long as wide (4.65 : 1) and just over 2.5 times beyond the lateral margin of the head; epistomal the length of the pedicel (2.72 : 1); pedicel longer margin indented and with a slight central prom- than the first funicle segment; the funicle seg- inence. Antenna (fig. 65) eleven segmented; an-

: tennal closer to ments approximately the same size (10 : 9 : 9 insertion the epistomal margin 9:9); all the funicle and club segments with one than to the vertex; scape almost five times longer row of long sensilla. Mandible (fig. 54) very ro- than wide (4.93 : 1) and 2.5 times the length of bust; bidentate tricuspidate, upper cusp of subap- the pedicel (2.55 : 1); pedicel longer than the first ical tooth larger than the apical tooth and lower funicle segment; first funicle segment slightly

cusp of subapical tooth approximately the same longer than the remaining four (19 : 16 : 16 : 16 length as the apical tooth, length 2.5 times the : 17); all funicle segments and the three seg- width (2.58 : 1). Maxillary palp (fig. 55) three mented club with one row of long sensilla. Mand- segmented (5 : 5 : 6); labial palp (fig. 56) two ible (fig. 62) bidentate, bicuspidate, subapical segmented (7 : 6). tooth clearly longer than the apical (a small pro- Thorax. Fore wing (fig. 59) 1.84 mm long; tuberance may be present on the inner margin of length 2.5 times the width (2.57 : I); marginal the subapical tooth); length four times the width vein slightly more than 1.5 times the length of (4.09 : 1). Maxillary palp (fig. 63) three seg- the stigmal (1.68 : 1); stigmal vein arising at an mented (11 : 17 : 14); labial palp (fig. 64) two acute angle to the wing margin; submarginal vein segmented (16 : 11). slightly longer than the marginal (1.22 : 1); Thorax. Fore wing (fig. 67) 1.98 mm long; membrane with dense microtrichia. Hind wing length almost three times the width (2.87 : 1); (fig. 60) 0.93 mm long; narrow almost 6.5 times marginal vein between 1.5 — 2 times the length

Figs. 53 —60. Diaziella alleni spec, nov., female. 53, head; 54, mandible, ventral aspect; 55, maxillary palp; 56, labial palp; 57, first five antennal segments; 58, hypopygium; 59, fore wing; 60, hind wing. Figs. 61 —68. Diaziella wiebesi spec, nov., female. 61, head; 62, mandible, ventral aspect; 63, maxillary palp; 64, labial palp; 65, first five antennal segments; 66, hypopygium; 67, fore wing; 68, hind wing. 140 Tijdschrift voor Entomologie, deel 130, 1987

Acknowledgements of the stigmal (1.72 : 1); stigmal vein arising at an acute angle to the wing margin; submarginal Ben Allen collected the insects in Borneo.

slightly longer than the marginal (1.29 : 1); Thanks also to I.C. Sharp and A. Pretorius for

membrane with dense microtrichia. Hind wing their assistance. Prof. J. T. Wiebes and Dr F. Gess (fig. 68) 0.93 mm long; narrow the length about for providing valuable comments and Dr D. Hill

seven times the width (6.89 : 1). Fore leg: tibia for the loan of material. with two dorso-apical teeth, one ventro-apical References toothh and one spur; tibia to tarsal ratio 1 : 1.07; Boucek, Z., A. Watsham & T. Wiebes, 1981. The fig tarsal segments in ratio 23 : 12 : 7 : 22. Hind leg: J. wasp fauna of the receptacles of Ficus thonningii tibia with two ventro-apical spurs (one much (Hymenoptera, Chalcidoidea). — Tijdschrift voor smaller than the other) ; tibia with a row of dorsal Entomologie 124: 149— 233. cones (usually six); tarsal segments in ratio 48 : Grandi, G., 1928. Un nuovo genere quattro nuove

44 : 30 : 19 : 27. specie di Imenotteri sicofili di Sumatra. — Bollet- Gaster. Hypopygium (fig. 66) 0.45 mm long, tino del Laboratorio di Entomologia del R. Istituto with many spines and clearly extending beyond superiore agrario di Bologna 26: 71 —89- the end of the abdomen. Pygostyle with four long Prinsloo, G. L., 1980. An illustrated guide to the families of African Chalcidoidea (Insecta: Hymenop- setae. tera). — Republic of South Africa Department of Total length 2.08 mm. Agriculture and Fisheries Science Bulletin 395: 1—66. Type material. — Holotype (slide mounted), T., fig $ Wiebes, J. 1974. The wasp genus Diaziella Brunei, Mt. Retak, iv.l981, leg. B. Allen, at light. Grandi (Hymenoptera Chalcidoidea, Torymidae Paratypes: 61 9. same data as holotype (coll. Sycoecini). — Proceedings of the Koninklijke Ne- dates 23—28.iv.1981). Additional material: 2 $, derlandse Akademie van Wetenschappen, Series C 77: 295—300. Sarawak, Mt. Pargon, 1850 m, 29.iv.1981, leg. B. Allen, at light. Holotype to be deposited in BM (NH) London, some paratypes in RMNH Leiden. Tijdschrift voor Entomologie 130: 141 — 175 Gepubliceerd 30 november 1987

EXTERNAL MORPHOLOGY OF ADULT SYRPHIDAE (DIPTERA)

M. C. D. SPEIGHT

Research Branch, Forest and Wildlife Service, Sidmonton Place, Bray, Co. Wicklow

Contents up a jungle of conflicting terminologies. Faced by the apparent inability of morphologists to Introduction 141 agree on the morphological terms which can be The head 142 applied to many of the taxonomically important Compound eyes 142 parts of adult flies, and also finding need to refer Head capsule: frontal, genal and vertical re- to parts which to morphologists seemingly had gions 144 no individual morphological identity, taxono- Head capsule: posterior surface 145 mists have frequently been forced to coin quasi- Antennae 145 morphological terms of their own. Workers on Mouthparts 146 Syrphidae have been particularly plagued by this The cervical region 148 problem since morphologists have concentrated The thorax 150 their efforts principally upon Nematocera and Prothorax 150 Calypterates, leaving Syrphid taxonomists to de- Mesothorax: mesonotum 152 cide for themselves which of the sclerites found Mesothorax: mesopleura 155 in Syrphidae could be identified with those Mesothorax: mesosternum 156 named in other Diptera. The resulting termi- Metathorax 156 nological chaos reaches an extreme in accounts Wings 158 of European Syrphidae, where writers in dif- Legs 161 ferent languages have developed partly indé- The abdomen 161 pendant terminologies but have also "borrowed" Male abdomen 163 terms (sometimes in translation and sometimes Male preabdomen 163 not) from each other, adding to this mélange Male postabdomen 164 a sprinkling of polysyllabic latinisms when all

Male postabdomen: components of the hypo- else failed. At this juncture, it is doubtful that pygium 166 all Europeans working on Syrphidae would adopt Female abdomen 168 in its entirety any proposed set of terms. The relationship between the Syrphidae and The present text does not attempt to select genera allied to Microdon 169 a definitive set! Certain principles have been em- Abbreviations used in figures 172 ployed here as follows: Bibliography 174

i) where the homology of a sclerite is generally Introduction agreed among morphologists and the scler- Dipteran morphology has been surprisingly ite is normally referred to in its entirety little studied. Recent reviews of the available when used by taxonomists the latin (or information are provided by McAlpine (1981) greek) morphological term for the sclerite and in the sections on Diptera in the volumes is used e.g. cercus; by Bitsch et al. (1973), Bitsch & Matsuda (1979) ii) where a sclerite has an agreed homology, and Matsuda (1965, 1970, 1976). From these morphologically, but is not normally re- texts conflicting theories as to the homologies ferred to in its entirety in taxonomie texts, of sclerites found in different Diptera are all the terms used by taxonomists for each in- too evident. And with the theories has grown dividual part of the sclerite are used if they

141 142 Tijdschrift voor Entomologie, deel 130, 1987

are rendered in latin or greek and do not by Gouin (1949) and Schiemenz (1957). Nayar imply an incorrect homology for the sclerite (1964) considered Episyrphus balteatus and

e.g. post-alar callus; Crampton ( 1942) figured the head of a N. Amer-

iii) where the homology of a sclerite is disputed ican Rhingia species.

but there is in existence for the sclerite a The major feature of the Syrphid head are latin or greek term which does not imply the compound eyes and the mouthparts, which

a particular homology that term is used e.g. occupy, respectively, the lateral and ventral sur- basale; faces of the head capsule. The front of the head,

iv) where the homology of a sclerite is disputed from mouth-edge to ocellar triangle, is rather and no suitable latin or greek term is avail- featureless, its main variation being in the extent

able for that sclerite, a neutral term derived to which it projects either ventrally to accom-

from some other language is used e.g. modate the mouthparts or dorsally to produce barrette. an antennal tubercle. The occipital region starts abruptly immediately behind the eyes, almost The literature search conducted as part of the without exception forming a sharp angle with present work revealed only one complete pub- the sides of the head. lished account of the external morphology of As Snodgrass (I960) has pointed out there an adult Syrphid — the unillustrated text by is a great difference between joints between Nayar (1965), on Episyrphus balteatus. Cramp- sclerites (Sutures) and intrascleritic invagina- ton (1942) considers the morphology of various tions of the exoskeleton (Sulci) that form sites Syrphids, but does not illustrate all the parts for muscle attachment, but these two phenomena of any one species. can be indistinguishable externally. In Syrphids

The present account is based on the morphol- the head capsule exhibits on its surface various ogy of the type species of the type genus of grooves marking the location of internal apo- the family, Syrphus ribesii L., augmented by com- demes and most authors referred to these parison with European representatives of the grooves as "sutures", thus not differentiating other two main subdivisions of the family, the them from features representing joint lines. In Eristalinae (exemplified by Eristalis tenax L.) order to make the necessary distinction, these and the Microdontinae (exemplified by Mi- cephalic grooves are here called sulci, following crodon mutabilis (L.)). Other species are men- Snodgrass's (I.e.) terminology. tioned in discussion of particular parts of the fly, so that some idea can be presented of the Compound eyes range of morphological variation found among In Syrphids the two compound eyes are large, Palaearctic Syrphidae. Illustrations are all based occupying most of the top and sides of the head on male specimens, except where specifically and making up two thirds or more of the width stated otherwise. In his massive work defining of the head capsule. There is a certain amount taxonomie subdivisions of the Syrphidae Hull of sexual dimorphism, the male eyes meeting (1949) illustrates a wide range of Syrphidae from in the mid-line between antennal insertions and different parts of the world, amply demonstrat- ocellar triangle (the holoptic condition) in a ma- ing the extremes of form exhibited by the family. jority of the genera, but always remaining se- His similar volume on fossil Syrphidae (Hull, parate (the dichoptic condition) in the females

1945) gives an impression of the morphology (figs. 1, 2). In addition, facets in the upper part of some of the family's antecedents. Studies of of the male eyes may be distinctly larger than individual morphological regions or features of those below (e.g. in Scaeva) while in the female particular hoverfly species, or groups of species, they are of very similar dimensions throughout. have been undertaken by various authors. For Short, straight hairs, inserted between the facets, instance, recently a favoured topic for consid- characterise some genera. These hairs may be eration has been the male abdomen and the mod- generally distributed over most of the surface ifications undergone by its terminal segments. (as in many Cheilosia species) or arranged in Such texts are mentioned in the relevant sections stripes of different density (as in Eristalis tenax) of the present account. or colour (as in Paragus species). In some genera the facets themselves iridesce in bands or spots The head of different colours (as in Eristalodes, Ortho- Detailed morphological studies of the head nevra and Eristalinus) which fade after death. capsule of Eristalis species have been conducted Some authors, e.g. Gouin (1949), have recognised Speight: Morphology of Syrphidae 143

\T 144 Tijdschrift voor Entomologie, deel 130, 1987 an externally visible ocular sclerite forming a each side of the lower frons, from the invag- rim to the eyes, but no external evidence of this ination of each anterior tentorial pit, and more plate has been encountered by the present au- or less parallel with the anterior eye margin, thor. is developed to a greater or lesser extent in many

Syrphid genera. This facial sulcus is almost uni- frontal, and Head capsule: genal versally well-developed in Cheilosia spp (fig. 6),

vertical regions where it reaches the level of the antennal in- Sulci precisely delimiting the frons are absent sertions. In this way, the part of the face between in Syrphidae, leaving the anterior ocellus, the the facial sulcus and the eye margin is cut off compound eyes and the anterior tentorial pits from the rest of the face and also frequently as the only reference points by which to define looks very different, due to differences in colour the extent of the frontal region of the head cap- and pilosity between it and the rest of the face. sule. Judged in this way, the "frons" is generally Some authors have sought to name these orbital regarded as extending from the ocelli to the an- strips as separate regions of the head capsule, terior rim of the buccal cavity. The clypeus, which labelling them genae or parafrontalia, etc. Here should intrude between frons and mouth-parts they are simply termed the orbital strips (figs. is taken either to have disappeared, or to have 3,5,6). been incorporated without trace into the lower At its ventral end the tentorial sulcus does areas of the frons, or (as in most recent liter- not terminate precisely at the anterior tentorial ature) to have been mostly incorporated into pit (usually detectable externally as a widening the buccal cavity. Most authors have failed to and deepening of the sulcus) and may bifurcate, refer to the area from the antennae to the edge one arm (buccal arm) continuing to the edge of the buccal cavity as the frons, restricting this of the buccal cavity, the other arm (ocular arm) term to the area between antennae and ocellar swinging round towards the antero-ventral angle triangle (the term ocellar triangle refers, as used of the eye, where it slopes at the eye margin. here, to the often raised triangular area con- Either or both of these arms may be incomplete. taining and demarkated by the ocelli). The lower In Syrphus (figs. 1, 2) both arms are incomplete, frons is then referred to as either the face (as but in genera where the ocular arm is complete in McAlpine, 1981) or the fronto-clypeus. In externally and the buccal arm is not in evidence some instances the areea immediately above the (e.g. in Cheilosia, fig. 6) the orbital strips are edge of the buccal cavity is termed the post- delimited precisely at their lower ends. In Er- clypeus, on the assumption that only the more istalis (figs. 3, 5), where the ocular arm is in- basal of the two clypeal plates has been absorbed distinct but the buccal arm continuous strongly into the frons. Since there are median sclerites to the edge of the buccal cavity, the buccal arms in the front part of the buccal cavity (see section divide the face into a median and lateral regions. on mouthparts) serving the functions which Schiemenz (I.e.) regards these lateral regions as sclerites of the clypeal origin might be expected the genae and in those Syrphidae in which both to serve and since the more basal of these is lower arms of the anterior tentorial sulcus are hinged to the upper edge of the buccal cavity, complete this use of the term would follow pre- it seems unreasonable to regard them as secon- cisely the definition of the genae given by Snod- dary sclerites and thus require the true clypeus grass ( I960). Where one or other, or both, lower to have been incorporated without trace into the arms of the anterior tentorial sulcus are missing frons. Following this logic the entire area be- the frontal and genal regions of the head capsule tween anterior ocellus and the upper mouth edge cannot, by definition, be precisely delimited. is here regarded morphologically as the frons. The antennal insertions are found on the However, there is taxonomically a need to dis- frons. Immediately above them a pair of arcuate tinguish between the frontal area above the an- raised areas is usually differentiated (absent in tennae and the frontal area beneath them, so Microdon) in Syrphidae. These are jointly known following popular usage the latter area is referred as the frontal lunule. That part of the top of to here as the face (this does not entirely accord the head capsule between the para-sagittal sulci with the definition of the face used in McAlpine, (fig. 7) and containing the ocellar triangle with

1981, where the face is taken to terminate below its three ocelli is most conveniently referred to the level of anterior tentorial pits) and the as the vertex. Crampton (I.e.) has discussed the former as the "frons". difficulties inherent in using this term in Diptera.

A dorsal-ventral sulcus continuing upwards on Here the vertex is taken to merge imperceptibly Speight: Morphology of Syrphidae 145

with the occipital region of the head capsule {post cranium of McAlpine, I.e.) is concave, mak- at the angle where the posterior face of the head ing a sharp angle with the sides of the head. In dips down. Gouin (I.e.) and Schiemenz (I.e.) used the case of Eumerus and Merodon this angle is the terms vertex to include the median part of extremely sharp, producing a distinct postcranial the occipital region referred to here as the post- carina (fig. 8). This angle makes a very conve- vertex. Other authors have used the terms "ver- nient marker for defining the outer edge of the tex" and "vertical triangle" as synonyms of the three major post-cephalic regions, referred to term "ocellar triangle" as employed here. It here as the occipital region, the post-genal region should be noted that in the males of the syrphid and the hypostomal region. Tentorial sulci, ra- species with holoptic eyes the vertex is frequently diating out from the centre, delimit these regions represented only by the area of the ocellar tri- one from another. angle, such that the terms vertex and ocellar More or less in the centre of the back of the triangle become interchangeable, giving rise to head is found the occipital fora?nen, or foramen some confusion. In both sexes of most species magnum. This is largely surrounded by the post- the ocelli are disposed in a roughly equilateral occipital sclerite, which merges below with a triangle. But in some genera e.g. Eumerus, spe- mid-ventral plate delimited laterally by the well- cies occur in which the anterior ocellus is placed marked hypostomal sulci. This mid-ventral plate further forward, some distance from the two lat- makes up the hypostomal region of the head and eral ocelli. is supposedly derived largely from lobes of the Dorsally, behind the eyes and lateral to the postgenae that have fused in the mid-line (Snod- vertex, the head capsule often projects to some grass. I.e.) giving it one of its names, the hypos- extent, before curving ventrally into the occipital tomal bridge. In Syrphus and its allies (fig. 7) a region. Whether or not there is a para-sagittal dorso-ventral, median, hypostomal suture prob- sulcus in evidence on each side of the ocellar ably indicates the fusion line. The posterior ten- triangle, thus precisely segregating the vertex torial pits occur in the hypostomal sulci and the from these post-ocular strips, they may conve- two lateral plates demarkated ventrally by the niently be termed the post-ocular orbits, -às inCoe hypostomal sulci make up the post-genal region (1953). The post-ocular orbits continue without (referred to simply as a part of the genae by interruption down the side of the head to merge Gouin and Schemenz) of the head. A more dorsal with the genal region. pair of rather incomplete sulci, referred to as the Between the antennae and the upper mouth transverse sulci, help to delimit the upper boun- edge a median protruberance is present in many dary of the postgenal region. The occipital region Syrphid genera. Although this bump has no dis- is then that part dorsal to the transverse sulci. Yet crete identity morphologically, it is important another pair of sulci, the parasagittal sulci, divide taxonomically and is usually referred to either as the occipital region into three fields. The two

û\e facial tubercle or facial prominence (figs. 1, lateral fields of the occipital region have usually 6). The area surrounding the antennal insertions been called the tempora but the dorsal, median, may also be drawn out to form a tubercle, which field has been subject to various appellations. in European Syrphids reaches its extreme of de- Here it is termed the post-vertex. In Merodon velopment in Ceriana (fig. 4). This has been and related genera the post-vertex is almost ob- referred to as the frontal prominence or frontal literated (fig. 8), the para-sagittal sulci swinging tubercle. into the mid-line at the post-cranial carina.

The surface of the frons is not infrequently Very prominent towards the centre of the thrown into a series of furrows or ridges (re- postgenal fields of Eristaline syrphids are large, ferred to as regulae) well illustrated by species of raised, dorso-ventral bands of sensillae — these Chryogaster and Orthonevra. Other genera ex- bands of sensillae are often very restricted in hibit highly polished concave or flattened median their extent in Syrphinae. In all Syrphids the fields contrasting markedly with the surrounding occipital foramen is so constructed just above surface, e.g. Neoascia and Neocnemodon, or mid-way as to be almost converted into two se- areas of procumbent, iridescent microhairs (dust- parate holes, due to the development of promi- ing) alternating with bare, shining patches, as in nent transverse cervical condyles for articulation many Syrphinae. with the cervical sclerites (figs. 7, 8).

Head capsule; posterior surface Antennae

In Syrphidae, the posterior surface of the head Syrphid antennae comprise three principal 146 Tijdschrift voor Entomologie, deel 130, 1987

F'g- 7, Syrphus ribesii, male, head, posterior view. Fig. 8, Merodon equestris, male, head, posterior view.

Fig. 9, Platynochaetus setosus, third antennal segment and arista, lateral view, outer side. Fig. 10, Syrphus ribesii, antenna, lateral view, outer side. Fig. 11, Callicera aenea, end of third antennal segment and arista, lateral view, inner side. Fig. 12, Syrphus ribesii, antenna, lateral view, inner side. Fig. 13, Microdon mutabilis, antenna, lateral view, outer side. Speight: Morphology of Syrphidae 147

segments and a more or less annulate arista which make up the mouth-parts proper (alterna- borne on the third segment. The third segment tive theories concerning the fate of the clypeus represents the first flagellar segment. The arista are mentioned under the section dealing with the represents the rest of the flagellar segments, one frons). or more of which may be distinct (figs. 9 — 13). Syrphid mouthparts comprise sclerites derived These aristal segments have been termed aristo- from the clypeus, located dorsally towards the meres (McAlpine, I.e.). Inserted between the base of the mouthparts complex; a modified la- eyes on the frons, the antennae occur at a point brum lying dorsally and distal to the clypeal scler- where there is a distinct change in the angle of ites; a lateral pair of maxillary stylets plus palps; slope of the frons. Not infrequently the frons a ventral hypopharynx and a partly membrane- projects at this point, producing in extreme cases ous and partly sclerotised labial complex lying a frontal tubercle (fig. 4). beneath the hypopharynx. The labial complex The antennal segments vary in their propor- provides a sheathing trough (the "labial gutter") tions between genera, but segment 3 is normally for the other mouthparts that lie distal to the the largest. Sensory pits are frequently discerni- clypeal sclerites. The entire apparatus, normally ble on either the inner or the outer face of seg- called the proboscis, is hinged to the head capsule ment 3. Cheilosia species often possess large at the upper mouth edge, via the basal clypeal numbers of small pits, while in many other sclerite, and hinged again at the junction between genera there is at least one large pit (figs. 9, H, the distal clypeal sclerite and the labrum. Because 13). The dorsal margins of segments 1 and 2 of this hinging arrangement, when the proboscis carry bristles in various genera. is retracted into the head it folds away such that The arista may be terminal or sub-dorsal, hair- the clypeal sclerites form a floor to the head less, pilose or plumose. In most genera it is bris- capsule in the anterior part of the buccal cavity tle-like, but it may be bulbous or strap-like. In and the labrum rests up against them with its tip

Platynochaetus it is spatulate (fig. 9). Crampton pointing forwards and surrounded by the fleshy (1942) discusses the merits of applying the term lobes at the end of the labial complex. The main "ceratostylate" to certain forms of the antennae labial sclerite, the premental sclerite, is then the with a terminal "arista" and the condition found plate seen bulging from the buccal cavity when in Ceriana would certainly be better termed ce- the head is viewed from beneath. rato-stylate rather than aristate were the term There is no obvious sexual dimorphism in the ceratostyle to come into common use. structure of syrphid mouthparts. There is no ex- To accord with general usage, the terms scape ternal indication of mandibular sclerites in any of and pedicel should be applied to the first and the genera examined during preparation of the second antennal segments, respectively. present text. Apart from in Microdon, two clypeal sclerites Mouthparts are present in Syrphidae: A postclypeus, articu- The modifications exhibited by the mouth- lating on its proximal margin with the anterior parts of Syrphidae, occasioned by these flies' spe- edge of the buccal cavity, and a more distal antec- cialised nectar and pollen-feeding habits, have lypeus. The proximal edge of the anteclypeus given rise to various published accounts of their articulates with the anterior edge of the postcly- structure. The most detailed study is that of peus. The distal margin of the anteclypeus is Schiemenz (1957), who deals with both external deeply concave, its lateral arms passing forwards and internal anatomy of the mouthparts of Eris- to articulate with the base of the labrum but talis arbustoruyn. Crampton (I.e.), Gilbert visible externally only as narrow sclerotised (1981), Gouin (I.e.), Holloway (1976) and Nayar strips (fig. 14). A solitary clypeal sclerite is pres- (1964) provide additional information. Gilbert's ent in Microdon, bearing a close resemblance to

