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DEEL 122

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD VAN DEEL 122

ACHTERBERG, C. van. — A revision of the species of Amicrocentrinae, a new sub-

family (Hymenoptera, Braconidae), with a description of the final larval instar of

Amicrocentrum curvinervis by J.R.T.Short ........................

ACHTERBERG, C. VAN. — A revision of the new subfamily Xyphozelinae (Hymeno-

pterasBraconidae) era ae tte n ein

ACHTERBERG, C. VAN. — A revision of the subfamily Zelinae auct. (Hymenoptera,

BA CONICA) teenie neh A N nr abs a

KLOFT, E. S., zie Kloft, W. J., etc. i

KLOFT, W. J., R. E. WOODRUFF & E. S. KLOFT. — Formica integra (Hymenoptera,

Formicidae) IV. Exchange of food and trichome secretions between worker ants

and the inquiline beetle, Cremastocheilus castaneus (Coleoptera, Scarabaeidae) . .

LITH, J. P. VAN. — The New World genus Pluto (Hymenoptera, Sphecidae, Psenini) . .

SAVTSHENKO, E. N. — Phylogenie und Systematik der Tipulidae (Translated and revis-

ed by B. Theowald en G. Theischinger) ..............:...........

WILKINSON, C. — A taxonomic study of the micro-lepidopteran genera Microcalyptris

Braun and Fomoria Beirne occurring in the United States of A merica (Lepidopte-

ramNepticulidae))e)ne wr. eZ Nari Rn RR E I

WOODRUFF, R. E., zie Kloft, W. J., etc.

1979

DEEL 39

ENTOMOLOGISCHE BERICHTEN

UITGEGEVEN DOOR

DE NEDERLANDSE

ENTOMOLOGISCHE VERENIGING

INHOUD

ACHTERBERG, C. VAN, Note on the homonymy of Macrocentrus brevicaudis Abdin-

bekova (Hymenoptera, Braconidae, Macrocentrinae) .................

ALDERS, K., Een kweek van Arctornis l-nigrum (Müller) (Lep., Lymantriidae) .....

ASSELBERGS, J. E. F., Depressaria emeritella Stainton, nieuw voor de fauna van Ne-

derland (ep? Oecophoridae) ere ee Ce ER eae en

BINK, F. A., Methods for mounting Aleyrodidae specimens

BLOM, W. L., Descriptions of new butterflies from Iran

——, New butterflies from Iran (Lep., Rhopalocera)

BOLLAND, H. R. Zie J. GUTIERREZ

BOSMAN, B. T., Hinderlijke of schadelijke mijten en insekten in en om gebouwen in

VE A ERA ae AIAR NE

BOTOSANEANU, L., Sur une nouvelle espèce d’Uenoa de l’Himalaya, et sur la remar-

quable manière dont les femelles protègent leur pont (Trichoptera: Uenoidae)

BRUGGE, BEN, Coleoptera verzameld van iepenschors

BUND C.F. VAN DE. Zie G. VAN ROSSEM

BURGE?, H.C. Zie G. VAN ROSSEM

COBBEN, R. H., A new Adarrus species from Austria (Cicadellidae, Homoptera Auche-

norrhyncha); i; loi Sn AE ROLO CI

EVENHUIS, H°H.,Vliegenuitmestvannertsen(Diptera) Re

FRANKEN !UYZEN, A. VAN, Waarnemingen aan Ectoedemia argentipedella (Zeller),

eenimineermotopiberki(lEep:SNepticulidae) re

FRANKENHUYZEN, A. VAN, en J. M. FRERIKS, Tischeria ekebladella (Bjerkander,

1795) (Lepidoptera, Tischeriidae)

1979

141

173

124

129

FRANKENHUYZEN, A. VAN, en TONNY WIJNEN, Een nieuwe vangmethode voor

Synanthedon myopaeformis (Borkhausen) (Lep., Sesiidae) ..............

FRERIKS, M. Zie A. VAN FRANKENHUYZEN

GIELIS, G., Capperia hellenica Adamczewski nieuw voor Spanje (Lep., Pterophoridae)

GOFFAU, L.J. W. DE. Zie G. VAN ROSSEM

GOOT, V. S. VAN DER, Enkele zweefvliegsoorten van Nederland (Dipt., Syrphidae) . .

GOUTBEEK, A., Een teratologische meikever (Melolontha melolontha (Linnaeus) . . .

GUTIERREZ, J., W. HELLE et H. R. BOLLAND, Etude d'une souche de Tetranychus

pierci (Acariens: Tetranychidae), d’Indonésie: redescription, caryotype et repro-

GUCHO TBE SLOT, SOR Seer ERE ee en

HARTEN, A. VAN, Sitobion hillerislambersi, a remarkable new aphid from Angola (Ho-

mopterawAphididae) wer ef AAA ee veren Gee ce oh à ca OU ju ran

HELLE, W. Zie J. GUTIERREZ

HEPPNER, JOHN B., Brachodidae, A new Family Name for Atychiidae (Lepidoptera:

Sestoidea) Er NEE N cee ele ee

JEEKEL, C. A. W., Duizend- en miljoenpoten uit Noordoost-Nederland. Aanvullende

EWEN OTIS) MA I MN, EEN ONRI ere MD een e LU, Mo

KANAAR, P., Naamlijst van de in Nederland en het omliggende gebied voorkomende

listenidae (Coleoptera) er aes VEN EEE. ern Gey ANT

——, Praktische wenken voor de studie van de Histeridae (Coleoptera) ..........

——, Notities over Nederlandse Histeridae (Coleoptera) ..................

KRIKKEN, J., Suralcis Machatschke, a remarkable genus of African ruteline chafers . .

LANGOHR, G. R., Nieuwe en minder gewone Lepidoptera voor de Nederlandse fauna

LEMPKE, B. J., A new form of Hymenia recurvalis (Fabricius) from ‚ne Canary Islands

BepmByralidae) a, Zaren. tod et Pee Aw soca ee ue Pl CU Le

MILK MING SMO OM, situa NE. heten MORENTE RATA TO OA LOLA RA

— —, Pyrausta cingulata (Linnaeus) en enkele opmerkingen over haar verwant Pyrausta

rectefascialis Toll (Lep., Pyralidae) ............................

— —, Interessante vormen van Lepidoptera uit Nederland ..................

—_ —, Tinea turicensis Müller-Rutz, nieuw voor de Nederlandse fauna ...........

LITH, J. P. VAN, Notes on palaearctic Psenini IX-XIII(Hymenoptera, Sphecidae)

NIESER, N., A new Hesperocorixa and notes on Parasigara from Portugal ........

NIEUKERKEN, E. J. VAN, De verspreiding van Hydrovatus cuspidatus (Kunze) in Ne-

derlandi(Coleoptera: Dytiscidae)! on Mate de en een ee et

— —, Faunistische notities over enkele soorten van het genus Hydroporus Clairville in

Nederland (Coleoptera: Dytiscidae) ...................,.,......

NIJVELDT, W., The synonymy of Rabdophaga clavifex (Kieffers) (Diptera, Cecidomyii-

CET) in I IRE te TE Ne

OOSTERHOUT, F. J. VAN, Vondsten van rupsen van Anticlea badiata (Denis & Schif-

fermüller) en Anticlea derivata (Denis & Schiffermüller) (Lep., Geometridae)

OOSTSTROOM, S. J. VAN, Voor Nederland nieuwe en minder algemene bladwespen

(Hym., Symphyta) (Mededelingen overSymphytano.9) ...............

PRESA, J. J., Mioscirtus wagneri maghrebi Fernandes in the Iberian Peninsula .....

RAMAKERS, P. M. J., Verdere gegevens over Chrysodeixis chalcites (Esper) (Lep.,

INOGtUIdale) parecer the ena II TER De tS

ROSSEM, G. VAN, C. F. VAN DE BUND, H. C. BURGER en L. J. W. DE GOFFAU,

Bijzondere aantastingen door insekten in 1978 .....................

SINGH, ASKET, A new species of Azarea Warov, 1926 from North West India (Ortho-

ptera, Acrididae, Gomphocerinae) ............................

TIMMER, JAN, Naamlijst van de in Nederland voorkomende soorten dazen (Dipter

Mabanid ae) le aaa er A eN EAD ne, Ge al aia de ue En dere. a

VECHT, J. VAN DER, Notes on Vespidae fromNepal ....................

VELTMAN, A., Zweefvliegen van het Amsterdamse Bos (Dipt., Syrphidae) .......

113

145

121

103

161

150

179

189

148

136

WILLEMSE, FER, Additional notes on the Sexavae of the Melanesian Subregion (Or-

thoptera, Tettigonidae, Mecopodinae) . .........................

——, A new species and new distributional data of Oropodisma Uvarov, 1942 from Gree-

ce(Orthoptera, Acrididae) Ri Was es RE

——, Stenobrothus (Stenobrotodes) spec. nov. from Greece (Orthoptera, Acrididae) . .

WIJNEN, TONNY. Zie A. VAN FRANKENHUYZEN

KORTE MEDEDELINGEN

BOUWER, ROBERT, Elateridae : 214240 Sige ee ana oe tgs cle anys el Ce

COBBEN, R. H., Macropterie bij oppervlaktewantsen ....................

EUPEDO; F', Te.koop;gevraagd ori

GOOT, V. S.VAN DER, Eristalis pertinax (Scopoli) en E. tenax (Linnaeus) (Dipt., Syr-

phidae) 5 ee A NEN RN

——, Brachypalpus laphriformis (Fallen) (Dipt., Syrphidae) en de Amerikaanse vogel-

kers(of,.bospest:’) (Prunus:serotina Ehrhart) o ee

——, Eristalis tenax (Linnaeus) en Episyrphus balteatus (De Geer) (Dipt., Syrphidae) elk

jaar als trekker vanuitihetzuiden? 016. be See

——, Eieren leggend 9 van Machimus atricapillus (Fallen) (Diptera, Asilidae) .....

——, Chrysochroma bipunctatum (Scopoli) (Dipt., Stratiomyiidae) weer in Nederland

Waärgenomen 1. u... se IO ae SO OR aE RS ae

——, Een goede vangplaats:Apeldoorn, omgeving Echoput en Aardhuis (Dipt., Syrphi-

dae, Conopidae, Asilidae en Coenomyüdae) "HE ne.

—, Over de talrijkheid van de Sphaerophoria-soorten van de ,,menthastri-groep”

(Dipt Syrphidae) u... ac ae I RC EEE

-—=, Spaerophoria.abbreviata Zetterstedt "2/22 a ee ee

JANSE,K.P.).;Aangeboden:‘.. … 5. An Aus tkn MIRO I et ee ie

LEMPKE, B. J., Eupithecia-rupsen op Lamsoor (Statice vulgare Mill.) ..........

——, Komt Aterpia corticana (Denis & Schiffermüller) wel in Nederland voor? (Lep.,

Tortricidae): Er de ed ENI RARI RO REA

——, De voedselplant van Pammene luedersiana (Sorhagen) (Lep., Tortricidae). . . ..

——, Hoe staat het met Lymantria dispar (Linnaeus) in Nederland? (Lep., Lymantriidae)

OUDE, J. E.DE, Nederlandse vertalingen van J. H. Fabre’s publicaties ..........

VELTMAN, A., Metasyrphus lundbecki (Soot Ryen) (Dipt., Syrphidae) . .........

WOLSCHRIJN, J. B., Nemapogon cloacella (Haworth) (Lep., Tineidae) .........

LITERATUUR

BLOMMERS, L. H. M., Proceedings of the 8th international congress of the Inter-

national Union for the study of Social Insects ......................

BOER, ?.J. DEN, Thiele, H. U., 1977. Carabid beetles in their environment .......

DIAKONOFF, A., A. K. Kuznetsov, A. A. Stokolnikov, I. L. Sukhareva & M. I. Falko-

vitch, 1978: Vepidoptera,partsl® . a 20 2 os serie ee EE IN

——,; Gozmany, L1978 Lecithoceridae : CRON oe eas Ce eee cue

——, Roe: -r, R. Ulrich & Peter V. Küppers, 1979. Beiträge zur Kenntnis der Insecten-

faunaSumatras re eee AAE EEn:

DUFFELS, J. P., Kinzelbach, Ragnar K., 1978. Strepsiptera . ................

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’ ELLIS, W.N., Schreiber, H., 1978. Centres of Sphingidae (Lepidoptera in the neotropi-

ELI STOTT 62 RE NER NE EN

——, Role, The, of arthropods in forest ecosystems ......................

——, Insects and other Arthropods of medical importance .................

——, Larsen, Sven Gisle, 1978. Baltic amber — a palaeobiological study .........

——, Proceedings of the second international symposium on Trichoptera.........

——, Merian, Maria Sibylla, 1976. Schmetterlinge, Käfer und andere Insekten; Lenin-

gradenStudienDuche ya Mae e ie

— ssBiosystematicsin.agrieulturen. one sl ee

——, Matthews, R. W. & J. R. Matthews, 1978. Insect behavior ..............

——, Evolution of insect migration and diapause, 1978 ....................

——, Endocrinologie (J. Lever & J. de Wilde, eds) .................,.....

——, Baker, R. R., 1978. The evolution ecology of animal migration............

MICA LOMA IMENtOMOlOPIE <a an. nenn fo sec lene) wala a sut à

EVENHUIS, H. H., Sluipwespen in relatie tot hun gastheer, Klomp H. en G. T. Wiebes

(GEHAG EISA NEDA A sec RSR An es tee Su RE

EYNDHOVEN, G. L. VAN, Van Bronswijk, J. E. M. H., Rijntjes, R. H., Garben, A. F.

M., Vos, H.; 1979. De teken (Ixodidae) van de Benelux-landen ...........

——, Davids, C., De watermijten (Hydrachnellae) van Nederland, Levenswijze en voor-

ECA ee e glo A tary age

GEEST, L. P. S. VAN DER, Biochemistry of insects .....................

GRAVESTEIN, W. H., Smit, F. G. A. M., 1978. Insects on stamps — a cross-referenced

CHE CRUS tere abos RESORT PRI CORO RIA LION Karl AE à

GUNST, J. H. DE, Klausnitzer, B., 1978. Ordnung Coleoptera (Larsen) ..........

HELSDINGEN,P.J. VAN, Savory, T., 1977. Arachnida ......... . ,. .. .. . ..

KREBS, B., Moller Pillot, H. K. M., 1978. De larven der Nederlandse Chironomidae

(Diptera) EEE A N EE ARI OE O io,

LAAN, P. A. VAN DER, te Pascal P., 1978. Diseases and pests of ornamental plants

——, Nayar, K. K., N. Ananthakrishnan & B. V. David, 1976. General and applied ento-

NOLO RV ENE RENE LEN LA es eee ee € ne waged eus id dia ard

LEMPKE, B. J., Traugott-Olsen, E. & E. Schmidt Nielsen, 1977. The Elachistidae (Lepi-

doptera) of Fennoscandia and Denmark .........................

——, Gepp, J. & M., 1977. Entomologia austriaca 1970-1974 ..:.............

—— , Russwurm, A. D., 1978. Aberrations of British butterflies ...............

——, Atlas provisoire des insectes de Belgique ........................

es Malicky.H., Entomologiein Österreich 4 ste. nn

——, Jeekel, C. A. W., 1978. Voorlopige atlas van de verspreiding der Nederlandse mil-

jJoenpotenDiplopoda)i, B. Ae TITLE AL er ade rt tenet Have

— —, Rougeot, P. C. & P. Viette, Guide des papillons nocturnes d'Europe et d'Afrique

GAUMONT 88.0 BNN NEN A AAR UGA de leen laeta

——, Kruseman, G., De kakkerlakken en bidsprinkhanen — Dictyoptera — uit de lan-

td enkvanidesBieneluxs nina ale dn du ene one à

MANTEL, W. P., Schliephake, G. & Klimt, K. H., 1979. Thysanoptera, Fransenflügler .

SCHULTEN, G. G. M., Coppel, H.C. & J. W. Mertins, Biological insect pest suppres-

HON: AE BE OPEL ENNE ted

——, The genetics and biology of Drosophila. Vol. 2a ....................

THEOWALD, Barendregt, A., 1978. Zweefvliegentabel ...................

——, C.R. Osten Sacken, 1903. Record of my life-work in entomology ..........

191

175

191

CONTENTS OF VOLUME 26 (1979)

ADESIYUN, A. A. Effects of intercrop on frit fly, Oscinella frit, oviposition and larval

survival on,oats pr ace RIINA RI

ALDRICH, J. R., BLUM, M. S., LLOYD, H. A., EVANS, P. H. & BURKHARD, D. R.

Novel exocrine secretions from two species of scentless plant bugs (Hemiptera:

R'hopalidae) 2 Haan REANO

ANDRE, H. & LEBRUN, Ph. Quantitative comparison of the funnel brushing methods

for extracting corticolous micro-arthropods .......................

BAI, M. K. & PRASAD, R. S. Role of dietary components in vitellogenesis in the rat

fleas Xenopsylla cheopsis and X. astia (Siphonaptera) ..................

BARBOSA, P., GREENBLATT, J., WITHERS, W., CRANSHAW, W. & HARRING-

TON, E. A. Host-plant preferences and their induction in larvae of the gypsy moth,

EEVIMANUF IG GISDOP me ne oe As a OI gee ET MC aE

BELTON, P. & COSTELLO, R. A. Flight sounds of the females of some mosquitoes of

WiesterniGanadaycn: esn koelde tne et tere whee cts, Con PROS ee

BERNAYS, E. A. The use of doppler actographs to measure locomotor activity in locust

NY Mp Hse Verh gets aera Aa Ciel Led Ag ORD I NE dyna ae wie MERE ee:

COCHRAN, D. G. A genetic determination of insemination frequency and sperm prece-

dence inthe. German CocKroach ew... eee ER Een ee

CREMA, R. Egg viability and sex determination in Megoura viciae (Homoptera: Aphidi-

(EV) ea ae 2 aa anny bate ISIN DIO I I AI RE E agua

DADD, R. H. & KLEINJAN, J. E. Vitamin E, ascorbyl palmitate, or propyl gallate pro-

tect arachidonic acid in synthetic diets for mosquitoes ................

DE LOOF, A., VAN LOON, J. & HADERMANN, F. Effects of juvenile hormone I, me-

thoprene and kinoprene on development of the hymenopteran parasitoid Nasonia

vitkipennis SOT onset ah Ok M IE A I AE nes EI

DOBIE, P., GREVE, J. VAN S., HOTHI, K. & KILMINSTER, A. M. Inability of storage

Bruchidae to infest winged beans (Psophocarpus tetragonolobus) ...........

FISHER, J. R. & O'KEEFFE, L. E. Seasonal migration and flight of the pea leaf weevil,

Sitona lineatus (Coleoptera; Curculionidae) in Northern Idaho and Eastern Was-

hington #73 4 Batse VIA RE RIE CORP RR eR

GREANY, P. D. & SZENTESI, A. Oviposition behavior of laboratory-reared and wild

Caribbean Fruit Flies (Anastrepha suspensa; Diptera: Tephritidae): II. Selected phy-

sical influenCes CERES IR

GRENIER, ©. Développement embryonnaire in vitro, en milieu artificiel défini de deux

parasite des ovolarvipares, Phryxe caudata et Luxophaga diatraeae (Diptera, Tachi-

nidae). — In vitro embryogenesis, on defined artificial media, of two ovolarviparous pa-

rasitoids, Phryxe caudata and Lixophaga diatraeae (Diptera, Tachinidae) ......

Entomologia

experimentalis

et applicata

208

323

GRIFFITHS, E. & WRATTEN, S. D. Intra- and inter-specific differences in cereal aphid

low-temperaturetolerance ..!.. 22.20.20 eee

HAYNES, J. W., WRIGHT, J. E. & MATTIX, E. Fractionated vs acute irradiation: the

effects of treating adult boll weevils (Coleoptera: Curculionidae) at different ages

HOY, M. A. Parahaploidy of the ,,arrhenotokous” predator, Metaseiulus occidentalis

(Acarina: Phytoseiidae) demonstrated by X-irradiation of males ...........

HOY, M. A. & SMILANICK, J. M. A sex pheromone produced by immature and adult

females of the predatory mite, Metaseiulus occidentalis (Acarina: Phytoseiidae)

JAENSON, T. G. T. Mating behaviour of Glossina pallidipes (Diptera, Glossinidae): dura-

tion of copulation, insemination and fecundity .....................

JOOSSE, E. N. G. & KOELMAN, T. A. C. M. Evidence for the presence of aggregation

pheromones in Onychiurus armatus (Collembola), a pest insect in sugar beet .

JORDENS-ROTTGER, D. The role of phenolic substances for host-selection behaviour

of the black bean-aphid, Aphis fabae ...........................

KAWADA, K. & TAMOTSU, M. Apterous males and holocyclic reproduction of Lipa-

Dinsterysimiinlapanı, nnee ee ee een een re ele seen

KENNEDY, G. G. & YAMAMOTO, R. T. A toxic factor causing resistance in a wild to-

mato to the tobacco hornworm and some other insects ................

LEWIS, A. C. Feeding preference for diseased and wilted sunflower in the grasshopper,

Melanoplussdifferentialis ete

LINLEY, J. R. Activity and motility of spermatozoa of Culicoides melleus (Diptera: Cera-

VOPSTOnIdae)r God ae ome HO ee en en erste:

LLEWELLYN, M. & EIVAZ, J. Abrasive dusts as a mechanism for aphid control ....

LLEWELLYN, M. & QURESHI, A. L. The energetics of Megoura viciae reared on differ-

ent parts of the broad bean plant (Viciafaba) ......................

MAGYAROSY, A. C., SINGH, P. & MITTLER, T. E. Rearing of the aphid Myzus persi-

cae on phloem exudate from Amsinckia douglasiana plants infected with curly top

ACLS so oe as hci Kiet rece nt

MONTGOMERY, M. E. & NIELSEN, D. G. Embryonic development of Otiorhynchus

sulcatus: effect of temperature and humidity ......................

NIELSEN, J. K., LARSEN, L. M. & SORENSEN, H. Host plant selection of the horsera-

dish flea beetle Phyllotreta armoraciae (Coleoptera: Chrysomelidae): identification

of two flavonol glycosides stimulating feeding in combination with glucosinolates

OGWARO, K. Seasonal activity of the sorghum shootfly Atherigona soccata (Diptera: An-

OTA) RA seth celeste ce ot Fuss pee ute i ett my e

OMER, S. M. Responses of females of Anopheles arabiensis and Culex pipiens fatigans to

air currents, carbon dioxide and human hands in a flighttunnel ...........

OLOO, G. W. & LEUTHOLD, R. H. The influence of food on trail-laying and recruit-

ment behaviour in Trinervitermes bettonianus (Termitidae: Nasutitermitinae)

PALANISWAMY, P., SEABROOK, W. D. & SIVASUBRAMANIAN, P. Effect of a ju-

venile hormone analogue on olfactory sensitivity of Eastern spruce budworm, Cho-

ristoneura fumiferana (Lepidoptera: Tortricidae) ....................

ROSE, D. J. W. & DEWHURST, C. F. The African Armyworm, Spodoptera exempta —

Congregation of moths in trees beforeflight .......................

RUMBO, E. R. Oxygen requirements of Lucilia cuprina during development within the

PUPIRUMEEREOS ee LS nn een

SHERLOCK, P. L. A method for mass-rearing the cutworm Agrotis segetum in the labo-

POM) og 4-0 0 50 à big o o 0 010 0/00 0 00010 D ee > 0 © D 0.0

SHOUKRY, A. & HAFEZ, M. Studies on the biology of the Mediterranean fruit fly Ce-

HALILISICADUOIA EN EN PIE REI CCC RO

SIVAPALAN, P. & GNANAPRAGASAM, N. C. Effects of varying proportions of dieta-

ry ingredients in meridic diets on the development of the tea tortrix, Homona cof-

Neanianınithellaboratonye ne... ee aie CCE + ae

161

142

267

175

346

SZENTESI, A., GREANY, P. D. & CHAMBERS, D. L. Oviposition behavior of labora-

tory-reared and wild Caribbean Fruit Flies (Anastrepha suspensa; Diptera: Tephriti-

dae): I. Selected chemicaliinfluencesy E

THIBOUT, E. Stimulation of reproductive activity of females of Acropeliopsis assectella

(Lepidoptera: Hyponomeutidae) by the presence of eupyrene spermatozoa in the

spermatheca.-...0.. a2. ra NRE ER

WHITE, P. F. Pot tests with methoprene and permethrin against the Mushroom Phorid

(Megaselia halterata) and the Mushroom Sciarid (Lycoriella auripila) ........

WOOL, D. & BERGERSON, O. Sperm precedence in repeated mating of adults of Tri-

bolium castaneum (Coleoptera, Tenebrionidae) ................... re

YOSHIDA, S. & HIDAKA, T. Determination of the position of courtship display of the

young unmated male Anisopteromalus calandriae (Hymenoptera, Pteromalidae) . .

YOSHIHARA, T., SOGAWA, K., PATHAK, M. D., JULIANO, B. O. & SAKAMURA,

S. Soluble silicic acid as a sucking inhibitory substance in rice against the Brown

Plant Hopper (Delphacidae, Homoptera)

“45. COMP. ZOOL,

LIBRARY

OCT 1 7 1980

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TIJDSCHRIFT

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UITGEGEVEN DOOR

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Tijdschrift voor Entomologie, deel 122, 1979

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INHOUD VAN DEEL 122

Achterberg, C. van. — A revision of the species of Amicrocentrinae, a new subfamily

(Hymenoptera, Braconidae), with a description of the final larval instar of

Amicrocentrum curvinervisbyJ.R.T.Short .......................

Achterberg, C. van. — A revision of the new subfamily Xyphozelinae (Hymenoptera,

Braconmidae) Wie cars OEE EL RI RR RE

Achterberg, C. van. — A revision of the subfamily Zelinae auct. (Hymenoptera, Braconidae)

Kloft, E. S., zie Kloft, W. J., etc.

Kloft, W. J., R. E. Woodruff & E. S. Kloft. — Formica integra (Hymenoptera, Formicidae) IV.

Exchange of food and trichome secretions between worker ants and the inquiline

beetle, Cremastocheilus castaneus (Coleoptera, Scarabaeidae) ............

Lith, J. P. van. — The New World genus Pluto (Hymenoptera, Sphecidae, Psenini)........

Savtshenko, E. N. — Phylogenie und Systematik der Tipulidae (Translated and revised by B.

MheowaldieniG-lheischingen as sss NM oe ne oe ee

Wilkinson, C. — A taxonomic study of the micro-lepidopteran genera Microcalyptris Braun and

Fomoria Beirne occurring in the United States of America (Lepidoptera,

Neen NAE) SE a cary mA ae cera ce hie Not: Sony nt aul, se

Woodruff, R. E., zie Kloft, W. J., etc.

127

MUS. COMP, ZOOL.

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9 LIBRARY

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UNIVERSITY

ii TIJDSCHRIFT

| VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

C. VAN ACHTERBERG. — A revision of the species of Amicrocentrinae, a new

subfamily (Hymenoptera, Braconidae), with a description of the final larval

’ instar of Amicrocentrum curvinervis by J. R. T. Short, p. 1—28, text-figs. 1—68.

€. VAN ACHTERBERG. — A revision of the new subfamily Xyphozelinae

_ (Hymenoptera, Braconidae), p. 29—46, text-figs. 1—45.

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A REVISION OF THE SPECIES OF AMICROCENTRINAE,

A NEW SUBFAMILY (HYMENOPTERA, BRACONIDAE),

WITH A DESCRIPTION OF THE FINAL LARVAL INSTAR

OF AMICROCENTRUM CURVINERVIS BY J. R. T. SHORT

C. VAN ACHTERBERG

Rijksmuseum var Natuurlijke Historie, Leiden

With 68 text-figures

ABSTRACT

A new subfamily is erected for the Afrotropical genus Amicrocentrum Schulz, 1911, hitherto included

in the Macrocentrinae. A key to the subgenera and species is given for the first time and the species are

redescribed. One new species, Amicrocentrum exilis, and one new subgenus, Platyxanion, are described.

The final larval instar of Amicrocentrum curvinervis (Cameron) is described.

INTRODUCTION

When revising the subfamily Macrocentrinae (the genus Macrocentrus s.s.

excluded) I had to deal with the aberrant genus Amicrocentrum Schulz, 1911. The

species are very conspicuous, large, mostly yellow or light brown in colour and

have many apomorphous (derived) character-states. Revision was difficult since

there was little material in most collections. Detailed study showed that

Amicrocentrum could not be retained in the Macrocentrinae because of the lack of

synapomorphous character-states. The characters of Amicrocentrum differ from

those of other subfamilies of Braconidae; a new subfamily is therefore erected for

the genus. The distribution of Amicrocentrum appears to be restricted to the

Afrotropical Region. An aberrant species which occurs in Malagasy is placed in a

new subgenus.

The small amount of relevant literature is listed in Shenefelt (1969: 141).

TERMINOLOGY

General terminology is given by Van Achterberg (1976a: 160—166). The name

of the postero-basal lobe of the hind wing needs explanation. Hamilton (1971: 429)

and Brothers (1975: 520) have pointed out that the use of the name anal or vannal

lobe of most authors is incorrect. This lobe is bordered anteriorly by a non-

functional fold (the plical furrow or fold, a “paleopterous fluting” according to

Hamilton (1971: 432)) and should therefore be called a plical lobe (Brothers, 1975:

D) TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

520). The main difference between an anal (vannal) fold and a plical fold is in its

position. The plical fold is situated in the submediellan cell (M + Cu in fig. 32 of

Brothers) and goes through the nervellus. The anal (vannal) fold is situated well

below the nervellus and the submediellan cell and functions in the wing-folding

process.

PHYLOGENY

The genus Amicrocentrum shows the following remarkable apomorphous

character-states:

1. The very large plical lobe is perpendicularly setose (fig. 8). The perpendicular

setosity of the lobe is unique in Braconidae, and in Hymenoptera as far as I know,

and is without doubt an apomorphous character-state. The well developed plical

lobe of Symphyta is a plesiomorphous character-state.

2. A medial basal hole is present in the first metasomal tergite. This depression

should not be confused with the much shallower depression which is situated more

basally where the adductor is attached (fig. 17). This hole is a unique development

in Hymenoptera and an apomorphous character-state within the order.

3. Deep depressions are present in the first and second tergites of the metasoma

of the males and these are more or less covered by fatty secretions between the

pubescence (fig. 15). I am not aware of such structures in other Hymenoptera,

apart from the males of Aleiodes excavatus (Telenga) (Braconidae, Rogadinae)

(Van Achterberg, 1975: 16, fig. 1) and the males of the striatula-group of

Parischnogaster Schulthess (Vespidae, Stenogastrinae) (Van der Vecht, pers.

comm.). The functions of these depressions and secretions are uncertain, but the

structures are an independently acquired apomorphous character-state in

Hymenoptera.

4. The maxillary and labial palpi are strongly reduced and both virtually one-

segmented (fig. 23). Reduction of the palpi is quite a common tendency in the

Braconidae (Van Achterberg, 1976b: 35), but the condition reached in

Amicrocentrum is extreme and a strongly apomorphous character-state. In

contrast, the palpi of Macrocentrinae are well developed and consist of 5 or 6 and

4 segments, respectively.

5. The prepectal, hypostomal and occipital carinae are absent. The presence of

these carinae is generally accepted as a plesiomorphous character-state. These

carinae are present in many groups of Hymenoptera which are not closely related

and have many other plesiomorphous character-states in common. In the

Macrocentrinae the occipital carina is absent but the prepectal and an

carinae are always present.

6. The dorsal carinae of the first tergite are absent (fig. 29). As shown by several

groups of Braconidae with other plesiomorphous character-states, the presence of

at least short basal dorsal carinae must be considered a plesiomorphous character-

state within the Braconidae.

7. The pedicellus is cup-shaped (figs. 28, 33). I have not seen this peculiar shape

in other Ichneumonoidea. It seems to have been derived from the common

cylindrical shape.

VAN ACHTERBERG: Amicrocentrinae 3

8. The second tergite of the metasoma is less setose than the posterior half of

the third tergite. Reduction in setosity is a common feature, but the pattern in

Amicrocentrum is remarkable and apomorphous.

9. The fore tibial spur is bare, stout, curved, rather wide and flattened (figs.

57—59). In Hymenoptera a slender, more or less cylindrical, rather narrow spur

with a narrow flange at the inner side is the common plesiomorphous character-

state.

10. The hind tibia and tarsus are long, the length of the hind tibia being 1.9— 2.4

times the hind femur (figs. 34, 56). This is a peculiar feature which is exceptional in

Hymenoptera. In Braconidae the hind tibia is generally 10—35% longer than the

femur.

It is striking that almost no synapomorphous character-states exist between the

Macrocentrinae and the genus Amicrocentrum. The most important apomorphous

character-states of the Macrocentrinae contain that the trochantelli are apically

toothed, the claws simple or with a lobe, the occipital carina absent and the n. rec.

far antefurcal. The shared character of reduction of the occipital carina is

insufficient for retaining Amicrocentrum in the Macrocentrinae.

Plesiomorphous character-states of Amicrocentrum are:

1. The n. rec. is postfurcal. In Hymenoptera there is a general trend towards

reduction of the apical veins with the veins becoming situated more basally. In this

perspective a postfurcal n. rec. is a plesiomorphous character-state.

2. The first transverse anal vein is present. This vein is weakly developed, but its

presence indicates a more complex and plesiomorphous state of venation.

3. A laterope is present. In the Ichneumonoidea this character is common in

groups which show several other plesiomorphous character-states. It is most likely

an early development and should be considered a plesiomorphous character-state

within the Ichneumonoidea.

4. The claws are bifurcate (fig. 13). As pointed out by Brothers (1975: 521)

simple bifurcate claws may be considered a plesiomorphous character-state in

Hymenoptera.

5. The costa and subcosta are more or less separated from each other (fig. 8). In

parasitic Hymenoptera there is usually no space between these veins or, at most, a

small space apically. Because there is a well-developed cell between these veins in

Symphyta, the presence of a narrow cell in the Amicrocentrinae is a

plesiomorphous character-state within the Braconidae.

6. The mesoscutum is without a lateral carina. A well developed lateral carina

ofthe mesoscutum seems to be a later development in the Braconidae.

7. The plical lobe is very large, as in the Symphyta.

The following are the more doubtful character-states of Amicrocentrum:

1. The ovipositor is long. An ovipositor as long as the forewing or somewhat

shorter is often associated with other plesiomorphous character-states in the

Braconidae. But the very long ovipositor of Amicrocentrum and some

Macrocentrinae may be a secondary (apomorphous) development. The ovipositor

of A. seyrigi is intermediate in length.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

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2. The metasoma is inserted above the hind coxae. This is probably a

plesiomorphous character-state, apart from the extreme conditions in the

Cenocoeliinae and Evanioidea. It occurs in several subfamilies of the Braconidae

which are not closely related (e.g., Helconinae, Macrocentrinae, Orgilinae,

Agathidinae) as well as in the Ichneumonidae (subfamily Labeninae (= Labiinae

sensu Townes)). |

3. The radiellan cell of the hind wing is somewhat widened apically. This is

probably a plesiomorphous character-state, if shown in a moderate manner as in

the Amicrocentrinae.

The Amicrocentrinae are an isolated and specialized group. As shown above, it

has little in common with the Macrocentrinae. It may be related to the Helconinae

sl. The genus Brulleia Szepligeti, in particular, shares some characters with the

Amicrocentrinae, such as the large plical lobe, the dilated radiellan cell, the short

second abscissa of the subcostella and the long nervellus. The tribe Trachypetini

of the Sigalphinae also shows some resemblance to the Amicrocentrinae, but the

nervellus is broken apically, the shape of the first metasomal tergite is quite

different, the occipital and prepectal carinae are present and the absence of well-

defined synapomorphous character-states indicate that the relationship is

superficial. The Amicrocentrinae may be treated provisionally as an early offshoot

of the Helconinae s.l. This concurs with the evidence of larval characters (figs.

2—5).

Table | gives the occurrence of the plesiomorphous and apomorphous states of

some characters within the genus Amicrocentrum. These indicate the pattern of

evolution within the genus. One of the two species which occur in Malagasy,

A. seyrigi, is isolated in showing a peculiar combination of apomorphous and

plesiomorphous character-states. A new subgenus is therefore erected for this

species. Of the remaining four species, A. flavipenne is also restricted to Malagasy.

This species is closely related to exilis and has probably developed from an

invasion of an exilis-like ancestor. Three species occur only in continental Africa.

Of these, concolor has the most plesiomorphous character-states and curvinervis,

which is relatively common, has an intermediate position. A. exilis is rather

specialized and shares some apomorphous character-states with flavipenne, such as

the flattened clypeus and the slender first tergite (figs. 49, 68). These characters

are absent in other species and indicate the close relationship shown in fig. 1.

BIOLOGY

At least two species of Amicrocentrum are larval parasites of large, boring

caterpillars of Lepidoptera. Amicrocentrum exilis spec. nov. has been reared from

Eulophonotus myrmeleon Felder (Cossidae, Cossinae). A. curvinervis (Cameron) has

been reared from the maize stalk borer Busseola fusca (Fuller) (Noctuidae) and

may therefore be of economic importance. The selection of boring hosts is a

further plesiomorphous character-state, since braconids are considered to be

derived from ectoparasites of boring coleopterous larvae.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

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VAN ACHTERBERG: Amicrocentrinae 7

LARVAL CHARACTERS OF AMICROCENTRUM CURVINERVIS (figs. 2—5)

(By J. R. T. Short, Department of Zoology, Australian National University,

Canberra, Australia). !)

The methods used in making slide preparations from final larval instar exuviae

are described in Short (1978: 4). Terminology, and its basis in comparative

morphology, is given in Short (1952). Material studied: | male final instar larva of

Amicrocentrum curvinervis (Cameron), ‘“Makarere, x.1969, Uganda, Dennis

Owen”, “Ex larva B. fusca” (TC).

Description. — Of the head sclerites (fig. 2), epistoma (e) with dorsal part

unsclerotized; pleurostoma (ps) very broad; anterior pleurostomal process (ap)

hook-like; posterior pleurostomal process (pp) in the form of a narrow rod;

hypostoma (hs) with median part only sclerotized and not extending laterally

(posteriorly) beyond lateral end of stipital sclerite (ss); hypostomal spur (hsp)

represented by broad and faintly sclerotized band between hypostoma and stipital

sclerite; each stipital sclerite (ss) a slender rod, with median end fitting into socket

in antero-lateral end of labial sclerite (lbs); each cardo (cd) represented by lightly

sclerotized oval plate; labial sclerite (lbs) with lateral parts slender and sclerotized

and ventral part broad and lightly sclerotized; maxillary (mp) and labial palps (Ip)

disc-shaped and each with one large sensillum and two very small sensilla; salivary

orifice (s) prominent; silk press (sp) broad and lightly sclerotized and with two

small sensilla on dorsal part; setae and sensilla present on maxillae, labium and

clypeo-labrum; prelabial and labral sclerites absent; mandible (m) with triangular

base and long, curved, slender blade with length about twice that of base and with

prominent teeth on median half of blade. Antenna (fig. 3) disc-shaped with

circumference lightly sclerotized and with two sensilla on membrane. Spiracle (fig.

4) relatively very large, with closing apparatus (ca) adjoining atrium (ar) and

closing apparatus with prominent sclerotized bands on wall. Skin (fig. 5) with small

setae and numerous small spines.

The cocoon of Amicrocentrum curvinervis is 20 x 4.5 mm, pale stramineous,

moderately thin and somewhat translucent but not porous, and with very little

loose silk on the surface. Emergence was by transversely cutting off one end.

Systematic position of Amicrocentrum. — Amicrocentrum shows characters

resembling those of the Macrocentrinae. I know the larval characters only of

Macrocentrus of the Macrocentrinae. Amicrocentrum, like Macrocentrus has the

lateral parts of the labial sclerite slender and sclerotized and the ventral part broad

and lightly sclerotized. In both genera the salivary orifice is prominent and the silk

press broad and lightly sclerotized. The stipital sclerite is also similar. However,

Macrocentrus differs from Amicrocentrum in that the lateral part of each hypostoma

is well sclerotized and the hypostomal spur is represented by a projection on the

hypostoma (Short, 1952: fig. 29). The mandibles of these genera differ in that,

although both are slender in form and have the blade toothed, in Amicrocentrum

!) This work was supported by the Australian Research Grants Committee.

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

Figs. 2—5. Final larval instar of Amicrocentrum curvinervis (Cameron). 2, anterior view of head with

sclerites flattened, ap = anterior pleurostomal process, cd = cardo, e = epistoma, hs = hypostoma,

hsp = hypostomal spur, lbs = labial sclerite, Ip = labial palp, m = mandible, mp = maxillary palp, pp

= posterior pleurostomal process, ps = pleurostoma, s = salivary orifice, s = silk press, ss = stipital

sclerite; 3, antenna; 4, spiracle, ar = atrium, ca = closing apparatus; 5, skin. (Drawing by J. R. T.

Short).

the blade is much more slender than the base and teeth are present only on the

median half of the blade. The disc-shaped antennae differ also in that the

circumference is lightly sclerotized in Amicrocentrum but not in Macrocentrus. The

spiracle is relatively very large in Amicrocentrum and distinctive in form. In

Macrocentrus, the closing apparatus of the spiracle is only as wide as the atrium and

does not adjoin the atrium. The skin also differs in these genera in that, although

there are numerous small spines present on the skin in both, setae are present in

Amicrocentrum but not in Macrocentrus.

Amicrocentrum, although keying out generally near the Macrocentrinae (see

Capek, 1970, 1973) must be considered to stand apart from this subfamily on the

differences listed above. It is therefore recommended that, on larval characters,

Amicrocentrum should be placed in a separate subfamily of the Braconidae.

VAN ACHTERBERG: Amicrocentrinae 9

Amicrocentrum is, as far as known, a solitary endoparasite of lepidopterous

larvae. The larval characters indicate that the genus is one of the less specialized

of the endoparasitic Braconidae and show a combination of generalized and

specialized characters. The hypostoma and hypostomal spur are reduced but the

antenna, although disc-shaped, shows a sclerotized circumference. The cardo is

present, the mandible is toothed and setae are present on the skin, all of these

being primitive characters. It appears that Amicrocentrum, like the

Macrocentrinae, is an endoparasite showing only some specialized characters.

Amicrocentrinae subfam. nov.

Diagnosis. — Length of body 9.0—27.3, length of fore wing 7.4—20.9 mm;

antennal segments of © 46—59, of ¢ 44—S3; apex of scapus truncate or nearly so;

pedicellus cup-shaped, narrowed basad (fig. 33); maxillary and labial palpi

reduced, very short, only visible from ventrad and both virtually one-segmented,

but their bases somewhat wider and slightly differentiated (fig. 23); face unevenly

convex; anterior tentorial pits large and deep (fig. 16); apical margin of clypeus not

differentiated, thick, and weakly concave or almost straight medially (fig. 22);

labrum not or narrowly visible; occipital and hypostomal carinae and occipital

flange absent; mandible large, with two sharp and stout teeth, the second tooth

much shorter than the first tooth (fig. 32); eye bare; pronope absent; propleural

lamella more or less developed (fig. 30); pronotum with postero-dorsal corner

somewhat protruding as does anterior part of mesopleuron; tegulae not reaching

anterior margin of mesopleuron (fig. 30); prepectal carina completely absent (fig.

18); precoxal suture not or shallowly impressed (fig. 42); mesoscutum without a

lateral carina in front of tegulae; metapleural flange or lamella absent; dorsal

surface of propodeum not differentiated from its posterior surface (fig. 42);

propodeum without areola and tubercle; antepropodeal suture narrow and deep;

propodeal spiracle large; scutellar suture rather deep, wide and rather short (fig.

66); scutellum without lateral carina; metanotum with large convex tubercle

posteriorly (fig. 27); metanotum with long pubescence latero-dorsally (this

probably connects the plical lobe of the hind wing with the metanotum); first

discoidal cell distinctly petiolate and rather wide anteriorly (fig. 25); cuqu 2

present; r 3 more or less curved towards metacarp (figs. 8, 25); n. rec. postfurcal; B

1 strongly widened apicad; cu 1 more or less weakly sinuate (figs. 8, 37); CU 1

smaller than CU 2; d 2 more or less roundly connected to s la (figs. 8, 25); nervulus

long and straight; nervellus long, departing submedially from the mediella; B 1

closed apically, s Ib present; fringe of wings short; parastigma large; a narrow

intercostal cell is more or less developed (figs. 8, 37); distal part of mediellan cell

bare; aqu | present as a faintly-brownish, pigmented stripe (fig. 37); aqu 2 and aqu’

absent; SM mainly bare except for some setae near the nervulus; CU | and base of

aqu | mainly bare; metacarp ends near apex of the radial cell; plical lobe very

large and with rather long and perpendicular arranged setae (fig. 25); radiellan cell

widened apicad; basella and second abscissa of subcostella short (fig. 46);

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

metacarpella rather short and more or less straight (fig. 25); discoidella absent;

hind tibia and tarsus comparatively long, length of hind tibia 1.9—2.4 times hind

femur; length of femur and tibia of hind leg 5.6—7.7 and 12.0—20.3 times their

width, respectively; hind tibial spurs short, straight, and setose; hind basitarsus

without ventral row of setae; all tarsal claws bifurcate, setose and without subbasal

lobe (fig. 65); fore tibial spur bare, curved, rather wide and flattened (figs. 57—59);

hind tibia without apical spines; trochantelli simple, without teeth; metasoma

inserted medially between dorsal surface of propodeum and insertion of the hind

coxae (figs. 42, 54); length of first metasomal tergite 3.0—6.3 times its apical width;

first tergite with a medio-basal hole (figs. 29, 41); large, deep, more or less

pubescent depressions in males in the first and second tergites (fig. 15); dorsal

carinae of first tergite absent (fig. 17); laterope deep, large, elliptical (fig. 42);

dorsope absent; first tergite convex, but medially somewhat flattened; second

tergite less setose than posterior third of third tergite; third and following tergites

densely setose; second tergite without a sharp lateral crease; second and following

epipleura with thyridia (fig. 6); length of ovipositor sheaths 1.10—2.01 times length

of fore wing and slender; ovipositor straight or nearly so, with an indistinctly

developed notch subapically, this notch being absent in Amicrocentrum seyrigi;

hypopygium large (fig. 18).

Distribution. — Contains only one genus, Amicrocentrum Schulz, with five

species. Two species are restricted to Malagasy and three species only occur in

continental Africa.

Key to subgenera and species of the genus Amicrocentrum

1. Second abscissa of mediella slightly curved (fig. 8); notauli absent (fig. 11);

ocelli comparatively small, OOL distinctly longer than diameter of ocellus (fig.

14); 3rd and 4th abscissae of cubitus with only yellowish pigment and not

sclerotized (fig. 8); 3rd (at least apically, fig. 15) and following tergites felty

setose (fig. 17) (Blatyxanionsubgen.nov.) . „2... v2 seyrigi Granger

— Second abscissa of mediella strongly sinuate (fig. 25); notauli narrowly

developed (figs. 27, 40); ocelli large, OOL subequal to diameter of ocellus (fig.

20) or shorter (fig. 38); 3rd and 4th abscissae of cubitus at least weakly

sclerotized (fig. 25); 3rd and following tergites only densely setose (fig. 41)

(subgenus AmierocentrumSchulzy TORRE RR 2

2. Clypeus distinctly convex and densely punctate (figs. 22, 32); pterostigma more

stout (figs. 25, 37); postero-dorsal corner of pronotum more or less densely

punctate;sometimes rather finely (figs 85730) eN 3

— Clypeus flattened and punctulate (figs. 47, 62); pterostigma slender (figs. 46,

55); postero-dorsal corner of pronotum almost smooth, punctulate (figs. 42,

SE RER NE ER DIL DIN IRE a RR RE Cr 4

3. Fourth metasomal tergite mainly dark brown; sides of 2nd tergite diverging

apicad; length of 2nd tergite 1.4—1.8 times its maximum width (fig. 29);

metasoma of © less compressed apicad; length of ovipositor sheaths 1.24— 1.50

times fore Wing Hr RAR RN ER RE concolor (Szépligeti)

VAN ACHTERBERG: Amicrocentrinae 11

— Fourth metasomal tergite yellowish; sides of 2nd tergite parallel; length of 2nd

tergite 2.0—2.8 times its width (fig. 41), exceptionally 1.8 times; metasoma of 9

strongly compressed apicad; length of ovipositor sheaths 1.60—2.01 times fore

mime usually 1.82.9 times: sions ann. curvinervis (Cameron)

4. Metacarpella well developed basally and connected with the subcostella (fig.

46); pterostigma somewhat more slender (fig. 46); medio-basal hole of Ist

tergite rather sharp posteriorly (fig. 49); length of malar space of 9 0.3—0.5

times basal width of mandible (fig. 47) ............... exilis spec. nov.

— Metacarpella mainly reduced, not connected with the subcostella (fig. 55);

pterostigma somewhat less slender (fig. 55); medio-basal hole of Ist tergite

rounded-truncate posteriorly (fig. 64); length of malar space of 9 ca. 0.6 times

basaliwidth of mandible (fig.62)'. .....».......... flavipenne Granger

Amicrocentrum Schulz

Platyxanion subgen. nov. (figs. 6—17)

Etymology: from “‘platys” (Greek for “broad, wide, flat’’) and “xanion” (Greek for ‘‘comb’’) because

of the rather flat and wide fore tibial spur and comb, which serves as a cleaning-device (figs. 57—59).

Gender: neuter.

Type-species: Amicrocentrum seyrigi Granger, 1949.

Diagnosis. — Ocelli rather small, OOL distinctly longer than diameter of ocellus

(fig. 14); mandible not twisted apically; notauli absent (fig. 11); second abscissa of

mediella slightly curved (fig. 8); third and fourth abscissae of radius with only

yellowish pigment and not sclerotized; nervulus postfurcal or interstitial (fig. 8);

radiella straight basally; nervellus distinctly reclivous (fig. 8); glymma reduced;

third (at least apically) and following tergites felty setose (figs. 15, 17).

Distribution. — Malagasy: one species.

Note. Apomorphous character-states of Platyxanion are: 1— notauli absent; 2—

third (at least partly) and following tergites felty setose; 3— reduction of

sclerotization of the radius. The plesiomorphous character-states are: I— second

abscissa of mediella slightly curved; 2— ocelli comparatively small; 3— nervulus

postfurcal or interstitial.

Amicrocentrum (Platyxanion) seyrigi Granger (figs. 6—17)

Granger, 1949, Mem. Inst. scient. Madagascar 2A: 374—375, fig. 380.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 141.

Lectotype, 9, length of body 26.1, of fore wing 20.9 mm.

Head. — Antennal segments 49, length of 3rd segment 1.2 times 4th segment,

length of 3rd and 4th segments 3.0 and 2.5 times their width, respectively, length of

both penultimate segments 1.4 and 1.8 times their width, respectively, and apical

segment with short spine (fig. 9); dorsal length of eye 1.1. times temple; temple

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

rugose, rounded behind, slightly wider than width of head at eyes (fig. 14); POL: &

ocellus: OOL= 5:7: 13; frons medially concave and rugose, laterally convex and

rugose as vertex; face finely punctate-rugose, depressions from antennal sockets

to anterior tentorial pits present (fig. 16); clypeus rather convex, finely and densely

punctate; epistomal suture indistinctly developed medially; length of malar space

0.7 times basal width of mandible; malar suture well-developed.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

mainly finely punctate, with some short carinae medially (fig. 6); epicnemial area

densely rugose-punctate; precoxal suture densely rugose-punctate, as its

surroundings (fig. 6); pleural suture deep and rather narrowly crenulate; episternal

scrobe narrowly elliptical; metapleuron coarsely reticulate-punctate, but more

rugose ventrally; scutellar suture with ca. 16 short carinae; scutellum convex,

finely and densely punctate; side of scutellum carinate; surface of propodeum

densely punctate; propodeal spiracle elliptical.

Wings. — Metacarpella complete; r 1:r2:r3 = 13:39: 115;d1:d2 =4: 73;

cuguili: 2 cuqu 2 — 2439.29

Legs. — Hind coxa densely and coarsely punctate dorsally, rather smooth

ventrally; length of hind tibia ca. 2.1 times hind femur (fig. 10); length of femur,

tibia and basitarsus of hind leg 5.0, 12.0 and 11.4 times their width, respectively;

length of spurs of hind tibia 0.34 and 0.28 times basitarsus, slightly curved (fig. 10).

Metasoma. — Length of first tergite 3.5 times its apical width, its surface

somewhat microsculptured medially, but mainly smooth (fig. 17); length of

ovipositor sheath 1.10 times fore wing; hypopygium somewhat roundly emargined

apically.

Colour. — Light brown; tibiae and tarsi more yellowish brown; wing membrane

yellowish.

Lectotype in MNHN: “Madagascar, Ranomafana” ‘Muséum Paris, X.38. A.

Seyrig”, “49”, “Type”. Lectotype herewith selected and labelled accordingly.

Paralectotypes: (MNHN) 10 & of which 6 examined, all from Malagasy, Bekily

and all collected in January or February. Antennal segments 44 (2), 45 (1) or 46 (1);

length of fore wing 16.0—18.5, of body 19—24 mm; Ist and 2nd tergites with the

typical depressions (fig. 15). Additional specimens examined, all from MNHN and

collected in Malagasy: 1 9, Vohémar, heavily damaged, nervulus interstitial; 1 9,

Fort Dauphin, cuqu 2 absent in right wing but in the left wing nearly complete; 1

3, Analandravaka, 25.111.1936, antennal segments 44, malar suture almost absent,

length of fore wing 19.5, of body 26 mm.

Note. This is a very distinctive species, which is isolated from other species of

Amicrocentrum. The species is interesting in its combination of plesiomorphous

character-states, such as the slightly curved second abscissa of the mediella, and

apomorphous character-states, such as the absence of notauli.

VAN ACHTERBERG: Amicrocentrinae 13

Amicrocentrum Schulz, subgenus (figs. 18—68)

Schulz, 1911, Zool, Annin 4: 88.

Szépligeti, 1904, Genera Insect. 22: 145 (Megacentrus; nec Heer, 1852).

Cameron, 1912, Annls Soc. ent. Belg. 56: 370 (Eiolo).

Type-species: Megacentrus concolor Szépligeti.

Diagnosis. — Ocelli large, OOL subequal to diameter of ocellus (fig. 20) or

shorter (fig. 38); mandible weakly twisted apically; notauli narrowly developed

(fig. 27); second abscissa of mediella strongly sinuate (fig. 25); 3rd and 4th

abscissae of radius at least weakly sclerotized; nervulus narrowly antefurcal (fig.

37); basally radiella weakly curved anterad (fig. 37); nervellus straight or weakly

reclivous (figs. 25, 37); glymma rather deep anteriorly; 3rd and following tergites

only densely setose, pilose (fig. 49).

Distribution. — Afrotropical: four species, one restricted to Malagasy and three

to continental Africa.

Note. Apomorphous character-states of Amicrocentrum s.s. are: 1— second

abscissa of mediella strongly sinuate; 2— nervulus narrowly antefurcal; 3— ocelli

large. Plesiomorphous character-states are: 1— notauli present; 2— 3rd and

following tergites only densely setose; 3— radius sclerotized.

Amicrocentrum (Amicrocentrum) concolor (Szépligeti) (figs. 18 —29, 57—59)

Szépligeti, 1904, Genera Insect. 22: 146, pl. 3, fig. 19 (in Megacentrus).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4 (1): 141.

Holotype, 9, length of body 18.1, of fore wing 16.0 mm.

Head. — Antennal segments 48, length of 3rd segment 1.2 times 4th segment,

length of 3rd and 4th segments 3.6 and 3.1 times their width, respectively, length of

both penultimate segments 1.8 and 1.6 times their width, respectively, and apical

segment without an apical spine (fig. 26); dorsal length of eye 2.4 times temple;

temple mainly smooth, somewhat punctate ventrally and subparallel behind eyes

(fig. 20); POL : @ ocellus : OOL = 14 : 8: 7; frons concave and coarsely rugose;

vertex punctate; face medially smooth, laterally coarsely reticulate-punctate, with

deep grooves from anterior tentorial pits to both sides of the antennal sockets (fig.

22); clypeus rather convex, punctate; epistomal suture obliterated medially; length

of malar space 0.2 times basal width of mandible; malar suture narrow, indistinctly

developed.

Mesosoma. — Length of mesosoma 1.4 times its height; ventral half of side of

pronotum smooth, posteriorly partly coarsely rugose, remaining area remotely

punctate and with a deep short crenulate suture (fig. 18); epicnemial area rugose-

punctate; mesopleuron dorsally finely and densely punctate but mainly smooth

near the pleural suture and ventrally more coarsely punctate; precoxal suture

coarsely reticulate-punctate; pleural suture narrowly and densely crenulate, rather

deep; episternal scrobe absent; metapleuron rather coarsely punctate and

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

ventrally rugose; scutellar suture with ca. 20 short carinae; scutellum convex,

punctulate, but medially mainly smooth; side of scutellum punctate-rugose;

surface of propodeum densely punctate-rugose, posteriorly more coarsely

sculptured and with a rather long medial carina; propodeal spiracle subelliptical.

Wings. — Metacarpella complete; r1:r2:r3 = 10:38:91;d1:d2=-2:51;

cugu 1: r2 -cuqu2 — 16-38. 21.

Legs. — Hind coxa densely punctate dorsally; hind tibia missing, but its.length

ca. 1.9—2.0 times hind femur in other specimens; length of femur, tibia and

basitarsus of middle leg 6.3, 9.6, and 14.0 times their width, respectively, these

measurements of the hind leg in the 9 from Urundi are 5.7, 13.0 and 8.6 times,

respectively; length of middle tibial spurs both 0.2 times their basitarsus, straight.

Metasoma. — Length of Ist tergite 3.3 times its apical width, its surface

punctate-rugose, but apically smooth (fig. 29); length of 2nd tergite 1.4 times its

maximum width; length of ovipositor sheath 1.24 times fore wing; hypopygium

somewhat concave medio-apically.

Colour. — Light brown (but other specimens usually more yellowish); apices of

mandibles and stemmaticum, blackish; pterostigma brownish yellow; wing

membrane brownish; 4th segment somewhat darker brown.

Holotype in TMA: “Africa or., Kilima-Ndjaro” “Kilimandscharo” (old hand

written label), “Holotype Megacentrus concolor Szepl., 1904, det. Papp ’67’’, “Hym.

Typ. No. 781, Mus. Budapest’.

Additional specimens examined: 18 ©, 5 G and | specimen without metasoma.

From Benin (Djougou Kouandé), Burundi (Usumbula), East Africa (Jombene

Range, I could not trace this locality), Zaire (Kivu: Kavimoira (Uvira), at light;

Kasenyi), Kenya (Tsavo N.P. (E.): Lion Hill near Voi (500—600 m, deciduous

orthophyll savanna, at light); Una (Nziu); Naivasha; Kenani (Mtito Andei)

Samburu Game Reserve (Public Camp Site 1, at light); Galana R. (2 ml E. of

Tsavo N.P.); Nakuru), Uganda (Karamoja); Ethiopia (Hawash, ca. 3500 ft.);

Namibia (Hohnung); S. Africa (Shilouvana (N. Transvaal)) (MNHN, TC, USNM,

MAC, BM, LH, RMNH, NMK). Variation: Length of fore wing 7.8—17.0 mm,

length of 2nd tergite 1.4—1.8 times its width, antennal segments 46—52, length of

ovipositor sheath 1.28—1.50 times fore wing, length of Ist tergite 2.4—3.5 times its

apical width, 4th tergite mainly dark brown, contrasting with other, yellowish

tergites. Collected in April (10), December (3) and July (2).

Amicrocentrum (Amicrocentrum) curvinervis (Cameron) (figs. 30—41)

Cameron, 1912, Annls Soc. ent. Belg. 56: 372 (in Eiolo).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4 (1): 141.

Holotype, 9, length of body 18.5, of fore wing 12.3 mm.

Head. — Antennal segments 25, but apical part absent, length of 3rd segment

1.1 times 4th segment, length of 3rd and 4th segments 3.6 and 3.2 times their width,

respectively; dorsal length of eye 2.9 times temple; temple punctate, except near

VAN ACHTERBERG: Amicrocentrinae 15

eyes, rounded behind and sides almost subparallel (fig. 38); POL : & ocellus : OOL

= 11: 11:6; frons medially concave, with some rugae; vertex finely and densely

punctate; face coarsely and densely punctate, depressions at the inner side of the

antennal sockets only (fig. 32); clypeus convex, densely punctate; epistomal suture

complete; length of malar space 0.4 times basal width of mandible; malar suture

narrowly developed.

Mesosoma. — Length of mesosoma 1.4 times its height; medio-anterior side of

pronotum with a deep crenulate furrow, medially and apically rugose-punctate,

dorso-apically densely and rather coarsely punctate, ventral half of pronotum

smooth but punctulate anteriorly (fig. 30); epicnemial area punctate; precoxal

suture coarsely and densely punctate, rest of mesopleuron more remotely

punctate; pleural suture narrowly and indistinctly crenulate; episternal scrobe

absent; metapleuron densely punctate; scutellar suture with 20, mainly rather

short, carinae; scutellum rather convex, punctate laterally, almost smooth

medially; side of scutellum punctate-rugose; surface of propodeum coarsely and

densely punctate; propodeal spiracle subelliptical.

Wings. — Metacarpella complete;r1l:r2:r3=7:32:71;d1:d2=-2:85;

enqul:r2:cuqu2 = 12:32: 16.

Legs. — Hind coxa punctate dorsally; length of hind tibia ca. 2.2—2.3 times

hind femur (fig. 34); length of femur, tibia and basitarsus of hind leg 6.0, 18.2 and

19.0 times their width, respectively; length of spurs of hind tibia 0.3 and 0.2 times

hind basitarsus, straight.

Metasoma. — Length of Ist tergite 4.0 times its apical width, its surface densely

and finely punctate, but apical and basal fifths mainly smooth; length of 2nd

tergite 2.3 times its maximum width (fig. 41), sides of the tergite parallel; length of

ovipositor sheath 1.77 times fore wing; hypopygium truncate apically.

Colour. — Brownish yellow; stemmaticum and tips of mandibles, dark brown;

wing membrane brownish.

Holotype in MAC: “Type”, Musée du Congo Belge, Kasai: Eiolo, 16—1—06,

Waelbroeck”, “R.Det., E., 189”, “Eiolo curvinervis Cam., Type” (in Cameron’s

handwriting), “Amicrocentrum concolor (Szpl.) (= Eiolo curvinervis Cam.), H. De

Saeger, det. 1942”.

Additional specimens examined: 31 9 and 2 &. From Zaire (1 © without precise

locality, “Megacentrus concolor Szepl., det. Enderlein, 1918”; Magalo (Ubangi);

Lisala Ter.; Libenge; Jaradje (Ituri, Kasima); Tuku (Haut Uelé); Moto (id.); Yebo

Moto (id.); Paulis (id.); Watsa (id.); Congo da Lemba; Bambesa; Bombona

(Ubangi); Gemena (id.); Bakere (id.); Kunzolo); Benin (?city); Uganda (Bwamba;

Zika Forest (= near Entebbe, “ex larva B. fusca’) (MAC, PAN, TC, LH, NMK,

RMNH). The variation is considerable: length of fore wing 10.0—18.0 mm, length

of ovipositor sheath 1.60—2.01 times fore wing, antennal segments 51—59, length

of Ist tergite 3.1—4.4 times its apical width (exceptionally in males 2.8 times),

length of 2nd tergite 2.0—2.5 times its maximum width (in males up to 2.8 times),

sides of 2nd tergite parallel, 4th tergite yellowish, not contrasting with other

tergites. This species, like exilis, seems to be more restricted to the tropical forest,

while concolor may be restricted to more savanna-like habitats.

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

Amicrocentrum (Amicrocentrum) exilis spec. nov. (figs. 42—53)

Holotype, 9, length of body 20.9, of fore wing 19.4 mm.

Head. — Antennal segments 53, 3rd segment equal to 4th segment, 3rd and 4th

segments both 4.0 times their width, both penultimate segments 2.5 and 3.0 times

their width, respectively, apical segment sharp apically, but spine not well

developed (fig. 51); dorsal length of eye 2.3 times temple; temple almost smooth,

indistinctly punctulate and rounded behind (fig. 50); POL : @ ocellus : OOL = 8:

13 : 7; medially frons concave, smooth except for a few short carinae; vertex

indistinctly punctulate; face below antennal sockets punctate-rugose, rest of face

remotely punctate, unevenly convex dorsally, flattened ventrally; clypeus

flattened, remotely punctulate; epistomal suture obliterated dorsally; length of

malar space 0.5 times basal width of mandible; malar suture indistinctly

developed.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

almost smooth, except for some crenulae medio-posteriorly and some

punctulation near the margins (fig. 42); epicnemial area finely striate-rugose

anteriorly, remotely punctate dorsally as surroundings of precoxal suture;

precoxal suture densely and finely rugose-punctate; pleural suture narrow, rather

shallow and indistinctly crenulate; episternal scrobe absent, except for an

indistinct longitudinal impression; metapleuron remotely punctate; scutellar

suture with 9 short carinae; scutellum convex, remotely punctulate; side of

scutellum punctulate and with some carinae; surface of propodeum densely and

rather finely reticulate-rugose, medially longitudinally depressed; propodeal

spiracle elliptical.

Wings. — Metacarpella complete; r 1:r2:r3 = 12:46: 137;d1:d2=-3:

100; cuqu Fr 2 :cuqu2 = 17 246: 23.

Legs. — Hind coxa finely and remotely punctate; length of hind tibia ca. 2.4

times hind femur (fig. 44); length of femur, tibia and basitarsus of hind leg 7.8, 18.2

and 15.8 times their width, respectively; length of hind tibial spurs 0.25 and 0.20

times hind basitarsus, slightly curved apically, but almost straight.

Metasoma. — Length of Ist tergite 4.4 times its apical width, its surface smooth,

except for some microsculpture laterally, only medially sparsely setose, and

medio-basal hole rather sharp apically (fig. 49); length of 2nd tergite 3.0 times its

maximum width; length of ovipositor sheath ca. 1.47 times fore wing; hypopygium

truncate apically.

Colour. — Brownish yellow; stemmaticum, tips of mandibles, flagellum (but

apically more yellowish), wing venation (but its basal half yellowish), dark brown;

pterostigma and ovipositor sheath brown; wing membrane rather hyaline.

Holotype in TC: “Zika Forest (= near Entebbe), Uganda, viii.23,’63, G.

Lancaster”. Paratypes: (9 © and 4 3): 1 ©, topotypic, ix—x.1963 (RMNH); 2 9

and 3 &, Zaire, Eala, xi.1935, J. Ghesquiere (MAC, RMNH); 1 9, Zaire,

Kunungu, 1938 (Nkele, coll. Schouteden ) (MAC); 1 9, Zaire, Riv. Busira, vi.1936,

J. Ghesquiere (MAC); 1 9, Sankuru: Katako-Kombe, 14.viii.1952, M. Fontaine

(MAC); 1 9, Zaire, N. Lac Kivu: Reankwi, 19.1x.1947, J. v. Leroy (RMNH); | ©,

Oyoko-Cacas Stn, Oyoka-Ghana, 24.ii-1959, parasite: Eulophonotus myrmeleon

VAN ACHTERBERG: Amicrocentrinae 17

(BM); 1 ©, Uganda, 7 mls from Entebbe, Zika Forest, iii—vi.1961, P. S. Corbet

(BM); 1 &, Tanzania, Ukerewe I., Father Conrad (NMK).

Variation: Antennal segments 55 (3); length of fore wing 14.0—21.0.mm; length

of ovipositor sheath 1.31—1.37 times fore wing; length of Ist tergite 4.4—5.1 times

its apical width, but in one male 3.2 times; length of 2nd tergite to 3.2 times its

maximum width; length of malar space 0.3—0.5 times basal width of mandible.

Amicrocentrum (Amicrocentrum) flavipenne Granger (figs. 54—56, 60—68)

Granger, 1949, Mem. Inst. scient. Madagascar 2A: 373—374.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4 (1): 141.

Lectotype, 9, length of body 27.3, of fore wing 20.9 mm.

Head. — Antennal segments 51, length of 3rd segment 1.4 times 4th segment,

length of 3rd and 4th segments 4.4 and 4.2 times their width, respectively, both

penultimate segments 2.4 and 2.8 times their width, respectively, apical segment

without spine (fig. 67); dorsal length of eye 1.8 times temple; temple punctate,

rounded behind (fig. 64); POL : @ ocellus : OOL = 5: 8 : 6; frons concave, almost

smooth; vertex remotely punctate; face rather flat, finely punctate, but below

antennal sockets aciculate-punctulate; clypeus flattened, shiny, punctulate;

epistomal suture absent dorsally; length of malar space 0.6 times basal width of

mandible; malar suture absent.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

mainly sparsely punctulate, shiny, almost smooth, but with some carinae medio-

anteriorly (fig. 54); epicnemial area almost smooth, but somewhat rugose

anteriorly; precoxal suture mainly finely punctate as its surroundings, but near

middle somewhat rugose (fig. 54); pleural suture narrowly crenulate; episternal

scrobe almost absent; metapleuron finely and remotely punctate; scutellar suture

with 9 short carinae; scutellum rather convex, remotely and finely punctate; side

of scutellum punctate-rugose; surface of propodeum densely rugose-punctate,

without a medial carina; propodeal spiracle circular, protruding.

Wings. — Metacarpella of hind wing scarcely developed, reduced and

disconnected from the 2nd abscissa of the subcostella (fig. 55);rl:r2:r3= 7:33

mod lad 2 =-2:69;cuqu l:r2:cuqu2= 10: 33: 11.

Legs. — Hind coxa almost smooth; length of hind tibia ca. 2.4 times hind femur

(fig. 56); length of femur, tibia and basitarsus of hind leg 7.7, 20.3 and 17.5 times

their width, respectively; length of spurs of hind tibia both 0.2 times their

basitarsus, subequal, straight.

Metasoma. — Length of Ist tergite 6.3 times its apical width, its surface smooth

but basally somewhat finely coriaceous, its medio-basal hole round apically (fig.

68); length of 2nd tergite ca. 2.9 times its maximum width; length of ovipositor

sheath ca. 1.34 times fore wing; hypopygium truncate apically.

Colour. — Brownish-yellow; tips of mandibles and stemmaticum blackish;

flagellum dark brown; wing membrane somewhat yellowish.

Lectotype in MNHN: ‘Madagascar, Rogez, Forêt cote est”, “Museum Paris,

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

iii.37, A. Seyrig”, “53”, “Type”. Herewith selected as lectotype. Paralectotype: 1

3 (MNHN), topotypic, ii.37. Length of fore wing 20.4, of body 26.0 mm; length of

Ist tergite 3.7 times its apical width.

ACKNOWLEDGEMENTS

I am much indebted to the following persons for the loan of types and/or gifts of

unidentified specimens. The abbreviations used for the collections are given in

brackets. Dr. J. Decelle, Musée Royal de l’ Afrique Central, Tervuren (MAC); Mr.

T. Huddleston, British Museum (Natural History), London (BM); Mr. K.

Kabuthia, National Museums of Kenya, Nairobi (NMK); Dr. S. Kelner-Pillault &

Mr. B. Sigwalt, Muséum National d’Histoire Naturelle, Paris (MNHN); Dr. E.

Kierych, Instytut Zoologii, Warsaw (PAN); Dr. P. Marsh, USDA c/o U.S. National

Museum, Washington (USNM); Dr. J. Papp, Zoological Department of the

Hungarian Natural History Museum, Budapest (TMA); Dr. H. K. Townes,

American Entomological Institute, Ann Arbor (TC); Drs. K. W. R. Zwart,

Laboratorium voor Entomologie, Landbouwhogeschool, Wageningen (LH);

(RMNH) = Rijksmuseum van Natuurlijke Historie, Leiden.

I wish to express my thanks to Dr. J. R. T. Short (Australian National

University, Canberra) for his description of the larva, his useful suggestions and

the correction of the English text.

LITERATURE

Achterberg, C. van, 1975. Een merkwaardige vondst in Voorne’s duinen (Hym., Braconidae). — Ent.

Ber., Amst. 35: 15—16, figs. 1—5.

—., 1976a. A revision of the tribus Blacini (Hym., Braconidae, Helconinae). — Tijdschr. Ent. 118

(7): 159—322, figs. 1—476, 2 tables.

, 1976b. A preliminary key to the subfamilies of the Braconidae (Hym.). — Tijdschr. Ent. 119

(3): 33—78, figs. 1—123, I table.

Brothers, D. J., 1975. Phylogeny and classification of the aculeate Hymenoptera, with special reference

to Mutillidae. — Univ. Kansas Sci. Bull. 50 (11): 483648, figs. 1—101, tables 1—7.

Cameron, P., 1912. On the Hymenoptera from Belgian Congo in the Congo Museum, Tervueren. —

Annls Soc. ent. Belg. 56: 357—401.

Capek, M., 1970. A new classification of the Braconidae (Hym.) based on the cephalic structures of the

final instar larva and biological evidence. — Can. Ent. 102 (7): 846—875, figs. 1—58.

—, 1973. Key to the final instar larvae of the Braconidae (Hym.). — Acta Inst. forest. Zvol.

259—268, | fig.

Granger, C., 1949. Braconides de Madagascar. — Mem. Inst. scient. Madagascar, Ser. A: 1—428, figs.

1—426.

Hamilton, K. G. A., 1971. The insect wing, part 1. Origin and development of wings from notal lobes.

— J. Kansas ent. Soc. 44 (4): 421—433, figs. 1—18.

Schulz, W. A., 1911. Zwei hundert alte Hymenopteren. — Zool. Annin 4: 1—220.

Shenefelt, R. D., 1969. Hymenopterorum Catalogus (nov. ed.). Part 4. Braconidae 1: 1—176. — Junk,

*s-Gravenhage.

Short, J. R. T., 1952. The morphology of the head of the larval Hymenoptera with special reference to

the head of the Ichneumonoidea, including a classification of the final instar larvae of the Bra-

conidae. — Trans. R. ent. Soc. Lond. 103: 27—84, figs. 1—34.

——., 1978. The final larval instars of the Ichneumonidae. — Mem. Am. ent. Inst. 25: 1—514, figs.

1—802. :,

Szépligeti, G. V., 1904. Hymenoptera, Fam. Braconidae. — Genera Insect. 22: 1—253, figs. 1—32.

VAN ACHTERBERG: Amicrocentrinae

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20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

Figs. 13—17. Amicrocentrum seyrigi Granger, lectotype (but 15 of paralectotype). 13, outer middle claw;

14, head, dorsal aspect; 15, Ist-3rd tergites, dorsal aspect; 16, head, frontal aspect; 17, Ist-3rd tergites,

dorsal aspect. 13: 5.0 x scale-line; 14—17: scale-line

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26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

Figs. 47—53. Amicrocentrum exilis spec. nov., holotype. 47, head, frontal aspect; 48, apex of hind tarsus,

outer aspect; 49, Ist—3rd tergites, dorsal aspect; 50, head, dorsal aspect; 51, apex of antenna; 52, apex

of hind tibia, outer aspect; 53, mesonotum, dorsal aspect. 47, 50: 2.0 x scale-line; 48, 51, 52: 5.0 x sca-

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28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1, 1979

Figs. 61—68. Amicrocentrum flavipenne Granger, lectotype. 61, apex of middle tarsus; 62, antenna; 63,

head, frontal aspect; 64, head, dorsal aspect; 65, inner middle claw; 66, mesonotum, dorsal aspect; 67,

apex of antenna; 68, Ist and 2nd tergites, dorsal aspect. 61: 2.0 x scale-line; 62, 64, 66, 68: scale-line;

63: 1.7 x scale-line; 65: 5.0 x scale-line; 67: 4.0 x scale-line

A REVISION OF THE NEW SUBFAMILY XIPHOZELINAE

(HYMENOPTERA, BRACONIDAE)

C. VAN ACHTERBERG

Rijksmuseum van Natuurlijke Historie, Leiden

With 45 text-figures

ABSTRACT

A new subfamily is erected for the genera Xiphozele Cameron, 1906, and Distilirella gen. nov.

Distilirella curvinervosa gen. et spec. nov., from New Guinea, and the species of Xiphozele are described

and fully illustrated.

INTRODUCTION

In the course of a revision of the Macrocentrinae s.l. I have tried to delimit the

subfamily Macrocentrinae with the aid of synapomorphous character-states. One

of the genera, which proved to be untenable in the Macrocentrinae because of the

lack of synapomorphous character-states is Xiphozele Cameron, and a genus with

even less apomorphous character-states has now been discovered in New Guinea.

The distribution of the Xiphozelinae is restricted to the South East Palaearctic and

Oriental regions and New Guinea. There are few specimens of Xiphozelinae in any

collection, though they are large, conspicuous insects. Despite this an attempt is

made to revise the group. For the scarce literature, see Shenefelt (1969: 174—175)

and for the general terminology, see Van Achterberg (1976a: 160—166).

PHYLOGENY

The new subfamily formed by the genera Xiphozele Cameron and Distilirella

‘gen. nov. possesses the following remarkable apomorphous character-states:

1. A deep and round laterope situated far from the base of the first metasomal

tergite (figs. 18, 28). This is a unique character-state, which, as far as I am aware,

does not occur in other subfamilies of the Braconidae; there is at most a rather

shallow, elliptical laterope far removed from the base of the tergite, e.g., in the

genus Zele Curtis (nec auct.). The presence of a laterope is itself a plesiomorphous

character-state.

2. The strongly inclivous nervellus of the hind wing, which is exceptional in the

Braconidae and probably an apomorphous character-state. Only in the genus

Brulleia Szépligeti does the same condition appear.

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

3. The short ovipositor, about equal to the apical height of the metasoma (fig. 1).

An ovipositor about as long as the fore wing or somewhat shorter is often

associated in the Braconidae with other plesiomorphous character-states.

Therefore the short ovipositor of the Xiphozelinae is considered to be an

apomorphous state.

4. The first recurrent vein is far antefurcal (fig. 9); it is a general tendency in the.

Braconidae that apical veins retreat towards the base of the wing. Thus an

antefurcal recurrent vein is an apomorphous character-state, while Brulleia with

its postfurcal first recurrent vein shows the plesiomorphous condition.

5. The presence of an “ophionoid facies” (Gauld & Huddleston, 1976), most

pronounced in the yellowish body colour, and the large ocelli and eyes, which are

an adaptation to the noctural activity of the parasites. This is probably induced by

the nocturnal activity ofthe caterpillars (Noctuidae!) wherein the egg is deposited. _

6. The absence of the dorsal carinae and the medio-basal depression of the first

metasomal tergite (fig. 8). As shown by several groups of Braconidae with many

other plesiomorphous character-states (e.g., Doryctinae, Helconinae), the

presence of at least short basal dorsal carinae and of a medio-basal depression has

to be considered to represent a plesiomorphous character-state. |

7. The spiracle is situated far from the base of the first tergite. This is a general

tendency in the Braconidae (Van Achterberg, 1976b: 36); instead of subbasal

spiracles (the plesiomorphous condition) the spiracles are situated more or less

submedially because of the petiolation of the first tergite. The slender posterior

half of the first tergite (fig. 19) in the Xiphozelinae is also unusual.

8. The reduction of the occipital carina. This carina is present in many not

closely related groups of Hymenoptera which show many other plesiomorphous

character-states.

9. The presence of the lateral carina of the mesoscutum. A well-developed

lateral carina seems to be a late development in the history ofthe Braconidae.

10. The claws possess a more or less developed ventral lamella (figs. 16, 35). As

pointed out by Brothers (1975: 521) simple bifurcate claws (without a lamella) have

to be considered a plesiomorphous character-state inthe Hymenoptera.

11. The long palpi; length of maxillary palp 1.8—2.3 times height of head. The

plesiomorphous condition of the maxillary palp in the Braconidae is a length about

equal to the height of the head.

The plesiomorphous character-states of the Xiphozelinae are:

1. The presence of the first transverse anal, the transverse anellan and the

transverse radiellan veins. These veins are weakly developed, but they indicate a

more complex (and plesiomorphous state of) venation. The same applies to the

presence of the second transverse cubital vein and the long radial vein of the fore

wing.

2. The presence of a laterope. In the Ichneumonoidea it is most likely an early

development and should therefore be considered a plesiomorphous character-

state within the Braconidae. It is common in not closely related groups, which

show several other plesiomorphous character-states.

3. The large plical (anal) lobe of the hind wing. A well-developed plical lobe

VAN ACHTERBERG: Xiphozelinae 31

such as present in the Symphyta is a plesiomorphous character-state.

4. The metasomal tergites are equally setose. The reduction of the setosity is a

common (apomorphous) condition, but it has not taken place in the Xiphozelinae.

5. The fore tibial spur is rather slender, more or less cylindrical and bears a

narrow flange on the inner side. This is the common plesiomorphous character-

state in the Hymenoptera-Apocrita.

6. The presence of the prepectal and hypostomal carinae. The presence of these

carinae is generally accepted to be a plesiomorphous character-state. The carinae

are present in many not closely related groups of Hymenoptera which have many

other plesiomorphous character-states in common.

7. The maxillary and labial palpi consist of 6 and 4 segments, respectively, and

are well-developed. There is a general tendency for reduction of the palpi (Van

Achterberg, 1976b: 35), but in the Xiphozelinae the plesiomorphous character-

state still occurs.

8. The first discoidal cell is shortly petiolate. A petiolate first discoidal cell is

generally considered to be a plesiomorphous character-state in the Hymenoptera.

More doubtful character-states are:

1. The metasoma is inserted above the hind coxae. Probably this character-state

is plesiomorphous, if it is not inserted extremely high as in the Cenocoeliinae and

the Evanioidea. A medially inserted metasoma occurs in several subfamilies of

Braconidae which are not closely related (e.g., Helconinae, Macrocentrinae,

Orgilinae, Amicrocentrinae, Agathidinae, and Xiphozelinae) as well as in the

Ichneumonidae (subfamily Labeninae (= Labiinae sensu Townes)).

2. The radiellan cell of the hind wing is more or less widened apicad (figs. 25,

37). Probably (if in a rather moderate manner) this is a plesiomorphous condition.

Less easy to answer is the question of the relationships of the Xiphozelinae.

Their inclusion in the Macrocentrinae as done by most previous authors is

untenable because of the synapomorphous character-states. The Macrocentrinae

have the following synapomorphous character-states: 1—trochantelli apically

toothed; 2—plical lobe rather narrow; 3—occipital carina completely absent;

4—claws simple or with a lamella; 5—first recurrent vein far antefurcal. Most of

these character-states are absent or at least are not completely present in the

Xiphozelinae. The unique apomorphous character-state of the Macrocentrinae,

viz., the apically toothed trochantelli, is absent in the Xiphozelinae. The other

characters show more or less a tendency to follow the developments in the

Xiphozelinae, but the Xiphozelinae have in the round and deep laterope, which is

situated far posteriorly, and the strongly inclivous nervellus their own

apomorphous character-states not shared by the Macrocentrinae. Of the other

apomorphous character-states of the Xiphozelinae the short ovipositor, the

‘ ophionoid facies, the reduction of the dorsal carinae and medio-basal depression

of the first tergite, and more posteriorly situated spiracle of the first tergite are

independently evolved in the Macrocentrinae, because the plesiomorphous

condition is common in the Macrocentrinae. The development of the lateral

carinae of the mesoscutum, the reduction of the occipital carina and the far

antefurcal first recurrent vein may indicate the same origin, but this conclusion is

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

uncertain because they are general tendencies in the Braconidae. The

plesiomorphous character-states not present in the Macrocentrinae are the

presence of the transverse anellan and radiellan veins and the large plical lobe of

the hind wing.

A candidate for the sister-group of the Xiphozelinae is formed by the

Helconinae s.l., because of the genus Brulleia Szépligeti. Brulleia has also a large

plical lobe, an apically dilated radiellan cell, a short second abscissa of the

subcostella and an inclivous nervellus. But only the latter is a probable

apomorphous character-state, while Brulleia has the first discoidal cell widely

sessile, maxillary and labial palpi 5 and 3 segmented, respectively, and the

transverse radiellan and anellan veins absent. Besides these apomorphous

character-states, it has the second transverse anal vein present, the first recurrent

vein postfurcal and a complete occipital carina, which are plesiomorphous

character-states not present in the Xiphozelinae.

In summary: the position of the Xiphozelinae is uncertain, they are not closely

related to the Macrocentrinae s.s. as suggested by other authors. There may be a

relationship with the Helconinae s.l., to which they are more closely related than

are the Amicrocentrinae.

BIOLOGY

Only in the case of Xiphozele compressiventris Cameron is something known

about the biology. I have examined two females from South India (Karwar,

Karnataka, 14.viii.1907, F. R. Bell (BM)), which were reared from the “pupa”

(probably cocoon) of Ophiusa simillima Guenée (Lepidoptera: Noctuidae).

Watanabe (1969: 327) reported a female bred from a lepidopterous larva feeding

on leaves of Quercus spec. at Sapporo. The host species seems to belong to the

Noctuidae. The parasite-larva left its host on September 7, 1966, and spun a

cocoon within which it passed the winter. The adult emerged from the cocoon on

March 23, 1967. The dates of capture of the 6 specimens from Japan are between

25.111 and 28.viii. The cocoon is 12—13 mm long, spindle-shaped, thick, dark

brown, and somewhat woolly. Because Noctuid larvae are usually active

nocturnally, it is likely that this induces the nocturnal activity of the adult

parasites.

XIPHOZELINAE subfam. nov.

Diagnosis. — Length of body 12.5—20.0, of fore wing 10.5—16.6 mm; antennal

segments of 9 53—56, of g 51—S4; pedicellus short, transverse and cylindrical

(figs. 1, 28); maxillary and labial palpi long, slender, 6- and 4-segmented,

respectively; apical segment of antenna with a long spine apically (fig. 2); ocelli

large (figs. 12, 24); anterior tentorial pits large, deep (fig. 10); epistomal suture

complete; occipital carina absent dorsally; eye bare; apical margin of clypeus thin

and differentiated from clypeus (figs. 22, 40); labrum visible frontally; occipital

flange narrowly developed; hypostomal carina present (fig. 7); mandible strongly

twisted, both its teeth sharp apically and second tooth much shorter than first

VAN ACHTERBERG: Xiphozelinae 33

tooth (figs. 7, 40); pronope and antescutal depression absent; pronotum convex

dorsally and with an upwardly directed lamella anteriorly; propleural lamellae

developed (fig. 1); dorso-apical corner of pronotum rounded and more or less

protruding posteriad (figs. 18, 28); tegulae almost reaching anterior margin of

mesopleuron (fig. 1); mesopleuron slightly protruding anteriorly (fig. 18); lateral

carina of mesoscutum present in front of tegulae; lateral and middle lobes of

mesoscutum equally convex; prepectal carina strongly developed (figs. 1, 28);

precoxal suture rather impressed (fig. 18); metapleural flange or lamella large, and

sharp apically (figs. 18, 28); dorsal surface of propodeum not differentiated from

its posterior surface (fig. 28); propodeum without areola and tubercles; propodeal

spiracle large, more or less elliptical (figs. 18, 28); antepropodeal depression rather

wide and deep (fig. 39); notauli present; mesoscutal lobes rather convex; scutellar

suture wide, long, deep and with one longitudinal carina (figs. 26, 43); scutellum

sculptured posteriorly (figs. 6, 26, 43); metanotum rather flat postero-medially

(figs. 18, 26); first discoidal cell shortly petiolate and sharp anteriorly (fig. 9); cuqu

2 present; r 3 more or less curved towards metacarp (figs. 25, 37); nervulus

interstitial or nearly so (figs. 9, 37); n. rec. far antefurcal; sides of B 1 parallel or

nearly so (figs. 9, 37); CU 1 larger than CU 2; nervellus very long, inclivous and

posteriorly curved basad (fig. 25); B 1 closed apically, s 1b present; fringes of wings

short; parastigma large (fig. 37); intercostal cell absent; aqu | and aqu’ present as

weakly pigmented stripes (fig. 25); aqu 2 absent; metacarp ends near apex of radial

cell (fig. 9); plical lobe very large and normally setose (fig. 37); radiellan cell

widened apicad after the completely developed rqu’ (figs. 25, 37); base of radiella

as sclerotized as basella; basella and second abscissa of subcostella short (fig. 9);

discoidella absent, but exceptionally a remnant is present (fig. 38); legs slender,

length of hind tibia ca. 1.3 times its femur; length of femur and tibia of hind leg

9.0—10.1 and 11.8—15.0 times their width, respectively; hind tibial spurs long,

straight, setose and sharp apically; inner hind tibial spur 0.6—0.7 times its

basitarsus; hind basitarsus without a ventral row of setae; tarsal claws of 9 witha

ventral lamella (figs. 15, 34); shape of inner hind claw equal to its outer claw; fore

tibial spur rather slender, more or less cylindrical and with a narrow flange or

lamella (figs. 20, 21); trochantelli without teeth, simple apically; apex of hind tibia

bristly (fig. 31) or with slender pegs (fig. 11); metasoma inserted medially between

dorsal surface of propodeum and the dorsal level of base of hind coxae (figs. 1, 18);

length of first metasomal tergite 5.5—6.4 times its apical width; sides of first tergite

(sub-)parallel; first tergite flat or convex medio-basally; dorsal carinae of first

tergite absent (fig. 39); laterope deep, round and large, at basal third of first

tergite, just in front of spiracle and far removed from the base of the tergite (figs.

18, 19); dorsope absent; metasoma evenly setose; second tergite smooth as

following tergites, with a weakly developed lateral crease (fig. 28); metasoma

strongly compressed apicad (fig. 19); ovipositor straight and with a shallow

subapical notch (figs. 1, 28); length of ovipositor sheath 0.05—0.07 times fore

wing; hypopygium large and truncate apically (fig. 28).

Distribution. — South East Palaearctic and Oriental regions and New Guinea.

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

Key to genera of the Xiphozelinae

l. Prepectal carina reaches anterior margin of mesopleuron (fig. 1); occipital

carina far removed from hypostomal carina ventrally (fig. 7); basella of hind

wing straight (fig. 5); nervulus abruptly bent distad and with a sclerome (figs.

17, 27); scutellum without a lateral carina (fig. 26); tarsal claws of © with only

one narrow.submediallamella (fig) 27.0 un, Xiphozele Cameron

— Prepectal carina remains far removed from the anterior margin of

mesopleuron (fig. 28); occipital carina reaching hypostomal carina ventrally

(fig. 32); basella strongly curved (fig. 45); nervulus straight (fig. 33), without a

sclerome; scutellum with a lateral carina (fig. 43); tarsal claws of 9 with two

lamellae, the 2nd lamella subapically attached to the submedial lamella (fig.

SO ioe scat er saline Re a ee ae Distilirella gen. nov.

Xiphozele Cameron (figs. 1—27)

Cameron, 1906, Entomologist 39: 204.

Enderlein, (1918) 1920, Arch. Naturgesch. 84A (11): 219 (Cerotopia).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 174—175.

Watanabe, 1969, Proc. ent, Soc. Wash. 71(3): 325—327, figs. 8—9.

Sharma, 1975, Oriental Ins. 9(2): 173—175, figs. 1—5.

Type-species: Xiphozele compressiventris Cameron

Diagnosis. — Apex of scapus rather roundly emarginated (fig. 18); occipital ca-

rina far removed from hypostomal carina (fig. 7); eyes not (fig. 22) or scarcely visi-

bly emarginated at inner side (fig. 10); teeth of mandible robust (fig. 10); apical

margin of clypeus more (fig. 22) or less (fig. 10) emarginated; prepectal carina

reaches anterior margin of mesopleuron (figs. 1, 18); episternal scrobe absent or

nearly so (fig. 18); scutellum without a lateral carina (fig. 26), widely sculptured

posteriorly (figs. 6, 26); cu 1 straight or nearly so (fig. 9); nervulus abruptly bent

distad, much narrower than surrounding veins, interstitial with basal vein and with

a sclerome (figs. 17, 27); SM more (fig. 17) or less (fig. 27) bare apically; basella

straight; metacarpella weakly curved or rather straight (figs. 5, 25), exceptionally

strongly curved (fig. 9 in Watanabe, 1969); tarsal claws of 9 setose and with a sub-

medial lamella (figs. 15, 16); tarsal claws of G with a large, somewhat inward

directed and apically sharp lamella (figs. 13, 14); laterope deep (fig. 8) or very deep

(fig. 19), more or less removed from each other; ovipositor sheath stout (fig. 1), but

unknown of burmensis.

Distribution. — Australian (New Guinea), Oriental and South East Palaearctic:

two species.

Note. — Apomorphous character-states of Xiphozele are: 1—nervulus abruptly

bent distad; 2— SM more or less bare; 3—sclerome present in fore wing; 4—claws

with a ventral lamella; 5—occipital carina absent ventrally; 6—ovipositor sheath

stout; 7—prepectal carina present antero-dorsaily, reaching anterior edge of me-

sopleuron. Plesiomorphous character-states are: 1—basella straight; 2—metacar-

pella weakly curved or almost straight; 3—scutellum without a lateral carina.

VAN ACHTERBERG: Kiphozelinae 35

Key to species of Kiphozele Cameron

l. Mesoscutal lobes brownish yellow; wing membrane hyaline; vertex punctulate

or smooth (fig. 12); clypeal margin weakly concave medially (fig. 10);

submedial cell mainly bare (fig. 17), exceptionally with ca. 30

EE err REA RETE PT compressiventris Cameron

— Mesoscutal lobes mainly dark brown; wing membrane more or less brownish;

vertex punctate (fig. 24); clypeal margin rather deeply concave medially (fig.

22); submedial cell mainly sparsely setose (fig. 27), usually with at least ca. 30

EEE UNE aa Ee oo EEM fee burmensis Sharma

Xiphozele compressiventris Cameron (figs. 1—17)

Cameron, 1906, Entomologist 39: 205.

Enderlein, (1918) 1920, Arch. Naturgesch. 89 A(11): 220, fig. 11 (Cerotopia corneimacula).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 174—175.

Watanabe, 1969, Proc. ent. Soc. Wash. 71(3): 325—327, figs. 8, 9.

Sharma, 1975, Oriental Ins. 9(2): 173, 175.

Redescribed after a 9 from Sri Lanka, compared with holotype. Length of body

15.0, of fore wing 11.5 mm.

Head. — Antennal segments 53, length of 3rd segment 1.3 times 4th segment,

length of 3rd and 4th segments 4.0 and 3.2 times their width, respectively,

penultimate segments 2.3 and 3.0 times their width, respectively, and apical

segment with a long spine (fig. 2); length of maxillary palp 2.2 times height of head;

dorsal length of eye 3.6 times temple; temple roundly receding (fig. 12); POL: 5

ocellus: OOL = 10: 9: 3; frons almost flat and smooth; vertex mainly smooth;

occipital carina absent, except for a lateral remnant at middle level of eyes (fig. 7,

in all other specimens examined there is at least a vague remnant present); face

convex, punctulate, shiny, but coriaceous near the medial convexity; clypeus

strongly convex, punctulate; apical margin of clypeus weakly concave (fig. 10);

length of malar space 0.6.times basal width of mandible; malar suture weakly

developed (fig. 10).

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, but medially and posteriorly somewhat crenulate and weakly striate

below the deep antero-dorsal depression (fig. 1); epicnemial area mainly weakly

punctate; precoxal suture coarsely reticulate-punctate and its surroundings

weakly punctate; pleural suture densely and narrowly crenulate, rather shallow

and narrow (fig. 1); metapleuron coarsely reticulate; notauli distinctly impressed,

but smooth (fig. 6); mesoscutal lobes indistinctly punctulate-coriaceous; side of

scutellum remotely crenulate; metanotum with one medial carina and a pair of

parallel carinae sublaterally (fig. 6); surface of propodeum rather finely and

closely reticulate, but anteriorly and posteriorly narrowly smooth, without a

medial carina, except for a weakly developed short part anteriorly.

Wings. — rl:r2:r3 = 16: 20: 52; cuqu | : r2: cuqu 2 = 17: 20: 12, r3 curved

anteriad; metacarpella curved (fig. 5, in some other specimens rather strongly);

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

SM with ca. 10 setae (in other specimens exceptionally as many as ca. 30 setae).

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

9.0, 11.8 and 10.6 times their width, respectively; length of hind tibial spurs 0.7 and

0.6 times their basitarsus.

Metasoma. — Length of Ist tergite 6.0 times its apical width, its surface mainly

smooth (fig. 8), but weakly transversely aciculate medially and weakly punctate

posteriorly; base of Ist tergite tube-shaped and with a pair of short ventral carinae

(fig. 1); whole Ist tergite convex, but basally weakly, and its spiracles not

protruding; length of ovipositor sheath 0.05 times fore wing.

Colour. — Brownish yellow; tips of mandibles, direct surroundings of ocelli,

wing veins mainly, more or less dark brown; apical half of metasoma somewhat

infuscated; base of Ist tergite, tibiae and tarsi rather whitish yellow; wing

membrane hyaline.

Holotype (only type-specimen) in BM; type-locality: Sikkim; no. 3.c.683.

According to Mr. T. Huddleston (in litt.), who was kind enough to examine the

type, this specimen has the submedial cell bare except for about 20 setae and the

mesoscutum is entirely without any dark brown coloration. The specimen figured

and redescribed is from Kandy: “E. Comber, Feb.’10, Kandy (Sri Lanka)”, “1910/

255”, “Cerotopia corneimaculata End., G. Nixon, det 1948” (BM). Additionally, 10

Bewani Mts.), Indonesia (Sumatra, Sukaranda, type of Cerotopia corneimacula

Enderlein), Sarawak (Kuching, at night, in house), Sri Lanka (Kandy, 2000 ft.),

India (Karwar, Karnataka, S. India, ex Ophiusa simillima Guenée), China (Kouy

Tcheou, Se Tchouen), Taiwan (Sunmoon Lake) and Japan (Wakayama; Hayatuki,

Toyama, Honsyu) (BM, EI, TC, USNM, MNHN, PAN, RMNH). Variation:

antennal segments of 9 56 (1 specimen), of & 51 or 53 (2 specimens), length of

ovipositor sheath 0.05 times fore wing; length of fore wing 10.5—16.6, of body

14.2—20.0 mm; length of Ist tergite 5.8—6.0 times its apical width; length of

metasoma 2.5—2.6 times length of metasoma.

Xiphozele burmensis Sharma (figs. 18 —27)

Sharma, 1975, Oriental Ins. 9(2): 173—175, figs. 1—5.

Holotype, 9 (according to the original description, but apical part of metasoma

and all claws lost), length of body (without apical half of metasoma): 12.0, of fore

wing 13.6 mm. |

Head. — Remnant of antenna consists of 8 segments (antennal segments of

paratype 50, of & from China 55), length of 3rd segment 1.2 times 4th segment,

length of 3rd and 4th segments 4.1 and 3.3 times their width, respectively,

penultimate segments absent but in G from China 2.0 and 1.6 times their width and

apical segment with a long spine; length of maxillary palp 1.8 times height of head;

dorsal length of eye 2.8 times temple; temple rounded behind (fig. 24); POL: @

ocellus : OOL = 28: 13: 15; frons mainly flat, smooth; occipital carina absent, but

dorso-laterally a weakly developed and short remnant is present (as in both other

specimens examined); vertex punctate (fig. 24); face convex, punctate, with a

VAN ACHTERBERG: Xiphozelinae 37

short medio-dorsal ridge (fig. 22); clypeus strongly convex, densely punctate;

length of malar space 0.6 times basal width of mandible; malar suture absent.

Mesosoma. — Length of mesosoma 1.2 times its height; side of pronotum

smooth dorsally, rugulose with some crenulae ventrally, and medially depressed,

with some crenulae (fig. 18); epicnemial area punctate and somewhat rugose;

precoxal suture coarsely rugose-reticulate; pleural suture indistinctly crenulate,

narrow, ventrally smooth except for some crenulae (fig. 18); metapleuron

reticulate-carinate; notauli only in posterior half distinctly impressed and

crenulate, anteriorly only rugulose (fig. 26); mesoscutal lobes punctulate; side of

scutellum somewhat indistinctly rugose; metanotum with 3 carinae medially and a

pair of submedial carinae (fig. 26); surface of propodeum coarsely transversely

reticulate-rugose, with a short carina anteriorly, situated in a weak depression.

Wings, —r l:r 2:r 3 = 29:49:139; cuqu 1:r 2: cuqu 2 = 38:49:26; r 3

weakly curved towards metacarp; metacarpella rather straight, weakly curved (fig.

25); SM with ca. 80 setae.

Legs. — Hind coxa weakly and remotely punctate; all tarsal claws absent;

length of femur, tibia and basitarsus of hind leg 10.1, 12.8 and 11.0 times their

width, respectively; length of spurs of hind tibia 0.6 and 0.5 times their basitarsus.

Metasoma. — Length of Ist tergite 5.5 times its apical width, its surface mainly

smooth in front of spiracle, behind spiracle rugose and with a medial crest-shaped

carina (fig. 19); whole Ist tergite distinctly convex; spiracles weakly protruding;

shape and length of ovipositor sheath unknown.

Colour. — Brownish yellow; tips of mandibles, apices of medial antennal

segments (of paratype), stemmaticum and its surroundings, wing veins and

mesoscutal lobes mainly, more or less dark brown; hind tarsus whitish yellow;

wing membrane somewhat brownish, more pronounced near sclerome (fig. 27).

Holotype in NR: “N. E. Burma, Kambaiti, 2000 m, 19/5, 1934, Malaise’’,

“Riksmuseum Stockholm”, “Holotype Xiphozele burmensis V. Sharma, 1974”. One

paratype: topotypic, NR, metasoma absent, length of fore wing 14.4 mm, SM with

ca. 30 setae, vertex coarsely punctate, colour as holotype, POL: @ ocellus : OOL =

12: 10:5, claws with a ventral lamella, somewhat more developed than in 9 of

compressiventris (fig. 16). Additionally examined 2 g from China: “Suifu, SZ.,

China, VI. 1—21, 1928, alt. 1000—1500, D. C. Graham Coll.” (USNM); tarsal

lamella enlarged in respect to the lamella of 9, about equally shaped as lamella of

& of compressiventris (fig. 13); length of metasoma 2.7 times length of mesosoma;

length of fore wing 13.8, of body 17.6 mm; metasoma infuscated apically, further

equally coloured as holotype; SM densely setose. Second male is from Shanghai

(1898, J. de Joannis, MNHN) with wings only weakly infuscate, SM with 24 setae,

clypeus less concave than in type, length of fore wing 12.1 mm and of body 16.1

mm.

Note. — X. burmensis is closely related to compressiventris and is rather variable.

The characters given by Sharma (1975: 175) are not suitable for separation of the

two species. Firstly because she confuses Cameron’s meaning of the term

“metanotum” (= propodeum) with the modern meaning of this term. Secondly

the length of the metasoma in respect to the mesosoma is variable; the male of

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

burmensis from China has a comparatively longer metasoma (2.7 times mesosoma)

even than several specimens of compressiventris (2.5—2.6 times). Finally the

differences in length of the body of parasites are usually unsuitable for species

separation, as proven again by the small series examined for this revision.

Distilirella gen. nov. (figs. 28—45)

Etymology: from “distantia” (Latin for “‘remoteness”) and “lirella” (Latin for a “small ridge’’),

because the prepectal carina is remote from the anterior edge of the mesopleuron. Gender: feminine.

Type-species: Distilirella curvinervosa spec. nov.

Diagnosis. — Apex of scapus slightly inclivous (fig. 28); occipital carina

reaching hypostomal carina ventrally, far above mandibular base (fig. 32); eyes not

emarginated, at most with a scarcely visible bend on the inner sides (fig. 40); teeth

of mandible rather slender (fig. 40); apical margin of clypeus straight medially;

prepectal carina remains far removed from anterior margin of mesopleuron (fig.

28); episternal scrobe deep and rather round (fig. 28); scutellum with a curved

lateral carina, which is absent posteriorly (fig. 43); scutellum narrowly sculptured:

posteriorly (fig. 43); cu 1 weakly sinuate (fig. 37); nervulus straight, only slightly

narrower than surrounding veins, slightly postfurcal and without a sclerome (fig.

33); SM setose, but in basal half less densely setose than in apical half; basella and

metacarpella strongly curved (fig. 45); tarsal claws of © indistinctly yellowish,

pectinate and with a submedial lamella at which another subapical lobe is situated

(figs. 34, 35); tarsal claws of ¢ bifurcate, without lamella (figs. 41, 42); laterope

very deep, almost touching each other (fig. 39); ovipositor sheath rather slender,

its sides subparallel (fig. 28).

Distribution. — Australian (New Guinea): one species.

Note. Apomorphous character-states of Distilirella are: 1—basella and

metacarpella strongly curved; 2—tarsal claws of 9 with double lamellae;

3—scutellum with a lateral carina. Additional plesio-morphous character-states

are: 1—nervulus straight and equally developed; 2—SM setose; 3—sclerome of

fore wing absent; 4—claws of ¢ bifurcate; 5—ventral half of occipital carina

present; 6—ovipositor sheath slender; 7—prepectal carina remains far removed

from anterior margin of mesopleuron.

Distilirella curvinervosa spec. nov. (figs. 28—45)

Holotype, 9, length of body 12.5, of fore wing 11.7 mm.

Head. — Antennal segments 47 (but apical segments absent), length of 3rd

segment 1.3 times 4th segment; length of 3rd and 4th segments 5.1 and 4.0 times

their width, respectively, and both penultimate segments absent (in allotype they

are 2.3 and 3.0 times their width and apical segment with a long spine (fig. 36));

length of maxillary palp 2.3 times height of head; dorsal length of eye 3.4 times

temple; temple roundly receding and punctulate (fig. 44); POL : @ ocellus : OOL

= 9:8:8; frons weakly concave and smooth; ventral half of occipital carina

completely present, reaching middle level of eye (fig. 32); face punctulate, weakly

VAN ACHTERBERG: Xiphozelinae 39

convex and with a small tubercle dorso-medially (fig. 40); clypeus convex,

punctulate; length of malar space 0.7 times basal width of mandible; malar suture

almost absent (fig. 40).

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

deeply depressed medio-anteriorly and remotely crenulate (fig. 28), its remaining

part mainly smooth; epicnemial area smooth, except for some crenulae; precoxal

suture with some spaced punctures, its surroundings indistinctly punctulate;

pleural suture narrowly crenulate, rather shallow and narrow; metapleuron

rugose-reticulate, but dorsally mainly smooth; notauli rather shallow, completely

and narrowly crenulate (fig. 43); mesoscutal lobes weakly punctulate; side of

scutellum smooth; metanotum with 2 parallel carinae submedially (fig. 43); surface

of propodeum mainly smooth between the carinae, with a long medial carina

anteriorly (fig. 39) and posteriorly with several more or less transverse carinae,

bordered by a carina latero-posteriorly.

Mines rr 23 = 17:20:60; d'1:d2:= 1:35; cuqu 1:1 2: cuqu'2 =

13 : 20: 12; r 3 strongly curved anteriad; radiella weakly curved basally (fig. 37).

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

9.9, 15.0 and 15.6 times their width, respectively; length of hind tibial spurs 0.6 and

0.5 times their basitarsus.

Metasoma. — Length of Ist tergite 6.4 times its apical width, its surface smooth,

flat in front of spiracles and convex behind spiracles; spiracles of Ist tergite slightly

protruding; length of ovipositor sheath 0.07 times fore wing.

Colour. — Brownish yellow; stemmaticum, ovipositor sheath and most wing

veins dark brown; flagellum and outer aspect of scapus infuscated; hind tarsus

whitish yellow.

Holotype in TC: “Wau, N. Guinea, October, 1969, P. Shanahan’’.

Paratype: 1 ¢ (allotype, RMNH): “Museum Leiden, Nieuw Guinea Exp.

K.N.A.G. 1939, Araboebivak, 6.X1.1939”. Antennal segments of allotype 54; frons

with some microsculpture laterally; nervellus with a short ramellus (d’), resulting

in a posteriorly broken nervellus (fig. 38), in left wing less developed than in right

wing; length of fore wing 11.8 mm, length of Ist tergite 6.1 times its apical width.

ACKNOWLEDGEMENTS

I wish to express my sincere thanks to the following persons for the loan of types

and/or gifts of unidentified specimens. The abbreviations used for the collections

are given in parentheses. Dr. V. K. Gupta, Dept. of Zoology, Delhi; Dr. K. J.

Hedqvist, Naturhistoriska Riksmuseet, Stockholm (NR); Mr. T. Huddleston,

British Museum (Natural History), London (BM); Dr. E. Kierych, Instytut

Zoologii, Warsaw (PAN); Dr. P. M. Marsh, USDA, U.S. National Museum,

Washington (USNM); Drs. M. Suwa & C. Watanabe, Entomological Institute,

Sapporo (EI); Dr. H. K. Townes, American Entomological Institute, Ann Arbor

(TC); Mr. B. Sigwalt & Dr. S. Kelner-Pillault, Muséum National d’Histoire

Naturelle, Paris (MNHN); (RMNH) = Rijksmuseum van Natuurlijke Historie,

Leiden.

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 2, 1979

LITERATURE

Achterberg, C. van, 1976a. A revision of the tribus Blacini (Hym., Braconidae, Helconinae). —

Tijdschr. Ent. 118 (7): 159—322, figs. 1—476.

——, 1976b. A preliminary key to the subfamilies of the Braconidae (Hym.). — Tijdschr. Ent.

119 (3): 33—78, figs. 1—123.

Brothers, D. J., 1975. Phylogeny and classification of the aculeate Hymenoptera, with special reference

to Mutillidae. — Univ. Kansas Sci. Bull. 50 (11): 483—648, figs. 1—101.

Cameron, P., 1906. On some Braconidae from the Himalaya. — Entomologist 39: 204—206.

Enderlein, G., (1918) 1920. Zur Kenntnis aussereuropaischer Braconiden. — Arch. Naturgesch. 84 (A,

11): 51—224, Figs. 111.

Gauld, I. D. & T. Huddleston, 1976. The nocturnal Ichneumonoidea of the British Isles, including a

key to the genera. — Entomologist’s Gaz. 27: 35—49, figs. 1—20.

Sharma, V., 1975. A new species of Xiphozele (Hym., Braconidae, Macrocentrinae). — Oriental Ins.

9 (2): 173—175, figs. 1—S. i

Shenefelt, R. D., 1969. Hymenopterorum Catalogus (nov. ed.) Part 4. Braconidae 1:1—176. — Junk,

’s-Gravenhage.

Watanabe, C., 1969. Notes on the genera Zele Curtis and Xiphozele Cameron, with special reference to

the species in Japan (Hym., Braconidae). — Proc. ent. Soc. Wash. 71 (3): 318—328, figs. 1—9

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INHOUD

_ W.J. KLOFT, R. E. WooDRUFF & E. S. KLOFT. — Formica integra (Hymenoptera:

Formicidae) IV. Exchange of food and trichome secretions between worker

ants and the inquiline beetle, Cremastocheilus castaneus (Coleoptera:

Scarabaeidae), p. 47—57, text-figs. 1—20.

RI Tijdschrift voor Entomologie, deel 122, afl. 3 Gepubliceerd 31-V-1979

FORMICA INTEGRA (HYMENOPTERA: FORMICIDAE) IV.

EXCHANGE OF FOOD AND TRICHOME SECRETIONS

BETWEEN WORKER ANTS AND THE INQUILINE BEETLE,

CREMASTOCHEILUS CASTANEUS (COLEOPTERA:

SCARABAEIDAE)

W.J. KLOFT', R. E. WOODRUFF? and E. S. KLOFT!

With 20 figures

ABSTRACT

Experimental evidence, using radioactive tracers, confirms the role of Cremastocheilus castaneus as a

predator in the ant nests, as well as proves the existence of trichomes on the beetle which supply

substances transmitted by ants through the social organization of the nest. A specific new ventral

trichome area was discovered on the beetle, using this technique. The trichome areas are illustrated

with scanning electron microscope photographs.

To conduct studies on biology, morphology, and foraging behavior of an ant,

Formica integra Nylander, we transferred two nests of this forest ant from its

natural habitat in Georgia (for description of locality, see Kloft et al., 1973) to the

Forest Insect Research Laboratory, University of Florida, Gainesville, Florida in

1972. This ant, of the subfamily Formicinae, is the southernmost representative of

the Formica rufa group in eastern United States. The studies about the possibilities

of an introduction of this ant into Florida forests could be continued; two more

papers are cited (Wilkinson et al., 1978, 1979, in press).

During our laboratory experiments, three specimens of a scarab beetle,

Cremastocheilus castaneus Knoch, flew from the nests established in open arenas in

a fully climatized laboratory (14 hrs light, 25 + 2° C, 50—80% rel. air humidity).

Possibly the change in environmental conditions between the natural colony and

the laboratory induced emigration of the beetles.

The genus Cremastocheilus Knoch has long been in need of revision. The most

recent general paper is that of Potts (1945), which provides a key to North

American species. Because of the numerous short scattered setae on the

pronotum and the geographic distribution, our specimens would key to

C. castaneus brevisetosus Casey. Potts (1945: 74) stated: “Casey describes

! Institute of Applied Zoology, University of Bonn, Germany.

? Bureau of Entomology, Division of Plant Industry, Florida Dept. of Agriculture, Gainesville, FL

32602 (Contribution No. 350).

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 3, 1979

brevisetosus from a specimen he records from Iowa. However, the only specimens

before me which agree are from Alabama, and I am led to suspect an erroneous

label on the Casey specimen. In the Alabama specimens the setae of the pronotum

are exceedingly broad and short and as the specimens are quite fresh, the

character is presumably a good one. This appears to me to be the most distinctive

race of any I have seen, and if this name is applicable to the southern specimens it

apparently represents a valid subspecies. The value of the other two names as

weak races of castaneae seems questionable to me. A long series from the Rocky

Mountains is not easily separable, although in the main, they most closely agree

with Casey’s pocularis.”

Figs. 1-2. Outline drawings of Cremastocheilus castaneus brevisetosus: 1) dorsal view; 2) lateral view:

a) anterior pronotal trichome area; b) location of prosternal apophysis with trichomes; c) posterior

pronotal trichome area; d) propygidial spiracle. (Scale line equals 2.5 mm).

The genus Cremastocheilus, which comprises more than 40 species, isendemic to

North America. All known species live as adults in the nests of ants of 11 genera;

C. castaneus has been recorded in association with Formica and Polyergus (Cazier

& Statham, 1962). Wheeler (1910, 1928) reported it from F. integra nests. The first

KLOFT, WOODRUFF & KLOFT: Cremastocheilus castaneus 49

monographic revision was given by Horn (1879); Casey (1915) described many

novelties and revised the North American Cremastocheilini together with other

groups of Cetoniinae. The species-level review by Potts (1945) has already been

mentioned. Howden (1971) and Krikken (1976), proposing new genera, presented

keystothe Western Hemisphere genera of Cremastocheilini.

Cazier & Statham (1962) pointed out that the true relationship between species

of the genus Cremastocheilus and their host ants is not yet known. In observations

of a western species (C. stathamae Cazier) in nests of the honey-ant, Myrmecocystus

mexicanus Wesmael, Cazier & Mortenson (1965) found these beetles were obligate

predators of the ant larvae. At the time this was the first recorded case of primarily

predatory behavior within the family Scarabaeidae. W. M. Wheeler (1910)

described the trichomes on the anterior and posterior prothoracic angles of the

beetles and presented a figure of a Formica integra worker gnawing on one of the

trichomes (see also this figure reproduced in Wilson (1971, fig. 20—4)). Alpert &

Ritcher (1975) stated that C. armatus Walker adults were predaceous on both ant

larvae and pupae, and that “Beetles were not disrupted from feeding even when

covered with attacking ants.”’

Some of the observations of Cazier & Mortenson (1965) appeared very

contradictory to them. As Wilson reported (1971: 390) the ants treated the beetles

sometimes as synechtrans and sometimes as symphiles. Most of the time, however,

they had the status of synoeketes; 1.e., they were ignored and allowed to wander

through the nest without interference.

Tracer studies on exchange of food and trichome secretions between

ants and beetles

As shown by Goesswald & Kloft (1958, 1960, 1963) radioisotopes are a useful

tool to elucidate interrelations within societies of social insects. Hoelldobler (1967,

1970) used 5P to demonstrate the physiology of guest-host relations (myr-

mecophily) in ants. We used radioactive food (20% sucrose solution + Na,H”PO,,

specific activity 0.5 mCi/ml) to trace social food exchange in worker groups of

Formica integra. All experiments were conducted in 1972, with dates shown.

EXPERIMENT I

7.vi: Worker ants were fed with **P honey and after 18 hrs checked under an

end-window G. M. tube. The detector was connected with a Berthold-Frieseke

Scaler-Timer System BF 22/25. Before checking, the insects were carefully

decontaminated with a “chaser” solution plus detergent (Kloft, 1977). The ants

had an average radioactivity of 20,000 CPM.

20.vi: At 12:00 noon one beetle (A) was put together with 10 radioactive-fed ants

of F. integra; no food was added. At 3:45 PM the beetle was activity besieged by

the ants which palpated it intensively with their antennae. Ants were sitting on and

underneath the beetle’s head, gnawing on the trichomes. Since we supposed food

transfer from the ants to the beetle, we checked the beetle at 3:50 PM for

radioactivity. The geometry was like that used for the measurements of the ants.

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 3, 1979

Result: After 3 hrs 50 min the beetle showed 3033 CPM. We checked at 5:30 PM

and the activity was 3124 CPM or about the same. All ants were in good condition.

21.vi: At 9: 00 AM the beetle was in good condition, but all ants were dead, the

gaster squeezed out. In some cases we found parts of the dead ants, presumably

dismembered by the beetle.

EXPERIMENT II

22.vi: One beetle (B) was put with 10 radioactive-fed ants. The beetle displayed

a death feint (Totstell-Verhalten), but was palpated by the ants. The palpation

itself did not lead to contamination, as shown by subsequent checkings.

23.vi: After 22 hrs the beetle showed a total activity of 1600 CPM, and all 10 ants

were in good condition. Presumably the beetle must have participated in the social

distribution of radioactive-labelled food. Four hours later, after a total time of 26

hrs, the beetle had about the same rate of counts. However, 2 ants were found

dead, and one of these appeared to be lacerated by the beetle.

In both experiments the ants were not only attracted to the trichomes on the

anterior and posterior prothoracic angle of the beetles, but also soliciting, by

antennal movements, certain parts of the sternum. These observations led to the

detection, for the first time, of ventral hair tufts (trichomes) on the prosternal

apophysis, described in the following part. As best we could observe, the beetle

lowered the anterior portion of the mentum, which completely covers the mouth

parts. The palpi could be seen moving in and out. We believe that by such

movements the regurgitation by the ant is released. The food transfer could not be

observed directly, but was evident by the increase of the beetle’s radioactivity long

before any ants were killed or squeezed out.

EXPERIMENT III

22.vi: Beetle (A) was carefully decontaminated and subsequently checked for

radioactivity. At 11: 45 AM it had 20.810 CPM. Certain inherent problems are

encountered in radio-isotope work. High counts are found when the radioactivity

is broadly distributed. When the radioactivity is localized, lower counts result,

even if the absolute radioactivity is the same. Equal distribution of the °P,

Table |. Transfer of radioactivity from a radioactive labelled Cremastocheilus castaneus to worker ants

of F. integra via the trichome secretions within 3 hrs.

COUNTS COUNTING DURATION CPM LI SORTE

(MIN) MIN

BACKGROUND 61 5 1252 + 3.49

BACKGROUND 43 5 8.6 + 2.93

ANT A 155 5 31 + 5.56

ANT B 121 5 AD + 4.92

KLOFT, WOODRUFF & KLOFT: Cremastocheilus castaneus 51

throughout the hemolymph, is probably responsible for this high count, although

body absorption could have affected the counts (Kloft, 1962). This complete

distribution within the body was a precondition for the following experiment: the

radioactive-labelled beetle (A) was put together with 10 nonradioactive worker

ants. The object was to determine if there were transfer of radioactivity, via

trichome secretions, from the beetle to the ants.

22.vi: The ants had been around the trichomes gnawing on the hair tufts. At 2: 45

PM, after a total elapsed time of 3 hrs, the ants were checked for radioactivity

(table 1). The ants definitely gathered radioactivity from the beetle, undoubtedly

derived from trichome secretions. The next question was whether these radio-

active-labelled secretions could be spread to other ants by social food distribution.

To determine this, the ant (A) was enclosed for 20 hrs in a small container (surface

1.5 cm?) with 2 ants of the same colony. During this time no food was added. The

results are shown in table 2. The donor ant, which was in direct contact with the

radioactive beetle, dispersed radioactive material by social food distribution. The

reason that the sum of counts of the donor ant (plus acceptor ants) is higher than

the original rate of the donor ant is a result of the technical problem discussed

earlier (Kloft, 1962).

Table 2. Transfer of radioactivity, gathered from a radioactive Cremastocheilus castaneus (table 1) to

further ants via regurgitation within 3 hrs.

COUNTS COUNTING DURATION CPM + SORT:

(MIN) MIN

BACKGROUND 56 5 11622 + 3.34

BACKGROUND 48 5 9.6 + 3.09

DONOR ANT

= ANT A FROM

TABLE I 97 5 19.4 + 4.40

1. ACCEPTOR

ANT 81 5 1662 + 4.02

2. ACCEPTOR

ANT 61 5 1252 + 3.49

Detection of further trichome areas on the venter of Cremastocheilus

Ever since Cremastocheilus has been known to inhabit ant nests, there has been

speculation about the glandular nature of several morphological structures. This

has usually centered around the large tufts of coarse setae (trichomes) located on

the anterior (figs. la, 2a) and posterior (figs. Ic, 2c) angles of the pronotum

(Hoelldobler, 1971). Other areas that have been previously suggested include the

enlarged and projecting propygidial spiracles (figs. 1d, 2d), although there is no

evidence for it.

It was therefore quite surprising when the structure located between the

anterior coxae (fig. 2b) was noted as attractive to the ants during this study. This

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 3, 1979

Figs. 3—8. Prosternal apophysis of C. castaneus brevisetosus (anterior legs removed): 3) postero-ventral

view (45 x); 4) enlargement of fig. 3 (95 x); 5) enlarged tip of apophysis (925 x ); 6) sensors on margin

behind apophysis, enlargement of area at arrow in fig. 3 (750 x); 7) lateral view of apophysis (100 x );

8) ventral or head on view of apophysis (100 x).

structure (figs. 3—8) has been termed a prosternal apophysis by Krikken (1976). It

projects forward toward the notch in the posterior margin of the mentum (fig. 19)

and would appear to fit there if the head or mentum were extended down or back.

The apophysis is fringed with very long sensors (figs. 3—5, 7—8) which appear to

be the trichomes sought by the ants.

During removal of the legs in preparation for the scanning electron microscope

studies, an additional group of sensors was located beneath the anterior coxae

(figs. 3 arrow, 6). Although previously undetected and well-hidden, the compli-

KLOFT, WOODRUFF & KLOFT: Cremastocheilus castaneus 53

Figs. 9—12. Anterior pronotal trichome area (fig. la) of C. castaneus brevisetosus: 9) dorsal view of right

side (120 x), area at arrow enlarged in fig. 11; 10) ditto, angle more dorsal (150x); 11) enlargement of

trichomes at arrow in fig.9(525x); 12) ditto (1000 x).

cated nature of the sensors (fig. 6) suggests that they serve an important function.

Unfortunately the exact behavior of the ant solicitations and the beetle’s feeding

are nearly impossible to observe carefully. Alpert & Ritcher (1975: 289) stated that

“when feeding, a beetle lowered its mentum and pierced an ant larva with its sharp

maxillae. The mandibles are greatly reduced and aid in the transport of fluid”.

The prominent propygidial spiracles (figs. Id, 2d) are a possible source of

secretions because of their elevated nature and the shape and size of the opening

(figs. 17, 18, 20). It is possible that attractant chemicals are produced, although

chemicals with a totally different function could be involved. Repellents would

certainly be advantageous to a commensal if the host ants became inhospitable

and agressive. Alpert & Ritcher (1975: 288) found that “If the beetle was violently

disturbed or turned over by the ants, a droplet of viscous fluid was released from

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 3, 1979

the anal opening. This fluid had an offensive odor and was effective in repelling

the ants”. They did not mention any observations on the nature of the propygidial

spiracles.

The pronotal trichomes are so prominent, and unique to the tribe Crema-

stocheilini, that their role cannot be overemphasized. The sensors, or individual

trichomes, appear to be gnawed upon by the ants (figs. 15—16). Only then is it

noticeable that they are hollow (fig. 16). The several shapes of trichomes nearly fill

the depressions surrounding them (figs. la, Ic, 2a, 2c, 10, 14). The anterior ones

(fig. la) are accessible to the ants from above; the posterior ones (fig. Ic) primarily

accessible from the side; and those on the prosternal apophysis are accessibly only

ventrally. i

Figs. 13—16. Posterior pronotal trichome area (fig. Ic) of C. castaneus brevisetosus: 13) ventral view of

left side (55x); 14) ditto (105x); 15) gnawed trichomes (535 x); 16) ditto, individual hollow seta of the

trichome (1050 x).

DISCUSSION

According to our results, Cremastocheilus castaneus seems to play a double role

within the Formica integra community. It is first a predator which kills ants,

squeezes out juices, and feeds on the contents of the gasters of workers. It also cuts

the ants into pieces, possibly feeding on these parts. Since we experimented only

with adult workers we couldn’t observe feeding on ant larvae, as described in

KLOFT, WOODRUFF & KLOFT: Cremastocheilus castaneus 55

detail by Cazier & Mortenson (1965) and Alpert & Ritcher (1975). But in spite of

being well protected in the sensitive mouth area by the anterior portion of the

mentum, which completely covers the mouthparts (fig. 19), the beetles should not

simply be regarded as synechtrans. They are, according to Wilson (1971),

symphiles (so-called “true” guests). We determined that they are not only

groomed but also participated in the social food exchange within the ant colony.

The additional trichome areas on the prosternal apophysis help in releasing the

food sharing behavior through the ants.

We obtained evidence of the transfer of radioactive material from the beetle to

the ants, presumably via trichome secretions. As we expected, an ant which was in

contact with a radioactive beetle also distributed the radioactivity, through social

food distribution, to other ants which were never in contact with the radioactive-

labelled beetles. Thus the “peace-making’”’ allomones (Pasteels, 1977) of the

beetle might be distributed to wider parts of the nest population. Possibly during

the reproductive activities of the beetles, the production of the attractive and

“peacemaking” secretions stop (or may be masked by sex pheromones which are a

deterrent for the ants). This would explain the observations of Cazier & Statham

(1962) that the beetles are pulled out of the nest and dumped in the refuse zone. In

Figs. 17—20. Cremastocheilus castaneus brevisetosus: 17) propygidial spiracle, lateral view (210x);

18) ditto, head on view (340 x ); 19) head, latero-ventral view (26 x ); 20) enlargement of propygidial spi-

racular opening of fig. 18 (850 x ).

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 3, 1979

the transitional stage, other workers of the same colony might attempt to pull the

beetle back in the direction of the nest. This dumping and pulling back lasts a

certain time, but finally the beetles fly away. Hoelldobler (1971) showed that

myrmecophiles communicate in the same chemical language as their hosts. He

defined trichomes as “... tufts of hairs that serve to increase the ability of well-

integrated ant guests to communicate chemically. They are located on the ventral

surface of highly modified anterior and posterior pronotal projections of

Cremastocheilus adults”. Mating and breeding areas are not yet known, but mating

outside the colonies would be advantageous for gene flow.

ACKNOWLEDGMENTS

We thank the following persons for their valuable assistance: Thelma Carlysle

(for the excellent SEM photos); Dr. J. Krikken (for reading the manuscript); Dr.

R. C. Wilkinson (for colony maintenance and other favours); and the Dept. of

Entomology, University of Florida (for travel funds and facilities for the senior:

author).

BIBLIOGRAPHY

Alpert, G. D., & P. O. Ritcher, 1975. Notes on the lifecycle and myrmecophilous adaptations of Cre-

mastocheilus armatus (Coleoptera: Scarabaeidae). — Psyche 82 (3—4): 283—291; 5 figs.

Casey, T. L., 1915. A review of the American species of Rutelinae, Dynastinae, and Cetoniinae. —

Mem. Coleoptera 6: 1—394.

Cazier, M. A., & M. Statham, 1962. The behaviour and habits of the myrmecophilous scarab, Cremas-

tocheilus stathamae Cazier, with notes on other species (Coleoptera: Scarabaeidae). — J. New

York Ent. Soc. 70: 125—149.

Cazier, M. A., & M. A. Mortenson, 1965. Bionomical observations on myrmecophilous beetles of the

genus Cremastocheilus (Coleoptera: Scarabaeidae). — J. Kansas Ent. Soc. 38: 19—44.

Goesswald, K., & W. Kloft, 1958. Radioaktive Isotope zur Erforschung des Staatenlebens der Insekten.

— Umschau 58: 743—745.

Goesswald, K., & W. Kloft, 1960. Neue Untersuchungen ueber die sozialen Wechselbeziehungen im

Ameisenvolk, durchgefuehrt mit Radioisotopen. — Zool. Beitr. N. F., 5: 519—559.

Goesswald, K., & W. Kloft, 1963. Tracer experiments on food exchange in ants and termites. In: Radia-

tion and radioisotopes applied to insects of agricultural importance, p. 25—42. — Int. Atomic

Energy Agency, Vienna.

Hoelldobler, B., 1967. Zur Physiologie der Gast-Wirt-Beziehungen (Myrmecophilie) bei Ameisen. I.

Das Gastverhaltnis der Atemeles- und Lomechusa-Arten (Coleoptera: Staphylinidae) zu Formi-

ca (Hym., Formicidae). — Z. Vergl. Physiol. 56: 1—21.

Hoelldobler, B., 1970. Die Physiologie der Gast-Wirt-Beziehungen (Myrmecophilie) bei Ameisen. II.

Das Gastverhältnis des imaginalen Atemeles publicollis Bris. (Coleoptera: Staphylinidae) zu

Formica und Myrmica (Hym., Formicidae). — Z. vergl. Physiol. 66: 215—250.

Hoelldobler, B., 1971. Communication between ants and their guests. — Scientific Amer., Mar., 1971:

86—91.

Horn, G. H., 1879. A monographic revision of the species of Cremastochilus of the United States. —

Proc. Amer. Philos. Soc. 18 (104): 382— 397; pl. 4.

Howden, H.F., 1971. Key to the New World Cremastocheilini, with notes and description of a new ge-

nus. — Proc. Ent. Soc. Washington 73 (2): 224-230,

Kloft, W., 1962. Technical problems of radioisotope measurement in insect metabolism. In: Symposi-

um (Bombay, 1960) “Radioisotopes and radiation in entomology”, p. 163—172. — Int. Atomic

Energy Agency, Vienna.

KLOFT, WOODRUFF & KLOFT: Cremastocheilus castaneus 57

Kloft, W. J., R. C. Wilkinson, W. H. Whitcomb, and E. S. Kloft, 1973. Formica integra 1. Habitat, nest

construction, polygyny and biometry. — Florida Ent. 56 (2): 67—76.

Kloft, W. J., 1977. Part V. Applied part. In: Laboratory manual on use of radioisotopes in entomology,

2nd ed., p. 141—220. — Int. Atomic Energy Agency, Vienna..

Krikken, J., 1976. New genera of New World Cremastocheilini, with revisional notes (Coleoptera: Ce-

toniidae). — Zool. Meded. 49 (25): 307—315; 16 figs.

Pasteels, J. M., 1977. Evolutionary aspects in chemical ecology and chemical communication. — Proc.

Int. Congr. Ent. 15: 281—293.

Potts, R. W. L., 1945. A key to the species of Cremastocheilini of North America and Mexico (Coleop-

tera, Scarabaeidae). — Bull. Brooklyn Ent. Soc. 40 (3): 72—78.

Wheeler, W. M., 1908. Studies on myrmecophiles. I. Cremastocheilus. — J. New York Ent. Soc. 16:

68—79.

Wheeler, W. M., 1910. Ants: their structure, development and behaviour. Columbia Univ. Press, New

York: xxv + 663 p.

Wheeler, W. M., 1928. The social insects: their origin and evolution: xviii + 378 p. — Kegan Paul

Trench, Trubner & Co., Ltd., London.

Wilkinson, R. C., A. P. Bhatkar, W. J. Kloft, W. H. Whitcomb, and E. S. Kloft, 1978. Formica integra II.

Feeding, trophallaxis, and interspecific confrontation behavior. — Florida Ent. 61 (3):

179— 187.

Wilkinson, R. C., A. P. Bhatkar, W. H. Whitcomb and W. J. Kloft, 1979. Formica integra III. Trial in-

troduction into Florida. — Florida Ent. (In press).

Wilson, E. O., 1971. The insect societies: 1—548. — Belknap Press, Harvard Univ., Cambridge, Mass.

Sn MUS. COMP, ZOOL.

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calyptris Benin and Fomoria Beirne occurring in ite United States ni America

Re dopter, Nepticulidae), p. 59—90, fig. 1—26.

Gepubliceerd 31-V-1979

A TAXONOMIC STUDY OF THE MICRO-LEPIDOPTERAN

GENERA MICROCALYPTRIS BRAUN AND FOMORIA BEIRNE

OCCURRING IN THE UNITED STATES OF AMERICA

(LEPIDOPTERA, NEPTICULIDAE)

CHRISTOPHER WILKINSON

Department of Animal Systematics and Zoogeography, Free University, Amsterdam

With 26 text-figures

ABSTRACT

The nearctic species of two genera of leaf-mining microlepidoptera are here revised as a prelude toa

revision of the North American Nepticulidae. The concept of Microcalyptris Braun is widened from

what was known as a monotypic genus to one having eight species.

Fomoria Beirne can no longer be regarded as solely Palaearctic for here the true relationship of two

species is shown in their transfer from ‘‘Nepticula’’ von Heyden. Generic and species diagnoses, key,

descriptions, and details of genitalia are given where necessary.

INTRODUCTION

In this revision eight species of Microcalyptris are discussed, of which two have

been transferred from “‘Nepticula’”’ von Heyden and five are new species, three of

which are named and described here. Until last year Microcalyptris was known as a

monotypic genus from a single specimen.

The genus Fomoria contains two species in North America and these, too, are

transferred from “Nepticula” von Heyden and presented as new combinations

here.

ABBREVIATIONS

L.A.Co.M. Los Angeles County Museum of Natural History, California, USA.

USNM United States National Museum of Natural History, Smithsonian

Institution, Washington D. C., USA.

ANS Academy of Natural Sciences, Philadelphia, USA.

MCZ Museum of Comparative Zoology, Cambridge, Massachussets,

USA.

CNC Canadian National Collection, Ottawa, Canada.

FUA Free University of Amsterdam, Netherlands.

The methods and abbreviations used are similar to those given in Wilkinson &

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Scoble (1979: 2), except that the letters ANS are used to mean the Academy of

Natural Sciences, Philadelphia, USA.

Scale lines on figs. are 0.1 mm unless stated otherwise.

Microcalyptris Braun, 1925

Microcalyptris Braun, 1925b: 224. Type species by monotypy: Microcalyptris scirpi Braun, 1925b: 225.

The genus shows one of the more simple patterns of venation found in the family

Nepticulidae. Other features characteristic of the genus are the peculiar lateral

arms associated with the vinculum and probably part of the gnathos; the very

slender valves; and the juxta, which is extremely complex in some cases. The

female genitalia often show complex sclerotisations of the ductus with the signa,

usually, as linear rows of plates or single spiculate cells.

Taxonomic History

The genus was described from a single specimen collected by Braun in Utah

(loc. cit.). Since that description there are no other published records of the genus

until now, even though some new species are described here from Braun’s ma-

terial.

Braun’s description of the venation of this genus is somewhat incomplete owing

to the fact that she was reluctant to damage the single specimen available to her.

However, Dr. D. Davis of the USNM and I have examined the specimen which

confirms Braun’s description in part but also shows that a vestige of the Cubitus is,

in fact, present on the forewing. This is also confirmed by our work on other

species. Braun was also unable to provide a description of the hindwing venation;

this information is also provided in this revision.

Generic description

External features: 4 ©. Head: palps extending well beyond labrum, pale grey or

white; antennae extending half the length of the forewing, fuscous; tuft on front of

head usually ochreous, sometimes white or brown, vertex concolorous in most

cases; eye-caps and collar ochreous, sometimes white or brown. Thorax pale white

or buff and sometimes irrorate with brown. Abdomen usually concolorous with

thorax, shining silver beneath. Venation as in fig. 11. Forewings: media coalescing

with Radius at base and anastomosing as far as the middle of the wing; Cubitus

vestigial; R, coincident with R,; Anal vein not reaching the margin. Hindwings:

Media single branched. Forewings: narrow and lanceolate, ground colour of

dorsal surface grey or buff often with each scale brownish at the tip; fringe greyish,

variously irrorate with wing scales apically; markings variable, usually with a single

fascia or two patches. Hindwings: narrow and lanceolate, half width of forewings;

usually grey and iridescent, sometimes with brightly coloured specialised scales on

both surfaces concolorous, with similar patches on ventral surface of forewings.

Legs: grey or brown, sometimes with scattered paler areas; proximal pair of spurs

on hind-tibiae below the middle.

WILKINSON: Microcalyptris and Fomoria 61

Figs. 1—10. External features. Fig. 1. Microcalyptris scirpi, male. Fig. 2. M. thoracealbella, male. Fig. 3.

M. postalatratus, male. Fig. 4. M. punctulata, female. Fig. 5. M. distaleus, male. Fig. 6. M. bipinnatellus,

female. Fig. 7. M. bicornutus, male. Fig. 8. M. tenuijuxtus, male. Fig. 9. Fomoria pteliaeella, male.

Fig. 10. F. hypericella, male.

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Male genitalia: vinculum always ring-shaped; tegumen fused with vinculum

dorsally and produced into narrow, tapering pseuduncus sometimes weakly

bilobed or bluntly rounded. Pseuduncus membraneous and uncus sclerotized in

the form of a bridge usually with medial process spatulate, sometimes narrowly

pointed or papillate. Gnathos with complex anterior and posterior projections and

unique sclerotised lateral arms of vinculum in all species except possibly distaleus

sp.n. Saccus usually markedly bilobed. Valves slender, usually bluntly rounded.

Transtillae forming an inverted U-shape; transverse bars continuous. Juxta often

complex and heavily sclerotised. Aedeagus usually long and slender with large

spine-like cornuti and anellus.

Female genitalia: Posterior apophyses at least as long as the ductus, sometimes

terminally sagittate. Colliculum usually funicular and weakly sclerotised. Ductus

spiculate, usually with complex sclerotisations in the form of fin-like plates. Bursa

copulatrix: large and irregularly pectinate; signum double, usually comprising

linear rows of spinose cells or plates and, in a single case, ovate patches of

reticulate cells.

i R043

A Cu

Fig. 11 Microcalyptris sp. Wing venation.

Generic differential diagnoses

Characters which differentiate North American genera of the family Nep-

ticulidae are given.

Microcalyptris Braun, 1925.

Venation: reduced; Media of forewing coalescing with Radius from base and

anastomosing to a point beyond the middle of the wing; R, coincident with Rs;

Cubitus vestigial; Media of hindwing single, unbranched. Ground colour of dorsal

surface of forewing usually pale and variously irrorate. Proximal pair of spurs on

hind tibia below middle. Male genitalia, with membranous pseuduncus and

WILKINSON: Microcalyptris and Fomoria 63

strongly sclerotised bridge-like uncus; sclerotised gnathos with complex anterior

and posterior projections; lateral arms of vinculum usually with associated

sclerotisations. Female genitalia with complex sclerotisations of the ductus;

posterior apophyses very long, longer than the ductus; signa usually comprising

linear row of spinose cells or plates. Larvae mining leaves.

Stigmella Schrank, 1802.

Venation: Media of forewing coalescing with Radius at base and anastomosing

to a point beyond the middle of the wing; R, coincident with R,; Cubitus arising

separately, approaching middle of the wing; Media of hindwing single. Forewings

usually uniform and dark in colour, with one or two complete fasciae or patches;

fringe with diffuse margin. Proximal pair of spurs on hind-tibiae above the middle.

Male genitalia usually with U-shaped vinculum; tegumen strap-like, articulating

with vinculum dorsally; uncus bilobed; juxta, if present, membranous; aedeagus

usually flask-shaped, vesica usually with many denticulate cornuti orientated in a

ridge and rarely with platelike cornuti at the anellus. Female genitalia with simple

ductus and accessory sac; bursa copulatrix usually without signum, but if present

often single and weakly sclerotised. Larvae mining leaves of trees and shrubs and

sometimes herbs.

Ectoedemia Busck, 1907.

Venation: Media of forewing coalescing with Cubitus at base, passing obliquely

to Radius at or beyond R,,, and anastomosing to a point beyond middle of wing;

R, and R, separate; Cubitus usually approaching margin; media of hindwing

single. Proximal pair of spurs on hind-tibiae sometimes in the middle. Male

genitalia with gnathos W- or V-shaped, may vary according to method of

mounting; vinculum ring-shaped and without associated lateral bars; tegumen

extended into tapering or lobed pseuduncus; uncus absent or weakly mem-

branous; valves inwardly curved distally sometimes with digitate setae; juxta

absent; aedeagus regular in shape with elaborate cornuti and usually anellar

spines. Female genitalia with or without complex sclerotisation of the ductus and

spiculate accessory lobe; apophyses shorter than ductus; signa comprising patches

of reticulate cells. Larvae may mine in, or form galls on leaves, petioles, bark or

cortex.

Fomoria Beirne, 1945.

Venation: Media coalescing with Cubitus from base, both passing obliquely to

Radius at R,,, and anastomosing to beyond middle of wing; Cubitus becoming

obsolete; R, and R, separate; Media of hindwing single, unbranched. Male

genitalia with membranous pseuduncus and uncus as a spatulate sclerotisation; Y-

or V-shaped gnathos; saccus weakly bilobed; valves sometimes with dorsal spine;

aedeagus regular in shape and usually with complex anellar spines and cornuti.

Female genitalia, colliculum with sclerotised funicular antrum or complex plates;

simple ductus; signa comprising rather linear patches of reticulate cells. Larvae

often recorded pupating within the leaf-mine.

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Obrussa Braun, 1915.

Venation: Media of forewing coalescing with Cubitus at base, both passing

obliquely to Radius at R,,, and anastomosing to beyond middle of wing; Media

and Cubitus separate terminally; R, and R, separate; Media of hindwing single.

Ventral surface of forewing and dorsal surface of hindwing in males with patch of

brightly coloured specialised scales. Proximal pair of tibial spurs below middle of

hind-tibia. Male genitalia with ring-shaped vinculum; tegumen extended into

tapering pseuduncus; convex saccus; valves blunted distally and each with large

dorsal arm projecting transversely to reach opposite side of capsule; vesica with

transverse plate expanded laterally. Female genitalia with plate-like sclerotisation

at the colliculum; signa comprising ovate reticulate patches. Larvae only known to

mine fruits of Acer spp.

Glaucolepis Braun, 1917.

Venation: Media of forewing coalescing with Cubitus at base, both passing

obliquely to Radius at R,,, and anastomosing to beyond middle of wing; Media

and Cubitus separate terminally; R, and R, separate; Media of hindwing bifurcate.

Hindwing of male with patch of brightly coloured specialised scales. Proximal pair

of spurs on hind-tibiae in the middle. Male genitalia with tegumen extended into

tapering pseuduncus; gnathos with large transverse arms and medial dorso-lateral

arms fusing terminally; valves markedly bifurcate distally; aedeagus with lateral

cornuti extending full length of vesica and digitate distally. Female genitalia with

simple ductus; signa comprising linear patches of rows of pectinations. Larvae

mining leaves.

Oligoneura Davis, 1978.!)

Venation: greatly reduced; only two branches of Radius present; Media

unbranched and arising from stem of R,,,; Cubitus absent; hindwing extremely

slender and Media unbranched. Forewing dark fuscous with a single, narrow, pale

golden yellow fascia at distal third. Proximal pair of spurs on hind tibiae near apex.

Male genitalia with uncus vestigial; gnathos well developed, complex, consisting

of two, largely separate, transverse sclerites of a highly irregular, but symmetrical

outline; vinculum broad, quadrate; aedeagus moderately short and stout, with a

relatively complex apex and no cornuti. Female unknown. Presumably a leaf

mining genus. The type-species mines Coccoloba uvifera (L.).

Artaversala Davis, 1978.

Venation: greatly reduced; Radius unbranched, terminating well short of apex;

Media unbranched, extending almost to apex; Cubitus shortened, indistinctly

present; hindwing extremely slender and Media unbranched. Forewings with a

pale yellow to whitish apex and a single, broad, median fascia. Proximal pair of

spurs on hind-tibiae near apex. Male genitalia with tegumen reduced to an

') The name Oligoneura is preoccupied by a Dipteran genus and must therefore be changed. In dis-

cussion with Dr. Davis it was decided to publish a replacement name in a subsequent volume of the

Florida Entomologist.

WILKINSON: Microcalyptris and Fomoria 65

extremely slender dorsal ring; uncus absent; vinculum well-developed and V-

shaped; valves deeply divided and aedeagus relatively simple, without cornuti.

Female genitalia with slender and elongate ductus; bursa copulatrix membranous;

signa absent. Larvae mining leaves.

Checklist to the species of Microcalyptris

Microcalyptris Braun, 1925b.

scirpi Braun, 1925. Type species by monotypy.

thoracealbella (Chambers, 1873),

= badiocapitella (Chambers, 1876).

specimen 11.

punctulata (Braun, 1910).

specimen 12.

bipinnatellus sp.n.

postalatratus sp.n.

distaleus sp.n.

bicornutus Davis, 1978.

tenuijuxtus Davis, 1978.

Microcalyptris. Key to species. Males and females

1. Forewing with background predominantly dark; may have pale fascia or

Patches dtt DELETE SMEG BIRD LOS SUR EMI DER BREED dI 2

— Forewing with background predominantly light, may have dark fascia or

OASI ee le ar AB ee >

ER orewingidark without fascia or patches ..........%......2..+.- 3

Eorewing dark with pale fascia or patches ........2..2...2.....2.. 4

3. Male genitalia with H-shaped juxta, deeply bifurcate saccus and flattened

Pseudumeus si)... «> INA ah ES bicornutus (p. 81)

— Male genitalia without H-shaped juxta, saccus only weakly bifurcate,

pseuduncus convex terminallyasinfig. 19 ......... postalatratus (p. 77)

4. Forewing usually with | complete pale fascia and | broken in centre leaving 2

light patches. Pseuduncus of male bifurcate and uncus not extending beyond

it, juxta mace-shaped with cornuti; 4th abdominal sternite with two patches of

long setae; as in fig. 13c. Female genitalia with one long pair and one short

pair of apophyses; signa less than 3 cells wide as in fig. 14 .............

ze 08 Be done RP ENS 2 RE MeL ca an Ne es thoracealbella (p. 67)

— Forewing usually with 2 large pale fasciae and terminal spot. Pseuduncus of

male convex and uncus extending beyond it, juxta not strongly developed; 4th

abdominal sternite without patches of long setae; as in fig. 17. Female genitalia

with two pairs of very long apophyses; signa more than 3 cells wide; as in fig.

N an Sii bipinnatellus (p. 75)

5. Forewing pale with one broad dark brown terminal fascia. Male genitalia

probably as in fig. 12, but see description ............... scirpi (p. 66)

— Forewing pale without dark fasciae but may be Horace n 6

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

6. Male genitalia with well developed T-shaped (inverted) juxta; gnathos without

central posterior process. Female genitalia without signa tenuijuxtus (p. 82)

— Male genitalia without well developed juxta; gnathos with central posterior

process. Female genitalia with signa®. =.) errang fl

7. Male genitalia with pointed valves; bifurcate pseuduncus; aedeagus without

very large anellar spines; as in fig. 20. Female genitalia with simple row of cells

in signa and unmodified ductus bursae as in fig. 21 ....... distaleus (p. 78)

— Male genitalia with rounded valves; simple convex pseuduncus; aedeagus with

very large annellar spines; as in fig. 16. See specimen 12. Female genitalia with

complex signa and ductus bursae as in fig. 15 ......... punctulata (p.71)

Microcalyptris scirpi Braun

(iss, 12)

Microcatyptris scirpi Braun, 1925b: 225 (Type species).

Microcalyptris scirpi Braun; McDunnough, 1939: 107 (no. 9790).

Description. External features: ¢ (fig. 1). Head: palps and antennae brownish

buff; tuft on front of head and vertex light brown; eye-caps and collar brown.

Thorax and abdomen shiny brown, probably light on living specimens. Forewings:

ground colour of dorsal surface buff with gold reflections; terminally dark brown

fascia extends onto fringe; fringe mainly buff, some dark brown at ends of fascia;

ventral surface also buff but edged in dark brown all round and fringe greyish buff

and in part dark brown. Hindwings: both surfaces and fringe greyish. Legs greyish

buff with metallic reflections.

---___--

(a) cb) , (c)

(===)

Fig. 12. Microcalyptris scirpi. Reconstructed male genitalia. a, genital capsule; b, aedeagus: c, valve.

WILKINSON: Microcalyptris and Fomoria 67

Female not known.

Wing expanse '): Holotype 5mm. _

Genitalia: & (fig. 12). Unfortunately this species is known only from a single

specimen and the abdomen has been damaged. Thus the genitalia are badly

broken and infused with some substance which does not dissolve in the usual

solvents. The figure and description are therefore reconstructions. Pseuduncus:

prolonged backwards, single and uncus in form of inverted Y. Gnathos: probably

Y-shaped. A pair of slender lateral support rods present, extending from base of

uncus to base of valves. Vinculum: lateral arms narrow, almost equal to length of

valves. Saccus broad, deeply bifurcate. Valves not reaching beyond pseuduncus,

tapered terminally. Transtillae: lateral arms narrow. Juxta appears to be a simple

weakly sclerotised lobe but may have two posteriorly directed spines. Aedeagus

probably as illustrated although specimen is in three pieces. There is, on the slide,

a broken “horn” which is possibly one of a pair belonging to the aedeagus

(dotted). However, this is not certain and they could be part of the juxta.

Female not known.

Host plant: Scirpus paludosus.

Mine: A lower surface ophionome.

Diagnosis. The buff colour of the forewings together with the single brown

terminal fascia should be sufficient to diagnose this species from others known at

present — see check list page 65. The posterior Y-shaped sclerotisation of the

gnathos is also characteristic as may be other parts when the structure is known for

certain.

Discussion. Braun (1925b) described this genus and species from a single

specimen. The specimen is still the only one representing scirpi although it should

not be too difficult to acquire fresh material. We have no knowledge of the female

and since the one male specimen has damaged genitalia we also need to be better

informed regarding the structure of the male.

Distribution. USA: Utah..

Material examined. & Holotype: USA: “B1142; Bear R. Bay, Utah, 1.22.vi.24.

Type collection of Annette F. Braun; Microcalyptris scirpi Braun Type”; slide no:

USNM 16785; in ANS.

Biology. Mine. Uniformly narrow tract, 8 cm, blackish. Not visible from upper

surface.

Cocoon. Brownish ochreous, ovoid, very convex above, lacking projecting rim,

presumably found in other Nepticulidae.

Microcalyptris thoracealbella (Chambers)

(figs. 2, 13, 14)

Nepticula thoracealbella Chambers, 1873: 127.

') The wing measurements represent the alar expanse in millimeters. The moths were measured from

the middle of the mesothorax to the wing-tip and this figure was doubled to give the full alar expanse.

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Nepticula thoracealbella Chambers; Chambers, in Hayden, 1878: 158.

Nepticula thoracealbella Chambers; Dyar, 1903: 547.

Nepticula thoracealbella Chambers; Braun, 1917: 189.

Nepticula thoracealbella Chambers; Braun in Forbes, 1923: 93.

Nepticula thoracealbella Chambers; McDunnough, 1939: 107 (no. 9763).

Microcalyptris thoracealbella (Chambers); Davis, 1978: 214.

Nepticula badiocapitella Chambers, 1876: 160 (syn. by Braun, 1917b: 189).

Nepticula badiocapitella Chambers; Chambers in Hayden, 1878: 157.

Nepticula badiocapitella Chambers; Dyar, 1903: 545.

Description. External features: & (fig. 2). Head: palps creamy white; antennae

pale greyish brown; tuft on front of head dark ochreous, vertex darker; eye-caps

shining white; collar dark ochreous. Thorax whitish, weakly lustrous. Abdomen

pale yellowish brown with metallic grey reflections on both surfaces. Forewings:

ground colour of dorsal surface pale brown with bronze reflections, each scale

AI MH

r———t[j

Fig. 13. Microcalyptris thoracealbella. Male genitalia. a, genital capsule; b, aedeagus: c, hair brushes on

abdomen segment 4.

greyish at the base producing an irrorate effect which becomes more prominent

distally; fringe variously irrorate with wing scales apically, whitish and iridescent

silver; single medial fascia, off white, broadening on dorsal margin, followed by

two marginal patches of the same colour, postmedial in position. Hindwings:

ground colour and fringe pale greyish white, shining silver. Legs pale yellow-

brown with scattered metallic grey reflections. A pair of hair brushes dorsally on

segment IV, lateral in position as in fig. 13c.

WILKINSON: Microcalyptris and Fomoria 69

Female. As ¢ except for a pair of convex external pockets ventrally on the third

abdominal segment, medial in position. Hair brushes absent.

Wing expanse: ¢: 4.6—5.2 mm (4 specimens); 9: 4.6—4.8 mm (2 specimens).

Holotype: 4.8 mm.

Genitalia: & (fig. 13). Pseuduncus extended, weakly bilobed posteriorly; uncus a

bridge-like sclerotisation with a single medial process. Gnathos: as in fig. 13a;

transverse ventral plate with a broad medial process; dorso-lateral arms broad and

blunted terminally. Vinculum: lateral arms narrow with associated weakly scle-

rotised bars; ventral plate very narrow. Saccus broader than the ventral plate,

markedly bilobed with each lobe twice as long as broad at the base. Valves not

reaching beyond the pseuduncus, straight and blunted terminally. Transtillae:

lateral arms short and narrow; ventral arms long and straight; transverse bars

lint 77

(b)

002mm

Fig. 14. Microcalyptris thoracealbella. Female genitalia. a, genitalia; b, detail of signum enlarged; c, de-

tail of ductus bursa enlarged.

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

fused to form a continuous narrow strap. Juxta a flask-shaped spinose plate as in

fig. 13a. Aedeagus: regular, equal to length of the capsule, with a medial orifice at

the point of entry of the ductus ejaculatorius; vesica well defined, with cornuti as

small denticles distally and with a cup-shaped plate of minute papillae; anellus

comprising a pair of very large canine tooth-like spines.

Female (fig. 14). Colliculum funicular and weakly sclerotised. Ductus ne

short, expanding medially and bearing three fin-like plates as in fig. 14a. Accessory

duct arising from area of dilation, spiral proximally. Bursa copulatrix: covered

proximally with scallop-shaped chains of pectinations as in fig. 14c; signum

double, comprising two equal length bands of spinose cells with a spinose margin

as in figs. 14a and b. Anterior apophyses short and broad. Posterior apophyses

narrow, approximately four times length of anteriores. Anal plate large and

constricted medially as in fig. 14a.

Host plant: Not known.

Mine: Not known.

Diagnosis. Resembles bipinnatellus sp.n. in the pattern of wing markings, but the

darker ground colour, smaller distal patches and the absence of the apical patch in

thoracealbella separates the two. The male genitalia also resemble those of

bipinnatellus sp.n. but are separated by the papillate pseuduncus, the single lobe of

the juxta, and the shorter aedeagus relative to the capsule, in thoracealbella. The

female genitalia most closely resemble those of punctulata: the scallop-shaped

pectinations on the ductus and the form of the cells of the signa in thoracealbella

distinguish the two.

Discussion. Originally described by Chambers (1873) froma ee specimen

captured in Kentucky. Chambers later (1876) described a similar species also from

a single Kentucky specimen, under the name badiocapitella. Braun (1917) regarded

these two as synonymous, stating that the descriptions were virtually identical and

that the range of variation found in thoracealbella is sufficient to include ba-

diocapitella. The material examined in this study, which is believed to represent

badiocapitella, has been checked with the type of thoracealbella, and found to be

similar. However there is no trace of the type of badiocapitella and this prevents a

complete comparison. Chambers’ description of badiocapitella could also be

applied to bipinnatellus sp.n. especially in the form of the forewing markings, but in

the interests of nomenclatorial stability badiocapitella is left in synonymy with

thoracealbella.

There is a single 9 from Arkansas, labelled specimen 10, which differs only in

that the tuft on the vertex and the forewing ground colour are dark chocolate

brown.

Distribution. USA: Kentucky, Pennsylvania, Ohio, Virginia, New York.

Material examined. & Holotype: USA: “Kentucky Chambers; thoracealbella;

Type 14952; labelled as N. thoracealbella, MCZ”; CNC slide no. 3513; in MCZ.

Other specimens: In USNM: Pennsylvania, Arendtsville; 19, 2.vii.1921, 24, 3

ex., 6.vii.1921 (Frost). Pennsylvania, Harrisburg; 14, 24.vi.1912. Virginia,

Mountain Lake; 13, 23.vii.1940 (Milne and Milne). In ANS: Ohio, Cincinnati;

1g, 17.vii.1903, 19, 1945, 19, 20.v.1945 (Braun). In L. A. Co. Museum: New York

WILKINSON: Microcalyptris and Fomoria 71

Sea Cliff; 1g 20.v.? Specimen 10: In USNM: Arkansas, Devil's Den St. Pk.,

Washington County; 1g, 19, 23.vi.1966 (Hodges); slide no. USNM 17298; wing

expanse 5.2 mm.

Biology. Immature stages unknown.

Voltinism. Bivoltine with adults on the wing in May and in late June and July.

Specimen 11

There is a single male with externals like thoracealbella except that the tufts on

the head are darker, the thorax and forewings are pale brown and the distal

markings of the forewing are triangular streaks rather than patches. The genitalia

are badly damaged but can be seen to differ from thoracealbella in the following:

the juxta is valve-like in form with a weakly curved apical hook; the spines of the

anellus are shorter than in thoracealbella and the cornuti comprise a transverse

papillate plate which is expanded laterally as in Obrussa spp.

This specimen is labelled as reared from Scirpus olneyi. Specimen 11: In USNM:

Maryland, Blackwater Refuge; from Scirpus olneyi; 13, 11.viii. 1943; slide no.

USNM 16273; wing expanse 4.0 mm.

Microcalyptris punctulata (Braun) comb. n.

(figs. 4, 15)

Nepticula punctulata Braun, 1910: 174.

Nepticula punctulata Braun; Braun, 1917: 192.

Nepticula punctulata Braun; McDunnough, 1939: 107 (no. 9769).

Description. External features: ¢ 9 (fig. 4). Head: palps whitish buff; antennae

pale brown; tufts on front of head very pale buff, vertex usually brownish; eye-

caps buff; collar buff. Thorax pale greyish buff, sometimes irrorate with brown.

Abdomen brown, shining metallic grey beneath. Forewings: ground colour of

dorsal surface greyish buff with scattered silver reflections, variously irrorate with

scales brownish at the tip; fringe grey, shining silver. Hindwings: ground colour

and fringe pale greyish buff. Legs grey-buff with scattered paler areas, shining,

metallic grey behind.

Wing expanse: g: 5.0 mm (1 specimen); 9: 4.8—6.0 mm (2 specimens).

Lectotype: 4.8 mm.

Genitalia: g': not known (the abdomen is missing from the 1g specimen but see

specimen 12). © (fig. 15): Colliculum membranous. Ductus bursae long, covered

with small denticles proximally and more elongate spicules distally, and with a

medial expansion bearing three fin-like sclerotisations as in fig. 15a. Accessory

duct arising from area of dilation, spiral distally. Bursa copulatrix: covered

proximally with irregular chains of pectinations; signum double, comprising two

equal linear bands of spinose plates as in figs. 15a and b. Anterior apophyses broad

basally, with associated papillae as in figure. Posterior apophyses as long as ductus

and straight.

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Fig. 15. Microcalyptris punctulata. Female genitalia. a, genitalia; b, enlarged detail of signum.

Host plant: Ceanothus cuneatus and Rhamnus californica.

Mine: an upper surface ophionome.

Diagnosis. Resembles distaleus sp.n. in externals but differs in the generally

darker ground colour in punctulata; the absence of fasciae or patches on the

forewing separates punctulata from thoracealbella and bipinnatellus sp.n. The female

genitalia most closely resemble those of thoracealbella; the spicules of the ductus

and the spinose plates of the signa in punctulata separate the two.

Discussion. A better evaluation of the affinities may be made when male

genitalia are known. The female genitalia relate closely to thoracealbella, compare

figs. 14 and 15. This species was originally described from two specimens bred

WILKINSON: Microcalyptris and Fomoria 73

from Ceanothus cuneatus in California. They were labelled as cotypes by Braun.

Both have been examined and are in extremely poor condition. They are

unsuitable for reference to the externals, however, the specimen designated as

lectotype is the one from which a genitalia slide has been prepared.

Distribution. USA: California.

Material examined. I designate as lectotype, a 9 syntype; USA: “B.422; Dutch

Flat, Placer Co., Cal. i. 22.11.1909; Type Collection of Annette F. Braun; Nepticula

punctulata Braun Cotype”’; on Ceanothus cuneatus; slide No. USNM 16212; in ANS.

Paralectotype: In ANS: same data as lectotype, | ex.

Other specimens: In ANS: California, Loma Linda; 19, 3.vi.1912, 19, 8.iv., 1

ex., 18.vi.1912, 1g, 25.vi.1912, (Pilate).

Mines examined: In ANS: California, Dutch Flat, Placer Co.; 2 mines on

Ceanothus cuneatus; 6.1.1909, B.422 (Braun).

Biology. Egg. Laid on the lower surface of the leaf.

Host. Braun (1917) reports that the species also mines leaves of Rhamnus

californica from Yosemite, although this material has not been traced.

Mine. A short linear mine not visible from the upper surface in the earlier stages

later becomes quite distinct and usually follows the margin of the leaf. The frass is

deposited as a central black line.

Larva. Emerges on the upper surface of the leaf.

Cocoon. Reddish brown in colour.

Voltinism. Trivoltine with adults on the wing in February, April and June. Braun

(loc.cit.) records that the mines on Rhamnus were collected in July.

Specimen 12 (fig. 16)

There is a single male which may be conspecific with punctulata:

Description. External features: ¢. As punctulata except that: tufts on front of

head and vertex creamy white; forewings dorsally lustrous white, each scale

darker at the tip producing a slight irrorate effect; forewings ventrally covered

with creamy white scales almost to the tip; hindwing dorsal and ventral surfaces

covered with creamy white scales extending almost to the tip.

Genitalia: ¢ (fig. 16). Pseuduncus long, with a single blunted lobe and uncus

with bridge-like sclerotisation and a broad, spatulate medial process. Gnathos:

transverse ventral plate with a long thin medial process; dorso-lateral arms broad

and blunted with short posteriorly-directed processes. Vinculum: lateral arms with

associated weakly sclerotised bars tapering distally; ventral plate reduced. Saccus

broad, markedly bilobed, each lobe as long as broad at the base. Valves not

reaching the pseuduncus, broad and constricted medially, rounded terminally.

Transtillae: lateral arms short and broad; ventral arms long and tapering; trans-

verse bars indeterminate. Juxta with a broad boat-shaped base as in fig. 16a, distal

portion indeterminate. Aedeagus: equal to length of capsule, broadening distally

and with a medial orifice; vesica well defined with cornuti in form of several long

denticles distally and a cup-shaped plate of minute papillae; anellus comprising a

pair of very long tooth-like spines.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

(=e

Fig. 16. Microcalyptris specimen 12. Male genitalia. a, genital capsule; b, aedeagus.

(b)

ee °

Fig. 17. Microcalyptris bipinnatellus. Male genitalia. a, genital capsule; b, aedeagus.

WILKINSON: Microcalyptris and Fomoria 75

Diagnosis. Resembles thoracealbella in the overall form of the male genitalia but

may be separated by the longer single lobe of the pseuduncus, the sharp medial

process of the gnathos and the more weakly bilobed saccus, in specimen 12. The

externals are generally more white than the females of punctulata and the absence

of scattered brown scales on the forewings separates it from the males of distaleus

sp.n.

Discussion. The similarity of this specimen to punctulata suggests that it may

represent the male of that species. The genitalia are close to those of

thoracealbella, a fact which correlates well with the relationship between the

female genitalia of punctulata and thoracealbella.

Material examined. Specimen 12: USA: Arizona, Flagstaff; 13, 17.vii.1939

(Braun); wing expanse: 5.2 mm; slide no. 149—PJN; in ANS.

Microcalyptris bipinnatellus sp.n.

(figs. 6, 17, 18)

Description. External features: 4. Head: palps greyish; antennae dark grey-

brown; tuft on front of head ochreous, vertex paler; eye-caps off-white, weakly

lustrous; collar sandy buff. Thorax sandy buff, irrorate with very pale brown.

Abdomen brown-grey with metallic reflections above, shining silver beneath.

Forewings: ground colour of dorsal surface very pale brown, each scale darker at

the tip, scattered blue reflections; fringe greyish white, becoming creamy white at

the apex; two creamy white fasciae, antemedial variable in width but usually

broadening on the dorsal margin, postmedial oblique and broken by a line of

brown scales centrally, followed by a creamy white apical patch, variable in

extent. Hindwings: ground colour and fringe greyish, shining silver. Legs dark

brown with paler areas, shining metallic grey behind.

Female (fig. 6). As ¢ except for a pair of convex, external pockets ventrally on

the third abdominal segment.

Wing expanse: &: 4.6—5.6 mm (2 specimens); 9: 5.0—6.4 mm (6 specimens).

Holotype: 6.8 mm.

Genitalia: & (fig. 17). Pseuduncus single lobed; uncus with bridge-like

sclerotisation and a large medial process. Gnathos: as in fig. 17a; transverse

ventral plate with large medial process; dorso-lateral arms narrow and tapering.

Vinculum: lateral arms with associated weakly sclerotised bars, ventral plate

reduced, with a convex medial expansion. Saccus broad, markedly bilobed with

each lobe longer than broad at base. Valves not reaching the pseuduncus, narrow

and tapering distally. Transtillae: lateral arms short and broad; ventral arms long

and straight; transverse bars indeterminate. Juxta trifurcate with central lobe

spinose. Aedeagus: longer than the capsule, broadening distally, basally quadrate

with large medial orifice at the point of entry of the ductus ejaculatorius; vesica

well defined with cornuti as small denticles distally and with a cup-shaped plate of

minute papillae; anellus comprising a pair of very large hook-like spines.

Female (fig. 18). Colliculum with weakly sclerotised funicular antrum and a pair

of fin-like plates. Ductus bursae long and weakly sclerotised, with a pair of fin-like

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

plates distally. Accessory duct arising distally. Bursa copulatrix: with weakly

sclerotised chains of pectinations proximally; signum double, comprising two

ovate cellular patches of equal size, as in figs. 18a and b. Anterior apophyses long

and narrow. Posterior apophyses broad and sometimes club-shaped distally,

reaching well beyond the anteriores. Anal plate ovate as in fig. 18a.

Host plant: Not known.

Mine: Not known.

Diagnosis. Similar to thoracealbella in externals but the paler ground colour, the

larger distal patches and the presence of the apical patch on the forewings of

bipinnatellus separate the two. The male genitalia also resemble those of

thoracealbella but they may be differentiated by the single lobe of the pseuduncus,

the tapering valves and the longer aedeagus relative to the capsule, in bipinnatellus.

The female genitalia are differentiated from other members of the genus, listed on

RATE

(a)

Fig. 18. Microcalyptris bipinnatellus. Female genitalia. a, genitalia; b, enlarged detail of signum.

WILKINSON: Microcalyptris and Fomoria 0]

page 65, by the complex ornamentation of the ductus and the ovate signa in

bipinnatellus.

Discussion. Perhaps most closely related to thoracealbella judging from externals

and male genitalia. This new species is represented by a well mounted and

prepared type series collected by Hodges comparatively recently.

Distribution. USA: Florida.

Material examined. & Holotype: “Florida Lake Placid Archbold Bio. Sta. 1—8

June 1964 R. W. Hodges”; slide no: USNM 17282; in ANS. Paratypes: In USNM:

Florida, Lake Placid, Archbold Bio. Sta.; 19, 1—7.v.1964, 39, 8—15.v.1964

(Hodges). Florida, Fisheating Cr. Palmdale; 19, 7—10.v.1964 (Hodges). Florida,

Parker Is., Highlands Co.; 14 26—29.v.1964 (Hodges). Florida, Roy. Palm State

Park; 1g, 39, 1.1930 (Jones, F. M.); slide nos: USNM 17240, 17241, 17253, 17281,

17302, 17303, 17427, 17428, 17247.

Biology. Immature stages unknown. Voltinism. Univoltine, with adults on the

wing throughout May and in early June.

Etymology. Pinna (Latin): fin.

Microcalyptris postalatratus sp.n.

(figs. 3, 19)

Description. External features: ¢ (fig. 3). Head: palps greyish; antennae pale

brown; tufts on front of head and vertex dark brown; eye-caps creamy white,

lustrous; collar dark brown. Thorax and abdomen buff with scattered brown

scales, abdomen shining metallic grey beneath. Forewings: ground colour of

(a)

Fig. 19. Microcalyptris postalatratus. Male genitalia. a, genital capsule; b, aedeagus.

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 1979

dorsal surface buff, with each scale dark brown at the tip; fringe greyish with

several purplish black scales at the base of the dorsal margin, otherwise shining

metallic grey; ventral surface with an oval patch of purplish black scales extending

to two thirds. Hindwings: dorsal and ventral surfaces covered with purplish black

scales extending almost to the apex; fringe greyish, shining metallic grey. Legs pale

greyish brown with metallic grey reflections behind.

Female. Not known.

Wing expanse: 4: 3.4 mm (paratype); holotype: 5.4 mm.

Genitalia: & (fig. 19). Pseuduncus with a single rounded lobe and uncus bridge-

like with a spatulate central process. Gnathos: transverse ventral plate with a large

medial process, tapering distally; dorso-lateral arms blunted with short posteri- :

orly-directed processes. Vinculum: lateral arms with associated weakly sclerotised

bars broad at the base; ventral plate reduced. Saccus narrow, bilobed with each

lobe as long as broad at base. Valves not reaching the pseuduncus, narrow and

tapering distally. Transtillae: broadly W-shaped as in fig. 19a, lateral arms long and

narrow, ventral arms short; transverse bars fused to form a continuous arcuate ©

strap. Juxta membranous. Aedeagus: broad and regular, equal to length of

capsule; vesica well defined with cornutus in form of many large denticles distally

and a cup-shaped plate of minute papillae; anellus comprising a pair and twor

single tooth-like spines. Female not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. The striking contrast between the forewing ground colour and the

purplish black scales of the hindwing separate this from all the other members of

the genus discussed here. The genitalia resemble those of other members of the

genus but may be differentiated by the short, rounded lobe of the pseuduncus, the

short lobes of the saccus and the rather broad, squat appearance of the aedeagus.

Discussion. Externally this species appears not closely related to any other in

the genus as it is presently understood but the male genitalia correspond well.

Further collection and the rearing of females especially, may make its affinities

within the genus more clear. Described from two specimens collected by Braun in

Arizona, but she failed to name them and identify them as belonging to her genus

Microcalyptris.

Distribution. USA: Arizona.

Material examined. 3° Holotype: USA: ‘Chiricahua Mts., nr. Portal, Arizona.

4.vii.1939, A. F. Braun”; slide no: 150— PJN; in ANS. Paratype: In ANS: Arizona,

Superior; 1g, 11.vii.1939 (Braun); slide no: 151—PJN.

Biology. Immature stages unknown.

Etymology. Atratus (Latin): dressed in black.

Microcalyptris distaleus sp.n.

(figs.5, 209211)

Description. External features: 4 9 (fig. 5). Head: palps whitish; antennae pale

brown, tuft on front of head whitish ochre, vertex whitish; eye-caps white; collar

WILKINSON: Microcalyptris and Fomoria 79

Fig. 20. Microcalyptris distaleus. Male genitalia. a, genital capsule; b, aedeagus; c, detail on vesica en-

larged.

creamy white. Thorax and abdomen creamy white, abdomen with metallic

reflections beneath. Forewings: ground colour of dorsal surface white, sparsely

irrorate with scales brown at the tip, shining silver; fringe whitish, shining silver.

Hindwings: ground colour and fringe pale greyish white with scattered darker

areas.

Wing expanse: ¢: 5.2 mm (1 specimen); 9: 4.2 mm (1 specimen). Holotype: 6.2

mm.

Genitalia: ¢ (fig. 20). Pseuduncus with broad single lobe, uncus bridge-like

bifurcate and papillate extending beyond pseuduncus. Gnathos: as in fig. 20a,

transverse ventral plate with large medial process; dorso-lateral arms long and

narrow. Vinculum: apparently lacking associated sclerotised bars; ventral plate

broad with medial excavation. Saccus as broad as ventral plate, very weakly

bilobed. Valves: reaching just beyond the pseuduncus, tapering markedly and

pointed terminally. Transtillae: with short, stout lateral arms; transverse bars

indeterminate. Aedeagus: markedly shorter than the capsule, regular in width;

vesica with cornuti as many small denticles orientated in a ridge laterally and with

a large medial spine as in fig. 20b and also with a cup-shaped plate of minute

papillae.

Female (fig. 21). Colliculum membranous. Ductus bursae long, expanding

distally. Accessory duct arising distally with a single patch of spines at the vesti- .

bule, spiral distally. Bursa copulatrix: large and covered proximally with short

chains of weakly sclerotised pectinations; signum double, comprising two equal

bands of spinose cells with whorls of pectinations as in fig. 21a and b. Anterior

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

apophyses short and broad. Posterior apophyses long and straight, markedly

longer than the anteriores.

Host plant: Not known.

Mine: Not known.

Diagnosis. Rather paler than the other members of the genus, listed on page 65,

except for Specimen 12; the scattered brown scales of the forewing in distaleus

separate the two. The male genitalia are easily differentiated by the markedly

tapering valves, the papillate sclerotisation of the pseuduncus and the orientation

of the cornuti. The female genitalia resemble those of thoracealbella and punctulata

in the form of the signa, but the absence of any sclerotisation on the ductus and the

spiculate accessory duct, in distaleus, are diagnostic.

(a)

Fig. 21. Microcalyptris distaleus. Female genitalia. a, genitalia; b, detail of signum enlarged.

Discussion. This species is rather atypical of the genus, because the male lacks

the characteristic lateral bars associated with the vinculum and gnathos, the

WILKINSON: Microcalyptris and Fomoria 81

complex form of the anellus and differs in the overall nature of the aedeagus. The

female is also atypical in the absence of an antrum and the associated sclerotis-

ation of the ductus which is found in the other known members of the genus.

Especially the wing venation and general aspects of both male and female genitalia

do, however, indicate that it is congeneric with thoracealbella.

Distribution. USA: Arizona; California.

Material examined. & Holotype: USA: “Flagstaff, Arizona, 18.vii.1939, A. F.

Braun”; slide no: 153—PJN; in ANS. Paratypes: In ANS: Arizona, Flagstaff; 19,

18.vii.1939 (Braun). California, Loma Linda; 1g, 14.vi.1912 (Pilate); slide nos:

152—PJN, 154—PJN.

Biology. Immature stages unknown.

Etymology. Talea (Latin): slender staff or rod; dis (Latin): without.

The two new species following are recently published by my colleague Dr. D. R.

Davis in a paper entitled “New Leaf-mining Moths of the Family Nepticulidae

from Florida.” Thus there is no need to treat them in full, but a diagnosis and other

important details are given for the sake of completeness.

Microcalyptris bicornutus Davis

(fig. 7)

Microcalyptris bicornutus Davis, 1978: 212.

Diagnosis. 4 ©. (fig. 7). Fully described by Davis (loc. cit.). Head yellowish.

Palps and forewings uniformly greyish brown, paler basally. Scales often tipped

with fuscous. Hindwings uniform lighter grey. In females the sternites of the

abdomen has a pair of indistinct fenestrae situated near anterior margin, on either

side of the mid-ventral line.

Male genitalia (see Davis, loc.cit., figs. 18—20) with blunt, apically flattened

pseuduncus. Uncus a bridge-like sclerotisation associated with pseuduncus and

gnathos, similar to that of postalatratus. Gnathos also similar. Vinculum with a pair

of characteristic slender apophyses projecting anteriorly. Valves simple, rounded

terminally. Juxta is large, complex and therefore diagnostic; comprising a broad

plate, deeply bifurcate anteriorly and posteriorly, producing rounded apodemes

directed forwards and pointed apodemes backwards. Aedeagus arcuate, without

cornuti, but having typical large canine tooth-like spine on anellus.

Female genitalia (Davis, loc.cit., fig. 32) have the usual very long posterior

apophyses. Bursa copulatrix membranous with a pair of asymmetrical signa; each

Signum mostly comprising a single row of 20—31 scalariform ‘cell’ elements with

thickened margin.

Discussion. The species is described from a large series of 48 types. They were

collected as adults so the host plant, mine and biology of the immature stages are

unknown. M. bicornutus is apparently univoltine with a flight period between

September and late November.

The long support rods extending from the gnathos in the male and the very long

posterior apophyses in the female are of particular interest in Microcalyptris spe-

cies.

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Distribution. USA: Only known from Florida Keys.

Material examined. ¢ Holotype: USA: “Key Largo, Monroe Co., Florida;

19.xi.1964. Mrs. Spencer Kemp”. & genitalia USNM 16938 Holotype USNM

72105 in USNM. Paratypes. In USNM: 403, 79, data as holotype, 8.x.—

20.x1.1964, 29.1x.1972.

Microcalyptris tenuijuxtus Davis

(fig. 8)

Microcalyptris tenuijuxtus Davis, 1978: 216.

Diagnosis. ¢ 9 (fig. 8). Described by Davis (loc.cit.); the following characters

separate this species from the others described here. Almost entire moth pale

yellowish white, forewing with some brown irrorations in basal two thirds and

increasing distally including fringe almost forming two brown fasciae in some

specimens (e.g., Holotype). Hindwing white to pale yellowish. Abdominal fene-

strae absent.

Male genitalia (see Davis, loc.cit., figs. 21—23). Pseuduncus characteristic with

a median lobe, a pair of spine-like lobes and a pair of rounded lobes arising from it;

median lobe (probably uncus) with four small apical setae and the pair of rounded

lobes, each with one seta. Gnathos horn-like and bearing the lateral support rods.

Vinculum with anterior apophyses moderately well developed. Valves slender and

rounded apically. Juxta in shape of inverted T with arms rounded and stem bifur-

cate at tip. Aedeagus characteristic with about three apical spines and rounded

lobes apically.

Female genitalia (Davis, loc.cit., fig. 33). Posterior apophyses again very long.

Bursa copulatrix without signa. Accessory duct present with spiral distally as in

bicornutus.

Discussion. This species is described from 22 specimens, but only one is female.

It is somewhat unusual in that the female has no signa on the bursa, and the male

has the curious development on the pseuduncus. The host plant, mine and

immature stages are unknown. The species is probably univoltine although the

range of collecting dates (early October to late November) may simply reflect the

periodicity of the collector.

Distribution. USA: Florida Keys.

Material examined. & Holotype: USA: “Key Largo, Monroe Co., Florida,

17.x1.1964. Mrs. Spencer Kemp.” ¢ genitalia USNM 1680, Holotype USNM 72106

in USNM. Paratypes. In USNM: 20 4, 1 © same data as holotype, 8.x.—

28.x1.1964.

Fomoria Beirne, 1945

Fomoria Beirne, 1945: 208. Type species by original designation: Fomoria weaveri (Stainton).

Taxonomic history

The genus was originally described by Beirne (1945) as a European genus with

WILKINSON: Microcalyptris and Fomoria 83

two species weaveri and septembrella (Stainton) transferred to it from ‘‘Nepticula”’

von Heyden, 1843 — a junior synonym of Stigmella Schrank, 1802 (see Wilkinson,

1978: 13).

The original generic description was based on the structure of the male

genitalia. Comment was made on the colour of the head and wings and also on

larval habits. However, venation and female structures were not discussed

although venation, at least, is characteristic.

The two species which are here transferred to Fomoria are the first North

American species to be recognized as belonging to this genus.

Generic description

External features: 4 Q. Head: palps extending well beyond labrum, pale in

colour; antennae approximately half the length of the forewing, pale or brown

annulate with paler areas; tuft on front of head brown or ochreous, vertex

concolorous; eye-caps white, sometimes with fuscous scales distally; collar as

vertex. Thorax dark brown to, black and iridescent. Abdomen brown and

iridescent gold or silver. Venation: as in fig. 22. Forewings: Media coalescing with

Cubitus at base and both passing obliquely to Radius at R,,,; Cubitus becoming

obsolete; R, and R, separate. Hindwings: Media single-branched. Forewings:

elongate and ovate in shape, ground colour of dorsal surface brown, sometimes

SC +R]

A Cu

Fig. 22. Fomoria sp. Wing venation.

with scales darker at the tips; fringe brown or grey and marked apically with a

band of wing scales; markings either absent or in the form of silver fasciae or

patches. Hindwings: narrow and lanceolate, brown. Legs: brown or ochreous,

sometimes annulate with paler areas; proximal pair of spurs on hind-tibiae below

the middle.

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

Male genitalia: vinculum always ring-shaped; tegumen fused with vinculum

dorsally, extending into long, bluntly pointed pseuduncus. Uncus sclerotised with

spatulate medial projection. Gnathos with medial arms fused to form posterior

central boss or tapering process, rarely with anterior medial projection, lateral

arms usually broad and straight. Saccus weakly bilobed. Valves tapering markedly

and in Nearctic species with large dorsal spine. Transtillae broadly W-shaped;

transverse bars continuous. Juxta present in Palaearctic species as complex,

heavily sclerotised plate; absent in Nearctic species. Aedeagus: regular or weakly

flask-shaped, usually equal to length of capsule; vesica usually adorned with small

denticles and with a complex plate of minute papillae; several pairs of heavily

sclerotised, sometimes elaborate, cornuti and anellar spines.

Female genitalia: apophyses approximately equal to length of ductus.

Colliculum with sclerotised funicular antrum or complex plates. Ductus denticu-

late and with spiral accessory duct and sac. Bursa copulatrix: large and variously

pectinate; signum double, comprising weakly sclerotised, linear, reticulate

patches. |

Mining habit: leaf mines; larvae often recorded as pupating within the mines.

Generic differential diagnosis. See page 63.

Fomoria pteliaeella (Chambers) comb.n.

(figs. 9, 23, 24)

Nepticula pteliaeella Chambers, 1881: 137; 1882: 276.

Nepticula pteliaeella Chambers; Dyar, 1903: 546.

Nepticula pteliaeella Chambers; Braun, 1917: 168.

Nepticula pteliaeella Chambers; Braun, in Forbes, 1923: 86.

Nepticula pteliaeella Chambers; McDunnough, 1939: 107 (no. 9721).

Description. External features: ¢. Head: palps greyish; antennae dark grey;

tufts on front of head and vertex brown; eye-caps shining white; collar very dark

brown. Thorax very dark brown to black, strongly iridescent silver. Abdomen dark

brown with scattered gold reflections above, shining metallic grey beneath.

Forewings: ground colour of dorsal surface dark brown with bronze reflections,

fringe greyish brown, shining silver at apex, with an apical band of dark brown

wing-scales; basal patch on dorsal margin, silver, followed by a single antemedial

fascia, shining silver and widening .on dorsal margin, two marginal streaks,

postmedial, almost touching in the middle, both shine silver but have dusted

appearance when viewed from certain angles. Hindwings: ground colour and

fringe greyish brown, shining metallic grey. Legs dark brown with metallic grey

reflections, yellowish behind (fig. 9).

Female. As 8 except for a pair of convex external pockets ventrally on the

third abdominal segment, medial in position.

Wing expanse: &: 4.0—5.0 mm (8 specimens); 9: 3.8—5.4 mm (11 specimens).

Genitalia: &. (fig. 23). Pseuduncus with a single tapering lobe and associated

sclerotisation with a medial spatulate process as in fig. 23a. Gnathos: an inverted V

as in fig. 23a; transverse ventral plate with a large, pointed medial process; dorso-

lateral arms broad and straight. Vinculum: triangular as in fig. 23a; lateral arms

WILKINSON: Microcalyptris and Fomoria 85

“il

Fig. 23. Fomoria pteliaeella. Male genitalia. a, genital capsule; b, aedeagus; c, valve.

very broad; ventral plate broad. Saccus narrower than the ventral plate, weakly

bilobed. Valves not reaching pseuduncus, tapering markedly and with dorsal spine

arising medially not reaching beyond the cuiller as in fig. 23c. Transtillae: W-

shaped as in fig. 23a; lateral arms short and narrow; ventral arms long, reaching

beyond the ventral plate; transverse bars fused. Aedeagus: broad and regular,

shorter than length of capsule; vesica with between five and ten very large cornuti

and with a plate of minute papillae as in fig. 23b; anellus comprising a pair of broad

lateral spines.

Female (fig. 24). Colliculum with a weakly sclerotised funicular pocket as in fig.

24a. Ductus bursae short and narrow with accessory duct arising medially,

becoming spiral distally. Bursa copulatrix: large and covered with short, heavily

sclerotised chains of pectinations proximally and heavily sclerotised denticles

distally, both of which are on striations of the bursa; signum double comprising a

pair of long cellular patches, unequal in length as in figs. 24a and b. Anterior

apophyses very long and slightly arcuate distally. Posterior apophyses straight and

narrow, not reaching the anteriores.

Host plant: Prelea trifoliata (Hop-tree).

Mine: An upper surface ophionome.

Diagnosis. Differs externally from hypericella in the strongly iridescent nature of

the thorax of pteliaeella and in the absence of wing markings in hypericella. Both

male and female genitalia resemble those of hypericella but may be separated by

the pointed median process of the gnathos, the relatively shorter aedeagus and the

larger and more numerous cornuti at the phallotreme in pteliaeella; the females

differ in that the funicular antrum lacks the heavy sclerotisation and the lance-

shaped plate found in hypericella.

Discussion. Obviously related to hypericella in the overall form of both male and

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

female genitalia and particularly in the large dorsal spine of the valves. First

described by Chambers (1881) from larva and mine and then later (1882) from the

adult he had reared.

Distribution. USA: Ohio.

(a)

Fig. 24. Fomoria pteliaeella. Female genitalia. a, genitalia; b, detail of signum enlarged.

Material examined. In ANS: Ohio, Cincinnati; on Ptelea trifoliata; | Gg,

Ovi 1917, 2-8 3) 9,7 ex., vi. 1917 1.0%, 1911917 B: 736) (Braun) MO hice

Clermont County; on Ptelea trifoliata; 1 3, 5.vi.1916, 1 &, 2.v.1921, 1 g, 2.v.1921,

boy 2lwl92l2 Alex, 232,192, 2,22 D BE evel OZ IEN

26.v.1921, B.736 (Braun). In USNM: Ohio, Clermont County; on Ptelea trifoliata; 2

®, 25, 26.v.1921, B.736 (Braun). In Zoölogisch Museum, Amsterdam. Ohio,

WILKINSON: Microcalyptris and Fomoria 87

Cincinnati, 3 &, 11.vi.1917, B.736 (Braun). Mines examined: in ANS: Ohio, 1 mine

on Ptelea trifoliata; date not certain, B.736 (Braun).

Biology. Egg. Laid on the lower surface of the leaf and, in the single case

examined, next to a vein.

Mine. An upper surface, much contorted linear mine with rather undulating

margins in the later portion. The frass is evenly scattered across the breadth of the

mine in the earlier portion, but is a continuous central line distally.

Larva. Emerges on the upper surface of the leaf prior to pupation.

Pupa. Cocoon dark brown (Braun) or yellowish green (Chambers).

Voltinism. Braun (1917) reports that larvae may be collected in July, August and

September, and that the species is bivoltine. The material examined here

represents a single generation, the adults being on the wing in late May and June.

This agrees with Chambers’ findings.

Fomoria hypericella (Braun) comb.n.

(figs. 10, 25, 26)

Nepticula hypericella Braun, 1925a: 17.

Nepticula hypericella Braun; McDunnough, 1939: 107 (No: 9768).

Description. External features: ¢ ©. Head: palps grey; antennae greyish brown,

faintly annulate with paler areas; tuft on front of head orange-ochreous, vertex

darker; eye-caps shining white, sometimes shading to fuscous distally; collar pale

(b)

_—__________aou

Fig. 25. Fomoria hypericella. Male genitalia. a, genital capsule; b, aedeagus; c, detail on vesica en-

larged.

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

ochreous. Thorax and abdomen dark brown, thorax with greyish lustre, abdomen

shining metallic grey beneath. Forewings: ground colour of dorsal surface greyish

brown, each scale darker at the tip, various reflections but predominantly grey and |

bronze; fringe greyish shining metallic grey, with an apical band of dark brown

wing-scales. Hindwings: ground colour and fringe brownish, shining metallic grey.

Legs greyish brown, paler and shining metallic grey behind, tarsi faintly annulate

with buff (fig. 10).

Wing expanse: &: 4.0—4.8 mm (8 specimens); 9: 3.8—5.0 mm (9 specimens).

Holotype: 5.0 mm.

Genitalia: &. (fig. 25). Pseuduncus with a single, tapering lobe and uncus

sclerotised with a medial spatulate process as in fig. 25a. Gnathos: an inverted V

with transverse ventral plate forming a broad and flat central boss; dorso-lateral

arms short. Vinculum: lateral arms broad; ventral plate narrow, with a medial

convex expansion. Saccus very narrow, bilobed. Valves just reaching the pseu-

duncus, tapering markedly into a broad point, with dorsal spine arising medially

just reaching cuiller as in fig. 25a. Transtillae: broadly W-shaped as in fig. 25a; °

lateral arms short and narrow; ventral arms blunted, not reaching the ventral

plate. Aedeagus: flask-shaped, approximately equal to length of capsule; vesica

with cornuti as many small spiculate plates and with a comma-shaped plate of

minute papillae; anellus comprising two pairs of very large tooth-like spines and

single spines.

Female (fig. 26). Colliculum with a weakly sclerotised funicular antrum and a

lance-shaped chitinous plate. Ductus bursae short and as broad as the bursa

copulatrix. Accessory duct arising from the area of sclerotisation, spiral distally.

Bursa copulatrix: long and narrow with short heavily sclerotised chains of

pectinations proximally and heavily sclerotised denticles distally, both of which

are on striations of the bursa; signum double, comprising an equal pair of long,

cellular patches as in figs. 26a and b. Anterior apophyses long and narrow.

Posterior apophyses straight and narrow, approximately equal to length of the

anteriores.

Host plant: Hypericum prolificum (St. John’s Wort).

Mine: An upper surface ophionome.

Diagnosis. The uniform colour of the forewings and the absence of any

markings separate this species from pteliaeella. The quadrate median process of

the gnathos, the larger aedeagus and the more complex anellar projections in

hypericella distinguish the male genitalia from those of pteliaeella, while the more

heavily sclerotised antrum of hypericella separates the females.

Discussion. Originally described from holotype and 18 paratypes all reared by

Braun.

Distribution. USA: Ohio.

Material examined. 9 Holotype: USA: “B. 1103; Eastwood O., i. 13.viii.1923,

Annette F. Braun; Type; Nepticula hypericella Braun Type”; on Hypericum

prolificum; slide no. 110—PJN; in ANS.

Paratypes: in ANS: data as Holotype: 1 &, 1 9, 6.viii.1923, 2 &,2 9, 8.viii.1923,

bg) 10.viii.1923, 579, 1.vu01923, 03 wen el 2 viii 923.01 GEV IOS MINOR

WILKINSON: Microcalyptris and Fomoria 89

20.viii.1923, B.1103 (Braun); slide nos: 111—PJN, 112—PJN, 113—PJN,

114—PJN, 115—PJN.

Mines examined: In ANS: Ohio, Eastwood; 2 mines on Hypericum prolificum;

20.vii.1923, B.1103 (Braun).

Biology. Egg. Laid on the lower surface of the leaf, adjacent to the midrib.

Mine. A very long, slender tract on the upper surface. The frass is deposited

centrally as a continuous line in the early portion but later more generally

scattered.

Larva. Emerges on the lower surface of the leaf. Braun reports that occasion-

Fig. 26. Fomoria hypericella. Female genitalia.

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 4, 1979

ally pupation occurs within the mine, with the anterior edge of the cocoon placed

at the slit in the leaf surface.

Pupa. Cocoon pale brown in colour.

Voltinism. Univoltine in Ohio, the adults being on the wing in August.

ACKNOWLEDGEMENTS

My thanks are due to Mr. Philip Newton who whilst my research assistant in

England helped considerably with dissections and drafting some species descrip-

tions. I am very grateful to Dr. Georgina Bryan (FUA) who assisted with final

checking and proof reading and to Dr. D. R. Davis (USNM) for his hospitality,

helpful discussions and for arranging the loan of specimens. I acknowledge Mr. G.

W.H. van den Berg and Mr. S. Paniry for their help with the illustrations.

REFERENCES

Beirne, B. P., 1945. The Male Genitalia of the British Stigmellidae (Nepticulidae) (Lep.). — Proc. R. Ir.

Acad., (B) 50 (9): 191—218, 81 figs.

Braun, A. F., 1910. New species of Tineina from California. — Ent. News. 21 (4): 171—179.

1915. New genera and species of Tineina. — Can. Ent. 47 (6): 188— 197, 6 figs.

——., 1917. Nepticulidae of North America. — Trans. Am. ent. Soc. 43 (762): 155—209, 4 pls.

—., 1923. Superfamily Nepticuloidea. Family 5. Nepticulidae. — In: Forbes, W. T. W. (ed.), Lepi-

doptera of New York and Neighbouring States. pp. 79—98, 10 figs.

——, 1925a. Some undescribed Microlepidoptera and notes on life histories. — Trans. Am. ent. Soc.

51: 13—17.

—., 1925b. Microlepidoptera of Northern Utah. — Trans. Am. ent. Soc. 51 (3): 183— 226.

Busck, A., 1907. New American Tineina. — Proc. ent. Soc. Wash. 16: 46—54, 143—150.

Chambers, V. T., 1873. Micro-Lepidoptera. — Can. Ent. 5 (7): 124—128.

——, 1876. Tineina. — Can. Ent. 8 (8): 158— 160.

———., 1878. Index to the described Tineina of the United States and Canada. — In: Hayden, F. V.

(ed.), Bull. U. S. geol. geogr. Surv. Territ. 4: 125— 167.

—, 1881. Further notes on some Tineid larvae. — Psyche 3: 135— 137, 147— 149.

, 1882. Nepticula pteliaeella n. spec. — Psyche 3: 276.

Davis, D. R., 1978. New Leaf-mining Moths of the family Nepticulidae from Florida. — Florida Ento-

mologist 61 (4): 209— 224, 36 figs., | map.

Dyar, H. G., 1903. List of North American Lepidoptera and key to the literature of this order of insects.

— Bull. U.S. natn. Mus. 52: xix + 723 pp. Washington.

Heyden, C. von, 1843. Amtliche Bericht der Versammlung der Naturforscher zu Mainz: 208 pp.

McDunnough, J., 1939. Checklist of the Lepidoptera of Canada and the United States of America, part

II. Microlepidoptera. — Mem. So. Calif. Acad. Sci. 2 (1): 1—171.

Schrank, F. von P., 1802. Fauna boica. 2 (2) 412 pp. — Nürnberg.

Wilkinson, C., 1978. On the Stigmella-Nepticula controversy (Lepidoptera). — Tijdschr. Entom. 121 (2):

13—22, pls. 1—2.

Wilkinson, C.& M. J. Scoble, 1979. The Nepticulidae (Lepidoptera) of Canada. — Mem. Canad. Ent.

107: 1—118, 61 figs., 10 pls.

È | LIBRARY

_ DEEL 122 AFLEVERING 5 a 1979

i JUN 4 9 1979

‘ HARVARD

3 UNIVERSITY

| TIJDSCHRIFT

| VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

_ E. N. SAVTSHENKO. — Phylogenie und Systematik der Tipulidae, p. 91126, fig.

. È I—5. Translated and revised by B. Theowald and G. Theischinger.

ijdschrift voor Entomologie, deel 122, afl. 5 Gepubliceerd 31-V-1979

PHYLOGENIE UND SYSTEMATIK DER TIPULIDAE

von

E. N. SAVTSHENKO

[Fauna Ukraini 14 (1), pp. 63—88 (1966)]

Deutsche Bearbeitung

von

B. THEOWALD

Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Plantage Middenlaan 64, Amsterdam

und

G. THEISCHINGER

Oberösterreichisches Landesmuseum, Museumstraße 14, Linz, Österreich

Mit fünf Abbildungen

VORWORT

In den letzten fünfzig Jahren ist von mehreren Spezialisten an der Taxonomie der Tipuloidea

gearbeitet worden, wodurch sich die Zahl der bekannten Arten vervielfachte. In diesem Zeitraum

erschienen auch die ersten Arbeiten über Anatomie, Vergleichende Morphologie, Physiologie,

Cytologie und Biologie der Tipuloidea, und Larven, Puppen und Fossilien einer Reihe von Arten

wurden bekannt. Unser Freund Eugen Nikolajewitsh Savtshenko hat als erster die Ergebnisse aller

dieser Diziplinen zur Nachprüfung der Stammesgeschichte der Tipuloidea benützt, und damit einen

wichtigen Beitrag zur Kenntnis der Phylogenie der Dipteren, von denen die Tipuloidea die älteste

Gruppe darstellen, vorgelegt.

Diese Arbeit erschien ursprünglich in ukrainischer Sprache. Das Institut für taxonomische Zoologie

der Universität von Amsterdam hat es Frau E. Hanicenko ermöglicht, sie wörtlich ins Holländische zu

übersetzen. Basierend auf dieser Übersetzung haben wir mit freundlicher Mithilfe des Autors eine

deutsche Bearbeitung fertiggestellt für diejenigen, die die ukrainische Sprache niet beherrschen.

Auf Vorschlag des Autors sind gegenüber der ursprünglichen Ausgabe einige kleine hauptsächlich

nomenklatorische Änderungen eingearbeitet worden, wodurch die Arbeit wieder up to date ist. Die

Figuren wurden von Herrn J. Zaagman (Institut für taxonomische Zoologie, Amsterdam) neu-

gezeichnet und für diese Ausgabe fortlaufend numeriert.

PHYLOGENIE UND SYSTEMATIK DER TIPULIDAE

Die Tipulidae gehören zusammen mit den anderen Tipuloidea (Trichoceridae,

Cylindrotomidae, Limoniidae, Tanyderidae und Ptychopteridae) zu der natür-

lichen Entstehungsgruppe der niederen Nematocera, die als monophyletisch gilt.

Es ist deshalb zweckmäßig diese Familien nicht gesondert, sondern in ihrem

Zusammenhang zu betrachten.

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

DIE PHYLOGENIE DER TIPULOIDEA

Die Tipuloidea gehören, wie alle anderen Dipteren, zu den Mecopteroidea oder

der Panorpoiden-Gruppe der Insekten. Ihren Ursprung haben sie in den meso-

zoischen Panorpiden (Tillyard, 1918; Handlirsch, 1939).

Die Tipuloidea und die heutigen oder schon ausgestorbenen Panorpiden haben

viele morphologische Merkmale gemeinsam, was auf nahe Verwandtschaft hin-

weist (Bodenheimer, 1924). Die Tipuloidea haben folgende ursprüngliche

Merkmale, wenn auch in etwas abgeänderter Form, von den Panorpiden

übernommen und behalten:

a. in beiden Geschlechtern dichoptische Augen und vielgliedrige Fühler mit etwa

gleichgeformten Fühlergliedern;

b. ein ziemlich kompliziertes Flügelgeäder, besonders in den Hauptstämmen der

Adern: Radius vierästig, eine m-cu-Querader, deutliche Reste eines zweiten

Cubitus in Form einer Falte entlang cu, und zwei Analadern, außerdem das

atavistische Vorkommen einiger anormaler Queradern — besonders in den

radialen und medialen Zellen der Flügelspitze, wo solche auch bei den Ur-

formen sehr lange gefunden wurden (Rohdendorf, 1960);

c. das sporadische, wahrscheinlich atavistische Vorkommen von Makrotrichien in

den Flügelzellen bei Arten von Artengruppen, deren andere Arten alle

unbehaarte Flügel haben (z.B. Nephrotoma quadristriata (Schummel) bei den

Tipulidae); eine Querzeichnung der Flügel, die charakteristisch ist für einige

Tipulidae (Tipula trifasciata Loew und andere), Limoniidae und

Ptychopteridae;

d. eine V-förmige Quernaht am Mesonotum, die mit Ausnahme der Tipulidae nur

bei wenigen anderen Gruppen von Nematoceren vorhanden ist (z.B. bei den

Mycetophilidae), und bei Spezialisation der Dipteren verschwindet;

e. eines der zwei Meronen, die für die Panorpiden charakteristisch sind, ist bei

vielen Limoniidae schon weitgehend reduziert und bei höher spezialisierten

Dipteren verschwunden (Edwards, 1938).

Die phylogenetische Verwandtschaft der Tipuloidea mit den Panorpiden wird

auch bestätigt durch das Vorhandensein von deutlich entwickelten Ozellen bei

archaischen Gruppen (Trichoceridae) und von wenigstens rudimentären Ozellen

(Slipka, 1950a, 1950b) bei höher spezialisierten Gruppen (Tipulidae); auch durch

die homologe Innervierung und Befestigung der Flügel am Thorax bei beiden

Gruppen (Zacwilichowski, 1933, 1934) sowie durch homologe Auswüchse an den

Tergiten des Abdomens der Larven (Theowald, 1957). Auch die Ernährungsweise

der Imagines der Tipulidae ist jener der Panorpiden ähnlich. Sie ernähren sich

vom Nektar der Blumen, von Honigtau und von normalem Tau (Stitz, 1926).

Auf Grund von vergleichend-morphologischen und palaeontologischen Ge-

gebenheiten (Martynowa, 1959; Rohdendorf, 1960, 1961, 1962 und 1964) sieht die

Entwicklung der mesozoischen Panorpiden bis zu den primitiven Formen der

Dipteren und von diesen bis zu den Tipulidae folgendermaßen aus.

Schon im Perm — vor wenigstens 225 Million Jahren — haben sich aus den

palaeozoischen Panorpiden, gehörend zu den Permochoristidae, die Para-

trichoptera entwickelt. Sie waren später in der Evolution des ganzen Panorpiden-

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 93

Abb. |: Geäder der Vorderflügel einiger Ahnen der Tipulidae (nach Rohdendorf): a Panorpide aus dem

Perm (Platychoristidae); b, c Panorpide aus dem Perm (Permochoristidae); d Panorpide aus dem Perm

(Permotipulidae, Gattung Permotipula); e, f Archaische Zweiflügler aus dem Trias (Archidiptera) der

Familie Dictyodipteridae; g Diplarchitipula Rohdendorf (Architipulidae); h Architipula Handlirsch (Ar-

chitipulidae).

Komplexes der Insekten von großer Bedeutung. Diese Gruppe, gekennzeichnet

durch eine große ökologische Plastizität, entwickelte sich schnell und breitete sich

über immer mehr Biotope aus (Martynova, 1959). In dieser Gruppe begann die

Reduktion der Hinterflügel, die schließlich zur Zweiflügeligkeit führte

(Dipterismus). Die Vorderflügel der Paratrichoptera unterscheiden sich von denen

der anderen palaeozoischen Panorpiden durch eine Anzahl von progressiven

Merkmalen wie Kostalisation der Hauptstämme des Flügelgeäders, wodurch sich

die Flügelmembran verschmälerte und die cubitalen Adern dicht aneinander zu

liegen kamen, Differenzierung der Basis des Radius und Reduktion der Anzahl

der Analadern bis auf eine oder zwei (Abb. 1:b, c). Im Gegensatz zu den

archaischen Panorpiden aus dem Perm hatten die Paratrichoptera viel weniger

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

oder gar keine Queradern in den radialen und medialen Zellen der Flügelspitze.

Gewöhnlich wurden aber wohl die m-cu Querader und die kleine Querader von

den Vorfahren vererbt.

Manche Paratrichoptera-Arten waren, was die Flügeladerung anbelangt, den

rezenten Tipulidae ziemlich ähnlich, ganz besonders die Panorpide Permotipula

patricia Tillyard aus Ablagerungen des Ober-Perm in Australien (Abb. 4: d). Erst

als festgestellt wurde, daß dieses Insekt zwei Flügelpaare hat (Tillyard, 1937),

wurde es nicht mehr den echten Tipulidae (Tillyard, 1929), sondern den

Panorpiden zugeordnet.

Wie auch die heutigen Panorpiden hatten die mesozoischen Arten der Para-

trichoptera wahrscheinlich Larven mit vollständig entwickeltem Kopfskelett

(eucephal), mit drei Paar gegliederten Thorakalbeinen und mit wenigstens einigen

Paaren von Stigmenöffnungen am Hinterleib (peripneustisch). Es gibt genügend

Evidenz anzunehmen, daß diese Insekten hygrophil waren und daß sie am Boden

in der Nähe von Gewässern unter ziemlich feuchten Umständen umherkrochen

(Martynova, 1959). Sie ernährten sich saprophag von verschiedenen schon in

Verwesung übergegangenen organischen Bestandteilen und von abgestorbenen

Moosblättchen, wenn möglich aber auch von frischem Moos, wie die Larven der

heute in Australien vorkommenden Panorpiden-Art Chorista australis Klug (Essig,

1942) und die Larven vieler rezenter Tipuloidea einschließlich Tipulidae.

Es ist nicht auszuschließen, daß schon bei den Paratrichoptera im Lar-

venstadium die Tendenz zu einer amphibischen Lebensweise bestand. Die Larven

krochen nicht mehr am Bodem umher, sondern in einer mehr oder weniger

verschlammten dünnen Substanz, was ihre Bewegungen erschwerte und

langsamer machte (Rohdendorf, 1964). Im Verlauf der weiteren Evolution der

ancestralen Formen der Tipuloidea könnte diese Tendenz — die sich in

bestimmten morphologischen Anpassungen manifestierte — die Ursache gewesen

sein, daß die Larven der Tipuloidea denen der Bibionidae ähnlich wurden; bei den

Bibionidae sind der Kopf noch eucephal und die Stigmenöffnungen noch

peripneustisch, gegliederte Thorakalbeine fehlen aber, weil sie beim Leben in

einer halbflüssigen Substanz funktionell nicht brauchbar sind.

Nach Martynova (1959) haben sich wahrscheinlich aus den Paratrichoptera —

nach Verlust der Hinterflügel in der Trias — die ersten primitiven Archidiptera

entwickelt, die dem Flügelgeäder nach den Tipuloidea ähnlicher waren als den

Paratrichoptera. Kennzeichnend für die Flügel der primitiven Diptera sind der

hohe Grad der Versteifung des Flügelvorderrandes, die Differenzierung der Dicke

der Hauptstämme der Längsadern, die weitere Reduktion und schließlich die

Änderung von cu, in eine cu, entlang laufende Falte (Abb. I: e, f).

Von den nur fossil bekannten Archidiptera ist, betreffend die Phylogenie der

Tipuloidea, die Familie der Dictyodipteridae aus der Trias von Issyk-Kul in

Kazakstan wohl am interessantesten. Rohdendorf (1960, 1964) meint, daß in der

Trias in dieser Familie die Grundlage gelegt wird für die echten Zweiflügler

(Eodiptera) mit an der Basis verschmälerten und länglichen Flügeln; in den ersten

Evolutionsstadien war, was die Flügeladerung anlangt, die Tendenz zur Reduktion

der Queradern am Ende des medialen Flügelfeldes kennzeichnend.

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 95

Die ersten mesozoischen Eodiptera hatten wie die Archidiptera wahrscheinlich

noch Larven des Bibioniden-Typs, die in sehr feuchtem, an organischen Stoffen

reichem Schlammboden und möglicherweise auch in seichtem Wasser lebten.

Rohdendorf (1962, 1964) sieht die überaus primitiven Eodiptera als Ahnformen

einerseits der Tipuloidea — diese entwickelten sich weiter, lieferten aber keine auf

höherem Evolutionsniveau stehenden Nachkommen — und anderseits der Bibio-

nidae, von denen alle höher entwickelten Familien der rezenten Diptera

abstammen.

Das Entstehen der Tipuloidea aus den primären Eodiptera fand in der Trias

statt. Ihre primitivste Familie sind die Architipulidae, von denen Rohdendorf

(1964) alle rezenten Tipuloidea ableitet. Die Architipulidae hatten wie alle heu-

tigen Tipuloidea die charakteristische Verschiebung von r,,, zu r, in Richtung

Flügelspitze und r,, der am Ende nicht mit dem Flügelrand sondern mit r,

verbunden war (Hennig, 1954). Überdies zeichneten sich die Architipulidae durch

einen ziemlich langen Radius aus und die primitiveren Formen durch überzählige

Queradern im medialen Spitzenfeld der Flügel und manchmal auch durch

Verzweigung der Adern m, und cu (Abb. |: g, h). Was die Aderung der Flügel

betrifft, unterschieden sie sich nur wenig von den rezenten Tipulidae; deshalb war

Handlirsch (1908), der diese Familie beschrieben hat, nicht ganz sicher, ob diese

Gruppe wohl den Status einer Familie verdiente oder nicht. Weil er nicht wußte,

welcher Gruppe der heutigen Tipulidae er sie am besten zuordnen sollte, hat er sie

als Familie aufgeführt (Handlirsch, 1925, 1939).

Schon in der Ober-Trias waren die Architipulidae von wenigstens drei Familien

mit einigen Arten vertreten (Rohdendorf, 1964) und im Unter-Jura — besonders

im Lias von Deutschland — mit zehn Familien und ziemlich vielen Arten (Hand-

lirsch, 1908, 1939), jedoch mit Ausnahme einiger Arten, die von Bode (1953)

beschrieben wurden, weil es zweifelhaft ist, ob diese Arten überhaupt zu dieser

Gruppe beziehungsweise zu den Tipulidae gehören.

Der bedeutenden systematischen Differenzierung der Architipulidae, die sie

noch vor dem Lias in der Ober-Trias erreicht haben, muß zweifellos ein lang-

fristiger Entwicklungsprozeß der Urformen dieser Familie vorausgegangen sein.

Es ist deshalb sehr wahrscheinlich, daß sich die Architipulidae schon viel früher

als eine gute Familie gebildet haben, als sie fossil belegt sind, wahrscheinlich

schon irgendwann in der Mittel-Trias. Darauf weist die hohe Spezialisation des

Flügelgeäders der Architipulidae hin (z.B. das Verschmelzen von m, und m, bei

einigen Formen); ihrem Flügelgeäder nach stehen manche Architipulidae sogar

höher als manche rezenten Tipulidae (Hennig, 1954).

Beim Entstehen der direkten Ahnen der rezenten Tipuloidea aus den Urformen

der Eodiptera, das mit den Anpassungen ihrer Larven an eine amphibische

Lebensweise Hand in Hand ging (Giljarow, 1949), muß die Morphologie der

Larven, besonders ihrer Atmungsorgane, wesentlich verändert worden sein. Aus

den ursprünglichen peripneustischen Formen sind wahrscheinlich amphi-

pneustische entstanden, die am Vorder- und Hinterende des Leibes ein Paar

Stigmenöffnungen hatten, und von diesen wieder metapneustische Formen mit

Stigmenöffnungen nur am Hinterende des Abdomens, was für alle rezenten

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Tipuloidea mit Ausnahme von einigen Limoniidae (Antocha) kennzeichnend ist.

Bei diesen ging die Anpassung an das Wasserleben noch weiter, indem apneus-

tische Larven mit völlig isoliertem Tracheensystem ohne Stigmenöffnungen

entstanden.

Daß die Larven der Stammformen der Tipuloidea amphipneustisch waren, zeigt

sich aus vergleichend anatomischen Untersuchungen (Slipka, 1952). Wie man

weiß, steht bei den Insekten mit jedem Paar funktioneller Stigmenöffnungen einer

Queranastomose in Verbindung, die in der Körperhöhle die zwei Hauptstämme

des Trachealsystems miteinander verbindet. Gleichzeitig mit der Reduktion der

Stigmenöffnungen verschwinden auch die Queranastomosen. Bei den meta-

pneustischen Larven der Tipuloidea bleibt nur die Queranastomose im analen

Abdominalsegment erhalten, wo sich auch noch funktionelle Stigmenöffnungen

befinden. Bei den Larven von Tipula maxima Poda, die zu den primitiveren

Formen der Tipulidae gehört, wurde aber noch ein zweite Queranastomose

festgestellt; diese Anastomose befindet sich im vordersten Teil des Leibes, gleich

hinter der Basis des Kopfes, d.h. gerade an der Stelle, an der sich bei den

primitiven Formen — soferne sie amphipneustisch waren — das vorderste Paar

der Stigmenöffnungen befunden haben sollte.

Als Nachkommen der peri- und amphipneustischen Landformen sollen die

metapneustischen Larven aller primitiven Tipuloidea als sekundäre Wasser-

organismen angesehen werden. Es gibt keinen Zweifel an ihrem sekundärem

Wasserleben, weil sie ein gut entwickeltes Trachealsystem haben, das an das

Atmen von atmosphärischer Luft angepaßt ist (Gilgarow, 1949).

Die Tatsache, daß die Tipuloidea als sekundäre Wasserorganismen als ein

gesondertes Taxon von höherem Rang gerade in der Trias entstanden sind, ist

ganz verständlich. Die Trias war eine der meist geokraten Perioden der

Erdgeschichte; in dieser Zeit fand die Regression des Meeres statt und in

Zusammenhang damit eine weitgehende Aridisation des Klimas am Festland, das

in dieser Zeit den größten Teil der Erdkugel bedeckte (Sinitzyn, 1962). Durch den

starken Rückgang einerseits der Luftfeuchtigkeit anderseits des

Grundwasserspiegels müssen für das Leben der ancestralen Formen der

Tipuloidea, die in allen ihren Entwicklungsstadien sehr hygrophil sind, äußerst

ungünstige Umstände entstanden sein. Gegen diese widrigen Umstände muß es für

die Vorfahren der Tipuloidea schon in der Unter-Trias Alternativen gegeben

haben: entweder sich an aride Umstände anpassen, zeitweise in kleinen

Überlebenszonen verbleiben, die noch feucht genug sind, oder aussterben. Die

Evolution der Ahnen der Tipuloidea aus der Trias lief wahrscheinlich in Richtung

einer Änderung und Anpassung des Organismus an die veränderten Umstände,

wobei sie kleine Überlebenszonen, in denen die ökologische Situation am

wenigsten verändert war, benützten.

Die ursprüngliche Richtung einer solchen Anpassung war bei den Para-

trichoptera aus der Trias sicher die Tendenz zur Entwicklung der Zweiflügeligkeit,

was zum Entstehen der zweiflügeligen Insekten führte. Nach Rohdendorf (1964)

hat die Zweiflügeligkeit die Möglichkeiten zum Fliegen bedeutend vergrößert und

dadurch die Überlebenschancen der spezialisierten Paratrichoptera und der

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 97

archaischen Zweiflügeligen vergrößert; die Zweiflügeligkeit hat die Emigration

von trockeneren zu feuchteren Biotopen und das Suchen nach den feuchtesten

Zonen — günstig für Eiablage und Entwicklung der praeimaginalen Stadien dieser

hygrophilen Organismen — auf den großen Flächen erleichtert.

Eine andere Erscheinung dieses Anpassungsprozesses, die sich wahrscheinlich

gleichzeitig vollzog, war die Tatsache, daß die Larven der Ahnen der Tipuloidea

die Bodenschichten wechselten, ein Prozess, der auch heute in unserer rezenten

Fauna bei Schwankung von Feuchtigkeit zu bemerken ist (Gilgarow, 1951). Die

Larven der ersten Zweiflügeligen und vielleicht auch ihrer direkten panorpiden

Ahnen sind vom Leben auf der Bodenoberfläche zum Leben innerhalb feuchter

Moorböden übergegangen und das ist so geblieben bis zur Trennung der ur-

sprüngliche Eodiptera in die Untergruppen der Tipuloidea und der Bibionidea und

bei den Bibionidea bis in unsere Zeiten.

Im ersten Stadium der Evolution der Tipuloidea hatten die Larven dieser

Untergruppen sich wahrscheinlich als besserer Form zum Benützen guter Über-

lebenszonen erst dem Leben und der Entwicklung in mit Wasser gesättigten

Moorböden angepaßt und erst später eine Lebensweise direkt an den Ufern von

seichtem Wasser angenommen; in Zusammenhang damit bildete sich die

Untergruppe der Tipuloidea mit den für sie charakteristischen Larven, die die

sekundäre Eigenschaft entwickelten, im Wasser zu leben, aber den Sauerstoff aus

der Luft zu atmen.

Es ist interessant, daß der Übergang vom Leben auf dem Lande zum Leben im

Wasser zusammen mit dem Entstehen von bestimmten hygromorphen Anpas-

sungen als Art des Überlebens bei ungünstigen ariden Umständen auch bei

manchen Wirbeltieren festgestellt wurde. Die Labyrinthodonten aus der Trias

wurden bei Mangel an Feuchtigkeit gezwungen das Land zu verlassen und ins

Wasser zurückzukehren, es erfolgte die Umänderung ihrer Gliedmaßen in Flossen

(Sinitzyn, 1962).

Die Bildung der Tipuloidea in der Trias kann als ein interessantes Beispiel dafür

dienen, wie — bei wesentlicher und dauerhaften Umänderung der Umwelt über

große Flächen der Erde, wobei der Widerspruch zwischen den Anforderungen der

Tiere an ihre Umwelt und den dort gegebenen Möglichkeiten verschärft wird —

eine tiefgreifende Verzerrung und Umbildung der genetischen Basis verursacht

wurde, und in der Folge davon im Evolutionsprozess der Tierwelt das Entstehen

von Taxen höheren Ranges und von qualitativen Umänderungen in der Fauna des

Festlandes stattfand.

Die PHYLOGENIE DER TIPULIDAE

Es gibt keinen Zweifel, daß die Tipulidae wie alle anderen rezenten Tipuloidea

von den mesozoischen Architipulidae abstammen (Rohdendorf, 1964). Es bleibt

aber eine offene Frage, ob es eine direkte phylogenetische Relation gibt zwischen

den Tipulidae und Architipulidae, oder ob sich aus letztgenannter Gruppe erst

eine neue uns noch unbekannte Gruppe entwickelt hat und später aus dieser die

der Tipulidae.

Die primitivsten Familien der heutigen Tipuloidea sind die Familien Tany-

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

deridae, Trichoceridae und Ptychopteridae, Edwards (1926) meint, die

Trichoceridae haben ziemlich viele Übereinstimmungen mit eventuellen

ancestralen Formen der Tipulidae. Diese drei Familien sind aber morphologisch

sehr isoliert von den Tipulidae und nach Rohdendorf (1964) schon viel früher aus

den Architipulidae entstanden als die Tipulidae (Abb. 2). Deshalb ist die Meinung,

daß sie vielleicht die direkten Ahnen der Tipulidae sind, unbegründet.

Die Limoniidae und die Cylindrotomidae sind näher miteinder verwandt; beide

werden manchmal zu derselben Gruppe, den Polyneura gerechnet. Von diesen

beiden Familien sind nach Lameere (1906) die Limoniidae die primitivere. Nach

dem Flügelgeäder stehen sie jedenfalls den ältesten Zweiflügeligen näher als die

Tipulidae. Fast alle Limoniidae haben z.B. die Ader sc,, die bei den Cylin-

drotomidae eine Tendenz zur Reduktion sehen läßt und bei den Tipulidae

meistens gar nicht vorhanden ist. Bei einigen primitiven Gruppen der Limoniidae

hat r, die Form einer langgestreckten Ader, und die primären Verzweigungen von

Abb. 2: Schema der phylogenetischen Beziehungen zwischen den Familien der Tipuloidea und ihren

direkte Ahnen. Geochronologische Perioden: T - Trias; J - Jura; K - Kreide; P - Palaeogen; N - Neo-

gen. Familien: Td - Tipulodictyidae; A - Architipulidae; Tr - Trichoceridae; C - Cylindrotomidae; L -

Limoniidae; T - Tipulidae; Ta - Tanyderidae; P - Ptychopteridae. (Nach Rohdendorf).

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 99

rs in r,,, und von dieser in r, und r, und auch die Verzweigung von r,,; in r, und r,,

die für die Flügel der Tanyderidae kennzeichnend, aber bei allen rezenten

Tipuloidea und insbesondere bei den Tipulidae nicht vorhanden ist, sind erhalten

geblieben. Manche Limoniidae (z.B. die Pediciinae) haben als Larven noch eine

normal entwickelte Kopfkapsel, die bei den Tipulidae immer primäre

Reduktionsmerkmale zeigt.

Wahrscheinlich auf Grund dieser Tatsachen sieht Alexander (1920) die Tipu-

lidae und auch die Cylindrotomidae als Seitenast an, der sich nicht direkt von den

archaischen Tipuloidea abspaltet, sondern irgendwo an der Basis des

genealogischen Stammes der Limoniidae. Hennig (1954) stellt die Hypothese auf,

daß die Tipulidae mit bestimmten Gruppen der Limoniidae verwandt sind, eine

Verwandschaft, die wahrscheinlich viel näher ist, als man heute allgemein

annimmt.

Die Verwandtschaft zwischen Tipulidae und Limoniidae zeigt sich aus zahl-

reichen vergleichend morphologischen Tatsachen. Es ist z.B. allgemein bekannt,

daß die primitiven Dolichopezinae nach ihrem Bau den Limoniiden ähnlich sind.

Nach Byers (1961) haben die Eilarven des primitiven Genus Oropeza fast

denselben Bau des analen Segmentes wie die Larven der Limoniiden-Gattungen

Limnophila und Pseudolimnophila. Bei den Larven der Tipulidae aus den tropischen

Gattungen Brachypremna und Megistocera sind die Scheiden der Maxillarpalpen

nicht hakenformig gekrummt wie bei fast allen Arten der anderen Gattungen

dieser Familie, sondern gerade wie bei den Limoniidae. Diese und zahlreiche

andere Tatsachen zeugen aber nicht so sehr fur die Abstammung der Tipulidae

von den Limoniidae, als vielmehr fur gemeinsame Ahnen. Die wesentlichen

Unterschiede zwischen Tipulidae und Limoniidae sind sicher nicht kleiner und

auch nicht größer als ihre gemeinsamen Merkmale, wobei gerade die

Unterscheidungsmerkmale für die Phylogenie wichtiger sind als die

Übereinstimmungen.

Die Entwicklung dieser beiden Familien ist ganz verschieden. Die Tipulidae

sind durch Anpassung nur Hygro-, Helo-, Geo- und Xylobionten geworden,

während die Limoniidae, für die ein breiter ökologischer Bereich typisch ist, fast

alle Möglichkeiten benützen, die die Umwelt ihnen bietet. Außer den

obengenannten ökologischen Typen, die sie mit den Tipulidae gemeinsam haben,

sind bei den Limoniidae auch die höchsten Formen ökologischer Spezialisation

bekannt, wie Entwicklung in den höheren Formen der Pilze (Lindner, 1958a), ın

Blättern von lebenden Pflanzen (Swezy, 1915), im Boden mit Salzgehalt bis 16%o

(Slipka, 1959) und sogar im offenen Meer (Tokunaga, 1940). Als Resultat dieser

breiten adaptiven Radiation ist wahrscheinlich einerseits die größere

morphologische Verschiedenheit, anderseits die größere Artenzahl der

Limoniidae in unserer heutigen Fauna, verglichen mit der Artenzahl der

Tipulidae, zu deuten.

Im Gegensatz zu den Eiern der Tipulidae, die — von einigen Ausnahmen ab-

gesehen — lackschwarz sind, mit strukturlosem Chorion und manchmal mit

fadenförmigen Anhang, was bei keiner der anderen Familien der Tipuloidea

vorkommt, sind die Eier bei den Limoniidae (Lindner, 1958) und auch bei den

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Cylindrotomidae (Peus, 1952) fast immer hell gefarbt, halbdurchsichtig und haben

oft ein netzförmiges Chorion, ererbt von den Panorpiden (Stitz, 1926).

Das anale Segment der Tipulidenlarven hat fast immer sechs fleischige Rand-

lappen um das Stigmenfeld, bei den Limoniidae sind es nicht mehr als funf.

Phylogenetisch ist dies besonders wichtig, weil es darauf hinweist, daß im

Larvenstadium, was die Struktur des Analsegmentes betrifft, die Tipulidae näher

PRU Puree Ee e

. Abb. 3: Aedeagus von der Seite: a, b Limoniidae; c, d Cylindrotomidae (nach Peus); e Tipula paludosa

Meigen.

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 101

dem Bibioniden-Typ stehen als den Limoniidae. Die Larven der Bibionidae haben

im Analsegment ebenfalls sechs fleischige Fortsätze, die pleuralen nicht

mitgerechnet (Theowald, 1957), und die Reduktion von sechs auf funf ist

zweifellos sekundär, was die direkte Abstammung der Tipulidae von den

Limoniidae wenig wahrscheinlich macht.

Nach Edwards (1938) sind die mannlichen Imagines der Tipulidae, was den Bau

der inneren Geschlechtsorgane betrifft, sowohl von den Limoniidae als auch von

den Cylindrotomidae verschieden; meistens wird dieser Tatsache nur wenig Auf-

merksamkeit geschenkt, obwohl der Bau des Hypopygs von großer Wichtigkeit ist

zur Feststellung von Verwandtschaften innerhalb die Familie Tipulidae. Bei den

Tipulidae, Prionocera Loew ausgenommen, hat die Samenblase keine feste

Verbindung mit dem Adminiculum und der fadenförmig gebogene Penis ist

besonders lang und wächst von der Samenblase nach vorne (Abb. 3: e). Die

Limoniidae und Cylindrotomidae aber haben eine feste Verbindung zwischen

Samenblase und Adminiculum und einen verhältnismäßig kurzen Penis, der von

der Samenblase nach hinten ragt (Abb. 3: a-d). Die Verbindung zwischen

Samenblase und Adminiculum wird durch Parameren, die an den Apodema der

Samenblase befestigt sind, bewerkstelligt, und nicht durch eine direkte

Verbindung der Äste des Adminiculums an den Seiten der Samenblase wie in der

Gattung Prionocera (Tjeder, 1948).

Nach interessanten Beobachtungen von White (1949) gibt es einen großen Un-

terschied zwischen Tipulidae und Limoniidae, was die Cytologie betrifft. Obwohl

es keinen Zweifel gibt über den monophyletischen Ursprung der Tipuloidea,

nehmen unter ihnen cytologisch die Limoniidae eine ganz eigene Stellung ein. Sie

haben mehr gemeinsam mit den Culicidae und Psychodidae, während die anderen

Tipuloidea — nach White — näher den neuropteroiden Ahnen der Zweiflügeligen

stehen.

Es gibt deshalb nur wenige Tatsachen, auf Grund derer wir die Tipulidae als

einen jüngeren und höher: spezialisierten Ast der Limoniidae ansehen können;

wohl aber müssen wir mit Hennig (1950) erkennen, daß die Tipulidae und die

Limoniidae wahrscheinlich zwei Schwestergruppen sind, die einander an der Basis

begegnen, d.h. an der Stelle, an der die Verzweigung ihres ancestralen gemein-

schaftlichen genealogischen Stamms stattfand.

Im phylogenetischen Schema der Tipulidae (Abb. 2) läßt Rohdendorf (1964) die

Tipulidae unabhängig von den Limoniidae und den Cylindrotomidae von den

Architipulidae entstehen, obwohl die Ursprünge dieser drei Familien sich nah

zusammen auf dem gemeinsamen Stamm befinden.

Die meisten Architipulidae aus der Ober-Trias und dem Jura haben sc am Ende

nicht mit r, verbunden, sondern via sc, mit dem Flügelvorderrand; sc, war bei

ihnen vermutlich gar nicht vorhanden, was für die Limoniidae charakteristisch ist,

und zwar im Gegensatz zu den Tipulidae, die sc, immer haben aber nur selten

überdies sc,. Deshalb sind als Ahnen der Tipulidae nur einzelne Arten der Gattung

Architipula Handlirsch möglich (z.B. A. clara Handlirsch aus dem Lias), die eine

zweiästige sc hatten. Aber auch diese Arten underscheiden sich von den heutigen

Tipulidae durch ein ziemlich spezialisiertes Flügelgeäder. Bei ihnen war z.B. die

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Zelle M, schon gestielt, während von den rezenten Tipulidae die Gattungen

Nephrotoma und Nigrotipula die primitivere ungestielte M,, begrenzt durch m, und

m, bewahrt haben.

Wahrscheinlich gibt es aber unter die Architipulidae auch noch andere nicht

aufgefundene Formen, aus denen sich die Tipulidae entwickelt haben können.

Es gibt leider kein palaeontologisches Material, auf Grund dessen das geo-

logische Alter der Tipulidae festgestellt werden kann. Die ältesten fossilen Arten

dieser Familie wurden in Europa entdeckt, im Übergang von Eozän und Oligozän

(Alexander, 1931; Theobald, 1937), aber schon in größerer Anzahl und ziemlich

heterogen, was ihre Merkmale und systematische Stellung betrifft. In dieser Zeit

war die Familie schon differenziert in Subfamilien und Gattungen, unter denen

auch heutige vorhanden sind (z.B. Tipula von den palaearktischen und Megistocera

und Brachypremna von den tropischen Gattungen). Aus dem ziemlich langsamen

Evolutionstempo der systematischen Gruppen von höherem Rang und auch aus

der Tatsache, daß sogar die Mehrzahl der heutigen Insektengattungen aus dem

Palaeozän herstammen (Handlirsch, 1913, 1939) kann geschlossen werden, daß die

Tipulidae aus dem Palaeozän das Resultat einer ziemlich langen historischen

Entwicklung dieser Familie sind, die nicht nur den Anfang des Tertiär umfaßte,

sondern auch einen beträchtlichen Teil des Mesozoikum. Auf mesozoisches Alter

der Tipulidae weisen auch zoogeographische Tatsachen hin, insbesondere die fast

universelle Verbreitung der Familie über die Welt und die Anwesenheit mancher

ihrer Gattungen (z.B. Dolichopeza) in Australien, das zu Anfang des Känozoikums

die Verbindung mit den anderen Kontinenten verlor. Das Nichtauffinden von

Tipulidae in palaeontologischem Material aus Jura und Kreide stellt ihr

mesozoisches Alter nicht in Frage, weil die fossile Dipteren-Fauna aus dem späten

Mesozoikum noch kaum studiert worden ist.

Nach dem phylogenetischen System von Rohdendorf haben sich die Tipulidae

nicht später als im Mittel-Jura (Dogger), vielleicht schon im Unter-Jura (Ober-

Lias), entwickelt. Letzteres ist am wahrscheinlichsten. Wenn das stimmt, dann

sind die Tipulidae schon wenigsten 160— 170 Millionen Jahre alt.

Das Mittel-Jura mit humidem, ziemlich feuchtem undifferenziertem Klima und

homogener Flora (Sinitzyn, 1962) war sehr geeignet für die Entwicklung und

Verbreitung der archaischen Tipuliden über die Welt. Die archaischen Tipulidae,

dendrophile und ziemlich hygrophile Insekten, haben damals wahrscheinlich die

feuchten subtropischen und tropischen Wälder bevölkert; als Larven lebten sie

wie ihre Ahnen der Familie Architipulidae an den Ufern von seichtem Wasser

oder in amphibischen Umständen. Die sekundär bodenbewohnenden Arten, die in

unserer heutigen Fauna den Großteil des Artenbestandes ausmachen (in Ukraine

z.B. 74%), waren damals wahrscheinlich noch gar nicht vorhanden. Unter Berück-

sichtigung der palaeontologischen Funde aus dem Jura von Kara-Tau, in welcher

Fauna die Nematocera 87% aller entdeckten zweiflügeligen Insekten sind, und

unter denen die Tipuloidea eine der individuenreichsten Gruppen waren

(Rohdendorf, 1947), kann man annehmen, daß die Tipulidae im Jura eine

bedeutende Entwicklung erreicht haben.

Der Prozess des sekundären Übergangs der Tipulidae von hydro- und amphi-

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 103

bionter zu geobionter Lebensweise hat wahrscheinlich nicht vor Ende des Ober-

Jura angefangen, als die humide Phase in der Erdgeschichte sich wieder in eine

aride änderte; dieser Prozess dauerte während der ganzen Unter-Kreide, als auf

der Erde eine aride Geokratie herrschte, die aber nicht so extrem war wie in der

Mittel-Trias.

Es ist nicht unwahrscheinlich, daß gerade in der Kreide sich die wichtigsten der

heutigen Formen der Tipulidae mit ihren sekundär amphibionten und geobionten

Larven entwickelten. Die Änderung des ökologischen Milieus der Larven und im

Zusammenhang damit ihre morphologischen Anpassungen — verursacht durch

Ausbreitung des Festlandes und eine übereinstimmende Ausbreitung des Land-

klimas — fand nach Giljarow (1949) als Gefolge des Austrocknens der kleinen

Wasserflächen und der seichten Stellen an den Ufern größerer Seen statt, wo sich

vorher die archaischen Tipulidae entwickelten. Beim Übergang der Larven der

Tipulidae, die bis heute das metapneustische Atmungssystem beibehalten haben,

zum Bodenleben fand einerseits eine Reduktion des Haarsaums am Rande des

Stigmenfeldes und damit parallel eine Reduktion der Analkiemen statt, anderseits

eine Entwicklung zur Anpassung an das Bewegen in mehr oder weniger festem

Substrat durch Bildung zum Teil sklerotisierter, größenmäßig differenzierter

Randlappen um das Stigmenfeld.

Es soll bemerkt werden, daß der Übergang der Tipulidae zum Leben im Boden

im Larvenstadium über eine amphibische Umwelt nicht der einzige Weg war,

sondern nur einer von vielen in der Evolution zur terrestrischen Lebensweise im

Mesozoikum. Einen anderen Weg, der wahrscheinlich auch bedeutungsvoll für die

archaischen Tipulidae war, kann man sich so vorstellen: erst im gesättigten Moos

im Wasser, dann zum Teil in etwas trockenerem Moos und zum Teil im Wasser,

dann im Moos auf dem Lande, und schließlich im Boden. Die große Bedeutung

der Moose einerseits in der Evolution der Landflora und anderseits in der Ver-

breitung vieler Wassertiere ist bekannt. Ein interessantes Beispiel dieser

Evolutionsfolge geben die heutigen Arten der Untergattung Savtshenkia, in der alle

Übergangsformen zu finden sind. Die Larven von Tipula rufina Meigen (Theowald,

1957) z.B. können direkt im Wasser leben, die Larven von T. simulans Savtshenko

und 7. cheethami Edwards leben in mit Wasser gesättigtem, untergetauchtem

Moos, aber sie sind auch imstande, sich in sehr feuchten Landmoosen zu

entwickeln. Die meisten Savtshenkia-Arten werden in mehr oder weniger feuchten

Landmoosen gefunden, aber einige von ihnen, wie alpium Bergroth und

benesignata Mannheims & Theowald können sich auch in ziemlich trockenem

Moos entwickeln (Theowald, 1957; Savtshenko, 1963). Schließlich bevorzugen

manche der hôchstspezialisierten Arten wie odontostyla Savtshenko trockene

Moose und leben auch unter Moos im Boden. Auch darin ist eine Tendenz zu

sehen zum Übergang zu einer geobionten Lebensweise (Savtshenko, 1964a).

Gleichzeitig mit dem Entstehen der terrestrischen Larven haben sich als zweite

Möglichkeit zur Lösung der Gegensätze zwischen Hygrophilie der Familie und

Aridisation der Umwelt in der Unter-Kreide die xylobionten Familiengruppen

entwickelt, die in faulendem Holz leben. Auf den ersten Blick scheint es nahe-

liegend, daß die xylobionten Arten sich aus den terrestrischen entwickelt haben,

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

weil es keinen prinzipiellen Unterschied gibt zwischen den letzteren und den

xylobionten, was ihre trophischen Bindungen betrifft (Brauns, 1953). Das

Vorhandensein eines rudimentaren Haarsaums um das Stigmenfeld bei den

Larven der ziemlich archaischen xylobionten Ctenophora-Arten aber weist darauf

hin, daß ihre Ahnen nicht im Boden gelebt haben, sondern amphibisch waren oder

sogar im Wasser lebten. Es ist möglich, daß die Geokratie der Unteren Kreide

infolge des schnellen Austrocknens von kleinen Wasserflächen — in denen sich

die Larven der archaischen Tipulidae entwickelten — diese zum Teil zwang, sich

nicht nur an neue Lebensumstände im Boden anzupassen, sondern auch nach

anderen lokalen Überlebenszonen zu suchen, wie Mikro-Wasserbehältern, die in

den Höhlen von Bäumen in tropischen und subtropischen Wäldern entstanden

(Rohnert, 1950). Es ist bekannt, daß sich in solchen Behältern eine interessante

Fauna entwickelt, zu der auch Tipulidae gehören. Anfangs suchten die

archaischen hydrobionten Tipulidae Versteck gegen Hitze und

Feuchtigkeitsmangel, versteckten sich in Baumhöhlen und fingen dort mit dem

Ablegen der Eier in Wasserbehälter an, in denen sich dann die normale

postembryonale Entwicklung vollzog. Mit Zunahme der Aridisation der

Umgebung trockneten die Behälter in den trockenen Perioden wahrscheinlich

schnell aus, wodurch die Larven der Tipulidae trocken lagen, bevor sie sich

verpuppten. Die Jüngeren von ihnen starben, die Entwicklung der Älteren jedoch

ging weiter, weil die Reste der Pflanzen, die in den Baumhöhlen waren, noch

einige Zeit feucht blieben. So wurden die hydrobionten Formen allmählich

saproxylobiont, wobei sich dieser Prozess in ökologischer Hinsicht kaum vom

Evolutionsprozess der terrestrischen Formen dieser Familie unterschied.

Die Bildung mancher geo- und saproxylobionter Gruppen dieser Familie in der

Kreide-Zeit ist gesichert durch systematische und zoogeographische Evidenz,

insbesondere durch das Vorkommen der gleichen Formen einerseits in jenen

systematischen Gruppen von Tipulidae, deren Verbreitung auf mesozoisches Alter

deutet (Dolichopeza und einige orientalische Ctenophorinae), anderseits bei den

Tipulidae von Südostasien, wo sich die Fauna im Mesozoikum kaum änderte.

Die Blütezeit der Tipulidae war in der ersten Hälfte des Tertiär, als in der

Landfauna die Nematocera allmählich von den Brachycera ersetzt wurden. Im

Ober-Eozän wie auch im Oligozän von Europa gab es ziemlich viele Arten, die

hauptsächlich zu rezenten Gattungen gehörten (Alexander, 1931; Theobald,

1937). Außer den ausgestorbenen Gruppen der Gattung Tipula (Electrotipula und

andere) waren im Paläogen wahrscheinlich auch schon Untergattungen da, die

heute zur Fauna von Südostasiens zählen (Savtshenko, 1961).

Im Paläogen hat die Familie der Tipulidae ihre primär hygrophilen Merkmale

beibehalten. Am Ende des Paläogens, im Ober-Oligozän von Zentraleuropa zum

Beispiel, waren die Biotope, in denen die meisten Tipuliden lebten, die Gegenden

mit feuchten moorartigen Wäldern oder Sträuchern (Statz, 1943— 1944).

Wichtige Änderungen, was die weitere ökologische Spezialisation anlangt, gab

es im Neogen, insbesondere am Ende des Miozäns und im Pliozän, als in den

gemäßigten Breiten von Eurasien Abkühlung und insbesondere Aridisation des

Klimas (Ekman, 1935; Wulf, 1944) Hauptfaktoren der Evolution waren. Seit der

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 105

zweiten Hälfte des Neogen entwickelten sich die Anpassungen der Tipulidae, wie

die aller Tipuloidea (Peus, 1952), beim Übergang von feuchten zu trockenen

Lebensumständen und anschließend daran ihre ökologische Spezialisation der-

gestalt, daß sie allmählich unabhängig von der Feuchtigkeit ihrer Umwelt wurden.

Resultat davon war einerseits eine erhebliche Verarmung der Familie — ihr

Anteil an der heutigen Fauna z.B. ist nur etwa 3.9% (etwa 3200 Arten) der

Gesamtanzahl der heute bekannte Diptera (81.000 Arten) — anderseits entstand

in der Familie eine Reihe spezialisierter Gruppen von meso- und xerophilem

Typus auf Kosten der hygrophilen, helobionten und geobionten Gruppen. Wie

zoogeograpische Tatsachen zeigen, haben sich fast alle mehr oder weniger

trockenheitsliebenden systematischen Gruppen der Tipulidae, zu denen zum

Beispiel die Untergattungen Pterelachisus, Oreomyza, Vestiplex, Lunatipula und

Odonatisca der Gattung Tipula gehören, gerade während des Neogen zu eigenen

Taxa entwickelt, und manche von ihnen, wie Lunatipula, erreichten ihren größte

Blütezeit erst am Ende des Tertiär oder vielleicht noch später, als nicht nur alle

rezenten Untergattungen, sondern auch schon eine Reihe von rezenten Arten der

Tipulidae vorkamen (Savtshenko, 1961, 1964).