(I.e.) account considers the functional implica- the anteclypeus of other Syrphidae (fig. 15). tions of variations in structure exhibited by a The labrum is a heavily sclerotised, highly pol- range of European species and also incorporates ished plate, in external appearance reminiscent a comprehensive bibliography. of an upturned canoe. In genera such as Rhingia, Although morphologically part of the head- with mouthparts adapted to nectar extraction capsule the clypeal sclerites have apparently from flowers with a deep corolla, the labrum is come to lie within the buccal cavity in Syrphidae, greatly elongate (fig. 16). so they are considered in this section of the pres- Roofed by the labrum, the food-channel is ent text together with the head appendages floored by the hypopharynx, a sclerotised, stylet- 148 Tijdschrift voor Entomologie, deel 130, 1987

like, concave sclerite, which projects forwards as like strips with strongly sclerotised tips such as far as the tip of the labrum. The pair of maxillary are found in Rhingia (fig. 16). sclerites, with their unsegmented^a/^x, lie in the In the literature reference is frequently made membrane lateral to the sclerites of the food to regions of the proboscis called the rostrum, and channel. Each maxillary sclerite is continuous haustellum. The rostrum is the largely mem- internally with a sclerotised rod interpreted as braneous basal section of the proboscis, as far as the stipes, which passes back into the head cap- the base of the labrum and labium. The haustel- sule. In most genera the maxillary sclerites sweep lum is then the most distal portion containing down from their lateral position towards the the mouthparts proper and including the label- mid-line, sheathing the hypopharynx ventro-lat- lum. erally. The maxillary palps, conversely, sweep upwards, hugging the outer surface of the labrum The cervical region so that distally they come to lie side by side along In Diptera, the cervical region or cervix, is its dorsal surface. In some general, e.g. Merodon, largely impacted onto the front face of the the maxillary palps are quite strongly sclerotised, thorax. Its structures comprise three pairs of but in the most cases they are membraneous. cervical sclerites (or cervicalia) and the cervical The identity of the maxillary sclerites, here organ complex (figs. 17 — 19, 21). The main fea- termed the maxillae, remains an unresolved ture of the cervical region is the pair of lateral problem: SeeMatsuda (1965). Currently, they are cervical sclerites. These flank the cervical cavity, most often regarded as the lacinia (e.g. in McAl- along their outer edges articulating internally pine, 1981). Microdon is exceptional among Syr- with prothoracic elements. Ventrally, the lateral phidae in possessing short, flange-like maxillary cervical sclerites meet in the mid-line over the

sclerites, orientated transversely rather than lon- . membraneous pocket containing the cervical or- gitudinally. Also, the maxillary palps are rudi- gan. Dorsally, arms from the lateral cervical mentary in Microdon (fig. 15). sclerites twist forwards, flanking the lower edge The most complex and varied structure of the of the cervical canal and articulating at their tips syrphid mouthparts is the labium and its appen- with the cervical condyles on the back of the head. dages. Matsuda (1965) summarises the informa- Resting alongside the tips of the lateral cervical tion available on the origin of dipteran labial sclerites is the pair of small, dorsal or anterior structures. Essentially, of the two vertical labial cervical sclerites. sclerites the more basal, the postmentum, is ab- In the cervical membrane along the lower edge sent. The other, the prementum, has become of the main body of the lateral cervical sclerites partly desclerotised but gave rise to the large is found the pair oi posterior cervical sclerites. In external, ventral plate of the Syrphid labial com- Syrphidae these are rather variable in appearance plex (figs. 14— 16) termed here the premental and in degree of sclerotisation. At their outer sclerite. The membraneous lobes lying distal to ends the posterior cervical sclerites appear to the premental sclerite, collectively known as the fuse with the prothoracic epistema. Their inner labellum, are apparently derived from the orig- ends bear sclerotised outgrowths, as in Syrphus inal labial palps. The terminal lobes of the orig- (fig. 21). inal prementum (glossae and paraglosae, to- According to Matsuda (1970), in Diptera the gether known as the ligula) have been lost. The lateral cervical sclerites are derived from pleural partly membraneous prementum containing the elements of the prothorax but the other cervical premental sclerite has become weakly resclero- sclerites are secondary sclerotisations. The lateral tised on its upper surface, producing a secondary cervical sclerites would most logically articulate sclerite (see fig. 16) known as the hypoglossa laterally with propleural elements, and since the (usually hidden from sight when the mouthparts antepronotum and proepisternum are indistin- are examined in side view). guishably fused somewhere in the region of the Within the labellum, the original two-seg- point of articulation of the lateral cervical scler- mented form of the labial palps has been mod- ites, the position of that articulation has been ified but is evidenced from the position of largely used by some authors as an indication of the internal strengthing rods, the furca and epifuraca dorsal extent of the proepisterna. However, Mat- (see Schiemenz, I.e.). In different Syrphids the suda (I.e.) has reduced the value of this argument labellar lobes exhibit a range of variation from by suggesting that in Tabanus the lateral cervical the voluminous, convoluted, membraneous flaps sclerites probably articulate with elements of the exhibited by Syrphus (fig. 14) to narrow, tongue- antepronotum. Speight: Morphology of Syrphidae 149

Fig. 14, Syrphus ribesii, mouthparts, lateral view. Fig. 15, Microdon mutabilis, mouthparts, lateral view. 150 Tijdschrift voor Entomologie, deel 130, 1987

Fig. 16, Rhingia campestris, mouthparts, lateral view.

The cervical organ complex has received scant results in deciding correct designations for tho- attention in Diptera, being entirely ignored in racic sclerites, since the number of Dipteran spe- reviews of Dipteran morphology such as those cies whose thoracic musculature has been com- conducted by Matsuda (1970), Bitsch & Matsuda prehensively investigated can almost be counted

(1973) and McAlpine( 1981). The only published ' on the fingers of one hand. The only syrphid review of its structure in Diptera is that incorpo- whose thoracic musculature has been reported on rated into Speight (1969). Among Syrphidae, the is Eristalinus megacephalus Rossi (as cervical organ is rather varied in its detailed Lathyrophthalmus obscuritarsis), in Maki structure, but is essentially a sensilla-bearing (1948). The degree of confusion existing at pres- plate, termed the sella, lying across the mid-line ent can be adduced from a comparison between in a membraneous pocket beneath the postero- the terminologies employed for the Dipteran median wings of the lateral cervical sclerites. The thoracic sclerites by Crampton (1942) and Mat- sensilla are directed upwards and slightly out- suda (1970): hardly a single part of the thorax is wards so that their tips touch the underside of the given the same name in the two texts. In the postero-median wings of the lateral cervical present account an attempt has been made to sclerites. The sella extends posteriorly in the follow the terminology employed by Matsuda mid-line as a sclerotised bar in the surface of the (I.e.), with certain modifications based on McAl- cervical membrane, where it articulates with, or pine (1981). Unfortunately, Matsuda's text is is fused to, a median sclerite which some authors based largely on a comparison between certain have identified as the presterum of the pro- Tipulids, Tabanus and a Nycteribiid, augmented thorax. Matsuda (I.e.), however, argues that this by occasional observations on Drosophila, and latter plate is more likely a secondary sclerotisa- syrphid thoracic morphology presents features tion of the cervical region and should be regarded departing significantly from what has been ob- as a cervical sclerite. Here, because of its evident served in these other flies. involvement in the functions of the cervical organ and its debateable origin, this plate is simply called the postsella. In many genera, (e.g. Prothorax Syrphus), at some point along its length the sella The most anterior of the three thoracic seg- expands laterally into a trapezoid, bearing at its ments, the prothorax, is in Diptera an insignif- lateral extremities a pair of sclerotised tubercles icant and incompletely delimited component of (fig. 21). the front end of the thorax, overhung by part of the mesonotum. The prothoracic tergum, the pronotum, occurs externally as three sclerites, The thorax the antepronotum and the paired postpronotal

sclerites. In Syrphidae, the antepronotum is dor- Even a small amount of original research on sally reduced to a narrow semi-circular plate, muscle origins would proably produce dramatic deeply notched in the mid-line, arching round the Speight: Morphology of Syrphidae 151 17

Fig. 17, Eristalis tenax, prothoracic region, anteroventral view.

upper edge of the cervical membrane. Laterally it the absence of information to the contrary, the widens out, passing ventrally to fuse indistingui- suture curving up, over the humeral calli poste- shably with the prothoracic epistema. Poste- riorly, from the antero-dorsal edge of the protho-

riorly, in the mid-line, the antepronotum dips racic spiracle, is here taken to delimit the junction down into the deep, pit-like antecostal suture between postpronotum and mesonotal elements. (fig. 20), from which the first thoracic phragma This suture curves round towards the antecostal

is invaginated. More laterally, the posterior edge suture but then usually fades out, so that postpon- of the antepronotum abuts onto the postprono- otum and mesonotum are incompletely demar- tum, a junction marked by a complete suture in kated from each other. most Syrphidae, e.g. Syrphus (fig. 18). The propleura comprise a proepisterum and a

The postpronotum is represented externally proepimeron. The proepisternum (propleuron by the pair of sclerites forming at least the major of taxonomists) is fused anterodorsally with the part of the prominent humeral calli of taxono- antepronotum and anteroventrally with the pos- mists. Unless the antecostal suture occurs be- terior cervical sclerites. Some authors (e.g. tween antepronotum and postpronotum in Syr- Thompson, 1972) have recognised anepisternum phidae, rather than between post-pronotum and and katepisternum in the proepisternum, but on

meso-notal acrotergite as is generally supposed, what basis is unclear. The position of the anterior the lateral elements of the postpronota are not edge of the proepimeron can be detected by the externally connected with each other in these line of the propleural suture, which divides flies. Instead, the postpronotal sclerites appear to proepisternum from proepimeron and runs from dip into the antecostal suture at its outer ends. In the rim of the fore coxal cavity to the prothoracic 152 Tijdschrift voor Entomologie, deel 130, 1987

Fig. 18, Syrphus rihesü, prothoracic region, anteroventral view. Fig. 19, Microdon mutabilis, prothoracic region, anteroventral view.

Spiracle. In Microdon (fig. 19) the posterior edge Syrphidae the probasisternum is clearly differen- of the proepimeron appears to be marked by a tiated along all its margins. It does not seem to suture for part of its length, but it is otherwise join with propleural elements to form a precoxal not differentitated from mesopleural elements. bridge in any members of this fly family. On its Anteroventrally, a projection of the proepis- anterior margin the probasisternum is in the ternum articulates with or fuses to the posterior mid-line contiguous with the sella or postsella of cervical sclerites. This projection is to a variable the cervical organ. extent marked off from the main body of the The profurcasternum is in syrphids a median, proepisternum by a suture and may or may not ventral, triangular sclerite (figs. 17 — 19), joined represent the remains of a propleural precoxale anteriorly to the probasisternum, from which it

— it is given no separate designation in this is delimited by a complex suture. It meets the account. Another sclerite which could be inter- mesothoracic presternum posteriorly, anterior to preted as either the propleural precoxale or a the point at which the latter plate disappears into trochantin lies free in the cervical membrane the mid-ventral thoracic suture. In its anterior along the anterior rim of the coxal cavity, be- angles the profurcasternum bears the furcal pits tween proepisternum and probasisternum. This from which the furcal arms are invaginated. sclerite is often poorly sclerotised and may be absent. Here it is regarded as of secondary origin and labelled simply as a secondary sclerite (figs. Mesothorax: mesonotum 17—19).

The sternum of the prothorax is represented In Diptera the mesothorax is greatly expanded externally by a basisternum and a furcasternum. and makes up the main bulk of the thoracic sur- The "presternum" of some authors is discussed face. This expansion is presumed to be due to the in that section of the present account dealing need to accommodate the flight musculature as- with the cervical region. The basisternum (pres- sociated with the fore wings, which are attached ternum of taxonomists) is a large median plate on this thoracic segment between pleural and interposed between the cavities of the fore coxae, tergal elements. Coincident with the expansion carrying a pronounced median sulcus, from of the musculature has been differentiation of a which is invaginated the basisternal carina. In complex endoskeleton, the 7neso sternal apophy- Speight: Morphology of Syrphidae 153

Fig. 20, M. mutabilis, prothoracic region, anterodorsal view. Figs. 21, 22, Syrphus besii, cervical region, anteroventral view (21) and thorax, dorsal view (22).

sis, for muscle attachment. In most Diptera — to be unrecognisable without first partially including Syrphidae — almost the entire meso- desclerotising the specimen. Even then they can- thoracic sternal region is invaginated into the not be seen in side view, being concealed beneath thorax as the mesosternal apophysis. the lateral edge of the mesoscutum. Viewed from above, the Syrphid thorax is The mesoscutum, generally known among tax- nearly all of mesonotal origin (fig. 22). The me- onomists as the mesonotum, is the main meson- sothoracic notum, or mesonotum, is made up of otal sclerite. Although its anterior and posterior five sclerites. The most anterior of these is the sutures are incomplete, it is well differentiated acrotergite, possibly visible on the antero-dorsal laterally and includes the morphological features face of the thorax in Microdon (fig. 20), but known taxonomically as the "notopleural area" otherwise seemingly not discernible in Syrphi- (or "presuturai area") and the "post-alar calli". dae. The acrotergite is followed by the The confusing term notopleural area (and asso- prescutum, which is reduced to a pair of lateral, ciated terms notopleural depression and noto- unconnected strips, running along the edge of the pleural callus) is not used here, since the part of mesonotum from just above the prothoracic spir- the thorax to which it refers is in origin meson- acle to as far as the transverse sulcus of the otal, not mesopleural. The alternative term of mesoscutum. These presentai strips are usually presuturai area (and associated terms presuturai just visible when the thorax is examined from the depression and presuturai calli) is used in its side (fig. 23), but are so reduced in Microdon as place. The mesoscutum carries a shallow trans- 154 Tijdschrift voor Entomologie, deel 130, 1987

Fig. 23, ^. ribesii, thorax, lateral view, left side.

verse gulley, just anterior to the wing base. A within the mass of the original mesoscutellum: similar feature occurs in many different Diptera (see Matsuda, (I.e.), so that some indeterminate and has been given various names: transverse portion of the hind part of the "mesonotum" of suture, lateral parapsidal suture, scuta! suture, taxonomists is scutellar in origin. In syrphids the transcutal suture. Since, according to Matsuda hind part of the mesoscutum differentiated by

(I.e.), this feature is intra-scutal and thus does the transscutellar sulcus protrudes from the pos- not mark a joint between sclerites, it should terior end of the dorsal thoracic surface as a well- ideally be termed a sulcus, following Snodgrass defined semi-circular lobe, generally called the (I960). Diptera may carry up to three distinct scutellum by taxonomists. Here, this feature is transverse sulci on the mesoscutum, but Syrphi- called the scutellar lobe. In some genera its outer dae luckily only exhibit one, so whatever its ho- (postero-dorsal) edge is tuberculate or crenulate. mologies (see McAlpine, 1981), it can conve- Ventral to the scutellar lobe is the median plate niently be called "the transverse sulcus" which is of the most posterior of the mesonotal elements, the term used for it here. At the outer ends of the the mesopostnotum (also known as the mediot- transverse sulcus of the mesoscutum. Microdon ergite, or subscutellum or postscutellum), possesses a pair of shelf-like, semi-circular, flanked by its two lateral sclerites (figs. 23, 24). sclerotised outgrowths of the mesoscutum, which These are the lateral post-nota of the mesono- do not seem to have an equivalent in other Syr- tum, also known as the laterotergites (or pleur- phids. The prescutal strips are beneath these otergites). Some authorities, e.g. Colless &: McAl- mesoscutal flanges. pine (1970), currently regard these postnotal Posteriorly, the suture marking the mesoscutal sclerites as derived from the acrotergite origi- junction with the succeeding mesonotal sclerite, nally interposed between mesonotal and metan- the scutellum, has been lost. The very deep and otal elements, rather than from the mesonotum. well-marked trans-scutellar sulcus is located The median postnotum is normally a convex Speight: Morphology of Syrphidae 155

Fig. 24, Microdon mutabilis, thorax, lateral view, left side.

plate in Syrphtis and its relatives (fig. 23), push- detached sclerites located in the axillary complex ing out into a distinct lobe. But in Microdon (fig. are believed to be derived from the mesonotum.

24) it is unusually flat. The lateral post-nota form These include the tegula, humeral plate, 4th ax- shallow calli just dorsal to the metathoracic spir- illary sclerite, and subalare. They are dealt with acles, but continue towards the mid-line beyond in that portion of the present text concerned with these calli to a point where an unobtrusive suture the wings. marks their junction with the median postnotum. Antero-dorsally, the lateral postnota border the posterior part of the wing-base complex of Mesothorax: mesopleura axillary sclerites. Postero-ventrally, the lateral The pleural sclerites of the second thoracic post-nota border the similar, but physically much segment exhibits in Diptera a bewildering array smaller, complex of sclerites round the base of of bumps, hollows and grooves, some of which the haltere. The massive second thoracic represent primary subdivisions of the pleura and phragma attaches directly to the hind margin of most of which have been employed as taxonomie the median postnotum and passes down inter- characters at some time or another. In Syrphidae, nally to form the sclerotised hind wall of the most of the side of the thorax is mesopleural in functional thorax. The second phragma stops origin. The largest recognisable entity is the just short of the floor of the thorax, immediately mesepisternum. The anterior margin of this above the membraneous strip between hind sclerite is marked by the posterior edge of the coxae and first abdominal sternite that acts as a prothoracic spiracle, from which an incomplete thoracic/abdominal flexion joint. Certain small, suture extends towards the fore coxae, the me- 156 Tijdschrift voor Entomologie, deel 130, 1987

sepisternum being fused with propieural ele- the meropleurite is probably located. The flat ments ventrally. In the mid-line ventrally, the part is regarded by McAlpine (1981) as the me- mesepisterna meet along the line of the mid- sepimeral anepimeron and the callus as the me- ventral suture, which terminates anteriorly in the sepimeral katepimeron. The callus is of taxo- mesofurcal pit. Posteriorly, the upper part of the nomie significance and by taxonomists has been edge of the mesepisternum is marked by the called the barrette, a term used for it here. The vertical membraneous strip of the pleural suture. meropleurite corresponds roughly with the "hy- The pleural suture continuous downwards as a popleuron" of taxonomists, though the term hy- well-marked groove, taking a zig-zag course to- popleuron has been applied in such a way that it ward the mid-coxae. The lowest (third) section of also includes metapleural elements. the suture is unusually complete in Syrphidae, being especially well-marked in Microdon (fig. Mesothorax: mesosternum 24). Dorsally, the margin of the mesepisternum Mesosternal elements are surprisingly well re- is indicated by the suture delimiting mesonotal presented externally in Syrphidae — in Schizo- elements and by the membraneous area round phora the mesosternum is, almost in its entirety, the wing-base containing the axillary sclerites invaginated into the thorax. In the three syrphids (some of which are mesepisternal in origin). illustrated here (figs. 26—28) the mesosternal Within the mesepisternum, various subdivi- presternum is clearly differentiated from the pre- sions may be recognised. From the ventral end of ceeding prothoracic furcasternum and invagi- the upper (first) section of the pleural suture the nated posteriorly into the mesofurcal pit at the incomplete anapleural suture proceeds forwards, beginning of the mid-ventral thoracic suture. dividing the mesepisternum into a dorsal Laterally, the meso-presternum meets mesepis- anepisternum (the "mesopleuron" of taxono- ternal elements, from which it is less clearly se- mists) and a ventral katepisternum (see McAl- parated. The two wings of the anterior pine, 1981, for reasons why the ventral area of mesosternal furcasternum each occur externally the mesepisternum delimited by the anapleural as a narrow strip along the anterior rim of the suture should be regarded as ^^/-episternum mesocoxal cavities, dipping down into the mid- rather than pre-episternum). The katepisternum line to reappear posteriorly as the ventral coxal is the "sternopleuron" of taxonomists. The anep- condyles, articulating with the median mesocox- isternum may be further differentiated into an ite (figs. 26— 28). Hhe posterior mesosternalfur- antero-dorsal flat area (the "anterior depressed casterna make a hind rim to the mid coxae. These portion of the mesopleura" of Coe, 1953, and the external furcasternal elements are continuous in- "anterior flat portion of mesopleuron" of Vock- ternally with the massive mesofurca, without in- eroth, 1969), and a shallow more posterior callus dications of sutures interposed, thus reducing the (fig. 23). These fields of the anepisternum are probability that they might be better interpreted frequently mentioned independently in taxo- as some vestige of a mesopleural trochantin. nomie texts but do not seem to have been named, other than by Speight (1980), where they were Metathorax referred to as mesopleurite 1 and mesopleurite 2. The metathorax is proably the least studied They would be better called anepisternite 1 and region of the external anatomy of Diptera and anepisternite 2, and these terms have been used apart from the remarkable paper of Young to denote them here. (1921) has been largely ignored in the literature. The mesothoracic epimeron is clearly delim- Whatever elements may have once been pres- ited anteriorly by the pleural suture. Posteriorly ent in the metanotum, it is represented in Syrphi- its edge is marked by the suture between it and dae solely by a narrow hoop-like sclerite evanes- the lateral postnotum of the mesonotum. Ven- cent medially in some genera, hugging the trally, it is fused with a plate regarded as the antero-dorsal face of the abdomen but invagi- mesothoracic meron, to produce a composite nated along its anterior edge into the suture sclerite known as the meropleurite. In Syrphidae marking externally the position of the second the mesepimeron and the meropleurite are thoracic phragma (figs. 41 —43). Due to its loca- hardly differentiated from each other by sutures. tion it is often only visible at its dorso-lateral The upper part of the mesepimeron corresponds corners, when the abdomen is viewed from

more or less with the "pteropleuron" of taxon- above. In side view it can be quite concealed on omists. It is flat above but exhibits a transverse the front of the abdomen, by a forward bulge of callus below, just above where its junction with abdominal tergite 1, as in Microdon (fig. 24). In Speight: Morphology of Syrphidae 157

mtb

SB SB

Fig. 25, Eristalis tenax, thorax, region surrounding the haltere, lateral view, left side. Fig. 26, Syrphus ribesii, meso and metathoracic sterna, ventral view. Fig. 27, Eristalis tenax, meso and meta thoracic sterna, ventral view. Fig. 28, Microdon mutabilis, meso and methatoracic sterna, ventral view.

Syrphus (fig. 41) the metanotum bears a distinct tum of the mesonotum behind the metathoracic transverse groove which could represent the su- spiracle (see Eristalis, fig. 25), but in others ture between two of the original metanotal scler- membrane of the haltere axillary complex inter- ites. Equally, this groove might be a secondary venes (see Microdon, fig. 24). Ventraliy, the me- feature. No attempt has been made to identify tepisternum may, in combination with the mete- subdivisions of the metanotum. Laterally, the pimeron, form a postmetacocal bridge (fig. 28). metanotum fuses with metapleural elements, the The metepimeron, as interpreted here, exhib- junction being in some instances, e.g. Syrphus, its some unlikely characteristics. That it is incom- distinctly marked. pletely delimited from the metepisternum has The metapleura comprise an episternum, an already been mentioned. Postero-ventrally, the epimeron and a precoxale. The met-episternum metepisternum is bordered by membrane inter-

is imperfectly demarkated from both mesotho- posed between it and abdominal sternite 1. In racic meropleurite anteriorly and metathoracic forms lacking a postcoxal bridge this membrane epimeron posteriorly and can only be detected is continuous with that of the meta-coxal cavities. clearly immediately posterior and ventral to the In some syrphids, the metepimeron bulges pos- metathoracic spiracle. In some genera the metep- tero-dorsally into a distinct callus on the front isternum appears to contact the lateral postno- margin of the first abdominal tergite (fig. 25), 158 Tijdschrift voor Entomologie, deel 130, 1987

from which it is separated by reflexed membrane. copic organs known as the haltères. As described

This metepimeral callus is frequently visible in preceding pages, the wings and the haltères from above as an angular projection on the outer, are located on the side of the thorax between anterior corner of tergite 1, and is especially vis- tergal and pleural elements. Externally, the junc- ible in Neoascia (fig. 44) where it is developed tion between wing and thorax is marked by a into a massive spine — illustrated in Stackelberg cluster of axillary sclerites and associated mem- (1965) as apparently part of abdominal sternite brane, which may be referred to as the axillary 1. The metepimeron frequently encloses the first complex. The haltere base has a corresponding abdominal spiracle at its lower edge (figs. axillary complex.