Es ist interessant, daß während des Tertiär und des Quartär, die zusammen

mehr als 150 Millionen Jahren währten, die Tipulidae sich verhältnismäßig wenig

differenzierten und morphologisch veränderten, ausgenommen eine Anzahl unbe-

deutender Merkmale, die in Imago oder Larve als Anpassungen an das Leben in

mehr oder weniger ariden Umständen entstanden. Dies deutet auf großen

Konservativismus, auf langsames Tempo und auf Einseitigkeit der Evolution der

Tipulidae hin, wahrscheinlich allgemeine Merkmale für die Mehrzahl von

archaischen und geologisch älteren Gruppen von Organismen. Es gibt aber auch

Grund zur Vermutung, daß die Tipulidae, wie alle Tipuloidea (Rohdendorf, 1947)

eine Restgruppe der Diptera sind, die sich mit Ausnahme von stark spezialisierten

Gattungen und Untergattungen heute in einem Zustand von Regression und

allmählichem Aussterben befinden. Darauf weist auch das Vorkommen einer

verhältnismäßig großen Anzahl von systematisch stark isolierten kleine Gattungen

mit sehr beschränktem Verbreitungsgebiet innerhald der Familie Tipulidae — in

gemäßigten wie in tropischen Zonen unserer Erdkugel — hin. Es verdient

Beachtung, daß die höchstspezialisierten xerophilen Untergattungen, die sich am

besten an die heutigen Lebensumstände angepaßt haben, heute die systematisch

meist differenzierten und artenreichsten Gruppen sind (Vestiplex, Lunatipula). Das

Beispiel der Tipulidae deutet darauf hin daß die Regel von Roz über progressive

Reduktion der Veranderlichkeit, nach der das Evolutionspotential der

Organismen mit dem Niveau ihrer Spezialisation nicht wächst, sondern absinkt,

nicht in allen Fällen stimmt.

INFRAFAMILIÄRE PHYLOGENETISCHE BEZIEHUNGEN

Die Verhältnisse innerhalb der Familie Tipulidae sind noch nicht klar. Edwards

(1926) — betreffend die Phylogenie der Nematocera — glaubt, daß die hydro-

bionten Nematocera sich aus den geo- und xylobionten entwickelt haben. Diese

Ansicht wird von ihm gestützt auf die Tatsache, daß die verhältnismäßig wenigen

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Gruppen von hydrobionten Formen unter den Nematocera meistens Merkmale

hoher Spezialisation zeigen, während die weniger spezialisierten Formen sich in

humusreichem Boden oder in faulendem Holz entwickeln.

Diese betreffend die Nematocera von Edwards unterstützte Annahme kann

nicht als Ausgangspunkt dienen zur Analyse der phylogenetischen Verhältnisse

der Tipulidae, weil sie mit vergleichend morphologischer Evidenz und auch mit

der Entwicklungsgeschichte der Familie im Widerspruch steht. Die Tatsache, daß

alle Tipulidae ohne Ausnahme, ungeachtet ihrer Ökologischen Spezialisation, im

Larvenstadium metapneustisch sind und außerdem die bei ihnen herrschende

Tendenz in der Evolution von humiden zu ariden Lebensumständen deuten darauf

hin, daß zu den weniger-spezialisierten und deshalb zu den Urformen in der Fa-

milie nicht die geo- und xylobionten sondern die hydro- und helobionten gehören.

Dies schließt nicht aus, daß manche heutige hydro- und helobionte Gruppen

dieser Familie nicht nur wenigspezialisierte sondern auch hochspezialisierte

Merkmale haben, und daß im Gegensatz dazu die geo- und xylobionten Formen

auch wenigspezialisierte Merkmale zeigen können. Das kommt, weil die Evolution

bei den verschiedenen Gruppen in dieser Familie sich erstens ungleichmäßig und

in ungleichem Tempo vollzog und zweitens nicht geradlinig sondern nach dem

Typ der dichotomen Verzweigung der Taxa.

Auf Grund eines Vergleichs der Gruppen der Tipulidae untereinander und auch

auf Grund von Vergleichen mit ihren panorpiden- und bibioniden-ähnlichen Ah-

nen können nachfolgende plesiomorphen und apomorphen Merkmale aufgestellt

werden. Die Terminologie plesiomorph und apomorph wird benutzt nach den

Auffassungen von Hennig (1950).

Merkmale

Plesiomorph Apomorph

Länglich Kurz, manchmal kugel- oder lin-

senförmig

Netzartiges Chorion Strukturloses Chorion

Kein Eifilament Eifilament vorhanden

Larve

Dorsal mit dunklen Streifen oder Dorsal einfarbig.

Flecken.

Fingerförmige, undifferenzierte Länglich konische, differen-

und beiderseits nur schwach zierte und mehr oder weniger

sklerotisierte dorsale und late- sklerotisierte (manchmal zu

rale Randlappen um das Stig- Stacheln modifizierte) dorsale

menfeld. und laterale Randlappen um das

Stigmenfeld.

Ein Haarsaum am Rande des Kein Haarsaum am Rande des

Stigmenfeldes. Stigmenfeldes.

Analkiemen vorhanden. Keine Analkiemen.

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 107

(Plesiomorph)

Deutliche Ausstülpungen um die

Analöffnung.

Zweites und drittes Makrochaet

links und rechts am Hinterrand

der abdominalen Tergite nah

zusammen.

Ziemlich viele kleine Dörnchen

an den Hinterrändern der letzten

abdominalen Sternite.

Ozellen rudimentär.

Geißel der Antennen mit zylin-

derförmigen Gliedern oder

gesägt.

Flügelfläche mit Mikrotrichien.

Ader sc, und/oder überzählige

Querader anwesend.

Flügel quergestreift oder ge-

fleckt.

Zelle M, ungestielt.

Lange dünne fadenförmige

Beine.

Beim Männchen sind 9. Tergit

und 9. Sternit deutlich getrennt.

Deutlich differenzierter Basi-

stylus.

Id einfach gebaut.

8. Sternit beim Männchen

undifferenziert.

Langgestreckter sklerotisierter

Ovipositor mit deutlich ent-

wickelten Hypovalven.

9. Tergit beim Weibchen normal

entwickelt.

(Merkmale)

Puppe

Imago

(Apomorph)

Kaum oder keine Ausstülpungen

um die Analöffnung.

Zweites und drittes Makrochaet

links und rechts am Hinterrand

der abdominalen Tergite weit

auseinander.

Eine konstante Zahl von

größeren Dornen an den Hinter-

rändern der abdominalen

Sternite.

Ozellen fehlen.

GeiBelglieder an der Basis

verdickt oder kammförmig ver-

zweigt.

Flügelfläche nackt.

Keine Ader sc, und keine

überzählige Querader.

Flügel einfarbig.

Zelle M, gestielt oder die Adern

m, und m, verschmolzen.

Beine kürzer und dicker.

Beim Männchen sind 9. Tergit

und 9. Sternit ohne Naht

ringförmig miteinander ver-

schmolzen.

Basistylus undeutlich oder re-

duziert.

Id kompliziert gebaut.

8. Sternit beim Männchen kom-

pliziert gebaut mit Anhängen

oder Ausstülpungen.

Verkürzter fleischiger Ovipositor

mit reduzierten Hypovalven.

Weibchen

9. Tergit beim

reduziert.

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

(Plesiomorph) (Apomorph)

Biologie

Hydro- und hygrophile Arten. Meso- und xerophile Arten.

Die Eier werden einzeln oder in Viele Eier werden zusammen in

Klumpen abgelegt. tieferen Bodenschichten abge-

setzt.

Die Larven entwickeln sich im Die Larven entwickeln sich in

Wasser oder in mit Wasser trockenerem Boden oder in

gesättigtem Boden (Moorgebie- faulendem Holz.

te, feuchtes Moos).

Die Larven ernähren sich von Die Larven ernähren sich von

faulenden organischen Stoffen lebenden höheren Pflanzen.

oder von Moos.

Der Entwicklungszyklus ist bi- Der Entwickiungszyklus ist mo-

voltin oder polyvoltin. novoltin.

Entwicklung ohne Diapause. Entwicklung mit Diapause.

Bemerkung: In Anbetracht der Tatsache, daß bei den Larven der Bibionidae

die dorsalen und lateralen Randlappen um das Stigmenfeld auch länglich-konisch

sind, glaubt Theowald (1957), daß diese Form bei den Tipulidae die primitivste

darstellt. Ich bin damit nicht einverstanden, weil die Form dieser Randlappen bei

den primären Bodenlarven der Bibionidae und den sekundären Bodenlarven der

Tipulidae nicht homolog sondern analog entstanden ist. Sie sind konvergent

entstanden als Anpassung an gleichartige Lebensumstände in einem dichten

Substrat.

Die meisten plesiomorphen Merkmale haben die Dolichopezinae, zu denen in

unserer Fauna die Gattung Dolichopeza gehört. In dieser Tribus findet bei den

Arten der Gattung Megistocera Wiedemann (tropisch) die ganze praeimaginale

Entwicklung im Wasser statt (Rogers, 1949). Die hygromorphen Larven von Me-

gistocera haben das Stigmenfeld am primitivsten gebaut, was sie vielleicht noch

von den archaischen Tipulidae aus dem Mesozoikum geerbt haben. Stigmenfeld

und Randlappen sind mit langen Haaren gesäumt. Die Flügel sind mit

Mikrotrichien bedeckt, wodurch sie gegen das naßwerden geschützt sind. Die

starke Beziehung zum Wassermilieu bei Megistocera wird auch aus der Tatsache

sichtbar, daß die Weibchen zum Absetzen der Eier und auch zur Ruhe auf der

Wasseroberfläche sitzen können, wobei sie sich mit den außerordentlich langen

Beinen stützen.

Als meist plesiomorphe Tribus der Familie haben die Dolichopezinae sehr viele

Merkmale mit den Limoniidae gemeinsam, was sich bei manchen von ihnen im

Bau des analen Segmentes der Eilarven, in den geraden statt hakenförmig ge-

krümmten Scheiden der Maxillarpalpen, im ziemlich einfachen Bau des Hypopygs

der Männchen und bei vielen Gattungen auch im Vorhandensein einer normal

entwickelten sc, zeigt.

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 109

Do Ti Ct

Abb. 4: Hypothetisches Schema der phylogenetischen Verhaltnisse zwischen den Unterfamilien und

Gattungen der Tipulidae: Die Namen der Perioden sind wie in Abb. 3 abgekurzt. Unterfamilien: Do -

Dolichopezinae; Ti - Tipulinae; Ct - Ctenophorinae. Gattungen: D - Dolichopeza; P - Prionocera; Ni -

Nigrotipula; T - Tipula; Ne - Nephrotoma; Di - Dictenidia; C - Ctenophora; Ta - Tanyptera.

Die zu unserer Fauna zahlende Gattung Dolichopeza gehort nicht zu den pri-

mitiven, sondern zu den mehr apomorphen Gruppen der Dolichopezinae; sie

zweigte irgendwann um Mitte Kreide ab und spezialisierte sich mehr in Richtung

Anpassen an aridere Umstande. Die larvale Entwicklung findet nicht mehr im

Wasser, sondern in feuchten Landmoosen statt. Die Larven haben längliche, fast

konische Randlappen und einen kurzen Haarsaum um das Stigmenfeld und

verhältnismäßig kurze Analkiemen und zeigen den apomorphen Typ der

Chaetotaxie; die Puppen haben kurze, am Ende gebogene Mesothorakal-Hörner,

wie sie bei den meisten Tipulidae vorkommen, und die Imago hat eine offene

Diskoidalzelle durch teilweise reduzierte m,. In vielen Fällen unterscheidet sich

Dolichopeza sogar durch eine größere Apomorphie von manchen anderen

Gattungen der Tipulidae in unserem Faunengebiet, was die Möglichkeit einer

direkten phylogenetischen Verbindung mit ihnen ausschließt (Abb. 4).

Ziemlich nah verwandt mit den primitiven Dolichopezinae ist die Gattung Prio-

nocera der Tipulidae. Die Larven leben an den Wurzeln von Wasserpflanzen

(Theowald, 1957) oder im Wasser (Nielsen, 1954). Der Bau ihres analen Segmentes

ist ebenso plesiomorph wie in der Gattung Megistocera. Die Puppen haben an den

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Hinterrändern der abdominalen Segmente viele kleine Dörnchen, die

charakteristisch sind für die plesiomorphen Formen der Tipulidae, und die Imago

hat rudimentäre Ozellen und ein ganz einfach gebautes Hypopyg (Tjeder, 1948).

Damit erinnert sie an die Dolichopezinae und auch an die Limoniidae. Die

Imagines haben aber auch apomorphe Merkmale. Sie haben zum Beispiel die für

die anderen Tipulinae typische Flügeladerung. Manche anderen Merkmale wie

die dunkle Färbung und die Behaarung des Leibes, die Entwicklung in moorigen

Biotopen des nördlichen Typs und die Verbreitung überwiegend in höheren

Breiten, wo die Fauna noch ziemlich jung ist, deuten auf verhältnismäßig hohe

Apomorphie der Gattung Prionocera, die wahrscheinlich nicht vor dem Ende des

Neogen entstand und möglicherweise erst im Pleistocän ihre Blütezeit erreichte.

Eine derartige Kombination von plesiomorphen und apomorphen Merkmalen bei

Prionocera muß als Nachweis gedeutet werden, daß diese Gattung phylogenetisch

mit einer sehr alten und primitiven, den Dolichopezinae nah verwandten Gruppe

dieser Familie, die schon in der Kreide von ihnen abzweigte, verbunden ist; sie

war schon im Palaeogen ein ganz unabhängiger Zweig und starb am Anfang des

Neogen aus, als eine neue geokrate Phase in der Erdgeschichte anfing. Es blieb

nur ein kleiner spezialisierter Seitenzweig übrig (Abb. 4), der sich zur Gattung

Prionocera, angepaßt an das Leben im kalten nordborealen Klima, entwickelte.

Von den primitiven Gruppen der Dolichopezinae zweigten wahrscheinlich

schon in der Kreide die ancestralen Formen der anderen Gattungen der Tipulinae

unseres Faunengebiets ab, wie die Ahnen von Nigrotipula, Tipula und Nephrotoma,

die zusammen einen ziemlich natürlichen genetischen Komplex bilden.

Die Gattung Nigrotipula hat bei den Larven plesiomorphe und apomorphe

Merkmale. Plesiomorph ist zum Beispiel bei den Larven das Vorhandensein eines

reduzierten Haarsaums am Unterrand des Stigmenfeldes; apomorph sind das

Fehlen einer hydro- oder hygromorphen Anpassung und die geobionte

Lebensweise. Die Imago hat noch eine ziemlich plesiomorphe Flügeladerung, für

die eine ungestielte oder nur kurz gestielte Zelle M, kennzeichnend ist; das

Hypopyg ist ganz einfach gebaut. All dies deutet darauf hin, daß der Ursprung

dieser Gattung in der Nähe einer primitiven alten Gruppe der Tipulinae, die

wahrscheinlich noch helobionte Larven hatte, zu suchen ist. Es ist interessant zu

wissen, daß die Nigrotipula-Arten die gleiche Kombination einer Reihe von

Merkmalen haben wie die Gattungen Tipula und Nephrotoma. Mit der ersteren

haben sie die Lage von sc, im Bezug auf die Basis von rs, die Anzahl von tibialen

Spornen (1-2-2) und den Bau des Prothorax der Larven gemeinsam, mit

Nephrotoma die kurze Schnauze, eine ungestielte Zelle M,, den mit

mikroskopischen Dörnchen versehenen Einschnitt am Hinterrand des 9. Tergits

bei den Männchen und die Anzahl der Dornen an den Hinterrändern der

abdominalen Tergite bei der Puppe. Es muß deshalb angenommen werden, daß

die ancestralen Formen, aus denen sich die Gattung Nigrotipula entwickelt hat,

vom gemeinsamen Stamm der Tipulinae vor dem Divergieren in die Gattungen

Tipula und Nephrotoma abzweigten und auch, daß diese ancestralen Formen, die

schon lange ausgestorben sind, keine apomorphen Merkmale hatten, wie sie

später während des Anpassungsprozesses der Annahme einer geobionten

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 111

Lebensweise gegenüber einer Anderung der palaeogenen Sumpfmoore im

Torfboden in der Gattung Nigrotipula entstanden sind.

Über den monophyletischen Ursprung der Gattungen Tipula und Nephrotoma,

die soviele Merkmale mit den Dolichopezinae, aber auch untereinander gemein-

sam haben, gibt es keinen Zweifel. Gemeinsam sind für sie sogar Merkmale wie

der Typ der Körperfärbung und die Stelle der Abzweigung der Ader r,, beides

spezifisch für beide Gattungen. Der lackglänzende Thorax ist nicht nur

charakteristisch für die Gattung Nephrotoma, sondern auch für manche auf Grund

anderer Merkmale in die Gattung Tipula gehörende Arten, zum Beispiel die in die

palaearktische Untergattung Dendrotipula gehörende nordchinesische T. hoi

Alexander und insbesondere für die Arten der nearktischen Untergattungen

Nephrotomodes und Nobilitipula. Die Stelle des Verzweigungspunktes der Ader

m,,, an der Basis der Diskoidalzelle ist nicht auf die Gattung Tipula beschränkt,

sondern kommt ausnahmsweise auch bei manchen charakteristischen Arten der

Gattung Nephrotoma vor (Edwards, 1928).

Die Gattung Tipula zweigte wahrscheinlich in der Kreide vom Hauptstamm der

Tipulinae und später, irgendwann am Ende des Palaeogens oder am Anfang des

Neogens, erreichte sie ihre größte Polymorphie und Differenzierung (Abb. 4).

Obwohl das Genus Nephrotoma denselben Ursprung hat wie die Gattung Tipula,

zweigte es wahrscheinlich nicht am Ende des Mesozoikums oder am Anfang des

Palaeogen direkt von Tipula ab, sondern von irgendeiner mehr plesiomorphen aus-

gestorbenen Zwischengruppe der Tipulinae, die wahrscheinlich nahe verwandt

war mit der von Alexander (1931) beschriebenen Untergattung Electrotipula,

gefunden im baltischen Bernstein, deren einzige bekannte Art (E. pinetorum

Alexander) im Flügelgeäder Merkmale sowohl von Tipula und als auch von

Nephrotoma zeigt.

Obwohl Nephrotoma einen ziemlich hohen Grad von ökologischer Spezialisation

erreichte, erlangte diese geobionte Gattung nicht den Umfang und den Grad der

Differenzierung der Gattung Tipula; dies kann einerseits durch das jüngere geo-

logische Alter, anderseits durch den engeren ökologischen Bereich ihrer

Anpassungen bedingt sein. Das fast völlige Fehlen von Fossilien, die zur Gattung

Nephrotoma gehören, ist wohl ein Hinweis darauf, daß in der Vergangenheit diese

Gattung nie artenreich war und wahrscheinlich nie eine derartige Blütezeit

erreichte wie in unserer heutigen Fauna.

Die einheimischen Gattungen Ctenophora und Tanyptera, die ein natürlicher

Komplex von Arten mit gekämmten Geißelgliedern sind, sind innerhalb dieser

Familie durch die höchste ökologische Spezialisation gekennzeichnet: ihre Larven

sind saproxylobiont, die der Tanyptera-Arten sogar fakultativ xylobiont. Im

imaginalen Stadium sind diese Gattungen überwiegend apomorph. In der Gattung

Ctenophora zum Beispiel ist der Bau der id sehr kompliziert apomorph und bei den

spezialisierten Arten der Gattung Tanyptera sogar das ganze Hypopyg.

Gleichzeitig zeigen die Arten dieser Genera auch viele plesiomorphe Merkmale:

kurze Schnauze, oft ungestielte oder kurzgestielte Zelle M, und einen ziemlich

primitiv gebauten Basistylus als Imagines, außerdem als Larven und Puppen einen

primitiven Typ von Chaetotaxie. Phylogenetisch hat ein plesiomorphes Merkmal

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

wie das Vorhandensein eines Haarsaums um das Stigmenfeld bei den Larven von

Ctenophora große Bedeutung. Es deutet darauf hin, daß die Gattung Ctenophora

und damit alle nah miteinander verwandten Ctenophorinae, nicht von den

apomorphen geobionten, sondern von den plesiomorphen helobionten Gruppen

dieser Familie abzweigten. Es ist wahrscheinlicher, daß sich die Ctenophorinae

aus einer primitiven amphibischen oder sogar aus einer im Wasser lebenden

ancestralen Form entwickelt haben, die im Mesozoikum von den Dolichopezinae

abzweigte und sich später sowohl zu Tipulinae als auch zu Ctenophorinae

entwickelte (Abb. 4).

Die frühesten archaischen Ctenophorinae hatten wahrscheinlich — wie die heu-

tigen Arten der Gattung Dictenidia — an der Basis und am Ende jedes

Geißelgliedes Fortsätze, die sich aus den proximalen und distalen Verdickungen

dieser Segmente, die kennzeichnend sind für viele Tipulidae, entwickelt haben. Es

gab dabei eine Verschiebung eines Teiles der Wirtelhaare zum Ende des unteren

Fortsatzes. Im weiteren Evolutionsprozess, in dem die Geißelglieder durch

Spaltung eines oder beider Fortsätze komplizierter wurden, entstanden die

Formen mit drei (Tanyptera) oder vier (Ctenophora) Fortsätzen an jedem

Geißelglied.

Am meisten plesiomorph und wahrscheinlich am ältesten unter den Cteno-

phorinae unseres Faunengebietes ist aber doch wohl die Gattung Ctenophora und

nicht die Gattung Dictenidia. Es gibt keinen Zweifel, daß Ctenophora irgendwo in

den gemäßigten Teilen von Ostasien zu Ende des Mesozoikums oder am Anfang

des Tertiär als unabhängiges Taxon abzweigte. Was die Gattung Dictenidia betrifft,

sie war wahrscheinlich ein “Seitenast’’ einer anderen mehr plesiomorphen, später

jedoch ausgestorbenen Gruppe der Ctenophorinae; sie zweigte von dieser Gruppe

bedeutend später ab, wahrscheinlich erst Ende Palaeogen oder Anfang Neogen.

Tanyptera zweigte wohl von derselben ausgestorbenen Gruppe ab, aber nicht

später als in Palaeogen. Was die Spezialisation in den Entwicklungsstadien

anlangt, hat Tanyptera sich aber deutlich weiter entwickelt als die anderen

Gattungen. Im Larvenstadium, in dem Tanyptera habituell an die Larven von

Bockkäfern erinnert, hat sie sich nicht nur an das Leben in weichem faulendem

Holz angepaßt, sondern auch an das Leben in hartem Holz.

Die phylogenetischen Beziehungen zwischen den Artengruppen innerhalb der

Gattungen der Tipulidae sind noch nicht klar. Die Gattung Dolichopeza hat in

unserem Faunengebiet (Ukraine) nur eine ziemlich spezialisierte Art. Die Männ-

chen dieser Art haben ein Adminiculum ohne stabförmige Verbindungen mit der

Vesica. Diese Verbindungen kommen noch wenig entwickelt vor bei der

nearktischen D. americana Needham und in schon ganz gut entwickelter Form bei

manchen Arten der Untergattung Oropeza, die für die Fauna von Ostasien und

Nordamerika charakteristisch ist (Byers, 1961).

Die meisten der ukrainischen Prionocera-Arten, die nur wenig deutlich gesägte

Antennen und einen ziemlich stark behaarten Körper haben, sind wahrscheinlich

ziemlich plesiomorph und damit ziemlicht alt.

Die einzige ukrainische Art der Gattung Nigrotipula ist ziemlich apomorph. Sie

hat eine dunkle Färbung, die bei den Tipuliden sekundär ist, und einen kom-

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 113

plizierten und spezialisierten Bau der id. Bei den verwandten Arten aus dem

Norden von Ostasien ist die Körperfärbung meistens braungelb und die id sind

einfacher gebaut.

Innerhalb der Gattung Tipula (Abb. 5) ist die Untergattung Platytipula wohl am

meisten plesiomorph. Die Flügelzeichnung und der Bau des Hypopygs erinnern an

die Dolichopezinae. Alexander (1926) meint, daß manche ostasiatischen Arten

dieser Untergattung, die sich vermutlich schon in der Kreide als unabhängiges

Taxon differenzierten (Savtshenko, 1961), direkt mit den Dolichopezinae ver-

wandt sind.

Als ziemlich apomorphe und junge Abzweigung der Untergattung Platytipula

oder einer mit ihr nahverwandten ausgestorbener Untergattung muß man wohl

Schummelia ansehen, die sich wohl nicht später als im Eozän entwickelt haben

dürfte. Flügeladerung und Eigentümlichkeiten im Bau des Hypopygs weisen auf

eine Verwandtschaft von Platytipula und Schummelia hin.

Die Untergattungen Acutipula, Yamatotipula und Tipula s.str. stammen von den

archaischen Zweigen der Gattung Tipula. Sie bilden zusammen einen genetischen

Komplex, der nach Theowald (1957) den Rang einer Gattung verdient. Sie sind

durch eine Reihe von plesiomorphen Merkmalen charakterisiert, zum Beispiel

durch helobionte und fakultativ hydrobionte oder ihnen sehr ähnliche primitive

geobionte Larven. Die Untergattung Acutipula ist die primitivste und älteste von

ihnen, sie stammt wohl aus dem Palaeogen. Von gemeinsamen ancestralen For-

men und parallel mit ihr entwickelte sich seit dem Oligozän die Untergattung

Yamatotipula. Und im Miozän zweigte vermutlich von Acutipula die Untergattung

P S Y A T Sal eM 8 Pt (o) 2 L Od D

Abb. 5: Hypothetisches Schema der phylogenetischen Verhältnisse zwischen den Untergattungen der

Gattung Tipula. Geochronologische Perioden: K - Kreide; P - Palaeozän; E - Eozan; O - Oligozan; M -

Miozän; PI - Pliozän. Untergattungen: P - Platytipula; S - Schummelia; Y - Yamatotipula; A - Acutipula; T

- Tipula s.str.; Sa - Savtshenkia; M - Mediotipula; B - Beringotipula; Pt - Pterelachisus; O - Oreomyza; V -

Vestiplex; L - Lunatipula; Od - Odonatisca; D - Dendrotipula.

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Tipula s.str. ab, die in diesem Komplex wohl am meisten apomorph ist

(Savtshenko, 1961). Alle Untergattungen dieses Komplexes haben nicht nur einen

ahnlichen Bau des analen Segments der Larven und eine ahnliche Chaetotaxie der

Puppen, sondern auch einen ahnlichen Bau des Hypopygs der Mannchen. Die

geobionte und apomorphe Untergattung Tipula ausgenommen, haben beide

anderen Untergattungen ihre Blütezeit wohl am Ende des Palaeogen oder im

Miozan erreicht; darauf weist die Verschiedenheit ihrer archaischen Formen in

der ursprünglichen Fauna von Südostasien hin.

Die Untergattungen Savtshenkia, Mediotipula und Beringotipula haben taxo-

nomisch eine sehr isolierte und phylogenetisch unsichere Stellung innerhalb der

Gattung Tipula. Das Vorkommen von rudimentaren Analkiemen und ein

rudimentärer Haarsaum um das Stigmenfeld bei den Savtshenkia-Larven die sich

in xeromorpher Richtung entwickelten, deuten auf eine Abstammung dieser

Untergattung von plesiomorphen, helo- und hydrobionten Gruppen aus der

Gattung Tipula. Der charakteristische Fortsatz am Ende der Hypovalven der

weiblichen Puppen (in unserer Fauna auch bekannt von den weiblichen

Dolichopeza-Puppen) bringt die Untergattung Savtshenkia sogar näher zu den

Dolichopezinae. Es ist interessant, daß dieser Fortsatz auch bei den weiblichen

Puppen von Mediotipula vorkommt, deren Arten — was den Komplex der anderen

Merkmale betrifft — ziemlich stark apomorph sind. Deshalb muß Mediotipula,

ungeachtet ihrer hohen Spezialisation, wahrscheinlich doch von den archaischen

Gruppen der Gattung Tipula abgeleitet werden. Dasselbe gilt auch für die

Untergattung Beringotipula, deren Larven nicht nur apomorphe Merkmale zeigen

wie eine starke Sklerotisation der Randlappen um das Stigmenfeld, sondern auch

plesiomorphe wie einen Haarsaum um das Stigmenfeld.

Die meistapomorphen Zweige der Gattung Tipula sind die geologisch ziemlich

jungen Untergattungen Prerelachisus, Oreomyza, Vestiplex, Lunatipula und Odo-

natisca, ein Komplex von Untergattungen, der möglicherweise auch als Gattung

qualifiziert werden kann (Theowald, 1957). Eine Reihe von Merkmalen deuten auf

die Verwandtschaft dieser Untergattungen, die wahrscheinlich irgendwann im

Oligozän oder Miozän anfingen sich in xeromorphe Richtung zu entwicklen, und

am Ende des Neogens eine beträchtliche Blütezeit erreichten — Lunatipula und

Odonatisca wahrscheinlich noch später —, und zwar Merkmale wie die

Flügeladerung und die Flügelzeichnung, der Bau des Hinterrandes des 9. Tergites

bei den Männchen, der geobionte Larventyp mit mehr oder weniger sklerotisierter

Oberfläche der dorsalen und lateralen Randlappen um das Stigmenfeld, die stark

entwickelten aber nicht zahlreichen Dornen auf dem Abdomen der Puppen u.s.w.

(Savtshenko, 1964). Diese Untergattungen, die ziemlich apomorph sind, zeigen

aber auch manche plesiomorphe Merkmale, zum Beispiel eine primitive

Chaetotaxie bei den Larven, die wahrscheinlich von ancestralen Formen, die, wie

anzunehmen ist, weniger spezialisierte und mehr archaische Gruppen der Gattung

Tipula waren, vererbt wurden.

Den ancestralen Formen am nächsten stehen wohl die Untergattungen Ptere-

lachisus und Oreomyza, von denen die zweite direkt von der ersten abzweigte.

Direkte Verwandtschaft zwischen beiden sieht man einerseits bei T. (Pterelachisus)

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 115

mutila Wahlgren und anderseits bei 7. (Oreomyza) trifasciata Loew, die viele

Merkmale gemeinsam haben, einerseits im Bau des Hypopygs der Männchen,

anderseits in der Flügeladerung (Savtshenko, 1964).

An einer Abzweigung der Untergattung Vestiplex von einer mit der Unter-

gattung Pterelachisus gemeinsamen Wurzel besteht kein Zweifel. Bis in unsere Zeit

findet man in der Fauna von Ostasien Übergangsformen zwischen beiden

Untergattungen (zum Beispiel T. ambigua Savtshenko), die manchmal nur

schwierig in einer dieser Untergattungen unterzubringen sind, weil sie nach dem

Bau des Ovipositors zu Vestiplex, nach dem Bau des Hypopygs aber zu

Pterelachisus gehören (Savtshenko, 1964).

Die Untergattung Lunatipula nimmt eine stärker isolierte Stellung ein. Die an-

cestralen Formen dieser Untergattung zweigten von der für den ganzen Komplex

gemeinsamen Wurzel wahrscheinlich noch vor der Abzweigung der

Untergattungen Pterelachisus und Vestiplex ab. Direkt aus Lunatipula differenzierte

sich im Neogen ohne Zweifel die Untergattung Odonatisca, die am meisten

xerophile und jüngste Untergattung dieses Komplexes. Die nahe phylogenetische

Verwandtschaft zwischen Lunatipula und Odonatisca wird bestätigt durch

gemeinsame Merkmale im Bau der Larven und Puppen. Die Larven von

Odonatisca zum Beispiel haben dieselben gänzlich sklerotisierten, hakenförmigen

dorsalen und lateralen Randlappen um das Stigmenfeld wie die Larven der /ivida-

Gruppe der Untergattung Lunatipula, und, was den Bau der Puppen betrifft, zeigen

sie nur wenige Unterschiede gegenüber den im Mittelmeergebiet vorkommenden

Arten der falcata-Gruppe der Untergattung Lunatipula. Der Anhang am

Hinterrand des 8. Sternits bei den Männchen von Odonatisca kann leicht von den

homologen, jedoch nicht so stark entwickelten Strukturen am Hinterrand dieses

Sternites bei den Arten der falcata-Gruppe abgeleitet werden.

Die stark apomorphe Untergattung Dendrotipula, die sich irgendwann im Oli-

gozän entwickelte, erreichte ihre Blütezeit im Miozän und entwickelte sich sehr

weit in Richtung Anpassung der praeimaginalen Stadien an eine Ernährung von

Verwesungsprodukten faulenden Laubholzes. Apomorphe Merkmale wie zum

Beispiel die Reduktion der dorsalen und lateralen Randlappen des Stigmenfeldes

der Larven und die Verkürzung der mesothorakalen Atmungsröhrchen bei den

Puppen finden wir auch bei den Ctenophorinae. Es besteht aber keine direkte

phylogenetische Verwandtschaft zwischen beiden, weil die Larven von

Dendrotipula keinen Haarsaum um das Stigmenfeld haben und mehr apomorph

sind als die Larven der Gattung Ctenophora, bei denen ein solcher Haarsaum

vorhanden ist. Theowald (1957) weist auf eine Verwandtschaft der Untergattung

Dendrotipula mit den Untergattungen Pterelachisus und Oreomyza hin, und zwar

auf Grund des Baues des analen Segments ihrer Larven und Puppen. Eine

Verwandtschaft dieser drei Untergattungen zeigt auch der plesiomorphe Typ der

Chaetotaxie ihrer Larven an. Darauf begründet, gewinnt die Hypothese, daß die

ancestralen Formen von Dendrotipula von einer mit den Untergattungen

Pterelachisus und Oreomyza gemeinsamen Wurzel abzweigten, und zwar noch vor

der Abzweigung von Lunatipula und Vestiplex (Abb. 5), an Wahrscheinlichkeit.

Nephrotoma ist im imaginalen wie im larvalen Stadium nur sehr undeutlich

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

gegliedert, weshalb auch sichere Aussagen uber die Verwandtschaftsbeziehungen

innerhalb dieser Gattung schwierig zu machen sind. Man kann nur annehmen, daß

unter den ukrainischen Arten dieser Gattung die Arten der scalaris-crocata-

Gruppe, die sich durch dunkle Leibespigmentierung und manche von ihnen durch

eine Tendenz zum Leben auf offenem Feld in ziemlich trockenen Umständen

(N. rossica Riedel) auszeichnen, am meisten apomorph sind.

Von Dictenidia gibt es in der ukrainischen Fauna nur eine ziemlich apomorphe,

dunkel gefärbte Art, deren Hypopyg komplizierter gebaut ist als das der anderen,

hell gefärbten Arten dieser Gattung aus Südostasien. Unsere Art repräsentiert

unter den Dictenidia-Arten deshalb vielleicht einen jüngeren und mehr spezia-

lisierten Zweig.

Innerhalb der Gattung Ctenophora sind auf Grund des Baues des praeimaginalen

Stadien die ukrainischen Arten der pectinicornis-guttata-Gruppe, deren Larven

deutlich entwickelte Randlappen um das Stigmenfeld haben, mehr plesiomorph

und die Arten der fastuosa-festiva-Gruppe mit reduzierten Randlappen mehr

apomorph. Nach imaginalen Merkmalen sind beide Gruppen verbunden durch die

elegans-Gruppe als Übergangsform. Es ist nicht klar erkennbar, wie die Evolution

vor sich ging: von der ersten zur zweiten Gruppe oder umgekehrt. Letzteres sieht

wahrscheinlicher aus, weil die fastuosa-festiva-Gruppe für die alte Fauna von

Südostasien charakteristisch ist und die pectinicornis-guttata-Gruppe für die

verhältnismäßig junge boreale Fauna.

Ganz sicher sind die phylogenetischen Verhältnisse innerhalb der Gattung

Tanyptera. Die ukrainischen Arten dieser Gattung sind der jüngste apomorphe

Zweig, dessen Männchen einen sehr stark spezialisierten Bau des Hypopygs

haben, das durch die nur kleine Öffnung und eine fast horizontale Lage der Styli

im Genital-Raum prinzipiell von demselben Organ der anderen Tipulidae ver-

schieden ist. Dieser Zweig entwickelte sich in den gemäßigten Breiten aus den

plesiomorphen ostasiatischen Arten der Gattung, die ein Hypopyg des normalen

Tipulidae-Typs haben, das heißt mit breiter Öffnung und mehr oder weniger

vertikaler Lage der Styli. Zwischen diesen beiden Gruppen finden wir in unserer

heutigen Fauna noch eine Übergangsform, die auf die Richtung der Evolution in

dieser Gattung hinweist. Es ist eine kleine und ziemlich seltene Art aus dem

Fernen Osten (T. parva Portschinsky), die in den Küstengebieten der Sovjet-Union

(Primorskij Kraj) und in Japan vorkommt.

Die Hauptblütezeit der Gattungen Crenophora und Tanyptera fand wahr-

scheinlich im Neogen statt, als sich die Laubwälder, an die diese Gattungen

ökologisch und trophisch gebunden waren, in den gemäßigten Breiten von

Eurasien im Norden und im Westen viel weiter ausdehnten als heute.

SYSTEMATIK UND KLASSIFIKATION

Die moderne Systematik der Tipulidae ist noch recht ungenügend bearbeitet

und ist hauptsächlich auf die Imagines und hier wiederum auf morphologische

Merkmale begründet. Erst in letzter Zeit sind die ersten Versuche gemacht

worden, auch die Larven und die biologischen Merkmale für ein neues System

SAVTSHENKO: Phylogenie und Systematik der Tipulidae 117

dieser Familie zu benützen (Theowald, 1957; Savtshenko, 1961).

Seit Latreille (1802), der die Familie Tipulidae für die Gattung Tipula von

Linnaeus errichtete, haben während des neunzehnten Jahrhunderts die meisten

Systematiker diese Familie ganz breit aufgefaßt, und neben den Tipulidae auch

die Limoniidae und Cylindrotomidae eingeschlossen, manchmal sogar auch die

Trichoceridae und die anderen Tipuloidea (Zetterstedt, 1842; Rondani, 1856;

Schiner, 1864; Osten-Sacken, 1878, und andere). Diese breite Auffassung der

Familie als Komplex von Tipulidae, Limoniidae und Cylindrotomidae, die dann

nur den Rang von Unterfamilien erhalten, akzeptieren auch die heutigen

englischen und amerikanischen Systematiker (Edwards, 1938; Alexander, 1942;

Coe, 1950; und andere).

Seit Kertesz (1902) sieht die Mehrzahl der europäischen Systematiker die

Tipulidae als eine unabhängige Familie neben Limoniidae und Cylindrotomidae

an, meistens jedoch ohne besonderere Begründung ihres Standpunktes (Hendel,

1928, 1936/37; Hennig, 1950; Mannheims, 1951; Theowald, 1957a; Savtshenko,

1961). Nur Peus (1952) bemerkt, daß eine derartige Einteilung und Qualifikation

mehr den Forderungen eines natürlichen Systems entspricht und überdies Vorteile

in Bezug auf Übersichtlichkeit hat. |

Die Tipulidae verdienen aber den Rang einer Familie nicht nur auf Grund von

Übersichtlichkeit, sondern auch aus mehr prinzipiellen Erwägungen. Wenn man

die höheren systematischen Gruppen nicht nur als etwas sieht, das seine Ursache

in dem Bedürfnis des Menschen zum systematisieren hat (Peus, 1952), sondern als

reelle Kategorien, die konkret die Entwicklungsgeschichte und die heutige Struk-

tur von bestimmten Gruppen von Organismen darstellen, dann wird der

Familienrang der Tipulidae keinen Widerspruch entfachen dürfen. Wie schon

früher bemerkt, unterscheiden sich die Tipulidae wesentlich von den am besten

mit ihnen übereinstimmenden anderen Gruppen der Tipuloidea, insbesondere den

Limoniidae und Cylindrotomidae, ganz zu schweigen von den Trichoceridae,

durch einen ganzen Komplex von Merkmalen: die Morphologie aller ihrer

Entwicklungsstadien, der Bau des männlichen Hypopygs, ihre Anforderungen an

die Umwelt, ihre trophischen Bindungen und andere, durch die sie als eine

systematische Einheit mit spezifischen, nur für sie kennzeichnenden Evolutions-

Tendenzen charakterisiert sind. Es muß hinzugefügt werden, daß die Tipulidae

sich als unabhängige Familie aus den schon längst ausgestorbenen Architipulidae

entwickelt haben, und daß Limoniidae und Cylindrotomidae sich aus einer

Schwestergruppe nicht der heutigen Tipulidae, sondern der Architipulidae

gebildet haben. Aus all diesen Gründen bleibt der Autor dieser Arbeit dabei —

wie in seinen anderen Arbeiten —, daß die Tipulidae eine unabhängige,

morphologisch, biologisch und evolutionell deutlich abgegrenzte Familie sind. Es

ist interessant zu bemerken, daß nach Hennig (1950) die Tipulidae als eine sehr

früh differenzierte Gruppe unter den anderen Familien der Diptera den Rang

einer Superfamilie verdienen.

Während des neunzehnten Jahrhunderts haben mehrere Systematiker die von

Linnaeus (1758) aufgestellte Gattung Tipula, die nicht nur Tipulidae sondern auch

andere Tipuloidea umfaßte, in kleinere natürlichere Gattungen aufgeteilt.

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

Der Klassiker der Dipterologie Meigen (1800) hat als erster von Tipula die Gat-

tungen Pales (Nephrotoma, 1803) und Flabellifera (Ctenophora, 1803) abgetrennt,

deren erste später von Macquart (1834) als Pachyrrhina zum zweiten Mal

beschrieben wurde. Durch eine Entscheidung der 1.C.Z.N. (1963) sind die Namen

Pales und Flabellifera unterdrückt worden.

Curtis (1825) trennte Dolichopeza, eine Gattung, die von Meigen (1830) als

Leptina und von Macquart (1846) als Apeilesis beschrieben wurde, von Tipula ab.

Loew (1844) beschrieb die Gattung Prionocera (Stygoropis Loew, 1863). Latreille

(1804) stellte einen Teil der Arten aus Meigens Gattung Ctenophora in Tanyptera,

eine Gattung, die 30 Jahre später von Brull& (1832) aufs neue als Xiphura

beschrieben wurde. Für eine der Arten aus der Gattung Ctenophora errichtete

Brulle (1833) die Gattung Dictenidia, die in Arbeiten späterer Autoren auch als

Ceroctenia (Rondani, 1856) und Dicera (Lioy, 1863) aufgeführt wird.

Von mehreren Autoren (Wiedemann, 1828; Macquart, 1838; Loew, 1869;

Osten-Sacken, 1869, 1886; Westwood, 1876; Skuse, 1890) wurden auch Gattungen

aus den Gebieten außerhalb der Palaearktis beschrieben. Infolge dieser schnelle

Erhöhung der Anzahl von Gattungen entstand das Bedürfnis nach höheren

Kategorien.

Als erster hat wahrscheinlich wohl Kertesz (1902) die Tipulidae in seinem

Catalogus in drei Unterfamilien aufgeteilt: Dolichopezinae, Ctenophorinae und

Tipulinae. Von unserer Fauna rechnet er Dolichopeza zur ersten Unterfamilie,

Dictenidia, Ctenophora und Xiphura (= Tanyptera) zur zweiten und Tipula (inkl.

Nigrotipula) und Nephrotoma zur dritten. Diese Einteilung, die auf unterschiedliche

Flügeladerung (Dolichopezinae und Tipulinae) und verschiedenen Bau der

Antennen bei den Männchen (Tipulinae und Ctenophorinae) begründet ist, wurde

von den meisten Systematikern übernommen, insbesondere von Enderlein (1936),

Hendel (1936/37) und Mannheims (1951, 1954).

In vielen seiner Arbeiten benützt auch Alexander diese Einteilung. Bei Ale-

xander und anderen Systematikern, die die Tipulidae als Unterfamilie

qualifizieren, werden dementsprechend die Unterfamilien von Kertesz als Tribus

aufgeführt.

Eine etwas andere Einteilung der Familie Tipulidae wurde von Pierre (1926)

aufgestellt. Aus nur formellen Gründen ohne Rücksicht auf phylogenetische

Aspekte, teilte er die Familie ein in zwei Hauptgruppen oder Divisionen, die sich

nur im Bau der Antennen voneinander unterscheiden. Die erste Division —

Tipulinae filicornae — umfaßt die größte Zahl der Gattungen dieser Familie. Sie

haben die Antennen nicht kammartig gebaut. Die zweite, kleinere Division —

Tipulinae ramicornae — hat, wenigstens bei den Männchen, charakteristische

kammförmige Antennen.

Masaki (1933) folgt Pierre und unterscheidet ebenso nur zwei Subfamilien:

Plusiomyini — Gattungen mit kammförmigen Antennen — und Tipulini — alle

anderen Gattungen. (Bemerkung: Plusiomyini nach der tropischen Gattung Plu-

siomyia Skuse.)

Es ist eindeutig klar, daß diese Einteilungen von Pierre und Masaki im Vergleich

mit der Einteilung in drei Subfamilien einen Schritt zurück bedeuten, weil nur auf

SAVTSHENKO: Phylogenie und Systematik der Tipuladae 119

den Bau der Antennen, nicht aber auf andere morphologische, biologische oder

geografische Merkmale geachtet wird. Diese Einteilung ist daher sehr künstlich.

Nach Masaki sind zum Beispiel die palaearktischen Ctenophorinae, von denen

nur die Männchen kammförmige Antennen haben, in einer und derselben Gruppe

zusammen mit einer Reihe von tropischen Arten, deren Männchen und Weibchen

kammförmige Antennen haben, aber nach dem Komplex aller anderen Merkmale

viel näher den Tipulinae oder Dolichopezinae stehen.

Auch Lameere’s (1906) Einteilung der Tipulidae in zwei Tribus ist mehr oder

weniger künstlich. Lameere unterscheidet die Dolichopezinae und die Tipulinae.

Charakteristisch für die Dolichopezinae ist das Fehlen von Tibialdornen und auch

das Fehlen einer Diskoidalzelle. Die Gattung Dolichopeza ausgenommen, haben

aber die meisten anderen Dolichopezinae eine Diskoidalzelle wie die Tipulinae.

Auf die Tatsache hinweisend, daß die Einteilung der Familie in drei Unter-

familien auf — auf den ersten Blick — sekundären und taxonomisch wenig

wichtigen morphologischen Unterschieden fußt, welche überdies bei den

Ctenophorinae sekundäres Geschlechtsmerkmal sind, erklärt sich Alexander

(1920, 1942) als Vertreter einer vollständigen Zurückweisung der Unterfamilien

und will zurück zur alten Einteilung in nur Gattungen. Seiner Meinung nach ist

dies zweckmäßiger, weil — mit weiter Kenntnis der exotischen Tipulidae — die

Grenzen zwischen den Subfamilien immer undeutlicher werden. Dasselbe wird

auch von Theowald (1957) vorgebracht, der ein neues System für die

westpalaearktischen Tipuliden aufstellt, begründet auf dem Studium von

imaginalen und praeimaginalen Merkmalen, wobei er in dieser Familie keine Taxa

von höherem als Gattungsrang aufführt.

In dieser Arbeit wird aber die allgemein akzeptierte Einteilung der Familie in

drei Unterfamilien erhalten und zwar nicht nur, weil sie das System der Tipulidae

mehr übersichtlich macht, was aus praktischen Erwägungen sehr wichtig ist, da sie

die Bestimmung erleichtert, sondern auch, weil sie vom phylogenetischen Stand-

punkt (Abb. 4) die realen Verhältnisse innerhalb der Familie erkennen läßt. Sie

zeigt unabhängige Gruppen von ungleichem Umfang, ungleicher morpholo-

gischer Differenzierung, ungleichem geologischem Alter und ungleichem zoo-

geographischem Wert. Im Gegensatz zu der Behauptung von Alexander gibt es

zwischen diesen drei Unterfamilien der Tipulidae — nach dem ganzen Komplex

von imaginalen und praeimaginalen Merkmalen — genügend und deutliche

morphologische Unterschiede, welche sich manifestieren in der Färbung und

Zeichnung des Körpers, in der Beinlänge, und insbesondere im Bau des Hypopygs

der Männchen wie auch im Bau des Hinterleibsendes der Larven. Schließlich,

jede dieser Unterfamilien zeigt eine bestimmte Richtung der Evolution innerhalb

dieser Familie, nämlich die Dolichopezinae hauptsächlich den Übergang einer

hydrobionten zu einer bryobionten Lebensweise auf dem Land, die Tipulinae (mit

nur wenigen Ausnahmen) den Übergang einer hydrobionten zur helobionten und

geobionten Lebensweise, die Ctenophorinae den Übergang einer hydrobionten

zur saproxylobionten Lebensweise. Rohdendorf (1964) bemerkt ganz richtig, daß

Taxa von höherem Rang nicht nur vom morphologischen Standpunkt, sondern

vielseitig, als eine Einheit oder eine Harmonie, gesehen werden müssen, die sich

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

durch bestimmte Existenzvoraussetzungen, die sich in gleichem Bau, gleichen

Funktionen und gleicher ontogenetischer Entwicklung zeigen, kennzeichnet.

Wenn man bei den Tipulidae — einer der meist archaischen Familien der

Diptera — dieselben taxonomischen Kriterien anlegt wie bei den höher ent-

wickelten Gruppen dieser Ordnung, dann kann auch das Fehlen von deutlichen

Grenzen zwischen manchen Unterfamilien nicht ein Grund zu ihrer Abschaffung

sein. Das Vorkommen von Übergangsformen zwischen einer Unterfamilie und der

anderen in solch einer archaischen Gruppe von Diptera wie die Tipulidae — in der

bis in unsere Zeit viele Elemente aus dem Palaeogen und sogar aus dem

Mesozoikum vorkommen — ist ganz natürlich. In diesem Zusammenhang

bemerken wir, daß ähnliche “Brücken” auch zwischen den Tipulidae und andere

Familie der Tipuloidea vorkommen, zum Beispiel zwischen Dolichopezinae und

Limoniidae, anderseits auch zwischen Gattungen innerhalb der Tipulidae, zum

Beispiel Nephrotoma und Tipula, zwischen den orientalischen Arten von Doli-

chopeza und Oropeza und auch zwischen den orientalischen Pselliophora’s und den

palaearktischen Ctenophora’s.

Meist werden im System der Tipulidae als erste Subfamilie die Dolichopezinae,

als zweite die Ctenophorinae und als dritte die Tipulinae aufgeführt, was dem

Standpunkt von Edwards (1926) entspricht, nach dem die xylobionten Arten

primitiver sind als die hydro- und helobionten. Dies steht aber im Widerspruch zu

den eigentlichen phylogenetischen Verhältnissen in der Familie der Tipulidae, wie

oben schon erwähnt wurde. Die Tipulinae sind mit den am meisten plesiomorphen

Dolichopezinae deutlich näher verwandt als mit den Ctenophorinae, die als

saproxylophage Arten zu der am meisten apomorphen und spezialisiertesten

Gruppe dieser Familie gehören. Deshalb müssen die Tipulinae im System der

Familie an zweiter und nicht an dritter Stelle stehen, was mit den interfamiliären

phylogenetischen Verhältnissen (Abb. 4) übereinstimmt.

Die Behandlung der Systematik innerhalb der Unterfamilie der Dolichopezinae

gehört nicht in diese Arbeit, da in der ukrainischen Fauna diese Unterfamilie nur

durch eine Gattung vertreten ist. Was die Unterfamilie Tipulinae anlangt, so hat

man bis heute nicht versucht, sie in Supergattungen oder Tribus einzuteilen. Es

besteht aber kein Zweifel, daß die ukrainischen Tipulinae sich morphologisch und

phylogenetisch leicht in zwei unabhängige Gruppen einteilen lassen, die zwei

verschiedene Evolutionsrichtungen darstellen. Die erste Gruppe sind die Arten

mit gesägten Antennen aus der Gattung Prionocera, die mit der Unterfamilie der

Dolichopezinae nahe verwandt sind. Die zweite Gruppe sind alle anderen Ti-

pulinae, die von denselben ancestralen Formen entstanden sind wie Prionocera,

sich aber parallel zu ihnen entwickelten. Nach dem phylogenetischen Schema

(Abb. 4) ist es zweckmäßig, beide Gruppen als unabhängige Stämme zu sehen: die

Prionocerini und die Tipulini. Der erste von ihnen ist in unserer Fauna vertreten

durch die Gattung Prionocera, der zweite durch Nigrotipula, Tipula und

Nephrotoma.

Auch die Unterfamilie Ctenophorinae kann in zwei Tribus eingeteilt werden:

Ctenophorini und Tanypterini, die deutlich in morphologischer und biologischer

Hinsicht verschieden sind. Die Ctenophorini sind als Imagines durch einen ziem-

SAVTSHENKO: Phylogenie und Systematik der Tipuladae 121

lich einfachen Bau des Hypopygs und eine fast vertikale Stellung der Styli, durch

einen kurzen Ovipositor, der immer kürzer ist als die Hälfte des Abdomens, und

durch kurze Hypovalven, die bei weitem nicht das Ende der Cerci erreichen,

gekennzeichnet. Die Larven der Ctenophorini haben mehr oder weniger

entwickelte Randlappen um das Stigmenfeld und manchmal einen rudimentären

Haarsaum am Rande des Stigmenfeldes. Die Puppen haben am Mesothorax

ziemlich lange Atmungshörnchen, die von normaler Form sind. Die Entwicklung

der Larven dieser Familie findet in Holz, das zu amorphem Mulm verfault ist,

statt. Für die Tribus Tanypterini ist nachfolgendes charakteristisch: das Hypopyg

der Männchen ist stark modifiziert, und die Stellung der Styli ist fast horizontal;

die Weibchen haben einen sehr langen Ovipositor (fast solang wie die Hälfte des

Abdomens, die langen Hypovalven erreichen fast das Ende der Cerci). Die

Randlappen um das Stigmenfeld der Larven sind fast vollständig reduziert; die

Puppe hat nur kurze, längliche, am Rande gerillte mesothorakale

Atmungshörnchen. Die Arten entwickeln sich in Holz, das noch nicht ganz zu

Mulm verfault ist. In der ukrainischen Fauna gehören die Gattungen Dictenidia

und Ctenophora zu den Ctenophorini und Tanyptera zu den Tanypterini.

Alexander (1936, 1954) und nach ihm Theowald (1957) sind der Meinung, daß

alle Gattungen der Ctenophorinae, die orientalischen Gattungen Plocimas und

Prionota ausgenommen, nur den Rang von Untergattungen haben sollen, weil

zwischen ihnen keine wirklichen Unterschiede sind, abgesehen vom Bau der

Antennen bei den Männchen, der zu den sekundären Geschlechtsmerkmalen

gehört. Die Gattungen Dictenidia, Ctenophora und Tanyptera werden deshalb von

ihnen als Untergattungen aufgeführt. Was die Gattung Tanyptera betrifft, soll diese

auf Grund von oben aufgeführten Unterschieden nicht nur Gattungsrang bei-

behalten sondern sogar als Tribus aufgeführt werden müssen. Solche wirklichen

Unterschiede gibt es auch zwischen den Gattungen Dictenidia und Ctenophora.

Erstens sind nicht nur die Antennen der Männchen, sondern auch die Antennen

der Weibchen auf andere Weise gebaut, weshalb es sich nicht nur um ein

sekundäres Geschlechtsmerkmal handeln kann. Überdies:

Dictenidia Ctenophora

Brustseiten immer nackt immer behaart

Flügel der & & oft mit Mikro- fast immer nackt

i trichien

9. Sternit und id einfach gebaut mehr oder weniger

kompliziert

Ovipositor der 9 © gerade etwas gebogen

Stigmenfeld der Larve ohne Haarsaum mit Haarsaum

Mesothorakale Atmungs- an der Vordersei- beiderseits des

hornchen der Puppen te des Körpers Körpers

‚Verbreitung nur palaearktisch holarktisch-orientalisch

Deshalb müssen Dictenidia und Ctenophora nicht nur als Untergattungen,

sondern als morphologisch und geographisch ziemlich stark voneinander

abgegrenzte Gattungen angesehen werden, was übereinstimmt mit den Prinzipien

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 5, 1979

der Einteilung der Tipulidae in infrafamiliare Taxa, welche Prinzipien in dieser

Arbeit beibehalten werden.

Besondere Aufmerksamkeit erfordert die Problematik der polymorphen Gat-

tung Tipula, zu der etwa 57% der Arten der Weltfauna und 71% der ukrainischen

Arten dieser Familie gehören. Schon Schummel (1833) — die Flügelzeichnung als

Basis nehmend — teilte die Gattung Tipula in eine Reihe von Artengruppen auf.

Von den späteren Bearbeitern hat Riedel (1913) diese Einteilung trotz ihrer

Künstlichkeit übernommen. Erst Edwards (1931) hat — nachdem er die Gattung

Tipula in Untergattungen gruppiert hatte — die taxonomischen Grenzen an Hand

von Komplexen imaginaler Merkmale wissenschaftlich motiviert. Einige richtig

motivierte Untergattungen wurden auch von Bezzi (1924) und Alexander (1924)

aufgestellt. Bis in letzte Zeit aber war die Systematik der Gattung Tipula sehr

dürftig, weil manche ihrer Untergattungen, wie auch zum Großteil die Gruppen

von Schummel und Riedel, morphologisch, ökologisch und phylogenetisch sehr

ungleiche und ungleichwertige Elemente vereinigten. Besonders die Untergattung

Oreomyza s.l. war sehr heterogen und enthielt nach Alexander (1935, 1942) und

Wu (1940) eigentlich eine zufällige Zusammenstellung von oft phylogenetisch weit

voneinander entfernten Arten, deren systematische Stellung in der Gattung aus

irgendeinem Grund unsicher war.

Auf Grund von vergleichend morphologischen Untersuchungen der imaginalen

und larvalen Stadien und auch in Anbetracht von Besonderheiten der Ökologie

und der Fortpflanzung (Hemmingsen, 1954— 1962) hat der Autor einige neue Un-

tergattungen unterschieden, in denen die /uteipennis—autumnalis-Gruppe,

rufina—obsoleta—signata-Gruppe, unca-Gruppe, variipennis—irrorata-Gruppe,

Juncea-Gruppe und flavolineata-Gruppe untergebracht sind, und den Umfang

einiger anderer Untergattungen näher präzisiert; diese letzten Untergattungen

waren schon von Edwards für die oleracea—paludosa-Gruppe, die variicornis-

Gruppe und die /ateralis—pruinosa-Gruppe aufgestellt worden (Savtshenko, 1961,

1964). Neue Untergattungen für die bidens—stigmatella-Gruppe und die bistilata-

Gruppe wurden in der letzten Zeit motiviert von Mannheims (Mannheims &

Pechlaner, 1963) aufgestellt, der auch die Richtigkeit der Namen mancher schon

früher beschriebener Untergattungen überprüfte. Alexander (1965) stellte eine

Untergattung auf für einige asiatische Arten, zu denen auch die europäische

saginata Bergroth paßt.

Der neuen Klassifikation entsprechend ist die Gattung Tipula in der ukra-

inischen Fauna vertreten durch 16 Untergattungen: Platytipula Matsumura,

Schummelia Edwards, Savtshenkia Mannheims, Yamatotipula Matsumura, Acutipula

Alexander, Tipula s.str., Mediotipula Pierre, Beringotipula Savtshenko, Pterelachisus

Rondani, Oreomyza Pokorny, Lindnerina Mannheims, Vestiplex Bezzi, Lunatipula

Edwards, Emodotipula Alexander, Odonatisca Savtshenko und Dendrotipula

Savtshenko.

Was die umfangreiche Gattung Nephrotoma betrifft, gibt es noch keine

motivierten Gründe zum Aufstellen von Untergattungen, weil jene Unterschiede

zwischen den Artengruppen, die bei den Imagines gefunden wurden, in den

praeimaginalen Stadien nicht bestehen oder noch nicht gefunden wurden.

SAVTSHENKO: Phylogenie und Systematik der Tipuladae 123

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THE NEW WORLD GENUS PLUTO (HYMENOPTERA,

SPHECIDAE, PSENINI)

J.P. VAN LITH +

Allard Piersonstraat 28c, Rotterdam

With 71 text-figures

ABSTRACT

A review of the genus Pluto Pate with key to the species is presented. Redescriptions of the forms

previously published, first descriptions of some opposite sexes and new distribution data are given. The

following new taxa are described: abbreviatus, alphitopus, araguensis, arenivagus cubanus, basifuscus,

biformis, castaneipes, colonensis, denticollis, depressus, duckei, emarginatus, evansi, facialis, fritzi, incarina-

tus, jugularis, marthae, medius zuliensis, menkei, metanus, nitens, obscurus, occipitalis, punctatellus, pyg-

maeus axillaris, rotundus, rufanalis, rugulosus, scytinus, simplicicollis, spangleri, spinicollis, stenopygidialis,

stramineipes, strigellus, trilobatus, zonatus.

The first review of the genus Pluto Pate (Psenia Malloch) was given by Malloch

(1933) in his study on the Psenini of North America. In 1901, Viereck included the

two North American species known to him, Mimesa tibialis Cresson and Psen

suffusus Fox in his new genus Neofoxia (type species Psen atratus Panzer) together

with two species now placed in the genus Psenulus, i.e. Psen frontalis Fox and Psen

trisulcus Fox. Malloch (1933) recognized two separate genera, Diodontus Curtis,

1834 (Neofoxia Viereck, now Psenulus Kohl, 1896) and a new genus Psenia, the

latter to receive those forms in which “the cubitus of the hind wing is distad of the

median transverse vein” (cu-a) as in Diodontus Curtis (Psenulus Kohl) but the

“occipital carina is not connected with the carina surrounding the mouth cavity”

(hypostomal carina).

Malloch in his diagnosis of Psenia rightly mentions the long, downward directed

bristly hair on the mid and hind coxa, but some of the other generic characters

enumerated by him apply to North American species only. He described or

recorded sixteen taxa, including atricornis from the West Indies, but not

argentifrons (Cresson) from Cuba. Pate (1937) changed the generic name into

Pluto, Psenia being preoccupied by Psenia Stephens, 1829 (a synonym of Psen

Latreille, 1796; type species Sphex atra Fabricius, 1793). In 1946, Pate furnished a

good redescription of Pluto argentifrons (Cresson) and pointed out that the females

from Cuba, which Malloch associated with the males of atricornis Malloch from

Puerto Rico, in fact belong to argentifrons. A few years later Krombein (1949)

described arenivagus as a new species from North America.

Thus far Mexico and South America had yielded very few species. Cameron

(1891) published Psen annulipes (female) from Mexico, Psen medius was described

127

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

by Smith (1856) and Psen Smithii by Fox (1898), both after males collected in

Brazil, and Psenulus (Neofoxia) Townsendi by Cockerell (1911) from Peru. Bohart &

Menke (1976) assigned these four species to the genus Pluto. To Brethes (1913) we

owe the descriptions of two species from Argentina, namely Psen Jörgenseni (male)

and the female of Gorytes pygmaeus. We should be grateful to Mr. M. A. Fritz,

Buenos Aires, who recognized this latter specimen as a Psenine wasp. He and Dr.

M. J. Viana of the Museum at Buenos Aires kindly enabled me to study the types

of both species.

Up till now little attention was paid to South American Pluto. It is therefore with

great pleasure that I express my gratitude to the many entomologists who allowed

me to study their Psenini. Over 1900 specimens of Pluto could be examined

including many from the Neotropical Region. This study resulted in the

description of 22 new species and two new subspecies from South America, 13 new

species from Central America, Mexico and the southern states of the United

States and one new subspecies from Cuba.

Most of the known species are redescribed and first descriptions of some

opposite sexes are also given. The genus Pluto now includes 59 species and three

subspecies.

The institutions and entomologists who entrusted me their material are

mentioned below, preceded by the abbreviations used for their collections:

AMNH — American Museum of Natural History, New York, N.Y., U.S.A.; J. G.

Rozen, Jr. and Mrs. M. Favreau

BISH — Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.; Miss A.

Manning, F. J. Radovsky

BM — British Museum (Natural History), Department of Entomology,

London, U.K.; L. A. Mound, C. R. Vardy

BSM — Zoologische Sammlung des Bayerischen Staates, Munich, Germany;

E. Diller

CAS — California Academy of Sciences, San Francisco, California, U.S.A.; P.

H. Arnaud, Jr., Mrs. Helen Court

CIS — California Insect Survey Collection, University of California,

Berkeley, California, U.S.A.; J. A. Powell

CNC — Canada National Collection, Entomology Research Institute, Ottawa,

Canada; J. Barron, L. Masner, C. M. Yoshimoto

CSC — California State Collection of Arthropods, Sacramento, California,

U.S.A.; M.S. Wasbauer

CU — Cornell University, Department of Entomology and Limnology,

Ithaca, N.Y., U.S.A.; L. A. Pechuman, A. C. Miller

FAG — Faculté des Sciences Agronomiques, Gembloux, Belgium; J. Leclercq

FSC — Florida State Collection of Arthropods, Gainesville, Florida, U.S.A.;

E. E. Grissell

HT — H. and M. Townes collection, American Entomological Institute, Ann

Arbor, Michigan, U.S.A.

IML — Instituto Miguel Lillo, Tucumän, Argentina; J. A. Haedo, A. Willink

VAN LITH: New World Pluto 129

KU — Snow Entomological Museum, University of Kansas, Lawrence,

Kansas, U.S.A.; C. D. Michener, G. W. Byers

KVK — Collection K. V. Krombein, now donated to the Smithsonian

Institution, Washington, D.C.

MACN — Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”,

Buenos Aires, Argentina; M. J. Viana, M. A. Fritz

MCZ — Museum of Comparative Zoology, Harvard College, Cambridge,

Massachusetts, U.S.A.; Ms. J.C. White, Ms. M. K. Thayer

MF — M.A. Fritz collection, Buenos Aires, Argentina

ML — Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands; J.

van der Vecht, C. van Achterberg, C. van Heijningen

NMW — Naturhistorisches Museum Wien, Vienna, Austria; M. Fischer

NCSU — North Carolina State University, Raleigh, N.C., U.S.A.; D. L. Stephan

OSU — Oregon State University, Entomological Museum, Corvallis, Oregon,

U.S.A.; G. R. Ferguson (including G. R. Ferguson collection)

RS — Naturhistoriska Riksmuseet, Stockholm, Sweden; S. Erlandsson

UCD — University of California, Department of Entomology, Davis, Califor-

nia, U.S.A.; R. M. Bohart

UFP — Universidade Federal do Parana, Departamento de Zoologia,

Curitiba, Parana, Brazil; J.S. Moure

USNM — National Museum of Natural History, Smithsonian Institution, Wash-

ington, D.C., U.S.A.; K. V. Krombein, A. S. Menke

UZM — Universidad del Zulia, Maracaibo, Venezuela; via A. S. Menke

ZMB — Zoologisches Museum der Humboldt Universität, Berlin, Germany; E.

Konigsmann

ZMC — Universitetets Zoologiske Museum, Copenhagen, Denmark; O.

Lomholdt

I am grateful to Mrs. Drs. Carol van Driel-Murray, Leiderdorp, for her

willingness to read and correct the English text of the introductory parts of this

article.

Pluto Pate, 1937

Psenia Malloch, 1933, type species Mimesa tibialis Cresson, 1872, original designation, nec Psenia Ste-

phens, 1829. i

Pluto Pate, 1937, new name for Psenia Malloch; Krombein, 1951; Gittins, 1969.

A good generic diagnosis has been given by Bohart and Menke (1976). Bohart

also refers to the distinctive downward directed long bristle on the lower hind

margin of the hind coxae; this bristle is most conspicuous in the female and Bohart

calls it the most diagnostic and unique feature of Pluto. The bristle on the mid coxa

is shorter. The clypeal margin is indeed generally relatively simple but the female

of the Mexican clavicornis (Malloch) has a peculiar snoutlike protruding clypeus

and the females of at least a few South A merican species (P. joergenseni (Brèthes),

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

P. facialis sp. nov.) have either strongly protruding lateral clypeal teeth or an

unusually deeply emarginate clypeal apex. The hypo-epimeral area, as well as the

mesopleura and the scutum may be coarsely rugose or smooth and shining.

The tyloidea, usually present on most of the antennal segments of the male, may

offer good differential characters. They are mostly long and linear, sometimes

more or less oval, rarely partly absent on some of the median segments or

indistinct on the whole flagellum. In P. clavicornis the antennae are clavate like the

antennae of a female and there are no distinct tyloidea.

Although the genitalia of the male have been insufficiently studied, the drawings

in this paper of the genitalia of a few species show that they are very uniform and I

do not expect that they will be of much use to facilitate identification.

The pygidial area of the female shows very little differentiation; its structure

leads to the conclusion that all Pluto species nest in the soil. Very little is known

with certainty about their biology. Most of the notes on the labels relate to the

flowers visited. Evans (1959) described the larva of P. albifacies (Malloch) and in

1968 the nests as well as the prey, which consisted of nymphs (%) and adults of the

small green leafhopper Opsius stactogalus Fieber. A few females of P. littoralis

(Malloch) and a female of P. punctatellus sp. nov. have been collected together

with their prey, which also consisted of small Jassids.

In the descriptions the term “‘intercarinal space” indicates the distance between

the ventral part of the occipital carina and the posterior part of the hypostomal

carina. In some species this space exceeds the width of the basitarsus of the fore

legs; usually, however, it is much narrower, rarely the carinae are almost touching.

In one or two species the occipital carina does not reach the midventral line, or

this carina, at least the lower part of it, is unusually high. The ratio between OOD

(distance between posterior ocelli and the oculi) and POD (distance between

posterior ocelli) has been roughly indicated. The tempora are usually finely striate,

sometimes very finely so. The term “pronotal collar” has been used for the

pubescent dorsal part of the pronotum behind the transverse carina. The anterior

scutellar suture is usually crenulate, rarely somewhat indistinctly so; in the group

of P. pygmaeus (pygmaeus (Brethes), pygmaeus axillaris subsp. nov. and facialis sp.

nov.) this suture is simple. The latero-dorsal rows of very short hairs on the petiole

are sometimes indistinct, laterally and ventrally the petiole has a number of long

erect hairs; the hind margins of the sternites show some long backward directed

bristles.

For the figures of the clypeal margin the position of the axis of the microscope

(x 30) was perpendicular to the surface of the clypeal disk. As the clypeus is more

or less convex, the apical margin may — when seen in ventral aspect — seem to be

-more emarginate than when seen in frontal aspect and the lateral corners or teeth

of the projecting median part may seem to be sharper.

RELATIONSHIPS

An attempt has been made here to split up the genus into species-groups. Many

of the species have a very uniform appearance and sometimes closely related

VAN LITH: New World Pluto 131

forms are not easily distinguished. A few groups, however, show distinct special-

ization. In the South American group of pygmaeus the axillae are not connected

with the disk of the scutellum and they project freely backwards. In P. joergenseni

the metanotum is peculiarly raised in the middle, especially in the male; the

anterior margin of the labrum of the female differs from that of other species. As

some species exhibit considerable sexual dimorphism and in many cases the

opposite sex is unknown, the arrangement of the species as given below can be no

more than a tentative one. Moreover, our knowledge of the geographic

distribution of some forms is very incomplete and any decision concerning their

status as a distinct species or geographic subspecies may be difficult.

In the enumeration of the species belonging to each group the numbers of

specimens examined for this study are also mentioned.

Group of longiventris

Characterized in both sexes by the coarsely rugose mesopleura, the female

moreover by the lateral pronotal lobes and the male by its large oval tyloidea.

longiventris (Malloch) — 19 9 and 65 Z; Arizona, California; Mexico

Group of angulicornis

Antennal tyloidea of males, in as far as they are known, short and mostly oval, in

P. angulicornis more or less angular in lateral view. In P. suffusus, which may belong

to a separate group, the tyloidea are small and oval. Clypeal margin of female

straight or rounded. Propodeum of female usually finely reticulato-carinate.

Mesopleura in both sexes usually reticulate alutaceous and punctate. Petiole of

female usually about half as long as first tergite, in a few species nearly as long as

this tergite. Intercarinal space of P. suffusus broad. Perhaps P. abbreviatus also

belongs to this group; the males of this species have short, somewhat linear

tyloidea.

angulicornis (Malloch) — 59 and 5 Z; Texas, Iowa, Louisiana, New

Mexico

pallidistigma (Malloch) — 29 and8 g; Arizona, California, Texas

basifuscus sp. nov. — 16 9 and 26 G'; Arizona, California, Texas,

New Mexico; Mexico

minutus (Malloch) — 29, d unknown; Texas

spangleri sp. nov. — 42 Q and 48 Z; Arizona, Texas; Mexico;

Guatemala

brevipetiolatus (Rohwer) — 29, unknown; California

rotundus sp: nov. — 19, unknown; Florida

suffusus (Fox) — 41 9 and 82 g; Arizona, California,

Florida, New Mexico, South Carolina,

Texas; Mexico

biformis sp. nov. — l g, 9 unknown; Mexico

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Group of texanus

Tyloidea indistinct or absent on antennal segments 8—10 of male. Clypeal

margin of female weakly bisinuate. Mesopleura reticulate alutaceous, punctate.

Hypo-epimeral area striate. P. littoralis seems to be the eastern relative of the

Texan species.

texanus (Malloch) — 29 and2 9; Texas

littoralis (Malloch) — 17 9 and 23 g; Florida, Maryland, North

Carolina, South Carolina

Group of sayi

Tyloidea of males, as far as known, linear. Clypeal margin of female weakly

emarginate or bisinuate. Mesopleura reticulate alutaceous and punctate, rarely

shining. Hypo-epimeral area finely punctate, often somewhat shining. Back of

propodeum, also in the males of some species (sayi, abbreviatus, depressus) finely -

reticulate. P. abbreviatus may have to be transferred to the group of angulicornis, as

its tyloidea are more elongate oval than linear.

sayi (Rohwer) — 87 © and 291 g; Alabama, Arizona,

California, District of Columbia, Florida,

Iowa, Kansas, Louisiana, Missouri, New

Mexico, South Carolina, Tennessee, Tex-

as, Virginia; Mexico; Cuba; El Salvador;

Nicaragua

stenopygidialis sp. nov. — 29, unknown; Arizona

Jugularis sp. nov. — 19, unknown; Brazil

punctatellus sp. nov. — 12 @ and 29 Z; Mexico; Costa Rica; El

Salvador; Guatemala

depressus sp. nov. — 6, 2 unknown; Mexico

abbreviatus sp. nov. — 39 and2 Z; Mexico

Group of aerofacies

Tyloidea of male linear. Clypeal margin of female slightly rounded or weakly

emarginate. Back of propodeum coarsely reticulato-carinate in both sexes.

Mesopleura reticulate alutaceous.

aerofacies (Malloch) — 6 9 and9 g; Texas; Mexico; Belize

evans! sp. nov. — 39 and l 3; Mexico

emarginatus sp. nov. — 19, & unknown; Mexico

Group of rufibasis

Tyloidea of male linear. Clypeal margin of female somewhat rounded. Scutum

and mesopleura very coarsely rugose and shining in both sexes.

VAN LITH: New World Pluto 133

rufibasis (Malloch) — 21 9 and 41 &; Florida, Georgia, Louisi-

ana, Maryland, Mississippi, North Caro-

lina, South Carolina, Virginia

Group of albifacies

Tyloidea of male linear. Clypeal margin of female somewhat rounded.

Mesopleura dull, moderately densely punctate. Hypo-epimeral area dull, densely

finely punctate.

albifacies (Malloch) — 149 and4 4; Iowa, Texas

Group of tibialis

Tyloidea of male linear, dark. Clypeal margin of female almost straight.

Mesopleura reticulate alutaceous, moderately densely punctate, of male of

arenivagus sometimes densely punctate. Hypo-epimeral area striate.

tibialis (Cresson) — 18 9 and 32 Z; Alabama, Florida, District

of Columbia, Louisiana, Missouri, South

Carolina, Tennessee, Texas, Virginia

arenivagus arenivagus Krombein — ll © and 6 g; Florida, Georgia, North

Carolina

arenivagus cubanus subsp. nov. — 1 9, unknown; Cuba

Group of argentifrons

Tyloidea of male linear. Clypeal margin of female weakly emarginate or

bisinuate. Mesopleura dull, usually densely or very densely finely punctate,

especially so in the males.

argentifrons (Cresson) — 24 © and 57 g; Cuba; Jamaica; Mexico;

Nicaragua

atricornis (Malloch) — 4 Q and 12 Z; Puerto Rico; Dominican

Republic; Virgin Islands; Leeward Islands

alphitopus sp. nov. — 19 and 5 4; Mexico

castaneipes sp. nov. — 2, 9 unknown; New Mexico, Texas

rugulosus sp. nov. — lg, 9 unknown; Texas

colonensis sp. nov. — 19, g unknown; Argentina

fritzi sp. nov. — 19 and 3 g; Ecuador

medius medius (F. Smith) — 5 9 and 17 Z; Argentina; Bolivia; Brazil;

Surinam

medius zuliensis subsp. nov. — 39 © and 2 Z; Venezuela; Curacao

scytinus sp. nov. — 19, unknown; Venezuela

stramineipes sp. nov. — 10 9 and 20 Z; Argentina; Bolivia; Brazil

strigellus sp. nov. — 29,3 unknown; Argentina

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Group of clavicornis

Antennae of male clavate, no distinct tyloidea. Clypeus of female strongly

protruding, snoutlike. Mesopleura very finely reticulate alutaceous, almost

shining, no distinct punctures.

clavicornis (Malloch) — 6 © and 141 3; Arizona; Mexico

Group of townsendi

Antennae of male relatively long and slender, tyloidea not always distinct. :

Clypeal margin of female weakly emarginate. Mesopleura somewhat dull,

indistinctly punctate. All species belonging to this group are South American.

townsendi (Cockerell) — 13 9 and 25 g; Peru; Ecuador

marthae sp. nov. — 8 Q and Il Z; Bolivia; Brazil; Colombia;

Ecuador; Peru

metanus sp. nov. — 39 and2 4; Colombia

Group of nitens

Tyloidea of male linear or absent, but most of the males are still unknown.

Thorax shining, smooth or very finely sparsely punctate, mesopleura rarely very

finely alutaceous. The females of P. occipitalis and P. spinicollis have sharp lateral

pronotal angles. The female of P. trilobatus is distinguished from other species by

the distinctly tridentate clypeal margin. The males of P. rufanalis, P. araguensis and

P. incarinatus have indistinct or no tyloidea. All species are South American.

nitens sp. nov. — 30 © and 30 Z; Argentina; Bolivia; Brazil;

Colombia; Ecuador; Paraguay; Peru;

Surinam; Venezuela

duckei sp. nov. — 29, unknown; Brazil

obscurus sp. nov. — 1 9, unknown; Argentina

zonatus sp. nov. — 2 9, unknown; Brazil

simplicicollis sp. nov. — 19, g unknown; Brazil

menkei sp. nov. —

occipitalis Sp. nov. —

spinicollis sp. nov. —

1 ©, & unknown; Venezuela

trilobatus sp. nov. — 6

©, g unknown; Peru

©, d unknown; Brazil; Panama

rufanalis sp. nov. — 5 © and 28 4; Peru

araguensis SP. NOV. — 39

incarinatus sp. nov. _

and 3 g; Venezuela

Group of annulipes

Large species, pronotal collar with large lateral lobes or projections. Tyloidea of

male linear. Clypeal margin of female weakly emarginate or weakly quadridentate.

VAN LITH: New World Pluto 135

Mesopleura smooth and shining. P. annulipes and P. smithii are very closely related

and may be conspecific; P. smithii seems to have many geographic forms.

annulipes (Cameron) — 21 92 and 9 Z; Mexico; Costa Rica; El

Salvador

smithii (Fox) — 30 © and 45 Z; Argentina; Bolivia; Brazil;

Colombia; Panama Canal Zone; Paraguay;

Surinam; Venezuela

denticollis sp. nov. — 1 9, unknown; Peru

Group of pygmaeus

Axillae posteriorly free, inner side not connected with scutellum. Anterior

suture of scutellum without distinct carinae (not crenulate). Tyloidea of male

linear. Mesopleura almost smooth. Clypeal margin of female weakly emarginate in

P. pygmaeus and its subspecies axillaris, in P. facialis the clypeus of the female is

deeply emarginate, the median part of the clypeus of the male is also raised and

protruding.

pygmaeus pygmaeus (Brethes) — 76 9 and 31 &; Argentina; Bolivia; Peru

pygmaeus axillaris subsp. nov. — 33 © and 19 g, Bolivia; Brazil; British

Guyana; Colombia; Surinam

facialis sp. nov. — 19 and9 g; Colombia

Group of joergenseni

Metanotum raised in the middle, forming two “wings”, notably in the male.

Tyloidea of male linear. Clypeus of female with large lateral teeth, those of the

male too are relatively large. Labrum of female with many small teeth on anterior

margin (fig. 68), labrum of male (fig. 69) with four blunt teeth only. Mesopleura

reticulate alutaceous, punctate.

joergenseni (Brethes) — 44 © and 46 3; Argentina; Bolivia; Brazil;

Paraguay

Key to the species of the genus Pluto

1. Females (unknown: biformis, castaneipes, depressus, incarinatus and

EIS ER EN AD IMs ROM. Mave Siac On acl dba elon. al pea 2

— Males (unknown: arenivagus cubanus, brevipetiolatus, colonensis, denticollis,

duckei, emarginatus, jugularis, menkei, minutus, obscurus, occipitalis, rotundus,

scytinus, simplicicollis, spinicollis, stenopygidialis, strigellus and zonatus) .. 60

2. Lateral pronotal angles with long elongate-lobular projection. Clypeal margin

almost straight. Mesopleura and scutum smooth, sparsely finely punctate.

Petiole about as long as tergite 1. Gaster black, at most hind margin of tergite

distiicthy reds Bengt lO 13mm Ue. LA LOA Ra ESS, 3

136

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Lateral pronotal angles not long elongate-lobular, at most with equilateral-

triangular projection and then clypeal margin not straight, or pronotal

projections small and tergites |—3 red. Smaller ............... 130

Basitarsus and second tarsal segment of hind legs whitish. At least outer side

of fore tibiae black, base ivory-white on outer side, foreside reddish-brown.

Base of hind tibiae largely whitish. Pronotal tubercles dark brown or black.

Gaster black. Face pale golden. Mexico; Costa Rica; El Salvador .......

A wich Elica ee FS eee ne annulipes (Cameron) (p. 224)

Hind basitarsus and segments 2—3 or 2—4 of hind tarsi whitish. Outer side of

fore tibiae entirely reddish or with dark streak on posterior surface only,

foreside often yellowish. Base of hind tibiae with many small, brownish thorns

(except in Argentine form C). Pronotal tubercles reddish or reddish-yellow,

rarely brown or black. Narrow hind margin of tergite 1 sometimes reddish.

Face golden. Argentina; Bolivia; Brazil; Colombia; Panama Canal Zone;

Paraguay; Surinam, Venezuelan won ern smithii (Fox) (p.226):

Upper part of mesopleura with some longitudinal rugae, lower part sparsely

punctate. Hypo-epimeral area rugoso-punctate. Scutum shining, sparsely

punctate. Lateral pronotal angles sharp in dorsal view; pronotal tubercles

whitish. Petiole about half as long as tergite 1. Intercarinal space as wide as

fore basitarsus. Gaster black, margins of tergites reddish transparent.

Underside of flagellum pale yellowish. Hind tarsi whitish. Face silvery.

Length 9—9.5 mm, Arizona, California; Mexico. Mer ee

RER SE ON ren Ia longiventris (Malloch) (p. 154)

Upper part of mesopleura not coarsely rugose, or mesopleura rugoso-

punctate and also scutum with coarse rugae or hind tarsi dark brown.

Smaller! o a al a aa NE DÌ

Clypeus with long snoutlike projection, apex bidentate. Lateral pronotal

angles sharp. Mesopleura very finely alutaceous, sparsely finely punctate.

Petiole about as long as tergite 1. Gaster black, pronotal tubercles whitish, all

tarsi yellowish. Arizona; Mexico ........ clavicornis (Malloch) (p. 206)

Clypeus:inotimuchisnoutlike protuuding REE 6

Clypeal margin widely emarginate, large lateral teeth. Intercarinal space

much broader than fore basitarsus. Mesopleura finely alutaceous, scutum

sparsely punctate, mesopleura more densely so, interstices here a few times

size of punctures. Metanotum laterally somewhat depressed. Petiole about as

long as tergite 1. Hind margin of tergite | and all of tergite 2 red. Hind tarsal

segments | —4 yellowish-white. Argentina; Bolivia; Brazil; Paraguay .....

A eRe ED EENS joergenseni (Brethes) (p. 235)

Clypeal margin at most with small teeth or lobes, sometimes more or less

deeply emarginate;it ir ts DR Fila. AEN ENEN 7

Axillae posteriorly not connected with scutellum. Back of propodeum

coarsely reticulato-carinate; enclosure with large median area. Petiole about

as long-as'tergitedl: Gasterblack fais hats sel 8

Axillae posteriorly normally connected with scutellum ........... 10

Clypeal margin thick, very deeply, almost rectangularly emarginate, exposing

VAN LITH: New World Pluto 137

labrum. Back of propodeum very coarsely reticulate, carinae high, enclosure

deep. Dorsal half of mesopleura finely alutaceous, sparsely finely punctate.

Hind tarsi brown. Face pale golden or yellowish-silvery pubescent.

Colom AREN AA ETE facialis sp. nov. (p. 234)

Clypeal margin not unusually deeply emarginate. Propodeal carinae

moral Ee air HO eol Nam Ba mm LRT en. LO GE, 9

Scutum on anterior part, scutellum and mesopleura slightly alutaceous.

Mesopleura and scutum also sparsely finely punctate, scutellum more densely

so. Back of head somewhat alutaceous. Clypeal margin slightly emarginate

with distinct small lateral angles. Hind tarsal segments except apices brown.

Face usually silvery, sometimes golden pubescent. Argentina; Bolivia;

Bet BEI te BANEN DRAN LE pygmaeus pygmaeus (Brethes) (p. 230)

Scutum and scutellum smooth and shining, sparsely finely punctate,

interstices on scutellum a few times size of punctures. Mesopleura slightly or

not alutaceous. Back of head smooth and shining. Clypeal margin almost

straight, no distinct lateral angles. Hind tarsi brown or yellowish-brown. Face

usually silvery, rarely golden. Bolivia; Brazil; British Guyana; Colombia;

SIRIA EB SIRIO. pygmaeus axillaris subsp. nov. (p. 233)

Face golden, rarely silvery. Tergite 2 or tergites 2—3 red. Clypeal margin with

small lateral teeth and more or less distinct median tooth. Vertex with broad

transverse depression between ocelli and oculi. Mesopleura finely alutaceous,

rarely shining, and very finely punctate. Propodeum coarsely reticulate.

Betiole as long asitergitenle titan Ua MN wal gio! Bang lei ae sek, Il

Notallthesecharacters combined least. 12312 25 ato 39228. 12

Face golden (silvery in females from Argentina and Bolivia). Median clypeal

tooth indistinct. Hind margin of tergite 1, all of tergite 2 and sides of tergite 3,

or all of tergites 2—3 red. Labrum reddish. Apical 2/3 of hind basitarsus

yellowish-white. Argentina; Bolivia; Brazil; Surinam ...............

EEE Peri a NA Prise) Viole medius medius (Smith) (p. 200)

Face deep golden. Median clypeal tooth more distinct. Tergite 1 except for

two black marks, all of tergites 2—3 and sides of tergite 4 red. Labrum black.

Hind tarsi almost entirely dark brown. Curacao; Venezuela ...........

Re RE ho GA. HO. Lets medius zuliensis subsp. nov. (p. 202)

Mesopleura including hind margin and hypo-epimeral area shining or

superficially alutaceous (x 30), hind margin of mesopleura not rugulose or

coriaceous, at most with some shining short rugae. Hind tarsal segments 1—4

brown or reddish-yellow, rarely whitish. Punctation of mesopleura very fine

and sparse or very indistinct and mesopleura densely pubescent; if punctation

more distinct then lateral pronotal angles very sharp and hind tarsal segments

whitish (cf. araguensis). Intercarinal space not wider than fore basitarsus. 13

Mesopleura usually distinctly alutaceous and/or punctate or coarsely rugoso-

punctate; if very finely punctate or somewhat shining, then punctation close

or hind margin rugulose or coriaceous, or lateral pronotal angles not very

sharp or hind tarsal segments 1—4 whitish. Intercarinal space sometimes

widenthamtoresbasitarsuss Hi 990 AMEDEE. EINEN. YO. SIs 28

138

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Clypeal margin more or less trilobate or tridentate, median tooth sometimes

very small. Occipital carina not connected with hypostomal carina. Scutum

and scutellum dull. Mesopleura smooth and shining, impunctate or

indistinctly punctate. Petiole as long as tergite 1 or slightly longer. Pronotal

tubercles and easter black \hind\tais brown EE <n eta ae 14

Clypeal margin not distinctly trilobate or tridentate, if indistinctly so then

gasterlargely reden. nr MERE RE ee 15

Clypeal margin with projecting shining median lobe and distinct triangular

lateral teeth. Pygidial area little longer than broad. Last gastral segment dark

brown. Face silvery. Ecuador; Peru; Surinam .. trilobatus sp. nov. (p.218) :

Median tooth of clypeal margin very small. Punctation of scutum weaker than

in preceding form. Pygidial area about 1.5 times as long as broad. Last gastral

segment reddish. Face silvery. Male unknown. Peru ................

aa da Lele Reg pean edie crate Ys Le ISIN sp. aff. trilobatus? (p. 220)

Clypeal margin with four distinct short lobes. Pronotal collar laterally with

large equilateral-triangular lobes, projecting forward. Mesopleura and

scutum smooth and shining. Petiole about as long as tergite 1. Underside of

flagellum orange-red. Gaster black. Pubescence of face golden, of dorsal side

of thorax brownish-golden. Male unknown. Peru denticollis sp. nov. (p. 229)

Clypeal lobes, if any, indistinct. Pronotum not with large triangular lobes,

angles at most sharp or with small triangular tooth or one or more of gastral

tergitesili 3 reden she eh Lost Res I ee REN: 16

One or more gastral tergites largely or entirely med) ay eee 17

Gaster black, at most hind margins of tergites and base of tergite 2 reddish or

yellowish transparent. Petiole about as long as or 4/5 length of tergite 1. Hind

tarsi, at least segments 2—5 brownish or reddish ............... 22

Lateral pronotal angles projecting as a sharp tooth or spine. Mesopleura

finely alutaceous, almost smooth, finely but distinctly punctate, hypo-

epimeral area shining, indistinctly punctate. Clypeal margin not distinctly

dentate. Petiole slightly over half length of tergite 1. Tergites I—3 except for

extreme base of tergite 1 red. Hind tarsi yellowish-brown. Male unknown.

Brazil: Panama au ER spinicollis sp. nov. (p.217)

Pronotum not with distinct lateral spines. Tergite 1 more extensively

darkened neten 22H EE EEN eee eee NEN: 18

Mesopleura and hypo-epimeral area smooth and shining .......... 19

Mesopleura relatively dull owing to fine reticulation and many, sometimes

indistinct, hairbearing punctures. At least tergites2—3red......... 21

Petiole about as long as tergite 1. Clypeal emargination about half total width.

Occipital carina not distinctly connected with hypostomal carina. Propodeum

with long oblique striae; enclosure shining, with large median area. Narrow

margin of tergite 1, all of tergite 2, basal half of tergite 3 and sternites 2—3

red. Hind tarsal segments 1—4 yellowish-red. Face silvery. Male unknown.

Brazil ins: er as ASR ene ICE duckei sp.nov. (p.212)

Petiole about 2/3 length of tergite 1. Clypeal emargination about 1/3 of total

width or indistinctly quadridentate. Occipital carina distinct. Propodeum

20.

22”

23;

24.

25.

VAN LITH: New World Pluto 139

reticulate, dorso-laterally with parallel carinae. Narrow margin of tergite 1, all

onnearrallofitergite 2'andistérnite 2ired u. 2 ea. 2 enke ao) 20

Face pale golden. Clypeal margin indistinctly quadridentate, this part about

half total width. Antennal segments 10—11 shorter than broad. Scutum

shining, very sparsely punctate. Hind tarsi brownish-red or yellowish-red.

Matcrnknown Brazile. Jt i zonatus sp.nov. (p.215)

Face silvery. Clypeal margin with weak emargination, about 1/3 of total

width. Antennal segments 10—11 about as long as broad. Scutum shining,

more densely punctate. Hind tarsi dark brown, apices paler. Male unknown.

MERE ey ieee ee PAS VS EE Pi menkei sp.nov. (p.216)

Petiole distinctly shorter than tergite 1. Propodeum dorso-laterally almost

smooth, lower part weakly reticulato-carinate. Tergite 2 entirely red, tergite 3

entirely or nearly entirely so. Hind tarsi brown. Ecuador; Peru .........

ER EEE RER ENDET ER! townsendi (Cockerell) (p. 207)

Petiole as long as or slightly longer than tergite 1. Back of propodeum

relatively coarsely reticulato-carinate, carinae dorso-laterally mostly parallel.

Apical half of tergite | and remainder of gaster red, bases of tergites 4—6

sometimes brownish. Hind tarsi brown. Bolivia; Brazil; Colombia; Ecuador;

RE tei ento ED an, marthae sp.nov. (p. 208)

Mesopleura and hypo-epimeral area somewhat dull, finely, rather densely,

punerate: Pronotaktubercles-whitish vu; sn moe. 2... ee ee 23

Mesopleura and hypo-epimeral area smooth and shining, or very indistinctly

alutaceous, sparsely very finely punctate. Pronotal tubercles whitish or

BE a el en retro MAUR No As 24

Lateral pronotal angles rectangular. Mesopleura and hypo-epimeral area with

indistinct, rather dense punctation. Scutum shining. Gaster black with reddish

hind margins of tergites and narrow red base of tergite 2. Hind tarsi brown.

PGi ER am MENE metanus sp.nov. (p. 210)

Lateral pronotal angles sharp, protruding laterally. Mesopleura finely

reticulate alutaceous, finely but distinctly punctate, interstices a few times

size of punctures. Hypo-epimeral area somewhat shining, indistinctly

punctate. Scutum finely reticulate alutaceous, punctures stronger than on

mesopleura, interstices a few times size of punctures. Hind tarsal segments

1—4 whitish. Male unknown. Venezuela .... araguensis sp.nov. (p.221)

Esenotaltüubercles brown. or blacktes 27 san Bande netti nen! 25

Pronotal tubercles whitish or yellowish-white. Frons somewhat raised, also in

Iondoftanterionocellusa. > A Ee. te Ban. Er; mer 271

Lateral pronotal angles obtuse. Median part of clypeal margin with distinct

narrow shining depression, very slightly widely emarginate. Back of

propodeum almost dull, with long parallel carinae; enclosure shining. Frons

not distinctly raised before anterior ocellus. Hind tarsi brown, at least

segments 2—5. Face silvery. Argentina; Bolivia; Brazil; Ecuador; Paraguay:

Ben2Surmanm:Venezuelanr tom ont aie Shon os nitens sp.nov. (p.210)

Lateral pronotal angles sharp or rectangular, not distinctly protruding.

Clypeal margin with four indistinct lobes or nearly straight. Frons distinctly

140

26.

27.

28.

Zon

30.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

raised around anterior ocellus. Back of propodeum somewhat shining,

coarsely reticulato-carinate, dorso-laterally with some oblique carinae . 26

Antennal segments 9—11 shorter than broad. Clypeal margin with four

indistinct lobes. Underside of flagellum orange. Hind tarsi reddish, segments

2—4 slightly darkened. Fore tibiae entirely yellowish-red. Face pale golden.

Male unknown Brazil 2... ayo er simplicicollis sp. nov. (p.215)

Antennal segments 9—11 distinctly longer than broad. Clypeal margin

straight. Flagellum except for underside of last segment blackish-brown.

Foreside of fore tibiae reddish-brown. Hind tibiae including base and all of

hind tarsi dark brown. Face silvery appressed pubescent and also with long

erect brownish hairs. Larger, about 9.5 mm. Male unknown. Argentina ....

MCDM Er Ae IS NO ORTI NIRO obscurus sp. nov. (p.214)

Lateral pronotal angles sharp, in antero-lateral view distinctly protruding.

Lateral parts of occipital carina broadened, transparent, in dorsal view as high

as pubescence of tempora. Median part of clypeal margin not depressed,

reddish. Hind tarsal segments 1—4 yellowish, apices somewhat brown. Face

silvery: Male:unknown. Perú Ut. ie va. occipitalis sp. nov. (p.217)

Lateral pronotal angles rectangular. Occipital carina of normal height.

Median part of clypeal margin distinctly depressed. Last tergite reddish. Hind

tarsal segments 1—4 whitish. Face pale golden. Peru ................

TOI RENE a MER ras ee rufanalis sp. nov. (p.220).

Mesopleura entirely distinctly and closely finely punctate, interstices smaller

than punctures, sometimes interstices below somewhat larger or middle part

finely striato-punctate. If mesopleura almost indistinctly punctate and striate

and tergite 2 red, cf. strigellus. Petiole between half and 3/5 length of tergite 1,

if longer, then cf. albifacies, No. 55. Face silvery ................ 29

Mesopleura with interstices at least on upper part largely about size of

punctures or larger, or upper part of mesopleura more coarsely punctate,

rugoso-punctate or rugulose, or face more or less golden or petiole

longer wii ae QR Tai. AEN Se EUD EEEN 34

Pronotal tubercles black. Face pale golden. Gaster black. Hind tarsal

segments I—4 yellowish-brown, segments |—3 with darker base. Mesopleura

dull, densely finely punctate, upper part also finely striate. Propodeum dull,

dorsal half with fine oblique carinae. Mexico .. alphitopus sp.nov. (p. 195)

Pronotal tubercles whitish. Face silvery eeN 30

Gaster black, at most sides and hind margins of tergites 2—3 and of sternites

2=S TOR. a ik er Go ae aes eae ee eon ANNE SAR EEE 31

Atleasttergite'2ientirelyirediet. terre er 32

Hind tarsi whitish. Mesopleura dull, distinctly finely punctate, interstices on

lower part often larger than punctures. Petiole about 2/3 length of tergite 1.

Argentina; Bolivia: Brazile sole ee oe stramineipes sp.nov. (p. 204)

Hind tarsi brown. Mesopleura very densely finely punctate, partly striato-

punctate. Scutum distinctly but finely, not sparsely punctate. Petiole about

half length tergite 1. Cuba; Jamaica; Nicaragua; Mexico .............

oats, oa an Re ep eke argentifrons (Cresson) (p. 192)

321

33.

34.

35.

36.

37.

38.

VAN LITH: New World Pluto 141

Scutum shining, central part sparsely finely punctate. Upper half of

mesopleura dull, very finely almost indistinctly punctate and extremely finely

longitudinally striate; lower half somewhat shining, interstices about size of

punctures. Apical half of tergite 1, all of tergite 2 and most of tergite 3 red.

Hind tarsi brown. Male unknown. Argentina ... strigellus sp. nov. (p. 205)

Scutum more densely or strongly punctate. Punctation of mesopleura fine but

SHC Strates. als tesa lege am arte nml 33

Scutum densely and regularly finely punctate. Clypeal margin slightly

emarginate, somewhat depressed with trace of median tooth. Mesopleura

dull, very closely and finely punctate. Hind tarsi yellowish, very slightly

brownish darkened. Hind margin of tergite 1 and all of tergites 2—3 red. Male

maknewn Venezuela: no catia; ar. scytinus sp. nov. (p. 203)

Scutum strongly punctate, hind margin rugulose. Clypeal margin weakly

emarginate, no trace of median tooth. Mesopleura dull, densely, somewhat

stronger punctate. Hind tarsi brown, apices paler. Hind margin of tergite 1, all

of tergite 2 and base of 3 red. Male unknown. Argentina .............

EEE Ann Hie ersten. alt colonensis sp. nov. (p. 198)

Scutum and scutellum with strong, shining, irregular rugae and also strongly

punctate. Mesopleura dull, upper half at least anteriorly coarsely rugoso-

punctate, punctures large, lower half not rugose and interstices larger than

punctures. Petiole red or black, nearly as long as tergite 1. At least tergite 2

and hind margins of following segments more or less red, sometimes tergites

1—2 entirely red. Tarsal segments 1—4 whitish-yellow. Face pale golden.

Florida, Georgia, Louisiana, Maryland, Mississippi, North Carolina, South

Carolin VAT SINE <P von dass our la rufibasis (Malloch) (p. 186)

Scutum not with strong rurae. Sculpture of mesopleura usually finer. Petiole

neven diensten we naive bemad cuit aah teen dal Ae 35

Hind tarsi largely brown or orange-brown, apices of segments often paler. If

basitarsus yellowish-white, then face golden .................. 36

Hind tarsi whitish or yellowish, usually last segment brown; if all tarsal

segments partly brownish, gaster entirely black, pronotal tubercles whitish

andipropodeumicoarsely-reticulate, cfevansii! vorsers al 47

Pronotal tubercles dark brown or black. Gaster black, at most hind margins of

REMPIPCSMEGGISH: ett tetes rig sy tamer it ne ei 37

Pronotal tubercles whitish. Gaster not always black ............. 40

Face dark brown-golden pubescent, leaving sculpture of clypeus well visible.

Punctation of mesopleura irregular, distinct on uppe iart, almost absent on

lower part. Back of propodeum very finely carinate, most carinae obliuué

iarallel. Petiole about half length of tergite 1. Pygidial area over 2.5 times as

long as broad. All tarsi entirely brown. Male unknown. Arizona ........

PRE onelioo tht cr) bg stenopygidialis sp.nov. (p. 176)

Face normally and appressed, densely silvery or golden pubescent. Fore and

mid tarsi yellowish-brown, yellowish or whitish, hind tarsi brown or orange-

brown Petolelonseremsen.betil..stenhmas.nécluniies alasli.uolalabon 38

Clypeal margin widely, shallowly emarginate, with distinct lateral teeth.

142

39:

40.

41.

42.

43.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Mesopleura and hypo-epimeral area indistinctly alutaceous, somewhat

shining. Mesopleura irregularly sparsely punctate, hypo-epimeral area more

densely punctate. Back of propodeum coarsely reticulate, carinae dorso-

laterally oblique. Petiole about 2/3 length of tergite 1. Face silvery. Male

unknown. Mexico: m... nn SE emarginatus sp.nov. (p. 185)

Median part of clypeal margin almost straight or slightly rounded.

Mesopleura more densely punctate, very finely alutaceous ......... 39

Mesopleura regularly, moderately densely punctate, hypo-epimeral area

finely alutaceous, very finely punctate. Scutum and scutellum finely

alutaceous, sparsely punctate. Back of propodeum dull, finely reticulate,

dorso-lateral carinae oblique. Petiole about 2/3 length of tergite 1. Hind tarsi

orange-brown. Face silvery, yellowish-silvery or golden. Costa Rica; El

Salvador; Guatemala; Mexico ........... punctatellus sp. nov. (p. 179)

Mesopleura stronger, more irregularly, moderately densely punctate; hypo-

epimeral area shining, very finely punctate. Scutum and scutellum smooth

and shining, moderately densely punctate. Back of propodeum shining,

coarsely reticulate. Petiole about as long as tergite 1. Face golden.

Ecuador; ir Ue TITLE IN ra fritzisp. nov. (p. 199)

Gaster black. Mesopleura on upper part obliquely rugulose, below finely

punctate, interstices there size of punctures, or larger. Scutum shining, finely,

not densely punctate. Intercarinal space almost as wide as fore basitarsus.

Puerto Rico; Dominican Republic; Haiti; Leeward Islands; Virgin

Islands: aan moie cheat Oe oe EREN atricornis (Malloch) (p. 194)

At least tergite 2 more or less red. Sculpture of mesopleura finer, not

distinctly: ruguloseorstriato-punctate 2 2... ais ie ee 41

Hypo-epimeral area punctate and with longitudinal striae or rugae,

sometimes most distinct from behind. Scutum shining, finely punctate. Petiole

2/3 to 3/4 length of tergite 1. Hind margin of tergite 1, all of tergite 2 and base

of:tergite 3red. Faceisilvery nan RE ER 42

Hypo-epimeral area not with distinct striae or rugae, more or less densely

punctate; Face sometimes golden. a. ine. eer a eee 43

Mesopleura indistinctly alutaceous, somewhat shining, finely punctate,

interstices a few times size of punctures, hind margin coriaceous and with fine

striae. Hypo-epimeral area ruguloso-punctate. Texas ...............

LIES eh Nari IP texanus (Malloch) (p. 170)

Mesopleura more distinctly alutaceous, distinctly punctate, interstices mostly

about size of punctures, hind margin granulose. Hypo-epimeral area with

coarse rugae, punctures finer. Florida, Maryland, North Carolina, South

Cardlinasst sh ei oo AAA littoralis (Malloch) (p. 171)

Petiole nearly as long as tergite 1. Intercarinal space broader than first

basitarsus. Mesopleura indistinctly punctate, interstices a few times size of

punctures. Punctation of hypo-epimeral area imperceptible with enlargement

x 30. Scutum and scutellum finely alutaceous. Back of propodeum

moderately finely reticulato-carinate. Hind margin of tergite 1 and all of

tergites 2—3 red. Face silvery. Male unknown. Brazil ...............

44.

45.

46.

47.

48.

VAN LITH: New World Pluto 143

PES CES CR GRENET ak. WI, RE jugularis sp. nov. (p. 178)

Petiole at most 2/3 length of tergite 1. Intercarinal space less broad than first

basitarsus. Back of propodeum finely reticulato-carinate, sometimes dorso-

laterallyawith tine oblique striae min ee ea nahe: 44

Hypo-epimeral area almost shining, distinctly finely punctate, interstices

about size of punctures. Scutum usually shining, medially sharply punctate.

Mesopleura dull, finely alutaceous and punctate. Petiole about 2/3 length of

tergite 1. Pygidial area about twice as long as broad. Hind margin of tergite 1,

‚all of tergite 2 and all or at least base of tergite 3 red. Alabama, California,

District of Columbia, Florida, Iowa, Louisiana, Mississippi, New Mexico,

North Carolina, South Carolina, Tennessee; El Salvador; Mexico; Nicara-

EUD se NME IRE Sa o ER BER TEE sayi (Rohwer) (p. 173)

Hypo-epimeral area dull, densely finely or indistinctly punctate. Scutum

finely reticulate alutaceous or pygidial area narrow, about three times as long

AIDROAU Are Ua uy EEL ME TLE OE ORI SRE ES oe ER 45

Hind margin of tergite | and all of tergites 2—6 red. Scutum sparsely

punctate. Mesopleura reticulate alutaceous, irregularly, partly closely, finely

punctate. Petiole about 2/3 length of tergite 1. Mexico: Baja Califor-

MIE sheet a RE FAY er Bo AUX STE, abbreviatus sp. nov. (p. 182)

Not tergites 4—6 entirely red. Petiole about half length of first tergite .. 46

Hypo-epimeral area dull, granulose, indistinctly finely and closely punctate.

Pygidial area narrow, about three times as long as broad. Scutum and

scutellum shining, indistinctly alutaceous, sparsely punctate. Mesopleura

finely alutaceous, anteriorly sparsely finely punctate. Intercarinal space

nearly as broad as fore basitarsus. Apex of tergite 2 and at least base of tergite

3 red. Hind tarsal segments 1—4 pale brown, apices paler. Male unknown.

California (cf. also No. 52) ............. brevipetiolatus (Malloch) (p. 164)

Hypo-epimeral area distinctly punctate, interstices mostly smaller than

punctures. Pygidial area about 2.5 times as long as broad. Scutum alutaceous.

Mesopleura finely alutaceous, punctate, interstices mostly a few times size of

punctures. Intercarinal space narrower. Tergite 2 usually entirely red. Hind

tarsal segments 2—4 more or less brown, basitarsus for greater part so.

Arizona, California, Texas, New Mexico; Mexico (for pale-legged form, cf.

AEO INC LE) AR SU A TRS ae basifuscus sp. nov. (p. 160)

Gaster more or less red, usually tergite 2 or 3 entirely so; if only apex of

tergite 2 or base of tergite 3 red, then petiole short or intercarinal space as

Baidesasctoneibasitarsustsnug Monroe Wiler gegetertarak 48

Gaster black, at most hind margins of tergites or sides of tergites 2—3 and

sternites 2—3 somewhat reddish or yellowish transparent, or last tergite

reddish. Intercarinal space not as wide as fore basitarsus .......... SI

Intercarinal space very narrow, occipital carina and hypostomal carina almost

touching. Petiole 2/3 or 3/4 length of tergite 1. Back of propodeum reticulato-

carinate, dorso-laterally some fine oblique parallel carinae. Median part of

elypealmargimmabeutistraight: „aurea she nei ette 49

Intercarinal space wider or petiole about half as long as tergite 1 or clypeal

144

49.

50.

Sle

52.

33.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

margindistinctly rounded’. #92 seat] en ea 50

Hypo-epimeral area dull, finely punctate, posteriorly indistinctly striate.

Mesopleura alutaceous, punctures moderately large, mostly contiguous, on

median part striato-punctate; punctures below somewhat finer, interstices

mostly smaller than punctures. Scutum shining, strongly punctate, interstices

mostly smaller than punctures. Fore and mid tibiae yellowish-red. Hind

margin of tergite 1, all of tergite 2 and at least base of tergite 3 reddish.

Florida, Georgia, North Carolina arenivagus arenivagus Krombein (p. 190)

Hypo-epimeral area entirely distinctly longitudinally striate and punctate.

Mesopleura somewhat more coarsely punctate. Punctation of scutum finer,

interstices mostly twice as large as punctures. Fore tibiae brown below, mid

tibiae dark brown below. Base of tergite 2 darkened, tergite 3 entirely red.

Male unknown. Cuba ........ arenivagus cubanus subsp. nov. (p. 192)

Petiole about 2/3 or 3/4 length of tergite 1. Back of propodeum reticulato-

carinate, carinae dorso-laterally somewhat parallel .............. 51

Petiole about half as long as tergite 1. Back of propodeum finely densely ©

reticulato-carinate, upper part finely:striate ie. verven m. ae 52

Intercarinal space as wide as fore basitarsus. Mesopleura with moderately

large, shallow punctures, interstices on upper part as large as or larger than -

punctures, punctures medially partly in rows, interstices below wider. Scutum

sparsely punctate. Clypeal margin almost straight. Apex of tergite 2 and

usually all of tergite 3 red. Arizona, California, Florida, New Mexico, South

Carolina, Texas: Mexicoyis: arene MAGEREN suffusus (Fox) (p. 166)

Intercarinal space about half width fore basitarsus. Mesopleura with regularly

placed small punctures, interstices mostly a few times size of punctures.

Scutum more densely punctate. Clypeal margin rounded, reddish. Tergites

1—3 red. Male unknown. Florida ........... rotundus sp. nov. (p. 165)

Hypo-epimeral area dull, granulose, indistinctly finely and closely punctate.

Pygidial area narrow, about three times as long as broad. Mesopleura finely

alutaceous, anteriorly sparsely finely punctate. Scutum and scutellum shining,

indistinctly alutaceous, sparsely punctate. Intercarinal space nearly as broad

as fore basitarsus. Apex of tergite 2 and base of tergite 3 red. Hind tarsal

segments 1—4 indistinctly brownish, apices paler. Male unknown. Califor-

NASE AER NN an Sa Te brevipetiolatus (Malloch) (p. 164)

Hypo-epimeral area distinctly punctate. Pygidial area less than three times as

long as broad) Tergite!2 usuallyentinely red’) PM ONE RENE EEE 53

Hypo-epimeral area somewhat glossy, finely punctate, interstices at least size

of punctures or larger. Mesopleura finely alutaceous, finely punctate, on

median part interstices mostly not much larger than punctures, on upper part

much larger. Scutum and scutellum shining, punctate, interstices mostly a few

times size of punctures. Hind tarsal segments 1—4 yellowish-white. Iowa,

Louisiana, New Mexico, Texas ........ angulicornis (Malloch) (p. 157)

Hypo-epimeral area dull, distinctly punctate, interstices mostly smaller than

punctures. Mesopleura finely alutaceous, punctate, interstices mostly a few

times size of punctures. Scutum finely alutaceous, sparsely punctate... 54

54.

SI.

56.

57.

58.

59:

VAN LITH: New World Pluto 145

All tarsi including last segment whitish; extreme base of hind basitarsus

brownish. Arizona, California, Texas .... pallidistigma (Malloch) (p. 158)

Tarsi whitish, base of hind basitarsus and last segment of hind tarsi brown (or

3/4 of hind basitarsus and bases of hind tarsal segments 2—4 brown; cf. No.

46). Arizona, California, Texas, New Mexico; Mexico ...............

EEP ne ep ln basifuscus sp. nov. (p. 160)

Petiole nearly as long as tergite 1. Mesopleura dull or weakly shining,

punctate, interstices about size of punctures. Propodeum reticulato-carinate

with fine laero-dorsal oblique striae. Hypo-epimeral area dull, densely very

finely punctate. Sides of tergites 2—3 and all of sternites 2—3 sometimes red

or reddish-brown. Face densely silvery pubescent. Iowa, Texas .........

MEME AM RARE ER albifacies (Malloch) (p. 187)

Petiole distinctly shorter than tergite 1 or mesopleura strongly punctate 56

Back of propodeum dull, with faint striae, lower part of back not distinctly

reticulate. Mesopleura finely alutaceous, punctures on upper part relatively

large and superficial, lower part very sparsely punctate. Hypo-epimeral area

densely superficially punctate. Petiole about half length tergite 1 or shorter.

Face silvery. Length 5 mm. Male unknown. Texas .................

REEN ETE AE Er PORC PP minutus (Malloch) (p. 162)

Sculpture of propodeum coarser or propodeum distinctly reticulate.

Mesopleura more regularly, sometimes sparsely punctate. Length 6—7

[TRIM 9 loi 6 at D E OE IDNR SER SR RSS SSSR See ee Sa a SÙ

Face brassy yellow or pale golden. Petiole about 2/3 of length of tergite |.

Mesopleura dull, on median part with shallow, partly moderately large

punctures, lower part with finer, widely placed punctures, below hypo-

epimeral area some very wide interstices. Hypo-epimeral area densely finely

punctate. Scutum strongly punctate, partly rugose. Back of propodeum

coarsely reticulato-carinate. Outer side of tibiae yellowish. Texas, North

CarolinaBelize; Mexico. =; m, sues aerofacies (Malloch) (p. 183)

Face silvery. Petiole about half length of tergite 1 or little longer ..... 58

Mid tibiae largely brown, not yellowish-white on outer side. Scutum and

scutellum finely alutaceous, sparsely distinctly punctate. Back of propodeum

coarsely reticulato-carinate. Mesopleura finely punctate, interstices a few

times size of punctures. Hypo-epimeral area very finely punctate, interstices

larger than punctures. Hind tarsi whitish, base of basitarsus and last tarsal

segment brown, sometimes also segments 3—4 darkened. Mexico .......

REPS EMER EBENEN: EEE N BERNER WREE de evansi sp. nov. (p. 184)

Mid tibiae whitish on outer side. Segments I—4 of hind tarsi whitish... 59

Mesopleura with large punctures, partly in longitudinal rows; interstices

partly as large as or larger than punctures. Intercarinal space linear. Scutum

shining, strongly punctate. Propodeal enclosure irregularly reticulate. Hypo-

epimeral area strongly longitudinally striato-punctate. District of Columbia,

Florida, Louisiana, Missouri, South Carolina, Tennessee, Texas,

VIT ARMOR LE RRQ SENEGAL A ME Lan el tibialis (Cresson) (p. 188)

Mesopleura with distinct but finer punctures, interstices mostly larger than

146

67.

63.

64.

65.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

punctures, below a few times size of punctures. Intercarinal space more than

half as wide as fore basitarsus, rarely narrower. Scutum finely alutaceous,

finely punctate. Propodeal enclosure usually shining, with large median area.

Hypo-epimeral area finely punctate, interstices about size of punctures.

Arizona, Texas, Mexico, Guatemala ......... spangleri sp.nov. (p.,162)

Lateral pronotal angles with long lobular projection. Large species ... 61

Lateral pronotal angles not with long lobular projection........... 62

Hind basitarsus, sometimes also mid basitarsus, whitish, following segments

dark. Antennal segments 5—11 or 5—12 with linear tyloides, at most half as

long as segment. Pronotal tubercles dark brown, at most with paler hind

margin. Base of tergite 2 black. Fore tibiae with black streak on outer side,

base yellowish-white. Scape ,of antennae black. Face silvery pubescent.

Mexico: Costa Rica lan RASS annulipes (Cameron) (p. 224)

Hind basitarsus and tarsal segments 2—3 or 2—4 whitish. Usually antennal

segments 4—13 or 5—13 with linear tyloides (if only segments 5—11 with

tyloidea, then cf. smithii form D). Pronotal tubercles reddish or yellowish-red.

Tergites 1—2 largely red or only base of tergite 2 red. Fore tibiae entirely

reddish. Scape of antennae black or reddish. Face silvery or pale golden.

Argentina; Bolivia; Brazil; Colombia; Panama Canal Zone; Paraguay;

Suninam: lat. ski ie smithii (Fox) (p. 226)

Antennae strongly clavate, segments 10—12 shorter than broad, no tyloidea.

Mesopleura, hypo-epimeral area and scutum shining, mesopleura indistinctly

alutaceous, sparsely punctate. Lateral pronotal angles sharp. Petiole about as

long as tergite 1. Gaster black, hind margins of tergites reddish. Hind tarsi

pale yellowish-brown. Arizona; Mexico .... clavicornis (Malloch) (p. 206)

Antennaenot;stronglyiclavate: a... sik a. Ae eee eee 63

Tyloidea partly oval, at most linear on segments 4—5 or 4—8, or tyloidea

interruptedionisegmentsg-- 10 hi) na DE ar Ne EEE 64

Tyloidea all linear or all tyloidea indistinct or absent (cf. also abbreviatus) 73

Mesopleura and hypo-epimeral area very coarsely rugose, punctures

indistinct. Scutum shining, a few coarse punctures or scutum rugoso-

punctate. Antennal segments 6—12 with large oval tyloides, a small one on

segment 13. Intercarinal space about as wide as fore basitarsus. Petiole about

as long as tergite 1. Hind tarsal segments 1—4 whitish. Arizona, California;

Mexico: Rn Dire longiventris (Malloch) (p. 154)

Mesopleura distinctly coarsely punctate and rugoso-punctate, tyloidea small

and intercarinal space wide, or mesopleura punctate only, mostly with distinct

interstices betweenipunctüres ata see EE RER 65

Mesopleura coarsely striato-punctate or rugoso-punctate, hypo-epimeral area

distinctly punctate. Scutum coarsely punctate with tendency to transverse

rugosity. Petiole about as long as tergite 1. Antennal segments 5—13 with

small oval tylidea. Intercarinal space about as wide as fore basitarsus. Gaster

black, hind margins of tergites reddish. Hind tarsal segments 1—4 whitish.

Arizona, California, Florida, New Mexico, South Carolina, Texas; Mex-

consi RETTEN IS Perea ea ae etal PESTE suffusus (Fox) (p. 166)

66.

67.

68.

69.

70.

VAN LITH: New World Pluto 147

Distinct interstices between punctures of mesopleura, at least on lower part.

Scutum less coarsely punctate. Antennae different, tyloidea often larger 66

Ventral part of occipital carina unusually high. Intercarinal space broader

than fore basitarsus. Antennal segments 6—12 with elongate tyloidea, oblong

and widening towards apex on segments 6—9, broadly oval on segments

10—11, narrower on segment 12. Mesopleura somewhat shining, coarsely

punctate, interstices smaller or slightly larger than punctures. Back of

propodeum including enclosure closely reticulato-carinate. Petiole as long as

tergite 1. Gastral tergite 2 and/or 3 or 2—4 red. Hind tarsi entirely yellowish-

white. Stigma pale yellowish-brown, central part sometimes darkened.

Arizona, California, Texas .......... pallidistigma (Malloch) (p. 158)

Occipital carina not unusually high. Stigma usually dark (cf. basifuscus).

Intercarinal space narrower than fore basitarsus or antennal segments 6—12

HO mmUniclongate tyloideaises amine air Im ee en 67

Tyloidea absent on segments 8—10 or much smaller than on preceding

segments, elongate on segments 4—7, short and almost linear on segments

11—12. Upper half of mesopleura coarsely punctate, interstices about size of

punctures. Hind margin of mesopleura and entire hypo-epimeral area

coarsely rugose. Petiole about as long as tergite 1. Hind margins of tergites

and bases of tergites 2—3 red. Hind tarsibrown ................ 68

Also antennal segments 8—10 with distinct tyloides ............. 69

Tyloides on segment 8 small but distinct, on segments 9—10 very small, still

distinct. Scutum shining, finely punctate. Texas . texanus (Malloch) (p. 170)

Tyloides small or absent on segment 8, absent or very indistinct on segments

9—10. Scutum shining, somewhat more strongly punctate. Florida, Maryland,

North Carolina, South Carolina .......... littoralis (Malloch) (p. 171)

Tyloidea on segments 4—5 linear and short or indistinct, on segment 6 long

oval, slightly raised, on segments 7—10 almost triangular, and angularly

raised, highest point close to middle of each segment; tyloidea on segments

11—12 linear, less distinct, a short tyloides on segment 13. Intercarinal space

narrow. Mesopleura slightly alutaceous, finely punctate, interstices on upper

part larger, on lower part size of punctures or smaller. Petiole about 2/3 length

of tergite 1, tergite 1 about 1.5 times as long as broad. Hind margin of tergite 1

and bases of tergites 2—3 more or less red. Hind basitarsi whitish, segments

2—4 yellowish-brown. Arizona, Iowa, New Mexico, Texas ............

EERE AN errr tht IOI IRB He A angulicornis (Malloch) (p. 157)

Antennal segments 5—12 or 6—12 with distinct tyloides, less strongly, not

angularly raised. Gaster black, more slender, tergite | longer ....... 70

First tergite over twice as long as broad. Antennal segments 5—6 or 5—7 with

narrow tyloides, shorter than segments, segments 7—12 or 8—12 with small

oval tyloides, sometimes a small point on segment 13. Mesopleura dull,

alutaceous, upper half anteriorly usually striato-punctate, below with distinct

interstices between punctures. Petiole somewhat shorter than tergite 1. Hind

tarsal segments 1—4 entirely whitish. Stigma dark brown. Arizona, Texas;

Mexico: Guatemala, dopu kai. ea ae spangleri sp.nov. (p. 162)

148

12%

13%

74.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

First tergite about twice as long as: broad). Aton Ss nn. 71

Antennal segments 5—6 with short narrow tyloides, sometimes absent on

segment 5, segments 7—12 with broad oval tyloides, segment 13 with small or

indistinct tyloides. Mesopleura slightly alutaceous, almost shining, strongly

punctate, interstices mostly as large as punctures, no striae. Usually basal

fourth of hind basitarsi brown or black, remainder of basitarsus and segments

2—4 whitish, or hind tarsus largely brown. Stigma usually whitish or pale

brown. Arizona, California, Texas, New Mexico; Mexico ............

nr e dot PRES Tadini ate tet a basifuscus sp.nov. (p. 160)

Antennal segments 6—8 or 7—8 with narrow tyloides or tyloidea distinctly

raised in lateral view. or gaster partly red. |. ves oen ee 12

Gaster black. Antennal segments 7—8 with long narrow tyloides (short one on

segment 6), tyloides on segment 9 widening towards apex, on segments 10—12

broad oval and shorter than segments; segments 8—12 in lateral aspect

angularly raised on apical 1/3. Antennal segments 10—12 about quadrate,

Mesopleura alutaceous, strongly, partly densely, punctate. Punctation of

hypo-epimeral area with distinct interstices. Back of propodeum coarsely

reticulato-carinate. Petiole about 2/3 length of tergite 1. Hind tarsal segments

1—4 whitish. Female unknown. Mexico ....... biformis sp.nov. (p. 169)

Hind margin of tergite 1 and all of tergites 2—-3 reddish. Antennal segments

5—8 with linear tyloides (on segments 6—8 as long as segments in lateral view,

in dorsal view shining part of tyloides not reaching base or apex of segment),

tyloidea on segments 9—12 shorter. Antennal segments 11—12 distinctly

longer than broad. Hypo-epimeral area dull, densely punctate, no distinct

interstices. Mesopleura finely alutaceous, irregularly punctate, interstices

mostly smaller than punctures. Back of propodeum finely reticulato-carinate.

Petiole about 3/4 length of tergite 1. Hind tarsal segments 1—4 yellowish-

brown, apices paler. Mexico: Baja California . abbreviatus sp. nov. (p. 182)

Antennae long and slender, segments 5—8 with long narrow tyloides, in

lateral view distinctly raised in the middle. Hypo-epimeral area and

mesopleura shining, indistinctly punctate. Scutum not depressed. Back of

propodeum dull, dorsal half obliquely striate. Petiole nearly 1.5 times length

of tergite 1. Black; pronotal tubercles and all tarsi brown. Ecuador; Peru;

Suriname En Bee ee trilobatus sp. nov. (p. 218)

More segments with narrow tyloides or all tyloidea indistinct or absent, or

scutum with deep depressions and antennal segments 4—8 with tyloides

Metanotum in the middle with two small raised ‘‘wings’’, each with

membraneous yellowish-transparent margin. Mesopleura alutaceous,

punctate, interstices larger than punctures. Petiole longer than tergite |.

Antennal segments 6—11 with narrow tyloidea. Intercarinal space wider than

fore basitarsus. Apical half of tergite 1 and greater part of tergite 2 red, or

only hind margins of tergites reddish transparent. Argentina; Bolivia; Brazil;

Paraguay av RENNER joergenseni (Brethes) (p. 235)

Metanotum normal... u... 0 tan oud oe RR 13

PD!

76.

lide

78.

19.

80.

VAN LITH: New World Pluto 149

Axillae projecting backward, laterally and posteriorly not touching scutellum.

Back of propodeum coarsely reticulato-carinate. Petiole about as long as

tergite 1. Intercarinal space narrow. Gaster black ............... 76

Axillae posteriorly normally touching scutellum ................ 78

Median part of clypeal margin in frontal view bluntly protruding, distinctly

raised, exposing labrum. Clypeal margin more shining than in pygmaeus.

Vertex, scutum, scutellum and mesopleura smooth and shining, sparsely finely

punctate. Antennal segments 6—12 with narrow linear tyloidea, segment 5

with indistinct tyloides. Hind tarsi largely brown. Colombia ...........

MER BI PUL USO AO. See, PEPE facialis sp. nov. (p. 234)

Median part of clypeal margin more rounded or pointed, hardly raised . 77

Axillae in dorsal view slightly longer than broad, apex rounded, black.

Scutum, at least anteriorly, slightly alutaceous, sparsely finely punctate.

Upper half of mesopleura slightly alutaceous, distinctly punctate. Vertex

behind ocelli very finely striate. Antennal segments 5—12 or 6—12 with

narrow tyloidea, segment 13 with indistinct tyloides. Hind tarsi largely brown.

Argentina; Bolivia; Peru ....... pygmaeus pygmaeus (Brethes) (p. 230)

Axillae in dorsal view more elongate, brownish. Scutum smooth, with bluish

shine, sparsely finely punctate. Upper half of mesopleura shining, not

distinctly punctate. Vertex behind ocelli smooth. Antennal segments 6—13

with narrow tyloidea, tyloides rarely absent on segment 13. Hind tarsi

yellowish-brown. Bolivia; Brazil; British Guyana; Colombia; Surinam ...

permease WOE NE BPI DE, pygmaeus axillaris subsp. nov. (p. 233)

Scutum, mesopleura and scutellum very coarsely rugose, no distinct

punctures. Back of propodeum coarsely reticulate, enclosure shining, with

large median area. Petiole about as long as tergite 1. Antennal segments 4—11

or 4—12 with long narrow tyloides. Intercarinal space narrow. Tergites 2—3,

in specimens from the southern states also petiole and first tergite, more or

less reddish. Tibiae reddish, rarely darkened. Hind tarsal segments 1—4

yellowish. Florida, Louisiana, Maryland, Mississippi, North Carolina, South

VO SEEN KOE ee rufibasis (Malloch) (p. 186)

Scutum much less coarsely rugose. Mesopleura smooth or more or less

IR EEE vr desta Mr ee achet 79

Mesopleura and hypo-epimeral area shining, smooth, mesopleura with fine or

indistinct scattered punctures. Scutum shining, finely sparsely punctate. Back

of propodeum coarsely reticulate, upper half with parallel oblique carinae,

enclosure shining with large median area. Antennae long and slender. Gaster

black, hind margins of tergites reddish transparent .............. 80

Mesopleura distinctly punctate or striato-punctate, if somewhat shining and

very finely, very densely punctate (x 50) then also scutum densely punctate,

or mesopleura shining and one or more tergites entirely red. Antennae not

RENS ONT CIO ow ce Sa ee ee CM The ne renee à 82

Tyloidea linear, distinct on segments 4—12, shorter on segments 11—12.

Petiole as long as or somewhat longer than tergite 1. Pronotal tubercles

brown. Fore and mid femora largely dark brown, tibiae yellowish-red, tarsi

150

81.

82.

83.

84.

85.

86.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

yellowish, hind tarsi brownish. Argentina; Bolivia; Brazil; Colombia;

Ecuador; Paraguay; Peru; Surinam; Venezuela ... nitens sp. nov. (p. 210)

Tyloidea indistinct. Petiole as long as or nearly as long as tergite 1. Pronotal

tubercles whitish. Hind tarsi yellowish-white or brownish-yellow ..... 81

Fore and mid femora and tibiae yellowish-red. Antennal segments 10—12

about 1.25 times as long as broad. Venezuela .. araguensis sp. nov. (p. 221)

Fore and mid femora largely brown or black, fore and mid tibiae reddish-

yellow. Antennal segments 10—12 at least about 1.5 times as long as broad.

Peru aten e ee ee rufanalis sp. nov. (p. 220)

Mesopleura densely distinctly punctate or ruguloso-punctate, interstices,

mostly narrower than punctures. If mesopleura partly striato-punctate and

interstices on upper and/or lower part somewhat larger than punctures cf.

tibialis and albifacies; if punctation moderately dense and hind tarsi orange-

brown or yellowish-brown cf. punctatellus. Tyloidea always distinct, at least on

segments:5—=l0- aia ods tpi ts iss tee debiti ae 83

Interstices on mesopleura mostly as large as or much larger than punctures, or

punctures close but fine and very superficial (townsendi), or punctation

indistinct (marthae and metaensis). Tyloidea sometimes indistinct, or only

segments 4—8 with tyloides or tyloidea entirely absent ........... 96

At least tergite 2 entirely red. Antennal segments 4—11 with distinct tyloides.

Scutum shining, strongly punctate. Mesopleura and hypo-epimeral area

densely finely punctate and striato-punctate. Back of propodeum coarsely

reticulate, enclosure with large median area. Petiole about as long as tergite

lecci ele Mer THe amen RTT MTC RITO 84

Gaster black, hind margins of tergites, rarely also bases of segments 2—3

reddish or transparent or scutum densely punctate. If hind margin of tergite 2

and all of tergite 3 red, then back of propodeum finely rugulose (rugulosus,

No ON ener eens Oe ee ee aE, LRE 85

Hind margin of tergite 1 and all of tergite 2 red. Mesopleura and hypo-

epimeral area dull (Brazil) or shining (Argentina, Surinam). Face silvery.

Argentina; Bolivia; Brazil; Surinam ..... medius medius (Smith) (p. 200)

Tergite 1 except for two large black marks, all of tergite 2 and all or nearly all

of tergite 3 red. Face pale golden. Curacao; Venezuela naar sen

ul a ONT EEE RA NEUE PT EOE PS oe medius zuliensis subsp. nov. (p. 202)

Petiole longer than tergite 1. Mesopleura densely punctate, partly striato-

punctate. Hypo-epimeral area shining, punctate, interstices about size of

punctures. Scutum shining, densely punctate, partly transversely or oblique

rugose. Antennae slender, segments 4—13 with linear tyloides, as long as

segments, short on segment 13. Hind tarsi brown or reddish-brown.

Pubescence of face pale golden, not appressed. Ecuador .............

eae ERR ARRE TI fritzi sp. nov. (p. 199)

Petiole.at most as long'asterpite li. CER 86

Antennal segments 7—13 with tyloides, segments 10—12 longer than broad.

Scutum shining, distinctly, relatively finely punctate, interstices mostly wider

than punctures. Mesopleura more or less shining, upper half rugoso-punctate,

VAN LITH: New World Pluto 151

lower part punctate with narrow interstices. Petiole about 3/4 length of tergite

1. Hind tarsi brown. Puerto Rico; Dominican Republic; Haiti; Leeward

IslandsaVarginiIslandsit. „orn. ri. oden atricornis (Malloch) (p. 194)

Antennal segments 4—5, 5—6 or 6 too with distinct linear tyloides .... 87

. Antennal segments 5—10 or 6— 10 with distinct lineartyloides ...... 88

Alsosesmentsilil#12orl3withtyloides to. MMA 2... Pao 89

. Antennal segments 5—10 with black tyloides, sometimes also a short tyloides

on segments 4 and 11. Intercarinal space narroa. Mesopleura dull, closely

coarsely punctate, interstices on upper or lower part partly larger t an

punctures. Petiole about 3/4 of length of tergite 1. Hind tarsal segments 1—4

whitish. District of Columbia, Florida, Louisiana, Missouri, South Carolina,

Tennessee, Texas; Virginia ove scot var tibialis (Cresson) (p. 188)

Antennae more slender, segments 5—10 or 6—10 with brown or black

tyloides. Intercarinal space nearly as broad as fore basitarsus. Mesopleura

much finer densely punctate, not distinctly striato-punctate. Petiole almost as

long as tergite 1. Bases of hind tarsal segments 2—4 pale brown. Argentina;

Broliviamsrazil paren. … aen à à stramineipes sp. nov. (p. 204)

Antennal segments 5—12 with tyloides, small on segment 12; segments 11—12

little longer than broad or almost quadrate. Mesopleura and hypo-epimeral

area densely ruguloso-punctate. Scutum shining, moderately densely

punctate, posteriorly mostly striato-punctate. Petiole slightly over half length

of tergite 1. Hind tarsi brown. Cuba; Jamaica; Mexico; Nicaragua .......

EPR Ee es AES. ii col, argentifrons (Cresson) (p. 192)

Antennal segments 4—11, 4—12 or 4—13 with tyloides. Mesopleura often

partly with distinct interstices or antennal segments at least 1.5 times as long

as broad or hind tarsi whitish. Petiole longer .................. 90

. All tarsi brown or dark brown. Petiole about 3/4 length of tergite 1.

Mesopleura and hypo-epimeral area densely punctate and striato-punctate.

Antennal segments 3—12 or 4—12 with distinct tyloides, segment 13

Sometumesmthvasinallsone team WA Ja al Alta MO 91

Hind tarsi whitish; if yellowish-brown or orange-brown then petiole as long as

tergite | and segments 4—11 with distinct tyloides .............. 92

Pronotal tubercles and gaster black. Underside of flagellum dark brown, last

segment reddish-brown. Antennal segments 4—12 with linear tyloides,

distinctly raised; segments below regularly convex (lateral aspect).

Mesopleura and hypo-epimeral area dull. Scutum dull, punctate, interstices

smaller than punctures. Back of propodeum coarsely reticulato-carinate.

Female unknown. New Mexico, Texas ..... castaneipes sp. nov. (p. 197)

Pronotal tubercles white. Tergite 3 red. Underside of flagellum orange-brown.

Antennal segments 3—12 or 4—12 with fine linear tyloides, not distinctly

raised. Mesopleura somewhat shining. Scutum very shining, coarsely

punctate, partly rugoso-punctate. Back of propodeum very finely rugulose.

Femaleunknowny Texasır van... toes oa. rugulosus sp. nov. (p. 197)

Antennae slender; tyloidea on segments 4—11 distinct, on segment 12

indistinct. Hind tarsi orange-brown, bases of segments more or less darkened.

152

98:

94.

95:

96.

IM:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Petiole about as long as tergite 1. Mesopleura alutaceous or shining,

moderately densely finely punctate, interstices size of punctures, partly

larger. Hypo-epimeral area faintly shining, finely punctate, interstices

distinct. Back of propodeum coarsely reticulate. Pronotal tubercles brown or

yellowish. Costa Rica; El Salvador; Guatemala; Mexico .............

Be ee EBER punctatellus sp. nov. (p. 179)

Antennal segments 4—12 or 4—13 with distinct tyloides. Hind tarsal segments

1==4 yellowishewhite En Mahe ee Pe eee ee 93

Petiole about as long as tergite 1. Antennal segments 4—12 with tyloides.

Pronotal'tubercles whitish i.) N. u We aes ee ee 94

Petiole 1/2 to 3/4 length of tergite 1. Antennal segments 4—12 or 4—13 with

tyloides. Pronotal tubercles brown or whitish ................. 95

Back of propodeum moderately finely reticulate. Punctation of mesopleura

fine, interstices on upper part larger than punctures, on lower part smaller.

Scutum dull, densely not coarsely punctate, punctures partly in rows.

Antennal segments 4—12 with black tyloides, short on segment 12. Iowa,

Texasa eat in he LEEDS ae albifacies (Malloch) (p. 187)

Back of propodeum very coarsely reticulate, enclosure with large shining

median area. Mesopleura and hypo-epimeral area dull, upper half or median

part of mesopleura moderately coarsely striato-punctate, interstices on lower

part may be partly wider than punctures. Scutum strongly punctate, with faint

transverse rugae. Hind tarsal segments 2—4 basally somewhat infuscated.

Texas, North Carolina; Mexico; Belize .... aerofacies (Malloch) (p. 183)

Pronotal tubercles black or brown. Petiole about 1/2 to 2/3 length of tergite 1.

Antennal segments 4—13 with tyloides, on segments 7—12 shorter than

segment, on segment 13 very short; segments 7—10 in lateral aspect raised

near the middle. Mesopleura dull, closely punctate. Upper half of back of

propodeum finely, mostly parallel, carinate. Intercarinal space narrow.

Mexico: verie het IRRE tara ER alphitopus sp. nov. (p. 195)

Pronotal tubercles whitish. Petiole about 3/4 length of tergite 1. Antennal

segments 4—12 with tyloides. Mesopleura with large, mostly contiguous

punctures, interstices, if any, lineolate, on upper part sometimes as large as

punctures. Occipital and hypostomal carinae almost contiguous. Back of

propodeum more coarsely reticulato-carinate. Florida, Georgia, North

Carolina etn AN arenivagus Krombein (p. 190)

Hypo-epimeral area distinctly punctate. Propodeal enclosure irregularly

carinate. Back of propodeum finely reticulato-carinate. Pronotal tubercles

whitish: Hind:tarsilargely brown fait a. 068 eee ee ee 97

Hypo-epimeral area almost smooth and shining, punctures indistinct.

Propodeal enclosure sometimes with large shining median area ...... 98

Antennal segments 5—12 with linear tyloides, short on segment 12. Hypo-

epimeral area dull, finely but distinctly punctate. Mesopleura finely

alutaceous, somewhat shining, distinctly punctate, interstices wide on upper

part (in Arizona specimens punctures on mesopleura coarser, interstices

about size of punctures or slightly larger and mesopleura and hypo-epimeral

98.

99.

100.

101.

VAN LITH: New World Pluto 153

area shining). Gaster black or tergites 2—3 entirely or partly red. Alabama,

Arizona, California, District of Columbia, Florida, Iowa, Kansas, Louisiana,

Missouri, New Mexico, North Carolina, South Carolina, Tennessee; Cuba; El

Salvador; Mexico; Nicaragua ...—...r..... sayi (Rohwer) (p. 173)

Antennal segments 4—8 with long linear tyloides. Hypo-epimeral area and

mesopleura shining, hypo-epimeral area finely punctate, mesopleura with

strong punctures, interstices about size of punctures. Scutum strongly

punctate, with longitudinal median depression and a triangular depression in

front of scutellar suture. Gaster black. Female unknown. Mexico .......

10 0.0 Au depressus sp. nov. (p. 181)

Antennae without tyloidea, segments 10—12 about 1.5 times as long as broad.

Mesopleura shining, finely punctate, interstices about four times size of

punctures. Hypo-epimeral area shining with a few minute hair-bearing

punctures. Back of propodeum coarsely reticulate. Petiole about as long as

tergite 1. Posterior half of tergite 1, all of tergite 2 red. Pronotal tubercles

whitish. Hind tarsal segments brown with paler apices. Female unknown.

Venezuelaanse De ne aes incarinatus sp. nov. (p. 222)

Antennae with fine, sometimes indistinct, linear tyloides. Mesopleura

somewhat dull, rarely shining, punctures indistinct or gaster black .... 99

Petiole shorter than tergite 1. Hind tarsi whitish or yellowish-white, at least

segments I—2, or bases of segments dorsally brown. Scutum shining, finely

puiciaie rer dn nee en a re a 100

Petiole somewhat longer than tergite |. Hind tarsi brown. Scutum shining,

strongly punctate. Back of propodeum, also laterally, coarsely reticulato-

carinate. Mesopleura somewhat dull, no distinct punctures ....... 101

Antennal segments 3—12 about twice as long as broad, segments 4—9 or

4—10 with fine linear tyloides, segment 13 also ventrally dark. Back of

propodeum dull, more closely reticulate, dorso-lateral carinae indistinct,

enclosure finely coriaceous. Mesopleura densely superficially punctate.

Gaster blackish-brown, hind margins of tergites reddish, often also base of

tergite 3. Hind tarsi yellowish or bases of segments dorsally brown. Ecuador;

Er. ae e ee townsendi (Cockerell) (p. 207)

Antennal segments 3—12 about 1.5 times as long as broad, segments 6—11

with very narrow pale tyloides, underside of flagellum orange-yellow. Back of

propodeum coarsely reticulato-carinate, with large shining median area.

Mesopleura more distinctly alutaceous, sparsely finely punctate. Gaster

black. Hind tarsal segments 3—5 pale brown. Mexico ...............

ee Se adele id. NA IN ae it arie evansi sp. nov. (p. 184)

Tergites 2—3 and at least sternites 2—3 reddish with more or less distinct

brown band before apical margin of tergites, remaining tergites with

transparent reddish hind margin. Antennal segments 4—10 or 4—11 with

indistinct, not shining, linear tyloides. Scutum, at least on anterior part,

weakly rugoso-punctate. Vertex behind ocelli dull, finely transversely striate.

Bolivia; Brazil; Colombia; Ecuador; Peru ..... marthae sp. nov. (p. 208)

Gaster black, segments with narrow reddish hind margins. Antennal segments

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

4—11 with distinct, shining, fine linear tyloides. Scutum more shining,

punctures separate. Vertex behind ocelli smooth and shining. Colombia

KOP AVE i ER D SL NER dn re a ae metanus sp. nov. (p. 210)

Group of /ongiventris

Pluto longiventris (Malloch)

(figs. 1—2)

Malloch, 1933: 52, 9 and g (Psenia longiventris; Arizona, California).

Krombein, 1951: 962 (Pluto longiventris, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length about 9 — 9.5 mm. Black. Apical 2/3 of mandibles yellowish-

red with darker tips. Underside of flagellum pale yellowish-white, base of third

antennal segment and underside of apical three segments more or less darkened.

Pronotal tubercles and tegulae yellowish-white. Veins of wings brown, stigma

paler. Foreside of fore tibiae with incomplete yellowish-white streak; base of mid

and hind tibiae and all tarsi except apex of last segment of hind tarsi yellowish-

white. Apical margins of tergites I—5 and of sternites reddish transparent.

Clypeus dull, median part of margin almost straight with small lateral teeth (fig.

2), before apical margin somewhat raised and shining. Distance between clypeal

teeth about half total width of margin. Frons before median ocellus densely

punctate, upper part of frons and vertex almost impunctate. Intercarinal space

about as wide as fore basitarsus (fig. 1). Third antennal segment over 2.5 times as

long as broad, segments 8—11 about quadrate, segment 12 over 1.5 times as long

as broad.

Pronotal angles in frontal view with short lobe, in dorsal view with sharp angle.

Scutum shining, punctures distinct, somewhat sparse, near fore and hind margins

more closely placed. Prescutal sutures continued to hind margin as a row of

punctures. Scutellum shining, sparsely finely punctate. Metanotum dull. Enclosed

area of propodeum shining, with large median area. Back of propodeum dull,

reticulato-carinate, carinae laterally behind enclosed area more parellel;

longitudinal groove narrow. Anterior oblique suture broad, coarsely foveolate.

Hypo-epimeral area coarsely longitudinally rugoso-punctate. Upper part of

mesopleura with long longitudinal rugae, posterior margin finely striate, lower part

reticulate alutaceous and with rather large sparse punctures. Gaster shining,

densely finely punctate. Petiole about half as long as first tergite. Pygidial area at

least twice as long as broad, elliptic, apex rounded. Second recurrent vein of fore

wings ending in third submarginal cell.

Pubescence of face silvery, of pygidial area brown and appressed, entire last

segment also with long erect brown hairs, pubescence of rest of body whitish.

Apical margins of sternites 2—5 with a few long erect pale hairs.

Male. — Length about 7—9 mm. Resembling female. Underside of flagellum

and all of last segment orange-brown. Clypeal margin rounded, medially with two

VAN LITH: New World Pluto 155

Figs. 1—2. Pluto longiventris (Malloch), 9; 1, head in ventral aspect; 2, clypeus. Figs. 3—4. P. angulicor-

nis (Malloch); 3, antenna of 3; 4, clypeus of 9. Figs. 5—8. Clypeus; 5, of P. pallidistigma (Malloch), 9,

allotype; 6, P. rotundus sp. n., 9, holotype; 7, P. suffusus (Fox), 9 ; 8, P. texanus (Malloch), 9, paratype.

Fig. 9. P. brevipetiolatus (Rohwer), 9, pygidial area. Fig. 10. P. littoralis (Malloch), 3, Cedar Key, left

paramere, dorsal aspect.

156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

indistinct teeth. Antennal segments 4—11 about 1.5 times as long as broad,

segment 12 somewhat shorter, segment 13 about twice as long as broad. Segments

6—12 with oval tyloidea, gradually increasing in size, on segment 12 about half as

long as segment; segments in lateral view angular. Segment 13 with short tyloides.

Lateral pronotal angles less sharp, in dorsal view about rectangular. Scutum

shining, coarsely punctate with short transverse rugae. Scutellum shining, a few

coarse punctures. Back of propodeum coarsely reticulato-carinate. Mesopleura

and hypo-epimeral area entirely coarsely rugose, mesosternum with fine separate

punctures. Gaster slender, petiole about as long as first tergite, this tergite over

twice as long as broad.

Material examined: 1 9, allotype, Gilbert, Arizona, 29 Z, paratypes, Higley and

Mount Superstition, Arizona; Lindsay, El Centro and Holtville, California

(USNM, type Nr. 44222); also 2 & from Higley and 1 © from Holtville without

type labels.

New records from USA: Arizona: 1 g, Atascosa Mts., Santa Cruz Co., 30 June

1953, R. S. Beal (CIS); 1 g, 10 mi southwest Patagonia, Santa Cruz Co., 13 Sept. |

1958, M. A. Cazier (CIS); 1 3, Tucson, 18 June 1938, R. H. Crandell (KVK); 1 &,

Yuma, 16 July 1953, on cotton, coll. Hago (USNM); I 9, Portal, 17 Aug. 1974, H.

and M. Townes (HT).

California: Butte Co.: | &, Chico, 13 July 1965, T. R. Haig (CSC); Glenn Co.: 2

3, Willows, 16 June 1970, D. Schult (CSC); Imperial Co.: 1 &, Calipatria, 14 Aug.

1916, F. A. McGregor; 1 9, Holtville, 25 Sept. 1916, F. A. McGregor; 5 9, 4—6

June 1912, 1 g, June 1911, 2 g, 29 May 1912, Experiment Farm, of which I © and

1 g on Helianthus annuus, “Vtg. gland. hairs of lvs” (visiting glandular hairs of

leaves), J. C. Bridwell (USNM); 1 3, Bard, 17 Aug. 1965, ex Gossypium hirsutum,

H. Ray, Pluto longiventris Malloch det. R. M. Bohart (CSC); Riverside Co.: 1 3,

Romaland, 17 Aug. 1946, J. W. MacSwain (USNM); San Mateo Co.: | 9, Palo

Alto, F. Grinnell Jr. (USNM); Stanislaus Co.: 1 ©, 10 mi southwest Turlock, 25

Aug. 1956, R. R. Snelling (CSC); Tulare Co.: 1 &, 24 May 1947, 5 g, 14—30 June

1947, Wood L., in Rotary trap, Norman W. Frazier (USNM).

First records from Mexico: Baja California: | g, Sierra de Juárez, Santa

Catarina foothills, 43 mi east Ensenada, 3600 ft, 18 Aug. 1962, on Adolphia

california, D. E. Breedlove (CAS). Guerrero: 2 9, Zumpango, 22 July 1963, F. D.

Parker and L. A. Stange (UCD). Jalisco: 1 9, 3 mi north Barra de Navidad, 100 ft

above Hotel Melanane, 14 Febr. 1966, D. Bolinger (OSU). Michoacan: 3 g, 11 mi

east Apatzingan, 20 Aug. 1954, E. G. Linsley, J. W. McSwain and R. F. Smith

(UCD). Oaxaca: | 9 and 1 Z, Tehuantepec, 15—16 July 1964, Paul J. Spangler

(USNM). Puebla: 8 3, 3 mi north Petlalcingo, 21 Aug. 1963, F. D. Parker and L.

A. Stange (UCD). Sinaloa: Mazatlan, 10 ft, 1 9, 19 July 1959, H. E. Evans (CU), 1

3, 10 May 1961, at light, Howden and Martin (CNC); 1 &, 5 mi north Mazatlan,

15 Aug. 1970, Malaise trap 10A-2P, M. Wasbauer and J. Chemsak (CIS). Sonora: |

3, 5 mi south Magdalena, 25 May 1962, F. D. Parker and L. A. Stange, 2 ©, 5 mi

east Navojoa, 9 Sept. 1970, R. M. Bohart (UCD).

The female of P. longiventris is easily recognized by the sharp angles of the

VAN LITH: New World Pluto 157

pronotum and the rugae on the mesopleura, the male by the coarsely rugose

mesopleura and the oval tyloidea.

The males from Mexico often have a more coarsely sculptured scutellum,

sometimes this is rugoso-punctate.

Group of angulicornis

Pluto angulicornis (Malloch)

(figs. 3—4)

Malloch, 1933: 58, © and ¢ (Psenia angulicornis; Texas).

Krombein, 1951: 962 (Pluto angulicornis, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length about 5 mm. Head and thorax black; mandibles and labrum

reddish. Underside of flagellum yellowish-red. Pronotal tubercles yellowish-white,

tegulae entirely reddish or partly yellowish-white. Fore tibiae and tarsi largely

reddish, tibiae with yellowish-white streak on outer side, first tarsal segments

yellowish-white. Mid tibiae reddish-brown with yellowish base and yellowish-

white streak on outer side, tarsal segments 1—3 yellowish-white, segments 4—5

brown. Basal third of hind tibiae and tarsal segments 1—4 yellowish-white, last

segment brown. Gaster largely black, hind margin of tergite 1, all of tergites 2—3

and sternites 2—4 red, or tergite 3 and sternites 3—4 largely brown.

Protruding median part of clypeal margin about 1/3 of total width of margin, this

part almost straight, lateral angles obtuse, no distinct teeth (fig. 4). Frons except

for lateral parts densely punctate. Vertex more or less shining, finely reticulate

alutaceous, distinctly punctate, interstices larger than punctures. POD about 1.5

times as long as OOD. Occipital carina fine but complete, also on ventral side of

head (Malloch: ‘“‘evanescent as it approaches central line of ventral surface”).

Intercarinal space narrow. Antennae clavate segments 9—11 much shorter than

long.

Lateral pronotal angles about rectangular. Scutum and scutellum shining, finely

punctate, interstices as large as or a few times size of punctures. Median part of

propodeal enclosure irregularly reticulate, back of propodeum rather finely

reticulato-carinate, dorso-lateral parts finely striate. Hypo-epimeral area

somewhat glossy, very finely but distinctly punctate, interstices about size of

punctures or larger. Mesopleura finely reticulate alutaceous, finely punctate,

interstices mostly not larger than punctures, interstices larger on upper part;

posterior part of mesopleura and of hypo-epimeral area finely coriaceous. Petiole

about half as long as first tergite, this tergite about as long as broad. Pygidial area

at most 2.5 times as long as broad. Second recurrent vein of fore wings ending in

third submarginal cell.

Face silvery, mostly appressed, pubescent, pygidial area with golden-brown

pubescence and also with long erect hairs, rest of body whitish to greyish

pubescent.

Male. — Resembling female. Hind margin of first tergite and bases of tergites

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

2—3 more or less red. Antennal segments 7—10 with dark and rather broad

tyloides, shorter than segments and almost angularly raised in the middle of the

segments (fig. 3); segments 4—5 with small linear tyloides which may be very

indistinct, segment 6 with long oval low tyloides, segments 11—12 with linear,

indistinct tyloides, a short one on segment 13. Petiole about 2/3 length of first

tergite, this tergite about 1.5 times as long as broad.

Material examined: Texas, 1 9, allotype and | 9, paratype, Plano, July 1907,

one in oatfield and one in wheatfield, E. S. Tucker, 1 9, paratype, Plano, August

1907, E. S. Tucker, 1 ¢, paratype, Brownsville, J. C. Bridwell (USNM, type No.

44229). The paratype from Neuecest, Texas, 28 April 1896, coll. Marlatt,

erroneously recorded by Malloch as a male, is a female and different from the

allotype. The punctation of the mesopleura is wider, the hypo-epimeral area is

more coarsely punctate, the scutum is distinctly alutaceous and the base of the

hind basitarsus is brown. Most likely it is a rather small specimen of P. basifuscus.

New records from USA: Texas: | 9, 16 May 1945, Laredo, inside airport:

building, with label “Pluto angulicornis Mall., det. Tow”. Iowa: 2 8, 25 Aug. 1935,

H. E. Jaques (CNC). Louisiana: 1 9 and I 3, Tallulah, 25 June 1948, Bug Catcher

Exp., R. C. Gaines (USNM). New Mexico: 1 &, 5.4 mi northeast Santa Rosa, 10

July 1929, on Salsola pestifer, V. E. Romney (USNM).

The female of P. angulicornis is very similar to that of P. basifuscus, but its

scutum and scutellum are shining and the punctation of the mesopleura is much

finer. The tyloidea of the males of the two species are distinctly different.

Pluto pallidistigma (Malloch)

(fig. 5)

Malloch, 1933: 52-53, © and Z (Psenia pallidistigma; Arizona, Texas).

Krombein, 1951: 962 (Pluto pallidistigma, comb. nov.).

Bohart & Menke, 1976: 171.

Male. — Length about 6—7 mm. Head and thorax black. Labrum and

mandibles yellowish-red. Antennae brown, underside of flagellum yellowish-red.

Pronotal tubercles, tegulae and stigma of wings whitish, stigma sometimes yellow-

ish-brown or central part brownish. Veins of wings pale brown, at base whitish,

wings hyaline. Fore and mid tibiae reddish-brown, outer side of tibiae and all of

tarsi yellowish-white. Basal 2/5 of hind tibiae and tarsal segments 1—5 whitish, last

segment slightly darkened. Hind margin of first gastral tergite, all of tergites 2—3

or 2—4 red, sometimes tergite 2 largely darkened. Ventral plate of petiole,

sternites 2—4 and apex of last segment reddish, often most of sternites darkened.

Median part of clypeus slightly protruding. Frons densely punctate, vertex more

sparsely so, more or less micro-sculptured, back of head somewhat transversely

striate. Lower part of occipital carina unusually high, directed obliquely back-

ward. Intercarinal space broader than first basitarsus. Antennae long and slender,

last segment 2.5 times as long as broad, segments 6—12 with distinct tyloides,

VAN LITH: New World Pluto 159

much shorter than segments, on segments 6—9 oblong, somewhat widening

towards apex, more broad oval on segments 10—11, narrower on segment 12.

Lateral pronotal angles about rectangular. Scutum and scutellum shining,

strongly punctate, interstices mostly a few times size of punctures. Propodeum

including enclosed area densely irregularly reticulato-carinate. Mesopleura

shining, coarsely strongly punctate, interstices partly smaller than punctures,

partly somewhat larger, hind margin coriaceous. Hypo-epimeral area shining,

punctures finer and closer, a few large interstices. Anterior plate of mese-

pisternum shining, densely coarsely punctate, oblique carina crenulate. Gaster

slender, petiole as long as first tergite, this tergite about twice as long as broad.

Second recurrent vein of fore wings ending in third submarginal cell.

Pubescence of head and thorax silvery, on face dense and mostly appressed,

gaster greyish-white pubescent.

Female (as associated with preceding male by Malloch). — Colour as in male.

Hind margin of first tergite, all of tergites 2—3 and sternites 2—3 red. All tarsi

entirely whitish, except for extreme base of hind basitarsus.

Median part of clypeal margin slightly rounded, almost straight (fig. 5), less than

1/3 of total width of margin. Lower part of occipital carina not unusually high,

intercarinal space narrower than first basitarsus. Antennae short, gradually

widening towards apex, segments 9—11 over 1.5 times as broad as long. Scutum

and scutellum somewhat shining, very finely reticulate alutaceous, sparsely

punctate. Propodeal enclosure somewhat shining, irregularly reticulate, back dull,

very finely and closely reticulato-carinate, dorsolaterally with oblique parallel

carinae. Mesopleura distinctly reticulate alutaceous, finely punctate, interstices a

few times size of punctures, hind margin coriaceous. Hypo-epimeral area dull,

densely distinctly punctate. Anterior plate of mesepisternum dull, finely punctate,

on lower part finely vertically striate. Petiole about half as long as first tergite, this

tergite longer than broad. Pygidial area about 2.5 times as long as broad, densely

golden-brown pubescent. -

Material examined: 1 &, holotype, Arizona, Mt. Superstition near Higley, 24

July 1917, E. G. Holt, labelled “ Psenia pallidistigma type det. J. R. Malloch’; 1 ©,

allotype, Texas, Cotulla, 11 May 1906, J. C. Crawford (USNM, type No. 44223).

New record from Arizona: 1 ©, Portal, 23-31 July 1959, on flowers of Acacia, K.

V. Krombein (KVK).

New records from Texas: 1 &, Big Bend National Park, Nine Point Draw, 3000

ft, 20 May 1959, 2 g, Fort Davis, Point Rocks, 5000 ft, 29—30 May 1959, W. R. M.

Mason (CNC).

First records from California: Imperial Co.: 1 g, Experiment Farm, 21 May

1912, v(isi)t(in)g gland(ular) hairs of I(ea)v(e)s Helianthus annuus, J. C. Bridwell

(USNM); Inyo Co.: 2 g, Ballarat, Panamint Valley, 10 June 1961, H. F. Howden

(CNC); San Bernardino Co.: 1 Z, 9 air mi south Baker Zzyzx Sprs., 20—21 April

1977, M. Buegler (CIS).

There is no proof, as yet, that the male and the female recorded from different

states were correctly associated by Malloch, although a similar female has now

160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

also been recorded from Arizona. The differences between the sexes are con-

siderable. I could not find any structural differences between the female studied

by Malloch and the females from Texas identified by me as basifuscus sp. nov.; the

latter were collected with some male specimens of basifuscus. The only

distinguishing character of the females seems to be the white colour of the hind

tarsi, including the last segment, of pallidistigma. This is not a very reliable

character, however, and possibly the true female of P. pallidistigma has not yet

been recognized. The males of P. pallidistigma and P. basifuscus are easily

separated.

Pluto basifuscus sp. nov.

Male. — Length about 6 mm. Black. Mandibles and labrum dark reddish.

Underside of flagellum brown, last segment reddish-brown. Pronotal tubercles and

tegulae whitish. Foreside of fore femora and of fore tibiae yellowish-brown, tibiae

with yellow outer streak, tarsi whitish, last segment brown. Mid tibiae brown, base —

and tarsal segments 1—4 whitish. Hind tibiae with narrow whitish basal ring, tarsal

segments I—4 whitish, basal 1/4 or 1/3 of basitarsus brown. Veins of wings brown,

stigma sometimes paler. Hind margins of tergites more or less reddish.

Median part of clypeal margin slightly rounded. Frons strongly punctate,

somewhat raised in front of posterior ocelli. Vertex shining or finely alutaceous,

sparsely punctate. POD much larger than OOD. Underside of head finely striate.

Ventral part of occipital carina of normal height, intercarinal space narrow.

Antennae slender, segments longer than broad; segments 5—6 with short narrow

tyloides, segments 7—12 with broad oval tyloides, about 1/3 of length of segment, a

smaller or indistinct tyloides on segment 13.

Lateral pronotal angles obtuse. Scutum shining, densely punctate, interstices

mostly larger than punctures. Scutellum shining, sparsely finely punctate.

Propodeal enclosure and back of propodeum irregularly finely reticulato-carinate.

Mesopleura reticulate alutaceous, almost shining, coarsely punctate, interstices

mostly about size of punctures, on upper part somewhat larger with also some

micropunctation. Hypo-epimeral area dull, densely punctate. Anterior plate of

mesepisternum striato-punctate. Gaster slender, petiole almost as long as first

tergite, this tergite about twice as long as broad. Second recurrent vein of fore

wings ending in third submarginal cell.

Pubescence of face silvery, mostly appressed, of rest of body whitish. Wings

whitish pubescent.

Female. — Normally basal 1/3 of hind basitarsus and greater part of tarsal

segment 5 brown. For further description cf. P. pallidistigma (Malloch).

U.S.A.: Texas: 10 mi west Fort Davis, Pt. Rocks, 5000 ft, 1 &, holotype and I 9,

allotype, 30 May 1959. Paratypes: 3 4 and 6 9, 29—30 May 1959; Fort Davis,

Limpia Cn., 5000 ft, 1 9, 28 May 1959, W. R. M. Mason (CNC); San Diego, 1 9,

24 May, Ashmead (USNM); Uvalde Co., 1 g, Speir Rch. 3 mi northwest Uvalde, |

May 1977, Malaise trap 10A—5P and | 4, Nueces Riv. 12 mi south Uvalde, 30

April 1977, Malaise trap 12M-4P, T. Eichlin and M. Wasbauer (CSC). Arizona:

VAN LITH: New World Pluto 161

2 &, Douglass, July 1940, D. G. Hale; 1 9, Cochise Co., San Bernardino Rch.,

10—11 June 1968, Menke and Flint (USNM); 2 © and 8 &, Portal, 10—25 Aug.

1974, 2 &, Parker Canyon Lk., 22—23 Aug. 1974, 1 9 and 1 &, McNeal, 24 Aug.

1974, all H. and M. Townes (HT). New Mexico: 3 ©, Hatch, 283—30 Aug. 1974, H.

and M. Townes (HT). California: 1 3, La Mesa, 8908 Lemon Avenue, 9 Aug. 1958,

F. X. Williams (CAS).

Mexico: | &, 17 mi north Fresnillo, Zacatecas, 16 July 1954, J. W. McSwain.

Chihuahua: 1 g, 10 mi north Jiménez, 21 Sept. 1970, R. M. Bohart. Oaxaca: 1 &,

coll. Crawford (UCD); this male has finer punctate mesopleura. Sinaloa: 1 &,

Mazatlan, 15—20 Aug. 1962, H. E. Evans (MCZ). Baja California: 1 9, San Pedro,

7 Oct. 1941, Ross and Bohart; I g, Big Canyon, Sierra Laguna, 13 Oct. 1941, Ross

and Bohart (CAS). All paratypes.

P. basifuscus is apparently a variable species. A female and a male from Texas

have a broad intercarinal space. In further respects they are identical with the rest

of the material and it does not seem likely that they belong to different species.

The gaster of the female from San Diego is darker, tergites 2—3 having a black

mark which covers the greater part of the disk. In three of the females and in one

of the males from the type-locality, as well as in the two females from Portal,

_ Arizona, also the hind tarsal segments 2—4 are more or less brown, the basitarsus

for the greater part so. In the two males from Arizona the underside of the

flagellum is orange-brown.

Because of the similarities between the basifuscus females with pale hind tarsi

and the single female from Texas, which Malloch associated with the male type of

pallidistigma from Arizona, further studies are much needed. That in Texas males

of pallidistigma were collected on the same spot and on the same date as basifuscus,

the latter species being represented by females as well as males, makes matters

even more confusing. Series of both sexes of pallidistigma, collected flying together

at the type-locality in Arizona, might enable us to solve this problem.

The male of P. basifuscus much resembles the male of P. spangleri, which has

smaller tyloides and a more slender gaster. Confusion of the male of P. basifuscus

with P. pallidistigma is hardly possible as the former has no high occipital carina

like P. pallidistigma. The specific name basifuscus refers to the dark base of the

hind basitarsi of both female and male.

Aberrant form.

A male from Mexico, Baja California, Aqua Caliente (San Carlos), 18.5 km east

of Maneadero, 6 July 1973, P. H. Arnaud (CAS) is extremely dark. The mid tibiae

have no white on their outer side; all tarsi are brown, hind tarsi dark brown with

paler tips. The tyloidea on antennal segments 5—6 are broader towards the apex of

the segments; tyloides on segment 7 long oval, all almost as long as segments,

tyloidea on segments 8—12 broad oval, about half length of segment. The

mesopleura are more finely punctate than in the other males; hypo-epimeral area

finely but distinctly punctate, interstices about size of punctures.

Although provisionally considered an aberrant form of P. basifuscus, this spec-

imen may represent a distinct species.

162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pluto minutus (Malloch)

Malloch, 1933: 59, 9 (Psenia minuta; Texas).

Krombein, 1951: 962, 9 (Pluto minutus, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length 5 mm. Head and thorax black; anterior margin of clypeus

dark reddish, mandibles and labrum yellowish-red, underside of flagellum orange-

brown. Pronotal tubercles and tegulae yellowish-white. Fore tibiae and tarsi

yellowish, underside of tibiae brown. Base and outer side of mid tibiae yellowish,

tarsi yellowish-white. Basal third of hind tibiae yellowish-white, tarsal segments

1-4 whitish. Gaster black. Veins of wings brown.

Median part of clypeal margin very slightly emarginate, almost straight, about

1/3 of total width of margin, lateral angles obtuse. Frons densely punctate. Vertex

finely reticulate alutaceous, sparsely finely punctate. POD larger than OOD.

Intercarinal space more than half as wide as first basitarsus. Underside of head

shining, sparsely punctate. Antennae clavate, segments 10—11 shorter than broad,

last segment about 1.5 times as long as broad.

Lateral pronotal angles sharp. Scutum and scutellum finely reticulate aluta-

ceous, finely punctate, interstices mostly a few times size of punctures. Propodeal

enclosure shining with large median area. Back of propodeum dull, finely

coriaceous and with extremely fine oblique striae, no distinct reticulation.

Mesopleura finely reticulate alutaceous, medially a few shallow relatively large

punctures, lower part sparsely finely punctate, extreme upper part almost im-

punctate. Hypo-epimeral area densely superficially punctate, interstices about size

of punctures. Anterior oblique suture crenulate, carinae somewhat continued on

anterior plate of mesepisternum. Second recurrent vein of fore wings ending in

third submarginal cell. Petiole slightly shorter than half first tergite, this tergite

about 1.5 times as long as broad. Pygidial area over 2.5 times as long as broad.

Face with silvery appressed pubescence and also with long erect hairs. Pronotal

collar appressed silvery pubescent, last tergite golden-brown pubescent, densely

so on pygidial area, rest of body whitish pubescent.

Male unknown.

Material examined: 1 9, holotype, Texas, San Diego, 16 May, collection

Ashmead, labelled: ““Psenia minuta type det. J. R. Malloch” (USNM, type No.

44230).

New record from Texas: | ©, Sabinal, May 1910, F. C. Pratt (USNM).

The back of the propodeum of this species is unusually finely striate, without

distinct reticulation.

Pluto spangleri sp. nov.

Female. — Length about 6 mm. Black. Middle part of mandibles, and labrum

reddish. Underside of flagellum orange-brown. Pronotal tubercles and tegulae,

foreside of fore tibiae and outer side of mid tibiae, basal 1/3 of hind tibiae and

VAN LITH: New World Pluto 163

segments I—4 of all tarsi whitish. Hind margins of tergites reddish transparent.

Veins of wings brown.

Median part of clypeal margin straight, about 1/3 of total width of margin, no

distinct lateral teeth, cf. fig. 5 (pallidistigma). Frons densely finely punctate. Vertex

reticulate alutaceous, sparsely finely punctate. Intercarinal space more than half

as wide as first basitarsus, rarely narrower. POD larger than OOD. Antennae

clavate, segments 9—11 shorter than broad.

Lateral pronotal angles sharp in dorsal aspect. Scutum finely reticulate

alutaceous, finely punctate, interstices larger than punctures, laterally much

larger. Scutellum with same microsculpture, sparsely finely punctate. Propodeal

enclosure almost shining, central area large, sometimes with an irregular carina.

Back of propodeum dull, with oblique and transverse fine striae and some

reticulation. Mesopleura finely reticulate alutaceous, punctures shallow, on

central part interstices mostly larger than punctures, on lower part and on

mesosternum a few times size of punctures. Hypo-epimeral area finely punctate,

interstices about size of punctures. Anterior oblique suture foveolate, rugae

continued on anterior plate of mesepisternum. Second recurrent vein of forewings

ending in third submarginal cell. Petiole about half as long as first tergite, this

tergite about 1.5 times as long as broad. Pygidial area about 2.5 times as long as

broad.

Face with silvery-whitish appressed pubescence and long erect hairs. Rest of

body silvery-whitish pubescent, on vertex and on scutum more yellowish-grey; last

tergite including pygidial area with backward directed brown hairs.

Male. — Resembling female, more slender. Length about 6 mm. Fore tibiae

yellowish-brown, mid tibiae largely black with yellowish-brown outer streak,

extreme base of hind tibiae yellowish-white. Veins of wings brown, stigma dark

brown.

Clypeal margin with two indistinct teeth, close together. Antennal segments

11—12 about 1.5 times as long as broad, segments 5— 12 with tyloidea, narrow and

shorter than segments on segments 5—6 or 5—7, small and oval on segments 7— 12

or 8—12, indistinct on segment 13. Punctation of mesopleura stronger and denser,

dorsal half anteriorly somewhat striato-punctate. Scutum shining, punctation

stronger than in female. Back of propodeum coarsely reticulato-carinate. Petiole

somewhat shorter than first tergite, this tergite over twice as long as broad; second

tergite about 1.75 times as long as broad.

Mexico: Oaxaca: Tehuantepec, 1 ©, holotype, 15—16 July 1964, 1 g, allotype,

23 July 1964, Paul J. Spangler (USNM). Paratypes: Oaxaca: Tehuantepec, | 9,

15—16 July 1964, 1g, 23 July 1964, Paul J. Spangler (USNM); 3 mi north

Huajuäpan de Léon, | 9, 8 Sept. 1959, R. H. and E. M. Painter (KU). Chihuahua:

29 and 2g, 10 mi north Chihuahua, 17 Aug. 1965, H. E. Evans (MCZ); 1 9, 11

mi west Gran Morelos, 11 July 1964, J. A. Chemsak (CIS); 1 9, 8 mi northeast

Hidalgo Del Parral, 13 July 1964, J. A. Chemsak and J. Powell, black and white

lights (CIS). Guerrero: 2 ¢, Xalitla, 1500 ft, 3—20 March 1959, H. E. Evans and D.

M. Anderson (CU); i 9, Zumpango, 22 July 1963, F. D. Parker and L. A. Stange;

1 ©, Zihuatanejo, 12 Dec. 1966, G. E. Bohart (UCD). Jalisco: 1 ©, 15 mi northeast

164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Guadalajara, 17 Sept. 1970, R. M. Bohart (UCD); 1 &, 15 mi north Guadalajara,

22 July 1965, H. E. Evans (MCZ). Michoacan: 5 3, 11 mi east Apatzingän, 20 Aug.

1954, E. G. Linsley, J. W. MacSwain, R. F. Smith (UCD, 1 g CAS). Morelos: | &,

Las Estacas, 3000 ft, 6 April 1959, H. E. Evans and D. M. Anderson (CU). Nuevo

Léon: 1 ©, 5 mi south Monterrey, 8 May 1968, Malaise trap, M. W. McFadden

(UCD). Puebla: 1 &, 3 April 1962, 1 9 and 2 g, 21 Aug. 1963, 3 mi northwest

Petlalcingo, L. A. Stange (UCD). Sinaloa: 1 ¢, 5 mi north Mazatlan, 15 Aug. 1970,

Malaise trap IOA-2P, M. Wasbauer and J. Chemsak (CIS). Sonora: 4 9 and 14 &,

10 mi southeast Alamos, 29 June 1963, F. D. Parker and L. A. Stange; | 9,

Alamos, 5 Sept. 1970, R. M. Bohart (UCD). Baja California: 1 9, La Laguna, 14

Oct. 1941, Ross and Bohart (CAS). All paratypes. i

U.S.A.: Arizona: Graham Co., east foot Pinalejo Mts., 0.9 mi along road to

Marijilda Canyon from Highway 666, 3860 ft, 2 3, 4 Aug. 1965, Hugh B. Leech;

Santa Cruz Co., Yank’s Spring, Sycamore Canyon, Tumacacori Mts., 4000 ft, 1 9,

28 July 1965, Hugh B. Leech; Santa Cruz Co., Pena Blanca Lake, 9.5 mi

westnorthwest Nogales, 4000 ft, 1 3, 20 Aug. 1974, on Kallstroemia grandiflora, T.

J. Zavortink (CAS); Portal, 16 9 and 11 g, 10 Aug.—7 Sept. 1974, H. and M.

Townes; Parker Canyon Lk., 4 ©, 20 Aug. 1974, H. and M. Townes (HT). Texas:

New Braunfels, | 3, 26 July 1942, E. S. Ross (CAS); Cristal City, 1 9, July 1945,

Weyrauch (IML); Big Bend, 5000 ft, 1 © and 4 &, 1 Aug. 1975, S. and J. Peck

(HT); Uvalde Co., Speir Rch. 3 mi northwest Uvalde, 1 9, 4 May 1977, Malaise

trap 9A—SP, T. Eichlin and M. Wasbauer (CIS). All paratypes.

Guatemala: | 9, nr. Guatemala City, 1932, C. N. Ainslie; | 9, El Rancho, 900

ft, 18 Febr. 1932, C. N. Ainslie (USNM). Paratypes. In the female from El Rancho

the thorns on the tibiae and on the whitish tarsi are conspicuously dark brown.

The female of P. spangleri closely resembles that of P. minutus (Malloch) but its

petiole is slightly longer, the punctation of the mesopleura medially stronger and

closer, the oblique striae on the back of the propodeum are coarser. It seems,

possible that P. spangleri is a subspecies of P. minutus, but Texan males of the latter

species are needed to obtain certainty. P. spangleri probably can also be dis-

tinguished by its larger size.

The female of P. albifacies (Malloch) is rather similar but it has a longer petiole

and its mesopleura are more densely punctate.

Pluto brevipetiolatus (Rohwer)

(fig. 9)

_Rohwer, 1910: 100—101, 9 (Psenulus (Neofoxia) brevipetiolatus; California).

Malloch, 1933: 54—55 (Psenia brevipetiolata).

Krombein, 1951: 962 (Pluto brevipetiolatus, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length about 8 mm. Head and thorax black; anterior margin of

clypeus and mandibles reddish, underside of flagellum yellowish-red. Pronotal

tubercles and tegulae yellowish-white. Fore and mid tibiae and tarsi yellowish-red,

VAN LITH: New World Pluto 165

outer side of tibiae and first tarsal segments yellowish-white. Basal third of hind

tibiae yellowish-white, tarsal segments yellowish, basal 2/3 of segments pale

brown. Gaster black, hind margin of first tergite, hind margin and sides of second

tergite and greater part of third tergite reddish, also broad margin of tergites 4—S,

last segment dark reddish. Veins of wings brown.

Median part of clypeal margin straight, lateral angles obtuse, no distinct teeth,

this part about 1/3 of total width of margin. Frons densely finely punctate, lateral

tubercles shining. Vertex and interocellar area finely reticulate alutaceous,

sparsely punctate. POD about 1.5 times OOD. Occipital carina complete, ventro-

laterally with angle or thickening. Intercarinal space nearly as broad as first

basitarsus. Antennae somewhat clavate, segments 10—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum and scutellum shining,

indistinctly alutaceous, finely punctate, interstices on scutum mostly a few times

size of punctures, also some micro-punctation; scutellum sparsely punctate.

Propodeal enclosure medially irregularly carinate, laterally with some oblique

carinae, enclosure not deep, posteriorly not sharply defined. Back of propodeum

dull, finely reticulato-carinate, dorso-laterally finely and closely obliquely striate.

Hypo-epimeral area dull, very finely and closely but indistinctly punctate,

appearing granulose. Mesopleura dull, reticulate alutaceous, irregularly finely

punctate, interstices on anterior part a few times size of punctures. Anterior plate

of mesepisternum finely obliquely rugose, rugae continuing into anterior oblique

suture. Petiole about half length of first tergite, this tergite somewhat longer than

broad. Pygidial area narrow, about three times as long as broad (fig. 9). Second

recurrent vein of fore wings ending in third submarginal cell.

Pubescence of face silvery, mostly appressed, of pygidial area and sides of last

segment dark golden-brown, of rest of body whitish to greyish.

Male unknown.

I have not seen the type, but I could examine the female, which Malloch (1933)

studied and compared with the type and which bears the following data:

California, Lindsay, 4 June 1923, on Asclepias, W. A. Davidson (USNM).

New record from California: | 9, Gavilan Hills, 1 Oct. 1952, A. L. Melander

(USNM).

P. brevipetiolatus probably belongs to the group of angulicornis, which is

characterized by the finely sculptured back of the propodeum and the short

petiole. P. brevipetiolatus differs from the other species of this group in having an

indistinctly punctate hypo-epimeral area and a longer and narrower pygidial area.

Unfortunately the male is unknown, so that the provisional placing into this group

cannot be confirmed by the shape of its tyloidea.

Pluto rotundus sp. nov.

(fig. 6)

Female. — Length about 7 mm. Head and thorax black. Mandibles yellowish-

red, labrum reddish, clypeal margin dark red. Apices of scape of antennae reddish,

underside of flagellum orange-brown. Pronotal tubercles yellowish-white. Tegulae

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

whitish to reddish. Veins of wings dark brown. Fore trochanters and foreside of

fore femora reddish-brown, fore tibiae yellowish-red with narrow yellow outer

streak, tarsi yellowish-red, basitarsi whitish. Mid femora brown, foreside reddish-

brown, tibiae reddish, tarsi whitish, last segment reddish-yellow. Hind femora

reddish-brown, tibiae reddish, basal 1/4 yellowish, tarsi whitish, last segment

darker. Tergites I—3, ventral plate of first sternite, sternites 2—4, hind margins of

tergites 4—5 and apical part of pygidial area reddish.

Anterior margin of clypeus (fig. 6) rounded, no distinct median part. Disk of

clypeus densely punctate, with narrow, shining, depressed margin. Frons medially

densely punctate, raised round anterior ocellus which is lying in a relatively deep

pit, frons laterally with large oblong, somewhat raised shining area. Vertex almost

shining, sparsely punctate, back of head finely striate, POD almost twice OOD.

Postocellar and interocellar area raised. Intercarinai space about half as wide as

fore basitarsus. Antennae regularly thickening towards apex, segments 10—11

about as long as broad.

Lateral pronotal angles rectangular. Scutum and scutellum finely reticulate

alutaceous, punctate, interstices mostly a few times size of punctures, scutum

posteriorly with fine short rugae. Propodeal enclosure shining, large median area.

Back of propodeum dull, reticulato-carinate. Mesopleura reticulate alutaceous,

finely punctate, interstices mostly a few times size of punctures. Hypo-epimeral

area dull, densely superficially punctate. Anterior plate of mesepisternum shining,

no distinct sculpture, anterior suture narrow, crenulate. Petiole about 2/3 length of

first tergite, this tergite about 1.5 times as long as broad. Pygidial area over twice

as long as broad. Second recurrent vein of fore wings ending in third submarginal

cell.

Head and thorax whitish pubescent, gaster with yellowish-grey hairs, face

silvery, mostly appressed pubescent, pygidial area dark brown pubescent.

Male unknown.

Florida: 1 9, holotype, Lake Placid, 20 June 1962, K. V. Krombein (USNM).

P. rotundus has a slender gaster and whitish hind tarsi, like P. suffusus. It differs

from the latter species in having a narrower intercarinal space, finer punctation of

the mesopleura, brighter colour of the gaster, and a rounded clypeal margin.

The specific name refers to the rounded anterior margin of the clypeus.

Pluto suffusus (Fox)

(fig. 7)

Fox, 1898b: 18, © (Psen suffusus; New Mexico).

Ashmead, 1899: 225.

Viereck, 1901: 342, 9 (Neofoxia suffusus Fox).

Smith, H. S., 1908: 66, 9 (Neofoxia suffusa (Fox); Nebraska).

Mickel, 1918: 41, © (Diodontus suffusa (Fox)).

Malloch, 1933: 51—52, © and ¢ (Psenia suffusa; New Mexico, Arizona, California).

Krombein, 1951: 962 (Pluto suffusus, comb. nov.; New Mexico).

Female. — Length about 7 mm. Black. Labrum reddish, mandibles orange-red

VAN LITH: New World Pluto 167

or yellowish-red, Clypeal margin more or less reddish. Underside of antennae

reddish-yellow. Pronotal tubercles and tegulae yellowish-white. Fore and mid

tibiae and tarsi yellowish-brown, outer side of tibiae, basitarsi and one or more of

following segments whitish. Basal 1/3 of hind tibiae and hind tarsi except apical

half of last segment yellowish-white. Apical half of second tergite, all of third

tergite, apical margin of second sternite and all of third sternite red. Veins of wings

dark brown.

Median part of clypeal margin almost straight, nearly half total width of margin,

laterally indistinctly defined (fig. 7). Lateral parts of frons somewhat raised,

densely distinctly punctate. Vertex with reticulate alutaceous microsculpture and

sparse punctation. POD almost twice OOD. Intercarinal space as broad as fore

basitarsus. Antennae clavate, segments 10—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum and scutellum finely

reticulate alutaceous, sparsely punctate. Back of propodeum reticulato-carinate,

not coarsely so, dorsal carinae more parallel. Propodeal enclosure triangular,

shining, irregularly carinate. Hypo-epimeral area dull, densely finely punctate.

Mesopleura dull, moderately coarsely punctate, on dorsal half interstices about

size of punctures, punctures sometimes in rows; on lower part interstices wider

than punctures, posterior margin finely coriaceous and rugulose. Mesosternum

densely very finely punctate. Petiole nearly as long as first tergite, this tergite

about 1.5 times as long as broad. Pygidial area about twice as long as broad.

Second recurrent vein of fore wings ending in third submarginal cell.

Pubescence of face, thorax and gaster silvery, on face mostly appressed, dense

on metanotum and propodeum. Pubescence of pygidial area brown.

Male. — Length about 7 mm. Colour similar to that of female; only basal 1/4 of

hind tibiae yellowish, tarsi more whitish, gaster black, hind margins of tergites

reddish transparent.

Median part of clypeal margin somewhat protruding, about 1/5 of total length of

margin. Intercarinal space about as wide as fore basitarsus. Frons more coarsely

punctate. Antennae slender, segments nearly twice as long as broad, last segment

longer, segments 5—13 with small and oval tyloides, smallest on segments 5 and

Lateral pronotal angles obtuse. Scutum except for fore margin shining, coarsely

punctate with tendency to transverse rugae, interstices mostly smaller than

punctures. Scutellum shining, sparsely punctate. Mesopleura with reticulate

alutaceous microsculpture, coarsely punctate, partly coarsely rugoso-punctate.

Hypo-epimeral area and anterior plate of mesepisternum less coarsely punctate,

mesosternum much finer punctate. Back of propodeum coarsely reticulato-

carinate. Gaster slender, petiole about as long as first tergite, this tergite about 2.5

times as long as broad.

The following specimens from the collections of the National Museum of

Natural History, Washington, which represent most of the material on which

Malloch (1933) based his descriptions, have been examined:

New Mexico: | 9, Las Cruces, Cockerell 5016, on Bigeloria wrightii, det. Psenia

suffusa (Fox) by J. R. Malloch; 1 9, Las Cruces, Cockerell 5017, USNM acc. no.

168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

30623, det. Psenia suffusa by Malloch; 1 g, Las Cruces, Cockerell 5105; 2 3,

Mesilla, 28 July, Cockerell (one ¢ labelled “Mimesa suffusa 3 Fox det. Roh.”); 1

©, La Luz Sta., 19 Aug. 1930, on L. alyssoides; 1 3,2 mi north Vado, 18 Aug. 1929,

on Salsola pestifer, lot 5823. Arizona: 22 3, Higley, 15 July 1917, E. G. Holt; 1 3,

Mt. Superstition nr. Higley, 24 July 1917, E. G. Holt; 1 9, Arizona, 2546, C. F.

Baker; 2 g, Arizona, 2572, C. F. Baker; 1 9, Tucson, collection Ashmead; I 9,

Phoenix, 16 Oct., on Baccharis, Cockerell. California; 3 &, Imperial Co., May

1911, J. C. Bridwell; 1 &, Redlands, F. R. Cole; 1 9, Antioch, 10 May 1932, E. O.

Essig; 1 9, Lindsay, 9 Sept., on Asclepias, W. A. Davidson; 1 9, June 1932, Fowler

(USNM).

New records from these states: Arizona: 1 ©, Phoenix, 8 Aug. 1950, R. S. Beal

(USNM); 1 9, Globe, 6 Aug., D. K. Duncan, K. V. Krombein det. (KVK); 1 9,

Tucson, May 1936, Bryant, 139; 1 ©, Box Canyon, Santa Rita Mts., 29 Aug. 1952,

B. Malkin and V. E. Thatcher (CAS); 2 9 and 4 g, Portal, 11—18 Aug. 1974, H.

and M. Townes (HT). California: Fresno Co.: 1 9, Fresno, 27 July 1968, T. R.

Haig; Imperial Co: 4 & and 1 9, Bard, 14—29 Aug. 1965, ex Gossypium hirsutum,

R. A. Flock, R. Haygood and H. Ray; 9 g, Chocolate Mts., Ogilby Rd. 3 mi south

Jct. Hwy. 78, 16—22 Oct. 1977, Malaise trap 8A—5P, M. Wasbauer (CSC); | 9,

Seeley, 25 June 1965, ex cotton, R. A. Flock (CSC); Riverside Co.: 2 9, Hemet, 24

July 1946, Helianthus, J. W. McSwain; 1 g, Hopkins Well, 29 April 1952, P. D.

Hurd; | 9, Palm Springs, 21 May 1953, A. L. Melander (USNM); San Diego Co.: 1

Q, Chula Vista, 2 Aug. 1961, on Atriplex semibacchata, F. X. Williams (CAS);

Sutter Co.: 1 g, 2 mi southeast Marysville, 28 June 1962, T. R. Haig (CSC). New

Mexico: Don Ana Co. 2 g, 13 July 1954, swept from cotton, R. E. Fye (USNM); 1

©, Las Cruces, 11 July 1956, H. and A. Howden, at light (CNC); 6 © and 6 g,

Hatch, 27—30 Aug. 1974, H. and M. Townes (HT); Hidalgo Co.: 1 9, Granite

Cap, 17 mi north Rodeo, 4 Sept. 1976, F. G. Andrews, collected on Flourensia

cernua, association Tachardiella cornuta, Lacciferiidae (CIS); Otero Co.: 1 9,

White Sands National Monument, 21 Aug. 1962, H. V. Weems Jr. (FSC).

First record from Florida: | 9, Highlands Co., Archbold Biol. Station, Lake

Placid, 21—28 April 1975, K. V. Krombein (USNM).

First records from South Carolina: 1 9, Aiken, 12 June 1957, W. R. M. Mason;

1 9, Hilton Head Is., 11—23 July 1965, H. F. Howden (CNC). The intercarinal

space of these females is somewhat narrower than usual.

First records from Texas: 1 9, Knippa, 24 July 1910, F.C. Pratt; 1 g, Fabens, 31

Aug. 1945, on cotton (USNM); 1 &, 10 mi west Ft. Davis, Pt. Rocks, 5000 ft, 29

May 1959, W. R. M. Mason (CNC).

First records from Mexico: Baja California: La Paz, 1 9 and I &, 29 June 1919,

G. F. Ferris, Stanford University (UCD), 3 9, 3—5 June 1921, E. P. Van Duzee; |

©, Mulegé, 14 May 1921, E. P. Van Duzee; Escondido Bay, | g, 14 June 1921, E.

P. Van Duzee; San Domingo, 6 g, 19 July 1938, Michelbacher and Ross; Sur San

José del Cabo, 2 &, 17 July 1971, H. G. Real and R. E. Main (CAS). Coahuila: 2 g,

Boquillas del Carmen, 1850 ft, 23 May 1959, Howden and Becker (CNC). Nayarit:

5 g, 2 mi east San Blas, 8 April 1963, G. W. Frankie (USNM). Sinaloa: | 9 and |

3, Guamúchil, 6 May 1953, R. C. Bechtel and E. I. Schlinger (UCD); I g, Villa

VAN LITH: New World Pluto 169

Union, 17 Aug. 1962, H. E. Evans (MCZ). Sonora: 1 © and | g, 10 mi southeast

Alamos, 29 June 1963, F. D. Parker and L. A. Stange (UCD). The tyloidea of the

males from Nayarit are dark.

Like Malloch I have not seen the types and I have followed him in his

conception of the species. The description of the female by Fox, who had five

specimens from Las Cruces and Rincon, New Mexico, before him, is not very

clear. He states that the hind tarsi are entirely whitish or ringed with whitish, that

either the greater portion of the gaster is black or that reddish prevails and that the

punctures of the mesopleura are large and sparse(?), striated posteriorly. A study

of his type material and designation of a lectotype is therefore wanted.

The sexual dimorphism is considerable. Although in the collections I have

studied the two sexes rarely originate from the same locality, I believe that they

have been correctly associated by Malloch.

The female of P. suffusus is easily distinquished by the broad intercarinal space,

the long petiole, the red tergites and the whitish tarsi. The male has coarsely

punctate mesopleura, small tyloidea on antennal segments 5—13 and whitish tarsi.

Confusion of the female with P. sayi by Fox, as suggested by Malloch, does not

seem to be very likely.

Pluto biformis sp. nov.

Male. — Length about 6 mm. Black; mandibles reddish-brown, apex of

flagellum orange-brown below. Pronotal tubercles whitish. Tegulae yellowish.

Fore femora largely orange-brown, fore and mid tibiae entirely so; basal 1/3 of

hind tibiae, tarsi except for last segment of hind tarsi yellowish-white. Veins of

wings dark brown. Hind margins of tergites reddish transparent.

Median 1/3 of clypeal margin somewhat depressed and protruding, almost

straight. Frons densely punctate. Vertex shining, behind ocelli densely punctate,

laterally with larger interstices. Back of head and tempora finely striate.

Intercarinal space narrow, occipital carina distinct. Antennae slender, segments

quadrate or slightly longer than broad, last segment about twice as long as broad,

segment 6 with short narrow tyloides near apex, segments 7—8 with long narrow

tyloides, segment 9 with long tyloides widening towards apex, segments 10—12

with broad oval tyloides, shorter than segments; tyloidea on segments 8—12

angularly raised on apical 1/3 when seen in lateral aspect.

Lateral pronotal angles sharp. Scutum shining, interstices often a few times size

of punctures. Scutellum shining, sparsely punctate. Median part of enclosed area

of propodeum irregularly carinate, back of propodeum irregularly, moderately

coarsely reticulato-carinate. Mesopleura alutaceous, coarsely punctate, on upper

part interstices larger than punctures, on lower part more densely punctate, on

median part punctures partly in rows. Hypo-epimeral area dull, densely punctate,

interstices distinct. Mesosternum densely finely punctate. Second recurrent vein

of fore wings ending well in third submarginal cell. Petiole about 2/3 length of first

tergite, this tergite about 1.5 times as long as broad.

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pubescence of face silvery, mostly appressed, of rest of body whitish. Wings

whitish pubescent (freshly emerged?)

Female unknown.

Mexico: 1 g, holotype, Durango, 6 mi east Durango, 6500 ft, 24 July 1964, L. A.

Kelton (CNC).

This male seems to be closely related to P. angulicornis (Malloch), but the

tyloidea are distinctly different.

The specific name refers to the tyloidea on the antennae of the male, which are

partly narrow, partly broad oval.

Group of texanus

Pluto texanus (Malloch)

(fig. 8)

Malloch, 1933: 56, 9 and g (Psenia texanus; Texas).

Krombein, 1951: 962 (Pluto texanus, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length about 6 mm. Head and thorax black, mandibles yellowish-

red with darker tips, labrum reddish. Antennal segments 4—12 yellowish-brown

below. Pronotal tubercles yellowish-white. Tegulae reddish transparent. Fore and

mid tibiae reddish-yellow with ivory-white outer streak. Basal third of hind tibiae

yellowish-white. Fore and mid tarsi whitish to yellowish-brown, hind tarsi dark

brown, apices of segments 1—4 yellowish-brown. Hind margin of first gastral

tergite, all of second tergite and sides of third tergite red. Hind margins of tergites

3—5 and greater part of sternites red. Veins of wings brown.

Apex of clypeus very slightly protruding, almost straight, median part slightly

protruding, laterally with distinct small tooth (fig. 8). Disk of clypeus closely

punctate. Frons densely finely punctate. Vertex almost shining, sparsely punctate.

Intercarinal space less wide than fore basitarsus. Antennae clavate, segments

8— 11 shorter than broad, last segment about 1.5 times as long as broad.

Lateral pronotal angles rectangular. Scutum shining, finely punctate, interstices

a few times size of punctures; narrow hind margin finely striato-punctate.

Scutellum shining, sparsely punctate, lateral and posterior margins finely rugoso-

punctate, no distinct median line. Propodeal enclosure somewhat shining, laterally

with oblique carinae, medially irregularly carinate. Back of propodeum

coriaceous and moderately coarsely reticulato-carinate, dorso-lateral carinae

somewhat parallel. Mesopleura weakly shining, regularly finely punctate,

interstices a few times size of punctures, posterior margin coriaceous with very

fine striae. Hypo-epimeral area with fine punctures and striae, posterior margin

with coarser rugae. Anterior plate of mesepisternum finely striate, oblique suture

crenulate. Petiole about 3/4 length of first tergite, this tergite somewhat longer

than broad. Pygidial area about twice as long as broad. Second recurrent vein of

fore wings ending in third submarginal cell.

VAN LITH: New World Pluto 171

Pubescence of face silvery, appressed and also with some long erect hairs,

tempora with short silvery pubescence. Pygidial area with reddish-brown

backwards directed hairs, vertex and rest of body greyish-white pubescent.

Male. — Slender. Length about 5 mm. Fore and mid tibae yellowish-red with

narrow yellow streak on outer side, tarsi whitish to yellowish-brown. Apical

margin of first tergite, second tergite, except for a black band in front of hind

margin and base of third tergite, red.

Apex of clypeus slightly protruding, almost straight. Third antennal segment

about 1.75 times as long as broad, following segments longer than broad, last

segment nearly twice as long as broad. Segments 4—7 with shining, black narrow

tyloidea, on segment 4 about 2/3 length of segment, on segments 5—7 gradually

decreasing in length, on segment 8 about 1/4 length of segment, on segments 9— 10

a very small but distinct shining point which is also visible in lateral view, tyloidea

on segments 11—12 linear, about 1/3 of length of segment on segment 11, a little

longer on segment 12.

Mesopleura strongly punctate, interstices on upper part about size of punctures,

on median part punctures more in rows. Hind margin of mesopleura and hypo-

epimeral area rugose. Scutum shining, punctate, interstices mostly about twice

size of punctures. Scutellum somewhat shining, irregularly punctate. Back of

propodeum coarsely reticulato-carinate. Gaster slender, petiole about as long as

first tergite, this tergite about 1.5 times as long as broad.

Material examined: 1 ©, paratype, and 1 J, allotype, Brownsville, Texas, 1921,

J.C. Bridwell (USNM, type No. 44226).

New records from Texas: 1 Z, Dallas, 421 (NMW); 1 ©, Olivia, 9 July 1956, D.

H. Habeck (NCSU).

The male of P. texanus was not included in Malloch’s key to the species. In his

brief description he did not pay attention to the irregular tyloidea, but he only

mentioned that there is “no evident sensory elevation on the second flagellar

segment”.

Pluto littoralis (Malloch)

(fig. 10)

Malloch, 1933: 56—57, 9 and ¢ (Psenia littoralis; Maryland).

Krombein, 1951: 962 (Pluto littoralis, comb. nov.); 1954: 233 (Florida); 1958: 189.

Bohart & Menke, 1976: 171.

Female. — Length about 6—7 mm. Head and thorax black. Mandibles

yellowish-red with darker tips, labrum dark reddish. Antennal segments 4—12

orange-brown below. Pronotal tubercles yellowish-white. Tegulae reddish trans-

parent. Fore and mid tibiae reddish-yellow with ivory-white outer streak, mid

tibiae below sometimes brownish. Basal third of hind tibiae yellowish-white. Fore

and mid tarsi whitish to yellowish-brown, hind tarsi dark brown, segments 1—4

with paler apex. Apical margin of tergite 1, all of second tergite, base or sides of

tergite 3 and sternites 2—3 red. Hind margins of segments 3—5 somewhat reddish

transparent, apex of last segment dark reddish. Veins of wings brown.

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Clypeal margin like that of P. texanus but median part almost straight, laterally

with distinct short tooth. Disk of clypeus closely punctate. Frons densely finely

punctate. Vertex sparsely punctate, posteriorly indistinctly transversely striate.

Intercarinal space nearly as wide as fore basitarsus. Antennae clavate, segments

7—11 shorter than broad, last segment about 1.5 times as long as broad.

Lateral pronotal angles almost rectangular. Scutum shining, distinctly punctate,

interstices a few times size of punctures; on central part punctures closer, narrow

hind margin striato-punctate. Anterior half of scutellum with two large impunctate

or nearly impunctate shining areas (in allotype scutellum slightly deformed),

lateral margins punctate, hind margin narrowly striato-punctate. Propodeal

enclosure somewhat shining, lateral parts obliquely striate, median part irregularly

carinate. Propodeum with reticulate alutaceous microsculpture and behind

enclosed area rather finely reticulate, posteriorly and laterally coarsely reticulate,

dorso-lateral carinae somewhat parallel. Greater part of metapleura shining,

upper third dull. Mesopleura with reticulate alutaceous microsculpture, distinctly

punctate, sometimes somewhat striato-punctate, interstices mostly as large as

punctures, partly larger, posterior margin granulose. Hypo-epimeral area with fine

punctures and more or less distinct longitudinal rugae, at least on posterior part.

Anterior oblique suture coarsely foveolate, anterior plate of mesepisternum very

finely striate. Petiole about 2/3 or 3/4 length of first tergite, about half as long as

hind trochanter and femora together. First tergite slightly longer than wide.

Pygidial area less than twice as long as broad. Second recurrent vein of fore wings

ending in third submarginal cell.

Pubescence of face silvery, appressed and also with some long erect hairs,

tempora with short whitish pubescence. Pygidial area with backward directed

reddish-brown hairs. Vertex and remainder of body greyish-brown pubescent.

Male. — More slender than female. Length about 6 mm. Mid tibiae not

distinctly whitish. Apical margin of first tergite, second tergite except for a broad

black band in front of hind margin, and base of third tergite red.

Median part of clypeal margin straight, slightly protruding. Intercarinal space

narrower. Third antennal segment about twice as long as broad, following

segments longer than broad, last segment over twice as long as broad. Segments

4—7 with narrow, shining black tyloides, on segment 4 about 2/3 length of

segment, on segments 5—7 gradually decreasing in length, small or indistinct on

segment 8, absent or very indistinct on segments 9—10, narrow and usually paler

on segments 11—12, on segment 11 about 1/3 length of segment, on segment 12

somewhat shorter than segment, indistinct on segment 13. Malloch’s description

of the antennae of the male does not agree with the antennae of the type which Dr.

A. S. Menke kindly sent me for study. Malloch states that the “sensory areas are

almost linear, black, entire, present on all but the basal and apical segments, and

sometimes rudimentarily so on apical one”, hereby overlooking that the tyloidea

on segments 8—10 are rudimentary or absent.

Scutum shining, strongly punctate, interstices on median part larger than

punctures. Punctation of mesopleura somewhat coarser than in female; hypo-

epimeral area and posterior margin of mesopleura coarsely rugose. Reticulation

VAN LITH: New World Pluto 173

on back of propodeum coarser than in female, carinae not parallel. Petiole about

as long as hind femur or as long as first tergite, this tergite about 1.5 times as long

as wide.

Base of genital apparatus (fig. 10) brown, middle part white and narrowed apex

dark brown. Inner margin of stipes triangularly emarginate.

Material examined: Maryland, Chesapeake Beach, 1 &, holotype, 3 July 1924, J.

R. Malloch, “Psenia littoralis type det. J. R. Malloch”; 1 ©, allotype, 2 July 1916,

W.L. McAtee (USNM, type No. 44227).

New records from U.S.A.: Florida: Placida, 1 9, 11 April 1952, G. S. Walley;

Punta Gorda, 1 &, 2 April 1953, K. V. Krombein, K. V. Krombein det. (KVK);

Cedar Key Levy Co., 8 9 and 12 &, 19 Aug., 16—28 Sept. and 12 Oct. 1975, two 9

with prey (small leafhoppers), E. E. Grissell (FSC); 1 © and I g, 19 Aug. 1975, E.

E. Grissell (USNM); Gulf Co., St. Joseph St. Park, 1 © and 3 Z, 1—3 May (1970),

W. W. Wirth (USNM); Cudjoe Key, Monroe Co., 3 9, 12 July 1971, W. H. Pierce

(FSC). North Carolina: Bogue Banks, 2 3, 14 Sept. 1959, Salicornia, L. Davis;

North R., 1 9, 5 Sept. 1959 and 1 g, 6 June 1960, L. Davis (USNM). South

Carolina: Fripp Island Beaufort Co., 1 9 and 2 g, 26 Sept. 1973, G. C. Steyskal

(USNM).

The three females from Cudjoe Island have somewhat weaker and more widely

placed punctures on the mesopleura. One of the males from Cedar Key has also on

the antennal segment 10 a distinct tyloides or at least a distinct tubercle.

P. littoralis and P. texanus are very similar. In the female of P. littoralis the

mesopleura are somewhat coarser and more densely punctate, the hind margin is

more coarsely rugose. The scutum is more densely punctate. The male of

P. littoralis has no distinct tyloidea on antennal segments 9—10, the allotype of

P. texanus has very small but distinct tyloidea on these segments.

Further studies may prove that P.littoralis is a geographic subspecies of

P. texanus.

Group of sayi

Pluto sayi (Rohwer)

(fig. 11)

Cresson, 1872: 227, 9 (misidentified as Mimesa pauper Packard; Texas).

Rohwer, 1910: 100, 9 (Psenulus (Neofoxia) sayi; Kansas, Texas).

Malloch, 1933: 55—56, 9 and g (Psenia sayi; Alabama, Arizona, California, District of Columbia,

Louisiana, New Mexico, Texas).

Krombein, 1951: 962 (Pluto sayi, comb. nov.; Austral zone east of California); 1967: 396 (California).

Bohart & Menke, 1976: 171.

Female. — Length about 5.5 mm. Head and thorax black; labrum reddish,

mandibles reddish with darker tips, underside of flagellum yellowish-red. Pronotal

tubercles and part of tegulae yellowish-white. Outer side of fore and mid tibiae,

basal third of hind tibiae and fore and mid basitarsi yellowish-white. Remainder of

174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

fore and mid tarsi brownish, hind tarsi except for apices of segments I—4 dark

brown. Gaster largely black; hind margin of first tergite, all of second tergite, great

part of third tergite and all of sternites 2—3 usually red (sometimes more tergites

and sternites), apex of last gastral segment dark reddish. Veins of wings dark

brown.

Median part of clypeal margin very weakly emarginate with short lateral teeth

(fig. 11). Frons raised, densely punctate. Vertex shining, indistinctly alutaceous,

sparsely punctate. POD larger than OOD. Occipital carina complete, intercarinal

space narrow. Antennae clavate, segments 10—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum shining, rather strongly

punctate, interstices mostly a few times size of punctures, medially smaller.

Scutellum sparsely finely punctate, hind margin dull. Propodeum dull, enclosure

without large median area, back irregularly reticulato-carinate, dorso-lateral

carinae mostly parallel. Mesopleura somewhat shining but distinctly reticulate

alutaceous, finely punctate, interstices larger than punctures or a few times larger

than punctures, interstices largest on upper part; hind margin finely coriaceous.

Punctation of mesosternum finer, interstices about size of punctures. Hypo-

epimeral area somewhat shining, finely alutaceous and punctate, punctures

smaller than on mesopleura, interstices about size of punctures. Petiole about 2/3

length of first tergite, this tergite about as long as broad. Pygidial area about twice

as long as broad. Second recurrent vein of fore wings ending in third submarginal

cell.

Pubescence of face silvery, mostly appressed. Hairs on pygidial area golden

brown, remainder of body whitish pubescent.

Male. — Length about 4.5—5 mm. Gaster black, hind margins of tergites

reddish, sometimes second tergite and base of third tergite red, especially in males

from Arizona and California. Antennal segments nearly 1.5 times as long as broad,

segments 5—12 with linear tyloides, shorter on segment 12, sometimes also an

indistinct tyloides on segment 4. Punctation of scutum and mesopleura coarser

than in female. Interstices on upper part mesopleura larger than punctures, on

median part smaller. Mesopleura sometimes shining, especially in males from

Arizona and California, which are often also smaller (3.5—4 mm). Hypo-epimeral

area dull, finely punctate, interstices larger than punctures, at least on lower part.

Propodeum somewhat coarser reticulate than in female. Petiole nearly as long as

first tergite, this tergite about 1.5 times as long as broad.

Material examined: | 9, paratype, Texas [Bosque Co., coll. G. W. Belfragel,

with labels ““Mimesa pauper Pack.” and ‘‘Neofoxia sayi Roh. paratype” (USNM,

type No. 12856).

Alabama: 1 © and 17 3, 2280, C. F. Baker. Arizona: | &, 2546, C. F. Baker, 43

4, Higley, 15 July 1917, E. G. Holt, 1 g, Higley, 25 June 1917, E. H. Holt, 1g

Sacaton, sweeping pomegranate. California: | g, Lindsay, 6 Sept. 1925, on

Helianthus, W. A. Davidson. Louisiana: 2 © and 6 g, 2336, 1 9 and 2 &, 2392, C.

F. Baker, 2 © and 1 Z, Opelousas, G. R. Pilate. New Mexico: | &, Albuquerque,

3240, Cockerell (USNM). This is probably all material studied and recorded by

Malloch.

VAN LITH: New World Pluto 175

New records from U.S.A.: Alabama: 9 © and 16 &, Montgomery, 23—27 June,

K. V. Krombein (KVK and ML). Arizona: Tucson, | &, 30 May 1920, F. X.

Williams (CAS), 3 © and 2 g, 9—18 June 1938, R. H. Crandall, det. K. V.

Krombein (KVK); 1 4, Sta. Rita Mts., 20 Sept. 1936, Bryant, Lot 41; 1 3, Oak

Creek Canyon, 6 June 1940, G. E. Bohart (CAS); 7 3, Benson, San Pedro River, 7

June 1968, Menke and Flint (USNM). California: Imperial Co.: 2 9, “Imperial

Co.”, May 1911, J. C. Bridwell (USNM); Imperial, 3 3°, 28 June—14 July 1965, ex

cotton and ex Gossypium hirsutum, Collins and R. Pineda; Seeley, 2 9 and 54 &, 2

July 1965, ex Gossypium hirsutum, L. Pineda and H. Ray, 1 9, 28 June 1965, ex

cotton, Collins, 1 g, 9 Aug. 1971, ex cotton, R. A. Flock and Pineda; Bard, 1 9

and 19 g, 22 June—17 July 1965, 2 © and 2 Z, 18 Aug.—8 Sept. 1965, ex

Gossypium hirsutum, H. Ray, R. Haygood and R. A. Flock; El Centro, | 9 and 6

gd, 12—13 July 1965, ex Gossypium hirsutum, R. A. Flock, 1 &, 13 July 1965, ex

cotton, R. A. Flock; Calexico, 3 Z, 8 June 1965, ex cotton, R. A. Flock; Brawley,

1 4, 14 June 1965, ex cotton, Pineda; Holtville, 3 &, 8 July 1965, ex Gossypium

hirsutum, R. A. Flock; Westmorland, 1 Z, 15 June 1965, ex Gossypium hirsutum, L.

Pineda (CSC); Westmorland, 3 &, 15 May 1974, H. and M. Townes, G. and C.

Townes (HT); Ventura Co.: Holtville, 3 ¢, 26 Sept. 1968, ex Sorghum, Cal. Dept.

Agric., No. 68J 9—17, R. A. Flock, Pluto sayi det. M. Wasbauer 1968 (CSC); San

Bernardino Co.: 4 ¢, Chino, 13 July 1965, ex Zea mays, Birdsall coll. (CSC);

Fresno Co.: Clovis, 2 ¢, 31 May 1974, ex almond, Peregrin coll.; Kerman, | &, 26

June 1972, ex alfalfa, Moré coll. (CSC); Sacramento Co.: American River, 1 9, 15

June 1966, M.S. Wasbauer (CSC); Stanislaus Co: 1 ©, Del Puerto Canyon, Frank

Raines Park, ca. 1100 ft, 4 July 1971, P. H. and M. Arnaud (CAS); Yolo Co.: Davis,

1 g, 5 July 1975, W. J. Pulawski (BM); Riverside Co: 1 © and 5 g, Blythe, E. P.

Van Duzee (CAS). District of Columbia: 3 9, Washington, 16—25 Aug. 1949, R.

Boettcher, D. G. Shappirio coll. (USNM). Florida: 1 &, Winter Park, 8 Aug. 1940,

H. T. Fernald (hind tarsi pale; with label “Pluto nr. tibialis’), 1 Q and 2 &,

Arcadia, 30 June—2 July 1962, K. V. Krombein, 1 g, Hialeah, 21 July 1965, C.

Stegmaier (USNM); 1 3, Fort Myers, Lee Co., 26 April 1967, P. P. Babiy (BSM); I

3, St. Joseph, Gulf Co., (T.H. Stone Memorial) State Park, 4 May 1973, C. KR.

Artaud (FSC). Iowa: 10 © and 2 g, Sioux City, 13 July 1927, 23 July 1931, 6 July

1933, 19— 30 July 1934, 1 © and | g, without date, C. N. Ainslie; 1 g, County, No.

47, 4 July 1932, Russell (USNM). Kansas: 1 9, Clay Co., Aug., Bridwell, labelled

“Neofoxia suffusus Fox 9 Bridwell det. 1905”; 1 &, Dickinson Co., Aug. 1901,

Bridwell, labelled ”Neofoxia suffusus Fox ¢ Bridwell det. 1905”; 1 © and 1 g,

Baldwin June, Bridwell; 1 &, Clay Co., Aug. 1901, Bridwell; 2 3, Manhattan, Ac.

4777 and 4782 Sp., Wm. P. Hayes (USNM); | 9, 7 mi east Washington, 8 Aug.

1973, E. S. Ross (CAS). Louisiana: 1 ¢, Tallulah, 8 July 1948, Bug Catcher Exp.

R. C. Gaines (USNM); 1 9, Rapides Parish, 1 June 1973, Peter Rush (HT).

Missouri: Kansas City, 1 g, 9 Aug. 1934, R. H. Crandall (KVK); 1 ©, Roaring

River State Park, 15 June 1954, J. Green (CAS); Columbia, Boone Co., 1 9 and 1

3, 17 Aug. 1966, 9 © and 12 3, 26 June—30 Aug. 1967, 2 9, 31 Aug. 1968, Malaise

trap, F. D. Parker (USNM). New Mexico: 1 &, Dona Ana Co., 17 Aug. 1954,

swept from cotton, R. E. Fye, 2 9, 3 mi southwest Las Cruces, 2 Aug. 1929, on

176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Salsola pestifer, V. E. Romney, one © labelled “Psen tibialis Cr. det. Malloch 1930”

(USNM). North Carolina: Raleigh, 2 9 and 1 g, 10 Sept. 1940, 2 3, 24 July 1941; 1

3, Faison, 9 June 1953, on celery, T. B. Mitchell; 1 ©, Hyde Co., 5 June 1959,

David A. Young; 1 ©, Robbinsville, 28 July 1959, David A. Young (NCSU). South

Carolina: 2 9 and 3 Z, Columbia, 8—19 Aug. 1951, L. and G. Townes (HT).

Tennessee: 2 9 and 17 g, Madison, Davidson Co., 9 June—3 Aug. 1967, P. P.

Babiy (BSM). Texas: 2 9, Presidio, 2 April—9 May 1941, W. L. Lowry, Box 1,

traps pink bollworm larvae installed, Lot No. 41—7893, Pluto sayi det. K. V.

Krombein (USNM); 1 4g, Dallas, No. 421 (NMW). Virginia: Clifton, | &, 12 June

1933, J. C. Bridwell (USNM); 1 9, 30 June 1935, K. V. Krombein; Arlington, 1 9

and 4 34, 7—22 June 1947, K. V. Krombein (KVK). |

First records from Mexico: Chihuahua: 1 Z, 9 mi south Hidalgo de Parral, 31

July 1967, R. C. Gardner, C. R. Kovacic and K. Lorenzen (UCD). Coahuila: 2 g,

Boquillas del Carmen, 1850 ft, 23 May 1959, Howden and Becker (CNC). Nayarit:

1 &, Acaponeta, 4 May 1953, R. C. Bechtel and E. I. Schlinger (CIS). Nuevo Léon:

1 9, 16 mi south Montemorelos, 19 July 1954, Univ. Kansas Mex. Exped. (UK).

San Luis Potosi: 1 ©, El Bonito, 7 mi south Ciudad Valles, 300 ft, 19 Dec. 1970, P.

H. and M. Arnaud (CAS). Sinaloa: 2 ©, 8 mi south Elota, 18 May 1962, F. D.

Parker (UCD). Veracruz: 1 9, Boca del Rio, 23 June 1961, Univ. Kansas Mex.

Exped., on flowers of Eupatorium (UK); | 9, Cordoba (Veracruz?), Mann (MCZ).

Tergite 2 of lastmentioned female is almost entirely dark brown.

First records from El Salvador: 1 &, San Andrés, 6 June 1958, O. L. Cartwright

(USNM) (this male has no tyloides on segment 12); 1 &, Usulutan, 50 ft, 12 July

1963, D. Q. Cavagnaro and M. E. Irwin (CAS).

First record from Nicaragua: 1 9, San Juan Managua, 30 May 1960, M.

Vaughan (USNM).

First records from Cuba: 2 ©, San Vicente, Pinar del Rio, July 1940, J. C.

Bradley (CU).

The mesopleura of specimens from Arizona, California, Texas and from Mexico

are more shining and somewhat more coarsely punctate, especially in the males

from Arizona. Females from these regions often have tergites 2—3, sometimes

also tergites 1 and 4, entirely red; in the males too the red colour on the gaster is

more extended than in those from Alabama and Louisiana. The mesopleura of the

females from Nicaragua and Cuba are dull, distinctly reticulate alutaceous. One of

the males from Florida has very pale, almost yellowish hind tarsi.

Pluto stenopygidialis sp. nov.

(figs. 12—13)

Female. — Length about 6 mm. Black: mandibles and underside of flagellum

reddish-brown; tibiae, tarsi and tegulae largely dark brown.

Median part of clypeal margin broad, almost straight, sharp lateral angles, in the

middle very weakly protruding, margin depressed (fig. 12). Clypeal disk convex,

closely finely punctate. Frons densely punctate, vertex alutaceous, finely punctate

and indistinctly transversely striate. Tempora finely striate. POD nearly twice

VAN LITH: New World Pluto 177

Fig. 11. Pluto sayi (Rohwer), 9, clypeus. Figs. 12—13. P. stenopygidialis sp. n., 9, holotype; 12, clypeus;

13, pygidial area. Fig. 14. P. jugularis sp. n., 9, holotype, head in ventral aspect. Fig. 15. P. punctatellus

sp. n., 9, holotype, clypeus. Fig. 16. P. aerofacies (Malloch), 3, allotype, clypeus. Fig. 17. P. evansi sp.

n., 9, holotype, clypeus. Fig. 18. P. emarginatus sp. n., 9, holotype, head in frontal aspect. Figs.

19—21. P. rufibasis (Malloch); 19, clypeus of © ; 20—21, left paramere of g, ventral and lateral aspect.

178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

OOD. Intercarinal space about half width first basitarsus. Antennae short,

segments 10—11 about as long as broad.

Lateral pronotal angles obtuse. Scutum shining, finely punctate, interstices

often a few times size of punctures. Scutellum almost shining, sparsely punctate.

Propodeal enclosure almost flat, carinae fine, medially irregular. Back of

propodeum dull, mostly finely obliquely striate. Mesopleura reticulate alutaceous,

upper half rather strongly irregularly punctate, partly striato-punctate, lower part

very finely very sparsely punctate, posterior margin finely coriaceous. Meso-

sternum densely finely punctate. Hypo-epimeral area finely punctate, interstices

at least size of punctures. Anterior oblique suture narrow, foveolate, anterior plate

of mesepisternum dull, sparsely punctate. Metapleura largely reticulate

alutaceous. Petiole about half length first tergite, this tergite somewhat longer than

broad. Pygidial area narrow, over 2.5 times as long as broad (fig. 13). Second

recurrent vein of fore wings ending in third submarginal cell.

Face dark brown-golden pubescent, not appressed, also a number of long erect

hairs. Vertex and scutum brownish pubescent, pubescence of mesopleura

yellowish-grey, of back of propodeum silvery-white, of pygidial area dark brown.

Male unknown.

Arizona: 1 9, holotype, near Roosevelt Lake, 29 April 1947, H. and M. Townes

(KVK); 1 ©, paratype, Parker Canyon Lk., 22 Aug. 1974, H. and M. Townes (HT).

This female keys out near P. punctatellus, but is easily distinguished by the

brown-golden, not appressed pubescence of the face and by the dark legs.

The specific name refers to the narrow pygidial area of the female.

Pluto jugularis sp. nov.

(fig. 14)

Female. — Length about 6 mm. Head and thorax black. Mandibles yellowish-

red with darker tips, labrum reddish. Underside of flagellum yellowish-red.

Pronotal tubercles yellowish-white, tegulae yellowish. Apices of fore and mid

femora, all of fore and mid tibiae and tarsi, basal 2/5 of hind tibiae and apices of

hind tarsal segments 1—4 yellowish. Tarsal claws of mid legs brownish. Hind

margin of tergite 1, all of tergites 2—3 and sternites 2—3 red, hind margins of

following segments reddish transparent. Veins of wings brown.

Clypeal margin resembling that of P. sayi (fig. 11), protruding median part over

1/3 of total width of margin, weakly emarginate, lateral teeth small. Disk of

clypeus densely finely punctate. Frons except for sides densely finely punctate.

Vertex reticulate alutaceous, sparsely finely punctate. Tempora almost smooth.

POD somewhat larger than OOD. Occipital carina complete, intercarinal space

broader than first basitarsus (fig. 14). Antennae short, clavate, segments 9—11

shorter than broad, segment 12 about 1.5 times as long as broad.

Lateral pronotal angles about rectangular. Scutum and scutellum finely

reticulate alutaceous, sparsely finely punctate, near hind margin more densely

punctate. Scutellar suture crenulate. Propodeal enclosure shining, with large

irregular median area. Back of propodeum coriaceous, moderately finely

VAN LITH: New World Pluto 179

reticulato-carinate, dorso-lateral carinae almost parallel. Mesopleura, hypo-

epimeral area and anterior plate of mesepisternum finely reticulate alutaceous,

mesopleura minutely punctate, hypo-epimeral area almost imperceptably (x 30)

punctate, interstices on mesopleura a few times size of punctures, its hind margin

dull, coriaceous. Mesosternum densely distinctly but finely punctate. Anterior

oblique suture finely crenulate. Petiole somewhat shorter than first tergite, this

tergite about as long as broad. Pygidial area about 1.5 times as long as broad.

Second recurrent vein of fore wings ending in third submarginal cell.

Face and pronotal collar with silvery, mostly appressed pubescence. Pubes-

cence of tempora and mesosternum silvery, of vertex, scutum and scutellum

brownish, of pygidial area reddish-brown, of rest of body yellowish-grey.

Male unknown.

Brazil: 1 9, holotype, Corumbà, Mato Grosso, 14—23 Dec. 1919 (CU).

P. jugularis resembles P. sayi but is easily distinguished by the broad intercarinal

space and the other characters mentioned in the key.

The specific name refers to the posterior part of the underside of the head with

broad intercarinal space.

Pluto punctatellus sp. nov.

(fig. 15)

Female. — Length about 7 mm. Black. Mandibles, labrum and underside of

flagellum reddish. Pronotal tubercles dark brown or pale brown. Fore tibiae and

tarsi yellowish-brown, tibiae more or less darkened; base and apex of mid tibiae

and entire tarsi yellowish-brown, last segment darker brown; hind tibiae and tarsi

brown, bases of tibiae and apices of tarsal segments paler. Veins of wings dark

brown. Hind margins of gastral segments reddish transparent, last segment dark

reddish.

Slightly protruding median part of clypeus almost straight, about half total width

of clypeal margin, laterally a weak tooth (fig. 15). Frons densely finely punctate,

lateral tubercles and vertex shining, almost impunctate. POD about equal to

OOD. Intercarinal space narrow. Antennae slightly clavate, segments 9—11 about

as long as broad.

Lateral pronotal angles almost rectangular. Scutum and scutellum very finely

alutaceous, almost shining, irregularly punctate, on lateral parts interstices a few

times size of punctures, hind margin with short fine rugae. Scutellum very sparsely

punctate. Propodeal enclosure shining, large median area. Back of propodeum

dull, finely coriaceous, reticulato-carinate with some dorso-lateral parallel oblique

carinae. Mesopleura finely reticulate alutaceous, regularly minutely punctate,

interstices larger than punctures, hind margin with short rugae; punctation of

hypo-epimeral area even finer, surface dull. Anterior plate of mesepisternum dull,

oblique suture narrow, crenulate. Petiole about 2/3 length of first tergite, this

tergite little longer than broad. Pygidial area about 1.5 times as long as broad.

Second recurrent vein of fore wings interstitial or ending in third submarginal cell.

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pubescence whitish, on vertex and scutum greyish-brown, on face silvery, rarely

golden, mostly appressed. Pygidial area with golden-brown, mostly appressed,

pubescence.

Male. — Resembling female. Apex of last antennal segment below dark brown.

Fore and mid tibiae almost entirely reddish-yellow. Hind tarsal segments 1—4

yellowish-brown, base of basitarsus paler. Pronotal tubercles somewhat yellowish.

Tergites with broad reddish hind margin.

Slightly protruding median part of clypeus about 1/3 of total width of clypeal

margin. Antennal segments about 1.5 times as long as broad, segment 13 over

twice as long as broad, segments 4—11 with reddish linear tyloidea. Vertex finely

transversely striate and sparsely punctate. Scutum more strongly punctate than in

female, posterior part rugoso-punctate. Mesopleura much more strongly punctate

than in female, partly striato-punctate, a few wide interstices beneath hypo-

epimeral area. Hypo-epimeral area reticulate alutaceous, finely punctate,

interstices as large as or larger than punctures. Back of propodeum coarsely

reticulato-carinate. Gaster slender, petiole about as long as first tergite, this tergite

about 1.5 times as long as broad.

El Salvador: Quezaltepeque, 1 9, holotype, 16 July 1963, 1 g, allotype, 17 July

1963; paratypes: 2 g, 5 mi north Quezaltepeque, 23 Aug. 1961, 2 G, 3—16 July

1963, 1 g, 6 Aug. 1963; Usulutan, 2 9 and 7 3, 12 July 1963, D. Cavagnaro and M.

E. Irwin (UCD).

Guatemala: | 4, paratype, Guazacapán, 11 Aug. 1952, R. H. Painter (CU).

Costa Rica: 1 g, paratype, Liberia, Guanacasta Prov., 6 Aug. 1964, G. E.

Eickwort (KU).

Mexico: Chiapas: 1 9, 20 mi south Tuxtla Gutiérrez, 12 Aug. 1963, F. D. Parker

and L. A. Stange (UCD); 1 &, 35 mi west Tuxtla Gutiérrez, 16 Aug. 1972, G. F.

and S. Hevel (USNM). Guerrero: 1 &, Almolongo, 6000 ft, 29 July 1962, H. E.

Evans; 1 g, 10 km east Chilpancingo, 5200 ft, 30 July 1962, H. E. Evans (MCZ); 1

&, 5 mi south, 2.5 mi east Chilpancingo, 3800 ft, 5 Aug. 1962, University Kansas

Mex. Exped. (KU); 1 9, 33 mi north Taxco, 5700 ft, 29 Aug. 1963, Scullen and

Bolinger (OSU). Jalisco: 1 9, 8 mi southwest San Juan de Los Lagos, 4 Aug. 1954,

J. W. McSwain; 1 © and 3 g, Guadalajara, Crawford (UCD); 1 9 and 3 6,

Guadalajara, 17—28 July 1965, H. E. Evans, 1 g, 9 mi south Guadalajara, 24 July

1965, H. E. Evans (MCZ). Morelos: 1 ©, Yautepec, 31 July 1963, F. D. Parker and

L. A. Stange (UCD). Oaxaca: 2 9, Temaxcal, 21 Sept. 1963, K. H. Janzen and 16

July 1966, J. S. Buckett, M. R. and R. C. Gardner (UCD). Sinaloa: 1 g, Mazatlan,

15—20 Aug. 1962, H. E. Evans; 2 &, Villa Union, 17 Aug. 1962, H. E. Evans

(MCZ). Yucatan: 1 9, Chichen Itza, 29 June (MCZ). All paratypes.

The last mentioned female is accompanied by a mature Homopteron and a

handwritten note “apparently an undescribed genus and species in the subfam.

Jassinae’’. The hind basitarsi of this female are pale brown with dark brown base,

the bases of the following segments are very pale brown.

One of the females from Mexico (Morelos) has a distinctly golden pubescent

VAN LITH: New World Pluto 181

face, in another female from Mexico (Guerrero, 33 mi north of Taxco) the face is

pale golden.

The mesopleura of the female of P. punctatellus are notably finely and regularly

punctate. It very much resembles the female of P. spangleri, which, however, is

distinguished by the somewhat stronger punctation of the mesopleura, the

yellowish-white outer side of the mid tibiae and the whitish hind tarsi. The males of

P. punctatellus and of P. spangleri have strongly different tyloidea.

The males of P. punctatellus from Mexico have more shining mesopleura than

the males from El Salvador and the scutum is not rugose.

The specific name refers to the minute punctures of the mesopleura.

Pluto depressus sp. nov.

Male. — Length about 4—6 mm. Black. Mandibles yellow with reddish tips.

Underside of flagellum brownish-yellow. Pronotal tubercles whitish. Tegulae

yellowish-brown. Fore and mid tibiae reddish-yellow, tarsi whitish, last segment

brown. Basal 1/3 of hind tibiae yellowish-white, tarsi brown with paler tips of

segments. Veins of wings dark brown.

Median part of clypeal margin indistinctly protruding, indistinctly bidentate.

Frons shining, distinctly punctate, somewhat raised around anterior ocellus.

Vertex shining, sparsely finely punctate. Intercarinal space narrow. POD

somewhat larger than OOD. Antennae slender, most segments over 1.5 times as

long as broad, last segment over twice as long as broad, segments 4—7 with narrow

tyloides, nearly as long as segments, segment 8 with much shorter tyloides,

sometimes segment 9 with small shining point.

Pronotum in dorsal view with oblique lateral angles. Scutum shining, strongly

punctate, medially distinctly longitudinally depressed, hind margin of scutum

strongly triangularly depressed. Scutellum sparsely punctate. Propodeal enclosure

shining, irregularly carinate. Back of propodeum dull, finely and closely

reticulato-carinate, with parallel dorso-lateral carinae. Mesopleura shining, with

strong separate punctures, Hypo-epimeral area shining, finely punctate, interstices

about size of punctures. Mesosternum densely finely punctate. Anterior plate of

mesepisternum dull, closely punctate. Gaster slender, petiole about as long as first

tergite, this tergite over twice as long as broad. Second recurrent vein of fore wings

interstitial.

Pubescence of face and tempora silvery, mostly appressed. Rest of body whitish

pubescent, hairs long on vertex, pronotal collar and pronotal tubercles.

Female unknown.

Mexico: 6 4, holotype and paratypes, Sinaloa, Villa Union, 17 Aug. 1962, H. E.

Evans (MCZ).

P. depressus seems to be very close to P. sayi and P. abbreviatus, all having a very

finely reticulate propodeum. The number of tyloidea is reduced as compared with

these two species, but they are not considerably shorter than the segments as in

P. abbreviatus.

The specific name refers to the median and posterior depressions of the scutum.

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pluto abbreviatus sp. nov.

Female. — Length about 5—5.5 mm. Head and thorax black. Mandibles

reddish. Labrum dark reddish. Underside of antennae yellowish-red. Pronotal

tubercles and tegulae yellowish-white. Fore and mid tibiae and tarsal segments

1—4 yellowish-white, underside of tibiae dark brown. Basal third of hind tibiae

yellowish-white, tarsi except for apices brown. Petiole and greater part of first

tergite black, hind margin of first tergite, all of tergites 2—6 and sternites 2—6 red.

Veins of wings dark brown.

Median third of clypeal margin straight, indistinct lateral angles. Frons densely

finely punctate, low lateral tubercles shining, much less densely punctate. Vertex:

finely reticulate alutaceous, sparsely finely punctate. POD about 1.5 times OOD.

Intercarinal space narrow. Antennae short, clavate, segments 10—11 shorter than

broad.

Lateral pronotal angles obtuse. Scutum and scutellum finely reticulate

alutaceous, sparsely punctate. Anterior plate of mesepisternum dull. Enclosed |

area of propodeum shining, back dull, finely reticulato-carinate, dorso-laterally

finely obliquely striate. Hypo-epimeral area dull, indistinctly superficially

punctate. Mesopleura dull, finely reticulate alutaceous, finely punctate, interstices

partly larger than punctures. Mesosternum densely very finely punctate. Petiole

about 2/3 length of first tergite, this tergite somewhat longer than broad. Pygidial

area about twice as long as broad. Second recurrent vein ending in third

submarginal cell.

Pubescence silvery-white, on face mostly appressed; hairs on pygidial area dark

brown.

Male. — Length 5 mm. Pronotal tubercles yellowish-white. Fore and mid tibiae

largely dark brown, narrow yellowish-white streak on outer side. Hind tarsi when

viewed from behind very pale brown. Hind margin of first tergite, all of tergites

2—3 and greater part of sternites 2—3 red.

Antennae long and slender, segments about 1.5 times as long as broad, segments

5—13 with tyloides, short on segment 5, linear and nearly as long as segment on

segments 6—7 or 6—8, narrow oval and about half as long as segment on segments

8—12 or 9—12, an indistinct tyloides on segment 13, which segment is over twice

as long as broad.

Scutum somewhat shining, densely punctate, interstices as large as punctures or

smaller. Punctation of mesopleura stronger than in female. Back of propodeum

finely reticulate. Petiole about 2/3 length of first tergite, this tergite over 1.5 times

as long as broad. Gaster slender.

Mexico, Baja California: 1 9 holotype and 1 4, allotype, 10 mi northwest La

Paz, 6 Oct. 1941, Ross and Bohart; 1 ©, paratype, San Pedro, 7 Oct. 1941, Ross

and Bohart (CAS); | &, paratype, Vircaino Desert, 10 Oct. 1959, F. E. Strong

(UCD). A 4, collected together with the paratype (CAS) is considered to belong

to this species, but it is coloured like P. basifuscus, i.e. the gaster is almost entirely

black; the hind tarsal segments 1—4, except for the base of the basitarsus, are

whitish.

VAN LITH: New World Pluto 183

P. abbreviatus is close to P. sayi as well as to P. basifuscus. The male is easily

recognized by the elongate tyloidea, most of which are considerably shorter than

the antennal segments. The female has an almost entirely red gaster. The

extension of the red colour may be reduced, however, and further studies are

needed. The punctation of the hypo-epimeral area is indistinct, as opposed to

P. sayi and P. basifuscus. The petiole is somewhat shorter than in P. basifuscus.

The specific name refers to the abbreviated tyloidea on the last antennal

segments of the male.

Group of aerofacies

Pluto aerofacies (Malloch)

(fig. 16)

Malloch, 1933: 49, © and g (Psenia aerofacies; Texas; Mexico).

Krombein, 1951: 962 (Pluto aerofacies, comb. nov.).

Bohart & Menke, 1976: 171.

Female. — Length about 7 mm. Head and thorax black; labrum and mandibles

dark reddish, underside of flagellum orange-brown. Pronotal tubercles whitish,

tegulae reddish, veins of wings brown. Fore and mid tibiae reddish, base and outer

side somewhat yellowish, underside more or less brownish, fore and mid tarsi

reddish, first two segments whitish. Basal third of hind tibiae and tarsal segments

1—4 yellowish-white. Gaster black, hind margins of tergites and apex of last

segment somewhat reddish transparent.

Slightly roundly protruding median part of clypeal margin about half total width

of margin, straight, laterally indistinctly defined. Frons between ocelli and

antennae dull, densely punctate, sides of frons shining. Vertex and interocellar

area reticulate alutaceous, sparsely punctate. Occipital carina complete,

intercarinal space narrower than first basitarsus. Antennae clavate, segments

10—11 somewhat shorter than broad.

Lateral pronotal angles rectangular. Scutum shining, rather strongly irregularly

punctate, interstices on lateral parts a few times size of punctures. Scutellum

shining, sparsely punctate, posterior margin somewhat rugose. Propodeal

enclosure deep and shining, large median area. Back of propodeum dull,

coriaceous, coarsely reticulato-carinate. Mesopleura dull, reticulate alutaceous,

median part rather coarsely striato-punctate, finer punctate just below hypo-

epimeral area and on lower half of mesopleura, interstices larger than punctures.

Mesosternum finely punctate. Hypo-epimeral area dull, densely finely punctate.

Anterior plate of mesepisternum dull, obliquely rugose, rugae continuing into

broad oblique suture. Petiole about 2/3 length of first tergite, this tergite longer

than broad. Pygidial area slightly over twice as long as broad. Second recurrent

vein of fore wings ending in third submarginal cell.

Face pale golden, mostly appressed pubescent. Hairs on vertex and on dorsal

side of thorax greyish-brown, pygidial area dark golden-brown appressed

pubescent, remainder of body with whitish or greyish hairs.

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Male. — Length about 6—7 mm. Similar to female. Hind tarsal segments 2—4

basally somewhat infuscated. Vertex finely transversely striate. Antennal

segments 11—12 longer than broad; segments 4—12 with reddish linear tyloides.

Median part of clypeal margin less than 1/3 of total width of margin (fig. 16).

Scutum transversely rugoso-punctate. Mesopleura somewhat stronger sculp-

tured, upper part rugoso-punctate, punctures on lower part also mostly in rows,

interstices between rows partly size of punctures, partly absent. Mesosternum with

finer, separate punctures. Back of propodeum very coarsely reticulato-carinate.

Petiole about as long as first tergite, this tergite about 1.5 times as long as broad.

Pubescence of face silvery.

Materal examined: 1 3, allotype, Victoria, Texas, 8 July 1907, on Acacia sp., J.

D. Mitchell; 1 g, paratype, No. 1785, Mexico, C. F. Baker (USNM, type No.

44219).

New records from Texas: 1 ©, Brownsville, 3 May 1904 (USNM); 1 &,

Brownsville, 20—30 June 1965, H. E. Evans (MCZ); 1 9, Victoria, 21 July 1911, on

L. natans, J. D. Mitchell; 1 3, Waco, 8 June 1948, on CSC weeds, P. A. Glick

(USNM); 2 &, Hidalgo, 17 July 1954, J. G. Chillcott (CNC); 1 &, 10 mi south

Kerrville, 22 June 1942, E.S. Ross (CAS).

First record from North Carolina: 1 9, Person Co., 19 Aug. 1964, H. D. Blocker

(NCSU).

New records from Mexico: Campeche: 1 9, 20 mi east Hopelchen, 10 ft, 16

Aug. 1963, Scullen and Bolinger (OSU). San Luis Potosi: 1 9, 3.4 mi northeast El

Naranjo, 800 ft, 5 Sept. 1962, Univ. Kansas Mex. Exped. (KU). Tabasco: 1 9,

Teapa, Febr., H. H. Smith (BM). Veracruz: | 3,2 mi south Panuco, 14 Aug. 1959,

L. A. Stange and A. S. Menke (UCD). Yucatan: 1 g, Quintana Roo (Territorio),

12 mi east Peto, 29 June 1966, Univ. Kansas Mex. Exped. (KU). The males from

Mexico have the lower part of the mesopleura closely striato-punctate, with very

small or no interstices.

First record from Belize (British Honduras): 1 &, Belize (UCD).

The female of P. aerofacies is easily recognized by the pale golden face, the

coarsely sculptured back of the propodeum and the length of the petiole, the male

by the striato-punctate mesopleura, the length of the petiole and the pale hind

tarsi. The male of P. tibialis, with which Malloch (1933) compared his male of

aerofacies, has no distinct striae or rugae on the mseopleura, although the

punctures are partly placed in rows.

Pluto evansi sp. nov.

(fig. 17)

Female. — Length about 5.5 mm. Black. Mandibles, labrum and underside of

flagellum reddish-brown. Pronotal tubercles yellowish-white. Foreside of fore

tibiae and last tarsal segments orange-brown, underside of fore tibiae with brown-

ish streak, tarsal segments 1—3 of fore and mid legs whitish, mid and hind tibiae

VAN LITH: New World Pluto 185

dark brown, bases yellowish-brown, hind tarsi except for last segment yellowish-

white, base of basitarsus brown, bases of segments 3—4 more or less darkened.

Tegulae brown, veins of wings dark brown.

Median part of clypeal margin slightly but distinctly emarginate, less than half

total width of margin, lateral angles little protruding (fig. 17). Disk densely

punctate, before margin somewhat shining and coarser punctate, narrow fore

margin slightly depressed. Median part of frons densely finely punctate, laterally

with interstices about size of punctures. Vertex slightly or indistinctly alutaceous,

punctate, interstices a few times size of punctures. Frons somewhat raised, a

depression on outer side of posterior ocelli. Occipital carina complete, intercarinal

space narrower than fore basitarsus. Antennae short, clavate, segments 9-11

shorter than broad.

Lateral pronotal angles about rectangular. Scutum finely reticulate alutaceous,

punctate, interstices a few times size of punctures. Scutellum with same micro-

sculpture, sparsely punctate. Metanotum densely punctate. Propodeal enclosure

shining, large median area, back of propodeum somewhat dull, coarsely

reticulato-carinate. Mesopleura and mesosternum finely reticulate alutaceous,

finely punctate, interstices a few times size of punctures. Hypo-epimeral area

somewhat shining, very finely, almost indistinctly punctate, interstices larger than

punctures. Anterior plate of mesepisternum dull, oblique suture crenulate, carinae

continued on anterior plate. Petiole slightly over half as long as first tergite, this

tergite longer than broad. Pygidial area over 1.5 times as long as broad. Second

recurrent vein of fore wings interstitial.

Pubescence of face and pronotal collar silvery, mostly appressed, of pygidial

area brown, of rest of body whitish or greyish.

Male. — Similar to female. Length about 5 mm. Fore and mid tibiae entirely

yellowish-brown. Hind tarsal segments 1—2 whitish, base of basitarsus and

segments 3—5 pale yellowish-brown.

Clypeal margin straight. Antennal segments about 1.5 times as long as broad,

segments 6—11 with fine linear tyloides, of same pale colour as underside of

flagellum, an indistinct tyloides on segment 5. Posterior 2/3 of scutum almost

smooth, punctation as in female. Gaster slender, petiole about 3/4 length of first

tergite, this tergite over 1.5 times as long as broad.

Mexico: Sonora: | ©, holotype, and 1 &, allotype, 10 mi southeast Alamos, 29

June 1963, F. D. Parker and L. A. Stange (UCD). Sinaloa: 1 9, paratype,

Chupaderos, 4 July 1963, F. D. Parker and L. A. Stange (UCD). Morelos: | 9,

paratype, Huajintlan, 2800 ft, 28 May 1959, H. E. Evans (CU).

Pluto emarginatus sp. nov.

(fig. 18)

Female. — Length about 7 mm. Black. Mandibles and underside of flagellum

reddish-brown. Pronotal tubercles and tegulae brown. Veins of wings dark brown.

Foreside of fore tibiae yellowish-brown, fore and mid tarsi and base of hind tibiae

pale yellowish-brown. Hind tarsi brown, apices of segments paler.

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Anterior margin of clypeus (fig. 18) emarginate, with distinct triangular lateral

teeth, distance between teeth about half total width of margin. Frons densely

finely punctate, raised lateral parts sparsely punctate. Vertex sparsely punctate.

POD wider than OOD. Intercarinal space about half width fore LEE EU

Antennal segments 9—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum very finely reticulate

alutacous, punctate, interstices mostly a few times size of punctures. Scutellum

with same sculpture, sparsely punctate. Back of propodeum coarsely reticulato-

carinate, carinae oblique on dorso-lateral parts; propodeal enclosure shining,

large median area. Mesopleura, hypo-epimeral area and mesosternum slightly

alutaceous. Mesopleura sparsely finely punctate, hind margin finely rugulose.

Hypo-epimeral area very finely punctate, interstices wider than punctures.

Anterior plate of mesepisternum dull, oblique suture crenulate. Petiole about 2/3

length of first tergite, this tergite about 1.5 times as long as broad. Pygidial area

about twice as long as broad. Second recurrent vein of fore wings interstitial.

Face and pronotal collar silvery, mostly appressed, pubescent. Pubescence of

pygidial area dark golden-brown, of rest of body greyish or whitish. Male

unknown.

Mexico: 1 9, holotype, 6 mi south Temixco, Morelos, 16 July 1963, F. D. Parker

and L. A. Stange (UCD).

P. emarginatus is distinguished by the distinctly emarginate clypeal margin and

the almost shining, sparsely but distinctly punctate mesopleura. It seems to be

closely related to P. evansi, which has a slightly emarginate clypeal margin, but in

this latter species the petiole is somewhat shorter and the pronotal tubercles and

hind tarsi are whitish.

The specific name refers to the emargination of the clypeal margin.

Group of rufibasis

Pluto rufibasis (Malloch)

(figs. 19—21)

Malloch, 1933: 53, 9 and @ (Psenia rufibasis; Georgia), 54, 9 and g (Psenia marginata; Louisiana,

South Carolina).