23 — 25). In Microdon and Ceriana this callus is The wing itself is a largely transparent sheet lacking, but the first abdominal spiracle is still of membrane, traversed by a series of sclerotised enclosed by the metepimeron in European spe- bars, the wing-veins, radiating from the wing- cies of these genera. In Eristalis, the first abdom- base. In syrphids there are six major wing veins. inal spiracle occurs partly in the metepimeron Those radiating into the middle area of the wing and partly in the adjacent abdominal membrane. branch one or more times before reaching the Zumpt & Heinz (1949) regarded the metepi- wing margin. Occasional cross-members are also meral callus of E. tenax as either a secondary present, termed cross-veins. The wing-mem- sclerite or a part of abdominal tergite 1. brane is in some genera banded or blotched with

The metacoxal precoxale is a narrow sclero- brownish pigment and may be all or in part tised strip clearly differentitated round the lateral covered by microtrichia. The most posterior rim of the metacoxal cavity. At its posterior end fields of the wing-membrane are largely separ- it reaches the lateral condyle of the metacoxa. ated from the main body of the wing, going to

Anteriorly, it joins with the metabasisternum in form the alula and, folded beneath the wing-base, most hoverflies, to form a composite sclerite the calypters (fig. 31). making a precoxal bridge round the metacoxae. Attempts have been made by morphologists It retains its identity independent of the metab- and taxonomists alike to homologise the main asisternum in Melanost07na and Sericomyia. fields of the wing-membrane and the main wing- The principal metasternal sclerite is the me- veins, throughout the various Orders of insects. tabasisternum, a mid-ventral plate located be- The wing-vein notations resulting have been lar- tween and anterior to the hind coxae. It is deeply gely incompatible, but nonetheless many authors grooved in the mid-line and frequently fused have adopted hybrid wing-vein terminologies with the metaprecoxale laterally, as described based partly on one theory and partly on another. above. The premetacoxal bridge thus developed Wootton (1979) provides an able review of the is frequently called the "metasternum". Its par- present chaos, which affects syrphid wing-vena- ticular make-up requires that it be given some tion notation along with that of other Diptera. morphological appelation such as metabasis- Goffe ( 1947) attempted to derive a stable system terno-precoxite — not a term likely to be met for naming syrphid wing-veins, but unfortu- with general approval! For want of a more ap- nately the notation he proposed is a confusing propriate alternative this composite structure is hybrid. Alternative notations, differing from one referred to here as the premetacoxite. With the another to a greater or lesser extent, can be found postmetacoxal bridge, the premetacoxal bridge in Coe (1953), Colles & McAlpine (1970), McAl- forms in some syrphids, e.g. Sphegina, a pro- pine (1981), Matsuda (1970), and Seguy (1959, nounced conical bump protruding postero-ven- 1961), etc. In the present account, Colless & trally from the underside of the thorax. McAlpine (1970) are followed, with modifica- The other external sclerite of the metasternum tions suggested by Wootton (1979). is the vestigial furcasternum, projecting from The nomenclature of the three most anterior within the thorax to form the ventral meta-coxal longitudinal wing-veins, named here the Costa, condyles, to either side of the mid-line just pos- Subcosta and Radius, is reasonably stable in Dip- terior to the tip of the metabasisternum. tera. However, the next vein, the Radial Sector, has anomalously usually been labelled as though its branches were branches of the Radial. That Wings convention is not adopted here, following Woot- In Diptera only the fore wings (hereafter re- ton's recommendation. The following three ferred to as "the wings") remain as organs of veins, here named the Median, the Anterior Cu- flight, the hind wings being modified into gyros- bitus and the Posterior Cubitus have been much Speight: Morphology of Syrphidae 159

confused, the appropriate designations for distal area of the wing posterior to the 1st Anal vein branches which could be derived from either the and including the alula is homologous with the Median or the Anterior Cubitus (or represent claval field recognised in other insects. The fugai cross-veins — see below) remaining today al- field is then represented by the calypters. The most a matter of personal opinion. The vein cleft between the alula and the main body of the

regarded here at the Posterior Cubitus has been wing is generally termed the axillary incision.

omitted from illustrations of syrphid wing-vena- The alula is a simple membraneous flap, its only tion by some authors, e.g. Oldroyd (1970). The venation a vein-stub at its postero-basal corner. First Anal vein as recognised here is labelled as The calypters hinge onto the base of the wing via such in most recent literature, but Wootton the 3rd and 4th axillary sclerites (fig. 31). The (1979) suggests the cubito-anal area of the wing upper calypter is folded over the lower when the may well have been misinterpreted by dipterists, wing is at rest, but when the wing is stretched and the true anal veins may well be confined to out, as in the illustration (fig. 31), their relation the alula in Diptera. Elucidation of that problem to the rest of the wing membrane can be better

awaits further work, as Wootton (I.e.) himself appreciated. The rim of each calypter is some- says, so the more traditional approach to the what sclerotised and bears a thick fringe of long identity of the First Anal has been adopted in the hairs. present text. The axillary complex of the Syrphid wing-base The more basal cross-veins of the syrphid has not been thoroughly investigated in the com- wing have generally received the same designa- pilation of this account. Suffice it to say that there tions, but there are differences of opinion as to are four principal axillary sclerites in Diptera. which of the more distal cross-veins are actually The 1st and 2nd have been tentatively identified transverseley deflected branches of longitudinal in fig. 31, following Matsuda (1970). The 3rd and veins. Thus the vein labelled here as an anterior 4th are easier to distinguish. Axillary sclerites branch (M.l) of the Median, has been called the 1 — 3 represent detached basal parts of the wing- "upper marginal cross-vein" and the vein la- veins (Matsuda, I.e.), while the 4th is supposedly belled marginal cross-vein has often been re- mesonotal in origin. On the front margin of the garded as a branch of the Anterior Cubitus. wing, two basal sclerites may be distinguished, The areas of membrane entirely enclosed by the humeral plate, and the tegula. These are ap- wing-veins comprise the "cells" of the wing. One parently homologous with the sclerites of the approach to naming wing-cells is to number same name found in other Orders and are seem- them from the anterior margin of the wing back- ingly of mesonotal origin. A detailed account of wards. An alternative approach is to designate the Dipteran wing-base complex is given by them according to their position on the general McAlpine (1981). wing-surface, a system which gives rise to costal A final important feature of the axillary com- cells, basal cells, marginal cells, etc. This system, plex is the subalare. Although located beneath as laid out in Oldroyd (1970) is employed in the the wing-base, according to Matsuda (I.e.) this

present account (fig. 29). Neither system is very sclerite is of mesonotal origin. In Syrphidae it satisfactory because of the problems of homology carries the plumule, a feature unique to the fam-

of wing-cells in fly families — like the Syrphidae ily. The plumule is a membraneous, finger-like — in which few wing-cells are present. appendage, thickly covered in long, fine, wavy

A venational characteristic of Syrphidae, men- hairs. It projects backwards from the posterior tioned almost universally in the literature, is the tip of the subalare and thus lies closely opposed vena spuria. This secondary strengthening of the to the underside of the lower calypter. It is an v^ing membrane lies between the Radial Sector easily seen feature in all syrphids except Neoascia

and Median veins and is more or less parallel and Microdontinae, where it is rudimentary and

with the latter. Almost invariably it crosses the represented only as a rather hairy tip to the

radio-median cross-vein into the posterior cell, subalare, and in Ceriana/Sphixim-orpha, where it

but most of its length is found in the first basal is not recognisable. In Myathropa it reaches an

cell where it may proceed almost as far as that opposite extreme of development, the entire cell's inner end. Although popularly regarded as outer rim of the subalare giving off small, plum- one of the most characteristic features of the ulate appendages. Syrphidae, the vena spuria is entirely absent in According to Bonhag (1949) and Mickoleit some species, for example, Syritta flaviventris. (1962), who worked on Tabanus and Tipula, re- According to Wootton (I.e.), in Diptera the spectively, the haltere base possesses clustered Entomologie, deel 130, 1987 160 Tijdschrift voor

stg

^^^3 + 4 + ^1

S9 CuA2+A

nbesH, axillary region tenax, right wmg. Fig. 31, Syrphus F.g. 29, Syrphus nbesü, right wing. Fig. 30, EnstaUs of right wing. Speight: Morphology of Syrphidae 161

around it miniaturised versions of the axillary The first tarsal segment is usually referred to complex sclerites found round the base of the as the basitarsus or (confusingly) as the metatar- fore wings, but the haltere base sclerites do not sus. The undersides of the tarsal segments, in seem to have been examined in any species of particular, carry arrangements of short, blunt Syrphidae and they have not been investigated bristles used in cleaning the body surface and the for the purposes of the present account. The hind basitarsi have a ventral brush of close- haltere itself is a roughly dumbell-shaped struc- packed bristly hairs used for this same purpose. ture broadening rather abruptly at the distal end Although the number of leg segments remains of its stalk into the head or knob and more grad- the same throughout the family, in a significant ually at its base. number of genera one or more of the leg segments are modified in form, the modifications in nearly all cases being more pronounced in the Legs males than in the females. In some genera, such

The legs of syrphids comprise the following as Platycheirus, in which the male fore tarsal elements; coxa, trochanter, femur, tibia, five tar- segments are flattened and expanded laterally, sal segments (or tarsomeres) ^nàpretarsus, typ- the vast majority of species exhibit the same only sin- ified by the fore leg of Syrphus shown in fig. 34. general type of modification. In others The fore and hind coxae are single sclerites, fully gle species may be affected — thus Sphegina mobile and articulated basally to the thorax, ap- platychira males possess flattened tarsal seg- ically to the trochanter. Thie mid coxa, however, ments reminiscent of those of species of Platycheirus, but the tarsi of other Sphegina spe- is made up of three separate sclerites or coxites cies are unmodified. Almost any part of the leg (fig. 36). The anterior mesocoxite is fused to the thorax, but the posterior and median mesocox- may be affected by such modifications in Syrphi- either fore ites are mobile. The median mesocoxite articu- dae, though usually segments of or lates with the mesothoracic furcasternum via the hind legs are involved. Rarely, as in coxal condyle and also with the other two mesoc- Neocnemodon latitarsis, all three pairs of legs oxites. The posterior mesocoxite articulates with have some segments modified. In the male of N. the anterior mesocoxite as well as with the me- latitarsis the fore basitarsi exhibit a large pit on dian mesocoxite. one surface, the mid-tibiae are expanded into a Almost universally in Syrphidae — Microdon leaf-like flange for about half their length and the hind legs carry pro- representing the exception — there is a long, trochanters of both and mid blade-like process projecting outwards from the nounced tines (fig. 39), as do the hind coxae. A antero-lateral end of the outer side of the poste- frequent modification of the hind legs is for the rior mesocoxite. This blade-like process, termed hind femora to become bulbous and carry some here the trochanteral process of the posterior arrangement of pegs and or tubercles and for the ridged (fig. mesocoxite (figs. 35, 36) fits into a shallow hol- hind tibiae to become angular and low on the surface of the mesotrochanter, when 40). the leg is in certain positions. The abdomen In all three pairs of legs the segments from the trochanter outwards (inclusively) are all individ- There is great sexual dimorphism in the form ual sclerites, essentially tubular and articulated to of the terminal segments (and occasionally in the each other. Only the pretarsus is more complex. form of some of the more anterior segments as

This, the terminal leg-joint, is attached within well) of the syrphid abdomen, requiring that the the concave end of the most distal (the fifth) male and female abdomen be given detailed con- tarsal segment. The pretarsus includes a central sideration separately. In the male the abdomen is process terminating in a bristle-like or peg-like divided into two distinct sub-regions, a largely empodium, flanked by a pair of membraneous unmodified "preabdomen" and a highly modified pads known as the pulvillae, which are them- "postabdomen". Equivalent terms are not used in selves attached to the central process via small relation to the female abdomen. Certain features sclerotised plates called the auxilliae (fig. 37). of the abdomen common to both sexes may be

The lower surface of each pulvillus is in syrphids considered at this juncture. densely covered in short hairs. Above the pulvil- As viewed from above, the overall shape of the lae are found a pair of simple clau'S, which attach syrphid abdomen exhibits considerable variation. to a median apical projection of the 5th tarsal It may be conical, parallelsided, ovate or petio- segment known as the unguifer process. late. These shape differences are due largely to 162 Tijdschrift voor Entomologie, deel 130, 1987

tre acx

Fig. 32, Microdon rtiutabtlts, right wing. Fig. 33 Ceriana sp., right wing. Figs. 34—36, Syrphus ribesit, left fore leg, anterior view (34), male, base of left mid leg, antero-lateral view (35) and diagrammatic representation of mid coxa to show inter-relation between the three meso-coxites. Fig. 37, Microdon mutabilis, pretarsus and last two tarsomeres of left hind leg, ventral view. Speight: Morphology of Syrphidae 163

intergeneric differences in the shapes of the vis- divided in two by a transverse membraneous ible abdominal tergites, rather than to differen- strip. The sclerotised anterior part of st. 2 then ces in the number or identity of the sclerites in appears in some genera, e.g. Microdon, as a se- view. The number of visible tergites is four or parate, narrow plate lying along the entire ante- five in the male and generally five or six in the rior edge of the main sclerite (fig. 48). In Eristalis female. and Syrphus (fig. 47) this anterior plate of ster- The first abdominal tergite (t.l) is largely nite 2 is a lunulate piece confined to the middle fused with the second (t.2) in Syrphidae, though half of the width of st. 2. In Sphegina clunipes the junction between the two sclerites is usually (fig. 45) the two sclerotised parts of sternite 2 are evident. The anterior margin of t.l is frequently separated from each other by an appreciable dis- complex, since in most syrphids the flexion line tance, though the intervening membraneous sec- between thorax and abdomen passes through it, tion of the sternite evidently connects them to as in Eristalis and Syrphus (fig. 41). In such cases each other.

there is a median area of t.l which reaches forwards to the metanotum, flanked by narrower Male abdomen

lateral areas of t.l which do not reach the metan- In Diptera the male abdomen is regarded as otum and are partly separated from the median comprising eleven segments (numbered from area of t.l by an incomplete, transverse mem- the thorax to the abdomen tip). In the Cyclor-

braneous cleft. Paragus (fig. 42) is an exception. rhapa in general there is extensive modification In this genus the anterior margin of 1. 1 is straight of segments 6— 10 in the male due to their incor- and membrane intervenes between it and the poration into the copulatory apparatus. Segment

metanotum across its entire width, leaving the 11 is rudimentary, represented only by the cerei abdomen noticeable more capable of flexion than (which flank the anus). The term "preabdomen"

in most other genera. A prominent feature of is used to denote the unmodified portion of the certain genera, such as Ceriana, is a pair of an- male abdomen and "postabdomen" to refer to the tero-laterial calli on t.l, giving the impression modified portion. Zumpf & Heinz (1949) have that an extra sclerite is present on the front argued that the postabdomen commences with

margin of the tergite (fig. 43). segment 5 in Eristalis, since it structure is also Tergite 2 is a large sclerite with the joint be- greatly modified by involvement in the copula- tween it and tergite 3 (t.3) usually clearly marked. tory apparatus. Thompson (1972) points out that The actual anterior margin of t.3 is concealed defined in this fashion the preabdomen com- beneath the hind margin of t.2, the two tergites prises but four segments throughout the Milesii-

being joined to each other by folded membrane. nae, but is found in its more usual 5-segmented This type of junction is common to succeeding form throughout the Syrphinae. The sclerites of tergites except (e.g. in Microdon) where two ter- the male postabdomen have become markedly gites are fused to each other: tergites 2 and 3 are asymmetrical, associated with a twisting or tor- fused in Paragus; tergites 3 and 4 are fused in sion of the segments on the long axis of the Microdon. In Triglyphus tergite 3 is greatly ex- abdomen and a progressive recurvature of them panded, occluding t.4 and t.5 from view. such that when at rest the terminal segments With the exception of the first, the abdominal now face the front of the fly. spiracles are always found in the membraneous The sclerites representing the abdominal seg- strip between tergites and sternites. The first ments distal to segment 8 are in male Syrphidae

abdominal spiracle is more often than not en- highly modified to form a genital capsule or closed within the metathoracic epimeron which hypopygium, incorporating a complex intromit-

is (as described earlier) in part functionally a tent organ and accessory structures, together component of the abdomen, together with the with the anus and its flanking cerei. The hypop-

metanotum. In Microdon it was only found pos- ygium is concealed beneath the terminal tergites sible to locate the 1st abdominal spiracle during of the preabdomen when at rest, in a hollow the course of this study. termed by Cole (1972) the genital pouch. The abdominal sternites are generally rather poorly sclerotised rectangular sclerites with Male preabdomen rounded corners, lying free in the membrane of In species of genera such as Melanostoîna and the underside of the abdomen. The 1st abdominal Neoascia there are distinct though minor differ- sternite may be largely desclerotised and much ences in abdominal shape between males and reduced. The second abdominal sternite (st. 2) is females, produced by differences in the propor- 164 Tijdschrift voor Entomologie, deel 130, 1987 tions of tergites 1 —4 (fig. 44). More subtle dif- pair would subsequently face outwards from the ferences in the overall appearance of male and same surface as the sternite of the more proximal female abdomen occur in other genera, produced segment, and vice versa. in the same way. Quite precise differences be- Griffiths' (I.e.) contention that a 360° torsion tween male and female occur in the form taken has occurred requires, in his view, a 180° twisting by abdominal sternites in some genera, notably between segments 7 and 8 and a further 180° Speghina, Neocnemodon and Eumerus, where twisting between segment 8 and the hypopy- particular sternites of the male preabdomen ex- gium. The more traditional approach of Zumpt hibit structural modifications. The modifications & Heinz (1949) contends that a 90° twist has may be in the form of sclerotised outgrowths occurred between segments 5 and 6 and a second along the mid-line, as in Neocnemodon latitarsis 90° twist between segment 8 and the hypopy- (fig. 46) or paired lobes developed on the distal gium. Lehrer (I.e.) suggests there has been 180° margin of some sternite, as in Eumerus. of torsion between abdominal segments 5 and 6, a further 90° of torsion between segment 8 and Male postabdomen the hypopygium and a reflexion of the hypopy- Literature accounts of the degree of torsion gium from the longitudinal axis along which the exhibited by the abdomen of male Syrphidae are segments up to and including segment 8 are confusing, since in most instances it is stated that aligned. He also points out that the hypopygium syrphids possess a hypopygiu-m inversum or h. has in the latter process been pushed to one side. retroversum, but more recently it has been In the present text, Griffiths' (I.e.) hyptothesis claimed they show the h. circumversum condi- of 360° torsion in the syrphid male postabdomen tion. The most explicit discussion of these con- is regarded as unproven and the views of Zumpt flicting views is found in Griffiths (1972: 56) as & Heinz (I.e.) and of Lehrer (I.e.) are regarded follows: Failure to appreciate the conceptual dif- as each in part correct. It is presumed that 90° of ference between rotation and deflexion has led to torsion has occurred between segments 5 and 6 some confusion in the literature on Syrphidae, and a further 90° between segment 8 and the which I illustrate from the work of Zumpt and hypopygium, leaving segments 6—8 on their Heinz (1949). Zumpt and Heinz state that in sides and the hypopygium inverted. Subsequent Eristalis, "we are dealing with a hypopygium reflexion of the postabdomen, progressively inversum, thus apparently contradicting the view more complete from segment 7 onwards, has

(which I hold correct) that all Cyclorrhapha pos- then led to the present state of the abdomen. The sess a hypopygium circumversion. However, if superfluity of complex — and different — lati- Zumpt and Heinz's arguments are followed nisms, each deemed to define precisely the same closely, it will become apparent that they have features of the male syrphid postabdomen, sug- confused rotational movement and deflexion. gests that the practice of deriving such terms is

The hypopygium of Eristalis is "inverse" in the a redundant exercise and no such term is em- sense that it is so strongly deflexed that it points ployed here. Detailed discussion of torsion of the anteriorly and its "dorsal" side has become ven- male abdomen in Diptera may be found in Grif- tral." fiths (1972) and McAlpine (1981). To indicate the complex derivation of the Abdominal segments 5 —8 in Milesiinae and orientation of the hypopygium found in Syrphi- 6—8 in Syrphinae are much modified by the dae, which his torsion theory demands, Griffiths torsion process, going to form the "stalk" at the

(I.e.) suggests the term hypopygium circumver- end of which is born the hypopygium. In both sum et reflexum be used to describe the condition subfamilies tergites 6—8 remain reasonably of the syrphid postabdomen. Lehrer (1971a) large, externally visible sclerites, but 'the other reached conclusions different again from those of plates involved are reduced to poorly sclerotised Griffiths, dubbing the condition of the syrphid transverse strips concealed on or at the base of male postabdomen as a hypopygium inverso- the "stalk". Zumpt & Heinz (1949) provide a transversum. clear illustration of these sclerites, suggesting Whether the post-abdomen has twisted that at the base of the stalk on the right side of through 360° or 180°, the twisting has occurred the abdomen an extra, secondary, spiracle-bear- anterior to the hypopygium. It is presumed to ing sclerite has been developed. This they term have occurred between segments, such that if a the "intersegmental sclerite". Lehrer (1971a) twist of 180° occurred between a given pair of demonstrates that this "intersegmental sclerite" segments, the tergite of the distal segment of the is part of tergite 5, joined narrowly to the other Speight: Morphology of Syrphidae 165

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elements recognised by Zumpt & Heinz (I.e.)- The main structure concealed within the ba- Lelirer refers to these two elements of t.5 as sale when the hypopygium is at rest is the theca. hemitergites. The theca is essentially a sclerotised tube contain-

ing the aedeagus, but it usually carries sclerotised Male postabdomen: components of the terminal lobes of some complexity which vary hypopygium considerably in their form from species to species Unfortunately, a favourite "sport" of syrphid and genus to genus. The theca articulates to the morphologists and ta5?onomists has been to de- inner edge of the basal rim of the basale. The velop their own individual theories on the ho- terminal lobes of the theca are the lingula and the mologies of the component sclerites and struc- paired superior lobes. In its simplest form the tures of the hypopygium, and to accompany each lingula is a digitate or pointed projection, as in theory with its own set of names for the sclerites Syrphus (fig. 49). It may also be absent, as in involved. For instance, the main external plate of Eristalis (fig. 51). The paired superior lobes may the hypopygium bears on its outer end a pair of be either articulated to or fused with the main appendages, to be seen on either side of, and just body of the theca. In many Syrphinae they artic- distal to, the cerei. These appendages have var- ulate to cuticular outgrowths of the theca known iously been called dististyli, gonopods, gonostyli, as the lateral arms (fig. 49). In Microdon the ninth coxites, paralobi, styli and surstyli. Matsuda theca is largely membraneous and carries no (1976) points out that attempts to homologise structure that could be homologised with either sclerites of the male ano-genital complex of one lingula or superior lobes (fig. 50). family of Diptera with those either of the geni- Three principal elements have been identified talia of other Diptera or of the original segments in the syrphid aedeagus. The most distal of these of the abdomen of Insecta in general are almost is the tubus, most often a weakly sclerotised, inevitably doomed to failure, because (a) they do trumpet-shaped sclerite, which protrudes beyond not arise ontogenetically from homologous pri- the end of the theca (fig. 49). The tubus may be mordia in different families, and (b) secondary entirely lacking (e.g. in Eristalis, fig. 51). The structures with no antecedents elsewhere are tubus passes down into the pyxis, a roughly ring- found repeatedly. It would thus appear that to shaped sclerite which sits in the mouth of the attempt to name all parts of the syrphid hypop- theca, with which its outer edge articulates. The ygium in such a way as to suggest precise homol- pyxis may itself carry a pair of sclerotised, rather ogies for the individual sclerites is by and large hook-like outgrowths known as the harpes, pointless. The names employed in this text for which often project from within the theca (fig. parts of the hypopygium are chosen from among 51). Beneath the pyxis and articulated to it is the existing terms which do not suggest particular aedeagal apodeme, the tip of which is external origins for sclerites. This approach is the con- and may be visible within the theca (fig. 49). verse of that employed by McAlpine (1981). Microdon again differs significantly from other When at rest tucked into the end of the pre- hoverflies in possessing a pair of simple, whip- abdomen (figs. 47, 48) the most obvious feature like sclerotised tubes as the sole aedeagal struc- of the hypopygium is the convex rim of a scoop- tures protruding from and contained within the shaped sclerite (behind which the other struc- theca. Metcalf (1921) recognised a structure deep tures are hidden from view) bearing the closely within the hypopygium of Microdon as homolo- opposed cerei in a membraneous cleft on its distal gous with the pyxis of other syrphids.

edge, which is oriented to point towards the This rather basic account of the principal com-

head-end of the fly. This scoop-shaped sclerite is ponents of the syrphid hypopygium falls far

referred to here as the basale, which is joined short of providing a reasonable indication of the

round its basal rim to the distal rim of tergite 8. range of variation they exhibit in different ho- At the distal end of the basale, more or less verflies. But in this one family of flies these struc- flanking the cerei, are attached a pair of appen- tures exhibit a quite remarkable range of forms dages here termed the styli. The styli are articu- and the male genitalia of many syrphid species lated to the basale rather than fused with it, and have been illustrated elsewhere recently: see on the inner wall of the basale are continuous Dusek & Laska (1964, 1967), Gaunitz (I960, with a weakly sclerotised plate here called the 1966, 1969), Glumac (I960), Goeldlin (1976), minis. They may be simple, thumb-shaped pie- Hippa (1968, 1978), Metcalf (1921), Thompson ces, as in Eristalis (fig. 51), or more complex (1972), and Vockeroth (1969). structures with more than one lobe. Speight: Morphology of Syrphidae 167

Fig. 44, Neoascia podagrica, male (left) and female (right), abdomen, dorsal view. Fig. 45, Sphegina clunipes, female, base of abdomen, ventral view. Fig. 46, Neocnemodon latitarsis, male, basal abdominal segments, lateral view, left side. Fig. 47, Eristalis tenax, male abdomen, ventral view. Fig. 48, Microdon mutabilis, male abdomen, ventral view. 168 Tijdschrift voor Entomologie, deel 130, 1987

Female abdomen designation tergite 9 is retained for this plate, Matsuda (1976) interprets the abdomen of fe- what are the small lateral selerites regarded as male syrphids as comprising discernible ele- tergite 9 by Lehrer? Authors other than Lehrer ments of nine segments, plus the cerei. An alter- have conveniently ignored any mention of these

native interpretation is provided by Lehrer lateral selerites. Neither of the terms epiproct (1971b), who follows Herting (1957) in identi- and proetiger can be used for the "tergite 9" of fying elements of ten segments plus the cerei. Matsuda and McAlpine because both have partic-