Krombein, 1951: 962 (Pluto rufibasis, comb. nov.), 962 (Pluto marginatus, comb. nov.); 1964: 18 (Pluto ru-

fibasis; Florida).

Bohart & Menke, 1976: 171 (Pluto marginatus), 171 (Pluto rufibasis).

Van Lith, 1976: 154—158 (Pluto rufibasis; Pluto marginatus syn. nov. of rufibasis).

For the redescription of P. rufibasis cf. Van Lith (1976). Clypeal margin of

female: fig. 19.

One of the paratype males (marginata Malloch) from Louisiana is only 6 mm

long, another male from this state has a length of 7 mm, the male from Mississippi

is not longer than 5.5 mm. Their normal length is 8—9 mm. Genitalia: figs. 20—21.

Material examined: | 4, allotype Psenia rufibasis Malloch, Tifton, Georgia,

VAN LITH: New World Pluto 187

Ashmead (USNM, type No. 44224). Psenia marginata Malloch: 1 3, allotype and 5

d, paratypes, Louisiana, 2336, 2337 and 2392, C. F. Baker; 1 9 and 1 &,

Opelousas, Louisiana, G. R. Pilate; | &, paratype, Buckfield Plantation, Yemas-

see, South Carolina, | Oct. 1926, J. T. Rogers (USNM, type No. 44225).

New records from USA: Florida: Lake Placid, 2 9, 20—26 Juni 1962, K. V.

Krombein (USNM), 3 © and 5 &, May 1967, G. Heinrich (HT); Archbold

Biological Station, Highlands Co., 3 9 and 1 g, 13—14 Oct. 1964, P. H. Arnaud,

Jr. (CAS); Fort Myers, Lee Co., 3 3, 5—20 May 1967, 1 © and 6 g, 19—25 April

1971, P. P. Babiy (ZSM); Weeki Wachi Springs, Hernando Co., 1 &, 16 Aug. 1968,

G. F. Hevel (USNM); Tall Timbers, 1 © and 2 g, 12 June—11 July 1971, R. H.

Arnett (HT). Louisiana: 2 © and 11 Z, Rapides Parish, 18 May—19 June 1973,

Peter Rush (HT). Maryland: Calvert Co., 1 9, 17 Aug. 1949, D. G. Shappirio

(USNM). Mississippi: Utica, 1 3, Ashmead (USNM). North Carolina: Wake Co., |

3, 6 Sept. 1951, H. and M. Townes (HT); Cumberland Co., Fort Bragg, 3 © and |

dg, 6—13 June, 16—25 Aug., 17—20 Sept. 1967, J. D. Birchim (CAS). South

Carolina: Greenville, 1 9, 7 June 1952, G. and L. Townes (HT); Seneca, 1 9, 15

June 1961, W. H. Anderson (USNM). Virginia: Montgomery Co., Westmoreland,

2 9,29 Aug. 1954, D. G. Shappirio (USNM).

The study of this material has confirmed my opinion (Van Lith, 1976) that

rufibasis and marginatus are conspecific, the species showing great tendency to

erythrization in the southern parts of Florida and in Georgia. The females from

Lake Placid, Highland Co. and the greater part of the specimens from Fort Myers

are largely red. The rest of the material from Fort Myers, the male from

Mississippi and the females from North and South Carolina, Virginia and

Maryland have a dark petiole and more or less darkened tergites and femora.

Group of albifacies

Pluto albifacies (Malloch)

(fig. 22)

Malloch, 1933: 50, © (Psenia albifacies; Iowa).

Krombein, 1951: 962 (Pluto albifacies, comb. nov.); 1967: 396 (Texas).

Evans, 1959: 140—141 (Pluto albifacies?, larvae; Texas); 1968: 1343— 1344 (Pluto albifacies; prey).

Bohart & Menke, 1976: 171.

Female. — Length about 6.5 mm (8 mm according to Malloch). Black.

Mandibles yellowish-red, labrum dark reddish. Underside of flagellum orange-red.

Anterior margin of clypeus dark reddish transparent. Fore and mid tibiae

yellowish-red, outer side whitish, tarsi whitish, last segment yellowish-red. Basal

third of hind tibiae and tarsal segments 1—4 yellowish-white, segment 5 brown.

Pronotal tubercles whitish. Tegulae yellowish or reddish. Hind margins of tergites

reddish transparent, sides of tergites 2—3 and all of sternites 2—3 sometimes

reddish or reddish-brown. Veins of wings dark brown.

Median part of clypeal margin straight, over 1/3 of total width of margin,

laterally with obtuse angle (fig. 22). Disk of clypeus dull, densely punctate. Frons

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

densely punctate, laterally with distinct almost impunctate tubercle. Vertex finely

reticulate alutaceous, sparsely finely punctate. POD about as large as OOD.

Occipital carina complete, latero-ventrally with thickening or angle; intercarinal

space about half width of first basitarsus.

Lateral pronotal angles rectangular. Scutum dull, reticulate alutaceous, punc-

tate, interstices on central part larger than punctures. Scutellum dull, anterior half

sparsely punctate, hind margin densely punctate. Propodeum dull, back finely

reticulato-carinate, dorso-laterally with fine parallel, oblique carinae; median part

of enclosure somewhat shining, usually irregularly carinate. Mesopleura dull or

faintly shining, finely punctate, interstices size of punctures or less. Mesosternum

very finely densely punctate. Hypo-epimeral area dull, densely finely, almost

indistinctly punctate. Anterior plate of mesepisternum dull, oblique suture

crenulate. Petiole nearly as long as tergite 1, this tergite somewhat longer than

broad. Pygidial area about twice as long as broad. Second recurrent vein of fore

wings ending in third submarginal cell.

Face densely silvery appressed pubescent, also some long erect hairs.

Pubescence of vertex and dorsal side of thorax greyish, of pygidial area dark

golden-brown, of rest of body whitish, dense on back and sides of thorax.

First description of male. — Length about 5 mm. Colour as in female. Antennae

slender, segments about 1.5 times as long as broad, segments 4—12 with black

linear tyloides, short on segment 12. Underside of flagellum yellowish-red.

Punctation of scutum rather dense, interstices mostly smaller than punctures,

punctures partly in rows. Lower 2/3 of mesopleura closely finely punctate, on

upper part a few interstices larger than punctures. Hypo-epimeral area finely but

distinctly densely punctate. Back of propodeum as in female. Petiole as long as

first tergite, this tergite about 1.5 times as long as broad. Pubescence of face very

silvery.

I have not seen the type (female) from Sioux City, Iowa, which was captured by

C. N. Ainslie on 13 July, 1929 (USNM, type No. 44220), but I could examine a

series of other specimens from Iowa, also collected by Ainslie: Sioux City, 12 9

and 3 4, July-Aug. 1925, 1926, 1927, 1928, 1931—1934, three of these females

without date; 1 © and I g, Sergeant Bluff, 10 July and 3 Aug. 1933 (USNM).

In addition I could study a female from Texas, Red River, Wilbarger Co., 4 Juli

1956, Biol. note No. 1184 (USNM). This is apparently one of the two adult

specimens from the nests containing the larvae described by H. E. Evans (1959:

140—141; total of females in this colony at least 200). In 1968 Evans presented

more details regarding the same nests and he also stated that their food consisted

of leafhoppers all belonging to the Cicadellid Opsius stactogalus Fieber. About 1/3

of the 50 examples were nymphs.

Group of tibialis

Pluto tibialis (Cresson)

Cresson, 1872: 227, 9 and g (Mimesa tibialis; Texas); 1916: 102, 9 (Amer. Ent. Soc. type No. 2048).

VAN LITH: New World Pluto 189

Dalla Torre, 1897: 355.

Fox, 1898b: 18, 9 (Psen tibialis; District of Columbia).

Ashmead, 1899: 225.

Viereck, 1901: 342, 9 and g (Neofoxia tibialis).

Malloch, 1933: 49 (Psenia tibialis; Texas, District of Columbia, Louisiana).

Krombein, 1951: 962 (Pluto tibialis comb. nov.; District of Columbia, Virginia, Tennessee, Louisiana,

Missouri, Texas); 1967: 396 (Alabama, South Carolina).

Bohart & Menke, 1976: 171.

Female. — Length 6—7.5 mm. Head and thorax black. Mandibles reddish;

underside of flagellum orange-brown. Pronotal tubercles whitish. Tegulae whitish

or reddish, veins of wings dark brown. Fore and mid tibiae and last tarsal segments

reddish or reddish-brown, outer side of tibiae and tarsal segments I—3 or 1—4

yellowish-white. Basal third of hind tibiae and tarsal segments 1—4 yellowish-

white, last tarsal segment brown. Gaster black, hind margins of tergites and last

segment reddish.

Apex of clypeus straight, median part less than half total width of margin,

laterally with indistinct teeth (cf. fig. 24. arenivagus). Central part of frons densely

punctate, vertex finely reticulate alutaceous, sparsely punctate, back of head also

finely transversely striate. POD larger than OOD. Occipital carina complete,

intercarinal space linear. Antennae somewhat clavate, segments 10—11 about

quadrate.

Lateral pronotal angles rectangular. Scutum shining, strongly punctate,

interstices often larger than punctures, also some very small punctures. Scutellum

shining, sparsely punctate, hind margin rugoso-punctate. Propodeal enclosure

medially irregularly reticulate, laterally some oblique carinae. Back of propodeum

dull, irregularly but not coarsely reticulato-carinate, with some fine oblique dorso-

lateral carinae. Hypo-epimeral area dull, longitudinally striato-punctate, striae on

lower part indistinct. Mesopleura dull, reticulate alutaceous, upper half moder-

ately strongly, partly coarsely punctate, somewhat longitudinally striate, lower

part with finer punctures in rows but no distinct striae, interstices about size of

punctures; hind margin finely rugulose. Mesosternum densely finely punctate.

Anterior plate of mesepisternum vertically striate, oblique suture crenulate.

Petiole about half as long as first tergite, this tergite somewhat longer than broad.

Pygidial area about 2.5 times as long as broad. Second recurrent vein of fore wings

ending in third submarginal cell.

Face with silvery appressed pubescence and long erect hairs. Vertex and dorsal

side of thorax greyish-brown pubescent, pubescence of pygidial area very dark

brown, of rest of body greyish-white.

Male. — Length 6—6.5 mm. Colour as in female. Antennae slender, segments

longer than broad; segments 5—10 with linear black tyloidea, on segment 10 often

shorter than segment, sometimes also a short tyloides on segments 4 and 11. Vertex

densely punctate, scutum strongly punctate, interstices mostly smaller than

punctures. Back of propodeum coarsely reticulato-carinate. Mesopleura stronger

and very closely punctate, punctation regular, sometimes on upper and/or lower

part interstices larger than punctures, no distinct rugae or striae, hind margin

coarsely rugulose. Hypo-epimeral area coarsely rugoso-punctate. Petiole about

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

2/3 or 3/4 length of first tergite, this tergite about 1.5 times as long as broad.

Material examined: Texas: 1 ¢, Belfridge (USNM, type No. 1718; no antennae);

1 ©, Paris, 26 Aug. 1905, F. C. Bishopp; 1 9, New Braunfels, 17 May 1906, J. C.

Crawford; 1 ©, Plano, July 1907, catfield, E. S. Tucker, “Neofoxia tibialis”

(USNM). Louisiana: 1 9, No. 2337, “compared with holotype of Mimesa tibialis

Cr. at ANSP (Academy of Natural Sciences, Philadelphia) 1950, K. V. Krombein”,

3 © and9 J, Nos. 2236, 2335, 2336, 2392, 2524, 2525, 2565, 2566, C. F. Baker; 2 9

and 3 Z, Opelousas, Pilate (USNM). Virginia: | 9, Clifton, 30 June 1935, K.V.

Krombein (KVK). Alabama: 2 © and 1 &, Montgomery, 23 June, K. V. Krombein

(KVK). District of Columbia: 1 3, 20 July 1879, 349, T. Pergande, ‘‘Mimesa tibialis

Cress., Cress. 80°’; this is probably the male to which Malloch (1933) refers when

he writes: “the District of Columbia male lacks a sensory area on the apical three

flagellar segments”. In his description of the male Malloch states that the tyloidea

are present on all but the apical and basal two segments of the flagellum and that

there is some variation in the presence or absence on the penultimate segment.

Unfortunately the abovementioned type from Texas has no antennae.

New records from Texas: Fodor, 1 © and 1 Z, 17 May 1898, Birkmann, “tibialis

Cr. det. Kohl” (NMW); Waco, 2 © and 5 Z, 16 June and 13 July 1948, 29 June

1949, on bluestem grass and on cotton, P. A. Glick. Louisiana: 1 ©, Tallulah, 1

July 1948, Bug Catcher Exp., R. C. Gaines (USNM).

First records from Florida: | 9, Fort Myers, 16 April 1971, P. P. Babiy (BSM).

Missouri: 2 &, 14 June and 17 July, Columbia, 4 pm — 7 am, Malaise trap, F. D.

Parker. South Carolina: 1 &, Seneca, 25 July 1962, on pine tree, R. D. Eikenbary.

Tennessee: | &, Cedar Glade Area, Mid Tennessee, 31 May, Adelphia Meyer,

““Psen tibialis (Cress.) det. Sandhouse” (USNM); 2 © and 6 4, Madison, Davidson

Co., 10 June—22 July 1967, P. P. Babiy (BSM). Virginia: 1 g, Clifton, 15—23 July

1933, J.C. Bridwell (USNM).

The records from Nebraska (Mickel 1918) seem to be doubtful and have

apparently not been accepted by Malloch (1933).

Pluto arenivagus arenivagus Krombein

(figs. 23—25)

Malloch, 1933: 48 and 58, 9 (Psenia angulicornis var.; Georgia).

Krombein, 1949: 268—269, © and g (Pluto arenivagus; North Carolina, Georgia); 1951: 962; 1953: 132

(North Carolina); 1954: 233 (Florida); 1958: 189 (Florida).

Bohart & Menke, 1976: 171.

Both sexes are characterized by the ventral part of the occipital carina and the

hypostomal carina being almost contiguous (fig. 23). The original description

(Krombein, 1949) can be supplemented with the following details. Clypeal margin:

fig. 24. Mesopleura densely and strongly punctate, in female partly striato-

punctate. Petiole about 3/4 length of first tergite. In female at least tergite 2

entirely red; gaster of male black with reddish hind margins of segments, rarely

also bases of tergites 2—3. Tarsi whitish to yellowish, last segment of fore and mid

VAN LITH: New World Pluto 191

tarsi yellowish-red, of hind tarsi brown. Second recurrent vein of fore wings ending

in third submarginal cell. Pygidial area of female: fig. 25.

BES N

i went

Fig. 22. Pluto albifacies (Malloch), 9, clypeus. Figs. 23—25. P. arenivagus arenivagus Krombein, 9, pa-

ratype; 23, head in ventral aspect; 24, clypeus; 25, pygidial area. Figs. 26—31. Clypeus; 26, of P. argen-

tifrons (Cresson), 9, Cuba; 27, P. atricornis (Malloch), 9, St. Croix; 28, P. colonensis sp. n., 9, holoty-

pe; 29, P. alphitopus sp. n., 9, holotype; 30, P. stramineipes sp. n., 9, holotype; 31, P. strigellus sp. n., 9,

holotype.

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Material examined: 1 &, holotype, Kill Devil Hills, North Carolina, 28 May

1948, K. V. Krombein (USNM, type No. 14058); 1 9 and | &, paratypes, same

locality, 30 May and 27 May 1948; 5 9, same locality, 30 May—5 Aug.

1950—1958, K. V. Krombein (KVK); I 9, paratype, Tifton, Georgia, Ashmead

collection, labelled “angulicornis var., paratype No. 44229" (USNM, paratype No.

59312); 1 9, Venus, Highlands Co., Florida, 17 April 1963, K. V. Krombein

(USNM).

New records from Florida: 1 &, Oneco, 29 March 1955, J. C. Martin (CNC); 2

3, Fort Myers, Lee Co., 28 April and 3 May 1967, 1 9 and I &, Fort Myers, 17

April 1971, P. P. Babiy (BSM); I 9, St. Joseph St. Park, Gulf Co., 1—3 May 1970,

W. W. Wirth (USNM); 1 9, Big Pine Key, Monroe Co., 23 June 1971, W. H.

Pierce (FSC).

P. arenivagus is closely related to P. tibialis, but the male is distinguished by the

presence of tyloides also on the antennal segments 11—12, the female by the

reddish second gastral tergite.

Pluto arenivagus cubanus subsp. nov.

Female. — Very similar to nominate subspecies. Fore and mid tibiae darker,

underside of fore tibiae brown, mid tibiae dark brown, apices and outer side

yellowish-white. Hind margin of first tergite, second tergite except for large basal

brown mark, third tergite and sternites 2—4 red.

Punctation of scutum finer, interstices wider, mostly twice size of punctures.

Punctation of mesopleura somewhat coarser. Hypo-epimeral area distinctly

longitudinally striato-punctate. Back of propodeum more coarsely reticulate.

Male unknown.

Cuba: | 9, holotype, Mantua, Prov. Pinar del Rio, June 1968, P. Alayo (UCD).

This is apparently one of the specimens recorded as “Pluto Sp. A” by P. Alayo

Dalmau (1973).

The differences between the holotype of cubanus and the continental form are

small. Further studies are needed to determine whether these differences are

constant.

Group of argentifrons

Pluto argentifrons (Cresson)

(fig. 26)

Cresson, 1865: 152, © and g (Psen argentifrons; Cuba).

Dalla Torre, 1897: 347 (Psen argentifrons; Cuba).

Ashmead, 1900: 305 (Cuba, Jamaica).

Cresson, 1916: 102 (Cuba).

Pate, 1946: 6—9 (Pluto argentifrons, comb. nov.; Cuba).

Alayo, 1973: 181 (Cuba, abundant).

Bohart & Menke, 1976: 171 (Cuba).

VAN LITH: New World Pluto 193

Pate (1946) furnished a detailed redescription. The following supplementary

remarks can be made. Intercarinal space less broad than first basitarsus. The back

of the head and the tempora are finely striate, the mesopleura of the female very

closely very finely punctate, anteriorly somewhat striato-punctate; in the male the

mesopleura are finely ruguloso-punctate. The second recurrent vein of the fore

wings ends in the third submarginal cell. Clypeal margin: fig. 26.

Malloch (1933) apparently confused P. argentifrons and P. atricornis (Malloch)

and he omitted to mention the former species. The authorities of the National

Museum of Natural History, Washington, D. C., kindly enabled me to examine the

two females from Cuba, incorrectly described by him as the female of P. atricornis

Malloch, 1933.

Pate (1946) suspected that the specimens reported by Ashmead on Jamaica

would be found to belong to another species, but I could not find any substantial

differences between the material from Cuba and the specimens from Jamaica.

Material examined: Cuba: | 9, Baraguä, 2 Dec. 1925, T.P.R.F., Ent. No. 346, at

light, C. F. Stahl; 1 9, Santiago de las Vegas, 21 June 1917, C. 19, P. Cardin,

labelled ““Mimesa argentifrons Cress., det Roh.”, both specimens misidentified by

Malloch as the female of P. atricornis from Puerto Rico (USNM, type No. 44228,

allotype and paratype). Jamaica: 1 g, Portland, labelled “‘Neofoxia argentifrons

Cress., Roh.” (USNM); 1 3, Placetas to Remedios, Sta. Clara, 20 July 1940, J.C.

Bradley (CU).

New records from Cuba: | 9, Baragua, 14 Sept. 1925, T.P.R.F. Ent. No. 346, on

grasses, C. F. Stahl, 1 g, Baragua, 2 Dec. 1925, T.P.R.F. No. 344, taken on

legumes, C. F. Stahl (MCZ); 2 9, Cabanas, Pinar del Rio, 5—8 Sept. 1913

(AMNH); 1 g N. Banks; 1 g, Central Soledad, Cienfuegos, 19 Aug. 1932, B. B.

Leavitt; 2 g, Soledad, 192. and 1925; 2 g, Soledad, Santa Clara Prov., 7—12 June

1939, C. T. Parsons; 2 &, Botanical Gardens, Central Soledad, Cienfuegos, 17

Aug. 1930, Richard Dow, with label “Psen (Psen) argentifrons”; 1 9, Belmonte,

Central Soledad, Cienfuegos, 10 Sept. 1930, Hoya Colorado, Hav. Pr., 23 Aug.

1917, H. Morrison; | 9, near Santiago, 31 Aug. 1917, H. Morrison (USNM); | &,

Soledad nr. Cienfuegos, 6—20 Aug., N. Banks; 1 9, Soledad, 24 Aug. 1927, C. 112,

on flowers of Kallstroemia maxima, J. G. Myers; 1 g, Havana, Baker 3614, P.

Cameron Coll. 1914—110, with label “Psen argentifrons Cr.” (BM); 1 9 and 1 2,

Havana, Baker, “Pluto argentifrons (Cresson) det. L. Stange” (IML); 2 © and 1 &,

Pinar del Rio, 16—29 May 1933, H. J. MacGillavry (MA); 2 © and 19 g, Soledad

nr. Cienfuegos, 6—20 Aug., Richard Dow, ‘‘Psen (Psen) argentifrons Cress.”; 1 3,

Banes, 4—11 June 1927, F. T. Baird; 2 g, Regla to Casa Blanca, 13 Aug. 1930

(MCZ); 2 &, Havana, Baker, Univ. of Kansas, Lot 940, ‘‘Psen (Psen) argentifrons”

(KU); 2 &, Baracoa, Aug. 1902, Aug. Busck; 1 ©, Havana, Baker, “Psen

argentifrons Cr.”;2 &, Havana, Baker; 1 4, Santiago de las Vegas, 3 July 1917, P.

Cardin; 1 9 and 3 g, Laquito Marianno Hab., March 1967, P. Alayo, “Pluto

argentifrons (Cress.) det. P. Alayo 1968” (USNM).

New records from Jamaica: 2 9 and 11 g, Liguanea Plain, Nov.—Dec. 1911, C.

T. Brues; 1 g, Newton, 3000 ft, Jan. 1912, C. T. Brues (MCZ); 1 9, Univ., Mona,

16 June 1970, A. Raw; 1 dg, St. Andrew Univ., 16 June 1971, Phyllanthus

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

(Euphorbiaceae), A. Raw (BM); 2 9, Trelawny, Good Hope, 22 Aug. 1966, H. F.

Howden; 1 ©, Trelawny, Duncans, 23 Aug. 1966, Howden and Becker (CNC); |

Q, Duckenfield, E. Jamaica, 12 July 1962, G. F. Mees (ML); 2 9, United Fruit

Farm, Spanishtown, 11 Sept. 1917, H. Morrison; 1 8, Kingston, 9 Sept. 1917, H.

Morrison; 1 &, Portland, Dover, 7 Aug. 1947, G. B. Thompson; 1 &, St.

Catharine,Amity Hall, Bushy Park, 20 March 1947, G. B. Thompson; 1 9,

St. Catharine, Old Harbour, 20 Oct. 1957, T. H. Farr; 1 © and 1 g, Trelawny

Parish, Windsor Estate, 29 July 1962, Farr, O. and R. Flint (USNM).

First records from Mexico: Yucatan: 1 9, Yaxcopoil, Oct. 1950, N. L. H. Krauss

(USNM); Chiapas: | &, San Cristóbal de las Casas, 7200 ft, 30 May 1969, W. R. M.

Mason (CNC). First record from Nicaragua: 7 &, Granada, Baker (UCD). |

P. argentifrons is easily distinguished by the very densely and finely punctate

mesopleura, more coarsely punctate in the male, and the brown hind tarsi. It

resembles P. atricornis which has a more eastern distribution, from Hispaniola to

Antigua (Leeward Islands). In the female of the latter species the sculpture of the

mesopleura is much coarser, distinct transverse rugae being present; the male of

atricornis has no tyloidea on the fifth and sixth antennal segments which

argentifrons has.

Pluto atricornis (Malloch)

(fig. 27)

Malloch, 1933: 57—58, ¢ only, 9 misidentified (Psenia atricornis; Puerto Rico).

Pate, 1946: 9—10, & (Pluto atricornis, comb. nov.).

Bohart & Menke, 1976: 171.

First description of female. — Length about 6.5 mm. Black; mandibles and

underside of flagellum reddish-brown. Tegulae yellowish-red. Pronotal tubercles

whitish. Foreside of fore tibiae, base and outer side of mid tibiae and basal third of

hind tibiae, fore basitarsi with next segment, and mid basitarsi an -white.

Veins of wings dark brown.

Protruding median part of clypeus less than half total width of margin, very

slightly emarginate with indistinct tooth in the middle (fig. 27) and small rec-

tangular lateral angles; disk densely punctate. Frons densely punctate.

Interocellar area sparsely punctate, vertex more densely so with tendency to

transverse striation. Intercarinal space about as wide as first basitarsus. Antennal

segments 10—11 shorter than broad.

Lateral pronotal angles sharp. Scutum and scutellum shining, scutum moder-

ately densely finely punctate, interstices partly larger than punctures, scutellum

sparsely punctate. Mesopleura finely alutaceous, upper half with coarse

longitudinal rugae and indistinct punctures, lower half punctate, interstices about

size of punctures, mesosternum finer and more widely punctate. Hypo-epimeral

area densely finely punctate. Anterior plate of mesepisternum with oblique rugae

which continue into anterior oblique suture. Enclosed area of propodeum shining,

back somewhat dull, densely reticulato-carinate, dorso-lateral carinae parallel.

Second recurrent vein of fore wings ending in third submarginal cell. Petiole about

VAN LITH: New World Pluto 195

half length of first tergite, this tergite little longer than broad. Pygidial area about

twice as long as broad.

Face pale golden or silvery, mostly appressed pubescent, pygidial area dark

brown or golden-brown pubescent, rest of body yellowish-grey.

Male. — More slender than female. Fore and mid tibiae yellowish-red.

Protruding median part of clypeus about 1/4 of total width of margin. Antennae

slender, segments about 1.5 times as long as broad, segments 7—12 with long

linear tyloides, segment 13 with short tyloides. Mesopleura more shining, on dorsal

half irregularly rugoso-punctate, on lower part rather densely punctate. Petiole

about 3/4 length of first tergite, this tergite about 1.5 times as long as broad. Face

silvery.

Material examined: Puerto Rico, | 9, Mayaguez, 13 May 1931, No. 8, Oramas,

with label “This is apparently the true 9 of atricornis Mall., det. Karl V. Krombein

1941” (USNM).

New records from Puerto Rico: 1 &, Coamo Sp’gs, 5—7 June 1915, Dept.

Invert. Zool. No. 22320; 1 4, Manati, 27—29 June 1915 (AMNH); 1 &, “L jas”,

Sept.—Nov. 1960, M. Santiago (USNM).

First records from Haiti: 3 4, Damien, 7 Aug. 1977; 1 © and I g, Léogane, 10

Aug. 1977, A. Pauly (FAG).

First records from Dominican Republic: 3 3, Santo Domingo, 8 mi up Macoris

River, 16 July 1917, H. Morrison; 1 g, San Christóbal, 26 July 1917, H. Morrison

(USNM).

First records from Virgin Islands: I 9, St. Croix, Christansted, by net No. 1296,

USNM Ins. No. 161551, Lot No. 41.20612, 1940, H. A. Beatty (USNM); 1 9, St.

Croix, East Point, 3—7 Febr. 1969, H. E. Evans, with label “Pluto atricornis Mall.,

Q, det. H. Evans 1969” (MCZ).

First record from Leeward Islands: 1 4, Antigua I., Body Ponds, 12 June 1965,

E. and S. Geijskes (ML). This male has a somewhat longer petiole, about 4/5

length of first tergite.

P. atricornis differs from other Pluto in having a broad intercarinal space, rugose

mesopleura, a short petiole and a black gaster, the male also in having no tyloides

on antennal segments 4—6.

Pluto alphitopus sp. nov.

(fig. 29)

Female. — Length about 6 mm. Black. Mandibles reddish, apical half of

underside of flagellum reddish-brown. Bases of fore and mid tibiae yellowish-

white, foreside of fore tibiae yellowish-brown. Fore and mid tarsi yellowish-white,

last segment brown. Base of hind tibiae yellowish-brown; basal 3/4 of hind

basitarsus brown, apex and segments 2—4 yellowish-brown with somewhat darker

bases. Tegulae yellowish-brown. Pygidial area dark reddish. Veins of wings dark

brown.

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Clypeal margin very indistinctly trilobate or bisinuate (fig. 29), disk convex,

densely punctate. Frons densely punctate, vertex finely punctate and finely

transversely striate. POD somewhat larger than OOD. Intercarinal space narrow.

Antennae short, slightly clavate, segments 9—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum shining, anterior margin

densely punctate, rest of scutum irregularly punctate, interstices often larger than

punctures. Scutellum shining, mostly sparsely punctate, posterior margin densely

punctate. Back of propodeum dull, coriaceous, mostly obliquely striate, enclosure

with a few oblique carinae and large median area. Mesopleura, mesosternum and

hypo-epimeral area dull, densely punctate, upper part of mesopleura also finely

longitudinally striate. Hypo-epimeral area dull, densely finely punctate. Anterior

plate of mesepisternum striate, oblique suture indistinctly crenulate. Petiole about

half length of first tergite, this tergite about as long as broad. Second recurrent

vein of fore wings ending in third submarginal cell.

Face pale golden, scutum greyish-brown, pygidial area dark brown, rest of body

yellowish-silvery or yellowish-grey pubescent.

Male. — Resembling female. Length about 6 mm. Fore tibiae, outer side of mid

tibiae and base of hind tibiae yellowish-brown. Tarsi pale yellowish-white except

for last segment and base of hind basitarsi, fore and mid basitarsi almost whitish.

Antennae slender, segments 11—12 about 1.5 times as long as broad, last

segment about 2.5 times as long as broad, segments 7—10 in lateral view raised in

the middle. Segments 4—13 with linear tyloides, on segments 4—6 as long as

segment, shorter on segments 7—12, very short on segment 13.

Scutum more closely punctate or somewhat rugoso-punctate. Scutellum densely

punctate, finely reticulate alutaceous. Lower part of propodeum more distinctly

reticulato-carinate. Punctation of mesopleura more superficial than in female and

punctato-reticulate, no distinct rows of punctures; hind margin finely coriaceous.

Punctation of hypo-epimeral area finer than on mesopleura. Gaster slender,

petiole about 2/3 or half length of first tergite, this tergite about 1.5 times as long as

broad. Face silvery, rest of body silvery or whitish pubescent.

Mexico: Guerrero: 1 ©, holotype, 20 mi west Acapulco, 10 Aug. 1962, 1 G (no

antennae), paratype, same locality, 11 Aug. 1962, Univ. Kansas Mexican Exped.

(KU). Jalisco: 1 &, allotype, Jocotepec, 5000 ft, 11 July 1959, H. E. Evans (CU).

Michoacan: 1 &, paratype, 11 mi east Apatzingan, 20 Aug. 1954, E. G. Linsley, J.

W. MacSwain, R. F. Smith (UCD). Oaxaca: 1 &, paratype, Crawford (UCD); hind

basitarsi entirely whitish. Puebla: 1 ¢, paratype, 30 mi northwest Acatlán, 14 Aug.

1972, G. F. and S. Hevel (USNM). I

The male of P. alphitopus resembles that of P. castaneipes, but its tyloidea are

different and the tarsi are whitish. The female of the latter species is still unknown.

The sculpture of the mesopleura of the female of P. alphitopus is very similar to

that of P. strigellus; unfortunately the male of latter species has not yet been found.

The specific name, derived from the greek nouns dAgitov androuo, refers to the

floury colour of the tarsi of the male.

VAN LITH: New World Pluto 197

Pluto castaneipes sp. nov.

Male. — Length about 6.5 mm. Black. Outer 2/3 of mandibles dark reddish.

Underside of flagellum of antennae brown, of last segment orange-brown. Apical

half of foreside of fore femora, fore tibiae and base of mid tibiae reddish-brown;

hind tibiae and all tarsi brown, base of hind tibiae paler. Veins of wings black.

Median part of clypeal margin somewhat protruding, about 1/4 of total width of

margin. Frons densely punctate, no interstices. Vertex transversely striate, punc-

tation indistinct. POD about 1.5 times OOD. Intercarinal space linear. Antennae

slender, segments 11—12 over 1.5 times as long as broad, segment 13 over twice as

long as broad. Segments 4—12 with linear tyloidea, as long as segments, on

segments 11—12 somewhat shorter, segment 13 with very small tyloides. Tyloidea

below much roundly raised, in lateral view highest point on segments 7—11 near

middle of segment.

Lateral pronotal angles about rectangular. Scutum shining, anteriorly dull,

densely coarsely punctate, interstices distinct, some superficial transverse rugae.

Scutellum shining, densely punctate, interstices mostly larger than punctures.

Propodeum dull, enclosure medially irregularly carinate, back coarsely reticulato-

carinate. Mesopleura dull, reticulate alutaceous, densely punctate and striato-

punctate, hind margin coarsely coriaceous. Hypo-epimeral area dull, densely

punctate. Anterior plate of mesepisternum dull, finely striato-punctate, oblique

suture narrow. Mesosternum somewhat shining, densely finely punctate. Second

recurrent vein of fore wings ending in third submarginal cell. Petiole about 2/3 or

3/4 length of first tergite, this tergite about 1.5 times as long as broad.

Face, tempora and pronotal collar silvery, mostly appressed, pubescent. Also

rest of body silvery-white pubescent, dense and somewhat appressed on

mesopleura.

Female unknown.

New Mexico, USA: 1 &, holotype, Rodeo, 4—5000 ft, 14 Sept. 1958, at

Asclepias, H. V. Weems Jr. (FSC).

Texas, USA: 1 3, paratype, Howard Co., 15 mi northwest Big Spring, US

Highway 87, 13 June 1963, D.C. and K. A. Rentz (CAS).

The long antennal segments with much raised tyloidea and the dark tarsi

distinguish P. castaneipes from other black species belonging to the group of P.

argentifrons; it differs from the next species by the sculpture of the propodeum and

the dark colour of the pronotal tubercles and of the gaster.

The specific name refers to the brownish colour of the legs.

Pluto rugulosus sp. nov.

Male. — Length about 6 mm. Head and thorax black; mandibles yellow with

reddish tips, underside of flagellum orange-brown. Thorax black, pronotal

tubercles yellow. Foreside of fore femora and fore tibiae yellowish-red, tarsi

yellowish-brown. Apex of mid femora and foreside of tibiae yellowish-red, tarsi

brown, Basal 1/4 of hind tibiae yellowish-white, tarsi dark brown, apices of tarsal

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

segments pale. Gaster black, narrow hind margin of first tergite, apical 1/3 of

tergite 2 and all of tergite 3 red. Veins of wings brown, base of stigma with whitish

spot.

Median part of clypeal margin straight, slightly protruding. Disk of clypeus dull,

very densely punctate. Frons and vertex densely punctate; frons convex and

somewhat shining, vertex behind ocelli finely transversely striate. POD nearly

twice OOD. Intercarinal space narrow. Tempora finely striate. Antennal segments

about 1% times as long as broad, last segment twice as long as broad, segments

3— 12 with fine linear tyloides, less distinct on segment 3, on most segments as long

as segment. .

Lateral pronotal angles obtuse. Scutum very shining, coarsely punctate, partly

rugoso-punctate. Scutellum shining, posteriorly densely punctate. Propodeal

enclosure entirely irregularly reticulate. Back of propodeum finely rugulose,

dorso-laterally indistinctly finely striate. Mesopleura shining, densely strongly

punctate, partly striato-punctate. Hypo-epimeral area somewhat dull, same punc-

tation. Anterior plate of mesepisternum dull, densely finely punctate. Anterior

oblique suture narrow, not distinctly crenulate. Petiole about 3/4 length of first

tergite, this tergite about 1.5 times as long as broad. Gaster slender. Second

recurrent vein of fore wings ending well in third submarginal cell.

Pubescence silvery-white, on face and pronotal collar mostly appressed, last

segment somewhat brownish.

Female unknown.

Texas: 1 ¢, holotype, Big Bend, 5000 ft, 1 Aug. 1975, S. and J. Peck (HT).

This species belongs to the group of argentifrons, is very close to P. castaneipes,

but differs in the tyloidea of the antennae, the sculpture of the propodeum and in

the colour of the gaster.

The specific name refers to the fine rugosity of the back of the propodeum.

Pluto colonensis sp. nov.

(fig. 28)

Female. — Very similar to P. argentifrons (Cresson). Underside of flagellum, and

fore tibiae and tarsi yellowish-red. Fore basitarsus white. Mid tibiae largely brown,

basitarsus whitish. Hind tibiae and tarsi dark brown, base of tibiae whitish.

Pronotal tubercles whitish. Hind margin of first tergite, all of second tergite and

base of third tergite reddish.

Clypeal margin (fig. 28) weakly emarginate, no median projection as in P.

argentifrons. POD slightly larger than OOD. Scutum shining, punctation coarser

than in P. argentifrons and more dense, especially on central part and on fore and

hind margins. Mesopleura dull, very finely and densely punctate, no distinct

longitudinal striae as in P. argentifrons. Back of propodeum dull, lower part finely

reticulato-carinate with large meshes. Petiole about half length of first tergite, this

tergite somewhat longer than broad. Pygidial area about twice as long as broad.

Pubescence of face silvery, of pygidial area brown, of rest of body whitish.

VAN LITH: New World Pluto 199

Male unknown.

Argentina: | 9, holotype, prov. Entre Rios, Dt. Colon, Zelich (MF).

Pluto fritzi sp. nov.

Male. — Length about 6—7 mm. Black. Mandibles dark reddish except for

basal 1/3. Underside of flagellum orange-brown. Foreside of fore femora, apex of

foreside of mid femora, fore and mid tibiae and tarsi orange-brown, basitarsi

somewhat paler. Hind tibiae and tarsi orange-brown, underside of tibiae, bases of

hind tarsal segments 1—3 and last segment of tarsi brown. Tegulae reddish-brown.

Wings somewhat smoky, veins black, fine pubescence of wings dark brown. Hind

margins of gastral tergites broadly reddish transparent.

Median part of clypeal margin straight, somewhat protruding, disk densely

punctate. Frons except for lateral margins densely rugoso-punctate. Vertex

densely punctate, interstices shining, back of head dull, transversely striato-

punctate. POD larger than OOD. Intercarinal space narrow. Antennal segments

longer than broad, last segment over 1.5 times as long as broad, segments 4—12

with linear tyloides as long as segment, tyloides on segment 13 short.

Lateral pronotal angles about rectangular. Scutum shining, strongly punctate,

partly transversely or obliquely rugulose. Scutellum shining, sparsely punctate,

hind margin densely punctate. Propodeal enclosure shining, large median area,

oblique carinae lacking or sparse. Back of propodeum very coarsely reticulato-

carinate. Mesopleura shining, densely rugoso-punctate. Hypo-epimeral area

shining, finely punctate, interstices about size of punctures. Anterior plate of

mesepisternum vertically striate, oblique suture broad, crenulate. Mesosternum

densely finely punctate. Second recurrent vein of fore wings interstitial. Petiole

somewhat longer than first tergite, this tergite longer than broad.

Pubescence of face yellowish-silvery, not distinctly appressed; pubescence

silvery, mostly appressed on tempora and pronotal collar, yellowish-grey on

scutum, scutellum and metanotum.

Female. — A separately collected female is very similar and provisionally

considered the female of P. fritzi. Length about 7.5 mm. Fore femora dark, base of

hind tibiae dark, hind tarsi brown, apices of segments paler.

Anterior margin of clypeus slightly rounded, disk densely punctate. Frons

densely punctate, lateral raised parts shining, vertex sparsely punctate. POD

slightly less than OOD. Intercarinal space about half width fore basitarsus.

Antennal segments 10—11 about as long as broad.

Scutum very shining, punctures sharp, interstices often larger than punctures.

Propodeal enclosure with a few oblique carinae, large median area; back of

propodeum coarsely reticulato-carinate. Mesopleura somewhat irregularly

densely punctate, hind margin with short rugae. Mesosternum densely punctate.

Hypo-epimeral area as in male. Petiole about as long as first tergite. Pygidial area

about twice as long as broad. Second recurrent vein of fore wings ending in third

submarginal cell.

Face golden pubescent.

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Ecuador: 1 &, holotype, Chone, May 1976, 1 &, paratype, Quevedo, May 1976,

M. Fritz (MF); 1 9, allotype, Guayaquil, 4 March 1964, R. O. Schuster (UCD); 1

3, paratype, Pto. Viejo-Quevedo, 400 m, 13 April 1965, L. Peña (MCZ). This latter

male has the nace pale golden.

P. fritzi seems to belong to the group of P. argentifrons which is characterized by

the densely punctate mesopleura. The petiole of the male is longer than that of any

other species of the group, the mesopleura and especially the hypo-epimeral area

are shining between the punctures.

This species has been named in honour of the Argentine hymenopterist Mr.

Manfredo A. Fritz.

Pluto medius medius (F. Smith)

(fig. 33)

Smith, 1856: 435, & (Psen medius; Brazil).

Dalla Torre, 1897: 348.

Bohart & Menke, 1976: 171 (Pluto medius, new combination by Bohart).

First description of female. — Length about 6.5 mm. Black. Underside of

flagellum orange-brown, labrum and mandibles reddish-brown. Pronotal tubercles

yellowish-white, tegulae yellowish-red. Hind margin of first tergite, all of second

tergite and sternite, sometimes also third tergite and sternite more or less red,

following segments with broad reddish transparent hind margin, last segment dark

reddish. Fore and mid tibiae and tarsi yellowish-red, basitarsi whitish. Basal 1/3 of

hind tibiae and basitarsi except base yellowish-white, segments 2—5 brown, with

paler apex. Veins of wings dark brown.

Median part of clypeal margin shining, less than half total width, medially very

slightly rounded and with small but distinct lateral teeth (fig. 33). Frons densely

punctate, upper part raised, a large, narrow, shining depression between posterior

ocelli and oculi. Vertex shining, sparsely punctate. Intercarinal space very narrow.

Antennae slightly clavate, not long.

Lateral pronotal angles rectangular. Scutum shining, interstices a few times size

of punctures, anterior margin dull. Scutellum shining, sparsely punctate. Pro-

podeal enclosure shining, with large pentagonal area, back dull, coarsely

reticulato-carinate, dorso-lateral carinae parallel. Mesopleura finely reticulate

alutaceous, upper part densely finely punctate, interstices on lower part as large as

or slightly larger than punctures, posterior margin finely coriaceous. Mesosternum

densely very finely punctate. Hypo-epimeral area somewhat shining, very finely

punctate. Anterior plate of mesepisternum finely striate. Second recurrent vein of

fore wings ending well in third submarginal cell. Petiole about as long as first

tergite, this tergite somewhat longer than broad. Pygidial area about twice as long

as broad.

Pubescence of face golden, mostly appressed. Vertex, scutum and pygidial area

brownish pubescent, mesonotum, mesosternum and gaster greyish pubescent,

propodeum whitish.

Male. — Hind basitarsi brown of largely yellowish. Sculpture of scutum coarser,

VAN LITH: New World Pluto 201

some short transverse rugae. Scutum laterally and anteriorly somewhat trans-

versely rugose, anterior margin moreover densely punctate, dull. Scutellum

posteriorly more rugose. Mesopleura dull or shining, very densely finely punctate,

partly striato-punctate. Petiole slightly longer than first tergite. Antennal segments

4—11 with linear tyloides, shorter than segment, on segment 11 much shorter.

Face silvery pubescent.

Fig. 32. Pluto scytinus sp. n., ©, holotype, head in frontal aspect. Fig. 33. P. medius medius (Smith), ©,

Brazil, clypeus. Fig. 34. P. medius zuliensis subsp. n., 9, holotype, clypeus. Figs. 35—36. P. clavicornis,

Mexico; 35, face of 9 in antero-dorsal aspect; 36, antenna of 3, dorsal aspect. Fig. 37. P. townsendi

(Cockerell), 9, Peru, clypeus. Fig. 38. P. metaensis sp. n., 9, clypeus.

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Material examined: Brazil: 1 g, Santarém, “medius Sm. Type”, BM type Hym.

21.815; 1 g, Santarém, “Psen medius Sm.” (BM).

New records from Brazil: Para, 1 9 and I Z, 25 July 1901, Ducke, No. 274 and

No. 206 (NMW); 1 &, Pará, Baker (UCD); Amazonas, | 9 and | g, Vista Alegre,

Rio Branco, 6 Sept. 1924 (CU); Mato Grosso, | 9 and 1 ¢, Diamantino, 10—14

Nov. 1965, S. Laroga (UFP); Roraima, 1 g, Surumu, Sept. 1966, M. Alvarenga

(HT).

First records from Argentina: Mendoza: | 9, Est Pedregal, 17 Nov. 1906,

Jensen-Haarup (ZMC); I &, 2 Dec. 1906, Jensen-Haarup (ZMB). Tucuman: 1 &,

Febr. 1947, J. Cordoba (IML). Salta: 1 Z, Tartagal, 11 Febr. 1950, R. Golbach

(IML). Yuto: 2 Z, 11 Jan. 1966, H. and M. Townes (HT). Catamarca: 3 g, Belen,

25 Nov. 1975, R. M. Bohart (UCD).

First records from Bolivia: Tarija, 1 © and 1 Z, Ing. Bermejo, 14—28 Febr.

1969, R. Golbach (IML).

First record from Surinam: | Z, Paramaribo, Agricult. Experiment Station, 21

Nov. 1963, G. van Vreden (ML).

In the males from Argentina the scutum is not rugose and the mesopleura are

shining and finer punctate; the females from Argentina and Bolivia have a silvery

pubescent face, the hind tarsi are entirely brown and the lower part of the

mesopleura as well as the mesosternum are shining, not distinctly alutaceous.

Perhaps these specimens represent a distinct subspecies.

Pluto medius zuliensis subsp. nov.

(fig. 34)

Female. — Resembling nominate subspecies. Length 6.5—8.5 mm. First tergite

except for two large black marks, all of tergites 2—3, sides of tergite 4, and

sternites 2—4 red. Hind tarsi dark brown, apices of segments yellowish-brown.

Median part of clypeal margin more sharply protruding than in nominate

subspecies (fig. 34). Scutum shining, with large impunctate areas. Petiole about 3/4

length of first tergite.

Face deep golden pubescent, pygidial area brown, rest of body whitish pub-

escent.

Male. — Similar to female. Length about 6.5 mm. First tergite except for two

large black marks, all of tergites 2—3, second sternite and base of third sternite

red. Hind basitarsus somewhat paler brown. Scutum with smaller impunctate

areas, no tendency to rugosity.

Face pale golden pubescent.

Venezuela: Zulia, Carrasquero, 1 ©, holotype and 1 Z, allotype, 15 June 1976,

38 9, paratypes, 29—30 May and 15 June 1976, A. S. Menke and D. Vincent

(holotype and allotype USNM, paratypes UZM and USNM).

Curacao : | @, paratype, Gr. Santa Martha, 24 Nov. 1963, on Sesuvium

portulacastrum, B. de Jong (ML).

VAN LITH: New World Pluto 203

This subspecies is on the whole somewhat larger than the nominate subspecies,

the gaster shows more red, the hind tarsi are dark brown and the face of the female

is deep golden pubescent, the face of the nominate subspecies being pale golden.

The clypeal margin is slightly different.

The subspecific name refers to the state of Zulia in Venezuela, where a large

series of females has been collected.

Pluto scytinus sp. nov.

(fig. 32)

Female. — Length about 5 mm. Head and thorax black; mandibles yellowish-

red, labrum reddish, underside of flagellum yellowish-red. Fore and mid tibiae and

tarsi, basal 2/5 of hind tibiae and hind tarsal segments 1—4 yellowish. Outer side of

fore and mid tibiae with ivory-white streak. Pronotal tubercles ivory-white.

Tegulae yellowish-red. Hind margin of first gastral tergite, all of tergites 2—3,

sides of fourth tergite and sternites 2—3 reddish. Apex of last segment dark

reddish.

Median part of clypeal margin slightly emarginate, indistinct median tooth,

lateral teeth small but distinct, distance less than half total width of margin (fig.

32). Frons very closely finely punctate, laterally with shining area. Vertex finely

alutaceous, finely punctate, interstices about twice size of punctures. POD nearly

twice OOD. Tempora finely striato-punctate. Intercarinal space about as wide as

first basitarsus. Antennae short, clavate, segments 10—11 shorter than broad.

Lateral pronotal angles about rectangular. Scutum shining, densely punctate,

interstices larger than punctures. Scutellum shining, sparsely punctate. Propodeal

enclosure almost dull, irregularly carinate. Back of propodeum dull, closely

reticulate, dorso-lateral parts with oblique parallel carinae. Mesopleura and hypo-

epimeral area closely very finely punctate, almost coriaceous, punctation of

mesosternum with distinct interstices. Anterior plate of mesepisternum dull,

oblique suture narrow. Petiole about half length first tergite. Pygidial area about

1.5 times as long as broad. Second recurrent vein of fore wings ending in third

submarginal cell.

Face and pronotal collar silvery, mostly appressed, pubescent. Pubescence of

scutum brownish, of pygidial area brown, of rest of body silvery-white.

Male unknown.

Venezuela: | 9, holotype, Zulia, Rosario, 4 June 1976, A. S. Menke and D.

Vincent (USNM).

P. scytinus belongs to the group of P. argentifrons. It differs from the South

American P. stramineipes in having reddish gastral tergites, from the Argentine

P. colonensis in having yellowish hind tarsi, from both species also in having finer

punctate mesopleura. |

The specific name refers to the leathery appearance of the mesopleura.

204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pluto stramineipes sp. nov.

(fig. 30)

Female. — Length about 6 mm. Black. Mandibles and labrum reddish.

Underside of flagellum orange-brown. Pronotal tubercles whitish. Fore and mid

tibiae and tarsi brownish-yellow, outer side of fore tibiae with narrow yellow line,

basitarsi yellowish-white. Basal 1/3 of hind tibiae and tarsal segments 1—4

yellowish-white. Hind margins of tergites somewhat reddish transparent. Pygidial

area more or less reddish. Veins of wings dark brown.

Protruding median part of clypeus almost straight, somewhat less than half total

width of margin, medially slightly protruding, lateral teeth small (fig. 30). Disk of:

clypeus dull, narrow shining anterior margin. Frons densely punctate, except for

shining low oval tubercle along eyes. Vertex finely transversely striate, sparsely

punctate, interocellar area shining. POD about equal to OOD. Intercarinal space

nearly as broad as first basitarsus. Antennae clavate, segments 10—11 much

shorter than broad. ‘

Lateral pronotal angles obtuse. Scutum and scutellum shining, finely punctate,

interstices mostly a few times size of punctures, punctation close on fore and hind

margin. Propodeal enclosure shining, back of propodeum dull, coarsely reticulato-

carinate, with fine oblique dorso-lateral striae. Mesopleura densely: finely

punctate, on median part striato-punctate, near lower margin interstices about as

large as punctures, hind margin finely rugulose. Mesosternum densely finely

punctate. Second recurrent vein of fore wings ending in third submarginal cell.

Petiole about 3/5 length of first tergite, this tergite slightly longer than broad.

Face and pronotal collar silvery, mostly appressed, pubescent. Pygidial area

golden-brown pubescent, thorax dorsally greyish, rest of body whitish.

Male. — Similar to female. Hind tarsal segments 2—4 more or less brown.

Antennae long and slender, segments about 1.5 times as long as broad, segments

5—10 with linear tyloidea, as long as segments. Vertex and scutum more coarsly

punctate. Propodeal enclosure with irregular, shining, large median area. Sculp-

ture of back of propodeum coarser than in female. Petiole subequal to first tergite,

this tergite about 1.5 times as long as broad.

Brazil: Santa Catarina: Nova Teutonia, 27°11’S., 52°23’W., 300—500 m, 1 ©,

holotype, 28 Jan. 1956, Fritz Plaumann (CU); 1 &, allotype, 14 Nov. 1964;

paratypes: 1 &, 17 Jan. 1964, 8 &, Oct.—Nov. 1968, 1 g, Nov. 1969, all collected

by Fritz Plaumann (UCD). Further paratypes from this locality: 2 9, 8 Nov. 1962

and | Febr. 1963, 5 8, Oct.-Nov. 1967, Fritz Plaumann (MCZ); 1 9, Nov. 1974,

Fritz Plaumann (BM). Mato Grosso: | 9, paratype, Aquidana, 11—13 Dec. 1919,

R. G. Harris, Cornell Univ. Exped. (CU). Parana: | 9, paratype, Falls Iguazu, Jan.

1962, Sakagami-Laroca (UFP).

Argentina: Misiones: | 9, Posadas, 15—24 Jan. 1920, Cornell Univ. Exped., Lot

569 (CU). Entre Rios: 2 ¢, Pronunciamento, 13—16 Febr. and 4—10 March 1965

(FAG); 1 g, Feliciano, Dec. 1972, M. A. Fritz (MF). All paratypes.

Bolivia: 3 9, Luis Calvo, Tiguipa, Jan. 1972, M. A. Fritz (MF). Paratypes.

The male of P. stramineipes differs from the other species of the group of

VAN LITH: New World Pluto 205

argentifrons in having tyloidea on segments 5—10 only and in having pale hind

tarsi.

The specific name refers to the straw-coloured hind tarsi, especially those of the

male.

Pluto strigellus sp. nov.

(fig.31)

Female. — Length about 5.5 mm. Black. Mandibles and labrum largely reddish.

Underside of flagellum reddish. Pronotal tubercles whitish, tegulae yellowish.

Fore tibiae and tarsi yellowish-red, base of tibiae and tarsal segments 1—3

yellowish-white. Mid tibiae brown, base and outer side of apex yellowish, tarsal

segments 1—3 yellowish-white. Hind legs brown, base of tibiae yellowish, apices of

tarsal segments yellowish-brown. Apices of first tergite, all of second tergite and

base of third tergite reddish. Veins of wings dark brown.

Protruding median part of clypeal margin slightly bisinuate, with distinct small

lateral teeth (fig. 31). Disk of clypeus densely finely punctate. Frons densely finely

punctate, vertex shining, sparsely punctate. POD slightly larger than OOD.

Intercarinal space narrow. Antennae slightly clavate, segments 10—11 about

quadrate, segment 12 about 1.5 times as long as broad.

Lateral pronotal angles obtuse. Scutum shining, distinctly, sparsely, punctate.

Scutellum shining, sparsely punctate, posteriorly some short rugae. Propodeal

enclosure shining, irregularly carinate. Back of propodeum coriaceous, dorso-

lateral parts obliquely striate, lower part irregularly reticulato-carinate. Meso-

pleura dull but lower part somewhat shining, densely finely punctate, interstices

about size of punctures, upper half finely, almost indistinctly, longitudinally

striato-punctate. Mesosternum shining, densely finely punctate. Hypo-epimeral

area dull, densely very finely punctate. Anterior plate of mesepisternum dull,

sculpture indistinct, oblique suture crenulate. Petiole about 2/3 of lenght of first.

tergite, this tergite about as long as broad. Pygidial area almost twice as long as

broad. Second recurrent vein of fore wings interstitial.

Face silvery, mostly appressed, pubescent. Vertex and dorsum of thorax

brownish pubescent, pygidial area brown, rest of body whitish pubescent, densely

soon mesopleura.

Male unknown.

Argentina: Santa Fe, Alberdi (near Rosario), strand, 890, | 9, holotype, 24 Nov.

1911; Alberdi, 1 ©, paratype, 4 Febr. 1917, J. Hubrich (BSM).

The sculpture of the thorax and the shape of the clypeal margin closely resemble

those of P. alphitopus; the clypeal margin point to close relationships with

P. medius as well, but in the latter species the front is raised before the posterior

ocelli, while this is hardly so in P. strigellus.

The specific name refers to the fine striae on the mesopleura.

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Group of clavicornis

Pluto clavicornis (Malloch)

(figs. 35— 36)

Malloch, 1933: 50—51, & (Psenia clavicornis; Arizona).

Krombein, 1951: 962 (Pluto clavicornis, comb. nov.).

Bohart & Menke, 1976: 171.

First description of female. — Length about 6.5 mm. Black. Mandibles, labrum

and underside of flagellum yellowish-red. Pronotal tubercles yellowish-white.

Tegulae reddish-brown. Fore and mid tibiae and tarsi largely yellowish-brown,

basitarsi whitish. Basal third of hind tibiae yellowish-white, tarsi yellowish-white or

largely brownish. Gaster black, hind margins of tergites and pygidial area reddish-

brown. Veins of wings dark brown.

Clypeal margin strongly snoutlike protruding, apex bidentate, laterally with

small triangular tooth (fig. 35); protruding part in ventral view flattened, finely

reticulate alutaceous. Disk of clypeus somewhat shining, indistinctly punctate.

Frons shining, densely superficially punctate, raised near anterior ocellus, a

longitudinal narrow depression between posterior ocelli; a shining raised area near

oculi. Vertex shining, sparsely finely punctate. POD somewhat smaller than OOD.

Ventral part of occipital carina high, intercarinal space about as wide as first

basitarsus. Antennae clavate, short, segments 11—12 shorter than broad.

Lateral pronotal angles sharp. Scutum and scutellum somewhat shining, very

finely reticulate alutaceous, sparsely finely punctate. Propodeal enclosure shining,

large median area with a median carina. Back of propodeum dull, coarsely

reticulato-carinate. Mesopleura, hypo-epimeral area and mesosternum somewhat

shining, very finely reticulate alutaceous, mesopleura sparsely finely punctate.

Anterior plate of mesepisternum dull, oblique suture broad, crenulate. Second

recurrent vein of fore wings interstitial or ending in third submarginal cell. Petiole

as long as first tergite, this tergite 1.5 times as long as broad. Pygidial area about

1.5 times as long as broad.

Face silverly to pale golden, mostly appressed, pubescent, pubescence on

thorax greyish, on pygidial area reddish-brown, on rest of body whitish.

Male. — Length about 6—6.5 mm. Very similar to female apart from the sexual

dimorphism. Clypeal margin almost straight, with two very small teeth at about 1/3

of total width. Antennae (fig. 36) short and clavate, no tyloidea. Face silvery.

Thus far recorded from Arizona only.

First records from Mexico: Guerrero: 4 &, Xalitla, 1500 ft, 20 March 1959, H. E.

Evans and D. M. Anderson (CU): 1 &, 9 mi south Tierra Colorado, 21 July 1963, F.

D. Parker and L. A. Stange (UCD). Morelos: 1 &, 3 mi north Alpuyeca, 3400 ft, 9

March 1959, H. E. Evans and D. M. Anderson; 6 3, same locality, 23 March 1959,

H. E. Evans (CU). Oaxaca: 1 &, Crawford (UCD). Sinaloa: 113 &, 8 mi southeast

Elota, 18—19 May 1962, F. D. Parker and L. A. Stange (UCD); 1 9 and | g, same

locality, 18 May 1962, F. D. Parker and L. A. Stange, with label “Pluto clavicornis

det. Stange” (IML); 2 9, Mazatlan, 10 ft, 20 July 1959, H. E. Evans (CU); 2 9,

VAN LITH: New World Pluto 207

Mazatlan, 6 Aug. 1964, W. R. M. Mason (CNC); 2 &, 11 mi north Culiacán, 20

May 1962, F. D. Parker (UCD); 1 © and 4 &, 21 mi east Villa Union, 1 Febr. 1964,

E. I. Schlinger (UCD, 1 g CAS). Sonora: 1 g, Alamos, 13 June 1961, F. D. Parker

(UCD). Veracruz: 7 3, July 1965, N. L. H. Kraus (USNM).

P. clavicornis is easily recognized by the conspicuous snout of the female, the

male by the clavate antennae.

Group of townsendi

Pluto townsendi (Cockerell)

(fig. 37)

Cockerell, 1911: 272, © and & (Psenulus (Neofoxia) Townsendi; Peru).

Bohart & Menke, 1976: 171 (Pluto townsendi, new combination by Bohart).

Female.—Length about 5.5 — 6 mm. Black. Mandibles and labrum reddish,

underside of flagellum yellowish-red. Pronotal tubercles and part of tegulae

yellowish-white. Apices of all femora, fore and mid tarsi and greater part of fore

and mid tibiae yellowish-red, fore basitarsi whitish, outer side of mid tibiae

whitish. Basal 1/3 of hind tibiae yellowish, hind tarsi largely brown. Narrow hind

margin of first tergite, all of second tergite, third tergite except for a black apical

triangle (rarely its base only), and sternites 2—3 red. Last gastral segment dark

reddish. Veins of wings brown.

Median part of clypeal margin very weakly emarginate, with short lateral teeth,

distance between these teeth about half total width of margin (fig. 37). Disk of

clypeus densely punctate. Frons somewhat shining, very finely punctate. Vertex

shining, punctation indistinct. POD about equal to OOD. Intercarinal space

narrow. Antennae clavate, segments 10—11 about quadrate, last segment about

1.5 times as long as broad.

Lateral pronotal angles obtuse. Scutum but for anterior margin shining, also

scutellum, sparsely punctate. Metanotum coriaceous. Back of propodeum and

enclosure dull. Enclosure with a few fine oblique carinae, back finely reticulato-

carinate with large meshes, dorso-lateral parts almost smooth with some fine

striae. Mesopleura and mesosternum somewhat dull, dorsal half of mesopleura

with very indistinct punctures, hind margin shining, very finely, almost indistinctly,

coriaceous; mesosternum densely, more distinctly punctate. Hypo-epimeral area

shining, no distinct punctures. Anterior plate of mesepisternum dull, finely

sculptured. Anterior oblique suture narrow, crenulate. Second recurrent vein of

fore wings interstitial. Petiole about 3/4 length of first tergite, this tergite about

1.25 times as long as broad. Pygidial area triangular, over 1.5 times as long as

broad.

Pubescence of face silvery, mostly appressed. Pubescence of vertex and scutum

somewhat greyish or brownish, of pygidial area golden-brown, of rest of body

whitish, rather dense on tempora, pronotal collar, metanotum and mesopleura.

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Male. — Similar to female, more slender, gaster darker. Length about 5 mm.

Gaster black or dark brown, tergites 2—3 with narrow red hind margin, some

specimens with apical half of second tergite and nearly all of third tergite red.

Basal third of hind tibiae whitish, all tarsi yellowish-white, last segment of hind

tarsi and base of hind basitarsus more or less darkened.

Clypeal margin slightly roundly protruding. Antennae about as long as head and

thorax together, segments 3—12 about twice as long as broad, segment 13 over

twice as long as broad. Segments 4—9 with long and fine linear tyloidea,

sometimes indistinct on segment 4, shorter on segment 9, often also segment 10

with a small tyloides. Scutum densely finely punctate. First gastral tergite about

1.5 times as long as broad. .

Material examined: | 4, Piura Apl. 28 (Twns), “Psenulus (Neofoxia) Townsendi

Ckil. Type 3” (USNM, type No. 14096). Piura is a town on the river Piura in Peru.

New records from Peru: 2 © and 20 g, Piura, Ostendorf S. G. (ZMB).

First records from Ecuador: 2 9, Guayas Prov., Guayaquil, 3—5 March 1964, P.

D. Ashlock (BISH); 8 9 and 2 3, Manabi Prov., Chone, May 1976 (MF); I &, 35

km southeast Bahia de Caräquez, 10 May 1975, Ashley B. Gurney (USNM). One

Q and one 4, labelled ‘“Puna, Kinb.” (RS) have been collected by Dr. J. G. M.

Kinberg, who joined the voyage around the world by the Royal Swedish frigate

Eugenie in 1851—1853 as a zoologist. According to Persson (1971) insects with

label “Puna” have been collected on the isle of Puna, Guayas Prov. on 27 March

1852 but he thinks that it is also possible that they were collected in or around

Guayaquil during the period 28 March—4 April 1852. Dr. S. Erlandsson,

Stockholm, kindly called my attention to this publication.

The group of townsendi is easily recognized by the weakly shining, almost

imperceptibly sculptured mesopleura of the female and the long antennae of the

male. P. townsendi is distinguished from the other two taxa belonging to this group

by its shorter petiole.

Pluto marthae sp. nov.

Female. — Lenght about 6 mm. Head and thorax black; mandibles yellowish-

red, labrum dark reddish. Underside of flagellum orange-brown. Pronotal

tubercles yellowish-white. Tegulae yellowish-red. Fore and mid tibiae and apices

of femora yellowish-brown, basitarsi whitish, mid part of tibiae and last tarsal

segment brown. Basal third of hind tibiae yellowish-white, hind tarsi brown with

yellowish-brown apices. Petiole and basal half of first tergite black, remainder of

gaster red, bases of some segments may be slightly darkened. Veins of wings dark

brown.

Median part of clypeal margin less than half total width, straight or in the middle

very slightly convex, shining, with two small lateral teeth; disk densely finely

punctate. Frons not distinctly punctate, central part dull. Vertex impunctate,

shining between ocelli and oculi, dull behind ocelli. Intercarinal space narrower

than first basitarsus. Antennae somewhat thickening towards apex, segments

9—11 about quadrate, last segment slightly over 1.5 times as long as broad.

VAN LITH: New World Pluto 209

Lateral pronotal angles nearly rectangular. Scutum shining, sparsely finely

punctate, anteriorly and near hind margin somewhat dull, owing to minute

hairbearing punctures. Scutellum shining, almost impunctate. Metanotum dull,

coriaceous. Propodeal enclosure triangular, with large pentagonal median area,

shining. Back of propodeum dull, with moderately coarse, reticulate carination,

carinal dorsally more parallel. Mesopleura, hypo-epimeral area and mesosternum

somewhat dull owing to indistinct reticulate alutaceous microsculpture, no distinct

punctation. Metapleura shining. Petiole about 1.25 times length of first tergite, this

tergite somewhat longer than broad. Pygidial area over 1.5 times as long as broad.

Second recurrent vein of fore wings interstitial.

Pubescence of head and thorax silvery, on face mostly appressed, dense on

mesopleura. Pubescence of gaster short, yellowish; pygidial area with golden-

brown short and appressed pubescence and a few long erect hairs.

Male. — Similar to female. Length 4—5 mm. Apical half of foreside of fore and

mid femora also yellowish-brown. Apical margin of tergite 1, all of tergites 2 and 3

red with an indistinct brownish band before hind margin, hind margins of

following tergites reddish transparant; sternites 2—4 red.

Antennae long and slender, segments 8—12 about twice as long as broad,

segment 13 over twice as long as broad. Segments 4— 10 or 4—11 with indistinct,

dull, linear tyloidea. Punctures of scutum stronger and more densely placed, partly

weakly rugoso-punctate. Back of propodeum more coarsely and more irregularly

reticulate than in female. Petiole about 1% times length of first tergite. In one of

the males the pubescence of the face is faintly yellowish.

Peru: 1 9, holotype, Loreto, Pucallpa, Lake Yarina Cocha, 180 m, 26—27 Aug.

1971, 3 3, allotype and paratypes, same locality, 25—27 Aug. 1971, C. and M.

Vardy (BM). Further paratypes: 2 Z, “Peru” (NMW); 2 9, Loreto, San Antonio,

13 Aug. 1965, Malaise trap, J. C. Hitchcock Jr. (USNM); | &, Puerto Bermudez,

Rio Pichis, 12—19 July 1920, Cornell U. Lot 569, Sub 256 (CU); 1 9, 25 mi west

San Jorge, 4 Oct. 1954; 1 4, Yurac, 67 mi east Tingo Maria, 350 m, 4 Oct. 1954, E.

I. Schlinger and E. S. Ross (CAS); 1 9, 15 mi northeast Tingo Maria, 700 m, 11

Nov. 1954, E. I. Schlinger and E. S. Ross (UCD).

Bolivia: | 9, paratype, near mouth Rio Mapiri, Sept., Mulford Bio Expl.

1921—22 (USNM).

Brazil: 1 9, “Brasil, Smith coll. pres. by Mrs. Farren White 99—303”(BM); 1 9,

Acre, Cruziero do Sul, Nov. 1963, M. Alvarenga (UFP). Paratypes.

Colombia: I 3, paratype, Caqueta, Rio Orteguaza nr. Rio Peneya, 14—18 Jan.

1969, Duckworth and Dietz (USNM).

Ecuador: Napo, 1 &, paratype, Coca on Rio Napo, May 1965, L. E. Pena

(AMNH); I 3, paratype, 42 km west Santa Cecilia, 16 May 1975, Ashley Gurney

(USNM); 1 &, Coca, Napo River, 250 m, 22—30 April 1965, L. Pena (CNC); the

| scutum of this latter male is much less coarsely punctate than in the other males,

| with distinct interstices between the punctures.

P. marthae is closely related to P. townsendi, which is also found in Peru, and

which has the same dull, indistinct sculpture of the mesopleura. It differs from the

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

latter species in the longer petiole and the coarser reticulation of the back of the

propodeum. The gaster of the female is more red.

P. marthae is named for Mrs. Martha Vardy, one of the collectors of the type

material.

Pluto metanus sp. nov.

(fig. 38)

Female. — Very close to P. marthae but differing in the following details. Length

about 5 mm. Bases of fore and mid tibiae and mid tarsal segments 1—4 yellowish-

white. Gaster black, hind margins of tergites and of sternites reddish.

Median part of clypeal margin less than half total width of clypeus, slightly but

distinctly emarginate, medially not protruding, with small lateral teeth (fig. 38).

Male. — Similar to female. Fore tibiae and tarsi more orange. Antennal

segments 4—11 with distinct, shining, fine linear tyloidea. Vertex behind ocelli

shining. Scutum with less tendency to rugosity than in the male of P. marthae.

Colombia: 3 ©, holotype and paratypes, 2 g, allotype and paratype, Meta, Rio

Duda, 8—12 March 1976, M. Cooper (BM).

This form may have to be ranked as a darker subspecies of P. marthae, although

the clypeal margins of the females seem to be slightly different. Further studies on

larger series are needed to assess whether these differences fall within the range of

specific variation.

Group of nitens

Pluto nitens sp. nov.

(fig. 39)

Female. — Length about 7 mm. Black. Mandibles pale yellowish-brown with

dark tips, labrum dark reddish. Underside of flagellum brown or reddish-brown.

Pronotal tubercles brown. Tegulae yellowish-brown. Knees, tibiae and tarsi of fore

legs yellowish, underside of tibiae slightly darkened. Apices of mid tibiae and

tarsal segments 1—4 yellowish. Basal 1 /3 of hind tibiae reddish-brown, tarsi brown

but greater part of basitarsi and apices of segments 2—5 brownish-yellow. Veins of

wings dark brown. Hind margins of gastral segments reddish transparent.

Median part of clypeal margin depressed, shining, very weakly emarginate with

indistinct lateral angles (fig. 39); disk convex, distinctly separated from depressed

margin, upper part of disk densely punctate, lower 1/3 shining, impunctate. Frons

medially with fine hair-bearing punctures, upper part and vertex shining, sparsely

punctate. POD smaller than OOD. Occipital carina fine, intercarinal space

narrow. Antennae relatively slender, gradually widening towards apex, segments

10—11 in frontal view longer than wide, segment 12 twice as long as broad.

Lateral pronotal angles almost rectangular. Scutum and scutellum shining,

VAN LITH: New World Pluto 211

almost imperceptibly reticulate alutaceous, sparsely punctate. Prescutal sutures

distinct on anterior 1/3. Propodeal enclosure shining, with long oblique carinae

and elongate pentagonal median area. Back of propodeum dull, with long parallel

carinae, lower part reticulato-carinate. Mesopleura and hypo-epimeral area

smooth and shining, sparsely minutely punctate, mesosternum more densely so.

Anterior oblique suture narrow, crenulate. Second recurrent vein of fore wings

ending in third submarginal cell, almost interstitial. Petiole about as longer as first

tergite, this tergite as long as broad. Pygidial area about 1.75 times as long as

broad.

Face with silvery appressed pubescence and many long, erect, silvery hairs.

Pronotal collar with some appressed silvery pubescence and many long erect hairs.

Vertex, scutum and scutellum greyish-brown or brown pubescent, pygidial area

and sides of last tergite golden-brown appressed pubescent and also with long

erect brown hairs, rest of body whitish pubescent.

Male. — Similar. Length 5-6 mm. Median part of clypeal margin slightly

protruding, margin narrowly depressed, shining. Antennae long and slender,

segments 3—12 about 1.5 times as long as broad, segment 13 over twice as long as

broad, segments 4—12 with fine linear tyloides, as long as segments, shorter on

segments |l and 12 or on segment 12, segment 13 sometimes with a short tyloides.

Petiole as long as or slightly longer than tergite 1, this tergite longer than broad.

Back of propodeum coarsely reticulate, upper part with some parallel oblique

carinae. Hind basitarsi largely brown, sometimes basitarsus yellowish-red. Face

silvery pubescent, sometimes golden, pubescence of head and thorax golden-

brown.

Brazil: Goiás: 1 9, holotype, 1 Z, allotype, 2 © and 1 g, paratypes, Jatai, Nov.

1972, F. M. Oliveira (HT); 1 9, paratype, same locality and date, F. M. Oliveira

(CNC). Further paratypes from Brazil: Mato Grosso: 2 9 and 2 g, Itaum, March

1974, M. Alvarenga (HT). Minas Gerais: | g, Vicoss, 1930, E. J. Hambleton (CU);

1 © and | g, Brazopolis, Dec. 1961, 4 4, Passos, 1—8 March and 24 Nov. 1962,

16—21 Oct. and 18—23 Nov. 1963, 1 9, Pratápolis, 5 Nov. 1963, 1 &, Araxa, 29

Nov. 1965, all collected by C. Elias (UCD). Sao Paulo: 1 Q, Sao Paulo, 18 March

1967, V. N. Alin (UCD); I 9, Sao Paulo, 12 Febr. 1968, V. N. Alin (USNM); I 9,

Ribeirào Préto, 1 July 1968, G. E. Bohart (UCD).

Two further females from Minas Gerais have brownish-orange fore and mid

tarsi and underside of flagellum. Their clypeus seems to be more flat: 1 9,

Perdizes, 8 April 1965, C. Elias; 1 9, Araxa, 22 March 1965, C. Elias (UFP). They

have not been labelled as paratypes.

The undermentioned specimens are all paratypes.

Argentina: Misiones: | g, Iguazu, 30 Jan.—13 March 1945, Hayward-Willink-

Golbach (IML).

Bolivia: Cochabamba: 1 g, Chapare, Cesar Zama, Jan. 1975, Martinez, 2 3,

Chapare, Chimore, Jan. 1972, M. Fritz (MF); I g, “Bolivia”(NMW).

Colombia: 1 g, Vaupés, Mitu, 11 May 1974, M. Cooper (BM). The hind tarsi of

this male are brownish-yellow. No antennae.

Ecuador: Napo Prov.: | © and I g, Tena, 17 Febr. and 24 March 1923, F. X.

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Williams (BISH); 2 ©, Coca, May 1965, Luis Pena (HT and MCZ); 3 © and 8 &,

Limoncocha, 250 m, 15—28 June 1976, S. and J. Peck (CNC); 1 &, Limoncocha, 9

June 1977, D. L. Vincent (USNM); 1 ©, Dureno Aguarico, 76° 31° west, 00° 4°

south, 29 May 1963, L. Pena (MCZ); 1 9, Pompeya, May 1965, Pena (MCD

Paraguay: | 9, Pirareta, Jan. 1972, L. Pena (MF).

Peru: Pasco Dep.: 1 9, Puerto Bermudez, Rio Pichis, 12—19 July 1920, Cornell

Univ. Exped., Lot 596, Sub. 256 (CU). Cuzco Dep.: 2 9, Quincemil, 15—30 Oct.

1962, Pena (CNC); 1 9 and 1 g, Quincemil, 750 m, 1—16 Nov. 1962 and 16—31

Oct. 1962, L. Peña (MCZ). Huánuco Dep.: 1 © and 1 &, Monson Valley, Tingo

Maria, 2 Nov. and 21 Oct. 1954, 1 g, Yurac, 67 mi east of Tingo Maria, 16 Nov.

1954, E. I. Schlinger and E. S. Ross (CAS); I 9, Tingo Maria, 20—27 Jan. 1968, A.

Garcia and C. Porter (MCZ). Madre de Dios Dep.: 1 ¢, Manu, Oct. 1962, Pena

(MCZ).

Surinam: | 9, Mapane Area, Camp 8, LBB, m.k. II, 30 May 1963, J. van der

Vecht (ML).

Venezuela: | 9, San Esteban, Falcon, 16 Jan. 1940, P. J. Anduze (CU).

The pubescence of the face of the males from Ecuador and Peru is golden,

except in the male from Manu (Peru), and the pubescence of the thorax is

somewhat golden-brown. The face of the females from these countries is very pale

golden.

The specific name refers to the shining mesopleura.

Pluto duckei sp. nov.

(fig. 40)

Female. — Length about 7.5 mm. Black. Mandibles and labrum orange-red;

underside of flagellum reddish-brown. Pronotal tubercles, tegulae, fore and mid

tibiae and tarsi yellowish-red. Basal 2/5 of hind tibiae and tarsal segments 1—4

yellowish-red. Hind margin of first tergite, all of second tergite and basal half and

hind margin of third tergite, hind margins of tergites 4—5, sternites 2—3 entirely

and hind margins of sternites 4—5 reddish. Veins of wings dark brown.

Median part of clypeal margin about half total width of margin, slightly

emarginate, no distinct lateral teeth (fig. 40). Anterior third of clypeal disk shining,

impunctate, somewhat raised, narrow margin depressed. Frons and vertex shining,

frons sparsely finely punctate. POD distinctly smaller than OOD. Occipital carina

somewhat thickened ventro-laterally, low and indistinct near hypostomal carina,

intercarinal space very narrow. Antennae gradually widening towards apex, last

segment about 1.75 times as long as broad.

Lateral pronotal angles obtuse. Scutum and scutellum shining, very finely

reticulate alutaceous, sparsely finely punctate. Propodeal enclosure shining, large

median diamond-shaped area, back of propodeum with widely placed oblique

parallel carinae, below somewhat reticulate. Mesopleura and hypo-epimeral area

shining, indistinctly alutaceous, mesopleura with very minute, hair-bearing

punctures. Mesosternum almost dull, densely very finely punctate. Anterior plate

of mesepisternum shining, oblique suture with few transverse carinae. Second

VAN LITH: New World Pluto 213

roa

45 46

Fig. 39. Pluto nitens sp. n., 9, holotype, clypeus. Fig. 40. P. duckei sp. n., 9, holotype, clypeus. Fig. 41.

P. obscurus sp. n., 9, holotype, clypeus. Fig. 42. P. menkei sp. n., 9, holotype, clypeus. Fig. 43. P. zona-

tus sp. n., 9, paratype, clypeus. Figs. 44—45. P. occipitalis sp. n., Q holotype; 44, clypeus; 45, prono-

tum. Fig. 46. P. spinicollis sp. n., 9, pronotum. Fig. 47. P. trilobatus sp. n., 9, paratype, head in frontal

aspect. Fig. 48. P. microlobatus sp. n., 9, holotype, clypeus.

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

recurrent vein of fore wings about interstitial. Petiole as long as first tergite, this

tergite as long as broad. Pygidial area about twice as long as broad.

Face, tempora and pronotal collar with dense, mostly appressed, silvery pub-

escence. Vertex, scutum and scutellum brownish pubescent. Pygidial area with

dark brown, backwards directed bristles. Rest of body whitish pubescent.

Male unknown.

Brazil: 1 ©, holotype, Pará, 25 Nov. 1899, Ducke (NMW); 1 9, paratype, Pará,

Baker (UCD).

This female differs from the female of P. nitens mainly in the colour of the gaster

and in the microsculpture of the thorax. The low ventral part of the occipital:

carina and the red second tergite are characteristic. Also close to P. zonatus which

has shorter antennal segments, different clypeal margin, less shining mesopleura

and shorter petiole.

Pluto obscurus sp. nov.

(fig. 41)

Female. — Length about 9.5 mm. Black. Mandibles dark reddish. Underside of

last antennal segment, foreside of fore tibiae and all of fore tarsi, apices of mid

tibiae and mid basitarsus reddish-brown. Hind tibiae and tarsi entirely dark brown,

hind tibial spurs reddish-brown.

Anterior margin of clypeus almost straight, no lateral teeth or angles, shining

margin very narrowly depressed (fig. 41). Disk of clypeus entirely with large

superficial punctures. Frons and vertex shining, sparsely punctate, a large low

shining tubercle near oculi, frons raised around anterior ocellus. POD somewhat

smaller than OOD. Intercarinal space narrow. Antennae slightly thickened

towards apex, third segment about 3.5 times as long as broad, following segments

about 1.5 times.

Lateral pronotal angles somewhat sharp, little protruding. Scutum and

scutellum shining, very finely reticulate alutaceous, sparsely punctate. Propodeal

enclosure shining, with few oblique carinae. Back of propodeum somewhat

shining, coarsely reticulate, dorso-lateral parts with some oblique carinae. Hypo-

epimeral area and mesopleura shining, very finely alutaceous, mesopleura sparsely

very finely punctate. Anterior plate of mesepisternum dull, with a few rugae,

oblique suture crenulate. Mesosternum densely superficially finely punctate.

Petiole about 4/5 length of first tergite, this tergite slightly longer than broad.

Pygidial area about 1.5 times as long as broad. Second recurrent vein of fore wings

ending in third submarginal cell. Posterior side of hind tibiae with longitudinal

rows of short pale brownish transparent thorns, each ending in a very short hair.

Face with silvery appressed pubescence and conspicuous long erect brown

hairs. Tempora and pronotal collar silvery, appressed, pubescent. Hind tibiae on

inner side at base and apex and inner side of hind basitarsi with conspicuous

golden pubescence. Pygidial area with dark brown pubescence. Rest of body with

greyish-brown hairs.

Male unknown.

VAN LITH: New World Pluto 215

Argentina: | 9, holotype, Misiones, Pto. Esperanza, Dec. 1976, M. Fritz (MF).

P. obscurus is apparently close to P. nitens, but is distinctly larger and darker, the

frons is distinctly raised, the thorax is very finely alutaceous, the antennal

segments are much longer and the long hairs intermixed with the appressed facial

pubescence are brown.

The specific name refers to the dark legs and the dark hairs of the clypeus.

Pluto zonatus sp. nov.

(fig. 43)

Female. — Length 7 mm. Black. Mandibles dark reddish. Dorsal side of

flagellum brown, underside orange. Pronotal tubercles dark brown. Tegulae

yellowish-red. Fore and mid tibiae and tarsi, basal 1/4 of hind tibiae and hind tarsal

segments 1—4 brownish-yellow or yellowish-red. Hind margins of first tergite

distinctly red, all or 2/3 of second tergite and sternite red. Veins of wings brown.

Median part of clypeal margin with three shallow emarginations, this part about

half total width of clypeal margin (fig. 43). Basal 2/3 of clypeal disk densely

strongly punctate, anterior 1/3 shining, almost impunctate. Frons and vertex

shining, a low elongate tubercle near the eyes; frons on either side of anterior

ocellus raised and dull, indistinctly punctate. POD nearly as large as OOD.

Intercarinal space narrow. Antennae thick, segments 10—11 broader than long,

last segment about 1.5 times as long as broad.

Lateral pronotal angles rectangular, anterior carina projecting laterally as a

minute tooth. Scutum and scutellum shining, sparsely finely punctate. Propodeum

shining, enclosure with large median pentagonal area, back coarsely reticulato-

carinate, dorso-lateral carinae close and parallel. Mesopleura and hypo-epimeral

area shining, indistinctly punctate, lower part of mesopleura slightly alutaceous.

Anterior oblique suture crenulate. Mesosternum somewhat reticulate alutaceous,

sparsely distinctly punctate. Second recurrent vein of fore wings ending in third

submarginal cell. Petiole about 2/3 length of first tergite, this tergite somewhat

longer than broad. Pygidial area about | % times as long as broad.

Face and tempora pale golden pubescent, vertex, scutum and scutellum

brownish-grey pubescent, pygidial area with appressed golden-brown pubescence

and long backward directed brownish hairs, rest of body whitish pubescent.

Male unknown.

Brazil: 1 ©, holotype, Sao Paulo (S.P.), 28 Jan. 1965, V.N. Alin (UCD); 1 ©,

paratype, Sao Paulo, Villa Americana, Febr. 1924, F. X. Williams (BISH).

The specific name refers to the red band of the gaster.

Pluto simplicicollis sp. nov.

Female. — Similar to P. zonatus, but smaller, length about 6.5 mm. Labrum

reddish. Gaster black with reddish transparent hind margins of tergites. Petiole

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

nearly as long as first tergite. Frons near anterior ocellus and lower part of

mesopleura more shining.

Male unknown.

Brazil: 1 9, holotype, Säo Paulo, Campinas, March 1924, F. X. Williams

(BISH).

The petiole being distinctly longer than in P. zonatus, I provisionally consider

this form a distinct species. The colour of the gaster may be subject to variation.

The specific name refers to the lateral pronotal angles which are not projecting.

Pluto menkei sp. nov.

(fig. 42)

Female. — Length about 5 mm. Head and thorax black. Mandibles yellow with

dark tips. Underside of flagellum reddish-brown. Pronotal tubecles whitish,

tegulae yellowish-red. Apices of fore and mid tibiae and base of hind tibiae

yellowish. Fore tarsi and mid basitarsi whitish, rest of mid tarsi yellowish-brown;

hind tarsi dark brown, apices of segments yellowish-brown. Gaster black, except

for hind margin of tergite 1, all of tergite 2, base of tergite 3, all of sternite 2 and

base of sternite 3, which are red. Veins of wings dark brown or black.

Median part of clypeal margin weakly emarginate, about 1/3 of total width of

margin, no distinct lateral teeth (fig. 42). Disk of clypeus densely punctate, with

narrow shining margin. Frons medially finely punctate, laterally sparsely punctate,

a small shining raised area along eyes. Vertex and interocular space shining, very

sparsely punctate. POD somewhat smaller than OOD. Intercarinal space narrow,

occipital carina distinct. Antennal segments 10—11 in lateral view about as long as

broad.

Lateral pronotal angles sharp in dorsal view but hardly toothlike projecting.

Scutum shining, sparsely punctate. Scutellum indistinctly alutaceous, sparsely

punctate. Propodeal enclosure shining with large median area; back of propo-

deum somewhat dull, rather coarsely reticulato-carinate, carinae on upper part

oblique, parallel. Mesopleura shining, mostly sparsely finely punctate, a few larger

punctures intermixed. Hypo-epimeral area shining, indistinctly punctate. Anterior

plate of mesepisternum somewhat dull. Anterior part of mesosternum densely

punctate. Petiole about 2/3 length of first tergite, this tergite somewhat longer than

broad. Pygidial area almost twice as long as broad. Second recurrent vein of fore

wings interstitial.

Pubescence of face and of pronotal collar silvery, mostly appressed, of vertex

and dorsal side of thorax greyish-brown, longer, of last gastral segment golden-

brown, of rest of body whitish.

Male unknown (cf. remarks P. incarnatus 3).

Venezuela: 1 ©, holotype, Aragua, 2 km north of Ocumare de la Costa, 21—22

June 1976, A. S. Menke and D. Vincent (USNM).

This small species may be closely related to P. zonatus but the clypeal margin,

VAN LITH: New World Pluto 217

the length of the antennal segments, the colour of the facial pubescence and that

of the hind tarsi are different.

Pluto occipitalis sp. nov.

(figs. 44—45)

Female. — Length about 6 mm. Black. Mandibles yellowish-red, apex of

clypeus and labrum reddish. Underside of flagellum yellowish-brown. Pronotal

tubercles yellowish-white. Fore and mid tibiae and tarsi brownish-yellow, basitarsi

paler. Basal third of hind tibiae and hind basitarsus straw-yellow, segments 2—4 of

hind tarsi pale brown, last segment dark brown. Hind margins of gastral segments

transparent. Veins of wings black.

Median third of clypeal margin slightly protruding and emarginate (fig. 44).

Frons dull, raised on either side of anterior ocellus; vertex shining, not distinctly

punctate, a low tubercle along the eyes. POD about equal to OOD. Lateral and

ventral parts of occipital carina extraordinarily high, in dorsal view as high as

pubescence of tempora. Antennal segments 10—11 about quadrate.

Anterior dorsal carina of pronotum high, laterally protruding as a sharp tooth

(fig. 45). Scutum and scutellum somewhat shining, very finely reticulate

alutaceous, sparsely punctate. Prescutal sutures about as long as 3/4 of scutum,

indistinct. Propodeal enclosure shining, with few lateral carinae and large

pentagonal median area. Back of propodeum finely coriaceous and coarsely

reticulato-carinate. Mesopleura and hypo-epimeral area shining, not distinctly

punctate. Anterior oblique suture narrow. Mesosternum densely finely punctate.

Second recurrent vein of fore wings ending in third submarginal cell. Petiole as

long as first tergite, this tergite as long as broad. Pygidial area about 1.5 times as

long as broad. Back of hind tibiae with relatively long yellowish-brown thorns.

Pubescence of face silvery, mostly appressed; pronotal collar silvery pubescent,

pubescence of vertex, scutum and scutellum greyish-brown, of pygidial area

brown, short and appressed. the latter area also with long brown hairs. Rest of

body with whitish hairs.

Male unknown.

Peru: 1 9, holotype, Loreto, San Antonio, 13 Aug. 1965, Malaise trap, J. C.

Hitchcock Jr. (USNM).

P. occipitalis is easily distinguished by the smooth mesopleura and hypo-

epimeral area, the high occipital carina and the sharp lateral pronotal angles.

The specific name refers to the unusually high occipital carina.

Pluto spinicollis sp. nov.

(fig. 46)

Female. — Length about 6.5 mm. Black. Clypeal margin and labrum dark

reddish. Mandibles brownish-yellow with reddish tips. Underside of flagellum

orange-brown. Pronotal tubercles yellow. Fore and mid tibiae and tarsi yellow,

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

back of mid tibiae blackish, base of tibiae and entire basitarsi whitish. Base ot

hind tibiae and hind tarsi except for last segment yellowish. Petiole dark brown or

black. Tergites I—3 and sternites 2—3 red, following tergites and sternites with

reddish transparent hind margins. Veins of wings dark brown.

Clypeal margin very slightly emarginate, almost straight (cf. duckei, fig. 40).

Disk of clypeus dull. Frons densely finely punctate. Vertex shining, sparsely

punctate. Intercarinal space narrower than first basitarsus. Antennae short,

clavate, segments 9—11 about quadrate, last segment about 1.5 times as long as

broad.

Lateral pronotal angles, in dorsal view, with a small sharp tooth (fig. 46). Scutum

and mesopleura shining, almost imperceptibly reticulate alutaceous and sparsely

very finely punctate. Hypo-epimeral area shining, indistinctly sparsely punctate.

Scutellum dull, finely reticulate alutaceous, with sparse distinct punctures.

Metanotum dull, densely punctate. Enclosed area of propodeum shining, with

oblique lateral carinae and pentagonal central area. Back of propodeum almost

shining, coarsely reticulate, dorso-lateral carinae about parallel. Anterior oblique

suture narrow, indistinctly crenulate, widened upper part smooth. Anterior plate

of mesepisternum dull, indistinctly sculptured. Scond recurrent vein of fore wings

interstitial. Petiole little over half length of first tergite, this tergite about as long as

broad. Pygidial area less than twice as long as broad.

Pubescence of face, frons and tempora silvery, below antennae mostly

appressed but also with long erect hairs. Pronotal collar with appressed silvery

pubescence. Last tergite with golden-brown hairs, longer and erect on sides; short,

dense and appressed on pygidial area; rest of body yellowish-grey pubescent.

Male unknown.

Brazil: Roraima, 2 9, holotype and paratype, Surumu, Sept. 1966, M. Alvarenga

(HT); 1 9, paratype, same locality and same date, M. Alvarenga (UCD).

Panama: | ©, paratype, Bella Vista, 6 July 1924, N. Banks (MCZ). In this female

the posterior half of the third tergite is darkened.

The shape of the pronotum and the red tergites easily distinguish P. spinicollis

from other Pluto with shining mesopleura.

The specific name refers to the lateral teeth of the pronotal collar.

Pluto trilobatus sp. nov.

(fig. 47)

Female. — Length about 6.5 mm. Black, gaster blackish-brown. Mandibles

yellowish or reddish-yellow with dark reddish tips; labrum reddish. Underside of

flagellum dark yellowish-brown. Pronotal tubercles dark brown to black.

Trochanters of fore legs, fore tibiae, foreside of mid tibiae and basal fourth of hind

tibiae yellowish-brown. Fore and mid basitarsi yellowish-white or brownish-

yellow, segments 2—5 dark brown. Hind tibial spurs whitish, hind tarsi brown.

Veins of wings including stigma dark brown.

Clypeal apex with distinct small lateral teeth, distance between these teeth

about half total width of margin; clypeal margin medially somewhat raised and

VAN LITH: New World Pluto 219

protruding (fig. 47). Disk of clypeus dull. Frons and vertex smooth and shining,

frons superficially finely punctate, vertex almost impunctate. POD distinctly

smaller than OOD. Occipital carina not continued to ventral carina, distance from

ventral carina and from hypostomal carina about length of first basitarsus.

Antennae somewhat clavate, segments 10—11 longer than broad, segment 12

twice as long as broad.

Lateral pronotal angles obtuse. Scutum and scutellum dull, finely reticulate

alutaceous and very finely punctate, interstices a few times size of punctures.

Prescutal sutures indistinctly continued to hind margin of scutum. Scutellar suture

except for distinct median longitudinal carina relatively indistinctly crenulate.

Propodeum including enclosure dull, very finely coriaceous, enclosure with large

pentagonal median area, dorsal half of back with oblique parallel carinae, lower

part reticulato-carinate. Mesopleura and hypo-epimeral area smooth and shining,

very indistinctly punctate. Mesosternum indistinctly finely punctate. Anterior

oblique suture indistinctly crenulate. Second recurrent vein of fore wings

interstitial. Mid tibiae with about seven slender thorns which are about as long as

4/5 width of tibiae. Petiole about length of first tergite, laterally flattened with

distinct upper and lower carina. First tergite longer than broad. Pygidial area

nearly triangular, about | % times as long as broad, sometimes less.

Face densely silvery, mostly appressed, pubescent and with long erect hairs.

Tempora with short silvery pubescence. Vertex, scutum and scutellum with

greyish-brown hairs. Pygidial area with dense golden-brown backwards directed

pubescence. Rest of body greyish pubescent.

Male. — Similar to female, but underside of antennae pale yellowish-brown.

Clypeal margin not raised, with three indistinct teeth. Antennae slender, most

segments about twice as long as broad, segments 5—8 with linear tyloides, as long

as segment, in lateral view distinctly roundly raised in the middle; segment 4 with

long narrow fine tyloides, segment 9 with very short fine tyloides. Petiole nearly

1.5 times as long as first tergite, this tergite about 1.5 times as long as broad.

Ecuador: Napo Prov., Limoncocha on Rio Napo, 1 ©, holotype and 2 &,

allotype and paratype, 250 m, 15—28 June 1976, S. and J. Peck (CNC); 2 2 and 1

d, paratypes, 9 June 1977, D. L. Vincent (USNM); 1 ©, paratype, 22 July 1974,

Malaise trap (FSC).

Peru: 1 ©, paratype, Madre de Dios, Avispas, 400 m, 10—30 Sept. 1962, L. Pena

(MCZ).

Surinam: | 9, paratype, Kabalebo River, Avanavero Falls, 5—12 April 1971, W.

Surinam Exp., D. C. Geijskes (ML).

The slightly raised and protruding median part of the clypeal disk of the female,

the small number of tyloidea of the male and the incomplete occipital carina in

both sexes of P. trilobatus are characteristic. The specific name refers to the

trilobate or tridentate clypeal margin of the female.

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pluto sp. aff. trilobatus

(fig. 48)

A female from Peru is very similar to P. trilobatus, but it is somewhat smaller and

differs also in the following details.

Length about 6 mm. Median part of clypeal margin weakly emarginate, small

lateral teeth, distance between these teeth about half total width of clypeal margin,

in the middle a very small protruding tooth (fig. 48). Occipital carina incomplete,

distance from ventral median carina about length of first basitarsus and from

hypostomal carina about half this length. Pygidial area about 1.5 times as lope = as

broad.

Peru: 1 9, Avispas, 30 m, nr. Marcapata, 27—30 Oct. 1962, Luis Pena (HT). As

the median clypeal tooth is smaller than in the preceding form and as the pygidial

area seems to be narrower, I had originally considered this female to belong to a

distinct species. Later I could examine another female from Avispas in Peru,

which is identical with the holotype of P. trilobatus. Therefore I do not exclude the

possibility that the first mentioned female is an aberrant form of P. trilobatus.

Pluto rufanalis sp. nov.

Female. — Length about 5 mm. Black. Mandibles, labrum and underside of

flagellum yellowish-red, dorsal side of antennae brown. Pronotal tubercles whitish.

Fore and mid tibiae yellowish, first tarsal segments whitish, outer side of mid tibiae

brownish. Basal third of hind tibiae and tarsal segments 1—4 yellowish-white.

Gaster black, hind margins of tergites and last segment largely or entirely reddish.

Veins of wings dark brown.

Median part of clypeal margin straight, narrow margin depressed and shining;

small lateral teeth, distance between these teeth less than half total width of

margin. Disk densely punctate. Frons densely superficially and finely punctate.

Vertex shining, almost impunctate. POD about equal to OOD. Intercarinal space

much narrower than first basitarsus. Antennae clavate, segments 10—11 shorter

than broad.

Lateral pronotal angles rectangular. Scutum and scutellum shining, sparsely

punctate. Propodeal enclosure shining, large median area. Back of propodeum

indistinctly coriaceous, coarsely reticulato-carinate. Mesopleura and meso-

sternum shining, slightly reticulate alutaceous, sparsely indistinctly punctate.

Hypo-epimeral area smooth. Anterior plate of mesepisternum dull, oblique suture

crenulate. Second recurrent vein of fore wings interstitial. Petiole nearly as long as

first tergite, this tergite somewhat longer than broad. Pygidial area about 1.5 times

as long as broad.

Pubescence of face yellowish-silvery, mostly appressed, of pronotal collar

silvery, appressed, of pygidial area golden-brown, of rest of body whitish or

greyish.

Male. — Length about 4.5—5 mm. Colour as in female but hind tarsi largely

pale brown.

Antennae slender, segments about 1.5 times as long as broad, no distinct

VAN LITH: New World Pluto 221

tyloidea, Mesopleura and hypo-epimeral area shining, mesopleura sparsely finely

punctate. Gaster slender, petiole about as long as first tergite, this tergite over 1.5

times as long as broad. Face silvery.

Peru: 4 9, holotype and paratypes and 2 g, allotype and paratype, Chancay,

River valley, 15 March 1951, Ross and Mickelbacher (CAS); 1 9, paratype, Piura,

Ostendorf (ZMB); 264, paratypes, Pariñas Vall., 7—8 April, Mrs. Frisell (MCZ).

P. rufanalis is closely related to P. nitens, but apart from the smaller size the

female differs in the yellowish-white pronotal tubercles and hind tarsi, the male in

the indistinct or absent tyloidea and the very slender gaster. It also much

resembles P. araguensis which, however, has sharp pronotal angles and more

densely punctate mesopleura.

The specific name refers to the reddish apex of the gaster.

Pluto araguensis sp. nov.

(figs. 49— 50)

Female. — Length about 5—6 mm. Black; mandibles yellowish-red with darker

tips, labrum reddish. Underside of flagellum reddish-brown. Fore and mid tibiae

and tarsi, base of hind tibiae and hind tarsal segments 1—4 yellowish, outer side of

fore and mid tibiae with narrow whitish streak. Pronotal tubercles whitish, tegulae

reddish-brown. Hind margins of gastral segments reddish transparent.

Median part of clypeal margin slightly emarginate, distance between the very

obtuse angles about half total width of clypeal margin (fig. 49). Frons densely

finely punctate, laterally with large, slightly swollen smooth area. Vertex shining,

sparsely punctate. POD somewhat smaller than OOD. Intercarinal space about

half width first basitarsus. Antennal segments about as long as broad.

Lateral pronotal angles very sharp (fig. 50). Scutum and scutellum almost

shining, weakly reticulate alutaceous, distinctly punctate, interstices mostly a few

times size of punctures. Propodeal enclosure shining, with large pentagonal

median area. Back of propodeum reticulato-carinate, dorso-lateral carinae

parallel. Mesopleura, mesosternum and hypo-epimeral area shining, very finely

alutaceous, mesopleura regularly finely punctate, interstices a few times size of

punctures, hypo-epimeral area indistinctly punctate. Anterior plate of mesepi-

sternum dull, oblique suture narrow, indistinctly crenulate. Petiole about 2/3

length of first tergite. Pygidial area about 1.5 times as long as broad. Second

recurrent vein of fore wings interstitial.

Pubescence of face and pronotal collar silvery, mostly appressed, of pygidial

area dark brown, of rest of body whitish or greyish.

Male. — Length about 5—5.5 mm. Underside of flagellum largely brown. Fore

and mid femora and tibiae and basal 1/3 of hind tibiae yellowish-red, fore and mid

tarsi yellowish-white, hind tarsi pale yellowish-brown or yellowish-white. Rest of

body of same colour as in female.

Median part of clypeal margin slightly protruding, about 1/4 of total width of

margin, disk shining, densely punctate. Antennae slender, segments 10—12 about

1.25 times as long as broad, tyloidea indistinct.

222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Lateral pronotal angles about rectangular. Mesopleura shining, sparsely punc-

tate; hypo-epimeral area shining, indistinctly punctate. Back of propodeum

coarsely reticulato-carinate, dorso-laterally with oblique parallel carinae; in the

allotype these carinae are not distinctly parallel. Petiole about as long as first

tergite, this tergite about 1.5 times as long as broad. Pubescence as in female.

Genitalia reddish-brown.

Venezuela: Aragua, 2 km north Ocumare de la Costa, 3 9, holotype and

paratypes, 1 g, allotype, 21—22 June 1976, A. S. Menke and D. Vincent

(USNM); 1 3, paratype, Puerto Caballo, 12 Febr. 1940, 1 3, paratype, San

Esteban, Jan. 1940 (CU).

P. araguensis, at least the female, seems to be very closely related to P. spini-

collis, although the pronotal angles are projecting laterally instead of obliquely

forwards. Its gaster is entirely black, that of P. spinicollis partly red. The pronotal

angles of the male of P. araguensis are less sharp than in the female and the

colour of the femora is much paler. However, as one of the males has been

taken together with three females they may safely be regarded as the same species.

Pluto incarinatus sp. nov.

Male. — Length about 5.5 mm. Head and thorax black; mandibles pale yellow

with dark reddish tips, labrum black, antennae black, underside of last antennal

segment reddish-brown. Pronotal tubercles whitish; tegulae light brown, wings

somewhat smoky, veins dark brown. Femora black. Fore tibiae almost entirely

and greater part of underside of mid tibiae yellowish-brown, rest of tibiae dark

brown; bases of mid and hind tibiae yellowish-white. Fore and mid tarsi whitish,

hind tarsal segments brown with paler apices. Tibial spurs of hind legs whitish.

Apical half of first tergite, all of second tergite and all of second sternite red, rest

of gaster black with bluish reflection.

Protruding median part of clypeal margin about 1/4 of total width, weakly

emarginate. Intercarinal space probably narrow. Median part of frons densely

punctate, vertex and interocellar area shining, sparsely punctate, tempora dull.

POD about equal to OOD. Antennal segments 10—12 nearly 1.5 times as long as

broad, segment 13 over twice as long as broad. No tyloidea.

Lateral pronotal angles sharp, in dorsal aspect. Scutum shining, very finely

reticulate alutaceous with widespread distinct punctures. Scutellum almost

shining, sparsely finely punctate, anterior suture coarsely crenulate. Back of

propodeum shining, coarsely reticulato-carinate; enclosure with shining median

area. Hypo-epimeral area shining with a few minute hair-bearing punctures.

Mesopleura and mesosternum shining, regularly finely punctate, interstices about

two to four times size of punctures. Mesosternum very finely reticulate alutaceous.

Anterior plate of mesepisternum dull, finely striate; oblique suture narrow,

crenulate. Legs stoutly built. Petiole about as long as first tergite, this tergite about

1.5 times as long as broad. Second recurrent vein of fore wings about interstitial.

Pubescence of face pale golden to silvery, mostly appressed. Tempora, pronotal

VAN LITH: New World Pluto 223

collar and metanotum pale golden pubescent, scutum brownish, rest of thorax pale

golden, propodeum scarcely pubescent. Gaster greyish-golden pubescent.

Female unknown.

a ap

Figs. 49—50. P. araguensis sp. n., 9, holotype: 49, clypeus: 50, pronotum. Figs. 51—53. P. annulipes

| (Cameron); 51, clypeus of 9 ; 52, pronotum of 9; 53, clypeus of &. Figs. 54—55. P. denticollis sp. n., 9,

| holotype; 54, clypeus; 55, pronotum.

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Venezuela: 1 ¢, holotype, Aragua, 2 km north Ocumare de la Costa, 12 June

1976, A.S. Menke and D. Vincent (USNM).

This male is very similar to P. menkei sp. nov. described after a single female

collected at the same locality but not on the same day. In view of the longer petiole

and the different ratio of POD : OOD, I provisionally consider it a distinct species,

also very closely related to P. araguensis, from which it differs in the colour of

gaster and legs.

The specific name refers to the absence of tyloidea or carinae on the antennae.

Group of annulipes

Pluto annulipes (Cameron)

(figs. 51—53, 56—57)

Cameron, 1891: 130—140, 9 (Psen annulipes ; Mexico).

Kohl, 1896: 292.

Dalla Torre, 1897: 347.

Ashmead, 1899: 225.

Bohart & Menke, 1976: 171 (Pluto annulipes, new combination by Bohart).

Female. — Length 10—12 mm. Black. Mandibles, base of flagellum below and

last antennal segment reddish-brown. Pronotal tubercles dark brown or black,

tegulae and veins of wings dark brown, stigma yellowish-brown, wings infuscated.

Foreside of fore tibiae reddish-brown, outer side of base ivory-white. Mid tibiae

dark brown or black, base ivory-white. Basal 2/5 of hind tibiae in lateral view

whitish, back of tibiae nearly entirely whitish. Tarsi brown; basitarsus of mid legs

and basitarsus and second tarsal segment of hind legs whitish, extreme base of hind

basitarsus dorsally brownish, sometimes also second tarsal segment of hind legs

pale brown. Hind margins of tergites somewhat reddish transparent.

Protruding median part of clypeal margin (fig. 51) about half total width of

margin, almost straight, lateral angles oblique. Frons shining, depressed between

anterior ocellus and oculi, lower part of frons medially with fine hairbearing

punctures. Ocellar region raised. Vertex shining, sparsely punctate. POD equal to

OOD. Intercarinal space about as wide as fore basitarsus. Occipital carina high,

mid-ventrally receding backward. Antennae slightly widening towards apex,

segments longer than broad.

Lateral pronotal angles with long lobular projection (fig. 52). Propleura

emarginate below. Scutum, scutellum, sides of thorax and mesosternum shining,

sparsely punctate. Scutellum impressed medially, anterior suture crenulate.

Propodeal enclosure shining, large pentagonal median area; back of propodeum

dull owing to microsculpture, coarsely reticulato-carinate. Petiole nearly as long

as first tergite, this tergite longer than broad. Pygidial area slightly over twice as

long as broad. Base of hind tibiae with many short, broad thorns. Second recurrent

vein of fore wings ending in third submarginal cell.

Pubescence of face pale golden, mostly appressed, pubescence of pronotal

VAN LITH: New World Pluto 225

collar more silvery, appressed, pygidial area with dark-brown to golden-brown

backward directed pubescence. Rest of body greyish pubescent.

Male (first description). — Length 9—10 mm. Very similar to female. Bases of

fore and mid tibiae may be dark. Mid and hind basitarsi whitish, second tarsal

segments not so. Clypeal margin slightly emarginate (fig. 53). Antennal segments

5—11 or 5—12 with linear tyloides, on segments 5—10 reaching at most middle of

segments, 11—12 about as long as 1/3 of segment, a very short tyloides on segment

13. Petiole somewhat longer than first tergite, this tergite 1.5 times as long as

broad.

Pubescence of face silvery.

Genitalia: figs. 56—57.

Material examined: Mexico: | 9, type No. 21.827, Guerrero, Rincon, 2800 ft,

September, H. H. Smith; 1 9, Guerrero, Chilpango, 4600 ft, October, H. H.

Smith, ‘’Psen annulipes Cam.”; 1 ©, Tabasco, Teapa, February, H. H. Smith,

Cameron Collection 1914—110 (BM). All these specimens are recorded by

Cameron (1891). I could not trace the female recorded from Amula (6000 ft). In

the Naturhistorisches Museum, Vienna, is a female from Mexico, Orizaba, 5 May

1871, Bilimek, labelled “annulipes Cam.” by Kohl.

New records from Mexico: Chiapas: 1 © and 1 g, Simojoval, 17 March 1953, E.

I. Schlinger; 1 9 and 2 g, 20 mi south Tuxtla Gutiérrez, 12 Aug. 1963, F. D.

Parker and L. A. Stange (UCD). Durango: 1 g, Tlahualilo, July 1905, Hunter No.

888, A. W. Morrill (USNM). Guerrero: 1 9, 8 mi north Taxco, 5500 ft, 19 June

1959, H. E. Evans (CU). Hidalgo: 1 © and 1 g, Jacala, 4500 ft, 31 Aug. 1963,

Scullen and Bolinger (OSU). Jalisco: 1 9, Guadalajara, Crawford; 1 9, 20 mi

north La Quemada, 27 July 1954, M. Casier and W. Gertsch Bradts (UCD).

Morelos: 4 ©, Yautepec, 31 July 1963, F. D. Parker and L. A. Stange (UCD); 1 9,

26 mi south Cuernavaca, 3150 ft, 28 Aug. 1963, Scullen and Bolinger (OSU).

Nuevo Léon: I &, Linares, 5 Oct. 1962, H. and M. Townes (HT). Oaxaca: | g, 104

mi south Acayucan, 750 ft, 18 Aug. 1963, Scullen and Bolinger (OSU). San Luis

Potosi: 1 9, El Salto, 1800 ft, 8 June 1961, on flowers of Kallstroemia hirsutissima,

Univ. Kansas Mex. Exped. (KU). Tamaulipas: 2 9, Gomez, Fariäs and vic.,

400—600 m, 20—24 July 1965, in Malaise trap, Cornell University Mexico Field

Party (CU). Veracruz: 2 9, Fortin de las Flores, 11 and 14—21 Sept. 1954, F. X.

Williams (CAS); 1 g, Cordoba, 20 July 1966, J. S. Buckett, M. R. and R. C.

Gardner (UCD).

The male from Oaxaca has black fore and mid legs, only foreside of fore tibiae

being somewhat brownish. The tyloidea are shorter, less than 1/4 length of

segment on segments 9—10, very small on segment 11, lacking on segments

12—13. The male from Simojoval has entirely brown tarsi. Also the females may

have their hind tarsi different from the usual form; sometimes the second tarsal

segment or even a part of the basitarsus are brown or these parts are paler than

usual, not only the basitarsus and the following segment being whitish, but also the

third tarsal segment.

First record from Costa Rica: 1 &, Liberia, 400 ft, 29 July 1963, Scullen and

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Bolinger (OSU). This male has dark hind basitarsi, the hind margins of the tergites

are more reddish, length about 8 mm. It may represent a different subspecies.

First record from El Salvador: 1 9, Quezaltepeque, 17 June 1963, D. Cavagnaro

and M.E. Irwin (UCD). The greater part of the hind basitarsi and all of the second

tarsal segment are brownish. 1

P. annulipes and P. smithii are the largest species of the genus Pluto. They are

readily distinguished by the long lateral projections of the pronotum. The females

of the two species differ in colour, mainly in that of the legs; the males have

different tyloidea.

Pluto smithii (Fox)

(figs. 58—60)

Fox, 1898a: 378—379, g (Psen Smithii; Brazil, Chapada (dos Guimaraes), Mato Grosso).

Bohart & Menke, 1976: 171 ( Pluto smithii, new combination by Bohart).

Male. — Much resembling the male of P. annulipes. Length about 10 mm.

Apical half of fore femora, all of fore and mid tibiae and tarsi reddish-yellow, mid

basitarsi paler. Basal 2/5 of hind tibiae whitish. Hind tarsi yellowish-white, last

segment largely brown. Pronotal tubercles and tegulae yellowish. Base of second

tergite red. Antennae including scape yellowish-red, dorsally partly brown. Stigma

of fore wings little paler than veins. Tyloides on antennal segments 5—6 about 3/4

length of segments, on segments 7—12 almost as long as segments, a very short

tyloides on segment 13.

Face and tempora pale golden, mostly appressed, pubescent. Thorax and gaster

with brownish or yellowish-silvery hairs, underside of hind femora with long

silvery-white hairs.

Genitalia: figs. 58—60.

First description of female. — Resembling male, structurally not different from

the female of P. annulipes. Length 11—13 mm. Gaster black, hind margins of

segments somewhat reddish transparent. Pronotal tubercles yellowish. Fore tibiae

entirely yellowish-red. Base of hind tibiae reddish, with many brown, flattened

thorns. Fore basitarsus usually reddish, mid basitarsus and mid tarsal segment 2 or

segments 2—3 whitish. Hind basitarsus and following two or three segments

whitish, basitarsus dorsally with a row of thorns, as on hind tibiae. Pubescence of

face darker golden.

New records from Brazil: Mato Grosso: 1 ©, Sinop, 12°31’S, 55°37’W, Oct.

1974, M. Alvarenga (HT). Minas Gerais: 1 9, Alpinopolis, Febr. 1961, C. Elias; 2

.d', Passos, 17—23 April and 7 May 1963, C. Elias (UFP). Para: 1 9, 10 Sept. 1901,

Ducke, 273 (NMW); 1 ©, Soure, Ilha de Marajò, O. Bertram (ZMB). Rio de”

Janeiro: | 9, Mangaratiba, April 1962, M. Alvarenga (UFP).

This species apparently replaces the Central American P. annulipes in South

America. Because of the different length of the tyloidea of the males it is likely

that P. smithii is a distinct species with many subspecies. Chapada, in Brazil, being

VAN LITH: New World Pluto 227

the type-locality I have provisionally considered the Brazilian material, two

females and two males from Minas Gerais and Rio de Janeiro, and a female from

Mato Grosso, to belong to the form as described by Fox. The identification of the

two females from Para is somewhat doubtful, their fore tibiae and their antennae

being darker than in the other Brazilian specimens. A series of males from

Paraguay is closely related to the Brazilian form; unfortunately I have seen no

females from Paraguay. The other forms are recorded below together with their

distinctive characters. A study of ample material from more regions could be

interesting.

Form A. Paraguay: 36 3g, San Bernardino, mid Febr. — early April, Fiebrig

(NMW). Length 8—10 mm. Tyloidea on segments 4—13 somewhat longer,

reaching nearly to end of segment on segments 5—6. Gaster more or less red,

sometimes all of tergites 1—2 and sternite | red, rarely only base of tergite 2 red.

Basal four segments of hind tarsi whitish. Stigma of fore wings somewhat paler.

Face and tempora silvery pubescent, vertex and dorsal side of thorax brownish.

Form A. Bolivia: 1 3, Buenavista near Santa Cruz, 1928, J. Steinbach (CU); 2

gd, Prov. Sara, Steinbach (MCZ). Face slightly golden, base of second tergite red.

Form B. Paraguay: I &, Caballero, Nov. 1971, Pena (MF). Base of tergite 2

black. Segments 4—S of hind tarsi dark brown. Stigma dark. Distinct tyloidea on

segments 5—13, about 3/4 length of segments. Face pale golden.

Form C. Argentina: Salta: 3 © and 1 g, Rio Pescado (Est. YPF), 19—25 Nov.

1967, C. Porter, E. Willink (MCZ); 1 ©, Prov. Salta, 1200 m (NMW); Yuto, 1 9,9

Jan. 1966, H. and M. Townes (HT). Salta-Orán: 1 &, Abra Grande, 10 Jan. — |

March 1967, R. Golbach (IML); 4 9, Abra Grande, 18 April—5 May 1969, C.

Porter (MCZ); 1 9, Jujuy (MF). Tucuman: 1 9, Horco Molle, Dec. 1968, C. C.

Porter (CNC). Scape of antennae and greater part of dorsal side of flagellum black

in both sexes. Fore tibiae of female sometimes with dark outer streak. Usually hind

basitarsus and next two or three segments whitish. Hind margin of tergite 1 often

reddish. Base of tergite 2 black. Pronotal tubercles yellowish-red. Face of male

pale golden, of female darker golden.

A series of females collected in the province of Entre Rios is very similar but the

scape of the antennae is entirely red in most specimens. Hind tarsi except for the

last segment whitish. Fore tibiae and tarsi entirely reddish. Base of hind tibiae

whitish. First gastral tergite in all females except for the specimen collected in

1964 with broad red hind margin. Argentina: Entre Rios: Pronunciamento, | 9,

March 1964, 1 9, 3—9 Febr. 1965, 6 9, 28 Febr. 1965 (FAG).

Form D. Argentina: 2 ¢, Tucumän, El Solidad, 11 km west Las Cejas, 13 Febr.

and 15 Dec. 1967, L. A. Stange (UCD). Tyloidea short as in P. annulipes. Face

silvery. Segments 1—3 of hind tarsi whitish. Apical half of tergite 1 conspicuously

red, also hind margins of following segments reddish. Scape of antennae reddish.

Form E. Panama Canal Zone: 1 9, Culebra-Arrijan Trail, 25 Dec. 1914, T.

Hallinan (AMNH); | 9, Barro Colorado Is., 16 March 1967, Roger Dakro (UCD).

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Figs. 56—57. Pluto annulipes (Cameron), genitalia of &, Acayucan, dorsal and lateral aspect. Figs.

58—60. P. smithii (Fox), genitalia of 3, Paraguay, dorso-lateral, lateral and dorsal aspect.

VAN LITH: New World Pluto 229

Pronotal tubercles dark brown with paler hind margin. Gaster black. Posterior

surface of fore tibiae brown. Base of hind tibiae with dark brown thorns. Segments

i—3 of hind tarsi yellowish-white. Stigma of fore wings paler than veins. Face pale

golden.

Form F. Surinam: | 9, NE Surinam (Exp. I), Mungatapu Wia Wia, 3rd camp

line km 14.9, “ritsenbos 1795”, 15 Oct. 1948, D. C. Geijskes c.s.; 1 ©, Republiek,

24—26 Sept. 1963, Malaise trap, D. C. Geijskes; 1 ©, Zanderij, Kreekbos, 18—21

July 1964, Malaise trap, D. C. Geijskes; 1 9, Mapane Area, Camp 8, LBB, 29 May

1963, J. van der Vecht (ML). Resembling the Brazilian form but scape of antennae

black and fore tibiae with black streak on outer surface. Gastral tergites 1—2 not

distinctly red. Pronotal tubercles reddish-yellow, stigmata of fore wings paler than

veins. Segments 1—3 or 1—4 of hind tarsi whitish.

Form F. Venezuela: | 9, Trujillo, Sabana Grande, 3 June 1976, 2 9, Aragua, 2

km north Ocumare De La Costa, 12 and 21—22 June 1976, 1 9, Zulia, 6 km west

La Concepcion, 18 June 1976, A. S. Menke and D. Vincent (USNM). Like

Surinam females but pronotal tubercles dark brown or black in three of the

specimens, in one female they are yellowish-brown.

Form F. Trinidad: 1 9, Maraccas Valley, 4 Aug. 1937, O. W. Richards (BM).

Form G. Colombia: 1 9, Meta, La Macarena, 20—29 Nov. 1976, M. Cooper

(BM). Differs from the Surinam females in having the antennae almost entirely

black, only segments 3—4 being somewhat reddish beneath. Pronotal tubercles

brownish with yellowish hind margin. Hind tarsal segments 1—4 whitish.

A male from Bogota, Lindig, 19704 (ZMB) may belong to the same form. Scape

of antennae partly red. Tergite 1 with narrow reddish hind margin. Fore tibiae

entirely reddish. Face pale golden. The tyloidea on segments 5—12 are relatively

short, about 2/3 length of segments; only basitarsus of hind legs whitish.

Pluto denticollis sp. nov.

(figs. 54—55)

Female. — Length about 8.5 mm. Black. Mandibles yellowish-red. Flagellum of

antennae orange-brown, apical half dorsally darker brown. Pronotal tubercles

dark brown. Tegulae yellowish-red. Fore and mid tibiae and tarsi, base of hind

tibiae, hind tarsi reddish-yellow. Hind margins of tergites and sternites reddish

transparent, pygidial area dark reddish. Veins of wings brown, stigma pale brown.

Median part of clypeal margin smooth and shining, four blunt lobes (fig. 54).

Upper 2/3 of disk densely punctate. Frons and vertex smooth and shining.

Intercarinal space narrower than width of fore basitarsus. Antennal segments

10—11 about quadrate, last segment about 1.5 times as long as broad.

Lateral pronotal angles in dorsal aspect protruding as an equilateral triangle (fig.

55) with rounded apex. Scutum and scutellum shining, sparsely finely punctate.

Prescutal sutures indistinct. Metanotum dull. Enclosed area of propodeum

shining, with sharp oblique lateral carinae and with large median pentagonal area.