Matsuda (I.e.) points out that the nine segments ular morphological connotations. Further, the he recognises cannot be assumed to be homolo- morphological definition of these terms varies gous in all dipteran families, because of the pos- with author, as a comparison of the definitions in sibility that the particular segment which has McAlpine and Tuxen (1970) demonstrates. been lost is not the same in all instances. Probably the most neutral of the available terms

Both Lehrer (i.e.) and Matsuda (I.e.) agree in that have been applied to this sclerite in Diptera numbering the first eight visible abdominal seg- is the supra-anal plate, so that is what it has been

ments of female syrphids from 1 to 8, sequen- termed here (see fig. 53 onwards, sap). tially, with each segment represented by a tergite The pair of lateral selerites regarded by Lehrer and sternite. The first 7 segments each carry a (I.e.) as derived from tergite 9 are not given any spiracle. The spiracles lie in the membrane be- particular designation here. They are indicated in tween tergite and sternite towards the anterior fig. 57 (fig. 57, Is). Lehrer was able to detect these end of each segment, except in the case of the 1st selerities in Ceriana, Volucella and Xylota, as well

abdominal spiracle, which is more frequently as in Eristalis. found entirely or partly enclosed within the pos- Considering the ventral abdominal selerites terior part of the metathoracic epimeron. distal to sternite 8, Eristalis appears to be excep- Beyond the eighth segment the situation be- tional in exhibiting the sclerotised strip regarded comes confused. Dorsally, Matsuda recognises a by Lehrer (I.e.) as sternite 9. In the other species tergite 9 flanked by the cerei. Matsuda's tergite 9 examined (including Volucella bombylans and V.

is Lehrer's tergite 10 and the epiproct or proc- pellucens) during preparation of this account the tiger of some other authors. For Lehrer, tergite most that is visible in this position is a median,

9 is identified with a pair of tiny lateral selerites membranous flap which serves as an egg guide. that are unconnected externally. These rudimen- Seeing the uncertain homology of this feature it tary plates are not mentioned by Matsuda. is referred to here not as sternite 9 but as the Ventrally, Matsuda identifies in his text "the ventral egg-guide (figs. 56 on, vg). definitive 9th sternum or the postgenital plate as A pair of lateral sclerotised or membraneous seen in Eristaiis". He thus interprets this "post- egg guides may also be present. They have been genital plate" as sternite 9- identified by Lehrer as elements of tergite 9. In the range of Syrphidae examined for pur- Since they are linked by a sclerotised bar passing poses of the present account, the number of seler- along the upper margin of the genital opening, ites noted distal to the eighth tergite and sternite these lateral egg guides may be of sternal origin. is not consistent. For instance, in Syrphus no The lateral egg guides are well exemplified by dorsal selerites distal to tergite 8 are found, al- Volucella (fig. 58, Ig). In Eristalis and Sericomyia though at least one additional dorsal sclerite dis- the ventral egg-guide is well-developed but the tal to tergite 8 occurs in many other genera. lateral ones are lacking, while in Syrphus no egg- Similarly, although in Eristalis a narrow sclero- guides are differentiated (figs. 56— 62). tised strip (st. 9 of Lehrer) occurs distal to ster- The terminal ventral sclerite of the abdomen nite 8 and basal to the genital opening, this is the sternite 9 or post-genital plate of Cramp-

sclerotised strip is not present in other genera ton (1942), Matsuda (1976) and McAlpine examined. Further, there are indications that se- (1981), which is sternite 10 of Lehrer (1971b). condary sclerotisation of membranous areas has This plate appears to be universally present.

on occasion occurred, as in Eumerus (fig. 55, ss). Since its homology is questionable it is referred What terminology to use for selerites in the to here as the sub-anal plate (fig. 56 onwards,

female abdomen posterior to tergite and sternite sup). The anus is located between the sub-anal

8 is problematic. Lehrer's (I.e.) argument that the plate and the supra-anal plate and is flanked by

so-called "tergite 9" of Matsuda and McAlpine is the cerei, which are generally recognised as being

in reality tergite 10 is persuasive. And if the derived from the original eleventh abdominal Speight; Morphology of Syrphidae 169

hp.

Fig. 49, Syrphus ribesii, male, hypopigium, lateral view. Fig. 50, Microdon mutabilis, male, hypopygium, lateral view. Fig. 51, Eristalis tenax, male hypopygium, lateral view.

segment. In Syrphus, where the supra-anal plate produced a "false segment" between segments 6

is lacking, the cerei come to occupy a mid-dorsal and 7 and another "false segment" between seg- position (fig. 60). ments 8 and 9- In the latter case a complete ring In many syrphids the abdominal segments of sclerotised membrane has formed (fig. 55, ss). posterior to segment 5 are hardly visible when at rest, being then retracted telescopically into seg- The relationship between the Syrphidae ment 5, but they do not really form a distinct sub- AND genera allied TO MiCRODON region of the abdomen as in the male. These terminal segments may be modified to form an Repeated references are made during course of ovipositor as long as or even longer than the first this account to differences existing between the five abdominal segments together, the elonga- morphology of Microdon and the morphology of tion being achieved primarily by widening the other syrphids. Various of these differences e.g. bands of intersegmental membrane, as in in Microdon's mouthparts, have not been alluded Eristalis and Sericomyia (figs. 53, 54). In to in other texts. Microdon and allied genera have

Eumerus a similar result is achieved by elonga- until recently usually been regarded as constitut- tion of the largely membraneous segment 8. Also ing a separate subfamily, the Microdontinae, in Eumerus, secondary sclerotisation of the atten- within the Syrphidae. uated bands of intersegmental membrane has Thompson (1969) reviewed the genera con- Entomologie, deel 130, 1987 170 Tijdschrift voor

sap. 53

sap

lateral view, right side. F.g. 53, ErtstaUs tenux, female abdomen, Fie 52 Syrphus ribem, female, abdomen, lateral abdomen, lateral view, right side. Fig. 55, Eumerus stngatus, view, right side. Fig. 54, Sencomyta stlentn, female, right side. female, distal abdominal segments, lateral view, Speight: Morphology of Syrphidae 171

8 ^

signed to the Microdontinae and provided a de- be: finition of the subfamily, based upon both adult and larval characteristics. He concluded that "the Microdontines should be considered the first div- ergence in the phylogeny of the family (Syrphi- dae)". He further remarked that the "strongly plesiomorphic nature of the subfamily suggests that the microdons might best be considered as a separate family". In a later paper Thompson (1972) separated the Microdontidae from the Syrphidae. The additional data provided in the present account would tend to support Thomp- son's action, but does only relate to the European genus Microdon. If the other microdontine genera differ from the rest of the syrphids in these same ways, there would seem little justifi- cation for retaining the Microdontinae within the Syrphidae. The incongruity attendent upon retention of the Microdontinae within the Syrphidae is high- lighted by bringing into consideration the degree of morphological difference found between other dipteran groups currently recognised as separate families. However, Thompson's recognition of the family Microdontidae has not been adopted by other authors. For that reason Microdon has been included in this account of syrphid morphology, despite the present author's doubt that the genus belongs in the family Syrphidae.

Abbreviations used in hgures a: arista of antenna A: first anal vein of wing aa: aedeagal apodeme of aedeagus of male genitalia ac: anteclypeus ae: aedeagus of Microdon male genita-

lia acs: anterior cervical sclerite acx: anterior mesocoxite of middle leg al: alula of wing am: aristomere an: anal cell of wing ans: anapleural suture of mesothoracic pleura as: antecostal suture at: acrotergite of mesothoracic notum att: anterior tentorial pit au: auxillia of pretarsus of leg axj, etc.: axillary sclerite 1, etc. bj, bj: first and second basal cells of wings ba: basale of hypopygium of male genitalia bat: buccal arm of anterior tentorial sulcus Speight: Morphology of Syrphidae 173

median vein of wing pis: pleural suture of mesopleura branches of median vein of wing pn: postpronotal sclerite of pronotum marginal cell of wing of thorax anepisternites of mesothoracic po: posterior cell of wing pleura pocb: post-metaxocal bridge mesofurcal pit ps: proepisternum of propleura median-cubital cross-vein of wing pu: prothoraeic furcasternum median mesocoxite of middle leg R: radial vein of wing mesepimeral sclerite of mesotho- rm: radial-median cross-vein of wing racic pleura Rsi_,: branches of the radial-sector vein of median hypostomal suture the wing katepisternum of mesothoracic scape of antenna pleura sa: subalare mesonotal prescutum sb: band of sensilla on postgena premental sclerite of the labium Sc: subcostal vein of wing mesosternal presternum sc: subcostal cell of wing meropleurite of mesothoracic sea: postalar callus of mesonotum of pleura thorax mesoscutum of mesonotum scr: subcostal-radial cross-vein of wing scutellar lobe of mesonotum se: sella of cervical organ median postnotal sclerite of me- sen: sensilla of cervical organ sonotum sep: sensory pit of 3rd antennal seg- basisternum of metathoracic ster- ment num si: superior lobe of theca of male geni- epimeron of metathoracic pleura talia metepimeral callus of metathoracic sm: submarginal cell of wing pleura sp: spiracle epimeral spine of the metathoracic sps: spiracular sclerite of Microdon pleura thorax metathoracic notum ss: secondary sclerite precoxale of metathoracic pleura ssu: parasagittal sulcus of head capsule episternum of metathoracic pleura stj, stj, etc. sternum of first abdominal seg- furcasternum of metathoracic ster- ment, second abdominal segment, num etc. anterior mesosternal furcasternum st2a: anterior sclerite of abdominal ster- posterior mesosternal furcaster- nite 2 num stg: stigma of wing mid-ventral thoracic suture sy; stylus of basale of hypopygium of maxillary stylet male genitalia

maxillary palp ti, t2, etc.: tergite of first abdominal segment, ocular arm of anterior tentorial sul- second abdominal segment, etc. cus ta^, ta,: tarsal segments of leg post-ocular orbits te: callus of 2nd tergite of abdomen orbital strip of face tg: tegula post-occipital sclerite th: theca of hypopygium of male geni- ocellar triangle talia pedicel of antenna tho: thorax

antepronotum of pronotum of ti: tibia of leg thorax tp: posterior tentorial pit of head cap- pc: postclypeus sule pcb: premetaxocal bridge tr: trochanter of leg pec: postcranial carina tre: trochanteral process of posterior pes: posterior cervial sclerite mesocoxite of middle leg pe: proepimeron of propleura ts: transverse sulcus

tt: of capsule Pg- postgena tempota head 174 Tijdschrift voor Entomologie, deel 130, 1987

tubus of aedeagus of male genitalia lia i tribusa. — Bulletin of the Museum of Natural pulvillus of pretarsus of leg History, Belgrade (B) 16: 69—103. Goeldlin de Tiefenau, P., 1976. Révision du genre vertex of head capsule Paragus (Dipt. Syrphidae) de la région palearc- vs: vena spuria of wing tique occidentale. — Bulletin de la Société entomo- vv: postVertex of head capsule logique Suisse 49: 79— 108. Goffe, R. E., 1947. The wing venation of Syrphidae Bibliography (Diptera). — Entomologists Monthly Magazine Termier, 1973. Bitsch, J.,J. R. Denis, E. Seguy, & M. 83: 225—239. Insects: tête, aile. - Traité de Zoologie 8 (1): 799 Gouin, F, 1949. Recherches sur la morphologie de pp. — Masson, Paris. l'appareil buccal de Diptères. — Memoirs du Mu- Insectes: thorax, abdo- Bitsch, J., & R. Matsuda, 1973. séum National d'Histoire Naturelle, Paris, N.S 28: men. — Traité de Zoologie 8 (2): 600 pp. — 167—269. Masson, Paris. Griffiths, G. C. D., 1972. The phylogenetic classifica- Bonhag, P. F., 1949. The thoracic mechanism of the tion of Diptera Cyclorrhapha, with special refer- adult horsefly. — Memoirs of the Cornell Univer- ence to the structure of the male postabdomen: 340 sity agricultural Experimental Station 285: 3 — 39. pp. — Junk, The Hague. Coe, R. L., 1953. Syrphidae. — Handbooks for the Herting, B., 1957. Das weibliche Postabdomen der identification of British insects 10 (1); 98 pp. — Calyptraten Fliegen (Diptera) und sein Merkmals- Royal Entomological Society, London. wert für die Systematik der Gruppe. — Zeitschrift Cole, F. R., 1927. A study of the terminal abdominal für Morphologie und Ökologie der Tieren, Berlin structures of male Diptera (two-winged flies). — 45: 429—461. Proceedings of the Californian Academy of Hippa, H., 1968. A generic revision of the genus Science, 4th Series 16 (14): 397—499. Syrphus and allied genera (Diptera Syrphidae) in Colless, D. H. & D. K. McAlpine, 1970. Diptera. In: the Palearctic region with descriptions of the male 1 The Insects of Australia: 656—740. — CSIRO, genitalia. — Acta entomologica Fennica 25 : —94. Canberra. Hippa, H., 1968. Classification of Xylotini (Diptera, Crampton, G. C, 1942. The external morphology of Syrphidae). — Acta Zoologica Fennica 156: the Diptera. In: Crampton, G. C, C. H. Curran & 1—153. C. P. Alexander. The Diptera or true flies of Con- Holloway, B. A., 1976. Pollen-feeding in hover-flies necticut, pt. 1: external morphology; key to fam- (Diptera: Syrphidae). — New Zealand Journal of ilies; Tanyderidae; Ptychopteridae; Trichoceridae; Zoology 3: 339—50. Anisopodidae; Tipulidae. — Connecticut Geolog- Hull, F M., 1945. A revisionai study of the fossil ical and Natural History Survey, Bulletin 64: Syrphidae. — Bulletin of the Museum of Compar- 10—174. ative Zoology, Harvard 95 (3): 251 — 355. P. Laska, 1964. contribution to distin- inter-relation- Dusek, J. & A Hull, F M,, 1949. The morphology and guishing the European species of the subgenus ship of the genera of Syrphid flies, recent and Syrphus Fabricius (Dipt. Syrphidae) according to fossil. — Transactions of the Zoological Society, male genitalia and larvae. — Acta Societas Ento- London 26 (4): 257—408. mologicae Cechosloveniae 61: 58— 70. Lehrer, A. Z., 1971a. Valeur morphologique des sclér- eines Dusek, J. & P. Laska, 1967. Versuch zum Aufbau ites abdominaux et homologie des terminalia naturlichen Systems mitteleuropaeischer Arten maies des Diptères Cyclorrhapha. 1. Fam. Syrphi- der Unterfamilie Syrphinae (Diptera). — Acta dae. — Bulletin de la Société d'Entomologie de scientiarum naturalium Academiae Scientiarum Mulhouse, Jan-Fev. 1971: 1—18. bohemoslovacae Brno 1: 349— 390. Lehrer, A. Z., 1971b. Morphologie et homologie des Gaunitz, S., I960. Syrphidenstudien III (Dipt.). — sclérites abdominaux femelles chez les Diptères Entomologisk Tidskrift 81 (1—2): 35—44. Cyclorrhapha. — Bulletin de la Société d'Entomol- Gaunitz, S., 1966. Syrphidenstudien V. Zur Kenntnis ogie de Mulhouse, Juillet- Août 1971: 59—67. vom Bau des Kopulationsapparates beim Männ- Maki, T., 1948. Studies of the thoracic musculature of chen einiger Helophilusarten. — Entomologisk insects. — Memoirs of the Faculty of Science and Tidskrift 87 (1—2): 69— 71. Agriculture of Taihoku imperial University 24: Gaunitz, S., 1966. Studien über die Unterfamilie Er- 1—343. istalinae. Der Bau des männlichen Genitalapparats Matsuda, R., 1965. Morphology and evolution of the (Dipt. Syphidae). — Entomologisk Tidskrift 90 insect head. — Memoirs of the American Entomo- (1—2): 79—99. logical Institute 4: 343 pp. Gilbert, F. S., 1981. Foraging ecology of hoverflies: Matsuda, R., 1970. Morphology and evolution of the morphology of the mouthparts in relation to feed- insect thorax. — Memoirs of the Entomological urban ing on nectar and pollen in some common Society of Canada 76: 431 pp. 245 262. species. — Ecological Entomology 6: — Matsuda, R., 1976. Morphology and evolution of the Glumac, S., I960. Priordan sisten sirfida (Syrphidae insect abdomen: 534 pp. Pergamon, Oxford and

Diptera) zasnovan no gradi genitalnog aparata i New York. nacinu razvitka earava so Karakteristikanna fami- McAlpine, J. F, 1981. Morphology and terminology: Speight: Morphology of Syrphidae 175

adults. In Manual of Nearctic Diptera 1: 9 —63- Speight, M. C. D., 1969. The prothoracic morphology Research Branch, Agriculture Canada, Monograph of acalypterates (Diptera) and its use in systemat- 27. ics. — Transactions of the Royal Entomological Metcalf, C. L., 1921. The genitalia of male Syrphidae: Society, London 121: 325 —421. their morphology, with especial reference to its Speight, M. C. D., 1980. The Chrysogaster species taxonomie significance. — Annals of the Entomo- (Dipt. Syrphidae) known in Great Britain and Ire- logical Society of America 14: 169— 214, 19 pis. land. — Entomologists Record and Journal of 'Va- Mickoleit, G., 1962. Die thorax musculatur von Tipula riation 92 (6): 145—50. vernalis Meigen. Ein Beitrag zum vergleichenden Stackelberg, A. A., 1965. New data on the taxonomy Anatomie die Dipterenthorax. — Zoologische of palaearctic hoverflies. — English language ver- Jahrbücher (Abteilung für Anatomie), Jena 80: sion in: Entomological Society of America, Scripta 213—244. Technica, Washington 4: 528— 537. Thompson, F. C, 1969. A new genus of Microdontine Nayar, J. L., 1964. External morphology of head capsule of Syrphus balteatus De Geer (Syrphidae, Dip- flies (Diptera: Syrphidae) with notes on the place- tera). — Indian Journal of Entomology 26: ment of the subfamily. — Psyche 76 (1): 74—85. 135—151. Thompson, F C, 1972. A contribution to a generic Nayar,J. L., 1965. Morphology of a hover fly Syrphus revision of the neotropical Milesiinae (Diptera: balteatus de Geer (Syrphidae, cyclorrhapha: Dip- Syrphidae). — Arquivos de Zoologia, Sao Paulo 23 tera). — Journal of Research of the Agra Univer- (2): 73-215. sity 14: 145—149. Tuxen, S. L. (ed.) 1970. A taxonomist's Glossary of Oldroyd, M., 1970. Diptera: Introduction and key to Genitalia in Insects. — Munksgaard, Copenhagen.

families. — Handbooks for the identification of 'Vockeroth, J. R,, 1969. A revision of the genera of the British insects 9 (1): 104 pp. — Royal Entomolog- Syrphini (Dipt. Syrphidae). — Memoirs of the ical Society, London. Entomological Society of Canada 62: 176 pp.

Schiemenz, H., 1957. Vergleichende funktionell-ana- 'Wootton, R. J., 1979. Function, homology and termi- tomische Untersuchungen der Kopfmuskulatur nology in insect wings. — Systematic Entomology von Theobaldta und Eristalis (Dipt. Culicid. und 4: 81—99. Syrphid). — Deutsche Entomologische Zeitung, Young, B. P., 1921. Attachment of the abdomen to the N. E 4 (5): 268— 331. thorax in Diptera. — Cornell University Memoirs Séguy, E., 1959. Introduction à l'étude morphologique 44: 255—82. de l'aile des insectes. — Memoirs du Muséum Na- Zumpf, F & H. Heinz, 1949. Studies on the sexual tional d'Histoire Naturelle A 21: 248 pp. armature of Diptera: a contribution to the study of Séguy, E., 1961. Diptères Syrphides de l'Europe occid- the morphology and homology of the male termi- entale. — Memoirs du Muséum National d'His- nalia of Eristalis tenax L. (Syrphidae). — Entomol- toire Naturelle A 23: 248 pp. ogists Monthly Magazine 85: 299— 306. Snodgrass, R. E., I960. Facts and theories concerning the insect head. — Smithsonian Miscellaneous collections 142 (1): 1—61.

Tijdschrift voor Entomologie 130; 177 —209 Gepubliceerd 30 november 1987

TAXONOMY AND BIOGEOGRAPHY OF ORIENTAL PRASIINI. 3. THE FATIWQUA AND PARVULA GROUPS OF THE GENUS LEMBEJA DISTANT, 1892 (HOMOPTERA, TIBICINIDAE)

M. R. DE JONG

Institute of Taxonomie Zoology (Zoölogisch Museum), University of Amsterdam, The Netherlands

Abstract

Tentative concepts for two species-groups of the genus Lembeja Distant, 1892, the fatiloqua and the parvula group, are presented. The widely distributed fatiloqua group incorporates ten species, viz., Lembeja fatiloqua (Stài, 1870) from Mindanao (Philippines) and North Borneo, L. consanguinea n.sp. from North Sulawesi, L. maculosa (Distant, 1883), L. fruhstorferi Distant, 1897, L. lieftincki n.sp., L. sanguinolenta Distant, 1909, and L. tincta (Distant, 1909) from South Sulawesi, L. roehli Schmidt, 1925 from Sumba, L. sumbawensis n.sp. from Sumbawa, and L. paradoxa (Karsch, 1890) from SE New Guinea, Torres Strait Islands and Cape York Peninsula. The parvula group is confined to Sulawesi with two species, viz., L. parvula n.sp. from South Sulawesi and L. wallacei n.sp. from North Sulawesi. Char- acters and character states are discussed in connection with the supposed monophyletic status of the species-groups. All species but L. paradoxa are (re)described and structures of taxonomie importance as well as the whole insects are depicted. A key to males and

females is presented.

Introduction sidered a synapomorphous character for the In previous publications (De Jong, 1985, 1986) species of the group. on the Oriental Prasiini, the genus Prasia Stal, Another highly characteristic feature of the 1863, and the /oliata group of the genus Lembeja group is the capability of the males to inflate Distant, 1892, have been defined and the species their abdomen in a probably unique manner (see incorporated have been (re)described. The pres- Moulds, 1975). The tergites 3 —6, when tele- ent studies of the fatiloqua and parvula groups scoping from under their preceding tergites, of the genus Lembeja are further contributions show clearly the, sometimes broad, intersegmen- to a revision of the oriental Prasiini (see also tal membranes. This feature is also displayed De Jong & Duffels, 1981; De Jong, 1982). For by the African genus Iruana Distant, 1905 a review on the history of the genus the reader (Boulard, 1975, 1981, 1985). This genus was for-

is referred to De Jong (1986). merly attributed to the Prasiini (Metcalf, 1963), but is probably more related to other genera Concepts of the fatiloqua and the than to those constituting the Prasiini. The tel- parvula groups escoping abdomen most probably developed in- (with notes on their relationships and dependently in the two groups. distribution) Species attributed to the fatiloqua group are L. consanguinea n.sp., L. fatiloqua (Stal, 1870), Monophyly of the fatiloqua group L. fruhstorferi Distant, 1897, L. lieftincki n.sp., L. maculosa (Distant, 1883), L paradoxa

The fatiloqua group is characterized by a mod- (Karsch, 1890), L. roehli Schmidt, 1925, L. san- erately to strongly developed, longitudinal me- guinolenta Distant, 1909, L. sumbawensis n.sp.

dial dent in the male tergite 1, which is con- and L. tincta (Distant, 1909).

177 178 Tijdschrift voor Entomologie, deel 130, 1987

Monophyly of the parvula group The small-sized species of the parvula group are very similar to one another.

The parvula group is characterized by the obliquely hindwards running edge of the pygofer Relationships between the fatiloqua and the par- between the lateral lobe and the caudodorsal vula groups beak, which is considered a synapomorphy for the species of the group. The male tergite 1 is medially slightly dented Furthermore, the wings have five apical areas near its proximal border in the parvula group, instead of the usual number of six. This character and moderately to strongly dented in the fati- state is also displayed by an undescribed species loqua group. On account of the more or less from New Guinea, belonging to the Oriental strongly developed medial dent the groups are Prasiini. Because this species is probably more tentatively regarded sister-groups. related to one, or more, of the other species- groups or genera, than it is to the parvula group, Distribution of the fatiloqua and parvula groups this character state cannot be used as a strong synapomorphy for the species of the parvula The fatiloqua group is by far the most wide- group. spread group of the genus. It is distributed from Finally, the male tergite 1 is only very slightly Mindanao (Philippines), North Borneo, North medially dented in the parvula group. and South Sulawesi, Nusa Tenggarah (Lesser Species attributed to the parvula group are Sunda Islands), SE New Guinea, Torres Strait L. parvula n.sp. and L. wallacei n.sp. Islands up to the Cape York Peninsula (North Queensland).

The parvula group is distributed in North and Relationships within the species-groups South Sulawesi.

Some character states suggest that the fati- Both groups show a distribution that seems loqua group can be divided into several sub- hard to reconcile with their supposed monophy- groups. As there is no evidence yet whether the letic origins. Both the fatiloqua group and the character states involved are to be interpreted parvula group show a peculiar disjunction in Su- as plesio- or apomorphic, a possible subdivision lawesi by their restricted occurrence in the most of the species-group is postponed until outgroup eastern part of the Minahassa Peninsula and in comparison provides more information on those the most southern part of the island; no repre- characters. These characters are: sentatives of these groups are found in Central Pigmentation of tegmina. — The males of Sulawesi. The collections made by the brothers five species have unspotted (sub)hyaline teg- Sarasin in Central Sulawesi at the end of the mina, viz., Lembeja fatiloqua, L. maculosa, L. 19th century, as well as recent collecting during roehli, L. sumbawensis n.sp. and L. tincta. The Operation Drake in Morowali N.P. in 1980 and P. males of four species have spotted tegmina: L. by Dr J. & Mrs M. J. Duffels, and Mr J. van consanguinea n.sp., L. lieftincki n.sp., L. paradoxa Tol in Lore-Lindu N.P. in 1985, did not provide and L. sanguinolenta. The remaining species, L. any material belonging to these two species- fruhstorferi, is known only from its female hol- groups. otype. The female tegmina are spotted, except L. paradoxa of û\e fatiloqua group has a widely in L. fatiloqua, L. roehli and L. sumbawensis n.sp. remote distribution in SE New Guinea, the Apical lobes of the aedeagus. — All species Torres Strait Islands and the Cape York Penin- with spotted tegmina in the males possess an sula of Australia; no other representatives of aedeagus with long, rounded apical lobes, where- the group are found in New Guinea. as those with unspotted male tegmina have an aedeagus with short, pointed to rounded, apical Depositories lobes or without lobes. However, one undes- The abbreviations given below have been used cribed species with spotted tegmina, only known in the list of material and throughout the text. from poor material, is provided with an aedeagus with short and pointed lobes. BIN Koninklijk Belgisch Instituut voor Na- Uncus. — The males of three species, viz., tuurwetenschappen, Brussel L. consanguinea n.sp., L. lieftincki n.sp. and L. BISH Bernice P. Bishop Museum, Honolulu sanguinolenta, have an uncus that is enlarged BMNH British Museum (Natural History), medially above the claspers. London M. R. DE Jong: Oriental Prasiini 179

DEI Deutsches Entomologisches Institut, I am indebted to G. Mr Verlaan and Mr J. Eberswalde Zaagman for technical assistance and to Mr L ESC Florida State Collection, Gainesville van der Laan for the photographs. of Comparative Zoology, Har- MCZ Museum The participation of Dr J. P. Duffels in the vard University, Cambridge "Project Wallace" expedition was supported by MW Institut Zoologique, Warszawa the Netherlands Eoundation for the Advance- MZB Museum Zoologicum Bogoriense, Bo- ment of Tropical Research (WOTRO: gor WR85— 197). NBM Naturhistorisches Museum, Basel The present investigations were supported by NHMW Naturhistorisches Museum, Wien the Eoundation for Fundamental Biological Re- NMWC Natural Museum Wales, Cardiff search (BION), which is subsidized by the NRS Naturhistoriska Riksmuseet, Stock- Netherlands Organization for the Advancement holm of Pure Research (ZWO). RMNH Rijksmuseum van Natuurlijke Histo- rie, Leiden Taxonomy SMD Staatliches Museum für Tierkunde, A short characterization of each species-group Dresden will preceed the descriptions of the species, in- SMN Staatliches Museum für Naturkunde, cluded in the group. Stuttgart All methods of investigation follow De Jong TMB Természettudomany Muzeum, Buda- (1985, 1986). The female genitalia will be de- pest scribed and discussed separately in a paper deal- USNM United States National Museum, ing with the female genitalia of the oriental Pra-

Smithsonian Institution, Washington siini. ZIM Zoologisches Institut und Zoologisches Museum, Hamburg Key to the species of the fatiloqua and parvula ZMA Instituut voor Taxonomische Zoologie, groups

Zoologisch Museum, Amsterdam 1. Wings with 6 apical areas. Medium-sized to ZSM Zoologische Staatssammlung, Mün- large species (body length $: 17.9 —21.8 mm, chen. $: 14.9 —24.5 mm). $: tergite 1 with a strong longitudinal medial dent fatiloqua group 2 Acknowledgements — Wings with 5 apical areas. Small-sized spe-

I very am obliged to Dr J. R Duffels, Dr R cies (body length $: 12.5 — 16.2 mm, $; 11.8 J. Oosterbroek and Prof. Dr J. H. Stock for critical — 13.7 mm). $ : tergite I with a weak, reading of the manuscript. proximal medial dent parvula group 19 The material studied was gratefully received 2. Tegmina (sub)hyaline, spotted 3 from Mr R. Detry (BIN); Dr E ]. Radovsky — Tegmina (sub)hyaline, unspotted, some- and Dr C. A. Samuelson (BISH); Dr W.J. Knight, times greenish or yellowish opaque .... 12 Mr M. D. Webb and Mr R S. Broomfield 3. Males 4 (BMNH); Prof. Dr H.J. Müller, Dr G. Petersen — Females 7 A. and Dr Täger (DEI); Dr J. B. Heppner (ESC) 4. Abdomen with large mediodorsal spines on Mrs M. K. Thayer (MCZ); Dr E. Kierych (MW) segments 4— 7. Papua New Guinea, Torres Dr S. Adisoemarto (MZB); Dr M. Brancucci Strait islands, Cape York Peninsula (NBM); Dr A. Kaltenbach es. (NHMW); Dr paradoxa M. E Claridge (NMWC); Dr P Lindskog (NRS); — Abdomen without such large spines. Su- P. H. van Dr Doesburg and Mr J. van Toi lawesi 5 (RMNH); Dr E. Emmrich (SMD); Mr E Heller 5. Body green; aedeagus provided with dorsal (SMN); Dr Z. Kaszab (TMB); Dr R. C. aedeagal appendage. North Sulawesi Eroeschner (USNM); Prof. Dr H. Strümpel consanguinea (ZIM) and Dr M. Baehr (ZSM). This paper is — Body brownish, reddish or ochreous; aedea- based in part on material collected whilst the gus without dorsal aedeagal appendage. author was a participant on Project Wallace, South Sulawesi 6 sponsored by the Royal Entomological Society 6. Medium-sized species (body length: of London and the Indonesian Institute of Sci- 21.4— 21.6 mm); tegmina, though spotted, ences (Results of Project Wallace No. 32). reasonably hyaline. Median uncus part amply 180 Tijdschrift voor Entomologie, deel 130, 1987

enlarged in the shape of a tube-like structure. length: 22.8—26.2 mm); aedeagus with South Sulawesi lieftincki hooked apex provided with two pointed — Large species (body length: 24.6 mm); teg- flaps. South Sulawesi tincta mina heavily pigmented with red. Median — Body greenish-yellow and medium-sized uncus part only slightly enlarged. South Su- (body length: 18.8—22.9 mm); aedeagus lawesi sanguinolenta straight without apical flaps. Sumba

7. Abdomen with three longitudinal fasciae: roehli one distinct, dorsal medial fascia and two 17. Tegmina hyaline. Mindanao, North Borneo, broad, lateral fasciae 8 Sumba 18 — Abdomen monochromous, or with faint fas- — Tegmina orange-greenish opaque. Sumbawa ciae 9 sum-bawensis 8. Body dark ochreous; tegmen areas finely 18. Large species (body length: 17.9— 20.9 mm); stippled; ovipositor sheath 0.4 —0.41 X as ovipositor sheath 0.22 —0.26 X as long as long as abdomen. South Sulawesi abdomen. Mindanao, North Borneo lieftincki fatiloqua — Body virescent to orange-yellowish, with — Small species (body length: 14.9— 16.1 mm); light pale ochreous; tegmen areas patchy ovipositor sheath 0.32 —0.33 X as long as stippled; ovipositor sheath 0.25 X as long abdomen. Sumba roehli as abdomen. South Sulawesi tincta 19. (5: body ochreous; tegmen areas unspotted,

9. Body and tegmina green; tegmen areas with veins with some markings; genitalia as in a faint stippling. North Sulawesi figs. 61 —64, 66. $: body brownish-ochreous; consanguinea abdomen with three longitudinal fasciae: one

— Body and tegmina variably coloured; tegmen - dorsal medial fascia and two broad lateral areas with a distinct stippling 10 fasciae. South Sulawesi parvula 10. Pigmentation along tegmen veins distinct — $: body green; 8th apical and 4th ulnar areas and dark-coloured. Papua New Guinea, of tegmen spotted; genitalia as in figs. Torres Strait Islands, Cape York Peninsula 68— 71, 73. $: body dark-brown, abdomen paradoxa usually without fasciae. North Sulawesi .... — Pigmentation along tegmen veins weak. wallacei South Sulawesi 11 11. Large species (body length: 25.9 mm); stip- The material studied contained several un- pling in tegmen areas patchy. South Sulawesi identified males and females belonging to the fruhstorferi fatiloqua group. Three females with pigmented — Medium-sized species (body length: tegmina, viz., two from North Sulawesi (a small 17.3 — 19-1 mm); stippling in tegmen areas one from Edwards Camp in the Dumoga-Bone

relatively fine. South Sulawesi . . . maculosa N.P. (RMNH), and a large one from Gunung 12. Males 13 Muajat, east of Kotamobagu (BMNH)), and a — Females 17 small one from South Sulawesi (Watampone 13. Tegmina hyaline 14 (MZB)); two females with opaque tegmina, one — Tegmina opaque; tymbal with 17 — 19 pairs from Flores (SMD) and one from Sumbawa of alternating ridges; genitalia as in figs. (NRS); a large female with hyaline tegmina from 18— 21. Sumbawa sumbawensis Sangihe island (RMNH). Furthermore, two 14. Aedeagus long and slender (figs. 6— 8, 23, males (representing one species) with pig- 26,28) 15 mented tegmina from North Sulawesi (Labua- — Aedeagus short and sturdy (figs. 12 — 14, nika (ZMA)); two males with hyaline tegmina, 29—31) 16 one of which from South Sulawesi (Assumpati 15. Body size: 17.9 —21.8 mm; tymbal with 15 (BMNH)), and one from North Sulawesi (Ton- pairs of alternating ridges; genitalia as in sealama (MCZ)); three males with opaque teg- figs. 6— 9. Mindanao, North Borneo mina, one from Lombok (Sapir (NHMW)), and .fatiloqua two males (representing one species) from Sum- — Body size: 22.3 —24.2 mm; tymbal usually bawa (SMD; BMNH). with 14 pairs of alternating ridges; genitalia as in figs. 23, 25, 26, 28. South Sulawesi The Lembeja fatiloqua group maculosa Head triangularly to obconically protruding in

16. Body orange-yellowish and large (body dorsal view. Antennal segment 1 long. Male M. R. DE Jong; Oriental Prasiini 181

Figs. 1 — 5. The L. fatiloqua group: 1, male abdomen, ventral view, Lembeja tmcta; 2, head and pronotum, lateral view, L. lieftincki; 3, left tymbal, lateral view, L. fatiloqua; 4, female femur, lateral view, L. fattloqua; 5, right tegmen and wing, L. fatiloqua. 182 Tijdschrift voor Entomologie, deel 130, 1987 opercula small, not covering tymbal cavities. Teg- together. Thorax and head together in males mina (sub)hyaline or opaque, with or without 0.63—0.82 X, in females 0.88—1.08 X as long as spots along the veins and inside the tegmen cells. abdomen. Greatest width of body in males at the

Abdomen in males moderately to strongly cari- level of abdominal segment 2 and 3, in females nate along tergites 3 — 7. Tergites strongly folded at lateral angles of pronotum collar. laterally, forming a ridge on each side of the Head. — Second antennal segment slightly sternites. Intersegmental membranes sometimes darker than 1st. Eye small, in dorsal view clearly exposed. Tergite 1 bulbous, usually with 0.49—0.57 X as wide as width of vertex between two short lateroproximal flaps, and provided eyes. Ocelli raised. Distance between lateral ocelli with a moderately to strongly developed, medial 1.0 — 1.31 X distance between lateral ocellus and longitudinal dent. Sternite 1 triangular and small. eye. Length of head 1.33 — 1.49 X as long as width Folded membranes and mirrors medium-sized, of vertex between eyes; width of head 1.98 —2.13 sometimes in an angle of 90° with one another. X as wide as width of vertex between eyes. Trans- Tymbals with 12 —20 long ridges, alternating verse ridges in the same colour as underside of usually with an equal number of short ridges. postclypeus. Rostrum with dark apex reaching Abdomen in females slender, carinate dorsally; in middle trochanter. lateral view convex from tergite 3 —8. Ovipositor Thorax. — Unicoloured. Fissures on prono- sheath extending beyond caudodorsal beak. Lat- tum not prominent. Pronotum collar 1.96 —2.47 eral lobes of pygofer usually short and concave on X as wide as length of head, 1.41 — 1.61 X as wide the outer surface. Claspers vary from short and as width of head head including eyes. Mesonotum curved to long and elongate; ventrally usually sometimes slightly darker than pronotum. Pa- concave. Median uncus part usually slightly com- ramedian obconical spots recognizable; lateral pressed, sometimes enlarged to a short to some- ones usually consisting of some dark coloured what longer tube, just above the claspers. Aedea- spots. gus long and slender to short and more sturdy; Legs. — Same colour as underside of body. apex rounded, or with two short to long lobes. Basal spine of fore femur blunt, provided with a Only one species (L consanguinea n.sp.) pro- small subapical spine. Middle and apical spines vided with an unsclerotized, dorsal aedeagal ap- acutely pointed. pendage. Tegmina and wings. — Tegmina and wings hyaline; venation whitish to virescent. In teg- Lembeja fatiloqua (Stal, 1870) mina transverse vein of 2nd ulnar area extending (figs. 6— 11,74, 75; map 1) into 3rd one. Corial fold recognizable. Node in Distant, xiv, Prasia fatiloqua Stal, 1870; 718; 1892: M3+4 present. Cuj and Aj forming a small triangle 146, PI. 6 figs. 2, 2a—b; Breddin, 1901: 153; Dis- at tegmen border. Third ulnar area 0.99— 1.15 X tant, 1905: 279; Distant, 1906: 184; Distant, 1909: as long as 1st one; 4th ulnar area 0.91 — 1.07 X as 394; Kato, 1932: 184; de Jong, 1985: 166; de Jong, 0.78 1986: 141. long as radial area. Third apical area —0.95 X Lembeja fatiloqua; Horvath, 1912: 609; Myers, 1928: as long as 4th one. Cuj and Aj in wings fused at 392, 460; Myers, 1929: 52, text fig. 24; Metcalf, 81 —98% from their origin. 1963: 430; de Jong, 1982: 182; de Jong, 1986: 142. Male: Operculum. — Small, more or less sickle- shaped; not reaching folded membrane. Long and The following reference was found to relate to pointed meracanthus broad at base. another species: Prasia fatiloqua; Lallemand, Abdomen. — Dorsally slightly carinate. Pale 1935: 677 (Sumba specimens belong to Lembeja ochreous to sometimes dark-brown. Tergite 1 with roeÂ//' Schmidt, 1925). two latero-proximal flaps. Folded membrane almost parallel with underside of thorax. Mirrors The male holotype of the species has been medium-sized. Triangular structure between the studied at the NRS, by kind permission of Dr P. folded membranes fairly large. Lindskog, in order to establish the identity of the Tymbals. — Medium-sized, provided with 15 species. long and 15 intercalary short ridges. Genitalia. — Lateral lobes small, convex inner Description. surface somewhat swollen, not reaching beyond Body pale ochreous, sometimes greenish; ab- anal valves. Caudodorsal beak short, only very domen sometimes slightly darker than remain- slightly pointed, reaching just beyond anal valves. ing part of body. Head and pronotum together Sturdy claspers short, curved and pointed apically. 0.92 — 1.07 X as long as meso- and metanotum Median uncus part narrow. Aedeagus long and M. R. DE Jong: Oriental Prasiini 183

Figs. 6— 11. Lembeja fatiloqua; 6— 10, $; 11, Ç. 6, pygofer, lateral view, Basilan island; 7, apex of aedeagus, laterodorsal view, Basilan island; 8, pygofer, ventrolateral view, Basilan island; 9, clasper, lateral view, Basilan island; 10, tergite 1, dorsal view, Davao; 11, sternite 7, ventral view, Davao.

slender. Apex of aedeagus with two short, hardly Abdomen. — Unicoloured; ovipositor sheath pointed flaps in the shape of a serpent's tongue. 0.22 — 0.25 X as long as abdomen. Female: Operculum. — Small, somewhat sickle- Measurements of the material studied: body shaped. length $: 17.9—21.8 mm, x = 19.7 ± 1.2 mm, $: 184 Tijdschrift voor Entomologie, deel 130, 1987

/ M. R. DE Jong: Oriental Prasiini 185

Figs. 12 — 17. Lembeja roehli; 12 — 16, $\ 17, $. 12, 13, pygofer, 12, ventrolateral view, Waingapu, 13, lateral view,

Waingapu; 14, apex of aedeagus, lateral view, holotype; 15, clasper, lateral view, Waingapu; 16, tergite 1, dorsal view, Waingapu; 17, sternite 7, ventral view, Meloio.

Tegmina and wings. — Tegmina hyaline; ve- folded membrane. Long and pointed meracanthus nation yellowish to whitish. Transverse vein of broad at base. 2nd ulnar area hardly extending into 3rd one. Abdomen. — Dorsally moderately carinate Corial fold hardly recognizable. Small node in from segment 4 — 7. Usually pale ochreous. Ter- Mj+4 present. Third ulnar area 0.91 — 1.1 X as long gite 1 broad. Mirrors relatively small. Abdomen, as 1st one; 4th ulnar area 0.85 — 1.04 X as long as when inflated, showing clearly the intersegmental radial area. Third apical area 0.79—0.98 X as long membranes; in normal position, underside curved as 4th one. Wings hyaline. Fusion of Cuj and A, upwards to the posterior in lateral view. at 81 —88% from their origin. Tymbals. — Provided with 12 long and 12 short Male: Operculum. —Small, round; just reaching intercalary ridges. 186 Tijdschrift voor Entomologie, deel 130, 1987

O roehli # SU mbawensis

116° 118° 120°

Map 2. Distributions of L. roehli and L. sumbawensis.

Genitalia. — Lateral lobes very small; hardly Lembeja roehli (MW); Kananggar, 700 m, E. swollen on convex inner surface, not reaching Soemba, v. 1925, Dammermann, 1 Q (MZB); La- beyond anal valves. Caudodorsal beak much less luku, E. Sumba, 4. vii, Dr Bühler & Dr Sutter, pointed than in L. fatiloqua. Claspers smaller, but Baeturia exhausta Guérin, det. V. Lallemand 1951, generally shaped as in L. fatiloqua; apically hardly 1 (5 (NBM); Laora, 100 m, N.W. Sumba, iv.l925, pointed. Median uncus part narrow. Aedeagus Dammermann, 3 $ (MZB); Melolo, E. Sumba, short and sturdy. Somewhat swollen apex of ae- 29.V.1949, Dr Bühler & Dr Sutter, Muda obtusa deagus almost round. Walk., det. H. Synave 1951, 1 9(NBM); Prai Ja- Female: Operculum. — Small. wang, E. Sumba, Rende Wai, 12.vi.l949, Dr Bühler Abdomen. — Unicoloured; ovipositor sheath & Dr Sutter, Baeturia exhausta Guérin, 1 $ 0.32 —0.33 X as long as abdomen. (RMNH); Waingapu, Ì.Ì9Ò2, Prasia fatiloqua Stal, Measurements of the material studied: body Lembeja fatiloqua Stal, 1 $ (NBM). Specimen

1 length (5 : 18.8—22.9 mm, x = 20.7 ± 1.7 mm, $: without labels: S (ZBM). 14.9— 16.1 mm; width of pronotum collar $: 5.3—6.1 mm, x = 5.6 ± 0.3 mm, $: 4.8—5.3 mm; Remarks. tegmen length $: 24.1—25.3 mm, x = 24.7 + 0.5 The taxonomie position of L. roehli within the mm, $: 20.6—22.0 mm. species-group is unclear. Though the species is much like L. fatiloqua, because of the coloration Distribution. — Sumba island (map 2). of the tegmina, some features suggest a close Material examined. — Indonesia, Sumba: affinity to L. paradoxa, viz. the telescoping of the Grelak, "Sumba/Grelak" (print, black cadre), "Ty- abdomen in the males, and the low number of pus" (print, red label, black cadre), "Lembeja/ ridges on the tymbal organ. The male genitalia, Roehli Schmidt/Edm. Schmidt/ 5'. determ. 1925" however, resemble those of L. sum-bawensis (partly print, partly handwritten) $ holotype of n.sp., and, to a certain extent, those of L. tincta. M. R. DE Jong : Oriental Prasiini 187

#1% /i%

Figs. 18— 22. Lernbeja sumbawensis; 18 — 21, holotype; 22, $ paratype. 18, 19, pygofer, 18, ventrolateral view, 19, lateral view; 20, apex of aedeagus, lateral view; 21, clasper, lateral view; 22, sternite 7, ventral view.

Lembeja sumbawensis n.sp. between eyes. Distance between lateral ocelli (figs. 18—22, map 2) 0.78— 1.29 X distance between lateral ocellus and eye. Length of head 1 .24 — 1 .4 1 X as long as width The description is made in comparison with L. of vertex between eyes; width of head 1.92 —2.11 fat. X as wide as width of vertex between eyes. Ros- trum with slightly darker apex. Description. Thorax. — Unicoloured. Fissures on prono- Body green to ochreous. Head and pronotum tum somewhat deeper than in L.fatiloqua. Pron- together 0.94— 1.07 X as long as meso- and me- otum collar 2.34 — 2.89 X as wide as length of tanotum together. Head and thorax together in head, 1.57 — 1.65 X as wide as width of head. males 0.76—0.79 X, in females 1.03 X as long as Obconical areas on mesonotum not discernable. abdomen. Legs. — Same colour as underside of body. Head. — Antennae unicoloured. Eye in dorsal Basal, pointed spine on fore femora relatively view 0.46—0.56 X as wide as width of vertex in shorter than in L. fatiloqua; very broad at base. 188 Tijdschrift voor Entomologie, deel 130, 1987

Tegmina and wings. — Tegmina opaque, 3, 3a—b; Karsch, 1890: 190; Mac-Lachlan, 1891: greenish to ochreous. Venation whitish to green- 320; Jacobi, 1903: 13. Lembeja maculosa; Distant, 1892: xiv, 147 (in par- ish. Transverse vein of 2nd ulnar area extending tim); Distant, 1897: 371; Jacobi, 1903: 12, 13; Dis- only shortly into 3rd ulnar area. Corial fold not tant, 1905: 279; Distant, 1906: 184; Kate, 1932: 0.88 as recognizable. Third ulnar area — 1.03 X 189; Metcalf, 1963: 428, 431; De Jong, 1986: 141, long as 1st one; 4th ulnar area 0.83 — 1.0 X as 142. long as radial area. Third apical area 0.83 —0.9 X Prasia maculosa; Breddin, 1901: 27, 113, 153. as long as 4th one. Wings subhyaline, fusion of The following reference was found to pertain Cu2 and Aj at 75 —85% from their origins. to L. distanti de Jong, 1986: Distant, 1892: xiv: Small, as in L. fatiloqua. Male: Operculum. — 147 (in partim), pi. 7, 13a—b. Meracanthus long and pointed.