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Back of propodeum reticulato-carinate, dorso-lateral carinae parallel. Sides of

thorax and mesosternum smooth, mesopleura almost impunctate. Anterior

oblique suture narrow, weakly crenulate, widened upper part smooth. Petiole

slightly longer than first tergite, this tergite about as long as broad. Pygidial area

less than twice as long as broad, densely punctate. Second recurrent vein of fore

wings interstitial.

Pubescence of face golden, mostly appressed. Tempora with short golden

pubescence. Frons and vertex with sparse, long golden-brown hairs. Pubescence

of scutum, scutellum and metanotum brownish-golden, pygidial area densely

golden-brown pubescent, last tergite laterally with long erect brown hairs;

remainder of thorax and of gaster yellowish-grey pubescent.

Male unknown.

Peru: 1 9, holotype, “Peru” (NMW).

As P. denticollis has conspicuous triangular projections of the pronotum and as

its scutum and mesopleura are smooth and its face is golden, this species is easily:

recognized. It seems to belong to the group of annulipes (Cameron) but differs in

the shape of the clypeal margin which has four small lobes instead of being

straight. .

The specific name refers to the lateral projections of the pronotal collar.

Group of pygmaeus

Pluto pygmaeus pygmaeus (Brethes) comb. nov.

(figs. 61—62)

Brèthes, 1913: 130, 9 ( Gorytes pygmaeus; Argentina: La Paz).

Bohart & Menke, 1976: 490 ( Ochleroptera pygmaea).

Female. — Length about 5.5 mm. Black, following parts orange-brown: mandi-

bles, underside of flagellum, fore and mid tibiae and tarsi, base of hind tibiae.

Labrum reddish. Hind tarsi brownish with paler apices. Pronotal tubercles and

tegulae yellowish-brown. Sternites with reddish hind margins. Last gastral segment

reddish-brown. Veins of wings brown.

Median part of clypeal margin slightly emarginate, almost straight, with small

lateral teeth or angles, distance between these teeth about half total width of

clypeal margin (fig. 62). Before narrow margin a low shining ridge. Disk densely

punctate. Median part of frons densely finely punctate, laterally a low shining

tubercle, vertex almost smooth, sparsely finely punctate. Back of head, especially

laterally, very finely reticulate alutaceous. Intercarinal space narrow. Antennae

short, clavate, segments 10—11 shorter than broad.

Pronotal angles sharp (fig. 61). Anterior part of scutum finely reticulate

alutaceous, rest shining, sparsely finely punctate. Scutellum entirely reticulate

alutaceous, more densely punctate, interstices once or twice as large as punctures.

Axillae triangular with rounded apex, on inner and posterior side free from

scutellum, reaching about halfway distance from base to scutellum (fig. 61).

VAN LITH: New World Pluto 231

Figs. 61—62. Pluto pygmaeus pygmaeus (Brèthes), 9, Argentina; 61, thorax in dorsal aspect; 62, clypeus.

Figs. 63—65. P. facialis sp. n.; 63, clypeus of 9, holotype; 64—65, head of &, allotype, ventral and fron-

tal aspect. Fig. 66. P. axillaris sp. n., Bolivia, genitalia of g, dorsal aspect. Figs. 67—71. P. joergenseni

(Brethes); 67, head of 9, Cordoba; 68, labrum of same, more enlarged; 69, labrum of g, Rio Negro,

same enlargement; 70, metanotum of &, Bolivia; 71, raised part of metanotum in lateral aspect.

232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Scutellar suture smooth, laterally open. Propodeal enclosure shining, large median

area; back of propodeum coarsely reticulato-carinate (fig. 61). Mesopleura and

mesosternum finely reticulate alutaceous, finely punctate, interstices on meso-

pleura a few times size of punctures, mesosternum densely finely punctate. Hypo-

epimeral area finely reticulate alutaceous, sparsely punctate. Anterior plate of

mesepisternum dull, anterior oblique suture crenulate, widened upper part with a

transverse carina. Second recurrent vein of fore wings interstitial. Petiole about as

long as first tergite, this tergite longer than broad. Pygidial area little more than 1.5

times as long as broad.

Face silvery, mostly appressed, pubescent; pygidial area golden-brown, rest of

body greyish-white pubescent.

First description of male. — Length about 5 mm. Similar to female, gaster more

slender. Clypeal margin medially somewhat protruding, clypeus before margin

somewhat thickened. Antennal segments 10—12 somewhat longer than broad,

segment 13 about 1.75 times as long as broad; segments 5—12 with long linear

tyloides, often short on segment 5, sometimes also one on segment 13. Occipital

carina and hypostomal carina very close. Vertex behind ocelli very finely striate.

Axillae with parallel sides, little longer than broad. Petiole about as long as first

tergite, this tergite 1.5 times as long as broad.

Material examined: 1 ©, holotype, with labels ““Mendoza’’, "No. 10401”, and

““Gorytes pygmaeus Br.” P. Jörgensen (MACN).

Brethes made no mention of the peculiar shape of the axillae and he wrongly

placed his species in the genus Gorytes. Mr. M. A. Fritz, Buenos Aires, discovered

that the type is a Psenid wasp and he was very helpful to have it sent to me for

study. He also informed me that La Paz, which locality is mentioned only in the

original description, not on the label pinned under the insect, is a small village in

the province of Mendoza (Argentina), where Jorgensen often collected insects.

New records from Argentina: Catamarca: 1 9, 6 km north Belén, 1240 m, 1—15

Jan. 1970, Malaise trap, Entomofauna Subandina, Willink-Teran-Stange (IML); 2

©, 16—30 Nov. 1968, 4 9, 1—31 Jan. 1969, Los Nacimientos de Abajo, Malaise

trap, Entomofauna Subandina, Willink-Terän-Stange (IML); 1 9, Belén, 25 Nov.

1975, R. M. Bohart, 1 9, La Cienega, 17 Dec. 1975, R. M. Bohart (UCD). Jujuy: 2

Q and 3 4, Porto Lozano, 28 Nov. — 2 Dec. 1967, 6 © and 9 g, Posta Lozano, 28

Nov. — 19 Dec. 1967, 4 © and 4 Z, Rio Lozano, 28 Nov. — 20 Dec. 1967, C. C.

Porter and E. Willink (MCZ). Mendoza: 16 9 and 5 4, 25 Nov. — 25 Dec. 1906, 1

Q, | Febr. 1907, Mendoza, Jensen-Haarup (ZMB); 10 © and 3 3, Mendoza

(NM W); 2 9, Mendoza, Chacras de Coria, 21 Febr. 1966, S. Stange (IML). Salta: 1

Q, Rio Pescado (Est. YPF), 19—25 Nov. 1967, C. Porter and E. Willink; 1 9,

Cachi, 20—22 Jan. 1966, 1 © and I &, Oran, Abra Grande, 18—25 Oct. 1968, 1 9,

Camp Jakülica, ca. Aguas Blancas, Oct. 1968, C. C. Porter (MCZ); 1 © and 2 &,

Tartagal, Nov. 1971, 1 9, Pocitos, Dec. 1971 (MF); 4 9, Angastaco, 7 Dec. 1968,

Entomofauna Subandina, A. Willink-Stange (IML). Santiago del Estero: 1 9,

Suncho Corral, 25 Dec. 1975, R. M. Bohart (UCD). Tucuman: San Pedro,

Colalao, 2 g, Foerster (MF); 2 ©, 8—9 Nov. 1969, L. A. Stange (UCD); 4 ©,

Quebrada Lules, 9 Dec. 1964, C. C. Porter (MCZ). Argentina: | ¢ (USNM).

VAN LITH: New World Pluto 233

First records from Peru: | ¢, 43 mi east Tingo Maria, 1200 m, 18 Nov. 1954, E. I.

Schlinger and E. S. Ross (CAS); 1 9, Madre de Dios Dep., Avispas, 400 m, 12—20

Sept. 1962, L. E. Pefia (CNC); 1 9, Madre de Dios dep., Avispas, 30 m, nr.

Marcapata, 1—15 Oct. 1962, Luis Pefia (HT). The face of the latter female is

distinctly golden pubescent.

First record from Bolivia: 1 9, Luis Calvo, Tiguipa, Jan. 1972 (MF). Face

golden.

The group of pygmaeus has very interesting axillae which are not connected

posteriorly with the scutellum. I have not seen this character in any other Psenini.

The females have the usual characteristic long stiff bristle on the hind coxae.

Pluto pygmaeus axillaris subsp. nov.

(fig. 66)

This form closely resembles the nominate subspecies but differs in the following

characters.

Female. — Vertex and back of head with bluish reflection. Median part of

clypeal margin very weakly emarginate, almost straight, with indistinct lateral

angles or teeth, distance between these teeth about half total width of clypeal

margin. Depressed margin of clypeus shining, behind this margin a weak, shining

ridge, disk of clypeus densely punctate. Frons and vertex shining, almost

impunctate, median part of frons dull, finely punctate; near oculi a large shining

tubercle; back of head smooth and shining.

Scutum and scutellum shining, sparsely finely punctate, interstices on scutellum

a few times size of punctures. Mesopleura and mesosternum finely reticulate

alutaceous, almost smooth, sparsely finely punctate. Hypo-epimeral area shining,

sparsely finely punctate. Anterior plate of mesepisternum almost shining. Second

recurrent vein of fore wings ending in third submarginal cell.

Male. — Head and scutum with bluish shine. Mandibles yellowish. Pronotal

tubercles brownish. Fore and mid tibiae, base of hind tibiae and all tarsi yellowish-

brown, bases of segments 2—5 of hind tarsi brown, often somewhat darker. Axillae

brown.

Antennal segments 6—13 with long linear tyloides, short but rarely absent on

last segment. Axillae with parallel sides and rounded apex, about 1.5 times as long

as broad. Genitalia: fig. 66.

Colombia: 1 9, holotype, Meta Cord., Macarena, 15—28 Febr. 1976, M.

Cooper (BM).

Bolivia: 18 g, allotype and paratypes, Huachi Beni, September, Mulford Bio

Expl. 1921—22, Wm. M. Mann (USNM); 1 9, paratype, Coroico (ZMB). Further

paratypes:

Brazil: Mato Grosso: | 9, Gallery Forest, 12°50’ south, 51°47’ west, 5 Oct. 1968,

Royal Soc. and Royal Geogr. Soc. Expedition, O. W. Richards (BM); Sinop,

12°31’ south, 55°37’ west, 9 ©, Oct. 1974 and 4 9, Oct. 1975, M. Alvarenga (HT).

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Pará: 1 9, Ipean, Belém, 1—4 Dec. 1969, J. M. and B. A. Campbell (CNC). Sao

Paulo: 1 g, Faz. Campininas, Moni Guacu, 1—8 Jan. 1970, J. M. and B. A.

Campbell (CNC). Minas Gerais: 1 9, Pedra Azul, Nov. 1970, F. M. Oliveira (HT).

Distrito Federal: 2 9, Estacao Florestal, Cabeca do Veado, 3600 m, 14—30 Oct.

1971, E. G., I. and E. A. Munroe (CNC).

British Guyana: 1 9, Essequibo River, Moraballi Creek, 3 Sept. 1929, Oxford

University Expedition (BM).

Surinam: 1 9, Republiek, 19 Oct. 1963, D. C. Geijskes; 1 9, Zanderij,

Kreekbos, 10—14 July 1964, Malaise trap, D. C. Geijskes (ML).

One female from Brazil has the pubescence of the face distinctly golden: Rio de

Janeiro, Baia de Guanabara, Floresta dos Macacos, April 1961, M. Alvarenga

(KU). In the female from Bolivia the face is pale golden.

The differences between axillaris and the nominate subspecies are very small.

The clypeal margin of the females seems to be slightly different but this also

depends upon the angle at which the insect is examined.

The subspecific name refers to the axillae which are somewhat longer than in

the nominate subspecies, at least in the male.

Pluto facialis sp. nov.

(figs. 63—65)

Female. — Length about 5.5 cm. Black. Underside of flagellum orange-brown,

labrum and mandibles reddish. Pronotal tubercles whitish, tegulae brown. Fore

and mid tibiae and tarsi and basal 1/3 of hind tibiae yellowish-brown, mid tibiae

partly darkened, hind tarsi dark brown, apices of gastral segments yellowish-

brown. Pygidial area dark reddish. Veins of wings brown.

Median part of clypeal margin straight, thickened, very deeply emarginate,

exposing labrum (fig. 63). Median part of frons raised, finely punctate. Vertex

shining, sparsely punctate. POD about equal to OOD. Intercarinal space narrow.

Mandibles long. Antennae short, clavate, segments 9—11 shorter than broad,

segment 12 about 1.5 times as long as broad.

Anterior carina of pronotal collar high, laterally protruding as a large tooth.

Scutum and scutellum shining, sparsely finely punctate. Axillae triangular with

rounded apex, almost reaching halfway distance from base of axillae to scutellum.

Scutellar suture not crenulate. Propodeal enclosure shining, with high posterior

carina, large median area and few oblique lateral carinae. Back of propodeum

somewhat shining, very coarsely reticulato-carinate, carinae high. Hypo-epimeral

area, mesosternum and upper half of mesopleura finely reticulate alutaceous,

lower half of mesopleura shining. Mesopleura and hypo-epimeral area sparsely

finely punctate, mesosternum densely finely punctate. Anterior oblique suture

crenulate, anterior plate of mesepisternum dull. Second recurrent vein of fore

wings interstitial. Petiole as long as first tergite, this tergite somewhat longer than

broad. Pygidial area slightly over 1.5 times as long as broad.

Pubescence of face pale golden, partly appressed. Tempora and pronotal collar

VAN LITH: New World Pluto 235

with short, yellowish-silvery pubescence, pygidial area ark brown, rest of body

yellowish-grey pubescent.

Male. — Length about 5 mm. Lower part of clypeus shining, raised (figs. 64, 65)

and protruding over emarginate margin. Antennae somewhat clavate, segments

11—12 longer than broad, segments 5—12 or 6—12 with linear, not very distinct,

tyloidea. Axillae with almost parallel sides, longer than broad, apex rounded.

Mesopleura, hypo-epimeral area and mesosternum smooth and shining. Face pale

golden.

Colombia: 1 9, holotype, | g, allotype, 8 & paratypes, Meta, Rio Duba, 8—12

March 1976, M. Cooper (BM).

P. facialis belongs to the group of P. pygmaeus (Brethes) and is easily recognized

by the deeply emarginate clypeal margin of the female and the raised clypeal disk

of the male.

The specific name refers to the peculiar shape of the clypeus, of the female as

well as of the male.

Group of joergenseni

Pluto joergenseni (Brethes) comb. nov.

(figs. 67—71)

Brèthes, 1913: 120, g (Psen Jörgenseni; Argentina: La Paz).

Bohart & Menke, 1976: 171 (Pseneo joergenseni).

The holotype, which was kindly sent to me by Dr. M. J. Viana, Buenos Aires,

has no head, but the characteristic shape of the metanotum does not leave any

doubt as to its identity. Bréthes recognized this peculiar shape of the metanotum

in his description as follows: “postscutello paulum elevato, postice christa in

media incisa acuta”. His reference to the pygidial area is confusing.

Male. — Length 5.5—7 mm. Head and thorax black. Mandibles dark reddish.

Underside of flagellum yellowish-red. Pronotal tubercles yellowish-white or

reddish-brown. Fore tibiae and tarsi yellowish-red, mid tibiae and tarsal segments

2—4 yellowish-red, basitarsus yellowish-white, last segment brown. Basal third of

hind tibiae and entire basitarsus yellowish-white, tarsal segments 2—4 pale brown

or yellowish-brown, last segment dark brown. Veins of wings brown. Gaster black,

hind margin of first tergite red, second tergite usually entirely red, sometimes also

base of tergite 3; or gaster black with reddish transparent hind margins.

Anterior margin of clypeus emarginate, distinct lateral teeth, distance between

these teeth about 1/3 of total width of margin. Labrum with irregular margin and

four distinct median teeth (fig. 69). Frons, except for lateral parts, dull, densely

punctate. Interocellar area and vertex reticulate alutaceous, sparsely punctate.

Occipital carina complete, below in the middle receding backward. Intercarinal

space broader than first basitarsus. Antennae slightly widening toward apex, not

distinctly clavate, segments 6—11 with narrow tyloides.

Lateral pronotal angles in frontal view with small, obliquely upward directed

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

tooth. Scutum and scutellum reticulate alutaceous, sparsely punctate. Metanotum

(figs. 70—71) in the middle with two small raised wings, their margins mem-

braneous, somewhat yellowish transparent. Propodeal enclosure shining, with high

oblique carinae and large median area, back of propodeum dull, coarsely

reticulato-carinate. Hypo-epimeral area rugose. Mesopleura and anterior plate of

mesepisternum finely reticulate alutaceous, irregularly and rather coarsely

punctate, punctures medially partly in rows, interstices on upper and lower part a

few times size of punctures. Anterior oblique suture crenulate. Petiole longer than

first tergite, this tergite about 1.5 times as long as broad. Second recurrent vein of

fore wings ending in third submarginal cell.

Face and pronotal collar silvery, mostly appressed, pubescent; pubescence of

rest of body whitish. |

Female. — Length about 7—8 mm. Colour mostly as in male, outer side of fore

and mid tibiae yellow, hind tarsal segments 1—4 yellowish-white. Microsculpture

as in male but punctation of mesopleura finer. Clypeal margin widely emarginate,

laterally with a large tooth, distance between teeth over half total width of clypeal

margin (fig. 67). Labrum different from that of other Pluto, margin having many

small teeth (fig. 68).

Median part of metanotum somewhat raised, margined by a sharp lateral carina,

no ““wings’’. Hypo-epimeral area dull, superficially punctate. Petiole about as long

as first tergite, this tergite about 1.5 times as long as broad. Pygidial area over 1.5

times as long as broad. Pubescence of face silvery, of pygidial area golden-brown.

Material examined: | &, holotype, with labels “Mendoza”, “No. 10405” and

“Psen Jörgenseni Br.”, coll. P. Jörgensen (MACN). Although the locality-label

only mentions Mendoza, it is evident from the original description (1913) that this

specimen has been collected in the small village La Paz (province of Mendoza, in

Argentina; information by letter from Mr. M. A. Fritz).

New records from Argentina: Catamarca: | &, San Antonio, 6—18 Febr. 1958,

R. Golbach (IML); 1 g, Tinogasta, 8 Nov. 1966, L. Stange (IML); 1 9, 2 km west

Cordobita, 30 Nov. 1968, Entomofauna Subandina, Stange - A. Teran (IML); 3 9

and 1 4, Belén, 25 Nov. 1975, R. M. Bohart (UCD). Cordoba: Cordoba, 1 9, Dec.

1940, Christensen (MF), 1 ©, Jan. 1947 (IML), 3 &, Febr. 1971, M. Fritz (MF);

Balnearia, 3 9, Fritz-Martinez (MF). La Plata: 1 g, Punta Lara, 13 Jan. 1970,

Malaise trap, Vardy and Arguindeguy (BM). La Rioja: 1 G', B. P. Clark (USNM); 1

©, Anguinan (Chilecito), 1—15 Febr. 1970, Entomofauna Subandina, Willink-

Terän-Stange, Malaise trap (IML); 1 g, Villa Mazan, 19 Dec. 1975, R. M. Bohart

(UCD). Mendoza: 1 9, Est. Pedregal, A. C. Jensen-Haarup (ZMC); 1 &,

Potrerillos, 4000 ft, 16—20 March 1920, Cornell Univ. Exped., R. G. Harris (CU);

I ©, Potrerillos, 1000 m, 20 Jan. 1947, Hayward-Willink (IML); 1 g, Agrela, 23

Febr. 1966, L. Stange (IML). Misiones: 1 4, Bemberg, Alto Paraná, 21—31 March

1934, K. J. Hayward (BM). Rio Negro: Lamarque, | 9, A. Baier, 7 9 and4 3, U.

and M. Fritz, 1 &, Febr. 1958, U. and M. Fritz, 1 g, Jan. 1974, M. Fritz (MF); 1 &,

Choele-Choel, Dto. Avellaneda, 24 Nov. 1946, Hayward-Willink; 1 9 and 2 &,

Isla Choele-Choel 14 Jan. 1968, J. and L. Stange (IML). Salta: 3 9 and 2 4,

Tartagal, Nov. 1971, M. Fritz (MF); 1 g, San Rafael (San Carlos), 7 Dec. 1968,

VAN LITH: New World Pluto 237

Entomofauna Subandina, Stange-A. Willink (IML); 1 &, Camp. Jakülica, Ca.

Aguas Blancas, Oct. 1968, 1 &, Oran, Abra Grande, 18—25 Oct. 1968, C. Porter

(MCZ). San Juan: 6 9, Pie de Palo, 11 March 1920, Cornell Univ. Exped., R. G.

Harris (CU). Santa Fé: 1 9, Alberdi, 685, 17 March 1912, J. Hubrich (BSM).

Santiago del Estero: 1 &, Fernandez, Jan. 1939, coll. Bosq (MF). Tucuman: 1 9,

Tucuman, Jan. 1947, J. Cordoba (IML).

First records from Bolivia: Luis Calvo: 5 9 and 12 Z, Tiguipa, Jan. 1972, M.

Fritz; Cordillera, 1 9, Rio Grande, 16 Febr. 1971, Fritz-Martinez, 1 9, Abapó,

Jan. 1972, M. Fritz (MF).

First records from Brazil: Mato Grosso: 1 © and 3 Z, Aguidauana, 11—13 Dec.

1919, Cornell Univ. Exped., R. G. Harris (CU). Minas Gerais: 1 9 and 2 5,

Pirapora, 11—13 Nov. 1919, Cornell Univ. Exped. (CU); 1 9, Santa Vitoria, Febr.

1970, F. H. Oliveira (AMNH). Sao Paulo: 1 9, Aguas Sao Pedro, 29 Jan. 1976, R.

M. Bohart (UCD); 1 ©, Municipio de Bertioga, 27—30 Nov. 1972, B.V. Peterson

(CNC).

First record from Paraguay: | g, Pirareta, Dec. 1971, L. E. Pena (MF).

The widely emarginate clypeus of the female and the remarkable structure of

the metanotum of the male are very characteristic.

REFERENCES

Alayo, D. P., 1973. Catalogo de los Himenopteros de Cuba: 1—218.

Ashmead, W. H., 1899. Classification of the entomophilous wasps, or the superfamily Sphegoidea. —

Can. Ent. 31 (8) : 212—225.

—, 1900. Report upon the Aculeate Hymenoptera of the Islands of St. Vincent and Grenada, with

additions to the parasitic Hymenoptera and a list of the described Hymenoptera of the West

Indies. — Trans. ent. Soc. London, part 2: 207—305.

Bohart, R. M. & A. S. Menke, 1976. Sphecid wasps of the world: I-X, 1—695. University of California

Press, Berkeley, Los Angeles, London.

Brethes, J., 1913. Himenöpteros de la América Meridional. — An. Mus. nac. Hist. nat. Buenos Aires

24: 35— 16S.

Cameron, P., 1891. Insecta. Hymenoptera (Fossores), fam. Mimesidae. — Biologia Centr.-Amer. 2:

129— 176.

Cockerell, T. D. A., 1911. A new Psenid wasp from Peru. — Can. Ent. 43: 272.

Cresson, E. T., 1865. On the Hymenoptera of Cuba. — Proc. ent. Soc. Philad. 4: 1—200.

, 1872. Hymenoptera Texana. — Trans. Am. ent. Soc. 4: 153— 227.

, 1916. The Cresson types of Hymenoptera. — Mem. Am. ent. Soc. 1: 1141.

Dalla Torre, C. G. de, 1897. Fossores (Sphegidae). — Catalogus Hymenopterorum hucusque descrip-

torum systematicus et synonymicus 8: 1— 749.

Evans, H. E., 1959. Studies on the larvae of digger wasps (Hymenoptera, Sphecidae) part V: con-

clusion. — Trans. Am. Ent. Soc. 85: 137—191.

——, 1968. Notes on some digger wasps that prey upon leafhoppers. — Ann. ent. Soc. Amer. 61:

1343— 1344.

Fox, W. J., 1898a. Contributions to a knowledge of the Hymenoptera of Brazil. No. 3. Sphegidae (sens.

lat.). — Proc. Acad. nat. Sci. Philad. 1897: 373— 370.

—, 1898b. The species of Psen inhabiting America North of Mexico. — Trans. Am. ent. Soc. 25:

1—18.

Gittins, A. R., 1969. Revision of the nearctic Psenini (Hymenoptera: Sphecidae). I. Redescriptions and

keys to the genera and subgenera. — Trans. Am. ent. Soc. 95: 49—76.

Kohl, F. F., 1896. Die Gattungen der Sphegiden. — Annin naturh. Mus. Wien 11 (3—4): 233—516, pls.

5—11.

238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 6, 1979

Krombein, K. V., 1949. An annotated list of wasps from Nags Head and the Kill Devil Hills (Hymenop-

tera Aculeata). — J. Elisha Mitchell scient. Soc. 65 (2): 262—272.

— _, 1951. In: C. W. F. Muesebeck, K.V. Krombein and H. K. Townes. Hymenoptera of America

North of Mexico. — U.S. Dep. Agric., Agric. Monograph 2: 1—1420.

, 1953. Kill Devil Hill wasps, 1952. — Proc. ent. Soc. Wash. 55 (3): 113—135.

1954. A list of wasps collected in Florida, March 29 to April 5, 1953, with biological annota-

tions (Hymenoptera, Aculeata). — Proc. ent. Soc. Wash. 56 (5): 225—236.

——, 1958. Hymenoptera of America North of Mexico. — U.S. Dep. Agric., Agric. Monograph 2

(first supplement): 1—305.

—— ., 1964. Results of the Archbold Expeditions. No. 87. Biological notes on some Floridian wasps

(Hymenoptera, Aculeata). — Amer. Mus. Novitates 2201: 1—27.

—, 1967. In.: K. V. Krombein & B. D. Burks. Hymenoptera of America North of Mexico. — U.S.

Dep. Agric., Agric. Monograph 2 (second supplement): 1—584.

Lith, J. P. van, 1976. Pluto rufibasis (Malloch) (Hymenoptera, Sphecidae, Psenini). — Ent. Ber. Amst.

36: 154—158.

Malloch, J. R., 1933. Review of the wasps of the subfamily Pseninae of North America (Hymenoptera:

Aculeata). — Proc. U.S. natn. Mus. 82: 1—60, pls. 1—2.

Mickel, Clarence E., 1918. A synopsis of the Sphecoidea of Nebraska (Hymenoptera). — Univ. Nebr.

Stud. (1917) 17: 342—456.

Pate, V. S. L., 1937. The generic names of the Sphecoid wasps and their type species nee

Aculeata). — Mem. Amer. ent. Soc. 9: 1—103.

—, 1946. On the Psenine wasps of Cuba (Hymenoptera: Sphecidae). — Mems. Soc. Cubana Hist.

nat. 18 (1): 3—10.

Persson, P. I., 1971. “Eugenies resa”. Localities, dates and labels of the insects collected during the

voyage around the world by the Swedish frigate “Eugenie” in the years 1851—1853. — Ent.

Tidskr. 92: 164—172.

Rohwer, S. A., 1910. Descriptions of new Psenid wasps from the United States (Hymenoptera; Pseni-

dae). — Proc. ent. Soc. Wash. 12: 99—104.

Smith, F., 1856. Catalogue of Hymenopterous insects in the collection of the British Museum, Part IV.

Sphegidae, Larridae and Crabronidae: 207—497.

Smith, H. S., 1908. The Sphegoidea of Nebraska. — Univ. Nebr. Stud. 8: 323—410.

Stephens, J. F., 1829. A systematic catalogue of British insects: XXXIV, 1—416, 1—388. Baldwin &

Cradock, London.

Viereck, H. L., 1901. New species of the subfamily Pseninae. — Trans. Amer. ent. Soc. 27: 338—342.

’

VAN LITH: New World Pluto

INDEX

The names of new species and subspecies are marked with an asterisk

* abbreviatus 182

aerofacies 183

albifacies 187

* alphitopus 195

angulicornis 157

annulipes 224

* araguensis 221

arenivagus 190

argentifrons 192

atricornis 194

* axillaris 233

* basifuscus 160

* biformis 169

brevipetiolatus 164

* castaneipes 197

clavicornis 206

* colonensis 198

* cubanus 192

* denticollis 229

* depressus 181

Diodontus 127 sqq

* duckei 212

* emarginatus 185

* evansi 184

* facialis 234

* fritzi 199

Gorytes 128 sqq

* incarinatus 222

joergenseni 235

* jugularis 178

littoralis 171

longiventris 154

marginatus 186

* marthae 208

medius 200

* menkei 216

* metanus 210

Mimesa 127 sqq

minutus 162

Neofoxia 127 sqq

* nitens 210

* obscurus 214

* occipitalis 217

Ochleroptera 230

pallidistigma 158

Pluto 129 sqq

Psen 127 sqq

Pseneo 235

Psenia 127 sqq

Psenulus 127 sqq

* punctatellus 179

pygmaeus 230

* rotundus 165

* rufanalis 220

rufibasis 186

* rugulosus 197

sayi 173

* scytinus 203

* simplicicollis 215

smithii 226

* spangleri 162

* spinicollis 217

* stenopygidialis 176

* stramineipes 204

* strigellus 205

suffusus 166

texanus 170

tibialis 188

townsendi 207

* trilobatus 218

* zonatus 215

* zuliensis 202

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A REVISION OF THE SUBFAMILY ZELINAE AUCT.

(HYMENOPTERA, BRACONIDAE)

C. VAN ACHTERBERG

Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands

With 900 text-figures

ABSTRACT

The subfamily Zelinae, as defined by Mason (1973) and Van Achterberg (1976b), is revised. The

genera Zele, Homolobus and Charmon are redefined, the genus Zele is separated from the Zelinae auct.

and is added to the Euphorinae-Meteorini as a senior synonym of Zemiotes. The remaining group of

genera is renamed Homolobinae, including two tribes, Homolobini tribus nov. and Charmontini tribus

nov. In the Homolobinae two new genera from the New World, Exasticolus and Charmontia, are

described. For the first time the genera Zele, Homolobus and Charmon are fully revised, keyed and

illustrated and a subgeneric division of Homolobus is proposed. Of the total of 61 valid species, 32 are

newly described, while in addition 32 new combinations and 34 synonyms are proposed.

CONTENTS

MON rg LEE te Ne he. fate Gian, REA OI SIE OE AME 242

LEGO oen ee ne Seen 242

SERIEN à 4 0 RO EA i LE AU 246

Ehylopenvatrn ts. den Deore En et ee data Se ie 250

LUNE) ce oren WA N En ARS SIOE LITIO ZONE 256

Key to the genera and subgenera of Homolobinae and Meteorini .................... 260

Saas oniolobinae NOM Move posh werner terr ee denn es En 261

Tics MOM ONIN OVE wt RUE NE 262

EENS OO NON EE 262

cis moos EIEN edad ai Rn en Neen WEN 263

RESTO MO ODINO I REN ERTL OMEIN a GE DEE A OAD 270

PRAMS ASIICOIUSINONA cazzoni AIS ler. A RE Maier Babe 271

esi onol0 bus Hoerstens ne re elle nie was eert LEGEN Man: 276

Bro enUstApattENd eil ener RARO ACI Mii 277

LOES CDO PONNE BEENS EEE Rn us 304

BE USM 0710 lo DUSIKOErStE AAE TE Ne N ede 311

SubpenusRhylaeterlReinharde Webs u.a SANS EET dE, Tat ig AN N AT tone 321

BEES OLO USN ONE Ro TL A ee 327

Key to the Palaearctic species of the genus Homolobus ........................... 350

BiexitotheNearetic species of the genus Homolobus. ‚ns en ene en 352

Key to the Neotropical species of the genus Homolobus ........................... 353

Key to the Afrotropical species of the genus Homolobus ........................... 354

Key to the Oriental and Australian species of the genus Homolobus .................... 356

EE uphorinaetRoerstere: . ns sanne modele nnn eee Gwe à à _ 358

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Tribus Meteorini:Cresson 130% onde ee EINEM 359

Genus Zele: Curtis’ eg vele Lela RR ee PI 359

Excluded! species ti. iI 383

Acknowledgements, 5e st EN 386

Literature. es ur NEE EE 387

Index of names used in the genera Charmon, Exasticolus, Homolobus, Zele and their synonyms .. 390

INTRODUCTION

When, in 1975, I started upon a revision of the Macrocentrinae (except for

Macrocentrus s.s.), I soon discerned the need for a revision of the Zelinae auctorum

because of the mixing of both groups and the lack of modern keys, even of so small

and common a genus as Charmon Haliday. An unexpectedly large number of

changes proved to be necessary to bring the taxonomy up-to-date. A total of 34

new synonyms and 32 new combinations are proposed. Some of the most

important changes are the synonymy of Zele Curtis with Zemiotes Foerster, its

removal from the Zelinae auctorum, its inclusion in the Meteorini of the

Euphorinae and the renaming of the Zelinae auct. as Homolobinae. The large

number of undescribed species in the genus Homolobus was unexpected, because

they are comparatively large insects. Their nocturnal habits may be one of the

main reasons of their rarity in collections, because hymenopterists usually do not

collect at light.

The number of valid species of Homolobus is in this paper increased from 18 to

43 species, while I have seen additional new species which have to remain

undescribed because of the lack of well prepared specimens. The number of new

species in the other genera is lower and the increase in the number of valid species

varies from 50—57%, while in the genus Zele Curtis a large number of new

synonyms are proposed. The total number of valid species in the groups treated in

this paper is increased from 29 to 61, or more than doubled.

I have illustrated all species in a comparative way to enhance the chance of an

unambiguous identification by workers without access to a reference-collection.

Any student has to be aware of possible artificial differences as a result of

illustrating from slightly different angles (as with, e.g., the frontal aspect of the

head), due to the different ways in which the specimens have been mounted.

Of interest to the biological control of pests is the large number of parasites of

pest species of Lepidoptera included in this revision, but more research has to be

conducted before a start can be made with their application. Because Shenefelt

(1965, 1969, 1970) has given excellent lists of the pertinent literature, I refer only to

the original publications, to Shenefelt’s catalogue and to the (usually) more recent

literature not included by Shenefelt. To facilitate identification per region, keys to

species of Homolobus per region were inserted after the descriptions.

TERMINOLOGY

The morphological terms used (figs. 1—18) largely follow Richards (1956, 1977)

and for the wing venation I have used a modified Comstock-Needham system

VAN ACHTERBERG: Revision Zelinae auct. 243

Figs. I—2, mesosoma of Blacus (Ganychorus) pallipes Haliday, 9, Netherlands, Wijster, legs and wings

removed. |, lateral aspect; 2, dorsal aspect. A = side of scutellum and axilla; B = precoxal suture; C =

episternal scrobe; D = pleural suture; E = epicnemial area; F = metapleural flange; Gl = anterior

part of metapleuron; G2 = posterior part of metapleuron; H = propodeum; I = metanotum; J = pro-

podeal spiracle; K = prepectal carina; L = lateral lobe of mesoscutum; M = middle lobe of mesoscu-

tum; N = notauli; O = mesopleuron; P = pronotum; Q = pronope; R = lateral carina of scutellum; S

= scutellum; TI + T2 = tegula and humeral plate, respectively; U = propleuron; V = mesosternum;

W = medial carina of propodeum; X = lateral carina of propodeum; Y = pleural carina; Z = scutellar

suture. 75 x

244. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

based on the proposals of Eady (1974) and Sigwalt (1977). For a comparison with

the modified Jurinean system I used before, see table 1. The following modifica-

tions of the Comstock-Needham system (as applied to the Braconidae by Eady

(1974)) are proposed:

i) the abbreviations of the longitudinal veins are completely written in capitals

(“CU” in stead of ‘“Cu”’), additional letters to indicate a certain part are in lower

case letters (e.g., CUla). The abbreviations of the true transverse veins are (as in

the original Comstock-Needham system) wholly in lower case letters.

ii) the abbreviations are derived from Latin names of the veins. Thus SR (from

“sectio radii’’) and not Rs (from “radial sector”) of Eady and others.

iii) the prefix ‘‘1-”, “2-”, “3-”, etc. is added to indicate the Ist, 2nd, 3rd, etc.

abscissa of a vein. This should not be confused with a cipher without minus-sign in

front of an abbreviation, this indicates which vein is involved. E.g., 2r-m is the

second transverse vein between the radius and the media, while 2-SR indicates the

second abscissa of the longitudinal vein SR. For the application of this system to

the genera revised in this paper, see figs. 15—18. Some abbreviations are extra

short for practical reasons: r-m of fore wing is actually 2r-m+SR2, except, with

certainty, in the Neoneurinae where SR2 branches off distad from 2r-m; an

apomorphous state as shown, e.g., by the hypothetical ancestral hymenopterous

wing proposed by Ross (1937). Normally Ir-m is absent in the fore wing of the

Braconidae, because it disappeared with the anastomosis of SR with M. But in,

e.g., the freak-type of fore wing of Alysia ridibunda (Say) (fig. 2 in Riegel, 1948) Ir-

m is still visible. This vein was wrongly interpreted by Riegel as 2r-m and resulted

in his anomalous nomenclature of r-m as 3r-m. The real transverse vein 3r-m is

absent in the Braconidae and its sister-group Ichneumonidae. Transverse vein r of

fore wing is actually 2r but lr is normally absent in the fore wing of the

Braconidae, while cu-a of hind wing is actually 1-CU +cu-a, but if it is no longer

recognizable (as in most subfamilies of Braconidae) it is abbreviated as cu-a.

For the following terms used, an additional explanation may be useful:

Antescutal depression: transverse depression between dorso-anterior part of

pronotum and middle lobe of mesoscutum (figs. 168, 316).

Diplope: distinct laterope and dorsope more or less touching each other (fig.

843).

Dorsope: antero-dorsal depression of the Ist metasomal tergite, more or less pit-

Figs. 3—8, Blacus (Ganychorus) pallipes Haliday, same specimen as in fig. 1. 3, frontal aspect of head; 4.

dorsal aspect of head; 5, apex of metasoma, lateral aspect; 6, fore tarsus, lateral aspect; 7, lateral aspect

of head; 8, Ist metasomal tergite, dorso-lateral aspect. A = anterior tentorial pit; B = basitarsus; C =

clypeus; CI = epistomal suture; C2 = clypeal margin; D = dorsope; E = eye; F = frons; G = face; H

= hypopygium; I = dorsal carinae; J = laterope; K = glymma; L = labrum; M = mandible; N = fron-

tal suture; O = ovipositor; P = posterior ocellus; Q = anterior ocellus; R = stemmaticum; S = ovipo-

sitor sheath; T = temple; Tl = antennal socket; U = occipital flange; V = vertex, W = occipital cari-

na; X = spiracle of Ist metasomal tergite; Y = tarsal claw; Z = telotarsus; AD = anterior muscle or

adductor of lst metasomal tergite; AN = annellus; MS = malar space; PA = postannellus or 3rd an-

tennal segment; PD = pedicellus or 2nd antennal segment; RA = radix; SC = scapus or Ist antennal

segment. 64 x

VAN ACHTERBERG: Revision Zelinae auct. 245

shaped, situated between the more or less developed dorso-lateral carina and the

dorsal carina (figs. 8, 852; Van Achterberg. 1974c: 213).

246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Height: maximum height, unless otherwise stated; for the height ofthe head, see

fig. 10.

Alone transverse vein of which the anterior end is nearer to the wing base

than its posterior end.

Laterope: antero-dorsal depression of the Ist metasomal tergite, more. or less

pit-shaped, situated in the glymma below the more or less developed dorso-lateral

carina (figs. 8, 815).

Length of 3rd antennal segment: for convenience’ sake the length of the

annellus is included.

Length of fore wing: measured from apex of humeral plate to apex of fore wing

(Van Achterberg, 1976a: fig. 15).

Length of Ist metasomal tergite: measured from posterior margin of anterior

muscle to its apex (fig. 14).

Length of malar space: shortest distance between eye and condylus of mandible

(fig. 9, M).

Length of ovipositor sheath: linear length of maximal visible part.

Malar suture: suture (usually shallow) between under edge of eye and base of

mandible.

Mesosoma: thorax of most authors, but because of the fusion of the Ist

abdominal segment (propodeum or epinotum) with the thorax, I prefer the term

“mesosoma”. The mesosoma is the part of the body between the Ist and 2nd

strong constriction.

Metasoma: following Michener (1944), it is defined as the part of the body after

the 2nd strong constriction in the Apocrita. It is usually called the abdomen or

gaster; for a discussion on the confusing nature of these terms in the parasitic

Hymenoptera, see Van Achterberg (1976a: 166).

OOL: distance between posterior ocellus and eye as measured in fig. 12.

Plical lobe or cell: anal (or vannal) lobe/cell of most authors (Brothers, 1975:

520).

Pronope: a medio-dorsal pit of the pronotum (fig. 2).

POL: distance between the posterior ocelli (fig. 11).

Reclivous: transverse vein of which the anterior end is further removed from the

wing base than its posterior end.

Width: maximum width, unless otherwise stated.

DISTRIBUTION

As shown in the tables 2 and 3, the genus Homolobus Foerster (comprising the

subgenera Apatia, Chartolobus, Homolobus, Phylacter, and Oulophus) is cosmo-«

politan, the genus Zele Curtis is widely distributed (but is absent in the Australian

and Afrotropical regions, and scarcely represented in the Oriental region), the new

genus Exasticolus is restricted to the New World, the new genus Charmontia is only

known from Chile, and the genus Charmon Haliday occurs in the Holarctic,

Afrotropical, Oriental, and Australian regions. According to Mason (1973: 215)

the genus Zele Curtis (Zemiotes in his paper) is restricted to the Holarctic region,

but as reported in this paper, the genus Zele is also present in the Neotropical and

247

VAN ACHTERBERG: Revision Zelinae auct.

= d ‘duly 1e][950-1e]n90 = 114

4 {snjj290 Jo119]SOd Jo Joyaweıp = q ‘snadjo Jo 1YyBIey = I ‘Joadse [esıop ‘a}1819} [ewose]aw JS] ‘pl ‘Joad

= H {998J Jo ıy3ıay =

-Se [81978] BWOsosau

‘

les

Sn M

<— d ——|-q

YIPIM = M Saul] Ie][2907s0d

O ‘aoueisip Ie[n90-o1IOJus} = N !a9eds IE|ew = W ‘yIZua] = T ‘99UEISIP [ELIO)U9]-193UI = | ysıay

yoodse jesiop ‘peay ‘ZI ‘wnorewwars ‘TI ‘joodse [eıoye] ‘peau ‘OI ‘399dse [ejuo1] ‘peau ‘6 716 ‘SSI

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= [| ‘189 jeseq = 9 ‘[[99 [eIso9 = ¢ ‘[[99 [eosipgns = p ‘]]99 [eosIip = € {]]99 Jeurdsewans = 7 ‘1199 jeurdrew = | :s][99 "ewänso1sjd

= jd ‘ewänseied = ed ‘uI9A [BIPaWI-O1pe1 asIOASULI] = W-I SUIDA [BIPEI ASIDASULI] = 1 ‘UI9A [BJIQNO-OIPAU asJ9ASUPI] = NI-W ;UIAA

[eue-oyqno ISIOASULI] = B-ND !UIDA [EUR ASI9ASULI] = B !IIPEI 011995 = YS :BIsOOqNs = DS ‘snipes = Y !elpaw = JW ‘snyiqnd = NI

‘ISO = J ‘sleuR = W ‘W9]SAS WRYPIdN-YOISWOD PATJIPOUI ay} 0} Zurp1099e UONEUSA BUIM dU} JO aınyejsuswou 'gI—S] "SLA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

: x e; =

ge va...

vz+tVvi Nes

INS

IND+W IND-1

248

H+9S+9 G

VAN ACHTERBERG: Revision Zelinae auct.

249

Table 1. Comparison of modified Jurinean system and modified Comstock-Needham system (figs.

15—18).

Modified Jurinean System

Modified Comstock-Needham System

Fore wing

costa

media

submedia

basalis

nervulus

vein for attachment of hamuli

Ist transverse anal vein

2nd transverse anal vein

Ist abscissa of discoideus

2nd abscissa of discoideus

basal abscissa of subdiscoideus

apical abscissa of subdiscoideus

parastigma

pterostigma

metacarpus

Ist abscissa of radius (rl)

2nd abscissa of radius (r2)

3rd abscissa of radius (13)

Ist transverse cubital vein

2nd transverse cubital vein

Ist abscissa of cubitus (cul)

2nd abscissa of cubitus (cu2)

3rd abscissa of cubitus (cu3)

4th cubitus (cu4)

abscissa of

radial cell

Ist, 2nd, 3rd cubital cells

Ist, 2nd discoidal cells

Ist, 2nd brachial cells

costal cell

medial cell

submedial cell

anal cell

Hind wing Abbreviations Fore wing Hind wing

costella/subcostella c C+SC+R C+SC+R/C/SC+R

mediella m M+CU1 M+CU/M

submediella sm 1A+2A/1A 1A

basella b 1-M Ir-m

nervellus nv cu-a cu-a

=== - 2A+3A mw

Ist transverse anellan vein aqu | 2A 2A

== aqu 2 a ==

nz d 1 1-CUI ==

=== de 2 2-CUI =

(sub)discoidella sla 3-CUl 2-CU

Sas Ss) 2 CUla 3-CU

=> s lb CUIb ==

n P pa ==

pterostigma (normally absent) pt pt pt

metacarpella mc : RI RI

transverse radiellan vein| rqu r r

2 3-SR

radiella r SRI { SRI

transverse cubitellan vein cuqu | 2-SR 2r-m

=== cuqu 2 r-m ==

1-SR+M

6 2-SR+M

cubitella cu 2-M 2-M

3-M

postnervellus pn == m-cu

radiellan cell R marginal cell marginal cell (1)

cubitellan cell CU Ist, 2nd, 3rd submarginal|cells submarginal cell (2)

discoidellan cell D Ist, 2nd discal cells discal cell (3)

= B Ist, 2nd subdiscal cells subdiscal cell (4)

costellan cell C costal cell costal cell (5)

mediellan cell M basal cell basal cell (6)

submediellan cell SM subbasal cell subbasal cell (7)

(v)anellan cell or lobe A plical cell plical cell or (8)

lobe

Oriental regions. The genus Zele is (as shown by the character-states of the

species) of Holarctic origin and its largest speciation has taken place there.

The situation in Homolobus is more complicated. There are two subgenera,

Phylacter Reinhard and Homolobus Foerster, which have restricted distributions.

Phylacter is restricted to the Palaearctic region, with one species in the

intermediate area between the Palaearctic and Oriental regions. The subgenus

Homolobus is restricted to the Palaearctic and Afrotropical regions, and,

considering the distribution of the apomorphous character-states among the

species, it has also a Palaearctic origin.

The subgenus Apatia Enderlein has its centre of speciation in the Afrotropical

region, where also the species with the largest number of plesiomorphous

character-states occur. The only species outside the Afrotropical region with a

peculiar plesiomorphous character-state (viz., the presence of vein r in the hind

wing) is H. (A.) elagabalus (Nixon) from the Oriental region. Three other species of

Apatia show a remarkably wide distribution. This distribution may have resulted

(partly) from human activities, but this seems unlikely, because, e.g., a species (H.

(A.) australiensis) may have evolved in Australia from H. (A.) ophioninus (Vachal),

itself probably originating from Africa. Another curious distribution is shown by a

species of the new subgenus Chartolobus; H. (C.) infumator (Lyle) occurs in the

Holarctic region, but has also reached the Neotropical (viz., the Andes) and the

Oriental regions. Or, if considered to originate from the Oriental regioa, it has

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Table 2, Number of revised species occurring in only one zoogeographical region.

% %

È à © si à

(Sub)genus = S a È a è

9 me rQ + à D e

‘A S) 9 DI O <= S S

+ "A + md 3 or È

. eN + iy © > S N È

Baal ie nn nn

Si x S mS À S 5 N =

Palaearctic (Himalayan = = = 3 3 5 = 3 n

area included)

Nearctic - - - - - 4 3 -

Neotropical 2 - - - - 4 - 2 l

Afrotropical - 9 - 4 = = È = =

Oriental (Himalayan =

area excluded)

- _ = 1 =>

Australian - ] 1

dispersed to the Holarctic and Neotropical regions. The origin of Chartolobus may

be in the Oriental region, but this is uncertain with the information available at

present. The new subgenus Oulophus is a relatively large group of species,

including species with comparatively large number of plesiomorphous character-

states, which occur in the South Nearctic and East Palaearctic areas. This may

reflect the original Holarctic primary speciation of this group. Unfortunately,

there are no fossil remains known of the Homolobinae to test the suggestions put

forward in this papier. The new genus Exasticolus contains two sparsely collected

Neotropical species and one widely distributed and rather common species, which

has penetrated the Nearctic as far as Canada from the Neotropical region.

The origin of Charmon Haliday is uncertain, but it may originate from the

Palaeotropics, from where it occupied the Palaearctic and, subsequently, the

Nearctic regions. Some support for this hypothesis lies in the absence of Charmon

in the Neotropical region and its presence in New Guinea. The new species from

New Guinea shows a plesiomorphous condition of the wing venation, if compared

with both other species. Interesting is the presence of the closely related new

genus Charmontia in Chile with a larger number of plesiomorphous character-

statesthan Charmon.

PHYLOGENY

In constructing a phylogenetic classification it is necessary to find

synapomorphous character-states. The terms apomorphous and plesiomorphous

character-states are here used to indicate, respectively, a comparatively high or

low degree of divergence from an ancestral state in respect to each other. The first

object has to be the defining of monophyletic groups and their sister-groups by

synapomorphous character-states. The interpretation of the relative apo- and

VAN ACHTERBERG: Revision Zelinae auct. 251

Table 3. Number of revised species occurring in more than one zoogeographical region.

È 9

S à

9 S Q

+) à 8 IS È S

(Sub)genus IS aS ES NR È

sii

ined region È = È N

pom Ê B = LS) © S N

Holarctic = = = 1 2

New World | = = I = =

Holarctic, Neotropical & Oriental = l 1 = = -

Holarctic & Afrotropical = = = = 2 =

Afrotropical, Palaearctic & Australian = | = = = =

Indo-Australian = = il =

Holarctic & Oriental = a = =

Palaearctic, Afrotropical & Oriental = 1 = =

+) The subgenera not mentioned are restricted to one zoogeographical region.

plesiomorphous character-states is based on the general hypotheses of the

evolution in the Hymenoptera (as compiled by Königsmann, 1976—1978) and in

the Braconidae (Van Achterberg, 1976b).

The synapomorphous character-states of the sister-groups Homolobinae and

Orgilinae combined (group E, Van Achterberg, 1976b: 51) are: 1—vein a of fore

wing absent; 2—vein m-cu of fore wing far antefurcal; 3—labial sclerite of larvae

transverse; 4—epistomal arch and hypostoma of larvae absent; 5—endoparasites

of larvae of Lepidoptera; 6—tendency to loose the dorsal carinae of the Ist

tergites.

The synapomorphous character-states of the Homolobinae (Homolobini and

Charmontini combined) are: I—antescutal depression present; a character-state

found almost exclusively in the Homolobinae as defined in this papier but see note

below about the Agathidinae; 2—Ist tergite more or less narrowed behind the

spiracles, a tendency also present in the Orgilinae (e.g., genus Microtypus); 3—|st

discal cell of fore wing (sub)sessile and vein 1-SR absent or nearly so, but shortly

developed in the genus Charmontia (fig. 892); 4—metapleural flange more or less

lamelliform and transparent; 5—prepectal carina (almost always) reaches the

anterior margin of the mesopleuron.

The apomorphous character-states of the tribe Homolobini, with regard to its

sister-group, the tribe Charmontini, are: 1—lateral carina of mesoscutum

lamelliform; the plesiomorphous condition in the Ichneumonoidea is probably a

weakly developed, non-lamelliform lateral carina; 2—apical segment of antenna

with a well developed spine; 3—vein 2A of hind wing absent; 4— mandible twisted

apically; 5—vein 2-R1 of fore wing absent.

252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

The apomorphous character-states of the tribe Charmontini, with regard to its

sister-tribe, are: 1—vein r-m of fore wing absent; 2—occipital carina reduced

medio-dorsally; 3—claws simple, without a subapical tooth; 4—precoxal suture

absent; as pointed out by Konigsmann (1977: 3) the presence of the precoxal

suture has to be considered the plesiomorphous condition in the Hymenoptera;

5—middle lobe of mesoscutum more or less truncate anteriorly and with a

transverse protruding horizontal part (fig. 60); this resembles the development in

the Macrocentrinae, but differences in other characters (e.g., prepectal carina,

trochantelli, pronope) indicate that this is very likely a convergent development;

6—marginal cell of hind wing narrowed distally; as shown, e.g., by the hind wing of

the saw-fly Macroxyela ferruginea (Say), the plesiomorphous character-state is a

medially widened marginal cell; the hind wing of M. ferruginea (Say) is one of the

most completely venated hind wings known in the Hymenoptera; 7—absence of

the lateral carina of the mesoscutum in front of the tegulae; in the

Ichneumonoidea a non-lamelliform carina probably is the plesiomorphous

condition; 8—apical margin of clypeus with a more or less developed row of

punctures.

The apomorphous character-states of the genus Charmon, with regard to its

sister-group Charmontia, are: 1—third segment of labial palp reduced;

2—scutellum smooth medio-posteriorly. Because the great majority of the species

of the Homolobinae have this area sculptured, it is likely that the reduction of the

sculpture is an apomorphous condition in the Charmontini.

The apomorphous character-states of the genus Charmontia, with regard to its

sister-group, are: 1—claws (except for the apical tooth) straight ventrally; 2— Ist

tergite slender (fig. 896); 3—propodeal spiracle situated submedially in

propodeum.

The apomorphous character-states of the genus Exasticolus, with regard to its

sister-group. Homolobus, are: l—inner aspect of apex of hind tibia with a comb of

bristles; 2—vein 1-SR+M of fore wing curved distad; 3—tarsi with a weakly

developed row of setae ventrally; 4—the selection of Lasiocampidae as hosts, a

group of Lepidoptera not attacked by the other Homolobinae according to the

available data.

The apomorphous character-states of the large diverse genus Homolobus, with

regard to its sister-group, are not well to define with the available set of characters.

There are some tendencies to apomorphous states (as depicted in fig. 19), which

made it possible to divide the genus Homolobus in five subgenera. The isolated

position of Exasticolus as a genus (and not as a subgenus of Homolobus) may be

caused mainly by the isolation of the parental stock of Exasticolus in South

America for a long period, combined with the change to another family of hosts.

In sharp contrast the apomorphous character-states of the genus Zele Curtis are

as follows: 1—Ist tergite petiolate; 2—spiracle of Ist tergite situated submedially;

3—mesopleuron more or less protruding antero-dorsally; 4—mandible with a pair

of (more or less) protruding, thin carinae; S—Ist subdiscal cell of fore wing

narrowly open postero-distally because of the reduction of vein CUIb;

6—mandible of larva without teeth, bare; 7—claws with a large submedial lobe;

VAN ACHTERBERG: Revision Zelinae auct. 253

8—vein m-cu of fore wing more or less antefurcal; 9—lateral carina of

mesoscutum lamelliform; 10—mandible twisted apically; 11—vein a of fore wing

absent; 12—metapleural flange more or less lamelliform.

It is clear, after comparing both lists of apomorphous character-states of Zele

Curtis (= Zemiotes Foerster) and of Homolobus Foerster, respectively, that Zele is

not likely to be a sister-group of Homolobus (= Zele auct.) as proposed by Mason

(1973). The first seven apomorphous character-states mentioned for Zele Curtis

are not shared by Homolobus, but are well matched in at least some of the species

of the genus Meteorus Haliday s.s. I do not hesitate to depart from Mason’s view

and consider the genus Zele Curtis a sister-group of the genus Meteorus Haliday,

both forming the tribe Meteorini Cresson of the subfamily Euphorinae. The

argument put forward by Mason (1973) is based on the faulty premise that he

considered “the possibility of Zemiotes being ancestral to Meteorus or vice versa”.

What really has to be considered, however, is the possibility that Meteorus Haliday

s.s. and Zele Curtis (Zemiotes of Mason) have a common ancestor. The group

defined by synapomorphous character-states has to include all descendants of this

common ancestor, to avoid defining para- and polyphyletic groups. Of the

characters used by Mason to unite Zele Curtis and Homolobus Foerster probably

not one is an apomorphous character-state! My view is supported by the existence

of species of Meteorus which are very close to Zele in all characters used to

separate both genera.

Mason (1973: 214) argues that because Meteorus and Zele have different

combinations of character-states they are not closely related. But, in my opinion,

the character-states he uses to separate the two genera are plesiomorphous and

cannot be used either to separate the genera upon phylogenetic grounds or to

justify their inclusion in different subfamilies. The existence of two closely related

genera with different sets of plesiomorphous character-states is easy to accept if a

different path of further evolution for both genera (as is most likely) is assumed.

The existence of intermediate Meteorus specimens connects both groups. As

pointed out by Huddleston (in litt.) at least males of the European Meteorus

abdominator (Nees) are intermediate in the degree of setosity of the metasomal

tergites. In this species rather wide bands of setae occur in the males, which are

almost equal in extent to the bands of setae in smaller species of Zele, e.g., Zele

caligatus (Haliday). Also the marginal cell of the hind wing of Meteorus

abdominator (Nees) and of a new species from India (DZD) is not distinctly

narrowed distad, also showing an intermediate character-state, indicating the

relationship of Zele and Meteorus. The anteriorly situated transverse carina of the

propodeum (fig. 824) in Zele occurs also in Meteorus s.s., but not in the

Homolobinae as defined in this paper.

Because of the synapomorphous character-states of the genera Exasticolus and

Homolobus on the one hand (the tribus Homolobini) and the genera Charmon and

Charmontia on the other (the tribus Charmontini) I consider the two tribes to be

sister-groups.

The placement of the genus Charmon by previous authors varied; Tobias (e.g.,

1976: 31) and Capek (1973: 264) included Charmon in the subfamily Mima-

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979.

gathidinae (a junior synonym of Orgilinae Ashmead). This was still rejected by

Capek in 1969 (p. 308), who included the genus Charmon (as Eubadizon) in his

Macrocentrinae basing his conclusion on a detailed study of the larval characters.

Additionally the emergence opening of the cocoon is regular in shape by removing

a cap at one end of the cocoon in the genera Homolobus and Charmon, while in the

Orgilinae s.s. the emergence opening is irregular (Capek, 1970: 853). Actually

Capek (1970: 850) considered Macrocentrus, Homolobus (as Zele auct.) and

Charmon (as Eubadizon auct. p.p.) to be closely related. As pointed out above, and

earlier (Van Achterberg, 1976b: 37), I agree with Capek’s view about a close

relationship between Homolobus and Charmon, but I have to disagree about a close

relationship with the Macrocentrinae s.s. Most of the complex of synapomorphous

character-states of the Macrocentrinae s.s. are not shared by the Homolobinae.

Thus the sister-group of the Homolobinae is more likely to be formed by the

Orgilinae s.s. (together forming group E of Van Achterberg, 1976b: 51, fig. 123).

Both have the larval labial sclerite transverse and the epistomal arch and dorsal

part of the hypostoma absent, or at least unsclerotized. The genus with most of the ©

plesiomorphous character-states in the Orgilinae is Microtypus Ratzeburg. The

adults share the following synapomorphous character-states with the

Homolobinae: 1—1st tergite somewhat constricted behind the spiracles;

2— reduction of the dorsal carinae of the Ist tergite; 3—reduction of the 3rd labial

palp segment; 4—endoparasites of lepidopterous larvae; 5—prepectal carina

reaching anterior margin of mesopleuron; 6—apex of antenna with a well

developed spine; 7— Ist discal cell of fore wing (sub)sessile or nearly so.

The inclusion of the Homolobinae as a tribe in the Helconinae (Watanabe, 1969:

319) is rejected because of differences of larval (Capek, 1970: 853) and adult

morphology (Van Achterberg, 1976b: 853), together with differences in their

biology.

Exceptionally a shallow and narrow antescutal depression, combined with a

weak constriction of the Ist tergite behind the spiracles is present in the

Agathidinae, but this seems to be a parallelism, because the plesiomorphous

character-states are also present in the Agathidinae. Additionally, no convincing

set of synapomorphous character-states has yet been found, a necessity to validate

the sister-group concept for the Agathidinae and Homolobinae, but further

research may reveal such a relationship.

The evolution within the tribe Homolobini is fairly complicated. The new genus

Exasticolus is easy to separate by two (for the subfamily unique or

autapomorphous) character-states, viz., the presence of a hind tibial comb and the

curved 1-SR + M in the fore wing. It may reflect the early isolation of the group on

the South American continent. Additonal apomorphous character-states are the

mainly smooth precoxal suture, the small 3rd labial palp segment and the short

ovipositor. Its sister-group, the genus Homolobus Foerster, consists of several

subgroups (subgenera) of which the subgenus Apatia Enderlein probably

originated first. Remarkable characteristics of Apatia are the simple claws and the

phenocline towards loosing the sharp apex of the hind tibial spurs of the males

(figs. 709— 713).

255

VAN ACHTERBERG: Revision Zelinae auct.

9H91898184 = d :[BIdONOON = N :[eoIdo}o1]y = y :suonerAssgge Futmo][oJ au} Aq poreorpur ase 99e]d uaye} aAey

few uoneroods Aiewurid ay} yorym ut suo1daı s1yde18098007 oy | "seurgojowod ay} ur sdiysuone]aI ajgıssod jo wei3801puap ‘61 ‘314

juasqe adouoad-

quasa1d uotsseidep [eInosogue+

quesqe (IsouJe) Sura pury Jo vz+

quesoid Burm 2107 Jo w-ı-

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3U31213S (ISOUTE) BUIM 9107 Jo W+NS-I-

quesqe quod TEIQII pury-

TUIZUOWLIEYI

quasa1d Zum puty JO VZ-

quasqe SuIM 2103 JO U-1+

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ATTRBIQUIA

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died TEIQEI pıg-

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= x3AU09 SMETI-

FO METO pury 19UUT peaano SUIM 2107 FO W+AS-I+ poonpez Juousas DAAUOUIPYD

queseid quod TEIQII PUIU+ died terqer pıg+

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panino ATHESM 10 3U31213S

Sura pury JO [4+9S pue ys-

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3U3I1213S Burm 2107 Jo VZ+VI- panino Sura pury

JO IH+9S PUE US+

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panino SUIM 9107 JO WZ+VI+

207007 hyd

snydo7no

snoydiowotsetd = -

“ce snoydıowode = +

SngO70740y)

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

The following separations seem to have taken place in the Palaearctic region.

Firstly the subgenera Phylacter Reinhard and Oulophus subgen. nov. Phylacter is a

small group of species, related to Oulophus and is characterized by the strongly

curved SC + RI of the hind wing and the more or less curved base of SR of the hind

wing. The new subgenus Oulophus is a large and rather diverse group of species,

the primary speciation of which seems to have taken place in the Holarctic region.

Secondly evolved the group of the subgenera Homolobus Foerster and Chartolobus

subgen. nov., which is characterized by a well-developed ridge on the 3rd—9th

antennal segments of the female and inequal hind claws of the female. The new

subgenus Chartolobus is the most peculiar subgenus of Homolobus, because of the

curved vein 1A +2A of fore wing, together with a lamelliform ridge at the base of

the antenna and peculiar inner hind claws of the female. The supposed relations in

the Homolobinae are depicted in fig. 19.

BIOLOGY

All the species treated in this paper are primary endoparasites of Lepidopterous

larvae; they pupate outside the host and construct a parchment-like, spindle-

shaped cocoon, which is covered by some loose silk. Allen (1977: 111) reported a

period of obligate ectoparasitism in Homolobus infumator (Lyle) after the host

(Campaea margaritata (L.)) has spun the cocoon. The ectoparasitic phase of the

final instar larvae of H. infumator lasted about 24 hours. During this phase the host

is almost entirely devoured (only the head capsule remains), thereafter the parasite

larva began to spin (slowly) its large white cocoon, taking 24—36 hours to

complete this. After 17—20 days under outdoor conditions the adults were bred

(Allen, in litt.). The absence of teeth at the mandibles of the final instar larvae of

the Euphorinae may indicate the absence of an ectoparasitic phase, because the

teeth on the mandibles of the larvae of the Homolobinae are presumably adapted

to the short ectoparasitic way of life.

Many species have an ophionoid facies (body largely yellowish, slender; eyes

and ocelli large and metasoma of 9 more or less compressed apically) and are

frequently captured at light. As pointed out by Gauld & Huddleston (1976: 35)

larvae of many Noctuidae and of some Lasiocampidae exhibit the habit to remain

concealed by day and coming out to feed at night. The nocturnal behaviour of the

parasites may be an adaptation to the activity-pattern of the hosts. There is a

profound difference in the host-selection between the species of the two tribes of

the Homolobinae. The tribe Charmontini contains parasites of small Lepidoptera

larvae with a hidden way of life, mainly in rolled and/or spun leaves, in stored

products or in mines of leaves. The species of the tribe Homolobini are parasites of

exposed living larvae of Lepidoptera, which may have a more nocturnal behaviour

as do the adult parasites. The differences are reflected in the shape and length of

the ovipositor: long and rather slender in the Charmontini, but usually short and

rather stout in the Homolobini. Some species of Homolobus (e.g., H. (O.) armatus

spec. nov.) have a long and slender ovipositor, which may indicate that more

hidden larvae are also used as prey. Unfortunately no host records are known for

VAN ACHTERBERG: Revision Zelinae auct. 257

these species. I consider a long ovipositor (and the associated selection of hidden

larvae as hosts) to be a plesiomorphous character-state, while the usually very

short ovipositor of most Homolobini is apomorphous.

The known hosts of Charmon cruentatus Haliday are sparse and comprise three

species of Tortricidae (Archips rosaceana Harris, Grapholitha molesta (Busck), and

Acleris variana (Fernald)), while of C. extensor (L.) many hosts are known. They

belong mainly to the Tortricidae (Acleris variana (Fernald), A. fuscana (?), A.

minutacinderella (?), A. oxycoccana Packard, Argyrotaenia pinatubana Kearfott, A.

tabulana (?), Choristoneura murinana (Hübner), C. fumiferana (Clemens), Epinotia

infuscana (?), Eucosma radicana Walsingham, and Grapholita molesta (Busck)),

Gelechiidae (Evagora spec. Eucordylea atrupictella Dietz, Recurvaria apicitri-

punctella (Clemens), R. canusella Chambers, R. milleri Busck, R. piceaella Kear-

fott, and R. starki Freeman), Coleophoridae (Coleophora ulmifoliella (?)), Oeco-

phoridae (Hoffmannophila pseudospretella Stainton), Geometridae (Operophthera

bruceata Hulst), and Pyralidae (Dioryctria reniculella (Grote)). Of the new genus

Charmontia no host records are known.

Of the new genus Exasticolus only one host is known, viz., Gloveria ballovi

Schaus, belonging to the Lasiocampidae. The hosts of the species of the subgenus

Apatia of the genus Homolobus mainly belong to the Noctuidae and Geometridae.

The Palaeotropical H. (A.) ophioninus (Vachal) is known to be a parasite of

Noctuidae (Spodoptera exempta Walker and Agrotis segetum (Denis & Schiff.)). The

closely related H. (A.) truncatoides spec. nov. is only known to have its host on

sugar beet (Beta). The only known host of H. (A.) elagabalus (Nixon) belongs to the

Noctuidae, viz. Selepa celtis Moore. The only species of this subgenus with many

host records is H. (A.) truncator (Say). The hosts belong to the Noctuidae (Agrotis

segetum (Denis & Schiff.), A. venerabilis Walker, Amathes smithii (Snellen),

Heliothis armigera (Hübner), Plusia gamma (L.), Porosagrotis orthogonia

(Morrison), P. tristicula (Morrison), Prodenia ornithogalli Guenée, Spodoptera

exigua (Hubner), and S. frugiperda Smith), Geometridae (Alsophia quadripunctata

Esper, Erannis bajaria (Denis & Schiff.), E. sorditana Hübner, Fidonia cebraria Tr.,

F. fasciolaria (Rottemburg), Hypagyrtis piniata (Packard), Lycia zonaria (Denis &

Schiff.), and Semiothisa bitactata Walker), Gelechiidae (Gnorimoschema operculella

Zeller), and Pyralidae (Margaritia sticticalis (L.)). An aberrant host spectrum seems

to be present in the Afrotropical H. (A.) huddlestoni spec. nov. All four hosts

belong to the Lymantriidae (Arctornis rubricosta Hering, Euproctis fasciata Walker,

E. rubricosta Fawcett, and E. sanguiguttata Hampson).

The only species of the subgenus Chartolobus with known hosts is H. (C.)

infumator (Lyle). They belong mainly to the Geometridae (Alcis repandata (L.),

Nepytia canosaria (Walker), Bupalus pinarius (L.), Campaea margaritata (L.),

Ectropis deodarae (?), Ematurga atomaria (L.), Lambdina fiscellaria (Guenée), L.

somniaria (Hulst), and Lycia zonaria (Denis & Schiff.); additionally it has been

reared from Oecophoridae (Agonopterix alstroemeriana (Clerck)), Noctuidae

(Orthosia stabilis (Denis & Schiff.)), and Pyralidae (Phycita roborella (Denis &

Schiff.)).

The only species of the subgenus Homolobus with host-records is H. (H.) discolor

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

(Wesmael). Probably it is a parasite of Geometridae (Alcis repandata (L.), Boarmia

spec., Cabera pusaria (L.), Ennomos spec., Eupithecia abietaria (Goeze), Geometra

alniaria L., Larentia spec., Odontopera bidentata (Clerck), and Thera variata (Denis

& Schiff.)) and Noctuidae (Acronycta aceris L., Lithocampa ramosa Esper, and

Polyphaenis sericata Esper). The host record of a Tortricid (Zeiraphera rufimitrana

(Herrich-Schäffer)) is probably incorrect.

The only species of the subgenus Phylacter with host records is H. (P.)

annulicornis (Nees). It seems to be mainly a parasite of Noctuidae (Apamea

unanimis (Hübner), Cosmia trapezina (L.), Enargia ypsillon (Denis & Schiff.),

Eupsilia transversa (Hufnagel), Lithophane lamda (F.), Mamestra brassicae (L.),

Mythimna obsoleta (Hübner), Naranga aenescens Moore, Orthosia populeti (F.), O:

stabilis (Denis & Schiff.), Panolis flammea (Denis & Schiff.), and Xestia triangulum

(Hufnagel)). Additionally reared from Pyralidae (Cnephalocrocis medinalis Guenée,

Margaritia sticticalis (L.), Phycita roborella (Denis & Schiff.)), Geometridae (Alcis

repandata (L.), and Alsophila aceraria (Denis & Schiff.)), and Tortricidae. The host

records of Tortricidae (Archips rosana L., and Tortrix viridana L.) are probably

incorrect, considering the size of the parasite.

The only species of the subgenus Oulophus with known host data, viz., H. (O.)

flagitator (Curtis), seems to be a specalized parasite of Geometridae (Alcis

repandata (L.), Campaea perlata (Guenée), Caripeta divisata Walker, Entephria

caesiata Lang (on Vaccinium myrtillus L.), Eupithecia annulata Hulst, E. harrisonata

MacK., E. longipalpata Packard, E. olivaceae Taylor, E. placidata Packard, E.

unicolor Hulst, Larentia citrata (L.), Melanolophia spec., Nyctobia limitata (Walker),

and N. nigroangulata Strecker).

The numerous host data of the genus Zele indicate a wide spectrum in some

species (Z. albiditarsus Curtis and Z. chlorophthalmus (Spinola)), while other

species (Z. niveitarsis (Cresson) and Z. caligatus (Haliday)) seem to be restricted to

one family of Lepidoptera. Zele caligatus (Haliday) is known from only one genus

of Geometridae, viz., Eupithecia (E. absinthiata (Clerck), E. expallidata Doubleday,

E. filmata Pears., E. goossensiata Mabille, E. indigata (Hùbner), E. luteata Packard,

E. palpata Packard, FE. satyrata (Hübner), and E. ?usurpata Pears.). The host record

of a Nymphalid (Euphydryas aurinia (Rottemburg)) is probably erroneous.

The known hosts of Zele niveitarsis (Cresson) belong mainly to the Pyralidae

(Acrobasis betulella Hulst, A. comptoniella Hulst, A. ostryella (?), A. rubrifasciella

Packard, A. sylviella (©), Meroptera pravella (Grote), Salebria contatella Grote, S.

subcaesiella (Clements), and S. virgatella (Clements)). Additional records, which

need verification are from Geometridae (Rheumaptera hastata (L.)) and Noctuidae

(Lithopane spec.).

The recorded hosts of Z. chlorophthalmus (Spinola) belong to the Pyralidae

(Acrobasis consociella (Hübner), A. tumidana (Denis & Schiff.), Eurhodope suavella

(Zincken), Eurrhypara coronata (Hufnagel), Margaritia sticticalis (L.), Nephopterix

adelphella (Fischer von Röslerstamm), N. hostilis (Stephens), Phlyctaenodes

turbidalis (Tr.), Phycita roborella (Denis & Schiff.), Poinea forficalis (L.), and Sylepta

ruralis (Scopoli)), Geometridae (Angerona prunaria (L.), Crocallis elinguaria (L.),

Ematurga atomaria (L.), Odontopera bidentata (Clerck), and Rheumaptera cervinalis

VAN ACHTERBERG: Revision Zelinae auct. 259

(Scopoli)), Noctuidae (Jaspidia pygarga Hufnagel and Metoponia koekeritziana

(Hübner)), Tortricidae (Cnephasia communana (Herrich-Schäffer), Laspeyresia

pomonella (L.), Tortrix viridana (L.)), Lasiocampidae (Malacosoma neustria (L.)),

Lymantriidae (Lymantria monacha (L.)), Arctiidae (Spilosoma urticae (Esper)), and

Limacodidae (Apoda avellana (L.)).

The numerous host data concerning Z. albiditarsus Curtis are grouped according

to the two colour-forms, to indicate their differences and similarities. The known

hosts of Z. albiditarsus Curtis f. deceptor (Wesmael) belong mainly to the

Geometridae (Anticlea badiata (Denis & Schiff.), Catarhoe cuculata (Hufnagel),

Chesias legatella (Denis & Schiff.), Chloroclysta truncata (Hufnagel), Colotois

pennaria (L.), Crocallis elinguaria (L.), Deileptenia ribeata (Clerck), Enypia moillieti

(?), Eupithecia indigata (Hübner), E. lariciata (Freyer), E. pseudotsugata MacK.,

Hydriomena caesiata (Denis & Schiff.), H. furcata (Thunberg), Ligdia adustata

(Denis & Schiff.), Nyctobia nigroangulata Strecker, Odontopera bidentata (Clerck),

Rheumaptera spec., Semiothisa continuaria (Walker), S. granitata (Guenée), S.

liturata (Clerck), S. unipunctaria perplexa (McDunnough), S. sexmaculata

(Packard), Spargania luctuata (Denis & Schiff.), Xanthorhoe fluctuata (L.)). Other

families of hosts represented are Noctuidae (Anarta myrtilli (L.), Hoplodrina alsines

(Brahm), /pomorpha retusa (L.), Lacanobia oleracea (L.), Lithacodia pygarga

(Hufnagel), and Syngrapha interrogationis (L.)), Pyralidae (Margaritia sticticalis (L.)

and Ostrinia nubilalis (Hübner)), Momphidae (Mompha contortella (?)), Tortricidae

(Acleris hastiana (L.), A. variana (Fernald), Epinotia solandriana (L.)), and

Saturniidae (Antheraea polyphenus Cramer).

Of the hosts of the nominate form of Z. albiditarsus (Curtis) comparatively more

hosts belong to the Noctuidae, but this may be because of the size of the parasite.

Large specimens of Z. albiditarsus usually belong to the nominate form and

Noctuidae are frequently larger than, for instance, Geometridae. The main part of

the known hosts belong to the Geometridae (Abraxas grossilariata (L.), Cidaria

pomonaria (Hübner), Eupithecia expallidata Doubleday, Macaria notata (L.),

Operophthora brumata (L.), Rheumaptera hastata (L.), and Thera obeliscata

(Hubner)), and Noctuidae (Anarta myrtilli (L.), Blepharita adusta (Esper), Dichonia

aeruginea Hubner, Dryobotodes eremita (F.), Hypena proboscidalis (L.), Lacanobia

oleracea (L.), L. suasa (Denis & Schiff.), Mamestria brassicae (L.), Orthosia cruda

(Denis & Schiff.), O. gracilis (Denis & Schiff.), O. miniosa (Denis & Schiff.), O.

stabilis (Denis & Schiff.), Panolis flammea (Denis & Schiff.), Polia nebulosa

(Hufnagel), and Zale spec.). Host records which need to be confirmed belong to

the Douglasiidae (Douglasia ocnerostomella (Stainton), Yponomeutidae

(Argyresthia brockeella (Hübner)), Lyonetiidae (Leucoptera scitella (Zeller)),

Arctiidae (Rhyparia purpurata (L.)), Gelechiidae (Aristotelia brizella (Treitschke),

and Caryocolum tricolorella (Haworth)), Conchylidae (Aethes francillana (F.), and

Falseuncaria ruficiliana (Haworth)), Pterophoridae (Adaina microdactyla (Hübner)),

Nymphalidae (Euphydryas aurinia (Rottemburg)), and Tortricidae (Semasia aemula

Schläg and Zeiraphera griseana (Hübner)).

260

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

KEY TO THE GENERA AND SUBGENERA OF HOMOLOBINAE AND METEORINI

First tergite petiolate and spiracles situated submedially (figs. 762, 775, 801);

antescutal depression absent; pronope more or less developed (fig. 874);

(tribus Meteorini) x... RA 9

First tergite sessile and spiracles situated subbasally (figs. 551, 729, 747);

antescutal depression present (figs. 31, 119, 168, 316); pronope completely

absent:(subfamily Homolobinae) sn eeN 2

Marginal cell of hind wing widened apicad (figs. 147, 161); vein r-m of fore

wing present (fig. 180); occipital carina present medio-dorsally (fig. 95); vein

2A of hind wing absent (fig. 85) or nearly so (fig. 107); lateral carina of.

mesoscutum.present; (tribus Homolobini) Le 3

Marginal cell of hind wing narrowed apicad (fig. 37); vein r-m of fore wing

absent (fig. 892); occipital carina reduced medio-dorsally (fig. 44); vein 2A of

hind wing present (figs. 63, 892); lateral carina of mesoscutum absent; (tribus

Charmontini) cc. otel e ENS),

Inner aspect of hind tibia without a comb apically (fig. 256); vein I-SR+M of

fore wing straight (fig. 258); tarsi without a ventral row of setae (Homolobus

Foersters Mise. tea Sexes cats Seka I O 4

Inner aspect of hind tibia with a well developed comb apically (figs. 98, 882);

vein 1-SR+M of fore wing curved distad (figs. 85, 94); tarsi with a weakly

developed row of setae ventrally ......... Exasticolus gen. nov. (p. 271)

At least fore tarsal claws with a minute subapical tooth or lamella (figs. 350,

394, 443); hind tibial spurs of g with a sharp, hyaline apex (figs. 710, 713) . 5

Tarsal claws simple or nearly so, without a tooth or lamella (figs. 123, 152,

212); hind tibial spurs of & sometimes without a sharp apex, spurs truncate

and pismented apically (igs 112,712 2 207 220 Apatia Enderlein (p. 277)

Submedially inner hind claw of © distinctly concave ventrally (figs. 35i, 439,

887, 888), its shape different from the outer claw (figs. 350, 443); antennal

ridge of 3rd—6th antennal segments of 9 strongly developed (figs. 349, 366,

424, BTT SHO) i i gion be aes OT O E 6

Submedially inner hind claw of 9 straight or convex ventrally (figs. 502, 679,

881), its shape similar to the outer claw (figs. 498, 678) or nearly so (figs. 570,

571); antennal ridge of 3rd—6th antennal segments of 9 usually absent (figs.

8719 880) ifpresentithenweakly developed ee fl

Vein 1A +2A of fore wing curved (figs. 343, 369, 380); basal third of vein SR of

hind wing curved and equally sclerotized as vein Ir-m of hind wing (figs. 349,

ZOT SSD ye pene a Ro cee eee Chartolobus subgen. nov. (p. 304)

Vein 1A +2A of fore wing straight (figs. 396, 419, 449): basal third of vein SR of

hind wing straight or weakly curved (figs. 396, 404, 425), much less sclerotized

than Le-m.( 999) e ee Homolobus Foerster (p. 311)

Basal third of vein SR of hind wing distinctly curved and sclerotized as vein Ir-

m (figs. 495, 507), if intermediate (fig. 482), then claws bifurcate (fig. 488); vein

SC + RI of hind wing (rather) strongly curved (figs. 482, 495, 507) ........

ee A Puen tin, Babes eS oc Phylacter Reinhard (p. 321)

Basal third of vein SR of hind wing straight (fig. 539) or rather curved (fig. 649)

VAN ACHTERBERG: Revision Zelinae auct. -261

and usually less sclerotized than Ir-m (fig. 655), if exceptionally well

sclerotized (figs. 618, 641), then SR of hind wing (almost) straight (fig. 641) and

claws with a small subapical tooth (fig. 643) or with a lamella (fig. 629); vein

SC+RI of hind wing straight (fig. 631) or moderately curved (fig.

COMME OIE. ANNE N Oulophus subgen. nov. (p. 327)

8. Length of 3rd segment of labial palp equal to length of 2nd segment (fig. 893);

Ist discal cell of fore wing petiolate (fig. 892); claws (except for the apical

tooth) straight ventrally (fig. 894); apical segment of antenna without spine

apically (fig. 897); scutellum weakly sculptured postero-medially (fig. 899); Ist

tergite slender, its length ca. 2.8 times its apical width (fig. 896) .........

NEEM ERINNERN, EI BRUDER Charmontia gen. nov. (p. 262)

— Third segment of labial palp absent or short, much shorter than 2nd segment

of labial palp (fig. 57); Ist discal cell of fore wing (sub)sessile (figs. 37, 63);

claws convex ventrally (figs. 40, 52); apical segment of antenna with spine

apically (fig. 53); scutellum smooth postero-medially (fig. 43); Ist tergite

comparatively stout, its length less than twice its apical width (figs. 31, 41,

THO) oie! ALLÉE RE RE een RE ERIN, Charmon Haliday (p. 263)

9. Marginal cell of hind wing widened apicad (figs. 784, 788); at least apical half

of 3rd and following tergites densely setose (figs. 783, 794); vein r of hind wing

present (fig. 788) or absent (fig. 758); dorsope more or less developed (figs.

TOA TOLD RUE BUBEN DONE WROTE PIELER I Zele Curtis (p. 359)

— Marginal cell of hind wing narrowed apicad or its sides are parallel; 3rd and

following tergites with a few rows of setae, exceptionally more extensively

setose; vein r of hind wing absent; dorsope variable .... Meteorus Haliday

Subfamily HOMOLOBINAE, nom. nov.

Syn.: Zelinae auct. p.p. |

Diagnosis. — Antescutal depression present; hypoclypeal depression absent; Ist

discal cell of fore wing (sub)sessile, 1-SR absent or nearly so, but in the new genus

Charmontia present (fig. 892); dorsope of Ist tergite absent; Ist tergite sessile, more

or less narrowed behind the spiracles, and spiracles situated in front of the middle

of the tergite; apical segment of antenna with a well developed spine, but absent in

the new genus Charmontia (fig. 897); occipital carina connected with the

hypostomal carina above the mandibular base; vein a of fore wing absent; pronope

absent; metapleural flange more or less lamelliform and transparent; prepectal

carina (almost always) reaching anterior margin of mesopleuron; hypostomal and

prepectal carinae present; lateral carina of scutellum absent; vein m-cu of fore

wing far antefurcal to 2-SR; subbasal cell of hind wing large (figs. 26, 161); lobes of

mesoscutum evenly convex; trochantelli simple, without teeth; scapus

(sub)truncate apically; veins CUlb, 2-SR, and 2A of fore wing present; Ist

subdiscal cell of fore wing closed distally; plical lobe of hind wing rather large;

laterope of Ist tergite deep, large and subbasal; mesopleuron not distinctly

protruding anteriorly; maxillary and labial palpi with, respectively, 6 and 4

segments, but 3rd labial palp segment often reduced, in the genus Charmon even

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

absent or nearly so (fig. 57); metasoma evenly setose; occipital carina present, at

least laterally; postpectal carina absent; hypopygium truncate apically, large to

medium-sized; antennal segments 37—55; ovipositor straight, or nearly so, and

with a small subapical notch.

Distribution. — Cosmopolitan. Contains two tribes: Charmontini and Homolo-

bini.

Tribus CHARMONTINI nov.

Diagnosis. — Occipital carina reduced medio-dorsally; tarsal claws without a

subapical tooth; anterior tentorial pits deep, medium-sized or large; precoxal

suture absent; middle lobe of mesoscutum more or less truncate anteriorly and

with a transverse protruding horizontal part (figs. 20, 60); vein 2A of hind wing

present; marginal cell of hind wing narrowed apicad; mandibles normal, not

twisted apically; vein 2-R1 of fore wing well developed (figs. 15, 892); hind tibial

spurs subequal, rather short, 0.2—0.4 times length of hind basitarsus; vein r-m of

fore wing absent (fig. 15); ventral margin of clypeus rather thick, not separated

from clypeus, and with a more or less developed row of punctures (fig. 898); eyes

bare and immarginate, medium-sized; frons and vertex smooth; face rather flat;

metapleural flange present as a narrow, rather thin and rounded ventral carina

(fig. 32); mesopleuron smooth or nearly so; Ist tergite concave medio-basally and

convex submedially; side of scutellum rugose (fig. 43); lateral carina of

mesoscutum absent; propodeum without a medial carina and areola, its posterior

part not separated from its antero-dorsal part; antepropodeal depression narrow;

fringe of wings short; vein 1A +2A of fore wing straight; vein 3-SR +SRI of fore

wing curved basally; 2nd tergite without a sharp lateral crease.

Distribution. — Cosmopolitan. Contains two genera: Charmontia gen. nov. from

the Neotropical region, and Charmon Haliday from the other regions.

Genus Charmontia nov.

Etymology: fantasy name based on the genus name Charmon, to which it is closely related. Gender:

feminine.

Type-species: Charmontia inopina spec. nov.

Diagnosis. — Length of body and of fore wing ca. 4 mm; apical segment of

antenna without spine apically (fig. 897); length of 3rd segment of labial palp equal

to length of 2nd segment (fig. 893); Ist discal cell of fore wing shortly petiolate (fig.

892); tarsal claws (except for the apical tooth) straight ventrally (fig. 894); anterior

tentorial pits deep and large (fig. 898); scutellum weakly sculptured postero-

medially (fig. 899); epicnemial area smooth, except for some rugae (fig. 890);

episternal scrobe well-impressed and elliptical (fig. 890); pleural suture moderately

crenulate and deep; length of Ist tergite ca. 2.8 times its apical width; length of

hind femur ca. 7.4 times its apical width; ovipositor sheath much longer than fore

wing; scutellum punctulate; propodeal spiracle round, small and situated

submedially in propodeum; parastigma rather large (fig. 892); vein cu-a of hind

VAN ACHTERBERG: Revision Zelinae auct. 263

wing almost straight and medium-sized (fig. 892); hind coxa punctulate, but

antero-dorsally rugulose and postero-dorsally striate (fig. 895).

Biology. — Unknown, but the long ovipositor suggests the same hosts as of the

genus Charmon.

Charmontia inopina spec. nov.

(figs. 889— 900)

Holotype, 9, length of body 4.4, and of fore wing 4.2 mm.

Head. — Antennal segments 41, length of 3rd segment equal to 4th segment,

length of 3rd and 4th segments both 7.3 times their width, both penultimate

segments each 2.5 times their width; length of maxillary palp 1.2 times height of

head; dorsal length of eye 1.9 times temples; temples directly narrowed apically

(fig. 900); POL : @ ocellus : OOL = 8: 3: 8; frons convex, but behind antennal

sockets flat and medially with a shallow suture; face punctulate, only finely striate

medio-dorsally (fig. 898); clypeus rather convex, protruding apically (fig. 890),

punctulate, and its margin straight medially; length of malar space 1.3 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.6 times its height; pronotal sides smooth,

but medially crenulate and anteriorly with some rugae (fig. 890); anterior half of

notauli (except posteriorly) distinctly impressed and crenulate (fig. 899);

mesoscutal lobes smooth and distinctly convex; metanotum striate medially;

surface of propodeum smooth, only medially reticulate-rugose.

Wings. — Fore wing: r : 3-SR+SRI : 2-SR = 6: 44: 9; 1-SR+M straight; cu-a

inclivous and longer than 1-CU1; 1-CU1 : 2-CUl = 5: 29. Hind wing: SC+RI

weakly curved; 1-M : cu-a = 1.1: 1 (fig. 892).

Legs. — Length of femur, tibia and basitarsus of hind leg 7.4, 13.6, and 11.7

times their width, respectively; length of spurs of hind tibia 0.2 times hind

basitarsus, subequal (fig. 895).

Metasoma. — Length of Ist tergite 2.8 times its apical width, its surface smooth,

but medially somewhat rugulose; dorsal carinae of Ist tergite weakly developed in

basal fifth and spiracles protruding (fig. 896); 2nd tergite smooth; length of

ovipositor sheath 1.88 times fore wing.

Colour. — Blackish-brown; pterostigma, 2nd and 3rd tergites, and hind coxa,

dark brown; rest of legs, palpi, tegulae, scapus basally, annellus and metasoma

ventro-basally, more or less brownish-yellow; tibiae and coxae somewhat

infuscated.

Holotype in CNC, Ottawa, 9: “Pichinahuel, Cord. Nahuelbuta, Arauco, Chile,

10—20.1.1959, L. Pefia’’, “New genus near Charmon, Det. W.R.M. Mason 76”.

Genus Charmon Haliday

Haliday, 1833, Ent. Mag. 1: 262.

Capek, 1969, Proc. ent. Soc. Wash. 71: 308 (as Eubadizon).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 228, 230.

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Capek, 1970, Can. Ent. 102: 850, 853, 868, 870, fig. 15.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 231.

Capek, 1972, Ent. Problémy 10: 133, 136.

Capek, 1973, Acta Inst. forest. zvol.: 264.

Van Achterberg, 1974b, Norsk ent. Tidsskr. 21: 110.

Mason, 1974, Proc. ent. Soc. Wash. 76: 237, 238.

Gauld & Huddleston, 1976, Entomologist’s Gaz. 27: 43, 47, fig. 18.

Van Achterberg, 1976a, Tijdschr. Ent. 118: 250.

Van Achterberg, 1976b, id. 119: 37, fig. 100.

Tobias, 1976, Opr. Fauna SSSR 110: 136.

Type-species: Charmon cruentatus Haliday.

Synonyms: Provancheria Ashmead, 1900; Cyclocormus Cameron, 1911; Eubadi-

zon auct. p.p.

Diagnosis. — Length of body 2.9—5.2, and of fore wing 3.3—5.9 mm; apical

segment of antenna with an apical spine (fig. 53); 3rd segment of labial palp absent

or shortly developed, much shorter than 2nd segment of labial palp (fig. 57); Ist

discal cell of fore wing (sub)sessile (figs. 37, 63); tarsal claws convex ventrally (figs.

40, 52); anterior tentorial pits deep and medium-sized (fig. 27); scutellum smooth

postero-medially; epicnemial area smooth; episternal scrobe narrow and linear;

pleural suture indistinctly and finely crenulate, narrow and shallow (fig. 20); length

of Ist tergite 1.3—1.7 times its apical width; length of hind femur 5.3—6.8 times its

width; length of ovipositor sheath 0.60—1.55 times fore wing; scutellum smooth;

propodeal spiracle round, small, and situated in front of middle of propodeum;

parastigma large (fig. 63); vein cu-a of hind wing long and straight; hind coxa

smooth or nearly so.

Biology. — Parasites of larvae of Lepidoptera with a hidden way of life.

Key to the species of the genus Charmon

1. Vein cu-a of fore wing much shorter than 1-CUI (fig. 37), resulting in a rather

transverse Ist subdiscal cell; basal half of vein M+CUI of fore wing scarcely

sclerotized; length of ovipositor sheath ca. 0.7 times fore wing; pterostigma

dark brown medially; Australian region ... brevinervis spec. nov. (p. 267)

— Vein cu-a of fore wing longer than 1-CUI, exceptionally subequal (figs. 26, 49,

75); M+CUI largely sclerotized; Ist subdiscal cell less transverse (fig. 63);

length of ovipositor sheath usually 0.82—1.55 times fore wing, if exceptionally

shorter, then pterostigma yellowish medially; Holarctic, Afrotropical, and N.

Oriental regionst. wu RI >

2. Length of ovipositor sheath 0.60—0.74 (forma brevicaudus (Hellen)) or

0.82—1.20 (nominate form) times fore wing, exceptionally longer; pterostig-

ma, apex of hind tibia, and hind tarsus yellowish, if intermediate, then middle

of hind tibia and tarsus similarly coloured ..... cruentatus Haliday (p. 268)

— Length of ovipositor sheath 1.21—1.55 times fore wing, exceptionally shorter;

pterostigma, apex of hind tibia, and/of hind tarsus infuscated, if intermediate,

then middle of hind tibia lighter coloured than apices of hind tarsal

sementi ee Ber SEE extensor (Linnaeus) (p. 265)

VAN ACHTERBERG: Revision Zelinae auct. 265

Charmon extensor (Linnaeus)

(fig. 20—31)

Linnaeus, 1758, Syst. nat., Ed. 10: 564 (as Ichneumon).

Provancher, 1880, Naturaliste can. 12: 171 (Eubadizon gracilis). Syn. nov.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 234, 237, 238.

Capek, 1970, Can. Ent. 102: 853, fig. 15.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 231.

Capek, 1972, Ent. Problemy 10: 133, 136 (re-identification needed).

Mason, 1974, Proc. ent. Soc. Wash. 76: 237, 238.

Tobias, 1976, Opr. Fauna SSSR 110: 136 (re-identification needed).

Fitton, 1978, Biol. J. Linn. Soc. 10: 377.

Note. Because of the presence of two species in the Palaearctic region the

existing literature (especially of Western European origin) deals, at least partly,

with Charmon cruentatus Haliday.

Redescribed after a © from Canada, Mobert; this © was compared with the

holotype of C. gracilis (Provancher) by Dr. W. R. M. Mason (Ottawa).

Head. — Antennal segments 44, its 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 5.7 and 5.0 times their width, respectiveiy, both penultimate

segments 1.7 and 1.8 times their width (fig. 25); length of maxillary palp 1.2 times

height of head; dorsal length of eye 2.3 times temples; temples directly narrowed

apicad (fig. 28); POL : @ ocellus : OOL = 5: 5: 7; frons flat; face almost smooth,

but with some aciculation near the antennal sockets (fig. 27); clypeus rather

convex, indistinctly punctulate, its margin straight medially; length of malar space

0.4 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; pronotal side smooth,

except for some indistinctly developed sculpture medially and posteriorly; notauli

absent; mesoscutum smooth, except for some punctures (fig. 30); metanotum

medially without well-developed carinae; surface of propodeum smooth, except

for some microsculpture medio-basally (fig. 31).

Wings. — Fore wing: r: SRI +3-SR : 2-SR = 11:81:21; 1-SR+M sinuate; cu-a

inclivous and longer than 1-CU1; 1-CUI : 2-CUI = 1: 10. Hind wing: SC+RI

weakly curved; 1-M : cu-a = 0.8: | (fig. 26).

Legs. — Femur, tibia and basitarsus of hind leg 5,9, 11.4, and 11.7 times their

width, respectively; length of spurs of hind tibia 0.4 times the basitarsus, subequal

(fig. 29).

Metasoma. — Length of Ist tergite 1.5 times its apical width, its surface finely

longitudinally striate; dorsal carinae of Ist tergite absent and its spiracles

indistinctly protruding (fig. 31); 2nd tergite smooth, but baso-laterally somewhat

aciculate; length of ovipositor sheath 1.46 times fore wing.

Colour. — Dark reddish-brown; pedicellus, annellus, patch between eyes and

ocelli, mandibles, prothorax, tegulae, metapleuron partly, metasoma ventrally and

legs, yellowish, but hind tibia (except base), middle and hind tarsi, infuscated; 2nd

and 3rd tergites more reddish; pterostigma rather dark brown.

Redescribed after 9 from CNC: “Ex Dioryctria reniculella, Mobert, Ont.”,

“Eubadizon gracile Prov., CWT, Det. W. R. M. Mason, (19)60”.

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Specimens additionally examined: 215 9 and 97 g. From the Nearctic region:

North West Territories (Yellowknife), Newfoundland (Blackhead), Nova Scotia

(St. Peters; Grand River; Halifax), New Brunswick (Kedgwick; Summit Depot;

Charlo; Green River (Lab.); Golden Ridge, Carlton Co.; Tobique; Upsalquitch),

Quebec (Montigny; Park Reserve, Kam. Co., 950 ft; Harrington Lake, Gatineau

Park; Lac Crescence; Duchesnay; Lac Mondor; Cascapedia River, Cape Rouge;

Berthierville; New Richmond), Ontario (Rush Biv., 15 mi. SE Kenora; Cedar

Lake; Chaplean; Bothwell; Arden; Hawk Lake; Wawa; White Falls; Thamesville;

Vivian; Rossport; nr. Lake Erie; Richmond; Simcoe; Mobert; St. Davids,

Normondale; Prescott; Ottawa, Holtyre; Stittsville; Chatterton; Dryden;

Porcupine; Siouxhook; Vermilion Bay; Belleville; Twin Elm; One Sided Lake;

South March), Alberta (Edmonton; Coleman; Eisenhower Jet, 4700 ft, Banff;

Orion; Granada, Jasper), Saskatchewan (Saskatoon; Great Sand Hills, W. of Swift

Current), British Columbia (Vancouver; Cowichan Lake; Victoria; Longford;

Keremeos; Terrace), New Hampshire (6 mi S. Gorham, Notch Road), New York

(Ithaca; Oneonta; Canadarago Lake), Massachusetts (Springfield), Michigan (Ann

Arbor), Montana (Missoula), Wyoming (Yellowstone National Park, Cathedral

Mt.), Oregon (Joseph), Colorado (Maybell; Nederland), North Carolina

(Highlands, 3800 ft; Clingman’s Dome), Virginia (Mountain Lake), Minnesota

(Kawishiwi Field Lab.), Florida (Gainesville; Torreya St. Park), Texas (Navasota),

Nevada (Lee Canyon, 38 mi. NW Las Vegas), California (Mono Co., Tom’s Place;

Oioville; San Francisco; Mill Valley, Marin Co.; Stanislaw Co., 5 mi. N. Turlock

Lake; Pt. Reyes, Marin Co.; Mineral, 7400 ft; Tanayon Lake), Mexico (Dgo, 9000

ft, 10 mi. W. of El Salto; Chis., 7200 ft, S. Crist. Las Casas) (CNC, CAS, UCA,

AMNH, RMNH, TC).

Specimens examined from the Palaearctic region: Finland (Korpo; Traskrias;

Jomala), Sweden (Eksharad), USSR (Pavilnys; Ilmen, S. of Leningrad), Japan (Mt.

Arakura, 1300 m; Nagano, 400 m; Shizuoka; Soranuma, Hokkaido; Mt. Gozaisho,

Mie Honshu; Sarobeto, Hokkaido), Netherlands (Wijster; Drijber; Heerde;

Herpen; Baarle-Nassau; Delft; Ede (Sijsselt)), West Germany (Steinbach am

Worthsee; Witzenhausen; Lippoldshausen), Czechoslovakia (Kufra), Austria

(Gampenjoch, Sudtirol, 1500 m; Bischofshofen; Flachgau, Zistelalm; Salm-Moos,

Salzburg; Judenbergalm; Sollheim Autobahn), Italia (Bolzano, Sarntal, 1250 m;

Campi, Riva s. Garda, 800 m; Castel Tesino, Trento, 1200—1500 m; Meran, 650

m) (RMNH, IZP, ZMH, ZSB, UZM, EI, HC, WHC).

Specimens examined from the Afrotropical region: Zaire (Lubumbashi (=

Elizabethville)) (CNC, RMNH) and from the Oriental region (or South

Palaearctic): India (Kashmir, Ladakh, Batalik, 2743 m) (DZD).

The holotype of Ichneumon extensor Linnaeus was examined by Mr. T.

Huddleston (London), who kindly supplied his notes. This reveals that Charmon

gracilis (Provancher) is actually a junior synonym of extensor. The holotype of

extensor is in the Linnean Collection of the Linnean Society at London; the

condition of this very dirty specimen is fairly reasonable. It bears two handwritten

labels, one by Linnaeus (‘34 extensor’) and one by Smith (“extensor 935”). The

pterostigma is very pale yellow but with a slightly infuscated border, the hind tibia

VAN ACHTERBERG: Revision Zelinae auct. 267

is infuscated apically, the hind tarsus is slightly infuscated, and the length of the

ovipositor sheath is 1.22 times fore wing.

The holotype of C. gracilis (Provancher) (PC) was not available for examination,

but the description of the colour by Provancher indicates its synonymy with

extensor.

The variation of C. extensor (L.) is considerable: length of fore wing 3.5—5.3

mm; length of ovipositor sheath 1.21—1.55 times fore wing, exceptionally shorter;

pterostigma and hind legs usually infuscated, but completely yellowish specimens

occur (as in cruentatus). Especially the African specimens are yellowish; the

occurrence of extensor and cruentatus in the Afrotropical region is in my opinion

the result of a recent (Quarternary) invasion of both species from the Palaearctic

region. The crossing of the Sahara was probably fairly easy during the last ice-age

(De Jong, 1976). The only stable difference I could find was the yellowish colour,

while colour is known to be a factor easily influenced by the temperature during

the development of the larva and/or pupa. Also in the (Nearctic) Sonoran region

C. extensor (L.) becomes more or !ess yellowish, probably also because of the

influence of the temperature. In Europe extensor seems to be most common

(compared with cruentatus) where a (sub)continental climate prevails, while

cruentatus seems to be most common in a more or less Atlantic climate. Some of

the specimens examined were taken at light.

Known hosts of examined specimens: Acleris variana (Fernald), A. fuscana (?), A.

minutacinderella (2), A. oxycoccana Packard, Argyrotaenia pinatubana (Kearfott), A.

tabulana (?), Choristoneura fumiferana (Clemens), C. murinana (Hubner),

Coleophora ulmifoliella (2), Dioryctria reniculella (Grote), Epinotia infuscana (?), E.

nigricana (?), Eucordylea atrupictella Dietz, Eucosma radicana (Walsingham),

Evagora spec. (on Elm, Hemlock, and Black fir), Hoffmannophila pseudospretella

Stainton, Laspeyresia molesta (Busck), L. (?) arboreus (?), Operophthora bruceata

Hulst, Recurvaria milleri Busck, R. starki Freeman, R. apicitripunctella (Clemens)

(on Tsuga canadensis), R. canusella Chambers, and R. piceaella Kearfott.

Charmon brevinervis spec. nov.

(figs. 32—44)

Holotype, 9, length of body and of fore wing 4.4 mm.

Head. — Antennal segments 30, but apical segments missing, its 3rd segment 1.1

times 4th segment, length of 3rd and 4th segments 5.6 and 5.2 times their width,

respectively (fig. 34); length of maxillary palp equal to height of head (fig. 32);

dorsal length of eye 1.5 times temple; temple roundly narrowed apicad (fig. 44);

POL : @ ocellus : OOL = 5: 6: 10; frons weakly concave; face indistinctly

punctulate and with some rugulosity near antennal sockets (fig. 42); clypeus

transversely convex, its surface almost smooth, and its apical margin almost

straight medially, with long setae (fig. 42); length of malar space 0.5 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; pronotal side smooth,

except for some crenulae medio-anteriorly and posteriorly (fig. 32); notauli

268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

completely impressed, but shallowly and indistinctly crenulate (fig. 43); mesoscu-

tum smooth; metanotum with one medial carina; surface of propodeum smooth,

but medially weakly rugulose.

Wings. — Fore wing: r : SR1+3-SR : 2-SR = 15: 111 : 22; 1-SR+M weakly

sinuate (fig. 37); cu-a short, almost straight, much shorter than 1-CUI (fig. 35); 1-

CUI : 2-CUl = 6: 15; M+CUI basally reduced (which is in other species

complete, in fig. 75 of Cyclocormus luteus Cameron dotted because of damage).

Hind wing: SC + RI weakly curved; 1-M : cu-a = 0.4: 1.

Legs. — Femur, tibia, and basitarsus of hind leg 5.3, 10.0, and 7.4 times their

width, respectively; length of spurs of hind tibia 0.3 times the basitarsus, subequal

(fig. 38).

Metasoma. — Length of Ist tergite 1.6 times its apical width, its surface rather

coarsely longitudinally striate; dorsal carinae of Ist tergite developed in basal 0.4

and spiracles weakly protruding (fig. 41); 2nd tergite smooth; length of ovipositor

sheath 0.68 times fore wing.

Colour. — Yellowish-brown; middle of pterostigma, wing veins in middle third

of fore wing, propodeum, metapleuron, and Ist tergite, more or less dark brown;

hind tarsus rather whitish-yellow.

Holotype in RMNH, Leiden, 9: “Neth. Ind.-American New Guinea Exped.,

Rattan Camp, 1150 m, ii.1939, L. J. Toxopeus”. For the location of this camp, see

Toxopeus (1940).

Note. Besides the rather diffuse character-differences such as the somewhat

longer malar space, the stouter mesosoma, the rather short vein I-M of the hind

wing, the rather transverse Ist subdiscal cell, and the long setae at the clypeal

margin, this species is characterized by the reduction of the basal half of vein

M+CUI of fore wing, the short cu-a of fore wing and the short ovipositor sheath,

combined with its coloration. It is surprising to see how close this species is to both

other species of Charmon.

Charmon cruentatus Haliday

(figs. 45—79)

Haliday, 1833, Ent. Mag. 1: 262.

Nees von Esenbeck, 1834, Hym. Ichn. affin. Mon. 1: 236 (Eubadizon pectoralis). Syn. nov.

Cresson, 1872, Can. Ent. 4: 230 (Eubadizon pleuralis). Syn. nov.

Cameron, 1911, Ann. Transv. Mus. 2: 209 (Cyclocormus luteus). Syn. nov.

Hellen, 1958, Soc. Fauna Flora Fennica 4: 29 (Eubadizon brevicauda). Syn. nov.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 228, 230, 235 (Eubadizon extensor auct.).

Van Achterberg, 1974b, Norsk. ent. Tidsskr. 21(1): 110.

Mason, 1974, Proc. ent. Soc. Wash. 76(3): 237, 238.

Gauld & Huddleston, 1976, Entomologist’s Gaz. 27: 43, 47, fig. 18.

Van Achterberg, 1976b, Tijdschr. Ent. 119(3): fig. 100.

Redescribed from the neotype of Eubadizon pectoralis Nees, 9, length of body

5.1, of fore wing 5.3 mm.

VAN ACHTERBERG: Revision Zelinae auct. 269

Head. — Antennal segments 44, its 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 4.8 and 4.2 times their width, respectively, both penultimate

segments 1.8 and 2.2 times their width, respectively (fig. 53); length of maxillary

palp 1.5 times height of head (fig. 45); dorsal length of eye 2.5 times temple; temple

directly roundly narrowed apicad (fig. 50); POL : @ ocellus : OOL = 6: 5: 6; frons

flat; face smooth, except for some striation near the antennal sockets (fig. 54);

clypeus convex, punctulate, its apical margin straight medially and crenulate;

length of malar space 0.4 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; pronotal side anteriorly

crenulate-striate and posteriorly crenulate-rugose, rest smooth (fig. 45); notauli

impressed and narrowly crenulate (fig. 56); mesoscutum smooth, only medially

punctulate; metanotum with one medial carina; surface of propodeum smooth,

but medially punctate-rugose.

Wings. — Fore wing: r : SR1+3-SR : 2-SR = 9 : 53 : 13; 1-SR+M almost

straight (fig. 49); cu-a long, inclivous, longer than 1-CU1; 1-CUI : 2-CUl = 2: 38.

Hind wing: SC + RI slightly curved; 1-M : cu-a = 0.8: 1.

Legs. — Femur, tibia, and basitarsus of hind leg 6.2, 12.0, and 10.8 times their

width, respectively; length of spurs of hind tibia 0.3 times its basitarsus, subequal

(fig. 51).

Metasoma. — Length of Ist tergite 1.7 times its apical width, its surface

longitudinally striate (fig. 58); dorsal carinae of Ist tergite developed in front of the

weakly protruding spiracles (fig. 58); 2nd tergite smooth, but subbasally

indistinctly microsculptured; length of ovipositor sheath 1.05 times fore wing.

Colour. — Dark reddish-brown; palpi, pedicellus dorsally, annellus, patch

between eyes and ocelli, tegulae, ptero- and parastigma, C+SC+R of fore wing,

metasoma medio-ventrally and legs, yellowish; mandibles, mesopleuron,

mesosternum and metapleuron, reddish; stemmaticum blackish.

Neotype in KBIN, Brussels, 9: “Coll. Wesmael”, “1879”, “& Eubadizon

pectoralis N. V. Es. 9, det. C. Wesmael”, “Type”. Neotype of pectoralis herewith

selected, and labelled accordingly. Because Wesmael is the first revisor and the

Nees types are lost, a neotype is chosen to fix this name to one of the Palaearctic

species. The lectotype of Eubadizon pleuralis (Cresson) (9, ANSP) from Missouri

is a genuine cruentatus, its ovipositor sheath is 1.06 times fore wing, while the legs

and pterostigma are yellowish.

The holotype of Eubadizon brevicauda Hellén (WHC: “Terijoki, Hellén’’) from

Finland is an aberrant form of cruentatus with very short ovipositor (figs. 60—71),

its sheath is 0.60 times fore wing. Additional specimens of this form have been

examined from Norway (1 9, Selva, nr. entrance of Trondheimsfjord, ca. 150 m),

from the Netherlands (1 9, Wijster), and from Canada. A series reared from

Acleris variana (Fernald) (Vancouver, British Columbia) showed that the short

Ovipositor occasionally occurs within a group of normal cruentatus specimens. The

length of the ovipositor sheath of the form brevicaudus varies from 0.60—0.74 times

fore wing. The cocoon of cruentatus is (as is that of extensor) a parchment-like

brownish cocoon, covered by a fine filamentous tissue.

The holotype of Charmon cruentatus Haliday (9, NMI) differs not from Charmon

270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

extensor auct. nec L. The type is rather dirty, bears an old handwritten label

“cruentatus”, has a rather short ovipositor (sheath ca. 0.85 times fore wing), with

legs and pterostigma yellowish.

The holotype of Cyclocormus luteus Cameron (9, TMP, figs. 72—79) from S.

Africa is a yellowish form of cruentatus as far as can be judged from the few

Afrotropical specimens available for study. The body is mainly yellowish, with the

apical antennal segments, mesopleuron, propodeum, Ist tergite, metasoma

apically, pterostigma, and ovipositor sheath, more or less infuscated;

stemmaticum black; length of ovipositor sheath 1.05 times fore wing and antennal

segments 37.

Specimens examined: 260 9 and 102 3. From the Nearctic region: North West

Territories (Judith Island, McKenzie River), British Columbia (Robson; Salmon

River; Cultus Lake; Vancouver; Parksville); Ontario (St. Davids; Kimborn; Sioux

Lookout; Orillia; Mer Bleue; Iron Bridge; Trenton); Quebec (Knowlton); North

Carolina (Cherokee; Highlands, 3800 ft); Virginia (Mountain Lake; Roanoake);

Utah (Longan); Illinois (Urbana); Kentucky (Mammoth Cave, Watl. Pk.); Florida

(Gainesville; Torreya St. Park); California (Pozo, S.L.O. C.; Kernvale, Kern Co.;

Eel River Rgr. Sta., Mendo Co., 1500 ft); Mexico (Baja California, 1 mi. E.

Mission Santa Domingo) (CNC, UCA, RMNH).

Specimens examined from the Palaearctic region: Finland (Helsinki;

Taivassalo; Vehkalahti; Lemland, Flaka); Denmark (no localities); Ireland (id.);

England (Wyck Rissington, Glos.); Netherlands (Wijster; Assel (nr. Zwolle);

Heerde; Putten; Otterlo; Naardermeer; Muiderberg; Overveen; Waarder;

Asperen; Den Haag; Delft; Meijendel, Kijfhoek, Bierlap; Oostvoorne, dunes;

Ouddorp; Oostkapelle; Valkenswaard; Schayk; Asselt), West Germany (Aachen;

Thüringen; Bramwald; Grainbach, 800 m; Wiershausen; Gröbenzeller;

Kottenforst (nr. Bonn); Rondorfer Tal, Siebengebirge; Unteres Ahrtal,

Rheinprov.; Geisenheim, Rheingau); Austria (Salzburg, Flachgau, Veitlbruch; St.

Peter, Ahrntal, Süd-Tirol, 1300 m; Obergurgl, Tirol, 1950 m); Italy (Campi, Rivas.

Garda, 1500 m); France (Agoz, Haute Pyr.); Bulgaria (Rodopi, Velinograd)

(RMNH, ZMH, HC, UZM, EI, ZMB, ZIL, CVR, CNC). From the Afrotropical

region: Ivory Coast (Bingerville) and S. Africa (Pretoria) (MAC, TMP).

Variation: Antennal segments 37—44; length of fore wing 3.3—5.9, and of body

2.9—5.1 mm; length of Ist tergite 1.3—1.6 times fore wing; length of ovipositor

sheath 0.60—1.20 times fore wing and exceptionally apex of hind tibia infuscated.

Known hosts of examined specimens: Acleris variana (Fernald), Archips

rosaceana Harris, and Grapholita molesta (Busck).

Tribus HOMOLOBINI nov.

Diagnosis. — Occipital carina completely developed medio-dorsally; 3rd labial

palp segment well-developed, although often small (fig. 82); tarsal claws with or

without a subapical tooth or lamella; anterior tentorial pits deep and large;

precoxal suture variable (figs. 105, 204, 616); middle lobe of mesoscutum more or

less rounded anteriorly and without a protruding horizontal part (figs. 92, 119);

vein 2A of hind wing absent; marginal cell of hind wing widened apicad; mandibles

VAN ACHTERBERG: Revision Zelinae auct. 271

twisted apically; vein 2-R1 of fore wing absent (fig. 85) or short (fig. 107);

scutellum narrowly sculptured medio-posteriorly (figs. 119, 359); fringe of wings

short; hind tibial spurs unequal and long, inner spur reaching middle of basitarsus

(fig. 99); lateral carina of mesoscutum and vein r-m of fore wing present (fig. 18).

Distribution. — Cosmopolitan. Contains two genera: Exasticolus gen. nov. and

Homolobus Foerster.

Genus Exasticolus nov.

Etymology: from “‘n§aotic’’ (Greek for“fringe”) and “x ov’ (Greek for “‘leg’’), because of the fringe-

like comb apically at the inner side of the hind tibia. Gender: masculine.

Type-species: Zele fuscicornis Cameron.

Diagnosis. — Length of body 7.1—10.5, and of fore wing 6.6—10.1 mm; ventral

margin of clypeus thin, separated from clypeus (fig. 91) and smooth; eyes bare,

large, and distinctly emarginate (fig. 96); temples roundly narrowed apicad (fig.

86); metapleural flange large, lamelliform (fig. 80); precoxal suture mainly smooth

(figs. 80, 105); antescutal depression medium-sized to rather large, deep (fig. 119);

3rd segment of labial palp small, length of 4th segment 6—12 times 3rd segment

(figs. 82, 97, 117); ocelli large (fig. 91); epistomal suture present; pleural suture

shallowly and narrowly crenulate (fig. 80); metapleuron mainly smooth (figs. 92,

105); episternal scrobe deep and small to medium-sized; notauli complete, and

narrowly impressed (fig. 119); scutellar suture deep, with one longitudinal carina;

scutellum smooth and convex; side of scutellum mainly smooth but posteriorly

crenulate; propodeum at most with some irregular carinae, mainly smooth and its

posterior part not separated from its antero-dorsal part; propodeal spiracle large,

(sub)elliptical (fig. 80); antepropodeal depression medium-sized; 1-SR+M of fore

wing curved distad (figs. 85, 94); Ist discal cell of fore wing subpetiolate; r of hind

wing absent; SR of hind wing straight; SC+ RI of hind wing rather straight; 2-

SC+R of hind wing short; 1 r-m of hind wing more (figs. 85, 94) or less (fig. 107)

curved distad; cu-a of fore wing long and straight; parastigma large (fig. 85);

1A +2A and SRI of fore wing mainly straight; tarsi with a weakly-developed row

of setae ventrally; inner aspect of hind tibia with a well-developed comb of bristles

apically (figs. 98, 882); claws with a rather small, slender subapical tooth; length of

hind femur 5.2—7.2 times its width; length of Ist tergite 2.2—3.0 times its apical

width; Ist tergite concave medio-basally, convex medially and apically rather flat;

2nd tergite with a sharp crease laterally (fig. 80); metasoma of 9 compressed

apically; length of ovipositor sheath 0.06—0.10 times fore wing.

Biology. — The only host record (of E. nigriceps (Enderlein)) indicates a relation

to the Lasiocampidae, which may suggests the function of the peculiar comb of the

hind tibia. The comb may facilitate walking on the webs of the hosts during

infestation. The Lasiocampidae are not known to be hosts of Homolobus, the

sister-group of Exasticolus.

Distribution. — New World, contains three known species.

272: TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Key to the species of the genus Exasticolus

1. Middle coxa with an antero-ventral tooth (figs. 80, 89); 2nd tergite behind its

middle rugulose-aciculate (fig. 90); hind basitarsus stout, its length 6.4—6.8

times its maximum width (fig. 733); length of maxillary palp of © 1.2—1.3

times heiehtof head: way ee tuberculatus spec. nov. (p. 272)

— Middle coxa without a tooth (fig. 128); 2nd tergite usually smooth, at most

anterior half somewhat rugulose or pimply (figs. 104, 111); hind basitarsus

slender, its length 9.1—10.4 times its width (fig. 99); length of maxillary palp of

O 152. times heightof head... ern Men ONE 2

2. Length of malar space of 9 0.4—0.6 times basal width of mandible (fig. 96);

face less coarsely rugose and yellowish (fig. 96); vertex with long rugae,

reaching stemmaticum (fig. 95), exceptionally reduced; hind leg of 9

yellowish, exceptionally partly dark brown .. fuscicornis (Cameron) (p. 273)

— Length of malar space of 9 0.2—0.3 times basal width of mandible (fig. 110);

face coarsely rugose and blackish-brown (fig. 110); vertex almost smooth, at

most with some short rugae which do not reach the stemmaticum (fig. 116);

hind leg of © partly dark brown ......... nigriceps (Enderlein) (p. 275)

Exasticolus tuberculatus spec. nov.

(figs. 80—91, 731—733)

Holotype, 9, length of body and of fore wing both 9.1 mm.

Head. — Antennal segments 47, 3rd segment 1.3 times 4th segment and with an

indistinctly developed ridge, length of 3rd and 4th segments 3.9 and 3.1 times their

width, respectively, both penultimate segments 2.7 and 2.3 times their width,

respectively (fig. 83); length of maxillary palp 1.3 times height of head; dorsal

length of eye 2.4 times temple; POL : @ ocellus : OOL = 4: 6: 4; frons smooth,

rather flat; vertex dull coriaceous, and with some rugae anteriorly (fig. 86); face

shiny coriaceous, weakly convex; clypeus convex, indistinctly punctate (fig. 91);

malar space 0.6 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

smooth, but medio-anteriorly crenulate, posteriorly and ventrally narrowly rugose

(fig. 80); epicnemial area and mesopleuron smooth; precoxal suture weakly

impressed, only anteriorly with some rugae (fig. 80); notauli narrowly crenulate

(fig. 731); mesoscutal lobes remotely and indistinctly punctulate; surface of

propodeum submedially with some transverse rugae and anteriorly with a weak

medial carina, remainder smooth.

Wings. — Fore wing: r: 3-SR : SRI = 8: 12 : 51; 1-CUI : 2-CUl = 2: 26; 2-SR:

3-SR : r-m = 13: 12:8; 2A only developed as a brownish stripe; area basally of 2A

mainly bare as basal third of subbasal cell. Hind wing: Ir-m curved distad (fig. 81).

Legs. — Hind coxa punctulate, with some striae apico-dorsally (fig. 80); middle

coxa with a well developed tooth antero-ventrally (fig. 89); claws indistinctly

yellowish pectinate basally, except the inner claw (figs. 87, 88); femur, tibia, and

basitarsus of hind leg 5.2, 9.5, and 6.8 times their width, respectively; length of

VAN ACHTERBERG: Revision Zelinae auct. 273

spurs of hind tibia 0.8 and 0.6 times basitarsus, somewhat curved, almost straight.

Metasoma. — Length of Ist tergite 2.2 times its apical width, its surface rather

coarsely rugose behind the spiracles (fig. 80); dorsal carinae of Ist tergite absent,

except for a basal remnant, and its spiracles protruding; anterior third of 2nd

tergite almost smooth, somewhat coriaceous, its posterior two-thirds distinctly

obliquely rugose-aciculate; 3rd tergite only pimply; length of ovipositor sheath

0.09 time fore wing.