Abdomen. — On the whole as in L. fatiloqua, This species is described in comparison with L. but somewhat more carinate. Lateral flaps of fatiloqua. The males and females of L. maculosa

tergite 1 less pointed. are differently coloured. Tymbal. — Provided with 17 — 19 long ridges, alternating with an equal number of short ridges. Description. Genitalia. — Lateral lobes very small, not ex- Body greenish- to yellowish-ochraceous in ma- tending beyond anal valves. Caudodorsal beak les, brownish-ochreous in females (holotype short and rounded. Claspers relatively small, green). Males larger than in L. fatiloqua. Females hardly pointed. Aedeagus short and sturdy, with spotted tegmina, tegmina in males hyaline. slightly incised at apex. Head and pronotum together 0.9— 1.08 X as Female: Operculum. — Very small, as in L. long as meso- and metanotum together. Head fatiloqua. and thorax together in males 0.64—0.76 X, in Abdomen. — Unicoloured; ovipositor sheath females 1.01 — 1.21 X as long as abdomen. 0.22 X as long as abdomen. Head. — Antennae unicoloured. Eye in dorsal Measurements of the types: body length $: view 0.47 —0.55 X as wide as width of vertex 17.9 — 19.5 mm, $: 14.9 mm; width of pronotum between eyes. Distance between lateral ocelli collar (5': 5.4 — 6.0 mm, $: 5.3 mm; tegmen length 0.72 — 1.0 X distance between ocellus and eye. $: 22.8—23.0 mm, ?: 22.7 mm. Length of head 1.28 — 1.44 X as long as width of vertex between eyes; width of head 1.84 — 2.1 X Distribution. — Sumbawa island (map 2). as wide as width of vertex between eyes. Dark Types. — Indonesia, Sulawesi: Holotype: apex of rostrum reaching middle coxae. "Soembawa/CoU. Noualhier 1898" (print), "Dis- Thorax. — Fissures on pronotum not very tant Coll./ 191 1—383" (print) 1 S holotype of deep. Pronotum collar 2.17 —2.57 X as wide as Lembeja sumhawensis (BMNH). Paratypes: length of head, 1.5 — 1.73 X as wide as width of Sumbawa, Tambora, W. Doherty 1903 — 31, 1 (5 head. Females with some dark brown patches on (BMNH). Sumbawa, without precize locality: ex areas between fissures and between fissures and coll. Fruhstorfer 1 $ (NHMW); same data but: pronotum collar; central fascia slightly darker Lembeja foliata (Walk) Jacobi det., coll. Breddin coloured than remaining part of pronotum. Ob- 1 (5 (DEI); coUectio Haglund, 266—84, 1 $ conical areas on mesonotum hardly discernable. (NRS). Female with median longitudinal dark brown Etymology. — The species is named after stripe on cruciform elevation. Sumbawa island. Legs. — In males with same colour as underside body; in females with darker patches. Basal Remarks. spine on fore femora long and pointed. Some- The species resembles, as far as its genitalia are times a small 4th, apically situated spine present. concerned, L. roehli very much. Three other re- Tegmina and wings. — Tegmina hyaline in lated species with opaque tegmina are recognized males, in females spotted regularly along and in in material from the Lesser Sunda Islands, but veins and faintly spotted within tegmen cells. description of these three species has not been Transverse vein of 2nd ulnar area extending into undertaken, because the material is too poor. 3rd ulnar area. Corial fold clearly distinct. Costa and veins whitish tinged in males, pale-ochreous Lembeja maculosa (Distant, 1883) in females. Third ulnar area 0.88— 1.08 X as long (figs. 23—28; map 3) as 1st one; 4th ulnar area 0.91 — 1.16 X as long Perissoneura maculosa Distant, 1883: 190, pi. 25, figs. as radial area. Third apical area 0.8 — 1.01 X as M. R. DE Jong: Oriental Prasiini 189

Figs. 23 — 28. Lembeja maculosa; 23 — 26, 28, $ Patunuang; 27, Ç Patunuang. 23, pygofer, ventrolateral view; 24, tergite 1, dorsal view; 25, clasper, lateral view; 26, apex of aedeagus, laterodorsal view; 27, sternite 7, ventral view; 28, pygofer, lateral view. 190 Tijdschrift voor Entomologie, deel 130, 1987

long as 4th one. Wings hyaline, extreme base and 1900, 12029, 1 9 (SMN); same data but: L. tincta veins whitish in males, red in females. Cu, and A, Dist, Prof. Dr A. Jacobi determ., H. Fruhstorfer fused at 70—89% from their origins. vend. 30.ix.1897, 1 S (ZIM); Samanga, S. Cele- Male: Operculum. — Small, hardly reaching bes, Nov. 1895, H. Fruhstorfer, Lembeja m,acu- folded membrane. Meracanthus long and slen- losa Dist. 9, 1 9 (SMD). Sulawesi, without fur- der. ther indication: "maculosa/Dht./iy^e" Abdomen. — Yellow to light brown; carinate. (handwritten), " maculosa/ (Dist)" (yellow label, In lateral view relatively larger than in L. fatilo- handwritten), "A.B. Meyer/Celebes 1871" qua. Mirrors relatively smaller than in L. fatilo- (print, yellow label), "coll. A. Jacobi" (print), "363" qua. Tergite 1 with relatively broader flaps. (handwritten) 9 ^oiotype oi Perissoneura Tymbals. — Provided with usually 14 (some- maculosa (SMD). times 15) long ridges with an equal number of short ridges. Remarks.

Genitalia. — Lateral lobes of pygofer broad, L. maculosa is very similar to L. fatiloqua in short and flat, concave on outer and convex on respect to the male genitalia. Furthermore, the inner surface. Caudodorsal beak short, bluntly tymbal in both species is provided with almost rounded. Claspers elongate, hardly curved and the same number of alternating ridges. A feature only slightly pointed. Median uncus part com- of distinction is the sexual dimorphism in color- pressed and narrow. Aedeagus long and slender, ation of body and tegmina of L. m-aculosa, which its two apical short lobes acutely pointed. is also found in L. tincta. Female: Operculum. — As in L. fatiloqua. Abdomen. — Darker coloured than remaining part of body. Three, usually faint, longitudinal Lembeja tincta (Distant, 1909) n. comb, fasciae, a dorsal, medial narrow one and two (figs. 29—34, 76, 77; map 3) lateral with fairly broad ones. Segment 9 two Prasia tincta Distant, 1909: 393; Gaedike, 1971: 319; broad longitudinal, dark coloured lateral stripes, de Jong, 1982; 182; Duffels & v. d. Laan, 1985: 313; just uniting in front of caudodorsal beak. Ovipo- de Jong, 1985: 166; de Jong, 1986: 141. sitor sheath 0.19—0.24 X as long as abdomen. Lembeja tincta; de Jong, 1986: 142. Measurements of the material studied: body length $: 22.3—24.2 mm, x = 23.2 ± 0.8 mm, This species is described in comparison with L. $: 17.3—19.1 mm, x = 18.4 + 0.6 mm; width of maculosa. The different coloration of males and pronotum collar $: 5.4 — 6.7 mm, x = 6.3 ± 0.4 females reminds that of L. maculosa. mm, $ = 6.1 — 6.7 mm, x = 6.4 ± 0.4 mm; tegmen length $: 24.9—27.9 mm, x = 26.6 ± 0.8 Description. mm, $: 25.2—28.5 mm, x = 26.7 ± 1.0 mm. Body virescent to orange-yellowish in males, brownish-ochreous to orange-yellowish in fe- Distribution. — South Sulawesi (map 3). male. Tegmina unspotted in males, spotted in Material examined. — Indonesia, Sulawesi: female. Head and pronotum together 0.86— 1.06 Makassar (= Ujung Pandang), leg. Dres. Sarasin, X as long as meso- and metanotum together. coll. Breddin, 1 ß (DEI); same locality and collec- Head and thorax together in males 0.61 —0.79 X, tors but: Lembeja fatiloqua (Stâl),Jacobi det., 1 $ in female 0.9 X as long as abdomen. (DEI); same locality and collectors but: fatiloqua Head. — Antennae dark-coloured from 2nd Stal, coll. A. Jacobi 1910—6, 1 $ (SMD); same segment to apex. Eye large, in dorsal view locality but: R Muir, Dec. 1908, 1 $ (BISH); 0.54—0.64 X as wide as width of vertex between Patunuang, S. Celebes, Jan. 1896, H. Fruhstorfer, eyes. Area between lateral ocelli brownish in fe- 4 (5 3 $ (NHMW); same data but: 1909—21, male only. Distance between lateral ocelli Type, 1 $ (BMNH); same data but: maculosa 0.74— 1.0 X distance between lateral ocellus and Dist., Type, Distant coll. 1911—383, 1 $ eye. Length of head 1.26— 1.43 X as long as width (BMNH); same data but: coll. A. Jacobi, 1910—6, of vertex between eyes; width of head 2.09—2.27 1 $ (SMD); same data but: Lembeja maculosa X as wide as width of vertex between eyes. Trans- Dist. 9; 1 9 (TMB); same data but: Prasia faticina verse ridges in the ground-colour in male, some- Dist, Cotypus!, tincta Dist. coll. A. Jacobi what darker than ground-colour, especially near 1910—6, 1 $ (SMD); same data but: Prasia sp.. base of rostrum, in female. Rostrum with black Dist. coll. 191 1—383, 7, 1 $ (BMNH); same data apex reaching middle trochanter. but: Cystosoma, C. paradoxa S. Celebes, Frühst. Thorax. — Pronotum collar 2.47 —2.84 X as M. R. deJong; Oriental Prasiini 191

Map. 3. Distributions of L. consanguinea, L. fruhstorferi, L. lieftincki, L. maculosa, L. sanguinolenta, L. tincta, L. parvula and L. wallacei. 192 Tijdschrift voor Entomologie, deel 130, 1987

Sj M. R. DE JONG: Oriental Prasiini 193

34 \

/ \

Figs. 31 — 34. Lembeja tincta, 31 — 33, (3; 34, Ç. 31, apex of aedeagus, laterodorsal view, Bua Kraeng; 32, tergite 1, dorsal view, Lompobatang; 33, clasper, lateral view. Bua Kraeng; 34, sternite 7, ventral view, Bua Kraeng.

21" slightly swollen; not reaching beyond anal val- Fruhstorfer" (print, black cadre), "1909— ves; hardly concave at the outer and convex on (print), "Type" (print, round label, red edged), inner surface. Caudodorsal beak medium-sized, "syn-type" (print, round label, blue edged), almost pointed. Claspers short, sturdy and cur- "Prasid/tincta/'Dist./Type" (handwritten), ved. Aedeagus medium-sized, subapically strong- "Brit. Mus." (print) $ holotype of Prasia tincta ly curved; apex with two short pointed flaps. (BMNH); same locality and collector but: syn- Female: Operculum. — Small, darker; edges type, Prasia tincta, 2 $ paratypes of Prasia tincta sometimes lighter. Meracanthus long and slen- (BMNH); same locality and collector, 1 (5 1 $ der. (BMNH) 1 S (NHMW); same locality and col- Abdomen. — Three dark-coloured longitudi- lector but: syntypus, Lembeja tincta (Dist) Jacobi nal fasciae situated as in L. maculosa, but more det., 1 $ (DEI); same locality and collector but: conspicuous. Coloration of segment 9 more con- Lembeja fruhstorferi Dist., 1 $ (TMB); Lompa- spicuous than in L. maculosa. Ovipositor sheath, Battau (= Lompobatang), 3000 ' Marz 1896, H. with dark apex, 0.25 X as long as abdomen. Fruhstorfer, vend. 30.ix.l897, Prof. Dr A. Jacobi Measurements of the material studied: body determ., 1 S (ZIM); same locality, 1600 m, length $: 22.8—26.2 mm, x = 24.7 + 0.9 mm vii.1936, L.J. Toxopeus, 5 S (MZB). $: 23.2 mm; width of pronotum collar $ 6.1 — 7.7 mm, x = 7.2 ± 0.3 mm, $: 7.1 mm tegmen length $: 32.5 — 34.3 mm, x = 33.6 + 0.6 Remarks. mm, $: 33.7 mm. L. tincta and L. maculosa are characterized by sexual dimorphism in the coloration of body and Distribution. — South Sulawesi (map 3). tegmina. The genitalia of L. tincta and especially Material examined. — Indonesia, Sulawesi: "S. the sturdy aedeagus, are more alike those found CeIebes/Bua-Kraeng/5000' Febr. 1896/H. in the species of the Lesser Sunda Islands. 194 Tijdschrift voor Entomologie, deel 130, 1987

M^ét ,jA^'\

Figs. 35 — 37. Lembeja paradoxa; 35,36,(5 Port Moresby; 37,9 Port Moresby. 35, tergile 1, dorsal view; 36, clasper, lateral view; 37, sternite 7, ventral view.

Lembeja paradoxa (Karsch, 1890) and thorax together in females 0.9 — 1.17 X as (figs. 35-37) long as abdomen, in males very variable because Perissoneura paradoxa Karsch, 1890: 191; Mac-Lach- of the telescoping abdomen, and therefore unre- lan, 1891; 320. liable. Lembeja paradoxa; Distant, 1892: xiv, 148; Distant Eye in dorsal view 0.40—0.49 X as wide as 1903: 13; Distant, 1906: 182: 1897b: 382;Jacobi, width of vertex between eyes. Distance between Schmidt, 1925: 43; Kato, 1932: 189; Metcalf, 1963 lateral ocelli 0.52 — 1.05 X distance between late- 432; de Jong, 1982: 175—179, 182—184, figs. ral ocellus and eye. Length of head 1.23 1.35 X 1_8, 17—19, 22—23; de Jong, 1986: 141, 142. — as long as width of vertex between eyes; width of Prasia paradoxa; Breddin, 1901: 153. Perissoneura acutipennis Karsch, 1890; 192; Mac- head 1.81 — 1.98 X as wide as width of vertex Lachlan, 1891: 320. between eyes. Prasia acutipennis; Breddin, 1901; 153- Pronotum collar 2.13 —2.53 X as wide as 1906: Lembeja acutipennis; Jacobi, 1903: 13; Distant, length of head, 1.42 — 1.68 X as wide as width of Kickaldy, 1907: 309; Schmidt, 1925; 43; 184; head. Burns, 1957: 669; Metcalf, 1963: 429; de Jong, Third ulnar area 1.19 — L65 X as long as 1st 1982: 175 — 177 (in synonymy of Lembeja para- one; 4th ulnar area 0.95 — 1.10 X as long as radial Lembeja brunneosa Distant, 1910: 418; Distant, area. Third apical area 0.89— 1.16 X as long as 1913: 601; Ashton, 1914: 356; Burns, 1957: 669, 4th one. Fusion of Cu2 and A, in wings at 670 (equals Lembeja australis and Prasia viticollis 71 —91% from their origins. (sic)); Metcalf, 1963: 430; Woodward, Evans and Tymbal provided with 13 — 14 long ridges, al- Eastop, 1970: 413; Moulds, 1975; 251—254, figs. ternating with an equal number of short interca- 1—6; de Jong, 1982: 175—177 (in synonymy of lary ridges. Lembeja paradoxa). Lembeja australis Ashton, 1912b: 77, pi. 7 fig. 3; Ovipositor sheath 0.13 —0.19 X as long as ab- Distant, 1913:601 Cm synonymy oi Lembeja brun- domen. neosa); Metcalf, 1963: 430 ditto; de Jong, 1982: 175—177 ditto. Lembeja fruhstorferi Distant, 1897 (map 3) No new material has been studied since the Lefnbeja fruhstorferi 'Distant, 1897 (part.): 371; Ja- redescription of Lembeja paradoxa (see de Jong, cobi, 1903; 13; Distant, 1906: 184; Kato, 1932: 189; 1982). In addition to the redescription, I add here de Jong, 1986: 141, 142. Prasia fruhstorferi; Breddin, 1901: 27, 153. all relevant measurements and some additional figures in order to facilitate comparison with other species of the jatiloqua group. The following reference was found to pertain to L. distanti àe ]ong, 1986: Distant, 1897: 371. Head and pronotum together 0.97 — 1.14 X as long as meso- and metanotum together. Head A very short description, made in comparison 7

M. R. DE JONG: Oriental Prasiini 195

with L. fdtiloqua, is given here, as the species is Lembeja sanguinolenta Distant, 1909 only known from its female holotype (figs. 38—41, map 3) Lembeja sanguinolenta Distant, 1909: 394; de Jong, Description of the female. 1982; 182; de Jong, 1986: 141, 142. Body dull-brown. Head and pronotum together 0.96 X as long as meso- and metanotum The description is based upon the only known together. Head and thorax together 0.92 X as specimen of the species, the male holotype. long as abdomen. Head. — Brownish ochreous. Eye in dorsal Description of the male. view 0.56 X as wide as width of vertex between Body for its greater part dull red. Head and eyes. Distance between lateral ocelli 0.87 X dis- body pilose. Head and pronotum together 0.91 X tance between lateral ocellus and eye. Length of as long as meso- and metanotum together. Head head 1.41 X as long as width of vertex between and thorax together 0.89 X as long as abdomen. eyes; width of head 2.12 X as wide as width of Greatest width of the body across the 2nd and 3rd vertex. Postclypeus and its transverse ridges uni- abdominal segment. form. Head. — Red, transverse ridges in the same Thorax. — Irregularly brown and ochreous. colour as underside. Eye large, in dorsal view 0.68 Pronotum with some irregular patches. Pro- X as wide as width of vertex between eyes. Dis- notum collar with two small medial light tance between lateral ocelli 0.92 X distance ocel- ochreous patches. Pronotum collar 2.54 X as lus and eye. Length of head 1.65 X as long as wide as length of head, 1.7 X as wide as width of width of vertex between eyes; width of head 2.35 head. Mesonotum with irregular patches in ob- X as wide as width of vertex between eyes. Ros- conical areas. trum with only slightly darker apex reaching Legs. — Same colour as underside of body. intermediate coxae. Basal spine on fore femora blunt and short. Thorax. — Median part of pronotum, includ- Tegmina and wings. — Tegmina subhyaline, ing central fascia and pronotum collar, red. Rest spotted along and in veins and in tegmen cells. of pronotum light ochreous with some brown Costa and veins light brownish. Extreme base patches. Pronotum collar 2.68 X as wide as light brownish. Transverse vein of 2nd ulnar area length of head, 1.88 X as wide as width of head. extending into 3rd one. Corial fold brown, in 4th Fissures fairly deep. Obconical areas on meso- ulnar area accompanied by a large brown patch. notum patchy. Third ulnar area 1.12 X as long as 1st one; 4th Legs. — Same colour as thorax. Basal spine of ulnar area 0.84 X as long as radial area. Third fore femur blunt. apical area 0.88 X as long as 4th one. Wings Tegmina and wings. — Tegmina subhyaline, hyaline, veins whitish tinged. Cuj and Aj fused at on the whole spotted with red. Costa and veins 92% from their origins. red. Transverse vein of 2nd ulnar area extending Operculum. — Very small. Small meracanthus into 3rd. Corial fold in 3rd ulnar area somewhat broad at base, just a little longer than operculum. darker, in 4th one indicated by a large red patch. Abdomen. — Brownish, underside a little Third ulnar area 1.02 X as long as 1st one; 4th paler. Hind margins of tergites 3 — 7 dull red. ulnar area 0.84 X as long as radial one. Third Ovipositor sheath 0.21 X as long as abdomen. apical area 0.97 X as long as 4th one. Wings subhyaline. Extreme base white with red. Fusion Measurements of the holotype: body length of Cuj and Aj at 78% from their origins. width 24.5 mm; of pronotum collar: 8.0 mm; Operculum. — Hairy. Hardly reaching folded tegmen length: 35.4 mm. membrane. Large mecanthus broad at base, pointed apically. Distribution. — South Sulawesi (map 3). Abdomen. — Reddish ochreous, tergite 3 — Material examined. — Indonesia, Sulawesi: "S. with red hind edges. Tergite 1 with small latero- Celebes/Bua-Kraeng/5000' Febr. 1896/H. proximal flaps. Mirrors large. Fruhstorfer" (print, black cadre), "fruhstorferi/ Tymbals. — Provided with 20 long and 20 Dist." (handwritten), "Type" (print, round label, short intercalary ridges. red edged), "Distant Coll./ 191 1—383" (print) $ Genitalia. — Pygofer large. Flat lateral lobes holotype of Lembeja fruhstorferi (BMNH). medium-sized, not reaching beyond anal valves. Remarks. Caudodorsal beak medium-sized, hardly pointed See L. sanguinolenta. apically. Edge of pygofer just below each lateral —

196 Tijdschrift voor Entomologie, deel 130, 1987

lobe somewhat protruding. Uncus narrow, a Legs. — Generally same colour as underside slight onset of a tube-like structure, just above thorax but with some dark patches, especially claspers, discernable. Curved claspers relatively basally and apically on tibiae and tarsi. Spines on small. Apex of sturdy aedeagus with two long, fore femora dark. more or less toothed, flaps. Tegmina and wings. — Tegmina hyaline, spot- Measurements of the holotype: body length: ted in and along veins in a regular pattern, and 24.6 mm; width of pronotum collar: 8.8 mm; in tegmen cells, especially in apical area 8. Corial tegmen length: 36.6 mm. fold recognizable by heavy pigmentation, especially in 4th ulnar area. Basal area infuscated.

Distribution. — South Sulawesi (map 3). Extreme base reddish. Venation green to red. Third ulnar area 0.97 — 1.0 X as long as 1st ulnar Material examined. — Indonesia, Sulawesi: "S. area; 4th ulnar area 1.0— 1.08 X as long as radial Celebes/Bua-Kraeng/5000' Febr. 1896/H. area. Third apical area 0.85 —0.96 X as long as Fruhstorfer" (print, black cadre), "Lembeja/san- 4th one. Wings hyaline. Fusion of the Cuj and Aj guinolenta/Type Dist." (handwritten), "Type" veins at 72 —84% from their origins. (round label, red edged, print), "1909—21" Male: Operculum. — Small. Meracanthus long (print) (5 holotype of Lembeja sanguinolenta and pointed. (BMNH). Abdomen. — Sometimes darker coloured. Mirrors large. Folded membranes nearly contin-

uous with underside of thorax. Tergite 1 laterally Remarks. slightly depressed near the small lateroproximal L.fruhstorferi and L. sanguinolenta have about flaps. the same body-size, but some body ratios are very Tymbals. — Provided with 19 long ridges, al- different. ternating with an equal number of short intercalary ridges. Lembeja lieftincki n.sp. Genitalia. — Lateral lobes of pygofer sturdy. (figs. 42—48, 78, 79; map 3) Lateral surface of pygofer with a short ridge-like

The description is based upon two males and structure running downwards from each lateral two females from South Sulawesi. lobe. Caudodorsal beak relatively long, apically rounded. Claspers short and curved, hardly Description. pointed. Median uncus part enlarged to a tube- Body green to dark ochreous. Head and pron- like structure. Apex of aedeagus with two re- otum together 0.96— 1.06 X as long as meso- and curved, long flaps; dorsally provided with a long metanotum. Head and thorax together in males subapical incision. 0.78—0.79 X, in females 0.84—0.95 X as long as Female: Operculum. — Small, but relatively abdomen. large compared to other species of the species- Head. — Green to reddish-brown. Antennae group. usually darker coloured. Area between lateral Abdomen. — Provided with three dark- ocelli sometimes darker coloured. Eye in dorsal coloured, longitudinal fasciae as in L. tincta, but view 0.55 —0.58 X as wide as width of vertex the two dark stripes on segment 9 narrower and between eyes. Distance between lateral ocelli not united in front of caudodorsal beak. Dark

0.8 1 —0.92 X distance between lateral ocellus and ovipositor sheath 0.40—0.41 X as long as abdo- eye. Length of head 1.33 — 1.39 X as long as width men. of vertex between eyes; width of head 2.09—2.16 Measurements of the types: body length $: X as wide as width of vertex between eyes. Un- 21.4—21.6 mm, $: 21.6—21.8 mm; width of derside of head usually slightly darker. pronotum collar $' 6.2 6.6 mm, $: 7.2 mm; Thorax. — Dark with black patches between tegmen length $: 27.5—28.0 mm, $: 29-0-30.0 fissures and between fissures and pronotum col-

lar. Central fascia in females darker coloured than remaining part of pronotum. Pronotum collar Distribution. — South Sulawesi. 2.31 —2.55 X as wide as length of head, Types. — Indonesia, Sulawesi: Holotype: 1.47 — 1.65 X as wide as width of head. Obconical "S.W. Celebes, 1100 m/Mt. Lompobatang/area, areas distinctly recognizable. Cruciform eleva- Malino, 2,/8— 10.VÌ.1982/M. A. Lieftinck" $ tion usually with median longitudinal dark stripe. holotype of Lembeja lieftincki (RMNH). Para- M. R. DeJonG: Oriental Prasiini 197

Figs. 38—40. Lembeja sanguinolenta, holotype. 38, apex of aedeagus; 39, tergite 1, dorsal view; 40, pygofer, lateral view. 198 Tijdschrift voor Entomologie, deel 130, 1987

Fig. 41. Lembeja sanguinolenta, pygofer, ventrolateral view, holotype.

types: same data as holotype, 2 (5 1 $ (RMNH); Description. Macassar (= Ujung Pandang), Rippon coll., 1 $ Body green. Tegmina greenish, tegmen areas (NMWC). usually faintly red mottled. Head and pronotum Etymology. — The species is named after its together 0.95 — 1.21 X as long as meso- and me- collector, the late Dr Lieftinck, odonatologist, in tanotum together. Head and thorax together in recognition of his major contributions to the bi- males 0.69—0.95 X, in females 0.92—1.10 X as ogeography of Indonesia and the Pacific region. long as abdomen. Head. — Eye in dorsal view 0.45 —0.56 X as n.sp. Lembeja consanguinea wide as width of vertex between eyes. Distance (figs map 49—56; 3) between lateral ocelli 0.71 — 1.04 X distance between lateral ocellus and eye. Length of head

This species is described after a large series of 1.26 — 1.56 X as long as width of vertex between specimens, collected by various participants to eyes; width of head 1.84 —2.12 X as wide as width the "Project Wallace" expedition. of vertex between eyes. Transverse ridges in the M. R. DE Jong: Oriental Prasiini 199

Figs. 42—46. Lembeja lieftincki, Q holotype. 42, pygofer, lateral view; 43, apex of aedeagus and mediane uncus part, dorsal view; 44, pygofer, ventrolateral view; 45, clasper and mediane uncus part, lateral view; 46, apex of aedeagus, lateral view. —

200 Tijdschrift voor Entomologie, deel 130, 1987

47 lieftincki; Al holotype; paratype. tergite dorsal view; Figs. —48. Lembeja , $ 48, 9, 47, 1, 48, sternite 7, ventral view.

same colour as underside postclypeus. Rostrum lobes; provided with unsclerotized dorsal aedea- with black apex reaching middle trochanter. gal appendage. Thorax. — Pronotum collar 2.02 —2.58 X as Female: Operculum. — Small. Meracanthus wide as length of head, 1.41 — 1.62 X as wide as long and pointed. width of head. Obconical areas of mesonotum Abdomen. — Green. Ovipositor sheath hardly or not discernable. 0.23 —0.29 X as long as abdomen. Legs. — Same colour as underside body. Basal Measurements of the types: body length $: spine on fore femora pointed. 18.9—22.1 mm, x = 20.9 ± 0.8 mm, $: Tegmina and wings. — Tegmina subhyaline, 16.7 —21.7 mm, x = 19-2 ± 1.1 mm; width of green; tegmen areas usually faintly mottled with pronotum collar $: 5.7 6.6 mm, x = 6.1 ± 0.2 red. Venation green to yellowish. Transverse mm, $: 5.7 — 7.2 mm, x: 6.4 + 0.4 mm; tegmen vein of 2nd ulnar area extending well into 3rd length $: 24.6—28.4 mm, x = 26.6 + 1.1 mm, one. Corial fold clearly recognizable. Third ulnar $: 25.0—29.4 mm, x = 27.1 ± 1.1 mm. area 0.88—^1.04 X as long as 1st one; 4th ulnar area 0.82 — 1.05 X as long as radial area. Third Distribution. — North Sulawesi (map 3). apical area 0.88— 1.02 X as long as 4th one. Types. — Indonesia, Sulawesi: Holotype: "stat. Wings hyaline. Fusion of Cu2 and A^ at 79—90% 3/Forest/margin" (print), "Toraut/Base Camp/ from their origins. 29— 30.Ì.1985/J. P Duffels" (print), "Indonesia/ Male: Operculum. — Very small. Meracanthus Sulawesi Utara/Dumoga-Bone N.P./Project fairly broad, long and pointed. Wallace" (print) $ holotype of Lenibeja consan- Abdomen. — Green, reddish near hind edges guinea (ZMA). Paratypes: Base Camp Toraut, of tergites. Tergite 1 laterally sometimes brown- same labels as holotype, 1 (^ 1 $ (MZB); same ish. Mirrors fairly large. locality but: st. 7, lowland rainforest, MV light- long ridges alternating trap, 1— Tymbals. — Seventeen 2.Ü.1985, J. P Duffels &J. D. Holloway, with an equal number of short intercalary ridges. 1 (5 (ZMA); same locality but: st. 8, river bank, Lateral lobes Genitalia. — Pygofer very large. lowland rainforest, 2.ÌÌ.1985, J. P Duffels, 1 $ flat, apically dark. Caudodorsal beak short and (MZB); same locality but: st. 16, lowl. rainforest, rounded. Lateral surfaces of pygofer with a ridge- light-trap site 1 + 2 understorey/canopy, like structure running downwards from each lat- 7— 14.ii.1985, H. S. Barlow, 4 (5 10 $ (ZMA); eral lobe. Gaspers short, curved apically, slightly same locality but: st. 24, lowl. rainforest, light- pointed. Two huge, broadly rounded parallel trap site 1, understorey, 17 —26.ii. 1985, H. S. shields elevating distally of the claspers, con- Barlow, 4 (5 3 $ (MZB) 4 (5 2 $ (ZMA); same nected distally by a medial, protruding flattened locality but: base camp, ii.l985, 1 $ (BMNH); lip, forming a tube-like structure. Aedeagus long same locality but: rothamsted light-trap site 1, and very slender, with two long, apically rounded 200 m, V.1985, H. Barlow, 4 $ (BMNH); same M. R. DeJong: Oriental Vrastini 201

Figs. 49— 53. Lemheja consanguinea, $ paratypes Toraut. 49, pygofer, lateral view; 50, clasper and mediane uncus part, lateral view; 51, aedeagus with dorsal aedeagal appendage, laterodorsal view; 52, 53 apex of aedeagus, 52, lateral view, 53, dorsal view. 202 Tijdschrift voor Entomologie, deel 130, 1987

data as previous but: vi.l985, 3^2$ (BMNH); 67). Claspers relatively long, ventrally concave. Median uncus part narrow and same locality but: 8—23.X.1985, J. B. Heppner, compressed. Ae- 1 (5 8 $ (FSC); same locality but: 19.x. 1985, M. deagus short; sturdy apex with slightly incised R. de Jong, 1 S (ZMA); Edwards Camp 664 m, dorsal projection. MV light-trap understorey/canopy, st. 27, Lembeja parvula n.sp. P. 1 19.ii.1985. J. Duffels & J. D. Holloway, S 3 (figs. 61 67; map $ (ZMA); same locality but: st. 29, — 3) 24— 25.ii.1985, 1 ? (ZMA); same locality but: iv.l985,J. H. Martin, 1 S (BMNH); Goeroepahi, This small species is described after one male Baeturia, 6.ÌÌ.1917, W. Kaudern, 1 Ç (RMNH); and one female specimen. Hog's Back Camp 492 m, s.-side of ridge, st. 15, lowl. rainforest understorey/canopy, MV light- Description. P. Body yellowish-green in male, brownish-och- trap, 12— 13.ii.1985, J. Duffels &J. D. Hollo- way, 1 S (ZMA); Kosingsolan, P. P. A. headquar- reous in female. Head and pronotum together ters, at light, René Dekker & Charlotte Ver- 1.0— 1.15 X as long as meso- and metanotum meulen, 23.iii.1985, 1 9 (ZMA); Minahassa, together. Head and thorax in male 0.72 X, in Prasîa foliata (Walk) Stal $}, det. MacGilL, coll. female 0.98 X as long as abdomen. D. MacGillavry, 1 $ (ZMA); Molosso Island, Head. — Green in male; brownish with paler underside in female. Antennae darker than re- station 100, xi.l985, R. Bosman & J. v. Stalle, 2 $ (BIN); Page Camp 302 m, st. 9, lowl. rainfor- maining part of head. Eye in dorsal view 0.42 0.49 as wide as width of vertex between est, MV light-trap, 4— 8.ii.l985, J. P Duffels & J. — X D. Holloway, 2 $ (ZMA). eyes. Distance between lateral ocelli 0.58 —0.71 Etymology. — The name is derived from con- X distance between lateral ocellus and eye. sanguinea (latin for "related"), for being closely Length of head 1.25 — 1.36 X as long as width of related to L. lieftincki, by the typically shaped vertex between eyes; width of head 1.84 — 1.97 X median uncus part. as wide as width of vertex between eyes. Rostrum with black apex reaching middle trochanter. Remarks. Thorax. — Pronotum in male unicoloured, in

I believe L. consanguinea and L. lieftincki to be female with dark-coloured central fascia, and very closely related. A strong synapomorphy provided with dark patches, especially between might be found in the shape of the median uncus fissures and pronotum collar. Pronotum collar part. 2.16—2.24 X as wide as length of head, 1.49— 1.52 X as wide as width of head. Meso- The Lembeja parvula group notum with the same colour as pronotum in Head triangularly to obconically protruding in male, in female dark-brown, though laterally

dorsal view. Antennal segment 1 long. Male somewhat lighter, and provided with a median, operculum small, not covering tymbal cavities. brown stripe on cruciforum elevation. Tegmina hyaline; a regular pattern of spots in Legs. — Same colour as underside thorax in veins, sometimes also in tegmen cells; Cuj and Ai male; in female with dark-brown patches, basally forming -a small triangle at tegmen border. and apically on femora, tibiae and tarsi. Armature Wings with 5 apical areas; Cuj and Ai fused. of fore femora as in fig. 60. Abdomen in males weakly carinate. Tergites Tegmina and wings. — Tegmina hyaline. strongly folded laterally, forming a ridge on each Veins in apical half of tegmen in male with side of the sternites. Abdomen in males in lateral brown spots, venation yellowish-green. Tegmina in female spotted spots in view more or less triangular shaped. Tergite 1, with veins, and with with two short and pointed lateroproximal flaps, tegmen cells in basal half of tegmen. Third ulnar only slightly swollen and faintly dinted near area 0.96— 1.12 X as long as 1st one; 4th ulnar proximal border. Tymbal provided with 18— 19 area 0.8 —0.81 X as long as radial area. Third long ridges, alternating with an equal number of apical area 0.81 —0.86 X as long as 4th one. short intercalary ridges. Abdomen in females Wings hyaline. Fusion of Cuj and A, at 63 —78% slender, dorsally carinate. Ovipositor sheath ex- from their origins. tending just beyond caudodorsal beak. Lateral Male: Operculum. — Small. Meracanthus lobes of pygofer short and fairly flat. Edge of slender and pointed. pygofer between each lateral lobe and caudodor- Abdomen. — Mirrors medium-sized. sal beak obliquely running hindwards (figs. 63, Tymbal. — Provided with 19 long ridges alter- M. R. DE Jong : Oriental Prasiini 203

Figs. 54 — 56. Lembeja consanguinea; 54, 56, $\ 55, $ paratype Toraut. 54, pygofer, ventrolateral view, paratype Toraut; 55, sternite 7, ventral view; 56, tergite 1, dorsal view, holotype. nating with an equal number of short intercalary Caudodorsal beak with dark-brown apex. Dark ridges. ovipositor sheath 0.2 X as long as abdomen. Genitalia. — Lateral lobes of pygofer fairly Measurements of the types; body length $: broad. Caudodorsal beak small. Claspers very 15.3 mm, $: 13.7 mm; width of pronotum collar long, apically only slightly recurved and weakly $: 4.3 mm, $ 4.5 mm; tegmen length $\ 20.3 pointed. Median uncus part triangularly com- mm. $; 19.6 mm. pressed. Dorsal projection of apex of aedeagus fairly long. Distribution. — South Sulawesi (map 3). Female; Operculum. — Small, basal half dark Types. — Indonesia, Sulawesi: Holotype; coloured. Meracanthus short and pointed. "Bantimoerang/552" (handwritten), "coll. A. Ja- Abdomen. — Three longitudinal dark-brown cobi" (print), "Staatl. Museum für Tierkunde fasciae, a dorsal, medial one, up to segment 8 and Dresden" (print) 1 $ holotype of Lembeja par- two lateral ones up to segment 7. Segment 9 with vula (SMD). Paratype: same data as holotype but; two broad laterodorsal, slightly darker fasciae, Lembeja sp., 1 $ (SMD). just not uniting in front of caudodorsal beak. 204 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 57 —60. The L. parvula group: 57, male abdomen, ventral view, Lembeja wallacei; 58, head and pronotum, lateral view, L. parvula; 59, right tegmen and wing, L. wallacei; 60, femur, lateral view, L. parvula.

Etymology. — The name is derived from par- females 0.91 — 1.02 X as long as abdomen. vula (latin for "small") as it is a small-sized spe- Head. — Uniformely green in males; in fe- cies. males dorsally irregularly covered with some light patches, postclypeus ventrally greenish. Antennae dark-brown. Eye in dorsal view Lembeja wallacei n.sp. 0.37 —0.53 X as wide as width of vertex between (figs. 68— 73, 80, 81; map 3) eyes. Distance between lateral ocelli 0.76— 1.0 X distance between lateral ocellus and eye. Length The species is described in comparison with L. of head 1.23 — 1.38 X as long as width of vertex parvula, after a series of specimens collected by between eyes; width of head 1.75 —2.05 X as participants to the "Project Wallace" expedition. wide as width of vertex between eyes. Rostrum as in L. parvula. Description. Thorax. — In males uniformely green, in fe- Body green in males, dark-brown in females. males darker coloured than in L. parvula. Pron- Head and pronotum together 1.08 — 1.24 X as otum collar 2.03 —2.25 X as wide as length of long as meso- and metanotum together. Head head, 1.35 — 1.58 X as wide as width of head. and thorax together in males 0.75 —0.93 X, in Legs. — Same colour as underside thorax in M. R. DE Jong; Oriental Prasiini 205

61 62

Figs. 61 —73. 61 —67, Lembeja parvula; 61 —66, $ holotype; 67, $ paratype. 61, pygofer, lateral view; 62, clasper, lateral view; pygofer, ventrolateral view; 64, apex of aedeagus, lateral view; 65, tergite 1, dorsal view; 66, apex of aedeagus, dorsal view; 67, sternite 7, ventral view. 68—73, Lembeja wallacei; 68, 70. Q paratype Page Camp, 69, 71, 73, S holotype; 72, $ paratype Edwards Camp. 68, apex of aedeagus, lateral view; 69, pygofer, ventrolateral view; 70, apex of aedeagus, dorsal view; 71, clasper, lateral view; 72, sternite 7, ventral view; 73, pygofer, lateral view. 206 Tijdschrift voor Entomologie, deel 130, 1987

ß f

Figs. 74—77. General facies. 74, Lenibeja fatiloqua S, Calian; 75, Lembeja fatiloqua $, Z. del Sur; 76, Lembeja tincta $, Bua Kraeng; 77, Lembeja tincta $. Bua Kraeng. M. R. DE JONG: Oriental Prasiini 207

Figs. 78—81. General faciès. 78, Lembeja lieftincki $, paratype; 79, Lembeja lieftincki $, paratype; 80, Lembeja wallacei S, holotype; 81, Lembeja wallacei $, paratype Edwards Camp. 208 Tijdschrift voor Entomologie, deel 130, 1987

males. In females fore legs with brown coxa, St. 27, lowl. rainforest, MV light-trap, understo- trochanter and femur, middle and hind legs with 20.ii.l985, rey/canopy, J. R Duffels &J. D. Hol- brown coxae, patches on femora, tibiae and tarsi. loway, 1 (5 1 $ (ZMA); same locality but: Tegmina and wings. — Tegmina hyaline. In 22.V.1985, 1 $ (BMNH); same locality but: light- males veins conspicuously spotted; tegmen cells sheet, 23.vi.1985, M. R. Wilson, 1 $ (BMNH); in basal half of tegmen provided with spots; Hog's Back Camp, 492 m, stat. 30A, lowl. rain-

yellowish green. Coloration in females forest, light-trap, 14. ii. venation MV canopy, 1985, J. D. more prominent than in L. parvula. Third ulnar Holloway, 1 $ (MZB); Page Camp, 302 m, stat. area 0.88—0.98 X as long as 1st one; 4th ulnar 9, lowl. rainforest, MV light-trap, 4— 8.ii.l985, J. radial one. Third area 0.85 —0.93 X as long as R Duffels & J. D. Holloway 4 ^ 3$ (ZMA). apical area 0.80—0.97 X as long as 3rd one. Etymology. — The species is named after Fusion of Cuj and A, in wings at 43 —69% from Alfred Rüssel Wallace, for numerous reasons. their origins. Male: Operculum. — As in L. parvula. References

Abdomen. — As in L. parvula, but with faintly The list presented here is additional to the

reddish hind edges of the tergites. references of Part 1 and 2 of the study "Taxon- Tymbals. — Provided with 20 (sometimes 19) omy and biogeography of oriental Prasiini" (de long ridges alternating with an equal number of Jong, 1986). short intercalary ridges. Ashton, H., 1914. Catalogue of the Cicadidae in the Genitalia. — Dark lateral lobes of pygofer ap- South Australian Museum; with descriptions of ically slender in L. parvula. Gaspers more than several new species. — Transactions of the Royal uncus part flat. somewhat more curved. Median Society of South Australia 38: 345 — 358, pi. 17. Apex of aedeagus more sturdy with short dorsal Boulard, M., 1985. Apparence et mimétisme chez les.

projection. Cigales [Hom. Cicadoidea. — ]. Bulletin de la So- Female: Operculum. — Dark coloured. Shape ciété Entomologique de France 90 (1 —4): 1016—1051. as in L. parvula. Burns, A. N., 1957. Check list of Australian Cicadidae. Abdomen. — Dark-brown. Segment 8 with a — Entomologische Arbeiten aus dem Museum side. pale lateral spot on each Hind edges of Georg Frey 8 (2): 609—678. tergites reddish. Dark ovipositor sheath Distant, W. L., 1910. Australian Cicadidae, with re- 0.22 —0.30 X as long as abdomen. marks on some recent disputation. — Annales de la Entomologique de Belgique 54: Measurements of the types: body length $: Société 415—420. 12.5—16.2 mm, x = 14.0 + 1.1 mm, $: Distant, W. L., 1913. Synonymical notes on some 11.8—13.6 x = 12.5 + 0.5 mm; width of mm, recently described Australian Cicadidae. — Pro- pronotum collar $: 3.8 —4.5 mm, x = 4.2 + 0.2 ceedings of the Linnean Society of New South mm, $: 3.8 —4.3 mm, x = 4.1 + 0.1 mm; tegmen Wales 37: 600—601. length (5: 18.2—20.1 mm, x = 19.2 + 0.6 mm, $: Gaedike, H., 1971. Katalog der in den Sammlungen 17.0—19.2 mm, x = 18.1 + 0.7 mm. des ehemaligen Deutschen Entomologischen In- stitutes aufbewahrten Typen-VI (Homoptera (ex- clusive Aphidina)). — Beiträge zur Entomologie Distribution. — North Sulawesi (map 3). 21 (3—6): 315—339. Indonesia, Sulawesi: Holotype: Types. — Horvath, G., 1912. Miscellanea Hemipterologica, xii. "Stat. U/Lowland/ rainforest" (print), "Toraut/ Adnotationes Synonymicae et systematical — bank of Tumpah R./( recreation area)/8.ii.l985/ Annales Historico-Naturales Musei Nationalis (print), "Indonesia/Sulawesi Hungarici 10: 607—609. J. P. Duffels" Utara/Dumoga-Bone N.P./Project Wallace" Jacobi, A., 1941. Die Zikadenfauna der Kleinen Sun- dainseln. — Zoologische Jahrbücher (Syst.) 74: (print) 1 holotype of Lembeja wallacei (ZMA). (5 277—322. Paratypes: Base camp Toraut, same data as hol- Jong, M. R. de, 1986. Taxonomy and biogeography of but: st. lowl. otype, 1 (5 (MZB); same locality 16, Oriental Prasiini. 2. The follata group of the genus rainforest, light-trap site 1 + 2, understorey/ Lembeja Distant, 1892 (Homoptera, Tibicinidae). canopy, 7— 13.ii.l985, H. S. Barlow, 1 $ (ZMA); — Tijdschrift voor Entomologie 129: 141 — 180. annotations to M. Dis- same locality but: lowl. rainforest, 200— 300 m, Kirkaldy, G. W., 1907. Some tant's recent Catalogue of Cicadidae [Hem.] (1). — 2 S (BMNH); same locality but: Toraut, M. R. Annales de la Société Entomologique de Belgique Wilson, vi.1985, 2 (BMNH); same locality but: $ 51: 303—309. forest, R. Wilson, malaise-trap Toraut M. Mac-Lachlan, R., 1891. The genus Perissoneura. — vi.1985, 1 S (BMNH); same locality but: Entomologische Nachrichten Berlin 17: 13.ii.1985, 1 (5 (BMNH); Edwards Camp, 664 m, 319—320. M. R. DE Jong: Oriental PrasUni 209

Moulds, M. S., 1975. The song of the cicada Lembeja Australia: 387—457, Figs. 26.1—26.74, PI. 3. — brunneosa (Homoptera: Cicadidae) with notes on Melbourne University Press. the behaviour and distribution of the species. — Journal of the Australian Entomological Society 14: 251 —254, figs. 1 —6. {Author's address: Institute of Taxonomie Zoology, T. E. V. Eastop, P.O. Box Woodward, & J. W. Evans & E 1970. 20125, , 1000 HC Amsterdam, The Nether- Hemiptera (Bugs, Leafhoppers, etc.). In: Insects of lands.)

Tijdschrift voor Entomologie 130: 211 —223 Gepubliceerd 30 november 1987

ON FOUR NEW PALAEARCTIC SPECIES OF THE GENUS CECIDOMYIA (DIPTERA, CECIDOMYIIDAE)

W. NIJVELDT

Gruttoweide 122, Wageningen, The Netherlands

Abstract

Larvae of the genus Cecidomyia live in resin of conifers, mainly pines. Twelve described species are recorded from North America, Europe and Asia. In this paper two new Euro- pean and two new Asiatic species are described and figured: Charmi, C japonica, C. phagwariae and C. sarae. A redescription of the type-species, C. pini, is given and a

neotype is designated.

Introduction European species are not known to cause eco- Larvae of the oldest gall midge genus Cecido- nomic damage but, according to Gagné (1978 myia are known to live in resin of conifers, b), the North American species promote brea- mainly pines. The first taxonomie revision of kage and secondary infection on twigs and Cecidomyia was published by Gagné (1978 b). branches of pines. C. bisetosa was the first spe- In his paper eleven species are recorded, eight cies found to feed on cones, inducing malforma- from North America and Cuba, two from Eu- tions of the scales and preventing release of the rope and one, or possibly two, from the Hima- seeds. The second record is that by Grijpma layan region. The twelfth species, C. bisetosa (1981). He found larvae of C. pini in resin exu- Gagné, was discovered too late to be included in dations between the scales of green cones on this revision (Gagné, 1978 a). C. mesastatica, P. sylvestris at Grubbenvorst, Province of Lim- very briefly described from two males and one burg, The Netherlands. However, it is not yet female by Mamajev (1971), was not included in known if seed production and seed release were Gagné's papers. This species was bred from res- affected by the feeding of the larvae. in on spruce in the Kirghizian Republic, USSR. Referring to the literature mentioned here it I have been unable to obtain specimens for may be concluded that Cecidomyia larvae feed study, but the description indicates that C. me- directly on the plant tissue and not on the resin sasiatica closely resembles C. magna (Mohn). Itself. It is more likely that the latter serves as a The genus Cecidomyia currently contains the medium for protection, though several hyme- following described species: C. bisetosa Gagné, nopterous parasites were reared from them in C. brevispatula Gagné, C. candidipes Foote, the past years. C. fortunactus Gagné, C. magna (Mohn), C. Pupation takes place in or outside the resin mesasiatica Mamajev, C. pini (De Geer), C. pi- masses. According to Gagné (1978 b), Nearctic niinopis Osten Sacken, C. reburrata Gagné, species that pupate apart from the resin have C. resinicola (Osten Sacken), C. resinicoloides distinctive dorsal abdominal lobes (fig. 13), Williams and C. tortilis Gagné. However, ex- which may be helpful in leaving the mass. Con- amination of material bred from resin masses, trary to this, C. magna, a European species, is taken from Pinus sylvestris L. in Europe, from recorded as pupating in the resin mass, unlike all P. thunbergii Pari, in Japan and from P. rox- the other species whose larvae have dorsal lobes burghii Sarg, in Pakistan showed that four new (Mohn, 1955). However, in the winter of 1984- species can be added. They will be described be- 1985 I found a mature larva which had left the low. resin to pupate in a white cocoon attached to a The larvae of these species of Cecidomyia live branch of a spruce tree near Wageningen. completely submerged in the resin, keeping The following description of the European their protruding hind spiracles free from it. The type-species is, as far as the adults are con-

211 212 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 1 —4. C. pini, 1: head; 2: third male flagellomere; 3: distal male flagellomere; 4: palp segments. Scale line of fig. 1: 0.25 mm, of figs. 2—4: 0.1 mm. Figs. 1 —4 after neotype.

cerned, based on material from Sweden, where 9 and two cocoons) were reared from Pinus pi- the type-locahty is situated. naster Ait. {P. maritima Mill.), SW. France, and

that it is not known if types exist or, if so, Cecidomyia pini (De Geer) where, but that females and cocoons are not di-

(figs. 1-14) agnostic. The type of C. pilosa Bremi is in poor TipulapiniDe Geer, 1776: 417 (Sweden). condition, covered with fungal hyphae, sex un- determined, "Bre. /Lindau" (nr. Zürich), in En- The following subjective synonyms of C. pint tomologisches Institut, Eidgenössischen Tech- are recorded by past authors: C pinimaritimae nische Hochschule, Zürich, Switzerland. This (as pini maritimae) Dufour, 1838: 294 and information was provided to Gagné by W. Saut- C. pilosa Bremi, 1847: 31, 61. That synonymy er of that Institute. Neither the type material was established when it was thought that only nor the original descriptions of C. pinimariti- one species occurred in Europe. Gagné (1978 b) mae and C. pilosa were adequate enough to use records that syntypes of C. pinimaritimae (two these names as subjective synonyms of C. pini W. NijVELDT: New Palaearctic Cecidomyia 213