Colour. — Brownish-yellow; stemmaticum and vertex around stemmaticum,

blackish; mesoscutum somewhat more dark brown.

Holotype in TC, Ann Arbor: “Nova Teutonia, Braz., Santa Catarina, x.4.48,

Fritz Plaumann’’. Paratype: 1 9, CNC, “Nova Teutonia, 27°11’S, 52°23’W, Brazil,

300—500 m, 25.1x.1948, Fritz Plaumann”. Paratype: length of fore wing 9.3 mm,

length of ovipositor sheath 0.10 times fore wing; length of maxillary palp 1.2 times

length of head; length of hind basitarsus 6.4 times its width; 2nd tergite medially

sculptured, but apical third mainly smooth.

Exasticolus fuscicornis (Cameron) comb. nov.

(figs. 92—104, 882—884)

Cameron, 1887, Biologia cent.-am., Hym. 1: 509, fig. 17—4 (as Zele).

Viereck, 1911, Proc. U.S. natn. Mus. 40: 478 (Zele rosenbergi). Syn. nov.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 224—226.

Holotype, 9, length of body 10.5, of fore wing 10.1 mm.

Head. — Antenna incomplete (but in 9 other 9 specimens 42—48), remaining

segments 13, 3rd segment 1.2 times 4th segment, length of 3rd and 4th segment 3.8

and 3.1 times their width, respectively; length of maxillary palp 1.7 times height of

head; dorsal length of eye 2.4 times temple; POL : @ ocellus : OOL = 4: 10: 4;

frons concave, and almost smooth; vertex finely coriaceous, with some coarse

rugae anteriorly, which reach the stemmaticum (fig. 95); face rather smooth and

flat, near antennal sockets and laterally weakly transversely striate (fig. 96);

clypeus convex, smooth, except for some punctures medially and some striae

laterally; malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

smooth, except for some medial and apical rugae (fig. 92); epicnemial area

smooth, except for some short rugae; mesopleuron superficially punctulate;

precoxal suture absent, except for a shallow, smooth depression; notauli almost

smooth anteriorly, medially superficially and finely crenulate (fig. 101);

mesoscutal lobes smooth; surface of propodeum mainly smooth, except for an

irregular transverse carina and some rugae between the dorsal and posterior

surface (fig. 92).

Wings. — Fore wing: r: 3-SR : SRI = 16: 11: 57; 1-CUI : 2-CUI = 3: 21; 2-SR:

3-SR : r-m = 15: 11: 7; 2A and surroundings as in tuberculatus. Hind wing: Ir-m

more or less curved distad (fig. 92).

Legs. — Hind coxa smooth, but dorso-apically striate (fig. 92); middle coxa

without tubercle; inner hind claw equal to its outer claw (but in some specimens

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

slightly different), setose; femur, tibia, and basitarsus of hind leg 6.2, 9.9, and 10.0

times their width, respectively; length of spurs of hind tibia 0.6 and 0.5 times

basitarsus.

Metasoma. — Length of Ist tergite 3.0 times its apical width, its surface

superficially and irregularly striate laterally (fig. 104); dorsal carinae of Ist tergite

superficially developed up to middle of tergite; 2nd tergite smooth; length of

ovipositor sheath 0.06 times fore wing.

Colour. — Brownish-yellow; antenna (as much as is present, but scapus, (except

for a longitudinal dark brown stripe at the outer side and pedicellus, yellowish),

stemmaticum and vertex, blackish-brown; hind basitarsus and spurs lighter than

hind tibia.

Holotype in BM, London: “Type, H.T.”, “B.M. Type, Hym. 3.c.862”, “B.C.A.

Hymen. I., Zele fuscicornis, Cam.”, “Paso Antonio, 400 ft, Champion”, “Zele

fuscicornis Cam., Type, B.C.A., ii.4.09.”” (in Cameron’s handwriting). The type-

locality is situated in Guatemala.

Note. The males are very similar to the females in coloration, at most the

metasoma apically and the mesonotum blackish. Sometimes the antennae of both

sexes are rather yellowish brown. Total specimens examined: 88 © and 168 &.

From the Nearctic region: Ontario (Rondeau Prov. Pk.), Michigan (Ann Arbor),

Maryland (Takooma Pk.), South Carolina (Wattacoo; Greenville), Georgia

(Forsyth), Florida (Ft. Myers). From the Neotropical region: Mexico (Ver.,

Minatittan; Palomares, Oaxaco; Teapa, Tabasco), Panama (Fortuna, Chiriqui,

8°44'N, 82°15’W, 1050 m, at light), Costa Rica (Monteverde), Colombia (Dept.

Magdalena, Pueblo Bello, 45 km W. Valledupar, Sierra Nevadade, S. Marta, 1100

m; Anchicaya Dam, 1200 (m?), 17 km E. Buenavanture; Colegio, Bolivar;

Cundinamarca, Finca Bella Vista, nr. Sasaina), Ecuador (Coca, Napo R., Napo,

250 m; La Toma, W. Loja, 1500 m; Batapamba, 700 (m?); Sto. Domingo, 680 m,

Pich. Prov.; Zambra), Surinam (Sipaliwini, at light), Peru (Loreto, Pucallpa; Tingo

Maria, 750 m), Bolivia (20 km W. Laranjeiras, Dept. Beni; Rio Itenez, Pampa de

Meio, Dept. Beni; mouth of Rio Baures; Rio Mamore, Dept. Beni, approx. 5 km

NW mouth of Rio Grande; Rio Mamore, Dept. Santa Cruz, 2 km N. mouth of Rio

Chapare; Dpto Santa Cruz, Estac. Experimental General Saavedra; Dpto Santa

Cruz, Buena Vista; Alto Beni, Inicua R., 1100 m), Brazil (Nova Teutonia, Santa

Catarina, 27°11’S, 52°33’W, 300—500 m; Sampaio, Teodora; Jatai, Goias; Bahia,

Encruzihada, 960 m; Manaus; Pedra Azul, M. Ger., 600 m; Caruaru, 900 m; Serra

do Caraca, S. Barbara, M. Ger., 1600 m; Jacareacanga, Para; Linhares, E. Santo;

Représa Rio Grande, Guanabare; Itatiaja Nat. Pk., Rio de Jan.; Surumu, Roraine;

Reserva Ducke, Manaus; Vilhena, Rond.), Paraguay (Escobar; Filadelfia, Fern.

Col. Chaco, at light), and Argentina (Misiones, San Pedro; Salta, Tartagal;

Corrientes, Las Narias, Camino Villa Virasoro; Misiones, Dos de Mayo; 11 km W.

Las Cejas, Tucuman; La Plata; Horco Molle, nr. Tucuman) (TC, CNC, RMNH,

IML, BM, USNM, TMA, MSU, AMNH, CAS).

Variation: Antennal segments 42—49; length of body 6.5—8.6, of fore wing

5.7—8.4 mm; length of ovipositor sheath 0.05—0.08 times fore wing; Ist tergite

2.2—2.9 times its apical width.

VAN ACHTERBERG: Revision Zelinae auct. 275

The holotype of Zele rosenbergi Viereck (9, USNM, Washington:

“Chanchamayo, E. Peru”, “Collection Rosenberg”, “Type No. 13797. U.S.N.M.”,

“Zele rosenbergi Vier. Type, Q”’) is a typical fuscicornis specimen. The length of the

malar space is 0.5 times basal width of mandible; claws absent; vein Ir-m of hind

wing somewhat more curved than in holotype of fuscicornis.

Exasticolus nigriceps (Enderlein) comb. nov.

(figs. 105— 111, 116— 119)

Enderlein, (1918) 1920, Arch. Naturgesch. 84A (11): 217.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 225.

Holotype, 9, length of body 7.8, of fore wing 7.3 mm.

Head. — Antennal segments 46, 3rd segment 1.3 times 4th segment, length of

3rd and 4th segment 3.8 and 2.9 times their width, respectively, its penultimate

segments 1.8 and 2.3 times their width; length of maxillary palp 1.7 times height of

head; dorsal length of eye 2.9 times temple; POL : @ ocellus : OOL = 6: 8: 4; frons

concave, smooth; vertex convex, largely smooth, slightly coriaceous (fig. 116);

face rather flat and mainly, rather coarsely, transversely rugose (fig. 110); clypeus

convex, smooth, except for some punctures; malar space 0.3 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

smooth, except for some medial and apical rugae (fig. 105); epicnemial area

smooth, but anteriorly with some short rugae; mesopleuron superficially

punctulate; precoxal suture absent, except for a smooth, weak depression (fig.

105); notauli finely and densely crenulate (fig. 119); mesoscutal lobes superficially

punctulate; surface of propodeum mainly smooth, only submedially with an

irregular transverse carina and some rugae (fig. 105).

Wings. — Fore wing: r: 3-SR: SRI = 12: 12: 58; 1-CUI : 2-CUI = 2: 25; 2-SR:

3-SR : r-m = 14: 12: 7; 2A present as a short remnant (fig. 107). Hind wing: Ir-m

straight; short remnant of 2A present.

Legs. — Hind coxa smooth, except for some striae dorso-apically (fig. 108);

middle coxa without tubercle; hind claws absent; femur, tibia, and basitarsus of

hind leg 6.2, 9.9, and 10.0 times their width, respectively; length of spurs of hind

tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.9 times its apical width, its surface

irregularly, longitudinally striate, with a smooth tubercle apically (fig. 111); dorsal

carinae almost reaching apex of Ist tergite; 2nd tergite smooth, but superficially

pimply; length of ovipositor sheath 0.06 times fore wing.

Colour. — Brownish-yellow; head (except for mandibles), 3rd-11th antennal

segments and apical 0.7 of hind tibia, blackish-brown; scapus and pedicellus,

partly reddish-brown; hind tarsus and palpi rather whitish-yellow.

Holotype in PAN, Warsaw: “Mexico, Chiapas, L. Conradt S., 15—11—07”,

“Type”, “Zele nigriceps Enderl., 9, Type, Dr. Enderlein, det. 1918”, “Mus. Zool.

Polonicum, Warsawa 12/45”.

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Number of additional specimens examined: 9 9 and 3 Z. Variation: length of

body 7.1—7.5, of fore wing 6.6—7.5 mm, length of Ist tergite 2.5—2.9 times its

apical width; length of ovipositor sheath 0.07—0.08 times fore wing; antennal

segments 44—48; length of maxillary palp 1.5—2.1 times height of head; dorsal

carinae of Ist tergite sometimes absent or nearly so; length of malar space of 9

0.2—0.3 times basal width of mandible (in males 0.3—0.4 times).

Additional specimens examined from: Costa Rica (San Pedro de Montes de

Oca), Ecuador (Loja; Coca & Napo Rivers; Playas de Montalro), Peru (nr.

Marcapata, 30 m), Bolivia (Cochabamba, 17 km E. Villa Tunari), (British) Guyana

(Upper Courantyne R., King Frederick, William IV Falls), and Brazil (Nova

Teutonia, Santa Catarina; Villa Vera, 12°30’S, 50°31’W; Sinop, M. Grosso,

12°31’S, 55°37’W) (USNM, BM, IML, TC, RMNH). Only one specimen (from

Costa Rica) was reared, the host being Gloveria ballovi Schaus, and belonging to

the Lasiocampidae (Lepidoptera).

Genus Homolobus Foerster

Foerster, 1862, Verh. naturh. Ver. preuss. Rheinl. 19: 256.

Capek, 1969, Proc. ent. Soc. Wash. 71: 308.

Watanabe, 1969, id. 71: 318—325, figs. 1—7.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 142.

Shenefelt, 1970, id. 5(2): 220— 227.

Capek, 1970, Can. Ent. 102: 851, 853, 868, 869.

Marsh, 1971, Ann. ent. Soc. Am. 56: 847.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 180, 230, 231.

Capek, 1972, Ent. Problémy 10: 133, 136.

Capek, 1973, Acta Inst. forest. zvol.: 262.

Mason, 1973, Proc. ent. Soc. Wash. 75: 213.

Kabasinskaite & Jakimavicius, 1973, Acta ent. Lituanica 2: 80, 86.

Papp, 1974, Fol. Ent. Hung. 27: 125—129, fig. 1.

Jakimavicius, 1974, Tr. AN. Lit. SSR B, 2(66): 97.

Capek, 1975, Biolögia 30(11): 819.

Gauld & Huddleston, 1976, Entomologist’s Gaz. 27: 47.

Van Achterberg, 1976b, Tijdschr. Ent. 119: 37, 39, 44, 50, figs. 103, 104.

Jakimavicius, 1976, Tr. AN Lit. SSR B, 2(74): 90, 93, 95.

Tobias, 1976, Opr. Fauna SSSR 110: 31, 131, 133, fig. 39: 1—10.

Type-species: Phylax discolor Wesmael.

Synonyms: Zele auct. nec Curtis, 1832; Phylax Wesmael, 1835, nec Dahl, 1823;

Phylacter Reinhard, 1863 (nom. nov. for Phylax Wesmael); Apatia Enderlein,

(1918)1920. Syn. nov.

Diagnosis. — Length of body 4.4—14.6, of fore wing 4.6—15.9 mm; ventral

margin of clypeus rather thin, not (fig. 132) or distinctly (fig. 360) separated from

clypeus, smooth; eyes bare, large, weakly emarginate at inner sides (fig. 572) or

almost immarginate (fig. 525); metapleural flange large, more (fig. 327) or less (fig.

416) lamelliform; precoxal suture variable; antescutal depression medium-sized to

large, deep, with 1—3 longitudinal carinae (figs. 250, 331) or only crenulate (fig.

332), 3rd segment of labial palp medium-sized to small, length of 4th segment

VAN ACHTERBERG: Revision Zelinae auct. 277

1.6— 7.0 times 3rd segment (figs. 164, 265); epistomal suture complete (fig. 311);

ocelli medium-sized (fig. 213) to large (figs. 127, 275); pleural suture rather

narrowly and shallowly crenulate (fig. 616); metapleuron smooth (fig. 263) to

coarsely sculptured (fig. 527); episternal scrobe deep, medium-sized (figs. 633,

647); notauli complete, rather narrowly (fig. 473) to widely (fig. 413) impressed and

crenulate; scutellum smooth or punctulate, and convex; side of scutellum

crenulate (fig. 332), rugose (fig. 454) or striate (fig. 413); propodeum mainly

smooth (fig. 128), coarsely areolate (fig. 400), or extensively rugose (fig. 414);

antero-dorsal part of propodeum not (fig. 120) or distinctly (fig. 513) separated

from its posterior part; propodeal spiracle medium-sized to large, elliptical or

rather round (figs. 527, 541, 577); antepropodeal depression rather wide to

medium-sized (figs. 250, 286, 661); 1-SR+M of fore wing straight (figs. 130, 368);

1st discal cell of fore wing sessile (figs. 196, 484) or subpetiolate (figs. 469, 455); r of

hind wing present (fig. 122) or absent (fig. 147); SR of hind wing variable (figs. 184,

243, 368, 435); SC + RI of hind wing straight (fig. 425) to curved (fig. 343); 2-SC+R

of hind wing rather long (fig. 196), short and quadrate (figs. 266, 267) to vertical

(figs. 506, 507); Ir-m of hind wing straight (fig. 122); cu-a of fore wing long,

inclivous (fig. 130), straight (fig. 147) or somewhat curved basad (fig. 494);

parastigma large (figs. 122, 484) to medium-sized (fig. 603); 1A+2A and SRI of

fore wing straight (fig. 147) or 1A +2A (figs. 368, 369) and SRI (fig. 258) curved;

tarsi without a ventral row of setae; inner aspect of apex of hind tibia without a

comb of bristles; claws very variable, simple and without a subapical tooth (fig.

252), with subapical tooth (figs. 350, 351), with a ventral lamella (fig. 392), or with

an enlarged lamella (figs. 393, 394); length of hind femur 4.7—8.1 times its width;

length of Ist tergite 1.7—4.8 times its apical width; Ist tergite concave medio-

basally and more or less convex medially (figs. 489, 615); 2nd tergite with a sharp

lateral crease (fig. 401) or with a rounded fold (fig. 120); metasoma of 9

compressed apically (fig. 231); length of ovipositor sheath 0.04—0.79 times fore

wing.

Biology. — The numerous host records indicate that the species of Homolobus

are parasites of caterpillars with more or less exposed way of life, mainly belonging

to the families Noctuidae and Geometridae.

Distribution. — Cosmopolitan. Contains five subgenera: Apatia Enderlein,

Chartolobus subgen. nov., Homolobus Foerster, Phylacter Reinhard, and Oulophus

subgen. nov.

Subgenus Apatia Enderlein stat. nov.

Enderlein, (1918) 1920, Arch. Naturgesch. 84A(11): 219.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 142.

Type-species: Apatia simillima Enderlein (= Bracon truncator Say).

Diagnosis. — Length of body 4.4—9.0, of fore wing 4.6—8.7 mm; antennal

segments 38—54; its 3rd—6th segments without a ridge at the inner side (figs. 879,

880); length of outer aspect of 4th segment of labial palp 1.6—5.5 times 3rd

278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

segment; length of maxillary palp 1.0—1.4 times height of head; length of malar

space 0.4—1.6 times basal width of mandible; temples roundly narrowed apicad

(figs. 127, 163); length of hind femur 4.7—7.4 times its width; claws simple or

nearly so, without subapical tooth or lamella (figs. 123, 152, 212); apices of hind

tibial spurs of ¢ truncate and pigmented (figs. 112, 712) or sharp and hyaline (figs.

710, 713); inner hind claw of 9 convex or straight basally, equal to its outer claw

(figs. 202, 203, 881); IA +2A of fore wing straight; basal third of SR of hind wing

mainly pigmented, not sclerotized (figs. 122, 130), straight (fig. 147) or weakly

curved (fig. 161); SC+RI of hind wing straight (fig. 146) or weakly curved (fig.

194); r of hind wing exceptionally present (fig. 122); length of Ist tergite 2.0—3.6

times its apical width; 2nd tergite smooth; length of ovipositor sheath 0.04—0.26

times fore wing; posterior part of propodeum not separated from antero-dorsal

part (fig. 128).

Distribution. — With 11 out of 14 species (or 78.5%) occurring in the

Afrotropical region, this is the main centre of speciation of this subgenus. Two

species are extremely widely distributed: truncator occurs in the Holarctic,

Neotropical and Oriental regions, while ophioninus is found in the Afrotropical,

Palaearctic, and Australian regions.

Key to the species of the subgenus Apatia

1. Vein r of hind wing present, at least posteriorly (fig. 122); vein SR of hind wing

weakly curved (fig. 122); lateral aspect of hind tibial spurs of g sharp apically;

Oriental ea Ge ecient cae shat CR ar elagabalus (Nixon) (p. 280)

— Veinr of hind wing absent (fig. 130); vein SR of hind wing, and tibial spurs of

& variable... ette dea See ERO 2

2. First tergite black or dark brown, strongly contrasting with the, at least partly,

whitish 2nd and 3rd tergites of metasoma; propodeum mainly smooth (fig.

128); or vein SRI of fore wing straight (fig. 147) and precoxal suture mainly

smooth (MR. 14a Sn ar A I NE 3

— Basal tergites of metasoma brownish-yellow, if more or less dark brown, then

2nd tergite yellowish, dark brown, or blackish and less contrasting;

propodeum sculptured posteriorly (fig. 158); vein SRI of fore wing more or

less curved towards RI (figs. 161, 184), if straight, then precoxal suture

extensively sculptured (fig 204) EN 4

3. Malar space comparatively short (fig. 132), 0.4—0.6 times basal width of

mandible; mesoscutal lobes smooth (fig. 136); veins SRI of fore wing and SR

of hind wing curved (fig. 130); claws without small prominence subapically

(fig. 133); apices of hind tibial spurs of @ truncate; Australian ..........

ee wel RONN australiensis (Nixon) (p. 282)

— Malar space comparatively long (fig. 151), 0.9—1.4 times basal width of

mandible; mesoscutal lobes punctulate (fig. 157); veins SR1 of fore wing and

SR of hind wing straight (figs. 146, 147); claws with a minute subapical

prominence (figs. 152, 153); apices of hind tibial spurs of G sharp; Afrotropical

(Malagasy, Grande'Comiore)) er ee albipalpis (Granger) (p. 283)

VAN ACHTERBERG: Revision Zelinae auct. 279

Length of outer aspect of 4th segment of labial palp 3.0—5.0 times the small

3rd segment (figs. 200, 222, 235), if intermediate, then vein cu-a of fore wing

antefurcal (fig. 206) and/or apical half of metasoma mainly dark brown or

eels ksi Singer eh rss ia ee fer Nas ker pe 5

Length of outer aspect of 4th segment of labial palp 1.6—2.8 times the

medium-sized 3rd segment (figs. 246, 257, 265); vein cu-a of fore wing more or

less postfurcal (figs. 266, 329); metasoma mainly yellowish apically ....... 9

Tarsal claws of @ without any prominence (figs. 202, 203); vein SRI of fore

wing curved towards RI (figs. 161, 196); vein SR of hind wing more or less

sinuate (figs. 184, 196); Malagasy or non-Afrotropical .................. 6

Tarsal claws of 9 with a tiny rounded subapical prominence (figs. 212, 225);

veins SRI of fore wing and SR of hind wing straight or nearly so (figs. 206,

PBA fricamicontinent ion ar Sanat el Sa Re Va Bea 7

Length of 4th segment of labial palp ca. 4—5.5 times the 3rd segment (fig.

185); upper part of mesopleuron and mesoscutum smooth (figs. 182, 186);

veins r and 3-SR of fore wing of equal width (fig. 161); vein SR of hind wing

moderately sinuate (fig. 184) or almost straight; Holarctic, Neotropical,

Oriental ARE OST ee elt heated truncator (Say) (p. 285)

Length of 4th segment of labial palp ca. 3.0—3.5 times 3rd segment (fig. 200);

upper part of mesopleuron and mesoscutum punctulate or punctate (figs. 191,

201); vein r of fore wing wider than vein 3-SR (fig. 196); vein SR of hind wing

rather strongly sinuate (figs. 194, 196); Malagasy rufithorax (Granger) (p. 289)

Length of malar space 1.2—1.6 times basal width of mandible (fig. 210); apical

third of metasoma blackish or dark brown ...... maculatus spec. nov. (p. 291)

Length of malar space 0.7—1.0 times basal width of mandible (figs. 229, 238), if

intermediate, then apical third of metasoma yellowish .................. 8

Vein r of fore wing longer than 3-SR (fig. 219); subapical prominence of claws

of 9 very small, scarcely visible at 80x (figs. 225, 226); head, antenna and hind

leg mainly dark brown; palpi, tegulae, fore and middle coxae yellowish-white;

Ne! nern mate oad NI nimble ali alternipes spec. nov. (p. 292)

Vein r of fore wing shorter than 3-SR (fig. 234), exceptionally of equal length;

subapical prominence of claws of 9 small, but at 80x easily visible (fig. 237);

head, antenna, hind leg, palpi, tegulae, fore and middle coxae brownish-

VEIlOWESHATLIEA, krieg ya en Bow. priapus (Nixon) (p. 293)

Vein SR of hind wing strongly sinuate (figs. 243, 258); marginal cell of hind

wing distinctly narrowed medially in respect to its basal width (fig. 254);

middle lobe of mesoscutum finely and densely punctate or punctulate (figs.

250, 262); scapus more or less dark brown; vein SC+RI of hind wing

Bo mparativelysshort (figs2254, 259) u. an. va... Os OR. mi. aha: 10

Vein SR of hind wing weakly sinuate (fig. 266); marginal cell of hind wing not

or weakly constricted medially in respect to its basal width (fig. 290); middle

lobe of mesoscutum smooth or weakly punctulate (fig. 300); scapus mainly

yellowish; vein SC + RI of hind wing somewhat longer (figs. 267, 307).... 11

. Marginal cell of hind wing constricted just after middle of the cell (fig. 243);

length of ovipositor sheath 0.24—0.26 times fore wing, the exserted ovipositor

ID.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

longer than 1.5 times length of Ist tergite (fig. 240); propodeum and Ist tergite

irregularly sculptured (figs. 240, 253); C. Africa lacteiceps spec. nov. (p. 294)

Marginal cell of hind wing constricted in front of middle of the cell (fig. 258);

length of ovipositor sheath ca. 0.14 times fore wing, the exserted ovipositor

slightly longer than Ist tergite (fig. 280); propodeum and Ist tergite evenly,

finely and densely rugulose (figs. 142,255), 8: Africa

tli ER RENNER: pulchricornis (Nixon) (p. 296)

. Vein 2-SC+R of hind wing transverse, longer than wide (fig. 290); length of

hind femur 5.6— 7.2 times its maximum width, usually comparatively slender

(fig. 291), if intermediate, then upper condyli of mandibles rather far below

lower level of eyes, or first tergite more slender, longer than 2.2 times its apical

width (fig. 311); lateral aspect of hind tibial spurs of 3 more or less truncate

apically:(figs: 296,297)... ne an FORO ORI 12

Vein 2-SC+R of hind wing vertical or quadrate (fig. 267); length of hind femur

4.6-5.8 times its maximum width, comparatively stout (fig. 269); upper condyli

of mandibles comparatively close to lower level of eyes (fig. 270); Ist tergite

stout (fig. 271), its length 1.7—2.2 times its apical width; lateral aspect of hind

tibial spurs of & sharp apically (figs. 272,273) huddlestoni spec. nov. (p. 297)

Frontal aspect of head comparatively long, trapezoidal (figs. 311, 330); upper

condyli of mandibles of 9 distinctly below lower level of eyes (figs. 311, 330);

length of malar space 0.8—1.1 times basal width of mandible, if exceptionally

shorter, then claws setose basally (figs. 313, 314)...................... 13

Frontal aspect of head comparatively short, transverse (fig. 301); upper

condyli of mandibles of 9 close to lower level of eyes; length of malar space

0.3—0.7 times basal width of mandible; claws yellowish pectinate basally (fig.

QI TA eers Ge eM A TEE Cu A AN ophioninus (Vachal) (p. 298)

. Vein SC+RI of hind wing somewhat curved and shorter (figs. 306, 307);

marginal cell of hind wing usually less widened apicad, its apical width

1.9—2.2 times its maximum basal width (fig. 306); length of fore wing 3.5—7.1

mm; claws only setose or indistinctly pectinate basally (figs. 313, 314);

Ovipositor sheath in undistorted position rather wide apically (fig.

303): EE AL RE EEN EBEN Ar truncatoides spec. nov. (p. 300)

Vein SC+RI of hind wing almost straight and somewhat longer (figs. 329,

337); marginal cell of hind wing more widened apicad, its apical width 2.4—2.6

times its maximum basal width (fig. 329); length of fore wing 7.0—9.5 mm;

claws distinctly pectinate basally (figs. 339, 340); ovipositor sheath somewhat

moresslender (104339) MN a eee pallidistigmus (Cameron) (p. 303)

Homolobus (Apatia) elagabalus (Nixon) comb. nov.

(figs. 120— 127, 284, 332, 333)

Nixon, 1938, Bull. ent. Res. 29: 417, fig. If (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 223, 224.

Holotype, 9, length of body 6.3, of fore wing 6.6 mm.

VAN ACHTERBERG: Revision Zelinae auct. 281

Head. — Antennal segments 33, but apical segments missing (44 according to

original description), 3rd segment 1.1 times 4th segment, length of 3rd and 4th

segments 3.2 and 3.0 times their width, respectively; length of 4th labial palp

segment ca. 3 times 3rd segment (fig. 125); length of maxillary palp 1.2 times height

of head; inner sides of eyes weakly emarginate (fig. 126); dorsal length of eye 3.7

times temple; POL : @ ocellus : OOL = 7: 8: 5; frons almost flat, smooth; vertex

largely smooth, somewhat punctulate by insertions of the setae; face rather flat,

finely rugose-punctate, only laterally more coriaceous (fig. 126); clypeus flattened,

almost smooth, somewhat punctulate; apical margin of clypeus convex medially

and not distinctly separated from clypeus; length of malar space 0.5 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.2 times its height; side of pronotum

almost smooth, except for some short crenulae medially and rugosity posteriorly

(fig. 120); epicnemial area largely smooth, somewhat superficially rugose near

subalar pit (fig. 120); precoxal suture scarcely impressed, superficially reticulate-

punctate; rest of mesopleuron mainly smooth; metapleural flange wide,

lamelliform, and rounded apically (fig. 120); metapleuron largely smooth,

reticulate-carinate ventrally; notauli rather narrow (fig. 332); mesoscutal lobes

smooth, except for some punctulation; surface of propodeum mainly smooth

anteriorly, with a short irregular medial carina, medially and posteriorly

transversely rugose (fig. 120).

Wings. — Fore wing: r: 3-SR: SR 1 = 10: 13: 60; SRI curved anteriad; cu-a

slightly inclivous and apically curved basad (fig. 122); 1-CUI : 2-CU1 = 2: 24; 2-

SR : 3-SR : r-m = 11 : 13 : 7; 2A well developed (fig. 122); area basally of 2A

remotely setose. Hind wing: r present, dividing the marginal cell into two subequal

parts; SC + Rl] and base of SR curved.

Legs. — Hind legs absent; fore and middle claws pectinate (figs. 123, 124);

length of middle tibial spurs 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.7 times its apical width, its surface smooth

(fig. 333); dorsal carinae of Ist tergite absent; length of ovipositor sheath 0.6 times

fore wing.

Colour. — Brownish-yellow; antenna (but apically lighter) and stemmaticum,

dark brown; pterostigma more transparent yellowish.

Holotype in BM, London: “Type”, “B.M. Type, Hym., 3.c.678”, “Zele

elagabalus Nixon, Type, 9, 1938”, “1938/16 slide”, “3836”, “pres. by Imp. Inst.

Ent. BM. 1939-92”, “Parasite on Selepa celtis”, “Dehra Dun, U.P., 26.x1.1935”,

“SNC Expt. No. 1294”. Paratypes: 10 3, belonging to the reared series from which

the holotype was selected. The tibial spurs are sharp and hyaline apically, number

of antennal segments 41—42, anterior half of vein r of hind wing absent; vertex

and temples punctulate; area basally of 2A of fore wing mainly bare, and precoxal

suture somewhat more rugose-striate than in holotype.

Additional specimens examined (7 ©) from Thailand (Bangkok, ex caterpillar

feeding on Sandoricum indicum) and Philippines (Manila) (BM, TMA, RMNH).

Variation: vertex and temple punctulate or finely punctate; length of malar space

0.4—0.5 times basal width of mandible; length of fore wing 5.6— 7.5 mm; length of

282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

ovipositor sheath 0.04—0.08 times fore wing. The 9 from Manila is slightly

aberrant: hind leg somewhat darkened, pterostigma and metasoma mainly dark

brown, and left hind wing with a short vein m-cu (fig. 284).

Homolobus (Apatia) australiensis (Nixon) comb. nov.

(figs. 128— 137)

Nixon, 1938, Bull. ent. Res. 29: 419 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

Holotype, 9, length of body 6.5, of fore wing 6.7 mm.

Head. — Antennal segments 37, but apical segments missing (according to

original description 46), 3rd segment 1.1 times 4th segment, length of 3rd and 4th

segments 3.6 and 3.2 times their width, respectively; length of 4th labial palp

segment 2.9 times 3rd segment (fig. 131); length of maxillary palp 1.2 times height

of head; inner sides of eyes weakly emarginate (fig. 132); dorsal length of eye 2.3

times temple; POL : @ ocellus : OOL = 6: 7: 6; frons rather flat, largely smooth,

with some striae near anterior ocellus and punctulate laterally (fig. 134); vertex

mainly smooth, except for some punctulation; face and clypeus flattened,

punctulate (fig. 132); apical margin of clypeus weakly convex and not distinctly

separated from clypeus; length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, but with some crenulae medially and apically (fig. 128); epicnemial area

smooth, but somewhat punctulate anteriorly; precoxal suture scarcely impressed,

rugose-punctate medially and indistinctly punctate anteriorly and posteriorly (fig.

128); rest of mesopleuron smooth; metapleural flange large, lamelliform and

truncate apically (fig. 128); metapleuron smooth, except for some carinae

ventrally; notauli medium-sized, strongly crenulate (fig. 136); mesoscutal lobes

smooth; surface of propodeum smooth, except for some indistinctly developed

rugosity posteriorly, without carinae (fig. 128).

Wings. — Fore wing: r : 3-SR : SRI = 10: 17: 74; SRI curved anteriad; cu-a

inclivous and apically curved basad (fig. 130); 1-CU1 : 2-CU1 = 1: 10; 2-SR : 3-SR

: r-m = ca. 14: 17: 10; 2A well developed (fig. 130); area basally of 2A remotely

setose. Hind wing: r absent; SC + Rl and SR weakly curved.

Legs. — Hind coxa smooth; claws simple, long setose and with some bristly

setae basally (fig. 133); length of femur, tibia, and basitarsus of hind leg 7.4, 11.5,

and 10.8 times their width, respectively; length of spurs of hind tibia 0.6 and 0.5

times basitarsus; outer side of hind tibia rather spiny (fig. 137).

Metasoma. — Length of Ist tergite 3.0 times its apical width, its surface smooth,

except for some punctulation; dorsal carinae of Ist tergite absent, except for a

vague remnant basally (fig. 135); length of ovipositor sheath 0.07 times fore wing.

Colour. — Brownish-yellow; antenna (except main part of scapus and

pedicellus), stemmaticum, hind leg and metasoma, blackish-brown, but 2nd tergite

laterally, 3rd tergite laterally and apically, metasoma baso-ventrally, apex of

hypopygium and ovipositor sheath, whitish-yellow; pterostigma, C+SC+R and

RI of fore wing, brown; wing membrane weakly infuscated.

VAN ACHTERBERG: Revision Zelinae auct. 283

Holotype in BM, London: “Type”, “B.M., Type, Hym., 3.c.680”. “Zele

australiensis Nixon, holotype, © ”, “MacKay, Queensland, 1909—45”, “MacKay,

2.94”, “912”. Total number of additional specimens examined: 27 © and 19 4, all

from Australia. Queensland: Bluff Range, nr. Biggenden; Mt. Crosby; Mt.

Cootha; Mt. Tamborine; Brisbane. New South Wales: Woodford; Willowtree; |

mi. W. Wombeyan Cayes; 2 mi. N. Boonoo Boonoo, nr. Tentifield; Mt. Brown.

Western Australia: Drummonds Cove, nr. Geraldton; Jacup; 19 mi. ENE. Perth.

Northern Territories: McArthur River, 14 km SW. Cape Crawford. Australian

Capital Territory: Blundell’s F.C.T.; Handmarsh Falls, at light; Canberra.

Southern Australia: Cape Jervis. Victoria: Nowa Nowa; Mt. Hotham. Tasmania:

Mt. Barrow; Coles Bay; Togari; Port Davey; Port Arthur (TC, BM, CNC,

RMNH, CSIRO).

Variation: Length of ovipositor sheath 0.06—0.07 times fore wing; length of

malar space 0.4—0.6 times basal width of mandible; length of fore wing 6.3—6.7

mm; length of 4th segment of labial palp 2.3—2.6 times 3rd segment; apices of

hind tibial spurs of G truncate and pigmented apically; middle of propodeum

finely rugose or rather coarsely sculptured; wing membrane rather hyaline to dark

brown; sometimes middle legs, propodeum and metanotum, blackish; 2nd and 3rd

tergites often mainly whitish-yellow, except for a narrow blackish longitudinal

stripe at middle of 2nd tergite, but exceptionally absent; head colour varies from

mainly brownish-yellow to dark brown or black. The (in the genus Homolobus)

unusual blackish/whitish colour markings may indicate that australiensis belongs to

a mimetic complex of species. The same colour pattern occurs in Homolobus also

in species from Malagasy, which are not closely related to australiensis.

Homolobus (Apatia) albipalpis (Granger) comb. nov.

(figs. 144—157)

Granger, 1949, Mém. Inst. scient. Madagascar 2A: 378, fig. 384 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

Redescribed after a 9 from Ranomafana (Malagasy), length of body 4.4, of fore

wing 4.6 mm.

Head. — Antennal segments 40, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 3.7 and 3.1 times their width, respectively, length of both

penultimate segments 1.6 and 1.8 times their width; length of 4th segment of labial

palp 4.0 times 3rd segment (fig. 154); length of maxillary palp 4.0 times height of

head; inner sides of eyes weakly emarginate (fig. 151); dorsal length of eye 2.0

times temple; POL : @ ocellus : OOL = 9: 9: 12; frons rather flat, with some striae

and punctulate laterally; vertex punctulate, rather flat; face weakly convex and

punctate; clypeus convex, remotely punctate; apical margin of clypeus rather

straight medially and not distinctly separated from clypeus (fig. 151); length of

malar space 1.1 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

almost smooth dorsally and ventrally, medio-anteriorly crenulate and posteriorly

284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

rugose (fig. 144); epicnemial area mainly smooth; precoxal suture absent, except

for a shallow depression, weakly punctulate, as rest of mesopleuron; metapleural

flange large, stout, rounded apically, bordered by a narrow carina; metapleuron

smooth, but with some rugae ventrally; notauli deep and widely crenulate

posteriorly, narrowed anteriad (fig. 157); mesoscutal lobes punctulate; anterior

surface of propodeum mainly smooth, but posterior half weakly reticulate-rugose

(fig. 144).

Wings. — Fore wing: r : 3-SR : SRI = 11 : 12: 78; SRI straight; cu-a almost

straight (fig. 147); 1-CUl : 2-CUl = 3: 31; 2-SR : 3-SR : r-m = 17: 12: 11; 2A

shortly developed (fig. 148); area basally of 2A mainly bare. Hind wing: r absent;

SR straight; SC + R1 weakly curved (fig. 146). .

Legs. — Hind coxa finely punctate and dorso-apically shortly striate; tarsal

claws with a scarcely visible subapical prominence (figs. 152, 153), setose basally;

length of femur, tibia, and basitarsus of hind leg 5.6, 9.8, and 7.2 times their width,

respectively; length of spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.3 times its apical width, somewhat

narrowed apically, its surface smooth (fig. 156); dorsal carinae of Ist tergite

absent; length of ovipositor sheath 0.11 times fore wing.

Colour. — Yellowish-brown; stemmaticum, metasoma (except for the whitish

2nd and 3rd segments and Ist sternite), black; 2nd and 3rd tergites yellowish-

white, but apical margin of 3rd tergite narrowly blackish; 1st-3rd segments of

labial palp, Ist and 2nd segments of maxillary palp, dark brown; 4th segment of

labial palp and 3rd-6th segments of maxillary palp, whitish-yellow.

Redescribed after © from Malagasy, Ranomafana, X.1938 (MNHN, Paris).

Holotype in MNHN, Paris, &: “Madagascar, Ankaratra, Alt. 1800 m”, “Museum

Paris, 11.38, A. Seyrig”, “Type”, “Zele albipalpis Gr., B. Sigwalt”. Antennal

segments 47; length of malar space 1.4 times basal width of mandible; length of

fore wing and of body both 6.4 mm; length of maxillary palp 1.5 times height of

head; pronotum and propodeum somewhat more sculptured than in figured

specimen; length of Ist tergite 2.7 times its apical width; apices of spurs of hind

tibia sharp and hyaline; whole 3rd tergite whitish; 2nd and 3rd segments of labial

palp whitish; head, all coxae, fore and middle tarsi, propleuron and pronotum

ventrally, infuscated to dark brown; hind tarsus yellowish.

Additional specimens examined from Malagasy and Grande Comore, 89 and |

& (Malagasy: Mandraka; Manjakatampo; Ankaratra-Antsasbatana, sous forêt,

0—5 h, 1970 m; Sakavondro, 40 m, Fort Dauphin; Andranotobaka, 1400 m,

Ambatolampy. Grande Comore: Convalescence, 1700 m; Nioumbadjoe, 505 m).

Variation: Length of fore wing 4.1—7.2, of body 3.6—6.8 mm; length of ovipositor

sheath 0.11—0.13 times fore wing; length of malar space 0.9—1.4 times basal width

of mandible. Length of 4th segment of labial palp 4.0—5.0 times 3rd segment;

length of Ist tergite 2.1—2.8 times its apical width; antennal segments 38—47;

colour of 3rd tergite completely white to mainly black, palpi mainly white,

yellowish, or wholly dark brown, except the 6th segment of maxillary palp

(MNHN, RMNH, MAC).

Note. In addition I have examined an aberrant series of 5 melanistic 9

VAN ACHTERBERG: Revision Zelinae auct. 285

specimens from Andranotobaka (Malagasy, 1400 m, Ambatolampy, MNHN,

RMNH). The 2nd and 3rd tergites are mainly blackish, at most are the margins

more or less whitish-yellow, the hind leg (except for the coxa and trochanter) and

palpi are mainly dark brown, and length of malar space 1.0—1.2 times basal width

of mandible. In the key they may run to maculatus, but maculatus differs by the

colour, the finely sculptured propodeum, and the longer malar space.

This species may be confused with the other almost similarly coloured species

from Malagasy, viz., cingulatus and inopinus. But the females of the latter two

species have an antennal ridge, the inner hind claw concave basally and the

precoxal suture more extensively sculptured. The males are less easy to separate,

but the posteriorly scarcely sculptured precoxal suture, the mainly white 3rd

tergite and usually whitish apical segments of the palpi of albipalpis may be

sufficient to separate them from cingulatus and inopinus.

Homolobus (Apatia) truncator (Say) comb. nov.

(figs. 112—115, 158—168, 174— 176, 178—190, 711, 879—881)

Say, 1828, Contrib. Maclur. Lyc. Philad. 2: 381 (as Bracon).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 227.

Wesmael, 1835, Nouv. Mém. Acad. Brux. 9: 161 (Phylax calcarator). Syn. nov.

Cresson, 1872, Trans. Am. ent. Soc. 4: 178 (Phylax melleus). Syn. nov.

Viereck, 1905, Trans. Kans. Acad. Sci. 19: 279 (Zele crassicalcaratus). Syn. nov.

Bengtsson, 1918, Acta Univ. Lund. (2)14(32): 42, fig. 18 (Phylacter fuscitarsis), Syn. nov.

Enderlein, (1918) 1920, Arch. Naturgesch. 84A: 218, fig. '0 (Zele unicolor). Syn. nov.

Enderlein, (1918) 1920, id. 84A: 219 (Apatia simillima). Syn. nov.

Nixon, 1938, Bull. ent. Res. 29: 420, fig. 1d (Zele chlorophthalma (nec Spinola, 1808!). Syn. nov.

Papp, 1971, Acta zool. Acad. Sc. hung. 17: 53.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 230.

Capek, 1975, Biologia 30: 819.

Tobias, 1976, Opr. Fauna SSSR 110: 131, fig. 39: 2—4.

Neotype, ©, length of body 6.8, of fore wing 6.3 mm.

Head. — Antennal segments 50, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 3.2 and 2.8 times their width, respectively, penultimate

segments 1.9 and 2.3 times their width (fig. 162); length of 4th segment of labial

palp ca. 4 times 3rd segment (fig. 164); length of maxillary palp 1.3 times height of

head; eyes weakly emarginate at the inner sides (fig. 165); dorsal length of eye 1.6

times temple; POL : @ ocellus : OOL = 12: 13: 16; frons almost flat, with some

superficial striae near antennal sockets (fig. 163); vertex smooth, but with some

microstriae near ocelli; face rather flat, superficially rugose below the antennal

sockets and punctulate ventrally; clypeus rather flat, punctulate; apical margin of

clypeus scarcely separated from clypeus, slightly convex medially (fig. 165); length

of malar space 0.5 times basal width of mandible; mandible rather weakly twisted

apically (fig. 165).

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely smooth, medially widely and posteriorly more narrowly crenulate (fig.

158); epicnemial area rugose-punctate; precoxal suture rather coarsely rugose-

punctate (fig. 158); rest of mesopleuron smooth; metapleural flange rather large,

286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

lamelliform, truncate apically and wide; metapleuron largely smooth, only rugose

ventrally; notauli deep, rather widely crenulate (fig. 168); mesoscutal lobes

smooth; surface of propodeum narrowly smooth anteriorly and with a short medial

irregular carina, medially and posteriorly reticulate-rugose (fig. 158).

Wings. — Fore wing: r: 3-SR : SRI = 8: 11:52; SRI somewhat curved anteriad

(fig. 161); cu-a slightly inclivous and curved; 1-CUI : 2-CU 1 = 2: 21; 2-SR: 3-SR:

r-m = 10: 11:7; 2A mainly present as an only pigmented brownish stripe (fig.

161); area basally of 2A mainly bare. Hind wing: r absent; SR and SC + RI weakly

curved (fig. 161).

Legs. — Hind coxa smooth, except for some microsculpture (fig. 167); tarsal

claws simple, setose basally, rather bristly (fig. 160); length of femur, tibia, and

basitarsus of hind leg 6.0, 9.5, and 8.5 times their width, respectively; length of

spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.2 times its apical width, its surface rather

superficially and irregularly punctate-rugose (fig. 166); dorsal carinae of Ist tergite

weakly developed in basal half of tergite; length of ovipositor sheath 0.07 times:

fore wing.

Colour. — Brownish-yellow; stemmaticum black; flagellum of antenna dark

brown; palpi and ovipositor sheath rather whitish-yellow.

Neotype in RMNH, Leiden: “U.S.A., Mich., Ann Arbor, 25—30.VIII.1976,

Malaise-traps, C. van Achterberg”.

Note. Males are very similar to the females, but the apices of the spurs of the

hind tibia are truncate and pigmented apically (fig. 112). Known hosts of examined

specimens belong to the Noctuidae (Amathes smithii (Snellen), Porosagrotis

orthogonia (Morrison), and P. tristicula (Morrison)) and to the Geometridae

(Hypagyrtis piniata (Packard), Lycia zonaria (Denis & Schiff.), and Semiothisa

bitactata (Walker)).

Total number of specimens examined: 324 © and 223 &. From the following

localities in the New World: British Columbia (3 mi. E. Lytton, 800 ft), New

Brunswick (Charlotte Co.), Nova Scotia (Mount Uniacke), Quebec (Kazubazua;

Queen’s Park, Aylmer; Wright; Kirks Ferry, light trap; Knigsmerg), Ontario

(Chaffeys Locks; Bothwell; Ottawa; Chalk River; Rondeau Park; Leamington;

Stratbury; Aylmer West, Malaise-trap), Saskatchewan (Beverley; 2 mi. Scout

Lake, 2600 ft; Sceptoe), Alberta (Brooks, Lethbridge; Rotlaw; Duchess),

Manitoba (2 mi. W. Stockton), Rhode Island (Westerly), New York (Ithaca, 6 Mile

Creek; Oneonta; Otsego L.; Troy; Poughkeepsie; Farmingdale; Elmire), New

Jersey (Moorestown), Michigan (George Res., Livingston Co.; Oakland Co.; Ann

Arbor, Malaise-trap; Crystal Falls, Iron Co.), Connecticut (Canterbury), Kansas

(Lawrence; Elwood; Douglas Co.), Kentucky (Golden Pond, Malaise-trap),

Louisiana (Rapides Parish; Shreveport; Bayon Chicot, Evangeline Co.), Montana

(Big Spring State Park; Williamsville, Malaise-trap), North Carolina (Black Mts.;

Murfreesboro; Faison; Raleigh; Wake Co.), South Carolina (Columbia;

Greenville; McClellanville; Table Rock; Wattacoo, Pickers Co.), Indiana (Posey

Co., Murphy’s Park, New Harmony), Arkansas (Hope), Maryland (Plummers

Island; Prince Georges Co., Beltsville; Takoma Pk.; Patuxent Ref., Bowie;

VAN ACHTERBERG: Revision Zelinae auct. 287

Laurel), Virginia (Galax; Charlottesville), Missouri (Cp. Shelby, nr Hattiesburg;

Mamon Co., Monroe City), Georgia (Forsyth; Waycross), Illinois (Wheaton,

Dupage Co.); Nebraska (Valentine Refuge; Thomas Co., Neb. Nat. Forest, 2.5 mi.

W. Halsey; 5 mi. NW. Harrison, 4400 ft), Colorado (Boulder; 6 mi. SE. Maybell,

6200 ft), Minnesota (Moorhead, Clay Co.; Big Fork), Wyoming (Douglass;

Sweetwater Co., 11.5 mi. S. Eden; 6 mi. N. Sage), Tennessee (Knoxville), Oregon

(Jackson Co., Mt. Ashland, 6500— 7000 ft), Idaho (Butte Co., Crater of Moon),

Florida (Subtrop. Exp. Sta., Homestead; Miami; Gainesville, Miachua Co., black

light; Lake Placid; De Funiah Spring; Highlands Co., Archbold Biol. Stat.; Tall

Timbers; Dunedin), California (Murray Kings Co., U.V. light trap; Walliston;

Potrero, S.D. Co.; Julian; Cedar Pass, 6000 ft, Warner Mts., Modoc Co.;

Cuyamaca; Andreas Cyn., Palm Springs; Orinda Village, Contra Costa Co., San

Pablo Ridge, below Eureka Peak, 1000—1200 ft, oak-chaparral zone; 6 mi. E.

Coalinga, Fresmo Co., U.V. light trap; Boyd Desert Res. Center, 4 mi. S. Palm

Desert, Rw. Co.; 9 mi. W. Lone Pine, Inyo Co.; Kern Co.; San Ignacio L.;

Catarina L.; Riverside Co., Agua Caliente, Ind. Res., Palm Canyon), Nevada

(Austin Summit, Lander Co., at black & white light; Washoe Co., 21 mi. SE.

Eagleville (Cal.); Golconda), New Mexico (Hatch, Raton, 6660 ft, Colfax Co.),

Texas (San Antonio, at light; Brownwood; Kerville; Ft. Davis, Limpia Cyn., 5000

ft; Big Band N. P., Panther Junction, 3500—4000 ft), Arizona (Baboquivari Mts.;

SW. Research Sta. of AMNH, Cave Creek Cyn., 5400 ft, Chiricahua Mts., Cochise

Co.; Indian Wash nr Martinez Lake, Yuma Co., at light; Tuczon; Pima Co., Organ

Pipe Cactus Nat. Mon., Williams Springs, flight trap; nr Roosevelt L.; Cochise

Co., Pinery Cyn., Chiricahua Mts., Portal; Ajo; Huachuca Mts., Sierra Vista; 3 mi.

W. Eager, 7100 ft, Pinon-Juniper zone; Ramsey Cyn., 6000 ft, 15 mi. S. Sierra

Vista, Huachuca Mts.), Mexico (Vera Cruz, Fortin; Baja California, Norte,

Diablito Cyn., East face Sierra San Pedro Martir, at light; id., Ensenada, at light;

id., 7 mi. W. Las Arrastras de Arriola; id., Bajada, 8 mi. E. of Ojos Negros, at light;

20 mi. E. Guasave, Sin.; Santa Clara Cyn., 5 mi. W. Parrita, Chih.; 2 mi. W.

Tlaxcala, Tlaxcala, swept from alfalfa; Chiap., Suchiapa; Oax., 4 mi. SE. Oaxaca;

20 mi. NNW. Obregon, Son., at light; Citlaltepetl, V. Cruz, 3000 ft; 10 mi. W. Vera

Cruz; Sonora, Bahia San Carlos; id., 83 mi. W. Sonoyta; Sinaloa, 25 mi. E. of Los

Mochis; Linares, N.L.; Teotihuacan, Pyr. Mex.; 15 km E. Sombrerete, Zac., at

light; Fortin de las Flores, 3400 ft, Malaise-trap; Boquillas del Carmen, Coah.,

1850 ft; 30 mi. SW. Tehuacan, Pue, 6800 ft; Chipinque Mesa, 5400 ft, nr.

Monterrey, N.L.; nr. Jame, 7500 ft, 31 mi. SE. Saltillo, Coah.; 5 mi. S. Monterrey,

N.L.; Dgo, 3 mi. E. El Salto, 8500 ft; Sin., 20 mi. E. Concordia, 3000 ft; Dgo, 24 mi.

W. La Cuidad, 7000 ft; Dgo, 6 mi. S. Durango, 6100 ft; Oax., El Paredon; Lake

Catemaco, Ver.; Dgo, 10 mi. W. El Salto, 9000 ft; Atlacomulco, 8500 ft; Orizaba,

Ver.), El Salvador (Quezaltepeque, 500 m), Panama (Cerro Punta, Chiriqui, 6500

ft), Costa Rica (Monte Verde), Guatemala (Sacapulas, 4500 ft), Cuba (Soledad,

Cienfuegos; Cuabitas, Stgo. de Cuba, Ote), and Venezuela (Cagua Edo, Aragua, at

light) (RMNH, CAS, UCA, TC, USNM, AMNH, CNC, PAC, PAN, MSU, ANSP).

From the Palaearctic and Oriental regions: Finland (Dickursby), Sweden

(Lund), Denmark (Emelsbu, Sgderjylland; Sondbg; Charlottenlund), East

288 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Germany (Thüringen; Berlin), West Germany (Rheingau, Gusenheim; Föhr;

Moosburg; Wurzburg; Garmisch, Ober-Bayern, Ettuler berg, ca. 700 m; id.,

Murnau, 700 m), Netherlands (Amsterdam; Kasteel Neercanne, St. Pietersberg, at

light), Belgium (nr. Charleroy), Czechoslovakia (Karlatejn; Pavlovake Kopce,

Bolni Véstonice), Poland (Gdansk), USSR (Moscow; Karagand, S. Tsj.-Arka,

pojma, Taldi-Manaka; Karagandinsk obl., 20 km W. Karkaralins; Sveuciouiliym,

Lit. SSR), Switzerland (Wallis), Austria (Sud-Tirol, San Martino di Castroza, 1444

m; Sansal Gebirge, 300—500 m, Styria; Nord-Tirol, Oberinntal, Kauns nr. Prutz,

1000— 1400 m), France (Cestas, Gironde; Vallon Pont d'Arc, Ardèche; Grignon;

Crimaud, Var), Jugoslavia (Hercegovina, Buna), Bulgaria (Mandrisa, Rodopi;

Karamansi, Rodopi), Hungary (Crepel; Baranya-Megya, Nagyharsany), Romania:

(Cibinsgeb., Transsylv. Alp.; Dannehl, id.), Spain (15 km NW. Tarifa; Albaracia),

Cyprus (Yermosoyia River; Skouili, Sapho Dist., nr. Limasol), Tunesia (Nefta),

Turkey (Priene, Asia Minor), S. India (Shevaroy Hills, 4500 ft, Yercaud),

Philippines (Baguio, Benguet), and Taiwan (Taihorinsho) (RMNH, MNHN, TC,

CNC, WHC, UZM, ITZ, HC, ZMH, ZMA, TMA, IZP, ZSB, BM).

Notes. The interpretation of this enigmatic species has long remained uncertain,

e.g., Muesebeck & Walkley (1951: 109) listed truncator as an “unrecognized

species” in Zele auct. The original description is very short, but is sufficient for

identification, despite the fact that the type is lost. The “much compressed,

truncate” metasoma, combined with the “body pale honey-yellow, polished,

impunctured” points to the genus Homolobus Foerster. The most commonly

captured species of Homolobus in the faunal area wherein the type-locality is

situated is H. melleus (Cresson, 1872), a junior synonym of H. calcarator (Wesmael,

1835). It is also the only species of this genus in the area that fits the original

description, e.g., antenna fuscous, but honey-yellow at base (viz., scapus is

yellowish), palpi whitish towards the tips (viz., the maxillary palp is often whitish

apically), the propodeum “slightly punctured” (which is actually finely rugose, but

in any case not areolated as in other species), and length of body, ca. 6 mm. Finally

in the AMNH collection (New York) there is a damaged specimen with an old,

handwritten label “Zele (Bracon) truncatus Say” (probably by Ashmead) which

belongs to this species. For the fixation of this interpretation I have selected a

neotype, which will be deposited in the RMNH-collection (Leiden). The original

type-locality is “Indiana”, while the neotype is from Ann Arbor, Michigan, but

both are situated in the Carolinian faunal region.

The holotype of Phylax calcarator Wesmael, 1835, was collected nr. Charleroy,

Belgium. According to the original description Wesmael had only one male at his

disposal, but in the Wesmael collection four specimens with type-labels are

present. Fortunately only one fits the original description, in possessing the back

of the metasoma ”entièrement d’un testacé comme le reste du corps” (Wesmael,

1835: 161). This specimen has been labelled holotype: the length of the body is 6.3,

of fore wing 5.6 mm, antennal segments 43, length of malar space 0.6 times basal

width of mandible; hind tibial spurs truncate apically, length of femur, tibia and

basitarsus of hind leg 4.9, 9.8, and 8.0 times their width, respectively, and length of

Ist tergite 2.9 times its apical width (KBIN, Brussels: “Coll. Wesmael”, “1875”,

“Phylax calcarator mihi, &, det. C. Wesmael”, “Type”).

VAN ACHTERBERG: Revision Zelinae auct. 289

The lectotype of Phylax melleus Cresson, 1872 (ANSP, Philadelphia: “Tex.”,

“Type, No. 1763”, ““Phylax melleus Cress.’’) is damaged, viz., the metasoma is

missing. The length of the fore wing is 6.5 mm, antennal segments 50, length of

maxillary palp 1.2 times height of head, and femur, tibia, and basitarsus of hind leg

6.1, 10.5, and 9.5 times their width, respectively. There are two specimens in ANSP

with the same printed locality-label, which have been labelled paralectotype.

The holotype of Zele crassicalcaratus Viereck, 1905 (SEM, Lawrence: “Aug.”,

“Douglas Co., Kansas, E. S. Tucker”, “Zele crassicalcaratus Vier., Type”, ‘‘617”’) is

a typical male of truncator (figs. 174—176, 178—181). The metasoma, hind tibia

and tarsus are absent, length of fore wing 5.3 mm, antennal segments 47, and

length of hind femur 5.5 times its width.

Finally Phylacter fuscitarsis Bengtsson, 1918, and Zele unicolor Enderlein, 1920,

have to be added to the synonyms of truncator, because no significant differences

could be detected after a thorough study of the available material. In the Thomson

Collection (Lund) are 7 specimens under Phylacter chlorophthalmus (sensu

Thomson, nec Spinola), which are part of the type-series of Phylacter fuscitarsis

Bengtsson, because in the original description the name fuscitarsis was proposed

for the species interpreted by Thomson as chlorophthalmus. Therefore one 9 (ZIL,

Lund: “Ilsp, 9/7”, “1977, 30”) is herewith selected as lectotype of Phylacter

fuscitarsis Bengtsson. The length of the fore wing is 5.7 mm, r : 2-SR : 3-SR : r-m =

10: ca. 13: 16: ca. 9, and the telotarsi are somewhat infuscated. Paralectotypes are

3 9,2 3, and one damaged specimen, of which the sex is unknown.

The examined type-series of unicolor (lectotype: figs. 112—115, 182—190)

consists of 23 © and 4 &. One © is herewith selected as lectotype (PAN, Warsaw:

“Costa Rica, H. Schmidt S.”, “Type”, “Zele unicolor Enderl., 9, Type, Dr.

Enderlein, det. 1918”, “Mus. Zool. Polonicum, Warszawa, 12/45”), which is a

quite normal specimen of truncator. Length of body 6.4, of fore wing 6.1 mm,

length of malar space 0.6 times basal width of mandible, length of maxillary palp

1.1 times height of head, SR of hind wing weakly sinuate, length of femur, tibia,

and basitarsus of hind leg 6.9, 10.3, and 8.8 times their width, respectively, length

of Ist tergite 2.5 times its apical width, and length of ovipositor sheath 0.08 times

fore wing. The examined holotype of Apatia simillima Enderlein, 1920 (chosen from

the same series as unicolor) is also a truncator, but has as an additional feature

the well developed notauli filled with glue (PAN, Warsaw).

Variation: length of body 5.8—8.1, of fore wing 5.3—8.2 mm; length of hind

femur 4.9—7.3 times its width; length of Ist tergite 2.5—3.4 times its apical width;

length of ovipositor sheath 0.06—0.08 times fore wing; antennal segments 44— 54;

length of maxillary palp 1.1—1.3 times height of head; length of malar space

0.5—0.7 times basal width of mandible; length of 4th segment of labial palp

4.0—5.5 times 3rd segment.

Homolobus (Apatia) rufithorax (Granger) comb. nov.

(figs. 191—203, 319, 320)

Granger, 1949, Mém. Inst. scient. Madagascar 2A: 377, 378, fig. 383 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 226.

290 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Lectotype, 9, length of body 9.0, of fore wing 8.7 mm.

Head. — Antennal segments 50, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 3.8 and 3.4 times their width, respectively, both penultimate _

segments 2.0 and 2.4 times their width; length of 4th segment of labial palp 3.5

times 3rd segment; length of maxillary palp 1.2 times height of head; eyes

distinctly emarginate (fig. 198); dorsal length of eye 2.0 times temple; POL : g

ocellus : OOL = 9: 13: 12; frons almost flat, with a weakly developed medial

carina (fig. 199); vertex punctulate; face and clypeus rather flat, punctulate; apical

margin of clypeus straight medially, not separated from the clypeus (fig. 198);

length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

crenulate medially, rugose-punctate medio-ventrally, punctate dorsally, and

rugose posteriorly (fig. 191); epicnemial area mainly rugose; precoxal suture

widely reticulate-rugose, surrounding part of mesopleuron punctate; metapleural

flange large, rounded and without carina apically; metapleuron weakly punctate

dorsally, rugose ventrally; notauli narrowly crenulate anteriorly, more widely

posteriorly, deep (fig. 201); mesoscutal lobes densely punctulate; surface of

propodeum coarsely rugose-reticulate, without carinae (fig. 191).

Wings. — Fore wing: r : 3-SR : SRI = 7: 14: 43; r wider than 3-SR; SRI

strongly curved towards RI (fig. 196); cu-a slightly inclivous, almost straight, and

shortly antefurcal; 2-M + CUI : CUI = 1: 23; 2-SR : 3-SR : r-m = 11: 14: 7; 2A

shortly developed; area basally of 2A remotely setose laterally (fig. 197). Hind

wing: r absent; SR sinuate (fig. 196); SC + R1 moderately curved (fig. 194).

Legs. — Hind coxa weakly punctate; tarsal claws simple, brownish yellowish —

pectinate basally, except inner hind claw (figs. 202, 203); length of femur, tibia,

and basitarsus of hind leg 6.6, 10.0, and 9.0 times their width, respectively; length

of spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.6 times its apical width, its surface weakly

reticulate-rugose medially, mainly smooth laterally (fig. 320); dorsal carinae of Ist

tergite absent; length of ovipositor sheath 0.04 times fore wing.

Colour. — Mesosoma, legs (but tibiae somewhat infuscated), Ist tergite

(mainly), and 2nd tergite laterally, reddish-brown; pterostigma dark brown; Ist

tergite medio-apically, 2nd tergite medially, 3rd and following segments mainly, —

blackish-brown; apical margins of tergites (except of Ist tergite) whitish-yellow;

dorsal half of clypeus, face, vertex anteriorly, and eye margins, yellowish-white;

remaining part of head and basal half of antenna, blackish-brown; apical half of

antenna brown.

Lectotype in MNHN, Paris: “Bekily, Reg. Sud de I’Ile’’, “Museum Paris, X1.36, —

A. Seyrig”, “50”, “Zele rufithorax Gr., B. Sigwalt’’. Lectotype herewith selected

From same locality 2 © paralectotypes (MNHN). Additionally examined a 9 from

Malagasy (Rogez, Forêt cote est, 1.31, A. Seyrig (MNHN)) with a type-label, ,

which cannot be a type-specimen because it is not mentioned in the original |

description. Length of malar space 0.6 times basal width of mandible; antennal |

segments 49; only frons and vertex anteriorly whitish-yellow; face light reddish-

brown; vein r of fore wing wider than 3-SR, length of Ist tergite 3.8 times its apical |

VAN ACHTERBERG: Revision Zelinae auct. 291

width, mainly smooth and wholly reddish brown; margins of 2nd and 3rd tergites

yellow; mesopleuron punctulate; length of fore wing 7.6 mm; length of 4th

segment of labial palp 3.0 times 3rd segment; hind tarsus mainly dark brown.

Homolobus (Apatia) maculatus spec. nov.

(figs. 204—215, 283)

Holotype, 9, length of body 5.7, of fore wing 5.6 mm.

Head. — Antennal segments 44, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 4.2 and 3.6 times their width, respectively, length of both

penultimate segments 1.3 and 1.5 times their width (fig. 211); length of 4th segment

of labial palp 3.5 times 3rd segment; length of maxillary palp 1.2 times height of

head; eyes indistinctly emarginate (fig. 210); dorsal length of eye 2.1 times temple;

POL : @ ocellus : OOL = 10: 9: 16; frons almost flat, medially mainly smooth,

laterally coriaceous (fig. 213); vertex rather convex, coriaceous; face weakly

punctate and convex; clypeus punctulate and with some lateral striae, convex;

apical margin of clypeus straight medially, not differentiated from clypeus (fig.

210); length of malar space 1.3 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

mainly smooth dorsally, crenulate medially and rugose ventrally (fig. 204);

epicnemial area weakly rugose; precoxal suture rather coarsely rugose-reticulate,

posteriorly more punctate; rest of mesopleuron punctulate; metapleural flange

round and wide apically, bordered by a rather narrow carina (fig. 204); notauli

rather widely crenulate posteriorly (fig. 214); mesoscutal lobes punctulate; surface

of propodeum narrowly smooth anteriorly and with a medium-sized medial carina,

rest of propodeum finely reticulate-rugulose.

Wings. — Fore wing: r: 3-SR : SRI = 10: 7: 45; r and 3-SR of equal width (fig.

206); SRI almost straight; cu-a antefurcal, straight; 2-M +CUI : CUI = 1: 19; 2-

SR : 3-SR : r-m = 11:7: 7; short part of 2A sclerotized (fig. 206); area basally of

2A mainly bare (fig. 208). Hind wing: r absent; SR straight; SC + RI weakly curved

(fig. 207).

Legs. — Hind coxa punctate and dorsally rugose (fig. 204); tarsal claws

pectinate basally and with a rounded, tiny subapical prominence (fig. 212); length

of femur, tibia, and basitarsus of hind leg 7.3, 10.8, and 9.2 times their width,

respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface behind

the spiracles reticulate-rugulose (fig. 215); dorsal carinae of Ist tergite absent

except for a short basal remnant (fig. 215); length of ovipositor sheath 0.09 times

fore wing.

Colour. — Yellowish-brown; apical half of antenna, vertex, stemmaticum,

middle of frons, pronotum partly, margin of mesoscutum, propodeum,

metanotum, Ist tergite mainly, base of 2nd tergite, apical margins of 3rd and 4th

tergites, Sth—8th tergites, Sth sternite, hypopygium partly, and tarsi mainly, more

or less blackish or dark brown; pterostigma brown; palpi slightly infuscated; wing

292- TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

membrane weakly brownish; apical margins of 3rd and following tergites

somewhat silvery.

Holotype in CNC, Ottawa: “N. Slope Mt. Elgon, Uganda, 2300 m,

17—26.x.i.1971, H.Falke”. Paratypes: 8 © and 4 &: 4 ©, topotypic (CNC,

RMNH); 1 &, “S. Tanganyika, Rungwe Mts., 2600 m, 5—10.xi.62 (allotype,

CNC); 1 9, “Afr. Orient. Ang., Kenya”, “Museum Paris, Nanyuki, VI.32, A.

Seyrig’ (MNHN); 1 ©, “Pietermaritzburg, XI—21—70, S. Africa, H & M.

Townes” (TC); 1 &, “Mt. Elgon, 8000 ft, IV.9.76, Kenya, lan Bampton” (TC); 1 &,

“Mpendle, Natal, XII-3-70, S. Africa, H. & M. Townes” (RMNH); 1 9 and 1 &,

“Kenya, Elgon Saw Mill, Mt. Elgon, Ver’est, (Camp II), 2470 m”, “Museum de

Paris, Mission de l’Omo, C. Arambourg, P.-A. Chappuis & B. Jeannel, 1932—33””

(MNHN); 1 ©, “Kenya, Kitale, Uasin Gishu, 2100 m”, “Museum de Paris,

Mission de l’Omo, C. Arambourg, P.-A. Chappuis & B. Jeannel, 1932—33”

(RMNH).

Variation: Length of fore wing 4.6—5.6 mm; length of ovipositor sheath 0.09

times fore wing; antennal segments 43—45; length of Ist tergite 2.5—2.6 times its

apical width; length of 4th segment of labial palp 3.2—3.7 times 3rd segment;

length of malar space 1.2—1.6 times basal width of mandible; vein cu-a of fore

wing antefurcal, interstitial, or shortly postfurcal; hind coxa sometimes only

punctulate; females with at least apical third of metasoma blackish.

Notes. Male essentially as female, but tarsal prominence more reduced, scarcely

visible at 80x ; apices of hind tibial spurs sharp and hyaline apically. For blackish

specimens from Malagasy, see note under albipalpis.

Homolobus (Apatia) alternipes spec. nov.

(figs. 216—230)

Holotype, 9, length of body and of fore wing both 5.0 mm.

Head. — Antennal segments 38, 3rd segment 1.3 times 4th segment; length of

3rd and 4th segments 4.9 and 3.8 times their width, respectively, length of both

penultimate segments 2.3 and 2.0 times their width (fig. 218); length of 4th segment

of labial palp 4.0 times 3rd segment; length of maxillary palp 1.1 times height of

head; eyes scarcely emarginate (fig. 229); length of eye 2.2 times temple; POL : 5

ocellus : OOL = 5: 5: 6; vertex rather flat, rugose near eyes (fig. 227); frons rather

flat, mainly smooth; face rather flat, punctate, and dorsally somewhat rugose;

clypeus convex, weakly punctate; apical margin of clypeus straight medially, not

differentiated from clypeus (fig. 229); length of malar space 0.7 times basal width

of mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

crenulate-rugose medially and posteriorly, coarsely rugose ventrally, and dorsally

narrowly punctulate (fig. 216); epicnemial area reticulate-rugose posteriorly;

precoxal suture densely rugose-reticulate; mesopleuron above precoxal suture

finely punctate; metapleural flange lamelliform, large, wide and rounded apically;

metapleuron dorsally mainly smooth, reticulate-rugose ventrally (fig. 216); notauli

VAN ACHTERBERG: Revision Zelinae auct. 293

largely narrowly crenulate, apically wider and more reticulate-rugose (fig. 228);

mesoscutal lobes densely punctulate; surface of propodeum coarsely reticulate-

rugose, but anteriorly narrowly smooth, with a short medial carina.

Wings. — Fore wing: r : 3-SR : SRI = 6: 5 : 42; SRI straight; cu-a straight,

postfurcal; 1-CUI : 2-CUl = 2: 16 ; 2-SR : 3-SR : r-m = 10: 5: 6; 2A shortly

sclerotized basally; area basally of 2A mainly bare (fig. 221). Hind wing: r absent;

SR straight; SC+ R1 weakly curved (fig. 220).

Legs. — Hind coxa coarsely punctate-rugose dorsally, more punctulate laterally

(fig. 223); tarsal claws slender, setose, with only a scarcely visible (at 80x)

subapical prominence (figs. 225, 226); length of femur, tibia, and basitarsus of hind

leg 5.8, 9.3, and 7.6 times their width, respectively; length of spurs of hind femur

0.6 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface largely

finely rugose, mediobasally and apically smooth (fig. 224); dorsal carinae of Ist

tergite absent; length of ovipositor sheath 0.09 times fore wing.

Colour. — Mesosoma brownish-red; head, antenna, pterostigma, parastigma,

wing veins, metasoma, and hind leg (except for the whitish trochanter and

trochantellus), more or less dark brown; palpi, fore and middle coxae, all

trochanters and trochantelli, tegulae, and (to a lesser degree) fore and middle

femora, yellowish-white; margin of 2nd tergite, mandibles and anellus, yellowish;

fore and middle tibiae and tarsi yellowish, but somewhat infuscated.

Holotype in MNHN, Paris: “Kenya, Nairobi, 1600 m”, ‘Muséum Paris, V1.32,

A. Seyrig”. Paratypes: (24), 1 &, “Meru, VI.32”, “Museum Paris, Kenya, A.

Seyrig” (allotype, MNHN). Length of malar space 0.8 times basal width of

mandible; r : 3-SR = 10: 7; antennal segments 39; apices of spurs of hind tibia

sharp and hyaline; Ist tergite mainly smooth; pronotum remotely rugose; length of

4th segment of labial palp ca. 4.5 times 3rd segment; mesopleuron above precoxal

suture punctulate; apical 0.4 of precoxal suture only punctulate; length of fore

wing 4.4 mm; claws without prominence. Second paratype without head, 1 &,

same labels as allotype (RMNH), pronotum only slightly punctate, precoxal suture

mainly rugose-reticulate, and above precoxal suture punctate.

Homolobus (Apatia) priapus (Nixon) comb. nov.

(figs. 138— 140, 231—239)

Nixon, 1938, Bull. ent. Res. 29: 418, 419, fig. 1b (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 225.

Holotype, 9, length of body and of fore wing both 7.2 mm.

Head. — Antennal segments 48, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 3.7 and 3.1 times their width, respectively, length of both

penultimate segments 2.0 and 2.3 times their width; length of 4th segment of labial

palp ca. 4 times 3rd segment; length of maxillary palp 1.4 times height of head;

eyes weakly emarginate (fig. 238); dorsal length of eye 2.3 times temple; POL : @

ocellus : OOL = 3: 8: 5; frons rather flat, anteriorly rugose; vertex punctate-

294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

rugose, rather flat (fig. 138); face mainly flattened, largely rugose, laterally

coriaceous; clypeus rather flat, punctulate; apical margin of clypeus straight

medially, narrowly differentiated from clypeus (fig. 238); length of malar space 0.8

times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely rugose, apico-dorsally and ventrally partly smooth (fig. 231); epicnemial

area reticulate-rugose; precoxal suture weakly impressed, coarsely rugose-

reticulate; rest of mesopleuron densely and finely punctate; metapleural flange

large, with a rather narrow lamella apically (fig. 231); metapleuron punctate, but

ventrally reticulate-rugose; notauli densely crenulate (fig. 239); mesoscutal lobes

densely punctulate; surface of propodeum largely reticulate-rugose, posteriorly

with a transverse carina, area behind it almost smooth, medial carina absent.

Wings. — Fore wing: r: 3-SR : SRI = 8:9: 43; SRI straight; cu-a inclivous, but

apically straight, postfurcal; 1-CUI : 2-CU1 = 1: 21; 2-SR : 3-SR : r-m = 11:9: 8;

2A well developed basally (fig. 234); area basally of 2A mainly bare. Hind wing: r

absent; SR straight; SC + R1 almost straight (fig. 232).

Legs. — Hind coxa finely and densely punctate, but postero-dorsally more

coriaceous (fig. 139); tarsal claws setose, with a (at 80 x) rather well visible, small

prominence (fig. 237); length of femur, tibia, and basitarsus of hind leg 7.0, 10.5,

and ca. 10 times their width, respectively; length of spurs of hind tibia 0.7 and 0.6

times basitarsus.

Metasoma. — Length of Ist tergite 3.2 times its apical width, its surface largely

reticulate-rugose, basally smooth (fig. 140); dorsal carinae of Ist tergite absent;

length of ovipositor sheath 0.08 times fore wing.

Colour. — Brownish-yellow; stemmaticum dark brown; apices of antennal

segments of apical half of antenna, labial palp and ovipositor sheath, somewhat

infuscated.

Holotype in BM, London: “Type”, “B.M. Type Hym. 3. c. 679”, “Zele priapus

Nixon, 9, Holotype’, “Cape Province, Somerset East, 10-22.xii.1930’’, “S. Africa,

R. E. Turner, Brit. Mus. 1931—37”. Paratypes: 1 9 (topotypic) and 7 g (Katberg

and Ceres, both S. Africa). Additional specimens examined (33 © and 18 @) are all

from South Africa (Jonkershoek, nr. Stellenbosch; Garies, Cape; Grahamstown;

Magoebaskloof, nr. Tzaneen; Kirstenbosch, nr. Cape Town; Deepwalls Forest,

Knysma, C. P.) (TC, HC, RMNH). Variation: length of fore wing 6.2—8.1 mm;

length of 4th segment of labial palp 3.3—5.5 times 3rd segment; length of malar

space 0.7—1.0 times basal width of mandible; antennal segments 44—48; whole

metasoma yellowish; only claws of male with a small subapical prominence; hind

tibial spurs of male sharp and hyaline apically; vein cu-a of fore wing antefurcal,

interstitial, or shortly postfurcal; length of vein r of fore wing equal to vein 3-SR,

or shorter.

Homolobus (Apatia) lacteiceps spec. nov.

(figs. 240— 254)

Holotype, 9, length of body 8.0, of fore wing 7.9 mm.

VAN ACHTERBERG: Revision Zelinae auct. 295

Head. — Antennal segments 49, 3rd segment 1.3 times 4th segment; length of

3rd and 4th segments 4.0 and 3.2 times their width, respectively, length of both

penultimate segments 2.1 and 2.7 times their width; length of 4th segment of labial

palp 2.4 times 3rd segment; length of maxillary palp 1.3 times height of head; eyes

weakly emarginate (fig. 245); dorsal length of eye 2.6 times temple; POL : @

ocellus : OOL = 7: 11 : 7; frons smooth, except for some lateral striae, rather flat

medially, convex laterally; vertex rather flat and smooth (fig. 242); face rather flat,

largely punctulate, dorsally weakly rugulose; clypeus convex, punctulate; apical

margin of clypeus straight medially, not differentiated from clypeus (fig. 245);

length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely smooth, medially and posteriorly crenulate-rugose (fig. 240); epicnemial

area almost smooth anteriorly, reticulate-rugulose posteriorly; precoxal suture

largely reticulate-rugulose, anteriorly and posteriorly only indistinctly sculptured

(fig. 240); rest of mesopleuron punctulate; metapleural flange small, with a narrow

rounded carina apically (fig. 240); metapleuron medially smooth, anteriorly

crenulate, and ventrally rugose; notauli narrowly crenulate (fig. 250); mesoscutal

lobes punctulate; surface of propodeum reticulate-rugose, laterally with some

more coarse rugae, medial and transverse carinae absent.

Wings. — Fore wing: r : 3-SR : SRI = 8: 11 : 41; SRI straight; cu-a strongly

inclivous, slightly bent basad apically (fig. 243), postfurcal; 1-CUI : 2-CUl = 5:

32; 2-SR : 3-SR : r-m = 20: 22: 11; 2A unsclerotized, only completely pigmented

(fig. 243); area basally of 2A setose (fig. 247). Hind wing: r absent; SR weakly

curved basally, strongly sinuate submedially; constriction of marginal cell distad

from its middle (fig. 243); SC+RI distinctly curved (fig. 254); 2-SC+R

subquadrate.

Legs. — Hind coxa in dorso-apical half striate, rest punctulate (fig. 240); tarsal

claws simple, yellowish pectinate basally (figs. 251, 252); length of femur, tibia, and

basitarsus of hind leg 6.0, 10.0, and 8.8 times their width, respectively; length of

spurs of hind tibia 0.6 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 2.7 times its apical width, its surface

anteriorly mainly smooth, posteriorly rugose, and medially finely rugulose (fig.

253); dorsal carinae of Ist tergite absent, except for a faint trace anteriorly;

exserted ovipositor longer than 1.5 times length of Ist tergite (fig. 240); length of

ovipositor sheath 0.25 times fore wing.

Colour. — Brownish-yellow; basal third of antenna and most wing veins, dark

brown; stemmaticum blackish, rest of head whitish.

Holotype in TC, Ann Arbor: ‘‘Zika Forest, Uganda, VIII.19.°63, G. Lancaster”.

Paratypes: (6 © and 1 g), all from Zika Forest, Uganda (1g (allotype), 23.viii. 1963

(TC); 2 9, 21.viii.1963; 2 9 (Mengo), 18.x.1963; 2 9 (Mengo, Entebbe),

13.iv.1964) (TC, RMNH). Variation: hind tibial spurs of male narrowly truncate

and pigmented apically; length of ovipositor sheath 0.22—0.26 times fore wing;

length of 4th segment of labial palp 2.2—2.4 times 3rd segment; antennal segments

48—52; length of fore wing 5.8—9.6 mm; length of malar space 0.5—0.6 times

basal width of mandible; area basally of 2A sometimes rather bare; middle lobe of

296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

mesoscutum punctate or punctulate; occiput sometimes with a dark bronze patch

behind the stemmaticum.

Homolobus (Apatia) pulchricornis (Nixon) comb. nov.

(figs. 141— 143, 255— 262, 279— 282)

Nixon, 1938, Bull. ent. Res. 29: 420, 421, fig. la (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 225.

Holotype, &, length of body 8.3, of fore wing 7.8 mm.

Head. — Remaining antennal segments 6, 3rd segment 1.1 times 4th eet,

length of 3rd and 4th segments 3.8 and 3.4 times their width, respectively; length of

4th segment of labial palp ca. 2.5 times 3rd segment; length of maxillary palp 1.3

times height of head; eyes rather emarginate (fig. 261); dorsal length of eye 1.8

times temple; POL : @ ocellus : OOL = 5: 7: 6; frons almost flat, smooth; vertex

flat, indistinctly coriaceous-punctulate (fig. 143); face rather flat, weakly and

finely punctulate-rugose medially; clypeus flattened, sparsely punctulate; apical

margin of clypeus straight medially, not well differentiated from clypeus (fig. 261);

length of malar space 0.6 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

rugose, dorsally and medio-ventrally smooth (fig. 255); epicnemial area crenulate

anteriorly, almost smooth dorsally; precoxal suture rather coarsely reticulate-

rugose; rest of mesopleuron somewhat superficially punctulate; metapleural

flange rather large, lamelliform, rounded apically (fig. 255); metapleuron remotely

punctate, only ventrally rugose; notauli densely crenulate (fig. 262); middle lobe of

mesoscutum densely and finely punctate, lateral lobes indistinctly punctulate;

surface of propodeum rather finely rugose, only anteriorly smooth (except

medially), carinae absent.

Wings. — Fore wing: r: 3-SR : SRI = 8: 12: 51; SRI curved towards RI; cu-a

inclivous, postfurcal; 1-CUI : 2-CU1 = 4: 20; 2-SR : 3-SR : r-m = 13: 12: 7; 2A

unsclerotized, only as a pigmented stripe (fig. 258); area basally of 2A remotely

and sparsely setose. Hind wing: r absent; SR distinctly curved basally and sinuate

medially; marginal cell constricted in front of middle of cell (fig. 258); 2-SC+R

subquadrate; SC + RI curved (fig. 259).

Legs. — Hind coxa mainly finely punctate-rugose, laterally almost smooth (fig.

141); tarsal claws simple, yellowish pectinate basally (fig. 260); length of femur,

tibia, and basitarsus of hind leg 6.0, 10.3, and 8.8 times their width, respectively;

length of spurs of hind tibia 0.6 and 0.5 times basitarsus, roundly truncate and

pigmented apically (fig. 256).

Metasoma. — Length of Ist tergite 2.8 times its apical width, its surface finely

and densely reticulate-rugose (fig. 142); dorsal carinae of Ist tergite absent.

Colour. — Brownish-yellow; stemmaticum, mesoscutal lobes partly, and

antenna (as far as present), more or less dark brown; behind stemmaticum a faint,

somewhat darker patch; pterostigma light yellowish.

Holotype in BM, London: “Type”, “B.M. Type Hym. 3.c.681”, “Zele

VAN ACHTERBERG: Revision Zelinae auct. 297

pulchricornis Nixon, 1938, Type, &”, “1938/18”, “Port St. John, Pondoland, July

10—31, 1923”, “S. Africa, R.E. Turner, Brit. Mus., 1923—398’’. One further

specimen examined (9, TC, allotype, “Gillitts, nr. Durban, XII-1-70, So. Africa,

H. & M. Townes”) on which the following addition is based. Antennal segments

47, length of both penultimate segments 2.1 and 2.5 times their width; length of 4th

labial palp segment 1.9 times 3rd segment; length of malar space 0.6 times basal

width of mandible; length of fore wing 9.1 mm; length of ovipositor sheath 0.14

times fore wing, slender, light yellowish; exserted ovipositor slightly longer than

Ist tergite (fig. 280); length of Ist tergite 2.7 times its apical width, its surface finely

and densely rugulose; antenna yellowish, except for the eight dark brown basal

segments; 2-SC + R shortly transverse (fig. 279); mesoscutum darkened anteriorly;

mesoscutum densely and finely punctate; face, frons and vertex light yellowish.

Homolobus (Apatia) huddlestoni spec. nov.

(figs. 263—277)

Holotype, 9, length of body and of fore wing both 7.5 mm.

Head. — Apex of antenna missing, remaining segments 24, 3rd segment 1.2

times 4th segment, length of 3rd and 4th segments 3.5 and 2.9 times their width,

respectively; length of 4th segment of labial palp 1.8 times 3rd segment; length of

maxillary palp 1.1 times height of head; inner sides of eyes moderately emarginate

(fig. 270); dorsal length of eye 2.6 times temple; POL: @ ocellus : OOL = 5:7: 3;

frons rather flat, striate (fig. 270); vertex narrow, concave near eyes, micro-

sculptured (fig. 275); face rather flat, dorsal half transversely and finely striate,

ventrally punctulate; clypeus rather flat, punctulate; apical margin of clypeus

straight medially, not differentiated (fig. 270); length of malar space 0.7 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

crenulate medially and apically, punctate dorsally, and mainly smooth ventrally;

epicnemial area reticulate-rugulose; precoxal suture reticulate-rugulose, weakly

impressed; metapleural flange large, rounded and lamelliform apically (fig. 263);

metapleuron almost smooth, only ventrally with some rugae; notauli almost

smooth anteriorly, crenulate-rugose posteriorly (fig. 274); mesoscutal lobes

punctulate; surface of propodeum densely and finely rugulose (fig. 263), without

carinae.

Wings. — Fore wing: r: 3-SR : SRI = 13: 17: 81; SRI weakly curved towards

R1; cu-a inclivous, postfurcal; 1-CUI : 2-CU1 = 1: 11; 2-SR : 3-SR : r-m = 18: 17:

10; 2A shortly sclerotized basally (fig. 266); area basally of 2A mainly remotely

setose (fig. 276). Hind wing: r absent; 2-SC +R quadrate; SC+R1 almost straight

(fig. 267); SR weakly sinuate (fig. 266).

Legs. — Hind coxa finely punctate-reticulate dorsally (fig. 269); tarsal claws

simple, and basally rather indistinctly yellowish pectinate, but inner hind claw only

setose (fig. 268); length of femur, tibia, and basitarsus of hind leg 5.3, 8.6, and 7.6

times their width, respectively; length of spurs of hind leg 0.6 and 0.5 times

basitarsus.

298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Metasoma. — Length of Ist tergite 2.0 times its apical width, robust (fig. 271), its

surface finely and densely rugulose and apically more striate; dorsal carinae of Ist

tergite absent, except for a weak basal remnant; length of ovipositor sheath 0.07

times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; flagellum and stripe on

outer side of scapus, brownish; pterostigma yellowish.

Holotype in BM, London: “Tanganyika, Ilonca, 1.10.1962, I: A. D. Robertson”,

“Ex pupa Euproctis sanguiguttata’’, ‘‘Zele sp., R. D. Eady det. 1964”.

The note that it emerged from the pupa of the host needs to be checked.

Paratypes: (89 and 7 3), 3 & (one allotype, BM): “E.A. Forest Insect Survey, J.

C. M. Gardner, Tanganyika, Mbulu, ex larva Euproctis fasciata WIk., No. 1238”,

“R. 662, 2.7.54" (BM, RMNH); 1 © “Ukiriguru T.T., Castor, 29.III.1958, J. A.

Robertson, Y. 129”, “Zele sp. nr. chlorophthalmus Nees, R. D. Eady, det. 1959”

(BM); 1 9, “Mtwapa, Kenya, Date 10.8.1971, No. 10718, B. R. Adams Coll., ex

larvae of PC 6599” (BM); 1 g, “S.Rhodesia, Fort Victoria, IV.1957, Min. Agric.”,

“Ex larva of Lepidopt. No. 8007”, “Larva No. 8007 = Euproctis rubricosta

Fawcett” (BM); 1 ©, topotypic (BM); 1 ©, “Jinja, Uganda, 26.11.1909, C. C.

Gowden No. 3” (RMNH); 1 9, “Uganda, Kampala, 5.x.1929, G. L. R. Hancock,

ex Arctornis rubricosta on cvHorn (?)”, “0356”, “Zele sp. n., Holotype, 9, R. D.

Eady, det. 1970” (RMNH); 1 9, “Entebbe, Mengo, V.11’64, Uganda, G. A.

Lancaster” (TC); 2 g, ‘Madagascar, Bekily, Reg. Sud de I’Ile’’, “Museum Paris,

IX.36, A. Seyrig” (MNHN); | 3, “Madagasc.: Fort Dauphin, A. Seyrig” (MAC);

1 ©, “Coll. Mus. Congo, Tanganika: Kamena, 1400 m (Riv. Kinga), H. Bomans,

1.1958” (MAC); 1 9, “Musée du Congo, Kibali-Ituri: Geti, 1934, Ch. Scops”

(RMNH). Variation: length of fore wing 5.3—6.8 mm; antennal segments 42 or 43;

length of both penultimate segments in figured apex of antenna (fig. 264) 1.6 and

1.9 times their width; length of 4th segment of labial palp 1.7—2.0 times 3rd

segment; length of malar space 0.5—0.7 times basal width of mandible; length of

hind femur 4.6—5.8 times its width, robust; hind tibial spurs of male sharp and

hyaline apically (figs. 272, 273); vein 2-SC + R quadrate or higher than wide; length

of Ist tergite 1.7—2.2 times its apical width; length of ovipositor sheath 0.04—0.08

times fore wing.

Notes. The hosts of this new species seems to be restricted to the Lymantriidae

(Lepidoptera), which is an aberrant choice within the genus Homolobus.

It is a real pleasure to dedicate this species to Mr. T. Huddleston (London);

without his spontaneous assistance this revision (and others) would be far less

complete.

Homolobus (Apatia) ophioninus (Vachal) comb. nov.

(figs. 278, 287—301)

Vachal, 1907, Revue Ent. 26: 122 (as Meteorus).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 83.

Holotype, 9, length of body 5.7, of fore wing 6.2 mm.

VAN ACHTERBERG: Revision Zelinae auct. 299

Head. — Antennal segments 46, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 3.4 and 2.8 times their width, respectively, length of both

penultimate segments 1.6 and 2.0 times their width; length of 4th segment of labial

palp 2.8 times 3rd segment; length of maxillary palp equal to height of head; eyes

weakly emarginate (fig. 301); dorsal length of eye 2.3 times temple; POL : @

ocellus : OOL = 8: 10: 9; frons flat, largely rugulose, medially mainly smooth (fig.

299); vertex convex, somewhat coriaceous; face rather flat, dorsally shortly

transversely rugulose and punctate, ventrally punctulate (fig. 301); clypeus rather

convex, punctulate; apical margin of clypeus almost straight medially, not

differentiated from clypeus; length of malar space 0.5 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

dorsally and ventrally smooth, medially and posteriorly crenulate-rugose (fig. 287);

epicnemial area finely reticulate-rugose as main part of precoxal suture, posterior

third of precoxal suture only punctate and rather flat; rest of mesopleuron

indistinctly punctulate; metapleural flange rather small, rounded and lamelliform

apically (fig. 287); metapleuron largely smooth, ventrally reticulate-rugose;

posteriorly notauli closely crenulate (fig. 300), anteriorly narrow and almost

smooth; surface of propodeum smooth anteriorly, rest mainly superficially

transversely rugulose-coriaceous; medial carina of propodeum shortly developed

anteriorly.

Wings. — Fore wing: r : 3-SR : SRI = 9: 18: 77; SRI rather curved towards RI

(fig. 289); cu-a inclivous, postfurcal, somewhat curved basad apically; 1-CUI : 2-

CUI = 2: 35; 2-SR : 3-SR: r-m = 15: 18: 11; 2A distinctly sclerotized basally (fig.

288); area basally of 2A sparsely setose (less than in figured specimen, fig. 288).

Hind wing: r absent; 2-SC + R transverse (fig. 289); basally SR less sclerotized than

Ir-m, weakly sinuate; SC + RI rather straight (fig. 290).

Legs. — Hind coxa mainly smooth, somewhat punctulate (but in figured

specimen finely reticulate-rugose dorsally, fig. 291); tarsal claws simple, only

basally indistinctly yellowish pectinate (figs. 294, 295); length of femur, tibia, and

basitarsus of hind leg 6.5, 10.5, and 9.0 times their width, respectively; length of

spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.4 times its apical width, its surface mainly

smooth, only posteriorly somewhat pimply-rugose (cf. fig. 298); dorsal carinae of

Ist tergite absent; length of ovipositor sheath 0.07 times fore wing, rather slender.

Colour. — Brownish-yellow; antenna apically, and outer side of scapus, slightly

infuscated; pterostigma (rather light) brown; stemmaticum blackish; wing

membrane subhyaline.

Holotype in MNHN, Paris: “Don de Mme Jaubert”, “Museum Paris, 1939,

Capit Quod”, ““G. Meteorus sp. ophioninus Vach., 9, Usumia”. The type originates

from New Caledonia.

Specimens additionally examined: 100 © and 42 Z. From the Australian region:

Australia: New South Wales (Narrabri; Maitland; Willowtree; 12 mi. NW.

Milton), Western Australia (Old Doongan; 10 mi. W. Mellewa; Yallingup;

Millstream; S. Coolgardie; 10 mi. S. Geraldton; 10 mi. W. Eucla; 5 mi. NW.

300 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Augusta; Wongan Hills; 13 mi. NEE. Caiguna; 21 mi. NE. Fraser Range; 19 mi.

NE. Mundrbilla), Southern Australia (9 mi. E. Cook; Mambray Creek; Leigh Co.;

35 mi. ESE. Morgan; 5 mi. S. Mungewarrie Sta.; 35 mi. E. Ceduna; Old Alton

Downs, Simpson Desert; Goyder Lagoon, Waterhole; 10 mi. ESE. Koonalda),

South Western Australia (Lake Magenta, at flowers of Eucalyptus), Australian

Capital Territory (2 mi. E. Mt. Coree; Canberra), Northern Territories (Tempe

Downs; 36 km SW. Borroloola). New Caledonia (Noumea). Norfolk Islands

(Burnt Pine, 370 ft; Duncombe Bay, 300 ft; J. E. Road, 200 ft) (RMNH, CSIRO,

CNC, BM, CAS, BPBM). From the Afrotropical and S. Palaearctic regions: Persia

(Beshire (? = Beshneh, S. Iran)), Ethiopia (Addis-Abbeba; Haut-Aduache,

Endessa; Karssa; Debra Zeit, 7200 ft), Kenya (S.W. Elgon, 6700 ft (specimen

figured); Naivasha; Nairobi; Muguga; Mau Escarpment, Molo, 2420 m; Mt.

Kenya, West side, lower zone, Ngaré Rungai, prairie river, 2000 m; Wa-Kikuyu,

Wambogo, 1750 m; Elgon Saw Mill, Mt. Elgon, 2470 m; Mt. Elgon, 2100 m;

Meru), Tanzania (Kilimandjaro, Kibonoto culture zone; id., Himo River, 1000 m,

lower zone), Malagasy (Ftanaransoa, Plateau Central; Bekily, Reg. Sud de lIle;

Tananarive; Banian, 70 m, Ankazoabo; Perinet; Andronotobaka, 1400 m,

Ambatolampy; Montagne d’Ambre, Les Roussettes, 1100 m; Ankasoka, 1130 m,

Route Lakete; Ankaratra, 1800 m; La Mandraka; Antsirabe), Ruanda

(Nyabikenke, Nyanza Terr., 1700 m; Sabiro, 1300 m), Zaire (Lomani, Kaniama;

Ituri, Blukwa; Lualaba, N’Zilo, N. Kolwezi, 1400 m; Kolwezi, Mulando;

Lubumbashi), Zambia (Welsley), and Rhodesia (Marandellas) (BM, MNHN, ZSB,

RMNH, MAC, TC, TMA, CNC, NR). Variation: Length of fore wing 5.8—9.0

mm; antennal segments 45—51; length of malar space 0.3—0.7 times basal width

of mandible; length of hind femur 5.0—6.9 times its width; spurs of hind tibia of

male truncate and pigmented apically (figs. 296, 297); length of 4th segment of

labial palp 1.8—2.8 times 3rd segment; length of Ist tergite 1.9—2.5 times its apical

width; length of ovipositor sheath 0.05—0.08 times fore wing; colour of

pterostigma varies from yellowish-brown to more or less dark brown; sometimes

mesosoma with dark patches (especially at the middle of the mesoscutal lobes);

2nd tergite rather whitish laterally; sometimes mesosoma with dark patches

(especially at the middle of the mesoscutal lobes); metasoma partly, apex of hind

femur, and main part of hind tibia and tarsus sometimes infuscated; vein 2-SC + R

of hind wing shortly transverse. Cocoon white and thin.

Notes. This species is closely related to australiensis, which differs mainly by the

presence of the black-and-white pattern of the metasoma. Known hosts of reared

specimens are Spodoptera exempta Walker and Agrotis segetum (Denis & Schiff.),

both belonging to the Noctuidae (Lepidoptera).

Homolobus (Apatia) truncatoides spec. nov.

(figs. 302—314, 324—326)

Holotype, 9, length of body 5.2, of fore wing 5.0 mm.

Head. — Antennal segments 42, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 3.2 and 2.8 times their width, respectively, length of both

penultimate segments 1.7 and 2.0 times their width; length of 4th segment of labial

VAN ACHTERBERG: Revision Zelinae auct. 301

palp 1.8 times 3rd segment; length of maxillary palp equal to height of head; eye

weakly emarginate (fig. 311); dorsal length of eye 2.1 times temple; POL : @

ocellus : OOL = 12: 10: 13; frons mainly flat and smooth, only behind antennal

sockets some sculpture (fig. 312); vertex convex, smooth; face rather flat,

transversely rugulose-punctulate (fig. 311); clypeus rather convex, punctate;

apical margin of clypeus straight medially, not differentiated from clypeus; length

of malar space 0.9 times basal width of mandible; upper condyli of mandibles

distinctly below lower level of eyes (fig. 311).

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, but medially crenulate and posteriorly reticulate-rugose (fig. 302);

epicnemial area and precoxal suture densely reticulate-rugose; rest of

mesopleuron smooth, except for some punctures near the pleural suture;

metapleural flange medium-sized, rounded and with a narrow carina apically (fig.

302); metapleuron smooth, only ventrally rugose; notauli crenulate (fig. 325);

‘mesoscutal lobes weakly punctulate; surface of propodeum densely and rather

finely reticulate-rugose, except for a narrow anterior part smooth and with a short

medial carina anteriorly.

Wings. — Fore wing: r : 3-SR : SRI = 6: 10: 47; SRI almost straight, but

slightly curved (fig. 306); cu-a weakly inclivous, postfurcal; 1-CUI : 2-CUl = 6:

37; 2-SR : 3-SR : r-m = 10: 10: 6; 2A sclerotized basally (fig. 306); area basally of

2A sparsely setose (fig. 304). Hind wing: r absent; SR weakly sinuate, scarcely

sclerotized (fig. 306); 2-SC +R transverse; SC + RI rather short and weakly curved

(fig. 307).

Legs. — Hind coxa largely punctulate, dorsally mainly rugulose; tarsal claws

simple, only yellowish setose, outer hind claw rather spiny setose (figs. 313, 314);

length of femur, tibia and basitarsus of hind leg 7.1, 10.2, and 8.8 times their width,

respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.2 times its apical width, its surface

longitudinally reticulate-rugose (fig. 326); dorsal carinae of Ist tergite absent,

except for a weak remnant; length of ovipositor sheath 0.04 times fore wing;

sheath truncate apically (fig. 303).

Colour. — Brownish-yellow; antenna (except inner side of scapus, pedicellus,

and anellus), all tarsi, middle of mesoscutal lobes, metanotum partly, Ist and 2nd

tergite, more or less brownish infuscated; wing membrane hyaline; pterostigma

light brown.

Holotype in ZMB, Berlin: “Egypten, Schmiedekn. S., 1897”, “Heliopolis bei

Cairo”, “28642”, ‘‘Phylacter nigricornis Walk., 2g” (in Schmiedeknecht’s

handwriting), “Zool, Mus. Berlin”. Paratypes: (40 © and 30 g) from the

Afrotropical region: 1 ¢, “Kenton-on-Sea, South Africa, XII.I—11.1970, Rex

Jubb” (TC); 1 4, “S. Africa, R. E. Turner, Brit. Mus. 1922—97”, “Mossel Bay,

Cape Province, Febr. 1922” (BM); 1 9, id., 15.iii—20.iv.1932 (RMNH); | 9, id.,

Febr. 1922 (BM); 1 9, ‘Madagascar, Tananarive, 6—13.x.1970”, “Coll. P.

Hammond, B.M. 1970-603” (BM); 1 ©, “Nyassa See, Langenburg, VI.98,

Fülleborn S.”, “Zele nigricornis Walk.” (in Szepligeti’s handwriting), “Zool. Mus.

Berlin” (ZMB); 1 &, “Coll. Mus. Congo, Madagascar: Ankaratra, IV-1944, A.

302 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Seyrig” (MAC); 2 g, “Museum Paris, Afrique occidentale, Konakry, A.

Chevalier, 1909”, “Décembre” (MNHN, RMNH); 1 ©, “Ilora, Nigeria, VIII’74,

W. State, J. T. Medler” (TC); 2 9, “Grahamstown, South Africa, I.17—31.71 &

II.15—22’71, Fred Gess” (TC, RMNH); 1 g, St. Lucia Estuary, XI.10, 70, So.

Africa, H. & M. Townes” (TC); 1 9, “Kenton-on-Sea, South Africa, XII.1.11,

1970 (RMNH); 1 &, id., January 1971 (TC); 2 2, id., H.1—19, 1971 (TC); 1 &, id.,

XI.15—30, 1970 (RMNH); 2 9, id., XI.1—14, 1970 (TC, RMNH); 2 9, id., March

1971 (TC); 1 9, id., April 1971 (TC); 2 2, id., June 1971 (TC); 1 9, “Madagascar,

Antsirabé”, ‘Muséum Paris, XI.36, A. Seyrig’’, “Zele chlorophthalmus Nees, B.

Sigwalt” (RMNH); 1 ©, “Madagascar, Ankaratra, Alt. 1800 (m)”, “Museum

Paris, 11.38, A. Seyrig”, “48”, “Zele chlorophthalmus Nees, B. Sigwalt” (MNHN); 1

©, “Museum Paris, Madagascar, Région du Sud-est, Forêt Dauphin, Ch. Allaud,

1901”, “Zele nigricornis Walker” (in Szepligeti’s handwriting), ‘Zele

chlorophthalmus Nees, B. Sigwalt” (MNHN); 2 9 and 1 &, “Museum Paris,

Zambéze, Nova Choupanga, pres Chemba, P. Lesne, 1929”, “Zele chlorophthalmus

Nees, B. Sigwalt”; 1 &, “Muséum Paris, Madagascar, Tananarive, R. Decary,

1921”, “Fevrier”, “Zele chlorophthalmus Nees, B. Sigwalt”” (MNHN); I 9,

“Museum Paris, Ethiopie Merid., Haut-Aduache, Endessa, Maurice de

Rothschild, 1905”, “Zele nigricornis Walk., V. Szépligeti, det. 1907” (TMA); 1 9,

“Madagascar, Perinet, XI-7-1959”, “E. S. Ross Collector” (CAS)). From the

South Palaearctic region: 3 g, “Aegyptus, Schmkn., 97”, “Memphis”, “Zele

nigricornis Walk., det. Szépligeti (TMA, RMNH); | g, “Alexandria, Egypt”, “Zele

chlorophthalmus (Nees), det. P. Marsh” (USNM); 1 g, “Spain, Almeria, El

Alquinan, 3 March 1966, Leif Lyneborg” (UZM); 4 9 and 4 g, “Museum Leiden,

Bär, Blôte, De Jong & Osse, Estepona, 3.X.1952, Spanje” (RMNH); 2 9 and 2 g&,

id., but 30 km ZW. Malaga, 4.X.1952 (RMNH); 1 g, id., but from Jerez de la

Frontera, 22.1X.1952 (RMNH); 1 3, “Islas Canarias, Tenerife, J. Wolschrijn”,

“Los Cristianos, 13/26.11.1977” (RMNH); 1 g, “Almunecar, Granada Prov.,

Spain, 0—30 m, J. R. Vockeroth, 16.VII.1960° (CNC); 1 &, “nr. Limassol,

X1.21’46, Cyprus Mavroumoustakis” (TC); 1 9 and 2 Z, “Cyprus, Yermasovia R.,

25.X1.66, 4.XI.1967, and 12.V.1966, respectively, Mavroustakis” (CNC, RMNH);

1 3, Italy, “Palermo, XI.63” (TC); 2 © and 1 g from Iraq: “Loc. Hindiya, 31/10/

1956”, “Host (on) Beta, Coll. S. Alyasiri”, “Zele cf. 9 calcarator Q”’, ““8/11/195.”,

“Host (on) sugar beet, Coll. D. Ahmad”, “Zele cf. calcarator” (HC), and “Iraq,

Diwanyye, 12-9-1954, light trap”, “Zele cf. calcarator &” (HC); 1 9 and 1 3, “El

Riyadh; Saudi Arabia, 9.X.1959, E. Diehl” (CNC, RMNH)). And from the

Oriental Region: 1 ©, ‘‘1400 ft, Coimbatore, South India, XI.1966, P. S. Nathan”

(CNC); 1 9, “India, Mysore, 10 mi. NW. Kittur, 15.11.1962, E. S. Ross & D. Q.

Cavagnaro” (CAS); 1 9, ‘“Malaya, Cameron Highlands, Mt. Brichang, 2— 7.1.59”

(BPBM); 2 9, “India, A. P., Warangal, A. R. S. Coll.” (DZD, RMNH); | 9,

“India, U.P., Dehra Dun, 600 m, 8.IV.1976, S. Biswas No. B13” (DZD). Specimens

excluded from the type-series: 1 g, “Kandy, Ceylon, W. Horn”, ““Co-type”,

“Macrocentrus ceylonicus Enderl., &, Type, Dr. Enderlein det. 1912”. A wrongly

identified specimen, still belonging to the type-series of Metapleurodon ceylonicus

(Enderlein) (PAN). Furthermore 1 9, “Jonkershoek, near Stellenbosch, X.8.70, S.

VAN ACHTERBERG: Revision Zelinae auct. 303

Africa, H. & M. Townes” (TC), excluded from the type-series because of the

colour and sculpture. Middle of frons, stemmaticum, middle of mesoscutal lobes,

antenna apically, pterostigma, propodeum, meso- and metapleura dorsally, Ist

tergite, base of 2nd tergite and tarsi, more or less dark brown; 2nd tergite is

somewhat rugulose basaily; 1 3, “Kenton-on-Sea, South Africa, April 1971, Rex

Jubb” (TC), excluded because the apices of the hind tibia are sharp and hyaline.

Variation: length of fore wing 3.5—7.1 mm, antennal segments 39—44; length of

4th segment of labial palp 1.6—2.5 times 3rd segment; length of Ist tergite 1.7—2.6

times its apical width; length of malar space 0.8—1.1 times basal width of

mandible; length of hind femur 5.6—6.3 times its width, exceptionally 4.9 times;

hind tibial spurs of male (sub)truncate and pigmented apically (figs. 308, 310);

length of ovipositor-sheath 0.08 times fore wing; claws at most indistinctly

yellowish pectinate basally; apical width of marginal cell 1.9—2.2 times its

maximum basal width.

Homolobus (Apatia) pallidistigmus (Cameron) comb. nov.

(figs. 327—331, 334—340, 709, 712)

Cameron, 1911, Ann. Transv. Mus. 2: 210 (as Macrocentrus pallidistigmas).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 166.

Holotype, &, length of body 9 (according to Cameron), of fore wing 8.9 mm,

metasoma and hind leg absent.

Head. — Remaining antennal segments 12, 3rd segment 1.1 times 4th segment,

length of 3rd and 4th segments 2.9 and 2.6 times their width, respectively; length of

4th segment ca. twice 3rd segment; length of maxillary palp about equal to height

of head; inner sides of eyes rather emarginate (fig. 330); POL : @ ocellus : OOL =

9: 9: 7; frons mainly rugose, flat (fig. 328); vertex superficially punctulate; face

densely punctulate, near eyes and near clypeus almost smooth, rather flat; clypeus

almost smooth, superficially and remotely punctulate, flattened; apical margin of

clypeus somewhat convex medially, not differentiated from clypeus (fig. 330);

length of malar space ca. 0.8 times basal width of mandible; upper condyli of

mandibles distinctly below lower level of eyes (fig. 330).

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

ventrally smooth, medially crenulate, posteriorly and dorsally closely punctate

(fig. 327); epicnemial area densely punctate; precoxal suture shallow, densely

punctate; rest of mesopleuron finely punctulate, almost smooth; metapleural

flange large, lamelliform, rounded apically (fig. 327); notauli deep and finely

crenulate (fig. 331); surface of propodeum densely punctate-rugose, but almost

smooth anteriorly, without carinae medially.

Wings. — Fore wing: r: 3-SR : SRI = 10: 17: 73; SRI curved (fig. 329); cu-a

subinterstitial, inclivous, and somewhat curved basad apically; 2-SR : 3-SR : r-m =

17: 17: 10; 2A sclerotized basally (fig. 329); area basally of 2A remotely setose.

Hind wing: r absent; 2-SC+R transverse; SR weakly sinuate, basally rather

sclerotized (fig. 329); SR+RI straight (fig. 337); apical width of marginal cell 2.4

times its maximum basal width.

304 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Legs. — Middle coxa smooth; middle spurs of hind tibia 0.3 times basitarsus,

subequal.

Colour. — Brownish-yellow; stemmaticum blackish; pterostigma mainly light

yellowish; according to the original description the apical half of the antenna is

black.

Holotype in TMP, Pretoria: “Rietf., 11.2.05, 11.” (= Rietfontein, Pretoria

District, S. Africa), “Macrocentrus pallidistigmus Cam., Type” (in Cameron’s

handwriting). Additional specimens examined (17 © and 14 @) from Kenya

(Karen, Nairobi; nr. Nairobi, 6000 ft), Uganda (Zika Forest, Mengo; Kampala;

Katona, Mujenje), Tanzania (W. Usambara Mts., 2100 m, Magamba; Mt. Meru,

1800 m; Chome, Pare Mts., 1800 m), and Zaire (Rutshuru) (TMA, MAC, RMNH,

CNC, TC).

Variation: Length of fore wing 7.0—9.5 mm; antennal segments 54 in one 9;

length of 4th segment of labial palp 2.6—2.8 times 3rd segment; length of malar

space 0.8—1.1 times basal width of mandible, exceptionally 0.6 or 0.7 times; length

of Ist tergite 2.8—3.4 times its apical width (fig. 334); length of hind femur 6.9—7.0 :

times its width (fig. 338); length of ovipositor sheath 0.04—0.06 times fore wing

(fig. 335); tibial spurs of hind leg of males truncate and pigmented apically, rather

slender (figs. 709, 712); claws simple, more or less pectinate basally (figs. 339, 340);

apical width of marginal cell of hind wing 2.4—2.6 times its maximum basal width;

antenna more or less yellowish-brown.

Notes. Homolobus (Apatia) pallidistigmus (Cameron) belongs to a group of four

species, which are sometimes difficult to separate. The two most closely allied

species being ophioninus and truncatoides, while huddlestoni is related to ophioninus

but rather easily recognizable, these three being usually somewhat smaller than

pallidistigmus. H. ophioninus is separable by the rather transverse frontal aspect of

the head, because of the highly situated upper condyli of the mandibles. While

truncatoides has a more trapezoidal frontal aspect of the head as in pallidistigmus,

the latter differs from truncatoides by a more widened marginal cell of the hind

wing, a (usually) more developed basal third of vein SR of hind wing, and a more

straight SC+RI. Because the variation is considerable, a careful examination is

needed to arrive at a reliable identification. The shape of the hind tibial spurs of

the males, for instance, may be useful; in pallidistigmus the spurs are rather slender

apically, while in ophioninus and usually also in truncatoides they are stout apically.

Because of the variation other characters have to be considered as well!

Macrocentrus pallidistigmus Cameron sensu Szépligeti belongs to ““Macrocentrus”

albitarsis Granger, 1949.

Subgenus Chartolobus nov.

Etymology: from ““charta” (Latin for “‘lamina’’) and “lobus” (Latin for “‘protuberance’’), because of the

more or less developed ventral lamella of the claws. Gender: masculine.

Type-species: Zele infumator Lyle.

Diagnosis. — Length of body 7.1—14.6, of fore wing 7.0—15.9 mm; antennal

VAN ACHTERBERG: Revision Zelinae auct. 305

segments 46—52, its 3rd—6th segments of 9 with a ridge at the inner side (figs.

348, 366, 877, 878); length of 4th segment of labial palp 3.0—4.0 times 3rd segment;

length of maxillary palp 1.5—1.7 times height of head; apical margin of clypeus

straight medially and more or less differentiated from clypeus (figs. 344, 374);

length of malar space 0.4—0.8 times basal width of mandible; temples roundly (fig.

347) or directly (fig. 375) narrowed apicad; length of hind femur 6.1—7.8 times its

width; claws with a subapical tooth or lamella (figs. 350, 364); inner hind claw of 9

concave and bare ventro-basally (figs. 351, 365, 888); hind telotarsus of 9 more or

less bare near base of inner hind claw (fig. 888); apices of hind tibial spurs of ¢

sharp and hyaline; 1A + 2A of fore wing curved (figs. 353, 369, 380); basal third of

SR of hind wing mainly sclerotized (figs. 349, 368, 379), curved (fig. 343) or almost

straight (fig. 379); SC+RI of hind wing curved (figs. 349, 382); r of hind wing

absent; length of Ist tergite 2.1—3.1 times its apical width; 2nd tergite smooth;

length of ovipositor sheath 0.04—0.07 times fore wing; posterior part of

propodeum more or less separated from antero-dorsal part by a transverse carina

(figs. 341, 352, 358, 729), both in about the same plane.

Distribution. — One of the three species has an immense distribution; it ranges

from the South Neotropical region (Argentina), through the Holarctic region as

far as the Oriental region (Indonesia). Both other species occur in the Australian

region, and H. undulatus occurs also in the Oriental region.

Key to the species of the subgenus Chartolobus

1. Vein 2A of fore wing widened basally and apically if compared with the

surrounding veins (figs. 368, 369, 379, 380); Indo-Australian ........ 2

— Vein 2A of fore wing slender, not or slightly widened if compared with

surrounding veins (figs. 343, 353); Neotropical, Holarctic, Oriental ......

se 0 6 AE e alia lie infumator (Lyle) (p. 305)

2. Vein SR of hind wing weakly curved basally (fig. 368); vein 2A of fore wing less

widened (fig. 369); face more shiny and punctulate (fig. 360); pterostigma light

brownish or yellowish brown; hind tarsus yellowish or whitish; Indo-

ES RAA CU NN undulatus spec. nov. (p. 309)

— Vein SR of hind wing straight basally or nearly so (figs. 379, 382); vein 2A of

fore wing strongly widened (fig. 380); face rugulose-coriaceous laterally and

rather dull (fig. 374); pterostigma and hind tarsus blackish; Australian

MEIN SA REN A Cel ee Ne es BT nigritarsis spec. nov. (p. 310)

Homolobus (Chartolobus) infumator (Lyle) comb. nov.

(figs. 171—173, 341—353, 877, 878, 888)

Nees von Esenbeck, 1834, Hym. Ichn. affin. Mon. 1: 202, 203 (as chlorophthalmus, nec Spinola, 1808!).

Lyle, 1914, Entomologist 47: 288, 289, figs. 2, 5, 9 (as Zele).

Bengtsson, 1918, Acta Univ. lund. (2)14(32): 39, 41 (Phylacter wesmaeli). Syn. nov.

Watanabe, 1932, Insecta matsum. 6: 135 (Zele testaceator f. japonica).

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319—324, fig. 7.

306 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 224.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 230, 231.

Capek, 1972, Ent. Problémy 10: 133, 136.

Kabatsjinskajte & Jakimavicius, 1973, Acta ent. Lituanica 2: 86.

Jakimavicius, 1974, Tr. AN Lit. SSR B2(66): 97.

Capek, 1975, Biolögia 30: 819.

Van Achterberg, 1976b, Tijdschr. Ent. 119: 73, figs. 103, 104.

Tobias, 1976, Opr. Fauna SSSR 110: 133, fig. 39: 5, 6.

Redescribed after the lectotype of H.(C.) wesmaeli (Bengtsson), &, length of

fore wing and of body both 7.1 mm.

Head. — Antennal segments 37, but apical segments absent, 3rd segment 1.3

times 4th segment, length of 3rd and 4th segments 4.0 and 3.2 times their width,

respectively; length of maxillary palp 1.5 times height of head; eyes weakly

emarginate (fig. 344); dorsal length of eye 1.6 times temple; temple rounded

apicad (fig. 347); POL : @ ocellus : OOL = 8: 7: 6; frons almost smooth and flat;

vertex smooth; face rather flat, transversely rugose-striate, but triangular area

above clypeus smooth (fig. 344); clypeus rather flat, superficially punctulate,

almost smooth; length of malar space 0.7 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

superficially punctulate, with some short crenulae medially (fig. 341); epicnemial

area almost smooth; precoxal suture anteriorly finely rugose, its posterior half

smooth (fig. 341); rest of mesopleuron indistinctly punctulate; metapleural flange —

large, lamelliform, rounded apically; metapleuron punctulate; notauli finely

crenulate (fig. 172); mesoscutal lobes punctulate; surface of propodeum smooth, .

except for some rugae medially and irregular medial and transverse carinae, with

an arc-shaped carina posteriorly, enclosing a small semicircular areola (cf.

fig.352).

Wings. — Fore wing: r: 3-SR : SRI = 7: 13: 54; SRI slightly curved (fig. 343);

cu-a shortly antefurcal, straight; 2-M +CUI : CUI = 1: 22; 2-SR : 3-SR: r-m = 12.

: 13:9; 2A sclerotized and slender basally (fig. 353); area basally of 2A bare except

for some setae basally. Hind wing: Basal third of SR sclerotized and curved (fig.

343); SC+RI strongly curved (fig. 349); marginal cell distinctly constricted (fig.

343).

Legs. — Hind coxa smooth; tarsal claws with a subapical tooth (fig. 345); length

of femur, tibia and basitarsus of hind leg 7.0, 10.9, and 9.0 times their width,

respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.4 times its apical width, its surface smooth.

(fig. 346); dorsal carinae of Ist tergite absent.

Colour. — Brownish-yellow; stemmaticum dark brown.

Lectotype in KBIN, Brussels: “Coll. Wesmael”, “1876”, “Phylax chloroph-

thalmus N.V.Es., 9, det. C. Wesmael”, “Type”. Lectotype of Phylacter wesmaeli

Bengtsson, 1918, herewith selected, and labelled accordingly. There are in the |

Wesmael Collection two other heavily damaged specimens with simple claws

which probably belong to truncator.

The type-series of Zele infumator Lyle consists of 11 9 and 13 g, of which

several are reared from Alcis repandata (L.) and one 9 from Agonopterix

VAN ACHTERBERG: Revision Zelinae auct. 307

alstroemeriana (Clerck), is in the BM collection (London). One ¢ (glued on a card

with a red dot, “type”, and “530”, with a whitish cocoon, “B.M. Type Hym.,

3.c.56”, “Zele infumator Lyle”, “G. T. Lyle, New Forest, 31.5.10, Ex B. (=

Boarmia) repandata’’, “G. T. Lyle Coll., B.M. 1930—579”’) is herewith selected as

lectotype. The propodeum is strongly areolate (fig. 352), the Ist tergite somewhat

reticulate-punctate and its length 2.2 times its apical width, and wings slightly

infumated apically.

The type of H. (C.) japonicus (Watanabe) is a normal specimen of infumator, with

rather whitish hind tarsi, a feature not uncommon in New World populations. The

holotype is a 9, housed in EI, Sapporo: “23.X.1924, Takao, Takeuchi”, “Zele

testaceator Curtis f. japonicus, Type”, “Type Hym. 22 No. 22”.

Additionally examined specimens: 209 © and 106 g. From the Neotropical

region: Argentina (S. Pedro d. Colalao, 1200 m; Horco Molle, nr. Tucuman; Villa

Nogues; Tafi del Valle), Peru (Dept. Lima, Matucana, 2389 m), Bolivia (Coroico,

Yungas La Paz; Coroico, 1800 m), and El Salvador (Monte Cristo, 2300 m). From

the Nearctic region: Mexico (Chis., 7200 ft, S. Crist. las Casas; Chis., 9600 ft,

Zontehuitz, nr. S. Crist.; Dgo., 30 mi. W. La Cuidad, 6500 ft; id., 24 mi. W. La

Cuidad, 7000 ft; Dgo., 9000 ft, 10 mi. W. El Salto), California (Skyline Blvd., San

Mateo Co.; Mill Valley, Marin Co.; Orinda Village, Contra Costa Co., San Pablo

Ridge, below Eureka peak, 1000—1200 ft, oak-chaparral zone; Julian; Lake

Wohlford; Forest Glen, 2300 ft, Trinity Co., black light), Nevada (Lee Cyn., 40 mi.