Figs. 5—9. C. pini, 5: wing; 6: tarsal claws; 7: male terminalia (dorsal); 8: distal female flagellomeres; 9: ovipo- 7 sitor (lateral). Scale line of fig. 5: 1 mm; fig. 6: 0.05 mm and figs. —9: 0.1 mm. Figs. 5— 7 after neotype. 214 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 10— 12. C. pini, 10: pronotum (anterior); 1 1 : head of last instar larva (dorsal); 12: sternal spatula. All scale lines 0.1 mm. or as available names for my newly described circumfila about 0.25 and length of setae 0.53 European species. times the length of third flagellomere. Distal node with two whorls of looped circumfila and Male. — Head without postvertical peak (fig. one whorl of setae; length of inferior circumfila

1). Antenna with twelve flagellomeres, binodal, of this node about 0.21, of superior circumfila first and second not connate; three circumfila 0.25 and of setae 0.35 times the length of third (fig. 2). Proximal node of third flagellomere: flagellomere. Nodes covered with microtnchia, length about 0.76 times its diameter; distal stalks bare, distal flagellomere tapering into a node: length 1.18 times its diameter. Length of short stalk-like process (fig. 3). Maxillary palps proximal stalk about 0.11 and of distal stalk 0.17 with four segments, about 0.54 times the height times the total length of third flagellomere. of head (fig. 4). Wing length about 2.5 mm and Proximal node with one whorl of looped cir- twice as long as wide; R 5 curved distally, join- cumfila and one whorl of long setae; length of ing C posterad of wing apex; C broken at June- W. NijVELDT: New Palaearcüc Cecidomyia 215

1 13 .---

Figs. 13 — 14. C. pint, 13; larval abdominal segment with dorsal tubercles; 14: larval terminal segment. Figs. 15 — 17. C. magna, 15: sternal spatula; 16: larval terminal segment; 17: larval head capsule. All scale lines 0.1 mm. 216 Tijdschrift voor Entomologie, deel 130, 1987

ture with R 5, Rs weak; M 3 + 4 fold present; comm.) and, in the interest of taxonomie stabili- Cu forked; R 5 about 2.2 times as long as R 1 ty, I here designate the following neotype of (fig. 5). Legs covered with brown scales; tarsal this species. Neotype: â, slide no. 5192, Swe- claws all simple, curved beyond midlength; em- den, Uppland near Stockholm, E. Sylvén, reared podia longer than claws (fig. 6). Abdomen elon- from resin on Finns sylvestris L., 1979. This des- gate cylindrical. Gonocoxite and gonostylus ignation stabilizes the usage by Barnes (1951), fairly stout; gonocoxite about 1.8 times as long Mohn (1955), Vockeroth (1960), Mamajev & as gonostylus, covered with setae except on in- Krivosheina (1965) and Gagné (1978 b). Para- ner side. Gonostylus , with apical tooth. Cerci types: 9, sHdes no. 5193-5196, with same data triangular, broadly rounded distally; hypoproct as holotype. All specimens in the Swedish Mu- shallowly emarginated. Aedeagus simple, short seum of Natural History, Stockholm. and rounded apically (fig. 7). Other material (larvae, collected from resin Female. — Antenna with twelve uninodal, on P. sylvestris in the Netherlands): slides no. stalked flagellomeres, first and second not con- 4537-4541, Grubbenvorst, October 1980, P. nate. Flagellomeres cylindrical with two cir- Grijpma, two year old cones; no. 4556—4566, cumfila connected by two longitudinal strands Ede, 26.X.1980; no. 4573—4575, Ede, October and with two whorls of setae (fig. 8). Node of 1980; no. 4576, Remmerden, 20.xi.l980; no. third flagellomere with a length of about 2.85 4577—4579, Leuvenum, October 1980; no. times its diameter, length of stalk 0.2 times the 5146—5147, Wageningen, 6.xii.l984; no. total length of third flagellomere. Length of 5158—5172, Grubbenvorst, 13.viii.l980, P. proximal whorl of setae about 0.38 and of distal Grijpma, two year old cones. Pupal skin, slide setae 0.47 times the length of third flagellomere. no. 2225, from resinous cocoon on needle of Nodes covered with microtrichia, stalks bare. P. sylvestris, Wageningen, 21. vii. 1960. 9, slide Distal flagellomere tapering with rounded tip. no. 3569, from resinous cocoon on needle of Maxillary palps with four segments and about P. sylvestris. Remmerden, 1960. Specimens de- 0.45 times height of head. Wing length about posited in the collection of the Instituut voor 3.2 mm and 2.4 times as long as wide; R 5 about Taxonomische Zoölogie (Zoölogisch Museum), 2.3 times as long as R 1. Abdomen elongate, Amsterdam. ovoid. Ovipositor short, retractile; cerei with two short apical sensoria and covered with setae Pupation of C. pini takes place apart from the (%9). resin mass in a white resinous cocoon. The fact Pupa. — Antennal horn ridged anteriorly; that larvae from C. barrisi and C. sarae, from pronotum as in fig. 10. which the adults are still unknown, have been Last instar Jarva. — Length about 5.5 mm. found together with those of C. pini in the same

Head capsule with apodemes of about 122 |i resin masses makes the exact identity of the re- (fig. 11). Sternal spatula light brown; total maining adult specimens in this collection rather length about 108 |J,; anterior end broadened, uncertain. Study of more material, reared from the cephalic margin somewhat convex (fig. 12). identified larvae, will be necessary to clarify this Pleural and dorsal abdominal papillae situated problem. on tubercles, two lateral pairs of dorsal papillae each on a forked lobe (fig. 13). Length of setae Cecidomyia magna (Mohn) of dorsal papillae from the left to the right: pro- (figs. 15-17) thorax: about 17, 20, 14, 14, 20, 17 \x; meso- Stelechodiplosis magna Mohn, 1955: 127— 151 (Ger- thorax: 20, 22, 14, 14, 22, 20 |x; metathorax: many).

22, 24, 15, 15, 26, 21 [i; 1st abdominal segment: 27, 34, 20, 20, 34, 27 \i; 7th abdominal This second European species was originally segment: 24, 27, 20, 19, 27, 24 ^i. Terminal described as Stelechodiplosis magna by Mohn papillae: two on each side; one with tapered seta (1955), but was included in Cecidomyia by of about 17 |x and one with peg-like seta of Vockeroth (1960). It was found in Germany in about 14 [Ji. Terminal spiracles bilaterally the resin of spruce in which, according to Mohn symmetrical, four caudal prongs with a length (1955), it also pupates. However, in the winter of about 10|i(fig. 14). of 1984— 1985 I found a mature larva which had left the resin to pupate in a white cocoon like Original type material of C. pini must be con- the other species whose larvae have two lateral sidered non-existant (Prof. E. Sylvén, pers. pairs of dorsal papillae on prolonged lobes. The W. NijVELDT: New Palaearctic Cecidomyia 217

Figs. 18 — 20. C. sarae, 18: sternal spatula; 19: larval terminal segment; 20: larval head capsule. Figs. 21 — 22. C. barrisi, 21: sternal spatula; 22: larval terminal segment. Fig. 23. C. japonica, male terminalia (dorsal). Scale line of fig. 20: 0.05 mm and of figs. 18, 19, 21, 22 and 23: 0.1 mm. 218 Tijdschrift voor Entomologie, deel 130, 1987

cocoon was attached to a branch of a spruce tree England, where it was found in 1980 by Dr K. near Wageningen, The Netherlands (Nijveldt, M. Harris (Commonwealth Institute of Ento- 1985). Mohn included illustrations of male ter- mology, London) in resinous wounds on P. syl- minalia, larval spatula and larval terminal seg- vestris in the Wisley Garden of the Royal Horti- ments in his description. Figs. 15 and 16 show cultural Society. It is not known whether the some larval characters. larvae leave the resin to pupate. A recent record of C. magna in England is: Shropshire, Cantlop near Shrewsbury, Cecidomyia harrisi n. sp. 18.x. 1984, in resin masses on Picea abies (figs. 21-22) (spruce). CIE Coll. A. 16422, four larvae in the I found the larvae in resin on Pinus sylvestris. British Museum (Natural History) (Dr K. M. Adult and pupa. — unknown. Harris, pers. comm.) Last instar larva. — Length about 3.3 mm.

Apodemes of head capsule about 68 \i (fig. Cecidomyia sarae n. sp. 20). Sternal spatula large, light brown, with a (figs. 18—20) total length of about 211 ^; anterior end

I found the larvae of this species in resinous broadened, the cephalic margin deeply cleft (fig. wounds and in resin lumps, caused by Retinia 21). Pleural and dorsal abdominal papillae with resinella L. (Lepidoptera) on Pinus sylvestris. short setae, situated on small tubercles. Lateral pairs of dorsal papillae not on prolonged lobes. Adult and pupa.—unknown. Length of setae of dorsal papillae from the left Last instar larva. — Length about 4.5 mm. to the right: prothorax: about 4, 7, 7, 7, 7, 4 Apodemes of head capsule about \12 \i (fig. H; mesothorax: 5, 7, 7, 7, 7, 5 n; metatho- 17). Sternal spatula dark brown, slender with a rax: 7, 7, 10, 10, 7, 7 \i; 1st abdominal seg- total length of about 108 jx; anterior end not ment: 7, 7, 10, 10, 7, 7 \i; 7th abdominal seg- broadened, the cephalic margin convex (fig. 18). ment: 7, 9, 10, 10, 9, 7 [1. Terminal papillae: Pleural and dorsal abdominal papillae with short three on each side; two with tapered seta of setae, situated on tubercles. No lateral pairs of about 14 n and one with peg-Hke seta of dorsal papillae on prolonged lobes. Length of about 14 \i. Terminal spiracles bilaterally setae of dorsal papillae from the left to the right: symmetrical, no caudal prongs (fig. 22). prothorax: about 10, 14, 8, 8, 14, 10 ^i; mesothorax: 12, 15, 10, 10, 15, 14 ^; metathorax: Holotype: one larva in resin on branch of

17, 20, 8, 7, 19, 15 fx; 1st abdominal segment: P. sylvestris. Ede (The Netherlands), October

20, 19, 10, 10, 18, 21 \i; 7th abdominal seg- 26.X.1980, W. Nijveldt, shde no. 4570. Para- ment: 17, 14, 7, 7, 14, 17 \}i. Terminal papil- types: three larvae with same data as holotype, lae: two on each side; one with tapered seta of sHdes no. 4569, 4571 and 4572. Specimens de- with peg-like seta of posited in the collection of the Instituut voor about 14 fj, and one about 13 ji,. Terminal spiracles longer mesally Taxonomische Zoölogie (Zoölogisch Museum), than laterally, four caudal prongs with a length Amsterdam. of about 20 n (fig. 19). I name this species in honour of Dr K. M. Harris (Commonwealth Institute of Entomolo- Holotype: one larva, in resin on branch of gy, London), who collected this species in 1980 P. sylvestris. Ede (The Netherlands), 26.x. 1980, for the first time from resinous stem wounds on W. Nijveldt, slide no. 4567. Paratypes: one lar- P. sylvestris in the Wisley Garden of the Royal va with same data as holotype, slide no. 4568; Horticultural Society in England. C. harrisi one larva in resin lump on a twig of P. sylvestris, closely resembles C. fortunactus, a species with caused by the larva of Retinia resinella, Wage- mainly plesiomorphic characters, as discussed ningen (The Netherlands), November 1960, W. by Gagné (1978 b) in his analysis. Nijveldt, slide no. 2271. Specimens deposited in the collection of the Instituut voor Taxono- Cecidomyia japonica n. sp. mische Zoölogie (Zoölogisch Museum), (figs. 23—28)

Amsterdam. Dr J. Yukawa (University of Kagoshima, Ja- I name this species after my wife Sara, in grat- pan) collected the larvae and reared the asso- itude for her great help during my study of the ciated adults from resin on Pinus thunbergii

Cecidomyiidae. C. sarae is also known from Pari, in Japan. W. NijVELDT: New Palaearctic Cecidomyia 219

Figs. 24—28. C. japonica, 24: pronotum (anterior); 25: larval head capsule; 26: sternal spatula; 27: larval abdominal segment with dorsal tubercles; 28: larval terminal segment. All scale lines 0.1 mm.

Male. — Head without postvertical peak. cumfila and one whorl of long setae; length of Proximal node of third flageliomere: length circumfila about 0.21 and length of setae 0.62 about 0.75 times its diameter; distal node: times the length of third flageliomere. Distal length about 1.7 times its diameter. Length of node with two whorls of looped circumfila and proximal stalk about 0.11 and of distal stalk 0.18 one whorl of setae; length of inferior circumfila times the total length of third flageliomere. of this node about 0.18, of superior circumfila Proximal node with one whorl of looped cir- 0.25 and of setae 0.46 times the length of third 220 Tijdschrift voor Entomologie, deel 130, 1987

flagellomere. Distal flagellomere tapering into a 22. vi. 1967, on P. thunbergii, J. Yukawa; pupal long stalk-like process. Maxillary palps with skins, slides no. 5207 and 5208 with same data. four segments, about 0.44 times the height of All specimens in the collection of the University head. Wing length about 2 mm and twice as of Kagoshima, Japan. long as wide; R 5 about 2.5 times as long as R 1. I name this species japonica because it is the Legs covered with brown scales. Abdomen first Cecidomyia species found in Japan. elongate cylindrical. Gonocoxite and gonostylus fairly stout; gonocoxite about 1.9 times as Cecidomyia phagwariae n. sp. long as gonostylus, covered with setae except (figs. 29—36) on inner side. Gonostylus with apical tooth. Gagné (1978 b) recorded an undescribed Ce-

Cerci rounded apically; hypoproct narrower, cidomyia species, which is known from a male, incised, with rounded lobes. Aedeagus simple, a female, and three larvae in the U.S. National short and broadly rounded apically (fig. 23). Museum (Natural History) in Washington. Female. — Node of third flagellomere with a They are from Pinus roxburghii in Pakistan. He length of about 3.25 times its diameter, length did not describe this species because the male of stalk 0.17 times the total length of third flag- genitalia are slightly distorted on the slide and it ellomere. Length of proximal whorl of setae was not known whether the larvae leave the res- about 0.45 and of distal setae 0.5 times the in mass to pupate. However, through the cour- length of third flagellomere. Distal flagellomere tesy of Dr K. M. Harris, I have studied another tapering with pointed tip. Maxillary palps with three males, four females and one larva of the four segments and about 0.56 times height of same origin from the collection of the British head. Wing length about 3.1 mm and 2.4 times Museum (Natural History) in London. Togeth- as long as wide; R 5 about 2.3 times as long as R er with the slides of the U.S. National Museum

1. Abdomen elongate, ovoid. of Natural History, kindly sent on loan by Dr

Pupa. — Antennal horn pointed anteriorly R. J. Gagné, they form the basis of the follow- (fig. 24). ing description. Last instar larva. — Length about 4.3 mm.

Apodemes of head capsule about 122 [i (fig. Male. — Head with postvertical peak (fig. 25). Sternal spatula light brown with a total 29). Antenna with twelve flagellomeres, bino- length of about 102 \i; anterior end broad- dal, first and second not connate; three circum- ened, the cephalic margin somewhat convex fila. Proximal node of third flagellomere: length (fig. 26). Pleural and dorsal papillae situated on about 0.81 times its diameter; distal node: tubercles; two lateral pairs of dorsal papillae length about 0.79 times its diamter. Length of each on a forked lobe (fig. 27). Length of setae proximal stalk about 0.16 and of distal stalk 0.25 of dorsal papillae from the left to the right: pro- times the total length of third flagellomere. thorax: about 17, 20, 17, 17, 20, 17 \i; meso- Proximal node with one whorl of looped cir- thorax: 24, 34, 14, 15, 34, 24 \i; metathorax: cumfila and one whorl of setae; length of cir-

24, 31, 20, 20, 31, 22 \i; 1st abdominal seg- cumfila about 0.33 and length of setae 0.41 ment: 34, 41, 20, 20, — , 34 \i; 7th abdominal times the length of third flagellomere. Distal segment: 31, — , — , 20, 34, 31 \i. Terminal node with two whorls of looped circumfila and papillae: two on each side; one with tapered seta one whorl of setae; length of inferior circumfila of about \2 \i and one with peg-like seta of of this node about 0.27, of superior circumfila about 14 \i. Terminal spiracles bilaterally 0.29 and of setae 0.66 times the length of third symmetrical, four caudal prongs with a length flagellomere. Nodes covered with microtrichia, ofabout 13.6|i(fig. 28). stalks bare. Distal flagellomere tapering into a stalk-like process. Maxillary palps with four

Holotype: â , slide no. 5203, Koga, Fukuoka- segments, about 0.48 times the height of head. pref., Kyushu, Japan, 6. v. 1965 em., on Pinus Wing length about 2.7 mm and 2.4 times as long thunbergii, reared by Yukawa. Paratypes: 9, as wide; R 5 curved distally, but not so strongly slide no. 5204, 5205 with same data as holotype; as in C. pini and C. japonica, joining C at wing S, slides no. 5197—5199, Mitoma, Fukuoka- apex; C broken at juncture with R 5, R s weak; pref. Kyushu, Japan, 4— 17. vii. 1966 em., on M3-I-4 fold present; Cu forked; R 5 about twice P. thunbergii, reared by Yukawa; 9, slides no. as long as R 1 (fig. 30). Legs covered with 5200—5202 with same data; one larva, shde no. brown scales; tarsal claws all simple, curved be- 5206, Hanami, Fukuoka pref., Kyushu, Japan, yond midlength and about as long as empodia W. NijVELDT: New Palaearctic Cecidomyia 221

Figs. 29—33. C. phagwariae, 29: head; 30: wing; 31: tarsal claws; 32: male terminalia (dorsal); 33: larval head capsule. Scale line of fig. 29: 0.2 mm; 30: 1 mm, 31 : 0.05 mm and figs. 32—33: 0.1 mm.

(fig. 31). Abdomen elongated cylindrical. Go- with microtrichia, stalks bare. Distal flagello- nocoxite and gonostylus stout; gonocoxite mere tapering with pointed tip. Maxillary palps about 1.6 times as long as gonostylus, covered with four segments, about 0.52 times the height with setae except on inner side. Gonostylus of head. Wing length about 3.6 mm and 2.5 with apical tooth. Cerci broadly rounded distal- times as long as wide; R 5 2.3 times as long as R ly; hypoproct shallowly emarginated. Aedeagus 1. Abdomen elongate, ovoid. Ovipositor short, simple, longer than hypoproct and rounded api- retractile; cerei with two short apical sensoria cally (fig. 32). and covered with setae. Female. — Antenna with twelve uninodal Pupa. — unknown. flagellomeres, which are cylindrical with two Last instar larva. — Length about 6.3 mm. circumfila connected by two longitudmal Apodemes of head capsule about 109 \i (fig. strands. Node of third flagellomere with a 33). Sternal spatula light brown with a total length of about 2.87 times its diameter, length length of about 204 \i, anterior end broad- of stalk 0.18 times the total length of third flag- ened, the cephalic margin nearly straight (fig. ellomere. Length of proximal whorl of setae 34). Pleural and dorsal abdominal papillae situ- about 0.76 and of distal setae 0.39 times the ated on tubercles, two lateral pairs of dorsal pa- length of third flagellomere. Nodes covered pillae each on a forked lobe (fig. 35). Length of 222 Tijdschrift voor Entomologie, deel 130, 1987

Figs. 34—36. C. phagwariae, 34: sternal spatula; 35: dorsal abominai tubercles; 36: larval terminal segment. Scale line of fig. 35: 0.2 mm, of figs. 34 and 36: 0.1 mm. setae of dorsal papillae from the left to the right: U.S. Museum of Natural History: Paratypes: prothorax: about 27, 34, — , 26, 34, 27 \i\ S, slide 974, with same data as holotype; 9, mesothorax: 38, 41, 30, 30, 41, 38 |i; meta- slide 973, Lathrar, on P. roxburghii, 9— 12— 70, thorax: 41, 48, 30, 27, 48, 41 \i; 1st abdomi- CIBC (Pakistan Sta.) 71—2291, SB, 12/70— nal segment: 57, 63, 31, 31, 63, 57 |i; 7th ab- 48B; one slide with three larvae with same data. dominal segment: 54, 54, 30, 27, 54, 54 |i. I name this species after its type locality in Terminal papillae: two on each side; one with Pakistan. tapered seta of about 14 ^ and one with peg- like seta of about 14 |a. Terminal spiracles bilaterally symmetrical, four caudal prongs with a Key to last instar larvae (except those of length of about 14 |i (fig. 36). C Mesasiatica). 1. Abdomen without dorsal lobes 2 The following specimens are in the British — Abdomen with dorsal lobes (figs. 13, 28, Museum (Natural History): Holotype: 6, slide 37) 3 no. 17.270, Phagwari, Pakistan, 3— 12—70, on 2. Spatula large, hght drown, cephalic margin P. roxburghii Sarg., CIBC, 5B 12/70, 47B 964, deeply cleft (fig. 21); terminal spiracles bi- BM, 1975 CIE coll. A 4502. Paratypes: 9, slide laterally symmetrical, no caudal prongs (fig.

no. 17.275,962 with same data as holotype; S , 22); from Pinus sylvestris slide no. 17.272,960 and 9 slide no. 17.274,961 C. barrisi Nijveldt with same data; 9, slide no. 17.273,965, Lath- — Spatula shorter, dark brown, cephaHc mar- rar, Pakistan, 4—12—70, on P. roxburghii, SB. gin convex (fig. 18); terminal spiracles lon- 12/70—48 B; three larvae, slides no. 5209— ger mesally than laterally, four rather long

5211. Lathrar, Pakistan, on P. roxburghii, caudal prongs (fig. 19); from P. sylvestris . . . 9.XÌÌ.1970, CIBC CIE A 4502. Specimens in the C. sarae Nijveldt —

W. NijVELDT: New Palaearctic Cecidomyia 223

3. Spatula short, light brown, cephalic margin References

somewhat convex, shaft long and slender, Barnes, H. F., 1951. Gall midges of economic impor- tapering slowly (fig. 12); terminal spiracles tance, 5. Gall midges of trees: 1 —270. — Crosby bilaterally symmetrical, four caudal prongs Lockwood & Son Ltd., London. of moderate length (fig. 14); from P. sylves- De Geer, C., 1776. Mémoires pour servire à l'histoire des insectes, 5(6): 1 523, 30 pis. tris C. pini (De Geer) — — Stockholm. Gagné, R. 1978 a. A. new species of Cecidomyia — Spatula shorter than in C. pini, light brown, J., injurious to cones of slash pine in Florida. — The cephalic margin somewhat convex, shaft Florida Entomologist 61 (3): 193— 196. broader and tapering quickly (fig. 27); ter- Gagné, R. J., 1978 b. A systematic analysis of the pine minal spiracles bilaterally symmetrical, four pitch midges, Cecidomyia spp. (Diptera: Cecido- caudal prongs of moderate length (fig. 29); myiidae). — Technical Bulletin, United States De- from P. thunbergii .... C. japonica Nijveldt partment of Agriculture 1575: 1 — 18. 4. Spatula long, light brown, cephalic margin Grijpsma, P., 1981. A new feeding site of Cecidomyia pini larvae nearly straight, shaft slender (fig. 36); ter- on Pinus sylvestris (Dipt., Cecidomyii- dae). Entomologische Berichten, minal spiracles bilaterally symmetrical, four — Amsterdam 41:145—148. caudal prongs of moderate length (fig. 38); Mamajev, B. M. & N. P. Krivosheina, 1965. Larvae of from P. roxburghii gall midges (Diptera: Cecidomyiidae): compara- phagwariae Nijveldt C. tive morphology, biology and identification tables — Spatula long, black, cephalic margin (in Russian): 1 —278. — Moscow, Izdatelstvo straight, shaft broad- (fig. 15); terminal spi- Nauka. racles bilaterally symmetrical, four caudal Mamajev, B. M., 1971. The use of resin as a habitat by prongs of moderate length (fig. 16); from insects, with a review of insects living in resin (in Picea abies C. magna (Mohn). Russian, with English summary). — Zurnal Obscej Biologli 32 (1): 501—507. Acknowledgments Mohn, E., 1955. Neue freilebende Gallmücken-Gat- tungen. — Deutsche Entomologische Zeitschrift The author wishes to thank Dr K. M. Harris 2:127—151. (London) and Dr R. J. Gagné (Washington) for Nijveldt, W., 1985. Nieuwe galmuggen voor de Ne- their kind suggestions, for critical reading the derlandse fauna (IX) (Diptera: Cecidomyiidae). — manuscript and for lending material of C. phag- Entomologische Berichten Amsterdam: 45: 85 wariae. Furthermore Prof, dr E. Sylvén (Stock- 88. Vockeroth, R., 1960. Taxonomy of the genus Ceci- holm) and Dr J. Yukawa (Kagoshima) for lend- J. domyia (Diptera: Cecidomyiidae) with special ref- ing material of C. pini and C. japonica. erence to the species occurring on Pinus banksiana Lamb. — Canadian Entomologist 92: 65— 79.

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Text List of references

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