NW. Las Vegas, Clark Co., 7400—7500 ft; Baker Creek Camp, 8 mi. W. Baker,

White Pine Co., 7700 ft), Arizona (5 mi. W. Portal, Cochise Co., 5400 ft; Huachuca

Mts., Cochise Co., Floor of Carr Cyn, 5400 ft; id., 15 mi. S. Sierra Vista, Ramsey

Cyn., 5000—6000 ft; Hidden Springs Cyn., 4875 ft, 12 mi. S. Sonoita; Canelo,

Santa Cruz Co.; Portal), New Mexico (Cimarron Cyn., 7900 ft, Sangre de Cristo

Mts., Colfax Co., black light; Ute Park, 7300 ft), Colorado (Saguache Co., Valley

View Springs, ca. 7 mi. E. of Mineral Hot Springs on W. foot of Sangre de Cristo

Range, ca. 8500 ft), Utah (Whiterock, 7300 ft), Florida (Waldo; Hawthorne),

Montana (Missoula, 3000 ft), Illinois (no locality), South Carolina (Wattacoo,

Pickens Co.), Minnesota (Big Fork), Washington (19 mi. NW. Newport, 2850 ft),

Maine (Dryden), New Brunswick (Charls Fork, N. Branch), Quebec (L. Expanse;

La Tugie; Otter Lake; Lake Mondor, Ste. Flore, at light), Ontario (Ottawa, Dow’s

Swamp; Sudbury), and British Columbia (Victoria; Squamish, Diamond Head

Trail, 3200 ft; Great Central L.; 28 mi. S. Radium Hot Springs, 2600 ft).

From the Palaearctic region: Finland (Helsinki; Somerniemi; Lapptrask;

Ruokolanti), USSR (Vilnius, Verkiat, Lit. SSR; Armenia, Tsav, Jabl. sad), Sweden

(Skane), Denmark (Veldes; Hannenvo, Falster; Egebjeggd, Nordfyn; Odense),

East Germany (Berlin), West Germany (Steinebach am Worthsee; Haffen;

Lüneburgerheide; Gräfelfing; Reither Alm, 850 m; Rheinhöhenweg im

Kottenforst (nr. Bonn); Mainz; Spessart, Lochmühle), Ireland (Drinnahilly, C.

Do.; Tollymore Pk., Co. Do.; Bansla Wd., Co. St.; Old Head, Co. Wm.; Lodge

Wds, Glengariff, Co. Wo.), England (Essex, Round Stone (in Curtis Collection,

under chlorophthalmus); Hants., New Forest, Minstead), Netherlands (Dorst, nr.

Breda; Valkenswaard; Harskamp; Amersfoort, Den Treek; Putten (Gld);

308 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

’t Harde; Hilversum; Hoog Soeren; Vierhouten; Heerde; Nunspeet; Crailo; Den

Dolder; Ede (Gld.); Loenen; Muiderberg; Naardermeer; Overveen; Melissant;

Chaam; Amsterdamse Duinwaterleiding, nr. Vogelenzang; Tegelen; Molenven (?

nr. Oisterwijk); Oostkapelle; Oploo; Venlo; Bergen op Zoom), France (Jura,

Baudiette), Spain (Huesca, Torla, 1035 m), Italy (Garda Lake, Malcesini, 300 m;

Süd-Tirol, Cortina d'Ampezzo, Pokol, 1527 m; Tirol, Leutaschstrasse nr.

Mittenwald, ca. 1000—1100 m; San Marino, Marche; Riva s. Garda, 250 m),

Austria (Styr., Podcetrtek; Nordkitte, 2000 m; Styria m., Sausal-Gebirge, Kitzeck,

300—500 m), Czechoslovakia (Radosima, 8 km SE. Piest’any, Povazsky Inovex;

Hostyn-okoli, Mor. or.), Romania (Transsylv. Alp., Cibins. Mts., Hohe Rinne),

Greece (Ellas, Kerkyra, Dassia, 5 km SE. Korakiana), Nepal (Ktmd., Pulchauki,

8000 ft), and Japan (Mie Honshu; Kyoto, Honshu; Nagano, Honshu; Sapporo;

Jokohama; Hirakura, Mie Honshu). From the Oriental region: Taiwan (Sunmoon

Lake), Philippines (Baguio, Benguet), India (U.P., 8000 ft, Chakrata), and

Indonesia (Java, Kenden(g) Ridge, 1500 m) (RMNH, ITZ, EI, CNC, BM, MSU,

AMNH, CAS, UCA, TC, UZM, HC, ZMH, ZMB, IZP, ZSB, USNM, NMV).

Variation: Length of fore wing 6.8—10.0 mm; antennal segments 46—SO;

flagellum of antenna often dark brown of blackish basally; cu-a of fore wing

shortly antefurcal, interstitial or shortly postfurcal; 2-SC +R sometimes quadrate;

cocoon whitish or yellowish; length of ovipositor sheath 0.05—0.07 times fore

wing; length of malar space 0.3—0.7 times basal width of mandible.

Known hosts of examined specimens: Lambdina fiscellaria (Guenée),

L. somniaria (Hulst), Nepytia canosaria (Walker), Alcis repandata (L.), Lycia zonaria

(Denis & Schiff.), Ematurga atomaria (L.) (all belonging to the Geometridae), and

Agonopterix alstroemeriana (Clerck), the latter belonging to the Oecophoridae.

Note. Nees (1834) has used the name chlorophthalmus in the genus Rogas

probably (at least partly) for this species and referred to the description of Bracon

chlorophthalmus Spinola, 1808, in this manner devaluing his description to merely a

misidentification of Spinola’s species, as pointed out under Zele chlorophthalmus in

this paper (p. 372). The misinterpretation was accepted by Wesmael (1835), whose

two specimens still exist. Through the action of Bengtsson (1918), who renamed

chlorophthalmus sensu Wesmael as wesmaeli, they became part of the type-series of

Homolobus (C.) wesmaeli (Bengtsson, 1918). In the Wesmael Collection there are

one ¢ and two damaged specimens, probably females. In his description Wesmael

stated that he possessed one 3 and one 9, of which the g had the metasoma

darkened dorsally. This male is selected as lectotype of wesmaeli in this paper; the

lectotype is a specimen of infumator, which is a senior synonym of wesmaeli. The

interpretation of chlorophthalmus sensu Nees (nec Spinola & Haliday) is uncertain

and his specimens are lost. I have rejected the interpretation by Nixon (1938),

because Wesmael is the first revisor of Nees’ interpretation and the lectotype of

wesmaeli fits well the description by Nees. The species which is named Homolobus

truncator (Say) in this paper, was named Zele chlorophthalmus sensu Nees by Nixon

(1938).

VAN ACHTERBERG: Revision Zelinae auct. 309

Homolobus (Chartolobus) undulatus spec. nov.

(figs. 358—369, 729, 730)

Holotype, 9, length of body 14.6, of fore wing 15.9 mm.

Head. — Antennal segments 52, ridge of 4th—6th segments undulate (fig. 366),

3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 3.5 and 2.8

times their width, respectively, length of both penultimate segments 2.0 and 2.4

times their width; length of maxillary palp 1.7 times height of head; eyes rather

emarginate (fig. 360); dorsal length of eye 2.2 times temple; temple directly

narrowed posteriad (fig. 363); POL : @ ocellus : OOL = 6: 13: 8; frons somewhat

concave medially, mainly smooth; vertex flat, somewhat punctulate and

coriaceous; face mainly flat, punctulate; clypeus weakly convex, punctulate (fig.

360); length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, except for some short crenulae medially and somewhat rugose posteriorly

(fig. 358); epicnemial area punctulate; precoxal suture crenulate antero-dorsally,

densely punctate antero-ventrally, and its posterior half finely punctulate, as rest

of mesopleuron (fig. 358); metapleural flange large, lamelliform, sharp apically,

and with a medial carina; metapleuron punctulate, ventrally with some carinae;

notauli rather narrow and mainly smooth (fig. 359); surface of propodeum smooth,

but medially carinate-rugose, with irregular lateral and transverse carinae (fig.

729).

Wings. — Fore wing: r: 3-SR : SRI = 11 : 22: 77; SRI weakly curved (fig. 368);

cu-a weakly inclivous, postfurcal; 1-CUI : 2-CU1 = 2: 27; 2-SR : 3-SR : r-m = 19:

22 : 12; 2A strongly widened and sclerotized basally (figs. 368, 369); area basally of

2A mainly bare. Hind wing: Basal third of SR sclerotized and curved (fig. 368);

SC+R1 strongly curved (fig. 367); marginal cell distinctly constricted.

Legs. — Hind coxa punctulate; tarsal claws with a rather large ventral lamella,

which is sharp apically, setose (figs. 364, 365); length of femur, tibia and basitarsus

of hind leg 7.8, 11.9, and 10.2 times their width, respectively; length of spurs of

hind tibia 0.5 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 3.1 times its apical width, its surface mainly

smooth, laterally and posteriorly somewhat rugulose (fig. 729); dorsal carinae of

Ist tergite weakly developed in front of spiracles; length of ovipositor sheath 0.04

times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; surroundings of stem-

maticum somewhat infuscated.

Holotype in RMNH, Leiden: “Neth. Ind.-Amer. New Guinea Exped., 2800 m,

Moss Forest Camp, 18.X.1938, L. J. Toxopeus leg.’’. For location of the camp, see

Toxopeus (1940). Paratypes: (10 9 and 8 4) from New Guinea (1 g, allotype,

“N.E. New Guinea, Eastern Highlands, Mt. Wilhelm, VI.1965, Research Station,

v. Balgooy”; 2 9, “Net. Ind.-Amer. New Guinea Exp., 1938, Lake Habbema,

3250—3300 m, ult. VII-ult. VIII., L. J. Toxopeus leg.”; 1 ©, “Museum Leiden,

Nieuw Guinea Exp., K.N.A.G. 1939, Paniai, 19.XI.1939”, (all RMNH); 1 6,

“New Guinea (NE), Morobe, Mt. Kaindi, 2350 m, X.1974”; 1 9, “New Guinea,

NE., Mt. Kaindi, 2350 m, 12.xi.1964” (both BPBM)), Australia (19 (TC), “Mt.

340 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Cootha, Qld., V.I—17, Australia”), Indonesia (1 © (RMNH), “Dammerman,

Idjen, 950 m, Blawan, VI.1924”; 1 3 (RMNH), “Museum Leiden, J. v. d. Vecht,

G. Bentang, III.1938”; 1 9 (RMNH), “Museum Leiden, J. v. d. Vecht, G.

Tjangkoedang, Djampang Wetan, X1.1938°), India (2 9 (CNC, RMNH),

“Anamalai Hills, Cinchona, India, 3500’ (ft), IV.1957, P. S. Nathan”; 2 g (CNC,

RMNH), “Anamalai Hills, Madras St., S. India, 3500’ (ft), V.1963, P. S. Nathan” 2

d (CNC), “Devala, Nilgiri Hills, 3200’ (ft), S. India, X.1960, P. S. Nathan”), and

Taiwan (1 © (TMA), “Formosa, Sauter”, ““Chip-Chip, (1)909, II’; 1 g (TC),

“Bukai, Formosa, VI—11°34’’, “L. Gressitt Collector”).

Variation: Length of fore wing 10.7—15.7 mm; antennal segments 47—52;

length of malar space 0.4—0.8 times basal width of mandible; length of maxillary

palp 1.5—1.7 times height of head; length of Ist tergite 3.0—3.1 times its apical

width, length of ovipositor sheath 0.05 times fore wing (in five specimens

measured); hind tarsus sometimes rather whitish-yellow or whitish, mesoscutum

partly and apex of metasoma sometimes infuscated; vein 2-SC+R sometimes

quadrate. |

Homolobus (Chartolobus) nigritarsis spec. nov.

(figs. 370— 384)

Holotype, 9, length of body 12.3, of fore wing 12.0 mm.

Head. — Antennal segments 51, 3rd—7th segments with a rather straight ridge

(fig. 381), 3rd segment 1.4 times 4th segment, length of 3rd and 4th segments 4.7

and 3.4 times their width, respectively, length of both penultimate segments 2.0

and 2.1 times their width; length of maxillary palp 1.5 times height of head; eyes

weakly emarginate (fig. 374); dorsal length of eye 2.6 times temple; temple directly

narrowed posteriad (fig. 375); POL : & ocellus : OOL = 11 : 10 : 8; frons concave,

smooth; vertex almost flat, indistinctly coriaceous; face almost flat, mainly

rugulose-coriaceous and rather dull, but medially punctulate and more shiny (fig.

374); clypeus convex, punctulate; length of malar space 0.4 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

medially and posteriorly crenulate, and dorsally somewhat punctate (fig. 370);

epicnemial area smooth, except fore some punctulation; anterior half of precoxal

suture shallowly crenulate dorsally and punctate ventrally, its posterior half

smooth dorsally and finely punctate ventrally; rest of mesopleuron smooth, except

for some punctulation; metapleural flange large, rounded apically, lamelliform,

with carinae (fig. 370); metapleuron smooth dorsally, reticulate ventrally; notauli

indistinctly crenulate, only posteriorly wider crenulate (fig. 384); surface of

propodeum coarsely and remotely reticulate, anteriorly and posteriorly narrowly

smooth.

Wings. — Fore wing: r: 3-SR : SRI = 13: 21 : 60; SRI almost straight (figs. 379);

cu-a almost straight, postfurcal; 1-CU1 : 2-CUl = 3: 25; 2-SR : 3-SR : r-m = 18:

21:12; 2A very wide and sclerotized basally (figs. 379, 380); area basally of 2A

bare, basally and ventrally brownish pigmented (fig. 380). Hind wing: Basal fifth of

VAN ACHTERBERG: Revision Zelinae auct. 311

SR sclerotized, rather straight (fig. 379); SC + RI rather curved (fig. 382);

marginal cell scarcely constricted.

Legs. — Hind coxa punctulate; tarsal claws with a sharp subapical lamelliform

tooth (fig. 378), setose, and basally indistinctly pectinate, except inner hind claw

(fig. 383); length of femur, tibia, and basitarsus of hind leg 6.1, 9.7, and 7.2 times

their width, respectively; length of spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.1 times its apical width, its surface

posteriorly and laterally partly rather weakly reticulate-rugose (fig. 372); dorsal

carinae of Ist tergite shortly developed basally; length of ovipositor sheath 0.06

times fore wing.

Colour. — Brownish-yellow; stemmaticum, antenna (but scapus and pedicellus

somewhat reddish), apical 0.6 of hind tibia, hind tarsus (but telotarsus rather

reddish), pterostigma, parastigma, and most wing veins, more or less dark brown;

basal half of wing membrane yellowish, its apical half infumate.

Holotype in RMNH, Leiden: “Neth. Ind.-American New Guinea Exped.,

Araucaria Camp, 800 m, 23.111.1939, L. J. Toxopeus”. For location of the camp,

see Toxopeus (1940). Paratypes: 1 9,2 8; 1g (TC), “Wau & Mt. Kaindi, 4—6500’

(ft), N. Guinea, June 17—22, 1962, Bernd Heinrich”; 1 & (TC), ‘“Wau, N. Guinea,

October, 1969, P. Shanahan’; 1 © (BPBM), “New Guinea (NE), Wau, Morobe

Distr., 1200 m, 11—18.x.1961”. Length of fore wing 9.1—11.1 mm, antennal

segments 49 (one 3), length of Ist tergite 3.1 times its apical width, body partly

rather whitish yellowish; antenna of 9 paratype baso-ventrally undulate and

lamelliform, as in undulatus.

Subgenus Homolobus Foerster

Foerster, 1862, Verh. naturh. Ver. preuss. Rheinl. 19: 256.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

Type-species: Phylax discolor Wesmael.

Diagnosis. — Length of body 5.1—11.5, of fore wing 5.3—10.6 mm; antennal

segments 42—48, its 3rd—6th segments of 9 with a longitudinal ridge at the inner

side (figs. 391, 424, 451); length of 4th segment of labial palp 3.5—6.0 times 3rd

segment; length of maxillary palp 1.2—1.7 times height of head; apical margin of

clypeus straight (fig. 388) or rather convex (fig. 411) apically, not (fig. 461) or

weakly (fig. 445) differentiated from clypeus; length of malar space 0.4—1.3 times

basal width of mandible; eyes weakly emarginate (figs. 388, 411); temples roundly

(fig. 409) or directly (fig. 429) narrowed posteriad; length of hind femur 5.8—6.9

times its width; claws with a small subapical tooth (fig. 426), bifurcate (fig. 406),

with a lamella (fig. 392), or double lamella (fig. 394); inner hind claw of 9 concave

and glabrous ventro-basally (figs. 427, 439, 452, 885—887); hind telotarsus of 9

more or less bare near base of inner hind claw (figs. 885—887); apices of hind tibial

spurs of Z sharp and hyaline (fig. 410); 1A +2A of fore wing straight (figs. 396,

436); basal third of SR of hind wing only pigmented, unsclerotized, straight or

nearly so (figs. 402, 446, 469); SC+ RI of hind wing straight (fig. 404) or weakly

curved (fig. 434); r of hind wing absent (fig. 396) or present (fig. 469); length of Ist

312 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

tergite 1.7—3.0 times its apical width; 2nd tergite smooth (fig. 400) or sculptured

(fig. 414); length of ovipositor sheath 0.06—0.39 times fore wing; posterior part of

propodeum not (fig. 444) or distinctly (fig. 401) separated from antero-dorsal part.

Distribution. — The subgenus Homolobus is restricted to the Palaearctic and

Afrotropical regions. Because the shape of the inner hind claw of the 9 is

unknown, the relationship of rugosus, and to a lesser degree of simplex, with the

other species of the subgenus is uncertain. The Palaearctic has three species: one

widely distributed Palaearctic species (discolor), and two East Palaearctic species,

of which simplex is an aberrant species, not closely related to dauricus.

There are four Afrotropical species, of which one (rugosus) is very aberrant, but

the other three are closely interrelated. One of them is restricted to the African

continent (ethiopicus), while two others are restricted to Malagasy.

Key to the species of the subgenus Homolobus

1. Claws bifurcate (fig. 406); 2nd tergite rugose (fig. 414); vertex punctate (fig.

409); vein SC + RI of hind wing short, RI mainly absent, and hamuli separated

from RI (bg..412)2Malagasy 72. poets a rugosus spec. nov. (p. 314)

— Claws with lamella (figs. 390, 394) or with a small subapical tooth (fig. 426);

2nd tergite smooth (fig. 428); vertex punctulate (fig. 429); vein SC + RI of hind

wing longer, RI present, short, and hamuli situated at RI (fig. 425) .... 2

2. Claws of 9 and fore claw of ¢ with a ventral lamella (figs. 390, 392), middle

and hind claws of & with a 2nd lamella situated on the Ist lamella (figs. 393,

394); propodeum coarsely areolate, with its surface mainly smooth (fig. 400);

East Palaearctic: ur. genes EEE simplex (Watanabe) (p.313)

areolate, or, if areolate, then surface densely rugose posteriorly (fig. 431) 3

3. Precoxal suture, its surroundings and hind coxa coarsely sculptured (figs. 416,

431); at least base of palpi infuscated; vein r of hind wing absent (fig. 418);

Afrotropieal oe. Ens im Tr CEO en ERE 4

— Precoxal suture (except anteriorly), its surroundings and hind coxa at most

punctulate, usually smooth (figs. 459, 467); palpi whitish or yellowish; vein r of

hind wing present (figs. 460, 469); Palaearctic .................. 6

4. Second tergite whitish; subapical tooth of tarsal claws of 9 scarcely visible at

80 x (fig. 443) or, if easily visible, then length of ovipositor sheath 0. 14—0. 17

times fore wing;)Malagasy 1.4 sleten oe SEE 5

— Second tergite dark brown and partly reddish- or yellowish-brown; subapical

tooth of tarsal claws of 9 easily visible at 80 x (figs. 452, 455); length of

ovipositor sheath 0.07—0.08 times for wing; African Continent .........

tert bake ey ee oer NE DE AE ethiopicus spec. nov. (p. 318)

5. Length of ovipositor sheath 0.14—0.17 times fore wing, about as long as apical

height of metasoma, slender as ovipositor (fig. 416); subapical tooth of tarsal

N ba eine ee Ga re Sn cingulatus (Granger) (p. 315)

— Length of ovipositor sheath 0.06—0.09 times fore wing, distinctly shorter than

VAN ACHTERBERG: Revision Zelinae auct. 313

apical height of metasoma, rather stout as ovipositor (fig. 431); subapical tooth

MR eke erkeer rebel ea inopinus spec. nov. (p. 316)

6. Length of ovipositor sheath 0.09—0.12 times fore wing, short (fig. 459);

propodeum without an areola, smooth, except for some rugae (fig. 459);

mesopleuron smooth (fig. 459); Palaearctic ... discolor (Wesmael) (p. 319)

— Length of ovipositor sheath 0.36—0.39 times fore wing, comparatively long

(fig. 467); propodeum with a suboval areola, surrounded by rugosity (fig. 467);

mesopleuron punctulate (fig. 467); East Palaearctic .................

Eer eh ete nader an RR A dauricus Shestakov (p. 320)

Homolobus (Homolobus) simplex (Watanabe) comb. nov.

(figs. 385— 400)

Watanabe, 1932, Insecta matsum. 6: 135, 136, fig. (as Zele).

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319, 324, 325, figs. 4, 5.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 226.

Holotype, 9, length of body 9.6, of fore wing 9.7 mm.

Head. — Remaining antennal segments 26, apical segments missing, 3rd

segment 1.3 times 4th segment, length of 3rd and 4th segments 4.4 and 3.5 times

their width, respectively; length of maxillary palp 1.5 times height of head; dorsal

length of eye 2.1 times temple; temple roundly narrowed posterially (fig. 387);

POL : @ ocellus : OOL = 4: 8: 5; frons almost flat and smooth; vertex dull, flat

and coriaceous (fig. 387); face rather flat, laterally rather dull, coriaceous,

medially rugulose; clypeus remotely punctate, rather convex; apical margin of

clypeus not differentiated, rather thick, and almost straight medially (fig. 388);

length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

rugose and crenulate medially, ventrally and dorsally mainly smooth (fig. 385);

epicnemial area rugose; precoxal suture largely reticulate, smooth apically; rest of

mesopleuron punctulate; metapleural flange large, sharp and _ narrowly

lamelliform apically (fig. 385); metapleuron largely punctulate, coarsely rugose

ventrally; notauli extensively crenulate (fig. 398); mesoscutal lobes punctulate;

surface of propodeum coarsely areolated, the enclosed areas smooth and with a

short medial carina anteriorly (fig. 400); posterior part of propodeum not well

separated from antero-dorsal part (fig. 385).

Wings. — Fore wing: r: 3-SR : SRI = 9: 15: 53; SRI almost straight (fig. 396);

cu-a inclivous, shortly postfurcal; 1-CUI : 2-CU1 = 2: 23; 2-SR : 3-SR : r-m = 13:

15 : 7; 2A sclerotized basally (fig. 396); area basally of 2A bare except for ca. 8

setae apically. Hind wing: r absent; SC + RI weakly curved (fig. 399).

Legs. — Hind coxa punctate dorsally, punctulate laterally; hind tarsal claws

absent; fore and middle claws with a rather large ventral lamella, yellowish

pectinate; length of femur, tibia and basitarsus of hind Jeg 6.2, 10.0 and 8.2 times

their width, respectively; length of spurs of hind tibia 0.8 and 0.6 times basitarsus.

314 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface

reticulate-rugose, medially weakly developed, with a smooth tubercle apically (fig.

400); dorsal carinae of Ist tergite absent; 2nd tergite smooth; length of ovipositor

sheath 0.06 times fore wing.

Colour. — Brownish-yellow; stemmaticum and apices of antennal segments

(except scapus and pedicellus), blackish.

Holotype in EI, Sapporo: “Hokkaido, Uchida/Jôzankei, 15/8-1925”, “Type”,

“Zele simplex Watanabe, 9, Type”, “Type Hym. No. 23”. One ¢ additionally

examined: (EI) “Sapporo, Hokkaido, 17.VII.1964, H. Takada’, “Zele simplex

Watanabe, &, Det. C. Watanabe, 1969”, with very peculiar middle and hind tarsal

claws (figs. 393, 394), ventrally with a double lamella. Fore claw as in 9, length of

fore wing 8.6 mm, length of Ist tergite 2.5 times its apical width, weakly rugulose,

further as holotype.

Homolobus (Homolobus) rugosus spec. nov.

(figs. 401—414)

Holotype. &, length of body 11.5, of fore wing 10.6 mm.

Head. — Remaining antennal segments 34, apical segments absent, 3rd segment

1.4 times 4th segment, length of 3rd and 4th segments 4.2 and 3.1 times their width,

respectively; length of maxillary palp 1.2 times height of head; dorsal length of eye

1.5 times temple; temple roundly narrowed posteriad (fig. 409); POL : @ ocellus:

OOL = 6: 8: 7; frons rather flat, smooth; vertex remotely punctate, rather flat (fig.

409); face rather flat, densely and coarsely punctate, with some striae dorsally;

clypeus rather flat, remotely punctate; apical margin of clypeus thin, not

differentiated, convex ventrally (fig. 411); length of malar space 0.4 times basal

width of mandible; mandible only slightly twisted apically.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

posteriorly and ventrally rugose, antero-medially with a crenulate groove, and rest

of pronotum mainly punctate (fig. 401); epicnemial area mainly smooth; precoxal

suture smooth; rest of mesopleuron mainly weakly punctate; metapleural flange

rounded apically, large, thick, rugose and not lamelliform (fig. 401); metapleuron

coarsely rugose ventrally, and punctulate medially, anteriorly widely impressed;

notauli rather narrowly crenulate (fig. 413); mesoscutal lobes punctulate; dorsal

surface of propodeum coarsely transversely rugose and medial carina absent; the

short posterior part of propodeum well separated from dorsal part (fig. 401),

mainly smooth except for some carinae, and with a narrow areola posteriorly (fig.

414).

Wings. — Fore wing: r: 3-SR: SRI = 12: 11:58; SRI almost straight (fig. 402);

cu-a postfurcal, almost straight; 1-CU1: 2-CU1 = 2: 26; 2-SR: 3-SR: r-m = 16:11:

9; 2A scarcely sclerotized (fig. 402); area basally of 2A mainly bare (fig. 407). Hind

wing: r absent; SC+RI short, straight, somewhat widened anteriorly (fig. 404);

hamuli separated from the mainly absent RI (fig. 412).

Legs.— Hind coxa weakly punctate, with some striae apically, and ventrally,

more coarsely punctate (fig. 410); tarsal claws bifurcate, because of a large sharp

VAN ACHTERBERG: Revision Zelinae auct. 315

subapical tooth, which is situated at the inner side of the claw, only fore claws

somewhat pectinate (fig. 406); length of femur, tibia and basitarsus of hind leg 5.8,

9.9, and 7.0 times their width, respectively; length of spurs of hind tibia 0.6 and 0.4

times basitarsus.

Metasoma. — Length of Ist tergite 2.9 times its apical width, its surface coarsely

rugose (fig. 414); dorsal carinae of Ist tergite present in its basal fifth; 2nd tergite

rugose (fig. 414).

Colour. — Brownish-yellow; face, ventral third of temple, and eye margins

dorsally, whitish-yellow; head dorsally, antenna basally (except annellus), apex of

metasoma, most wing veins, dark brown; middle and hind tarsi (except telotarsi),

white; wing membrane hyaline; pterostigma brown.

Holotype in MNHN, Paris: “Madagascar Est, Marojejy, res. nat. int. XII,

Anjanaharibe S., 1600 m, III.1961, P. Soga”.

Note. — Because the female is unknown, the inclusion of this species in the

subgenus Homolobus is only tentative. The other possibility is the subgenus

Oulophus, but because it does not fit well in there and the subgenus Oulophus is

unknown from the Afrotropical region, I prefer to include it in the subgenus

Homolobus. H.rugosus is a peculiar species because of the remarkable

combination of apomorphous character-states (e.g., the shape of SC+RI, the

reduced RI, the separated hamuli, the large ocelli and antescutal depression, and

smooth precoxal suture) and of plesiomorphous character-states (e.g., the

bifurcate claws, the separated areolated posterior part of propodeum, the thick,

non-lamelliform metapleural flange, the presence of the dorsal carinae of the Ist

tergite, and the sculptured 2nd tergite).

Homolobus (Homolobus) cingulatus (Granger) comb. nov.

(figs. 415—430)

Granger, 1949, Mém. Inst. scient. Madagascar 2A: 378, fig. 382 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 223.

Lectotype, 9, length of body and of fore wing both 6.6 mm.

Head. — Remaining antennal segments 29, apical segments absent, 3rd segment

1.1 times 4th segment, length of 3rd and 4th segments 4.1 and 3.7 times their width, _

respectively; length of maxillary palp 1.3 times height of head; dorsal length of eye

2.3 times temple; temple directly narrowed posteriad (fig. 429); POL : @ ocellus :

OOL = 6: 9: 8; frons rather flat, with some short crenulae; vertex punctulate,

rather flat (fig. 429); face rather flat, punctate, dorsally somewhat rugose; clypeus

punctulate, rather flat; apical margin of clypeus thin, not differentiated, convex

ventrally (fig. 430); length of malar space 1.1 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

crenulate antero-medially, rugose ventrally and posteriorly, and punctate dorsally

(fig. 416); epicnemial area rugose; precoxal suture widely punctate-rugose; rest of

mesopleuron finely punctate (fig. 416); metapleural flange large, thick, rugose,

narrowly lamelliform apically, rather round apically (fig. 416); metapleuron

punctulate medially, rugose ventrally and posteriorly; notauli rather narrowly

316 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

crenulate (fig. 421); mesoscutal lobes punctulate; dorsal surface of propodeum

almost smooth anteriorly, reticulate-rugose medially and posteriorly areolated,

enclosed areas mainly smooth; posterior part of propodeum not separated (fig.

416); medial carina of propodeum absent.

Wings. — Fore wing: r : 3-SR : SRI = 9:15:71, SRI straight; cu-a interstitial in

left wing, shortly postfurcal in right wing (fig. 418) and 1-CUI : 2-CUI = 1: 15; 2-

SR : 3-SR : r-m = 16: 15: 11; 2A shortly sclerotized basally (fig. 418); area basally

of 2A mainly bare, except distally (fig. 419). Hind wing: r absent; SC + RI weakly

curved (fig. 425); hamuli at RI.

Legs. — Hind coxa finely rugose dorsally, laterally and ventrally punctate; tarsal

claws with small subapical tooth, which is somewhat more developed than in

inopinus and well visible at 80x (fig. 426), pectinate basally, except inner hind claw

(fig. 427); length of femur, tibia and basitarsus of hind leg 6.9, 10.5 and 8.8 times

their width, respectively; length of spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.2 times its apical width, its surface smooth

basally and its posterior half rugulose (fig. 428); dorsal carinae of Ist tergite

absent; 2nd tergite smooth; length of ovipositor sheath 0.17 times fore wing,

slender, and almost as long as apical height of metasoma (fig. 416).

Colour. — Light reddish-brown; palpi (except for 6th segment of maxillary

palp), tarsi, apices of tibiae and trochanters partly, hind coxa apically, more or less

infuscated; pterostigma and wing veins, brown; metasoma black, except for the

2nd tergite and base of the 3rd tergite, which are white.

Lectotype in MNHN, Paris: “Madagascar, Ankaratra, alt. 1800 [m]”, “1/1’40”,

“Museum Paris, A. Seyrig”, “44” (= antennal segments), “Type”. Lectotype

herewith selected and labelled accordingly. Paralectotypes examined: | 9 and 8

&, of which at least the 9 belongs to the new species inopinus. Additional

Ambatolampy; Ampitameloka, 840 m, Sud Moramanga) (MNHN, RMNH).

Variations: Length of fore wing 5.3—6.7 mm; length of 4th segment of labial palp

4.3—5.0 times 3rd segment; length of ovipositor sheath 0.14—0.17 times fore wing;

some males have the flagellum dark brown.

Note. H. cingulatus as defined in this paper is easily recognizable if females are

present, because of the comparatively long and slender ovipositor sheath,

combined with the whitish 2nd tergite and rather long malar space. The male of

inopinus is unknown and may be confused with the males of cingulatus, but

cingulatus may be separated by the somewhat longer 4th segment of the labial palp

compared with the 3rd segment (length of 4th segment of labial palp 4.35.0 times

its 3rd segment in cingulatus, and 3.5—4.0 times in inopinus).

Homolobus (Homolobus) inopinus spec. nov.

(figs. 431—443)

Holotype, 9, length of body and of fore wing both 7.1 mm.

Head. — Remaining antennal segments 37, but apical segments absent, 3rd

segment 1.2 times 4th segment, length of 3rd and 4th segments 4.4 and 3.8 times

VAN ACHTERBERG: Revision Zelinae auct. 31197

their width, respectively; length of maxillary palp 1.5 times height of head; dorsal

length of eye 2.4 times temple; temple directly narrowed posteriad (fig. 442); POL:

@ ocellus: OOL = 3: 5 : 5; frons weakly concave, with some rugae; vertex rather

flat, punctulate; face rather flat, weakly punctate; clypeus convex basally, weakly

punctate; apical margin of clypeus not differentiated, straight medio-ventrally (fig.

437); length of malar space 1.1 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

crenulate antero-medially, punctulate dorsally, and rugose posteriorly and

ventrally (fig. 431); epicnemial area mainly rugose; precoxal suture rather coarsely

and widely reticulate-rugose; rest of mesopleuron densely punctulate, more

punctate near pleural suture; metapleural flange rather large, rounded and

lamelliform apically (fig. 431); metapleuron rugose ventrally, punctulate dorsally;

notauli narrowly crenulate anteriorly, widely crenulate posteriorly (fig. 438);

mesoscutal lobes finely and densely punctulate; surface of propodeum reticulate-

rugose anteriorly, only medially remotely reticulate posteriorly, with a short

medial carina anteriorly, its posterior part not separated (fig. 431).

Wings.— Fore wing: r : 3-SR : SRI = 11 : 16: 74; SRI straight; cu-a straight,

postfurcal; 1-CUI : 2-CU1 = 1: 17; 2-SR : 3-SR : r-m = 18: 16: 12; 2A shortly

sclerotized basally (fig. 435); area basally of 2A mainly bare (fig. 436). Hind wing: r

absent; SC +R1 weakly curved (fig. 434); hamuli at RI.

Legs. — Hind coxa coarsely (but basally rather weakly) punctate; tarsal claws

with a minute subapical tooth, scarcely visible at 80 x (figs. 439, 443); length of

femur, tibia and basitarsus of hind leg 6.7, 9.7, and 9.2 times their width,

respectively; length of inner spur of hind tibia 0.7 times basitarsus.

Metasoma. — Length of Ist tergite 2.9 times its apical width, its surface rather

shallowly reticulate-rugose, but medially and basally smooth (fig. 441); dorsal

carinae of Ist tergite absent; 2nd tergite smooth; length of ovipositor sheath 0.06

times fore wing, somewhat widened apicad, distinctly shorter than apical height of

metasoma (fig. 431).

Colour. — Light reddish-brown; tarsi and tibiae somewhat infuscated; antenna

(except scapus, pedicellus, and apex of antenna), palpi, hind trochanters, and

pterostigma, more or less dark brown; Ist tergite, 3rd tergite mainly and following

posterior part of metasoma, black; basal half of metasoma (except Ist tergite)

yellowish-white.

Holotype in MNHN, Paris: “Madagascar, Bekily, Reg. Sud. de l'Ile”, “Museum

Paris, IV.38, A. Seyrig”, “Zele cingulatus Gr., B. Sigwalt’’. This specimen is also a

paralectotype of cingulatus; I have selected the other 9 as lectotype of cingulatus

because it agrees better with the original description (“tarière aussi longue que le

metatarse postérieur”). Paratype: 1 9 (RMNH), “La Mandraka, 1250 m,

Manjakandria, 30.X.56, A.R.”; length of fore wing 5.3, of body 6.1 mm; length of

ovipositor sheath 0.09 times fore wing; precoxal suture extensively rugose-

punctate; length of 4th segment of labial palp 4.0 times 3rd segment; length of

malar space 1.2 times basal width of mandible; antennal segments 46; length of

both penultimate segments of antenna 1.9 times their width; propodeum only

weakly sculptured.

318 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Homolobus (Homolobus) ethiopicus spec. nov.

(figs. 444—457)

Holotype, 9, length of body 7.2, of fore wing 6.9 mm.

Head. — Antennal segments 44, but apical segments missing, 3rd segment 1.2

times 4th segment, length of 3rd and 4th segments 4.4 and 3.6 times their width,

respectively; length of maxillary palp 1.4 times height of head; dorsal length of eye

3.5 times temple; temple directly narrowed posteriad (fig. 456); POL : @ ocellus :

OOL = 6: 10: 11; frons weakly concave and somewhat rugose; vertex almost flat,

punctulate-coriaceous (fig. 456); face weakly convex, coriaceous, dorsally and

medially rugose clypeus remotely punctate, rather convex; apical margin of

clypeus thin, weakly differentiated from clypeus, and weakly convex ventrally (fig.

445); length of malar space 0.8 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

densely reticulate-rugose, medio-anteriorly crenulate and dorsally punctulate (fig.

444); epicnemial area rugose-punctate; precoxal suture coarsely punctate-.

reticulate; rest of mesopleuron punctate; metapleural flange large, narrowly

lamelliform and rather sharp apically (fig. 444); metapleuron punctate medially,

ventrally reticulate, and dorsally more punctulate; notauli anteriorly narrowly,

and posteriorly widely crenulate (fig. 454); mesoscutal lobes densely punctulate;

surface of propodeum largely densely reticulate-rugose, anteriorly almost smooth,

medial carina absent, and its posterior part not separated (fig. 444).

Wings. — Fore wing: r : 3-SR : SRI = 6: 8: 39; SRI straight; cu-a almost

straight, shortly postfurcal; 1-CUI : 2-CUl = 1: 17; 2-SR : 3-SR : r-m = 10:8: 6,

basal half of 2A sclerotized (fig. 446); area basally of 2A bare (fig. 449). Hind wing:

r absent; SC + RI somewhat curved (fig. 448); hamuli at RI.

Legs. — Hind coxa densely coriaceous-rugose dorsally, laterally punctate; tarsal

claws with a small subapical tooth, well visible at 80 x (figs. 452, 455), setose;

length of femur, tibia and basitarsus of hind leg 6.4, 9.2, and 8.7 times their width,

respectively; length of spurs of hind tibia 0.7 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.0 times its apical width, its surface rugulose

(fig. 457); dorsal carinae of Ist tergite absent; 2nd tergite smooth; length of

ovipositor sheath 0.07 times fore wing.

Colour. — Reddish-brown; antenna (but annellus and apical half of antenna

more light brown), mandibles, base of palpi, propleuron, pronotum ventrally, face

sublaterally, stemmaticum, vertex medially, occiput (except near eyes), temples

posteriorly, metasoma (but Ist tergite, except for its apex, more reddish-brown;

ventral half of metasoma and sides of 2nd notum, yellowish-brown), tegula

medially, pterostigma, wing veins, fore and middle coxae, trochanters, and

femora, middle tibia, telotarsi, hind coxa posteriorly, hind trochanters, femur, and

tibia, more or less dark brown; palpi except for their bases, yellowish.

Holotype in CNC, Ottawa: “Tanganyika, W. Usambara Mts., 1600 m, Lushoto,

1—12.i1.1933”, “S.Africa, Ri E. Turner, Brit. Mus... 1933-1982: Ome):

“Mpendle, Natal, XII-3-70, S. Afr., H. & M. Townes”; 1 & (TC, allotype),

topotypic with 9 from Natal. Variation: Length of fore wing 6.2—6.9 mm; length

VAN ACHTERBERG: Revision Zelinae auct. 319

of malar space 0.8—1.3 times basal width of mandible; length of Ist tergite

2.4—3.0 times its apical width; length of ovipositor sheath 0.07—0.08 times for

wing; hind and middle claws of & virtually without subapical tooth (as in

cingulatus); sometimes only coxae, propleuron (partly), pronotum ventrally and

metasoma (except for Ist tergite), dark brown.

Homolobus (Homolobus) discolor (Wesmael) comb. nov.

(figs. 169, 170, 458—466, 885—887)

Wesmael, 1835, Nouv. Mém. Acad. Brux. 9: 162 (as Phylax).

Snellen van Vollenhoven, 1858, in Herklots: Bouwstoffen Fauna Nederland 2: 282 (Phylax aestivalis).

Syn. nov.

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319, 320, fig. 3.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 223.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 230, 231.

Capek, 1975, Biologia 30(11): 819.

Tobias, 1976, Opr. Fauna SSSR 110: 131, fig. 39:1.

Head. — Remaining antennal segments 38, apical segments missing, 3rd

segment 1.1 times 4th segment, length of 3rd and 4th segments 4.0 and 3.6 times

their width, respectively; length of maxillary palp 1.7 times height of head; dorsal

length of eye 2.4 times temple; temple directly narrowed posteriad (fig. 458); POL

: @ ocellus : OOL = 6: 6: 6; frons almost flat and smooth, with some striae

laterally; vertex smooth, weakly convex; face rather flat, coriaceous laterally and

superficially transversely rugose (fig. 461); clypeus rather flat, smooth, except for

some punctulation; apical margin of clypeus not differentiated, thin, slightly

convex ventrally (fig. 461); length of malar space 0.6 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, except for some medial crenulae and rugae posteriorly (fig. 459);

epicnemial area smooth; precoxal suture smooth, mainly absent; metapleural

flange medium-sized, lamelliform, round apically (fig. 459); metapleuron almost

smooth; notauli indistinctly crenulate (fig. 170); mesoscutal lobes indistinctly

punctulate; surface of propodeum mainly smooth, except for some short carinae

posteriorly and a short medial carina anteriorly, its posterior part not separated

from antero-dorsal part (fig. 459).

Wings. — Fore wing: r: 3-SR : SRI = 10: 22: 91; SRI almost straight (fig. 460);

cu-a weakly inclivous, postfurcal; 1-CUI : 2-CU1 = 2: 21; 2-SR : 3-SR : r-m = 23:

22 : 11; 2A shortly sclerotized basally (fig. 460); area basally of 2A bare. Hind

wing: r present; 2A faintly indicated by pigmentation (fig. 460); SC+ RI rather

curved (fig. 460); hamuli at R1.

Legs. — Hind coxa smooth; tarsal claws with a small, sharp, subapical tooth (fig.

465), setose; length of femur, tibia, and basitarsus of hind leg 6.9, 10.3, and 9.0

times their width, respectively; length of spurs of hind tibia 0.6 and 0.5 times

basitarsus.

320 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Metasoma. — Length of Ist tergite 1.9 times its apical width, its surface smooth,

except for some microsculpture laterally (fig. 466); dorsal carinae of Ist tergite

faintly indicated in front of spiracles; 2nd tergite smooth; length of ovipositor

sheath 0.10 times fore wing.

Colour. — Dark brown; body more or less yellowish-brown ventrally; mandibles

(mainly), palpi, legs (except apical 0.6 of hind tibia), and upper hind corner of side

of pronotum, yellow; dorso-apical 0.6 of hind tibia brown.

Lectotype in KBIN, Brussels: “Coll. Wesmael”, “1877”, ““Phylax discolor mihi,

Q, det C. Wesmael”, “Type”. The type-locality is the surroundings of Brussels.

Lectotype herewith selected and labelled accordingly. There are two

paralectotypes, 9, with the same labels as the lectotype. There is a further 9 in

the Wesmael Collection which does not belong to the type-series (“4 JI. 1852,

from Finland (Helsinki; Degerö, Hels.), Sweden (Höör, Skane), Denmark

(Klaekket; Mglbaek; Bukke, Skov.; Tandby; Satruphole; Sondby), England

(Dorking, Surrey; Box Hill, Surrey), Ireland (Saggart, Co. Du.; Carrowgarry, Co.

SI.; Trawalua, Co. Sl), Netherlands (Plasmolen, Z.L.; E. bank Wijde Aa, nr.

Woubrugge; Wijster; Oisterwijk; Asperen; Naardermeer, at light; Crailo, at light;

Asselt, at light; Ede; Flevopolder), West Germany (Wiesen, Spessart;

Geierlambach; Reither Alm, 850 m; Mainz; Ellmau, ca. 1050 m; Tegernsee; nr.

Reval), East Germany (Thüringen), Austria (Styr., Podéetrtek), Switzerland

(Mülenen, 800 m), USSR (Irkutskaja obl., garden), Japan (Kamikochi; Kyoto,

Honshu) (RMNH, HC, CNC, TC, ZMH, ZMB, ZSB, UZM, ITZ, EI, ZIL, USNM,

WHC). Variation: Length of fore wing 7.2—8.9 mm, antennal segments 43—48;

length of Ist tergite 2.1—2.2 times its apical width; length of ovipositor sheath

0.09—0.12 times fore wing; anterior half of vein r of hind wing sometimes absent;

cocoon whitish. The only known host of the specimens examined is Odontopera

bidentata (Clerck), which belongs to the Geometridae (Lepidoptera).

Note. The type of Snellen van Vollenhoven’s aestivalis is lost (Van Achterberg,

1974a: 23), but as already indicated in the original description it is morphologically

close to discolor. Knowing the Dutch fauna, and considering the ability of Snellen

van Vollenhoven to identify correctly despite a lack of literature, I do not hesitate

to synonymize aestivalis with discolor.

Homolobus (Homolobus) dauricus Shestakov

(figs. 467—480)

Shestakov, 1940, Ark. Zool. 32A: 18.

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319, 320, figs. 1, 2 (p.p.).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 223.

Holotype, 9, length of body 7.6, of fore wing 7.4 mm.

Head. — Antennal segments 46, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 3.9 and 3.2 times their width, respectively, length of two

penultimate segments 2.0 and 2.4 times their width; length of maxillary palp 1.4

VAN ACHTERBERG: Revision Zelinae auct. 321

times height of head; dorsal length of eye 2.0 times temple; temple directly

narrowed posteriad (fig. 474); POL : & ocellus : OOL = 3: 7: 4; frons rather flat

and smooth; vertex rather flat, punctulate-coriaceous (fig. 474); face rather flat,

laterally coriaceous, dorso-medially punctate-rugose and ventro-medially punc-

tate; clypeus weakly convex, remotely punctate; apical margin of clypeus not well

differentiated, thin, almost straight medially (fig. 479); length of malar space 0.4

times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

coarsely crenulate medially and posteriorly, and striate ventrally (fig. 467);

epicnemial area crenulate; precoxal suture punctulate as rest of mesopleuron,

mainly absent; metapleural flange large, lamelliform, wide, and round apically;

metapleuron punctulate, but rugose-carinate; notauli coarsely and narrowly

crenulate (fig. 473); mesoscutal lobes densely punctulate; surface of propodeum

smooth anteriorly, medially coarsely transversely rugose and with a suboval areola

and costal carinae present (cf. fig. 352), its posterior part not separated from

antero-dorsal part (fig. 467).

Wings. — Fore wing: r: 3-SR : SRI = 5: 11:40; SRI slightly curved (fig. 469);

cu-a somewhat inclivous, but posteriorly curved basad (fig. 469), postfurcal; 1-

CUI : 2-CUl = 1: 17; 2-SR : 3-SR : r-m = 10: 11 : 6; 2A well-developed and

sclerotized basally (fig. 469); area basally of 2A mainly bare. Hind wing: r present;

SC +RI evenly curved (fig. 480); hamuli at Ri.

Legs. — Hind coxa punctulate; tarsal claws with a small subapical tooth (figs.

475, 476), indistinctly yellowish pectinate, except inner hind claw; length of femur,

tibia, and basitarsus of hind leg 5.9, 10.8, and 8.8 times their width, respectively;

length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 1.7 times its apical width, its surface smooth,

but posterior half weakly and irregularly rugose (fig. 478); dorsal carinae weakly

developed in front of spiracles; 2nd tergite smooth; length of ovipositor sheath

0.39 times fore wing.

Colour. — Dark reddish-brown; pterostigma dark brown; palpi, dorso-apical

corner of pronotum, and tegulae whitish-yellow; fore and middle legs, light

yellowish; eye margin dorsally, vertex, metapleuron mainly, and hind leg mainly,

brownish-red; trochanter yellowish and apical two-thirds of hind tibia dark brown;

hind tarsus and spurs, reddish; apical third of antenna light brown.

Holotype in NR, Stockholm: ‘Vladivostok, Sedanka, Malaise/ 10/8.30”,

“Homolobus dauricus sp. n. typ., det. Shestakov”, “402, 77”, “Riksmuseum

Stockholm”. Additional specimens examined: (1 9 and 2 &, EI) from Japan

(Hirakura, Mie Honshu; Hikosan, Kyushu; Sapporo, Hokkaido). Length of fore

Subgenus Phylacter Reinhard

Wesmael, 1835, Nouv. Mém. Acad. Brux. 9: 159 (as Phylax nec Dahl, 1823).

Reinhard, 1863, Berl. ent. Z. 7: 248 (nom. nov. for Phylax Wesmael).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

322 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Type-species: Rogas annulicornis Nees.

Synonym: Phylax Wesmael, 1835 nec Dahl, 1823.

Diagnosis. — Length of body 6.0—10.9, of fore wing 6.3—11.5 mm, antennal

segments 46—55, its 3rd-6th segments of 92 without a longitudinal ridge; length of

4th segment of labial palp 2.6—3.1 times 3rd segment; length of maxillary palp

1.5—1.6 times height of head; apical margin of clypeus rather convex, not

differentiated from clypeus (figs. 487, 500, 509); length of malar space 0.5—1.0

times basal width of mandible; eyes weakly emarginate (figs. 487, 500); temples

slightly roundly narrowed posteriad (figs. 321, 490, 499); length of hind femur

7.1—8.1 times its width; claws with a rather wide subapical tooth (fig. 491), or with

a posteriorly sharp ventral lamella (figs. 498, 510); inner hind claw of 9 convex

and setose basally (figs. 491, 502); hind telotarsus of 9 setose near base of inner

hind claw; apices of hind tibial spurs of & sharp and hyaline apically (cf. fig. 317);

1A +2A of fore wing straight (figs. 485, 496); basal third of SR of hind wing more

(fig. 494) or less (fig. 484) sclerotized and curved (fig. 507); r of hind wing absent;

SC+RI distinctly curved (figs. 482, 495, 507); length of Ist tergite 2.6—4.8 times

its apical width; length of ovipositor sheath 0.12—0.25 times fore wing; posterior

part of propodeum not or slightly separated from antero-dorsal part of propodeum

(figs. 481, 492, 503).

Distribution. — The subgenus Phylacter is only known with certainty from the

Palaearctic region, if the Himalayan area is considered to be an extension of the

Palaearctic region. One species has a mainly more northern distribution

(annulicornis), while the other two species are restricted to the South of the

Palaearctic region. I have examined a Z from Indonesia (Idjen) which may belong

to a species close to annulicornis.

Key to the species of the subgenus Phylacter

I. Tarsal claws bifurcate, the subapical tooth large and in Q truncate apically

(figs. 488, 491); length of ovipositor sheath 0.12—0.16 times fore wing (fig. 481);

South East,Palacarctic en. eae bifurcatus spec. nov. (p. 322)

— Tarsal claws with a subapical sharp and tooth-shaped ventral lamella (figs. 498,

510); length of ovipositor sheath 0.17—0.25 times fore wing (figs. 492, 503) 2

2. Vein 2-SC+R of hind wing transverse, longer than wide (fig. 495) or quadrate;

base of hind tarsus more yellowish basally than medially, 2nd—4th segment

whitish, contrasting with hind tibia; mainly North and Middle Palaearc-

ti nl LIO annulicornis (Nees) (p. 324)

— Vein 2-SC+R of hind wing vertical, wider than long (fig. 507); hind tarsus

equally whitish-yellow, only weakly contrasting with hind tibia; South West

Palaearctice oe e PE meridionalis spec. nov. (p. 326)

Homolobus (Phylacter) bifurcatus spec. nov.

(figs. 285, 286, 481—491)

Holotype, 9, length of body 9.7, of fore wing 9.6 mm.

Head. — Antennal segments 41, but apical segments missing, 3rd segment 1.3

VAN ACHTERBERG: Revision Zelinae auct. 323

times 4th segment, length of 3rd and 4th segments 4.1 and 3.2 times their width,

respectively; length of maxillary palp 1.5 times height of head; dorsal length of eye

2.2 times temple; POL : @ ocellus : OOL = 5: 5: 6; frons en vertex almost flat and

smooth; face mainly flat and slightly punctulate; clypeus convex and punctulate

(fig. 487); length of malar space 0.7 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

dorsally smooth, medially and postero-ventrally crenulate-rugose (fig. 481);

epicnemial area weakly rugose; precoxal suture largely crenulate-rugose,

posteriorly almost smooth (fig. 481); rest of mesopleuron smooth; metapleural

flange large, lamelliform, wide and rounded apically (fig. 481); metapleuron

largely smooth, rugose ventrally and crenulate anteriorly; notauli closely crenulate

(fig. 286); mesoscutal lobes punctulate; surface of propodeum rugose posteriorly

and laterally, antero-medially mainly smooth, with a medium-sized carina; this

antero-medial carina is divided posteriorly, enclosing a triangular area, its base

formed by a lamelliform transverse carina apically.

Wings. — Fore wing: r : 3-SR : SRI = 9: 15 : 52; SRI curved (fig. 484); cu-a

almost straight, postfurcal; 1-CU1: 2-CU1 = 3: 23; 2-SR : 3-SR : r-m = 12: 15:9;

2A shortly sclerotized basally (fig. 484); area basally of 2A remotely setose (fig.

485). Hind wing: SR rather shortly sclerotized basally (fig. 482); 2-SC+R

transverse (fig. 484).

Legs. — Hind coxa punctulate; tarsal claws with a large and truncate subapical

tooth (figs. 488, 491), outer claws yellowish pectinate basally, inner claws only

setose basally; length of femur, tibia, and basitarsus of hind leg 7.1, 10.7 and 10.4

times their width, respectively; length of spurs of hind tibia 0.6 and 0.4 times

basitarsus.

Metasoma. — Length of Ist tergite 4.8 times its apical width, its surface mainly

smooth, laterally somewhat microsculptured (fig. 489); dorsal carinae of Ist tergite

present in front of spiracles; length of ovipositor sheath 0.14 times fore wing.

Colour. — Brownish-yellow; stemmaticum and its surroundings, and areola,

dark brown; hind tarsus evenly yellowish-white, contrasting with hind tibia;

pterostigma light brown.

Holotype in NR, Stockholm: “‘N.E. Burma, Kambaiti, 2000 m, 21/5.1934,

Malaise”, “Riksmuseum Stockholm”. Paratypes: (4 9 and 4 ¢) from Burma (1 9

ft (RMNH); 1 4, 8.V.1934, 7000 ft (NR)), India (2 &, “United Prov., India, 1949,

F. Bianchi” (TC); 1 9, “India, H.P., Kalalop, 2138 m, 7.VI.1971, Kamilko, DH

79” (DZD)), and Nepal (2 9, “27°58’N, 85°00’E, Nepal, 11100 ft, 7 June 1967,

Can. Nepal Exp.”, “Zele Det. W. R. M. Mason” (CNC)). Variation: Length of fore

wing 9.0—11.5 mm; antennal segments 49—54; length of malar space 0.7—1.0

times basal width of mandible; length of ovipositor sheath 0.12—0.16 times fore

wing; length of Ist tergite 3.7—4:7 times its apical width; frequently middle of

mesoscutal lobes dark brown or blackish; 2-SC+R quadrate or transverse;

324 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Homolobus (Phylacter) annulicornis (Nees) comb. nov.

(figs. 315—318, 492—502)

Nees, 1834, Hym. Ichn. affin. mon. 1: 201 (as Rogas).

Haliday, (1835) 1836, Ent. Mag. 3:141, 142 (as testaceator (nec Curtis, 1832!)).

Wagner, 1928, Verh. Ver. naturw. Unterh. Hamb. 20: 10.

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319, 320, 323, fig. 6.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 226.

Papp, 1970, Israel J. ent. 5: 65 (needs confirmation).

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 230, 231.

Capek, 1972, Ent. Problémy 10: 133, 136.

Papp, 1973, Acta Mus. Mac. Sc. Nat. 14: 9.

Kabasinskaité & Jakimavicius, 1973, Acta ent. Lituanica 2: 80, 86.

Jakimavicius, 1976, Tr. AN Lit. SSR B2 (74): 90, 93.

Gauld & Huddleston, 1976, Entomologist’s Gaz. 27: 43, fig. 20.

Tobias, 1976, Opr. Fauna SSSR 110: 133, fig. 39: 7—10.

Neotype, 9, length of body and of fore wing both 9.7 mm.

Head. — Antennal segments 45, but apical segments missing, 3rd segment 1.1

times 4th segment, length of 3rd and 4th segment 3.5 and 3.2 times their width,

respectively; length of maxillary palp 1.6 times height of head; dorsal length of eye

1.7 times temple; POL : @ ocellus : OOL = 5: 6: 5; frons rather flat, with some

microstriae laterally (fig. 499); vertex rather flat, smooth; face punctulate-

rugulose, rather flat; clypeus rather flat, punctulate (fig. 500); length of malar

space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

crenulate medially and posteriorly, striate ventrally and almost smooth dorsally

(fig. 492); epicnemial area reticulate-rugose; precoxal suture reticulate-rugose, but

almost smooth posteriorly (fig. 492); rest of mesopleuron punctulate; metapleural

flange large, lamelliform, wide en rounded apically (fig. 492); metapleuron

punctulate, with some crenulae ventrally; notauli almost smooth, somewhat

crenulate posteriorly (fig. 316); mesoscutal lobes punctulate; surface of

propodeum finely rugose, almost smooth anteriorly, medial carina absent, except

for a short part anteriorly.

Wings. — Fore wing: r: 3-SR : SRI = 10: 14: 55; SRI weakly curved (fig. 494);

cu-a weakly curved (fig. 494), postfurcal; 1-CUI : 2-CUI = 1:11; 2-SR:3-SR:r-m

= 13: 14: 8; 2A scarcely sclerotized basally (fig. 494); area basally of 2A remotely

setose (fig. 496). Hind wing: 2-SC+R transverse; basal third of SR wholly

sclerotized (figs. 494, 495).

Legs. — Hind coxa punctulate; tarsal claws with a subapical sharp, tooth-

shaped lamella (figs. 498, 502), yellowish pectinate basally, except inner hind claw

(fig. 502); length of femur, tibia, and basitarsus of hind leg 8.0, 11.2, and 10.4 times

its width, respectively; length of outer spur of hind tibia 0.4 times basitarsus (in a Q

specimen from Denmark (Klaekket) both spurs 0.5 and 0.6 times basitarsus (fig.

317)).

Metasoma. — Length of Ist tergite 3.5 times its apical width, its surface mainly

smooth, but somewhat rugulose laterally (fig. 315); dorsal carinae of Ist tergite

absent; length of ovipositor sheath 0.18 times fore wing.

VAN ACHTERBERG: Revision Zelinae auct. 325

Colour. — Brownish-yellow; stemmaticum blackish; hind tarsus (except its base

and apex) contrasting with its tibia; subapical antennal segments indistinctly

infuscated apically; pterostigma yellowish.

Neotype (9!) in KBIN, Brussels: “Coll. Wesmael”, “1874”, “4 Phylax 9

annulicornis N.V. Es., det. C. Wesmael”, “Type”. Collected in Belgium, in

Charleroi or near Brussels, as stated by Wesmael. Because the type of Nees is lost

and Wesmael is the first revisor, I designate here this specimen as the neotype of

Rogas annulicornis Nees, 1834, the type-species of Phylacter Reinhard.

Öland, Gärdby), Denmark (Ordrüp, Lynabg; Thali; Humlebaek; Amager; Hinde;

Klaekket, Haas; Vorso; Moens Fyr, nr. Borre; Moesgaard; Braband; Bukke,

Skov.; Aakiaer; Silkehorn; Bogg; Rus; Satruphlg; Schelde; Sathò; Aarò;

Dyrehavn; Copenhagen; Markskel, 100 m, N. Kohavegärd, Ostjylland, S. Vejle),

West Germany (Steinebach am Wörthsee; Hüll nr. Wolnzack; Gambach, nr.

Würzburg, at light, beech wood; Wiesen, Spessart; Mainz; Kiel; Worms, Rosg.;

Weisskirchen, Mähren; Goslar; Niederadenau, Eifel; Fahnersche Höhe;

Tübingen; Bodensee, Ueberlingen; Eisenberg; Goslar a. H., Grauhöfer Holz;

Oelber a. W.; Barenkopf; Park Allee, Eichst.; Gräfeling, Bayern; Günzburg a. D.;

München; Furstwied; Harz, Eikntal; Tegernsee), East Germany (Thüringen,

Blankenburg; Brünshäupten; Berlin; Ebersdorf), Austria (Leitha Mts.,

Donnerskirchen; Wien; Rainberg; Steinbruch, Salzburg; Styria, Pagaska, Slatina,

228 m), Switzerland (Glion, 800 m), Czechoslovakia (nr. Prague), Poland

(Gdansk), USSR (Moscow; Kaunas; Vilnius, Jaansali; Azerbaijdzhan SSR,

Kalajbugurt, wood), Hungary (Budapest), Ireland (Finglas, Co. Du.; Knather,

Bundoran dt., Co. Ed.; Strangford, Co. Do.; Trawalua, Co. SI.), England (Coomb

Wood), Netherlands (Tegelen, De Holtmühle, Zuid-Limburg; Driebergen;

Venray; Oosterbeek; Neerijnen, Waardenburg; Asperen; Venlo; Otterlo), France

(La Bégude de Mazenc, Drôme), Italy (Garda L., Malcerni, 300 m), Romania

(Transsylv. Alps, Cibins. Mts.), China (Manchuria, Maderschan), and Japan

(Kamikochi; Kyoto, Honshu) (RMNH, ITZ, HC, CNC, TC, UZM, ZSB, ZMB,

IZP, NMV, ZIL, USNM, ZI, WHC, EI, KBIN).

Variation: Length of fore wing 6.3— 10.3 mm; antennal segments 48—52; length

of Ist tergite 2.9—3.5 times its apical width; length of ovipositor sheath 0.17—0.22

times fore wing; metasoma sometimes infuscated apically; vein 2-SC+R of hind

wing transverse or quadrate; cocoon whitish, with a more or less developed white

transverse medial band. Known hosts of examined specimens: Lithophane lamda

(F.), Enargia ypsillon (Denis & Schiff.), Xestia triangulum (Hufn.), and Orthosia

spec., all four belonging to the Noctuidae, Lepidoptera.

Note. This species is often named Zele testaceator Curtis, 1832 (e.g., Nixon,

1938), but examination of the type, the original description, and the figures given

by Curtis revealed its synonymy with Zele albiditarsus Curtis, 1832 (formerly

placed in Zemiotes or Meteorus). That the correct name for this species is

annulicornis was stated in 1918 by Bengtsson. He based his opinion solely on the

original description, which is clear enough to show testaceator of Curtis is the

female sex of albiditarsus, of which Curtis described only the male. In the Curtis

326 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Collection under testaceator, there are 2 9 of Zele albiditarsus Curtis (of which one

is the type of testaceator, see note under albiditarsus) and 3 3. One male without a

label and two males collected after the publication of testaceator; these males

belong to annulicornis, and cannot be types of testaceator.

Homolobus (Phylacter) meridionalis spec. nov.

(figs. 321—323, 503—512)

Holotype, 9, length of body 9.9, of fore wing 10.3 mm.

Head. — Antennal segments 41, but apical segments missing, 3rd segment 1.3

times 4th segment, length of 3rd and 4th segments 4.1 and 3.1 times their width,

respectively; POL : & ocellus : OOL = 4: 6: 5; length of maxillary palp 1.5 times

height of head; dorsal length of eye 2.0 times temple; frons rather flat, largely

smooth, with some microsculpture (fig. 321); vertex rather flat, punctulate; face

punctulate, somewhat rugulose near antennal sockets, rather flat (fig. 509);

clypeus convex, punctulate; length of malar space 0.5 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

smooth dorsally, rugulose-crenulate medially and striate ventrally (fig. 503);

epicnemial area reticulate-punctate; precoxal suture largely reticulate-punctate,

posteriorly almost smooth (fig. 503); rest of mesopleuron punctulate; metapleural

flange large, rather thick, and sharp apically; metapleuron mainly smooth,

ventrally rugose; notauli narrowly and indistinctly crenulate (fig. 323); mesoscutal

lobes punctulate; anteriorly surface of propodeum mainly smooth, posteriorly

coarsely reticulate (fig. 511), without medial carina.

Wings. — Fore wing: r: 3-SR : SRI = 15: 30: 118; SRI curved (fig. 506); cu-a

inclivous, slightly curved posteriorly, postfurcal; 1-CUI : 2-CUI = 2: 25; 2-SR : 3-

SR: r-m = 25 : 30: 17; 2A distinctly sclerotized basally (fig. 506); area basally of

2A sparsely setose (fig. 504). Hind wing: 2-SC+R vertical (fig. 507); basal third

wholly sclerotized (fig. 506).

Legs. — Hind coxa punctulate; tarsal claws with an apically sharp, tooth-shaped

lamella (figs. 510, 512), indistinctly yellowish pectinate basally, except inner hind

claw; length of femur, tibia, and basitarsus of hind leg 8.1, 12.6, and 10.0 times

their width, respectively; length of spurs of hind tibia 0.6 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 3.0 times its apical width, its surface largely

smooth, laterally with some carinae and striae (fig. 511); dorsal carinae of Ist

tergite absent, only behind the spiracles with a dorso-lateral carina; length of

ovipositor sheath 0.20 times fore wing.

Colour. — Brownish-yellow; stemmaticum black; hind tarsus, base of hind tibia,

fore and middle legs mainly, more or less whitish-yellow; hind tarsus only weakly

contrasting with hind tibia.

Holotype in CNC, Ottawa: “15.V.1960, Oued Tisquite, 1650 m, 2 km NO Itrano,

“Spain, Granada, Sierra Nevada, Alb. Universitaria, 6 May 1966, 2600 m, Lyveb.-

Matin-Langem.”), Cyprus (1 & (CNC), “24.4.53, Cypern, Pera Pcd.,

VAN ACHTERBERG: Revision Zelinae auct. 327

Lempke & K. Straatman”; collected between 11-19.X.1971). Variation: length of

fore wing 7.8—10.0 mm; antennal segments 46—50; length of ovipositor sheath

0.20—0.22 times fore wing; length of Ist tergite 3.0—3.1 times its apical width;

propodeum sometimes with an irregular medial carina.

Note. This new species seems to have escaped attention because it occurs in

spring (April, May), early summer (beginning of June), late autumn (October), or

winter (December), while annulicornis is most frequently captured in July and

August, and less frequently in May, June, and September, although I saw some

exceptional captures of annulicornis from April, October and November.

H. meridionalis seems to replace the closely related annulicornis in the

Mediterranean Region.

Subgenus Oulophus nov.

Etymology: From “où” (Greek for “‘not’’) and “.opos” (Greek for “ridge”), because the antennal rid-

ge of the 9 is not distinctly developed. Gender: masculine.

Type-species: Homolobus armatus spec. nov.

Diagnosis. — Length of body 5.8—9.9, of fore wing 5.4—11.5 mm; antennal

segments 40—54, its 3rd—6th segments of 9 usually without a longitudinal ridge,

if exceptionally present (fig. 720), then rather weakly developed; length of 4th

segment of labial palp 2.2—7.0 times 3rd segment; length of maxillary palp

1.2—1.9 times height of head; apical margin of clypeus rather convex (fig. 525) or

almost straight (fig. 549) medially, not (fig. 525) or distinctly (fig. 550)

differentiated from clypeus; length of malar space 0.3—1.3 times basal width of

mandible; eyes weakly emarginate (figs. 525, 572, 645, 675); temples directly (fig.

677) or weakly roundly narrowed (fig. 722) posteriad; length of hind femur

5.8—7.6 times its width; claws with a large (fig. 524) or small (fig. 534) subapical

tooth, or with a narrow, apically sharp, tooth-shaped lamella ventrally (figs. 522,

570); inner hind claw of 9 convex or straight and setose basally (figs. 524, 534,

560), exceptionally bare and slightly concave (fig. 571); hind telotarsus of 9 setose

near base of inner hind claw; apices of hind tibial spurs of G sharp and hyaline

apically; 1A +2A of fore wing straight (figs. 515, 591); basal third of SR of hind

wing only pigmented, not sclerotized (fig. 516) or sclerotized (figs. 618, 635),

straight (fig. 631) or weakly curved (fig. 641); r of hind wing present (figs. 590, 603)

or absent (figs. 516, 635); SC+R1 more or less curved (fig. 683 versus figs. 539,

555) or almost straight (figs. 631, 725); length of Ist tergite 1.8—3.9 times its apical

width; length of ovipositor sheath 0.07—0.77 times fore wing; posterior part of

propodeum distinctly (figs. 513, 527) or not (figs. 541, 564) separated from antero-

dorsal part of propodeum.

Distribution. — This largest and rather diverse subgenus of Homolobus is

widespread, but is unknown from the Afrotropical and Australian regions. In total

ten out of 16 species occur in the New World, including four species restricted to

328 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

the South West Nearctic (Sonoran) area, where they seem to have evolved. In the

Palaearctic region (including the Himalayan area) six species occur, one of which

is Holarctic in its distribution. In the Oriental region (excluding the Himalayan

area) only one species is Known.

Key to the species of the subgenus Oulophus

1. Malar space comparatively long (figs. 525, 532), its length 1.0—1.3 times basal

width of mandible; hind coxae extensively and more or less coarsely rugose

(figs. 523, 527); both penultimate segments of antenna of 9 stout (figs. 518,

531), their length 1.2—1.6 times their width; Ist tergite somewhat narrowed

apicad (figs. 526, 537); antero-dorsal part of propodeum distinctly separated

from its posterior pant; Palaearctic: E 2

— Malar space comparatively short (figs. 549, 554, 595), its length 0.3—0.8 times

basal width of mandible; hind coxae smooth or punctulate, at most somewhat

rugose dorsally (figs. 553, 596); both penultimate segments of antenna of 9

more slender, their length 1.7—2.9 times their width; antero-dorsal part of

propodeum variable, but in most species not distinctly separated from its

posteriorpart (Mes 553. 311) TERRE ee N 3

2. Subapical tooth of tarsal claws comparatively stout, rather blunt and subequal

to the apical tooth, resulting in sub-bifurcate claws (figs. 522, 524); body

mainly black; medially propodeum mainly coarsely reticulate; vein r of hind

wing absent (fig. 516) or only present as a short remnant (fig. 519); pterostigma

dark brown; hind basitarsus basally black or brownish, and at least apical half

of basitarsus white; vertex smooth or punctulate (fig. 520) .............

EELDE GUE, SAL LÄNGE HERE da ea RE carbonator (Shestakov ) (p. 330).

— Subapical tooth of tarsal claws slender, sharp, much shorter than the apical

tooth (figs. 531, 536); body yellowish and/or brownish; propodeum, except for

the carinae, only indistinctly sculptured (fig. 537); vein r of hind wing present,

but anterior half usually indistinctly developed (fig. 539), exceptionally mainly

absent; whole hind basitarsus and pterostigma yellowish; vertex coriaceous

(fio: 338), SER patie, PUS REVERE bohemani (Bengtsson) (p. 332)

3. Vein r of hind wing present (figs. 543, 555), at least posteriorly present as a

brownish pigamented'striper. 2410) OEE ee EE rae eee 4

— ;Veinr completely absenti(figs) 618,635), set o 9

4. Ovipositor sheath short (figs. 541, 553), 0.08—0.16 times fore wing;

mesonotum more orless/)brownish-yellow vatten TEAR ENE 5

— Ovipositor sheath comparatively long (figs. 577, 588, 602), 0.25—0.52 times

fore wing; mesonotumimainly black nnie eN 7

5. Pterostigma and parastigma of 9 unicolorous, yellowish; hind tarsus whitish-

yellow; vein r of hind wing comparatively long and strongly reclivous (figs.

525, 569); vertex smooth (fig, 568) Re OO ee 6

— Pterostigma and parastigma of 9 bicolorous, yellowish and dark brown; hind

tarsus brownish-yellow; vein r of hind wing short and comparatively straight

(fig. 540); vertex coriaceous (fig. 550); South Nearctic ua rn. SE N ee

VAN ACHTERBERG: Revision Zelinae auct. 329

ana ia arts lem INNERE bicolor spec. nov. (p. 333)

Precoxal suture with some rugae antero-dorsally (fig. 553); subapical tooth of

tarsal claws comparatively slender, claws weakly concave medio-ventrally (fig.

558, 560); vein cu-a of fore wing parallel to 3-CUI (fig. 555); palpi, fore and

middle legs more or less whitish-yellow; costulae of propodeum at least partly

developed: (fie..553); Holarctiex 0.0 case ies flagitator (Curtis) (p. 334)

Precoxal suture smooth (fig. 564); subapical tooth of claws lamelliform, rather

wide (figs. 570, 571); claws straight medio-ventrally (fig. 571); vein cu-a of fore

wing more inclivous than 3-CUI (fig. 567); palpi, fore and middle legs,

brownish-yellow; costulae of propodeum absent (figs. 564, 661); South

Neareticand Neotropical: 5» sent ir 0m acares spec. nov. (p. 336)

Apical two-thirds of hind tibia dark brown or blackish (fig. 596); vertex

smooth (fig. 594); propodeum more or less rugose medially (fig. 588);

pterostigma unicolorous, dark brown; surroundings of the veins 1-M and 1-

SU ottiore wingshyaline: Palaearctic» 419544. Hate mue eee. 8

Hind tibia completely brownish-yellow; vertex coriaceous (fig. 578); propo-

deum smooth, except fore some indistinctly developed sculpture postero-

medially (fig. 577); pterostigma bicolorous, basally yellowish and medially

dark brown; surroundings of the veins 1-M and 1-CUI infuscated (fig. 580);

Neetropicalubyitsiag tenga nie Oa occidentalis spec. nov. (p. 337)

Ovipositor sheath subequal to length of metasoma (fig. 588), its length

0.51—0.52 times fore wing; length of vein 3-SR of fore wing 1.7—2.0 times

vein r of fore wing; face blackish; length of malar space 0.6—0.7 times basal

width of mandible; East Palaearctic ...... nipponensis spec. nov. (p. 338)

Ovipositor sheath much shorter than metasoma (fig. 602), its length 0.25—0.26

times fore wing; length of vein 3-SR of fore wing 1.0—1.3 times vein r of fore

wing; face reddish-brown; length of malar space 0.4—0.5 times basal width of

mandibles South Palaearctici.. enr toe nepalensis spec. nov. (p. 340)

Basal quarter of vein SR of hind wing equally sclerotized as vein 1-M (figs.

618, 635); area basally of vein 2A of fore wing mainly bare (figs. 619, 636);

precoxal suture widely sculptured (figs. 616, 633); Oriental and South

PA RESOURCE D ET LE 10

Basal quarter of vein SR of hind wing only pigmented, not sclerotized as vein

1-M (figs. 649, 692); area basally of 2A variable, but if mainly bare, then

precoxal suture mainly smooth (figs. 714, 734); New World ......... 11

. Vein 2-SC+R of hind wing long, transverse (figs. 618, 631); base of vein SR of

hind wing straight; tarsal claws with a ventral lamella (fig. 629); antenna, hind

tibia and body yellowish; Oriental ........ crenulatus spec. nov. (p. 341)

Vein 2-SC+R of hind wing short, vertical (figs. 635, 641); base of vein SR of

hind wing weakly curved; tarsal claws with a subapical tooth (fig. 638);

antenna mainly dark brown, but with a medial white or yellowish ring; body

and apical 0.7 of hind tibia mainly brownish-black; South Palaearctic .....

SEES an at Ar eI A bce IENA CONAI or psn te annulatus spec. nov. (p. 342)

. Vein SC+RI of hind wing curved (figs. 655, 669, 683); area basally of vein 2A

of fore wing sparsely setose (figs. 648, 671); vein cu-a of fore wing antefurcal or

330

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

interstitial with vein 1-M, exceptionally postfurcal (figs. 649, 670); precoxal

suture extensively sculptured, at least dorsally (figs. 647, 680) ........ 12

Vein SC+RI of hind wing straight (figs. 716, 739); area basally of vein 2A of

fore wing bare (figs. 719, 738); vein cu-a of fore wing distinctly postfurcal (fig.

716); precoxal suture usually mainly smooth (figs. 714, 734) ........ TS

. Length of ovipositor sheath 0.08—0.36 times fore wing, comparatively short

(figs. 667, 680) or medium-sized (fig. 694); subapical tooth of tarsal claws

medium-sized (fig. 678) or rather large (fig. 699); hind tarsus more whitish

yellow, more strongly contrasting with the brownish tibia .......... 13

Length of ovipositor sheath 0.68—0.79 times fore wing, long (figs. 646, 649);

subapical tarsal tooth comparatively large (fig. 656); hind tarsus and its tibia

almost equally coloured, not or only weakly contrasting; South

Nearctich Hu RER O E a armatus spec. nov. (p. 343)

. Head and antenna brownish-yellow; base of vein SR of hind wing weakly

curved (fig. 682); vein cu-a of fore wing antefurcal (fig. 698) or subinterstitial;

propodeum rather coarsely and rather remotely rugose or almost smooth (figs.

680,694): South Nearcticr tr De. ASI RE NT SR: 14

Head and basal half of antenna (except both basal segments partly), dark

brown; base of vein SR of hind wing almost straight (fig. 669); vein cu-a of fore

wing interstitial with 1-M (fig. 670) or postfurcal; propodeum partly finely and

densely reticulate-rugose (fig. 667); Neotropical obscurus spec. nov. (p. 344)

. Length of ovipositor sheath 0.12—0.15 times fore wing, comparatively short

(fig. 680); length of vein 3-SR of fore wing less than twice vein r of fore wing

(fii GSD) soa ea RT se eee antefurcalis spec. nov. (p. 345)

Length of ovipositor sheath 0.29—0.36 times fore wing, medium-sized (fig.

694); length of vein 3-SR of fore wing more than twice vein r of fore wing (fig.

698): EN Coad mr ORGE AREE AEN mesoxiphius spec. nov. (p. 346)

. Length of fore wing of 9 ca. 1.3 times length of body (compare fig. 716 with

fig. 714); inner aspect of 3rd—8th antennal segments with a rather weakly

developed ridge (fig. 720); subapical tooth of claws comparatively stout (figs.

723, 734); length of ovipositor sheath ca. 0.07 times fore wing ...........

POOR A MENE macropterus spec. nov. (p. 347)

Length of fore wing of 9 1.0—1.1 times fore wing (compare fig. 737 with fig.

734); antenna of 9 without ridge; subapical tooth of claws comparatively

slender (fig. 740); length of ovipositor sheath 0.13—0.14 times fore

WINE. NME REI E LENA eee rectinervis spec. nov. (p. 348)

Homolobus (Oulophus) carbonator (Shestakov) comb. nov.

(figs. 513—526)

Shestakov, 1940, Ark. Zool. 32A: 17 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 222.

Holotype, 9, length of body 7.5, of fore wing 6.1 mm.

Head. — Antennal segments 46, 3rd segment 1.4 times 4th segment, without

antennal ridge, length of 3rd and 4th segments 2.9 and 2.1 times their width,

VAN ACHTERBERG: Revision Zelinae auct. 331

respectively, both penultimate segments 1.4 times their width (fig. 518); length of

4th segment of labial palp 4.0 times 3rd segment; length of maxillary palp 1.5 times

height of head; dorsal length of eye 2.3 times temple; temple directly narrowed

posteriad (fig. 520); POL : @ ocellus : OOL = 4: 5: 6; frons mainly flat, with some

striae near antennal sockets (fig. 520); vertex almost flat and smooth; face weakly

convex, punctulate medially, punctate laterally and punctate-striate dorsally;

clypeus rather flat, remotely punctulate; apical margin of clypeus weakly convex

ventrally, not differentiated (fig. 525); length of malar space 1.3 times basal width

of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

smooth dorsally, coarsely crenulate medially and posteriorly, superficially rugose

ventrally (fig. 513); epicnemial area punctate-rugose; precoxal suture rugose

dorsally, reticulate-rugose medially and punctate ventrally; rest of mesopleuron

punctulate; metapleural flange large, wide, thick, without carina, rounded apically

(fig. 513); metapleuron largely reticulate-rugose, more punctate medially and

almost smooth dorsally; notauli well impressed, wholly crenulate, anteriorly rather

narrowly; mesoscutal lobes punctulate; surface of propodeum largely coarsely

rugose-reticulate, anteriorly almost smooth, medial carina present in anterior

third, its posterior part well separated from somewhat shorter antero-dorsal part,

and with weak tubercles laterally (fig. 513); spiracle of propodeum rather large

(fig. 513).

Wings. — Fore wing: r : 3-SR : SR1=6 : 6: 40; SRI straight; cu-a straight,

postfurcal; 1-CU1=1 : 14; 2-SR : 3-SR : r-m=32 : 24 : 23; 2A shortly sclerotized

basally (fig. 516); area basally of 2A sparsely setose (fig. 515). Hind wing: r absent,

but remnant present in right wing (fig. 519); 2-SC+R transverse; SC +R1 weakly

curved (fig. 519); basal third of SR weakly curved and unsclerotized (fig. 516).

Legs. — Hind coxa coarsely rugose-reticulate (fig. 513); tarsal claws with a

rather blunt, large subapical tooth (fig. 522, 524), yellowish and inconspicuously

pectinate; length of femur, tibia and basitarsus of hind leg 6.3, 10.3, and 7.6 times

their width, respectively; length of spurs of hind tibia 0.7 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 3.1 times its apical width, its surface

narrowed posteriad, reticulate-rugose (fig. 526); dorsal carinae of Ist tergite

absent; length of ovipositor sheath 0.15 times fore wing.

Colour. — Black; mandibles, palpi, antenna, antennal sockets, legs (except

coxae, hind femur and tibia (except their bases), and hind tarsus), metasoma

ventrally, 2nd tergite laterally (fig. 526), wing venation, and tegulae, brownish;

hind tarsus (except the black basal half of basitarsus and brownish telotarsus) -

white; pterostigma dark brown; wing membrane hyaline.

Holotype in NR, Stockholm: “Vladivostok, Sedanka, Malaise/ 18/7, 30”, “Zele

carbonator sp. n. typ., det. Shestakov”, “401, 77”, “Riksmuseum Stockholm”.

“N.E. Burma, Kambaiti, 1800 m, 17/6 & 16/6, 1934, Malaise, Riksmuseum

Stockholm”. Variation: wing membrane somewhat brownish, base of hind tarsus

only narrowly brownish; subapical tooth of tarsal claws smaller than in holotype,

332 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

still larger than in bohemani; vertex punctulate; vein r of hind wing completely

absent; length of malar space 1.2 times basal width of mandible.

Homolobus (Oulophus) bohemani (Bengtsson) comb. nov.

(figs. 354—357, 527— 539, 707)

Bengtsson, 1918, Acta Univ. lund. (2)14(32) : 39, 44, 45 (as Phylacter).

Watanabe, 1969, Proc. ent. Soc. Wash. 71: 319, 323 (as geminator).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

Holotype, 9, length of body 7.1, of fore wing 7.5 mm.

Head. — Remaining antennal segments 18, apical segments absent, 3rd segment

subequal to 4th segment, without ridge, length of 3rd and 4th segments 3.0 and 2.9

times their width, respectively; length of 4th segment of labial palp 7 times 3rd

segment; length of maxillary palp 1.4 times height of head; dorsal length of eye 1.8

times temple; temple directly narrowed posteriad, coriaceous (fig. 538); POL : &

ocellus : OOL= 9: 9: 14; frons flat, striate (fig. 538); vertex rather flat, coriaceous;

face weakly convex, transversely and finely striate, especially laterally coriaceous

(fig. 532); clypeus convex, punctulate; apical margin of clypeus weakly convex

ventrally, thin, well differentiated; length of malar space 1.1 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely densely reticulate-rugose, dorsally smooth and crenulate medio-anteriorly

(fig. 527); epicnemial area reticulate-rugose; precoxal suture coarsely reticulate-

rugose; rest of mesopleuron largely reticulate-rugose, posteriorly almost smooth

(fig. 527); metapleural flange large, rather thick lamelliform apically (fig. 527);

metapleuron largely rugose-reticulate, antero-dorsally smooth; notauli finely and

narrowly crenulate (fig. 535); mesoscutal lobes almost smooth; surface of

propodeum coarsely areolated, the area between the carinae irregularly and rather

weakly rugose, medial carina anteriorly shortly present; posterior part of

propodeum well separated (figs. 527, 537).

Wings. — Fore wing: r : 3-SR : SR= 5: 11 : 39; SR straight (fig. 529); cu-a

mainly straight, apically curved basad, postfurcal (fig. 529); 1-CU1 : 2-CUl=1:9;

2-SR : 3-SR : r-m=13: 11: 7; 2A absent, except for a faintly pigmented trace; area

basally of 2A medially bare, posteriorly somewhat setose (fig. 533). Hind wing:

posterior half of r present (fig. 529); 2-SC +R transverse; SC+R1 weakly curved

(fig. 539); basal third of SR straight basally and unsclerotized.

Legs. — Hind coxa densely and rather finely reticulate-rugose (fig. 527); tarsal

claws with small subapical tooth, slender, sharp, and much shorter than apical

tooth (figs. 534, 536), setose basally; length of femur, tibia, and basitarsus of hind

leg 6.0, 11.3, and 5.7 times their width, respectively; length of spurs of hind tibia

0.5 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 3.2 times its apical width, its surface coarsely

reticulate-rugose, scarcely narrowed apicad (fig. 537); dorsal carinae of Ist tergite

present in basal third; basal half of 2nd tergite somewhat pimply; length of

ovipositor sheath 0.10 times fore wing.

VAN ACHTERBERG: Revision Zelinae auct. 333

Colour. — Brownish-yellow; palpi, tegulae, pterostigma, metasoma baso-

ventrally, and hind tarsus, light yellowish.

Holotype in NR, Stockholm: “Sm. [=Smaland]”, “Bhn [=Boheman]”,

“Phylacter Bohemani Bgtn” (handwritten in pencil), “407.77”, “Riksmuseum,

Stockholm”. Additional specimens examined: (119 and 14 g) from Finland

(Perikkala; Länsi-Teisko; Lemland, Flaka; id., Apelholm), Sweden (Ljungby),

West Germany (Drensfeld; Niederaudorf, Obbay., 1000 m), Kurile Islands

(Uruppu), Nepal (27°58’N, 85°00’E, 11100 ft), and India (Kumaon Hills, Dhakuri,

2612 m; H.P. Dhenkund (or Dainkund), 2743 m; Ahla, H.P., 2286 m; H.P.,

Kalatop, 2438 m) (CNC, WHC, ZMH, EI, HC, TC, DZD, RMNH).

Variation: length of fore wing 6.3—7.8 mm; antennal segments 43—44; length of

both penultimate segments 1.2—2.3 times their width, but in 9 more stout (fig.

531), 1.2—1.6 times their width; length of malar space 1.0—1.3 times basal width

of mandible; length of Ist tergite 3.0—3.9 times its apical width; length of

ovipositor sheath 0.10—0.11 times fore wing: vein r of hind wing sometimes

complete or only posteriorly weakly developed (fig. 357); sometimes only anterior

half of precoxal suture and mesopleuron distinctly rugose; antenna frequently

dark brown, except for both basal segments; sometimes hind coxa, Ist tergite, hind

femur, and mesosoma mainly, rather dark brown.

Homolobus (Oulophus) bicolor spec. nov.

(figs. 540— 552, 706)

Holotype, 9, length of body 6.0, of fore wing 7.0 mm.

Head. — Antennal segments 40, 3rd segment 1.3 times 4th segment, without

ridge, length of 3rd and 4th segments 4.4 and 3.4 times their width, respectively,

length of both penultimate segments 1.8 and 2.1 times their width; length of 4th

segment of labial palp 3.3 times 3rd segment; length of maxillary palp 1.4 times

height of head; dorsal length of eye 2.4 times temple; temple directly narrowed

posteriad (fig. 550); POL : @ ocellus : OOL= 7: 11 : 8; frons slightly concave,

mainly smooth; vertex rather flat, coriaceous; face punctulate and indistinctly

aciculate, rather flat; clypeus convex, punctulate; apical margin of clypeus thin,

straight medially, differentiated (fig. 549); length of malar space 0.6 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

smooth, but medio-anteriorly crenulate, posteriorly weakly rugose, and ventrally

somewhat punctulate (fig. 541); epicnemial area smooth; precoxal suture, as rest

of mesopleuron, smooth, except for some punctulation; metapleural flange rather

large, wide, rounded apically (fig. 541); metapleuron smooth, except for some

short carinae ventrally; notauli anteriorly mainly smooth, only apical third

crenulate (fig. 548); mesoscutal lobes punctulate; surface of propodeum smooth,

except for some rugosity laterally, medial carina and areola absent; posterior part

of propodeum not separated from antero-dorsal part (fig. 541).

Wings. — Fore wing: r : 3-SR : SR1=8 : 21: 88; SR! straight; cu-a inclivous,

postfurcal; 1-CU1 : 2-CUl=1 : 7; 2-SR : 3-SR : r-m=23 : 21 : 12; 2A well

334 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

developed and sclerotized (fig. 543); area basally of 2A bare (fig. 542). Hind wing: r

present, comparatively short and rather straight (fig. 540); 2-SC+R transverse;

SC+RI rather curved (fig. 543); basal third of SR weakly curved and

unsclerotized (fig. 540).

Legs. — Hind coxa punctulate; tarsal claws with a rather small subapical tooth,

setose basally (figs. 545, 552); length of femur, tibia and basitarsus of hind leg 5.8,

10.7, and 7.8 times their width, respectively; length of spurs of hind tibia 0.6 and

0.5 times basitarsus.

Metasoma. — Length of Ist tergite 1.8 times its apical width, its surface basally

smooth, posterior half rugulose (fig. 551); dorsal carinae of Ist tergite absent;

length of ovipositor sheath 0.13 times fore wing.

Colour. — Brownish-yellow; palpi, fore and middle legs (except tarsi),

ovipositor sheath, tegulae, dorso-posterior corner of pronotum, whitish-yellow;

stemmaticum, and veins, dark brown; parastigma and pterostigma bicolorous:

base and posterior margin of parastigma and apical 0.6 of pterostigma dark brown,

rest of para- and pterostigma yellowish; wing membrane slightly brownish,

especially near veins.

Holotype in CNC, Ottawa: “Mex., Chis., 7200 ft, S.Crist. las Casas, 17 June

1969, Malaise trap”. Paratypes: (9 9), topotypic (CNC, RMNH). Variation: length

of fore wing 6.3—7.5 mm; antennal segments 39—43; length of ovipositor sheath

0.14—0.16 times fore wing.

Homolobus (Oulophus) flagitator (Curtis) comb. nov.

(figs. 553—563)

Haliday, 1835 (1836), Ent. Mag. 3: 142 (as chlorophthalmus (nec Spinola, 1808, and Nees, 1834)).

Curtis, 1837, Guide Br. Insects: 119 (Zele flagitator nom. nov. for chlorophthalmus Haliday).

Lyle, 1914, Entomologist 47: 289, 290 (Zele geminator nom. nov. for chlorophthalmus Haliday).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 224.

Redescribed after a 9 from Ireland, length of body 7.3, of fore wing 8.3 mm.

Head. — Antennal segments 45, 3rd segment 1.2 times 4th segment, without

ridge, length of 3rd and 4th segments 3.8 and 3.2 times their width, respectively,

length of both penultimate segments 1.8 and 2.0 times their width; length of 4th

segment of labial palp 4.0 times 3rd segment; length of maxillary palp 1.7 times

height of head; dorsal length of eye 1.9 times temple; temple directly narrowed

posteriad (fig. 556); POL : @ ocellus : OOL= 3: 6: 5; frons almost flat and smooth;

vertex rather flat, almost smooth (fig. 556); face rather flat, densely and finely

coriaceous, with a medial tubercle (fig. 554), shiny; clypeus strongly convex,

remotely punctate; apical margin of clypeus weakly convex, differentiated (fig.

554); length of malar space 0.7 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, except for some crenulae medially and some micro-striation posteriorly

(fig. 553); epicnemial area almost smooth; precoxal suture smooth, except for

some rugae anteriorly (fig. 553); rest of mesopleuron smooth; metapleural flange

large, lamelliform, wide and rounded apically; metapleuron smooth, but ventrally

VAN ACHTERBERG: Revision Zelinae auct. 335

rugose; notauli rather finely crenulate (fig. 562); mesoscutal lobes smooth; surface

of propodeum smooth anteriorly, medially and posteriorly superficially rugose,

with a rather long medial carina anteriorly, areola weakly developed and costulae

present (fig. 553); posterior part of propodeum not separated from antero-dorsal

part.

Wings. — Fore wing: r : 3-SR : SRI=7 : 12 : 55; SRI straight; cu-a strongly

inclivous, postfurcal, parallel to 3-CUI (fig. 555); 1-CUI : 2-CUl=1: 10; 2-SR : 3-

SR; r-m= 14 : 12: 7; 2A absent except for a minute basal remnant (fig. 555); area

basally of 2A sparsely setose. Hind wing: r present, comparatively long and

reclivous (fig. 555); 2-SC+R transverse; SC + RI evenly curved; basal third of SR

slightly curved and unsclerotized (fig. 555), scarcely posteriorly curved distad of r.

Legs. — Hind coxa weakly punctate-rugose dorsally; tarsal claws with a

comparatively slender subapical tooth, medio-ventrally weakly concave (figs. 558,

560), setose basally; length of femur, tibia and basitarsus of hind leg 7.4, 12.8, and

10.0 times their width, respectively; length of spurs of hind tibia 0.5 and 0.4 times

basitarsus.

Metasoma. — Length of Ist tergite 2.8 times its apical width, its surface

shallowly rugulose submedially (fig. 563); dorsal carinae present in front of

spiracles; length of ovipositor sheath 0.09 times fore wing.

Colour. — Brownish-yellow; stemmaticum and apical half of antenna, dark

brown; palpi, fore and middle legs, hind tarsus slightly more whitish-yellow.

Holotype probably lost, not present in the Haliday Collection, Dublin, nor in

the Royal Scottish Museum, Edinburgh, nor in the National Museum of Victoria,

Melbourne. Redescribed after 9 from Stelfox Collection, USNM: “45”,

“Amongst logs!, Drinahilly, Co. Do., A.W.S., 2.11.65”, “geminator 9 A.W.S.”,

“A.W. Stelfox Collection 1966”. Because the description of Haliday is sufficient

and confusion with another Palaearctic species is unlikely, the designation of a

neotype is not necessary. Additional specimens examined: 74 9 and 51 ¢. From

the Palaearctic region (including the Himalayan area): Sweden (Vmld, Ekshäred),

England (Aviemore), Ireland (Hallyfort, Co. Wx.; Alerlow, Co. St.; Drinnahilly,

Co. Do.; Tollymore Park, Co. Do.), West Germany (Ober-Harz, Torfhaus, ca. 800

m; Ober-Bayern, Ellmau, ca. 1050 m; Reither Alm, 850 m; Schladwohg, 1250 m),

Nepal (Pulchauki, Ktmd, 8000 ft; 28°00/N, 85°00’E, 10500 ft; 27°58’N, 85°00’E,

11100 ft; 27°56’N, 85°00’E, 9900 ft), and India (H.P., Narkanda, 2700 m; Kumaon

Hills, Phurkia, 3504 m; Simla Hill, H.P., Narkanda) (ZSB, USNM, HC, CNC, AC,

DZD).

From the Nearctic region: Alaska (Spenard; Tsaina R.; Casio Core, Attu,

Aleut.), British Columbia (Hixon; Miskatla Inlet; Falkland; Port Renfreu;

Galiano Isl.; Coleman Cr.; Klemtu; Cowichan L.; Lagoon Rd.; Sooka; Beechy

Head; Cumshavalnld; Alliford Bay; Terrace, airport area; Wellington; Tofino; S.

Pender Isl.; Johnson L.; Mt. Arrowsmith; Ootsa; Ocean Falls; Port San Juan;

Green Inlet; Metchosin; Gualicum; Murtle R.; S. Gote Y.N.P.; Zeballos R.),

Alberta (Smith; Pocohontas; Brule; Hinton), Ontario (Pass Lake; Beardmore;

Nakina), Quebec (Sac à l’Ours; Ct. Pope; St. Vianney; Indian House L.; Sapin),

New Foundland (South Branch), Colorado (Phantom Vy., RMNP, 9400 ft), New

336 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Hampshire (Pinkham Notch), North Carolina (Devil’s Court House, Blue Ridge

Parkway; Clingmans Dome, 6600 ft; Mt. Pisgah, 5000—5749 ft; Pisgah Nat.

Forest, Haywood Co., Chestnut Bald, 5900 ft; Highlands, Whiteside Mt., 4900 ft,

and California (Marin Co., Mill Valley; Arcata, Humboldt Co., black light trap;

Lily Pond Alpine Lk., Marin Co., 1500 ft) (CAS, UCA, BM, CNC, TC, RMNH).

Variation: length of fore wing 5.4—9.0 mm; antennal segments 45—49; length of

4th segment of labial palp 4.0—4.5 times 3rd segment; length of malar space

0.7—0.8 times basal width of mandible; length of Ist tergite 2.5—2.9 times its

apical width; length of ovipositor sheath 0.08—0.10 times fore wing; 3rd segment

of antenna of 9 sometimes with a weakly developed ridge at the inner side;

sometimes middle of mesoscutal lobes infuscated. Cocoon rather thin and whitish. |

Known hosts of examined specimens all belong to the Geometridae: Eupithecia

longipalpata Packard, E. placidata Packard, E. unicolor Hulst, E. annulata Hulst, E.

olivaceae Taylor, E. harrisonata MacK., Nyctobia limitata (Walker), N.

nigroangulata Strecker, Oporinia pulchraria (Minot), Melanolophia spec., Campaea

perlata (Guenée) on Betula papyrifera, id. on Salix sp., Caripeta divisata Walker,

Entephria caesiata Lang, and Alcis repandata (L.) on Vaccinium myrtillis L.

Homolobus (Oulophus) acares spec. nov.

(figs. 564— 574, 661—663)

Holotype, 9, length of body 9.2, of fore wing 10.2 mm.

Head. — Antennal segments 45, 3rd segment 1.3 times 4th segment, without

ridge, length of 3rd and 4th segments 4.3 and 3.4 times their width, respectively,

length of both penultimate segments 2.3 and 2.4 times their width; length of 4th

segment of labial palp 4.0 times 3rd segment; length of maxillary palp 1.7 times

height of head; dorsal length of eye 2.6 times temple; rather directly narrowed

posteriad (fig. 568); POL : & ocellus : OOL= 5: 12 : 9; frons smooth, slightly

concave medially; vertex flat, smooth; face rather flat, smooth laterally,

indistinctly rugulose-punctate medially (fig. 572); clypeus strongly convex,

punctulate; apical margin of clypeus straight medially, thin, differentiated; length

of malar space 0.6 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

crenulate medio-anteriorly, rugose posteriorly, ventrally somewhat punctulate,

and dorsally smooth (fig. 564); epicnemial area smooth; precoxal suture smooth,

rest of mesopleuron punctulate; metapleural flange large, rather thick, wide and

rounded apically; metapleuron smooth, except for some ventral carinae; notauli

largely smooth, posteriorly somewhat crenulate (fig. 663); mesoscutal lobes

indistinctly punctulate; surface of propodeum mainly smooth, except for some

rugae (fig. 661), with a short medial carina anteriorly, without costulae and areola;

posterior part of propodeum not separated from antero-dorsal part (fig. 564).

Wings. — Fore wing: r: 3-SR : SR1=17: 29: 110; SRI weakly curved (fig. 567);

cu-a more inclivous than 3-CUI (fig. 567), somewhat curved basad apically,

postfurcal; 1-CU1 : 2-CU1=1 : 9; 2-SR : 3-SR : r-m=28 : 29 : 14; 2A shortly

sclerotized basally (fig. 567); area basally of 2A mainly bare (fig. 566). Hind wing: r

VAN ACHTERBERG: Revision Zelinae auct. 337

present, comparatively long and strongly reclivous (fig. 569); 2-SC + R transverse;

SC+R1 rather curved; basal third of SR rather straight, distad of r rather abruptly

curved posteriad (fig. 567).

Legs. — Hind coxa punctulate, dorso-apically with some weak striae (fig. 662);

tarsal claws with apically sharp, tooth-shaped lamella (fig. 570), somewhat

yellowish pectinate basally, but inner hind claw bare and slightly concave basally

(fig. 571); length of femur, tibia and basitarsus of hind leg 6.0, 9.6, and 7.8 times

their width, respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.7 times its apical width, its surface smooth

anteriorly, posterior half rugulose (fig. 661); dorsal carinae of Ist tergite absent;

length of ovipositor sheath 0.10 times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; hind tarsus whitish-yellow;

wing membrane somewhat infuscated; wing veins partly dark brown.

Holotype in RMNH, Leiden: “Museum Leiden, N. Panama, Boquete, Alto

Lino, 1300 m, 8°48’N-82°26’W, 21.1.1977, H. Wolda, at light”. Paratypes: (20 9

and 10 ¢), 2 & (CNC, RMNH), “Mex., Dgo., 9000’ (ft), El Salto, 10 mi. W., 2—6

(RMNH); 1 9 (CNC), “Mex., Dgo., 24 mi. W. La Ciudad, 7000’ (ft), 25 July 1964,

W. R. M. Mason”; 1 & (CNC), id., 8 Aug. 1964, W. R. M. Mason; 1 9 (RMNH),

id., 12 Aug. 1964; 19 id., 2 July 1964 (CNC); 1 9, “Mex., Dgo., 30 mi. W. La

Cuidad, 6500’ (ft), 25 July 1964, W. R. M. Mason” (CNC); 1 9, “Mex., Dgo., 3 mi.

E. El Salto, 8500’ (ft), 10 July 1964, W. R. M. Mason” (RMNH); 1 9, id., 4 July

1964 (CNC); 1 9, “Ramsey Cyn., 6000’ (ft), 15 mi. S. Sierra Vista, Huachuca Mts.,

Ariz., Sternitzky, 29.IX.67° (CNC); 3 &, id., 29.X.1967 (CNC); 3 9, id., 29.X.1967

(CNC); 1 9, id., 28.XI.1967 (CNC); 1 &, id., 9.IX.1967 (RMNH); 1 9, id.

30.IX.1967 (CNC); 1 9, id., VII.1968 (CNC); 1 9, “Omilteme, Guerrero, 8000 ft,

“Rustler Park, 8500 (ft), Chiricahua Mts., Ariz., VII-20-72, at light, J. Powell”

(UCA); 1 9, id., VIII-20-72 (UCA).

Variation: length of fore wing 7.5—10.2 mm; antennal segments 45—48; length

of ovipositor sheath 0.08—0.12 times fore wing; inner side of 3rd segment of

antenna sometimes with weakly developed ridge; vein SR of hind wing sometimes

as in flagitator, scarcely curved distad of r. I exclude from the type-series | ¢ from

Bolivia (Coroico, 1800 m (CNC)), with white hind tarsus and the tarsal lamella less

developed than in holotype.

Homolobus (Oulophus) occidentalis spec. nov.

(figs. 575— 586)

Holotype, 9, length of body 5.8, of fore wing 7.1 mm.

Head. — Antennal segments 20, but apical segments missing, 3rd segment 1.3

times 4th segment, without ridge, length of 3rd and 4th antennal segments 4.4 and

3.4 times their width, respectively; length of 4th segment of labial palp 3.8 times

338 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

3rd segment; length of maxillary palp 1.4 times height of head; dorsal length of eye

2.0 times temple; directly narrowed posteriad (fig. 578); POL : @ ocellus : OOL= 6

: 9: 12; frons almost flat and smooth, but laterally somewhat rugose; vertex rather

flat, coriaceous; face rather flat, coriaceous, but medio-ventral triangle smooth

(fig. 585); clypeus convex, punctulate; apical margin of clypeus almost straight

medially, thin, differentiated; length of malar space 0.8 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, but medio-anteriorly shortly crenulate and posteriorly rugose (fig. 577);

epicnemial area smooth; precoxal suture absent; mesopleuron slightly punctulate;

metapleural flange rather small and rounded apically (fig. 577); metapleuron

slightly punctulate; notauli narrowly and indistinctly crenulate (fig. 584);

mesoscutal lobes weakly punctulate; surface of propodeum smooth, except for

some short rugae medio-posteriorly, without medial carina and areola, and its

posterior part not separated from antero-dorsal part.

Wings. — Fore wing: r: 3-SR : SR1= 10: 14: 80; SRI straight; cu-a postfurcal,

inclivous; 1-CUI : 2-CU1= 3: 17; 2-SR : 3-SR : r-m= 19: 14: 10; 2A sclerotized

basally (fig. 580); area basally of 2A bare (fig. 576). Hind wing: r present (fig. 580);

2-SC +R transverse; SC +RI curved and rather short (fig. 575); basal third of SR

weakly curved, unsclerotized (fig. 580).

Legs. — Hind coxa indistinctly punctulate; tarsal claws with a small subapical

tooth (figs. 579, 582), at most somewhat yellowish pectinate basally; length of

femur, tibia and basitarsus of hind leg 5.8, 11.4, and 9.4 times their width,

respectively; length of spurs of hind tibia 0.5 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 1.8 times its apical width, its surface smooth,

but laterally behind spiracles and posteriorly somewhat rugulose (fig. 586); dorsal

carinae of Ist tergite shortly developed in front of spiracles; length of ovipositor

sheath 0.43 times fore wing.

Colour. — Brownish-yellow; stemmaticum, vertex and frons medially,

C+SC+R (except basally), antenna (as far as present, and without annellus),

pronotum anteriorly, mesonotum and ovipositor sheath, more or less blackish;

face and most wing veins, infuscated; pterostigma medially dark brown; base and

apex of pterostigma yellowish; wing membrane hyaline, but near the veins 1-M

and 1-CUI infuscate (fig. 580).

Holotype in TC, Ann Arbor: ‘“Unduavi/Corioco, Yungas La Paz, Bol., 1.2.76,

3000 m, Luis Pefia”.

Homolobus (Oulophus) nipponensis spec. nov.

(figs. 587—600)

Watanabe, 1969, Proc. ent. Soc. Wash. 71(3): 319, 320, figs. 1, 2 (Zele daurica p.p.).

Holotype, 9, length of body 7.1, of fore wing 7.5 mm.

Head. — Antennal segments 42, 3rd segment 1.2 times 4th segment, without

antennal ridge, length of 3rd and 4th segments 3.8 and 3.3 times their width,

respectively, length of both penultimate segments 1.7 and 2.1 times their width;

VAN ACHTERBERG: Revision Zelinae auct. 339

length of 4th segment of labial palp ca. 5 times 3rd segment; maxillary palp

incomplete; dorsal length of eye 1.9 times temple; temple directly narrowed

posteriad (fig. 594); POL : @ ocellus : OOL= 7: 9: 12; frons flat, almost smooth;

vertex almost flat and smooth; face rather flat, smooth medio-ventrally, dorsally

rugose-punctate and coriaceous laterally (fig. 595); clypeus weakly convex,

shallowly punctate; apical margin of clypeus weakly convex, rather thick, and not

differentiated (fig. 595); length of malar space 0.7 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

aciculate ventrally, crenulate medially and posteriorly, dorsally and submedially

smooth (fig. 588); epicnemial area more or less reticulate-rugose; precoxal suture

mainly absent, only anteriorly somewhat depressed and with some rugae; rest of

mesopleuron remotely punctulate; metapleural flange large, wide and rounded

apically (fig. 588); metapleuron reticulate-rugose ventrally, almost smooth

dorsally; notauli narrowly crenulate (fig. 599); mesoscutal lobes densely

punctulate; surface of propodeum with areola and costulae, its surroundings

almost smooth, somewhat rugulose, medial carina absent, except for a short part

anteriorly; posterior part of propodeum not separated from antero-dorsal part (fig.

588).

Wings. — Fore wing: r: 3-SR : SR1= 6: 12: 42; SRI slightly curved (fig. 590);

cu-a somewhat inclivous, postfurcal; 1-CU1 : 2-CU1= 1: 15; 2-SR : 3-SR : r-m=

11 : 12 : 6; 2A shortly sclerotized basally (fig. 590); area basally of 2A laterally

sparsely setose, medially bare (fig. 591). Hind wing: r present, rather long (fig.

590); 2-SC +R transverse; SC + RI curved, rather short (fig. 592); basal third of SR

slightly curved and unsclerotized.

Legs. — Hind coxa remotely punctulate; tarsal claws with a small subapical

tooth (figs. 597, 598), setose basally; length of femur, tibia and basitarsus of hind

leg 6.6, 10.6, and 8.4 times their width, respectively; length of spurs of hind tibia

0.6 and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 2.1 times its apical width, its surface rather

superficially rugulose (fig. 600); dorsal carinae of Ist tergite present in basal

quarter of tergite; length of ovipositor sheath 0.51 times fore wing, somewhat

longer than metasoma (fig. 588).

Colour. — Blackish-brown; patch at vertex near eyes, apico-lateral corner of

mesoscutum, and hind leg largely, reddish-brown; apical 0.7 of hind tibia dark

brown (fig. 596); palpi, fore and middle legs, tegulae, dorso-apical corner of

pronotum, margin of hypopygium, yellowish; pterostigma dark brown; apex of

antenna brownish; wing membrane hyaline.

Holotype in EI, Sapporo: “Tsu Mie, Honshu, 9.XII.1962, M. Matsuura”, “Zele

daurica (Shestakov) 9, Det. C. Watanabe, 1969”. Paratypes: (1 9 and 2g), 1g

(EI, allotype), ‘“Hirakura, Mie Honshu, 18.X1.1963, M. Matsuura”, “Zele daurica

(Shestakov) &, Det. C. Watanabe, 1969”, antennal segments 41, length of fore

wing 7.1 mm, length of malar space 0.6 times basal width of mandible, and length

27.VIII.1965, M. Miyaz”, “Zele daurica (Shestakov) 9, Det. C. Watanabe, 1969”,

length of fore wing 7.8 mm, length of ovipositor sheath 0.52 times fore wing, length

340 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

of maxillary palp 1.5 times height of head, length of malar space 0.6 times basal

width of mandible, and length of 2-SR 1.7 times r of fore wing; 1 ¢ (EI), topotypic

with allotype, 11.XI.1963. The males of nipponensis may be easily confused with

males of dauricus, but are recognizable by the rugose epicnemial area uu more

strongly developed propodeal carina of nipponensis.

Homolobus (Oulophus) nepalensis spec. nov.

(figs. 601—615)

Holotype, 9, length of body 6.4, of fore wing 7.0mm.

Head. — Antennal segments 42, 3rd segment 1.2 times 4th segment, without

antennal ridge, length of 3rd and 4th segments 4.0 and 3.4 times their width,

respectively, length of both penultimate segments 1.8 and 2.2 times their width;

length of 4th segment of labial palp 1.2 times 3rd segment; length of maxillary palp

1.2 times height of head; dorsal length of eye 2.1 times temple; temple directly

narrowed posteriad (fig. 608); POL : & ocellus : POL= 3: 5: 5; frons almost

smooth and shallowly concave; vertex rather flat, smooth; face flat, punctulate,

but near antennal sockets punctate-rugose; clypeus convex, punctulate; apical

margin of clypeus rather thin, not differentiated, and weakly convex medially (fig.

601); length of malar space 0.4 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

medially and posteriorly crenulate, rest mainly punctulate (fig. 602); epicnemial

area punctulate; precoxal suture almost absent, punctulate as rest of mesopleuron;

metapleural flange rather large, lamelliform, and rounded apically (fig. 602);

metapleuron shallowly punctate, and with some carinae ventrally; notauli

narrowly crenulate, but anteriorly almost smooth (fig. 612); mesoscutal lobes

almost smooth, somewhat punctulate; surface of propodeum smooth, except for

the costulae and a rather irregular medial carina, posteriorly with an irregularly

defined, elliptical areola, and posterior part of propodeum not separated from

antero-dorsal part (fig. 601).

Wings. — Fore wing: r : 3-SR : SR1= 13: 13: 88; SRI almost straight (fig. 603);

cu-a almost straight, narrowly postfurcal; 1-CUI : 2-CU1= 1: 15; 2-SR: 3-SR : r-

m= 18: 13: 8; 2A shortly sclerotized basally (fig. 603); area basally of 2A mainly

bare (fig. 606). Hind wing: r present; 2-SC +R transverse; SC+ RI rather curved

(fig. 607); basal third of SR almost straight, unsclerotized (fig. 603).

Legs. — Hind coxa punctulate, with some striae dorso-apically; tarsal claws

with a medium-sized lamelliform subapical tooth (figs. 613, 614), setose basally;

length of femur, tibia and basitarsus of hind leg 6.6, 10.3, and 8.0 times their width,

respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 1.8 times its apical width, its surface

indistinctly and remotely rugulose (fig. 615); dorsal carinae of Ist tergite absent,

except for a pair of short stubs basally; length of ovipositor sheath 0.26 times fore

wing, much shorter than metasoma (fig. 602).

Colour. — Blackish-brown; head, surroundings of eye, temple, antenna (but

basal half infuscated), hind trochanters, hind femur, basal third of hind tibia, its

VAN ACHTERBERG: Revision Zelinae auct. 341

spurs, and hind tarsus, more or less reddish-brown; fore and middle tibiae and

tarsi, Ist and 2nd epipleura, brownish-yellow; palpi, fore and middle coxa and

femora, dorso-posterior corner of pronotum, tegulae, and base of hind wing,

yellowish-white; pterostigma and wing veins, dark brown; hind coxa dark reddish-

brown; wing membrane hyaline.

Holotype in CNC, Ottawa: “27°58’N, 85°00’E, Nepal, 11100 ft, 31 May 1967,

Can. Nepal Exped.”, “Homolobus, Det. W.R.M. Mason”. Paratype: | 9,

topotypic, 7 June 1967 (RMNH), length of fore wing 7.5 mm, length of ovipositor

sheath 0.25 times fore wing; length of 3-SR 1.3 times r of fore wing; antennal

segments 41, and length of malar space 0.5 times basal width of mandible.

Homolobus (Oulophus) crenulatus spec. nov.

(figs. 616—632)

Holotype, 9, length of body 9.9, of fore wing 9.6 mm.

Head. — Antennal segments 47, 3rd segment 1.1 times 4th segment, with a

rather distinctly developed ridge at the inner side (fig. 628); length of 3rd and 4th

segments 4.2 and 3.8 times their width, respectively, length of both penultimate

segments 2.3 and 2.6 times their width; length of 4th segment of labial palp 4.6

times 3rd segment; length of maxillary palp 1.5 times height of head; dorsal length

of eye 2.6 times temple; temple directl6 narrowed posteriad (fig. 625); POL : 5

ocellus : OOL= 8 : 11 : 8; frons almost flat and smooth; vertex rather flat,

coriaceous, dull; face rather flat, punctulate-coriaceous, rather dull (fig. 620);

clypeus rather convex, punctulate; apical margin of clypeus thin, straight medially

and not separated (fig. 620); length of malar space 0.5 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

smooth, but medially and posteriorly remotely crenulate (fig. 606); epicnemial

area crenulate anteriorly, rugose posteriorly; precoxal suture deeply impressed,

anteriorly widely and coarsely crenulate, medially coarsely crenulate-rugose, and

posterior third mainly smooth (fig. 606); metapleural flange large, rather slender,

lamelliform, narrowly rounded apically (fig. 606); metapleuron punctulate,

ventrally with some coarse carinae; notauli anteriorly narrowly and posteriorly

widely crenulate (fig. 632); mesoscutal lobes punctulate; surface of propodeum

coarsely areolate (fig. 626), enclosed areas mainly smooth, with a medial carina

anteriorly and well-developed costulae submedially; posterior part of propodeum

not separated from antero-dorsal part (fig. 616).

Wings. — Fore wing: r: 3-SR: SR1= 22: 27: 115; SRI almost straight (fig. 618);

cu-a almost straight, interstitial; 2-SR : 3-SR : r-m = 29: 27: 16; 2A shortly

sclerotized basally (fig. 618); area basally of 2A bare (fig. 619). Hind wing: r absent;

2-SC +R transverse; SC +R1 rather straight and long (fig. 631); basal third of SR

straight and sclerotized (fig. 618).

Legs. — Hind coxa punctulate, but postero-dorsally punctate; tarsal claws with

a ventral lamella (figs. 624, 629), setose basally; length of femur, tibia, and

basitarsus of hind leg 7.0, 9.9, and 8.1 times their width, respectively; length of

342 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

spurs of hind tibia 0.8 and 0.6 times basitarsus.

Metasoma. — Length of Ist tergite 3.1 times its apical width, its surface

indistinctly rugulose (fig. 625); dorsal carinae of Ist tergite absent; length of

ovipositor sheath 0.09 times fore wing.

Colour. — Brownish-yellow; stemmaticum black; hind tarsus somewhat

whitish-yellow; wing membrane hyaline.

Holotype in BM, London: “B.N. Borneo, Mt. Kinabalu, Lumu Lumu, 5500 ft,

8:4:1929/ H.M. Pendlebury, coll. F.M.S. Museums”, “Ex F.M.S. Museum, B.M.

1955-354”.

Homolobus (Oulophus) annulatus spec. nov.

(figs. 633—645, 708)

Holotype, 9, length of body 8.2, of fore wing 7.7 mm.

Head. — Antennal segments 44, 3rd segment 1.1 times 4th segment, without

ridge, length of 3rd and 4th segments 3.7 and 3.3 times their width, respectively,

length of both penultimate segments 2.3 and 2.9 times their width; length of 4th

segment of labial palp 4.0 times 3rd segment; length of maxillary palp 1.4 times

height of head; dorsal length of eye 2.5 times temple; temple directly narrowed

posteriad (fig. 639); POL : & ocellus : OOL= 9: 10: 10; frons almost smooth,

except for some rugae (fig. 639), flat; vertex rather flat and coriaceous; face rather

flat, punctate, coriaceous laterally, rugulose medio-dorsally and latero-ventrally

(fig. 645); clypeus weakly convex, remotely punctate; apical margin of clypeus thin

and straight medially, not differentiated (fig. 645); length of malar space 0.3 times

basal width of mandible.

Mesosoma. — Length of mesosoma 1.2 times its height; side of pronotum

crenulate medially, rugose ventrally, posteriorly and dorsally mainly smooth (fig.

633); epicnemial area reticulate-rugose; precoxal suture widely reticulate-rugose,

except posteriorly (fig. 633); rest of mesopleuron punctulate; metapleural flange

lamelliform, large, wide and rounded apically; metapleuron smooth, but

posteriorly reticulate-rugose; notauli coarsely crenulate (fig. 642); mesoscutal

lobes punctulate; dorsal surface of propodeum mainly smooth laterally, medially

and posteriorly with strongly developed carinae, medial carina absent and costulae

lamelliform; posterior part of propodeum with an areola, well separated from

antero-dorsal part (fig. 633).

Wings. — Forewing: r: 3-SR : SR1= 8: 11 : 46; SRI almost straight (fig. 635);

cu-a weakly inclivous, far antefurcal; 2-M +CUI : 1+2CU1= 3: 37; 2-SR : 3-SR:

r-m= 10: 11:5; 2A shortly sclerotized basally (fig. 635); area basally of 2A bare

(fig. 636). Hind wing: r absent; 2-SC+R short, vertical (fig. 635); SC +R1 rather

straight (fig. 641); basal third of SR weakly curved and main part sclerotized (fig.

635, 641).

Legs. — Hind coxa punctulate, dorso-anteriorly punctate, and dorso-posteriorly

rugose (fig. 633); tarsal claws with a medium-sized, sharp subapical tooth (figs. 638,

643); length of femur, tibia and basitarsus of hind leg 6.1, 9.7 and 7.8 times their

width, respectively; length of spurs of hind tibia 0.7 and 0.5 times basitarsus.

VAN ACHTERBERG: Revision Zelinae auct. 343

Metasoma. — Length of Ist tergite 2.2 times its apical width, its surface smooth

anteriorly except for some rugae, and posterior half rugose (fig. 644); dorsal

carinae of Ist tergite present in basal quarter of tergite; length of ovipositor sheath

0.09 times fore wing.

Colour. — Brownish-black; 13th—19th segments of antenna, base of C+SC+R

and hind tarsus (only telotarsus yellowish), white; scapus, pedicellus, and apex of

antenna, fore and middle legs, Ist and 2nd tergites, hind coxa, hind femur,

ovipositor sheath, tegulae, vertex laterally, palpi, and all trochanters, more or less

brownish-yellow; bases of all tibiae and basal half of metasoma, yellowish-white;

pterostigma and wing veins, mainly dark brown.

Holotype in DZD, Delhi: “India: H.P., Ahla, 2286 m, 4.viii-16.1x.1971, M.trap,

No. tr. I”. Paratype: 1 &, allotype (DZD), “India, Simla Hill, Sangla, 2743 m,

16.vi.1972, Girish, No. G14”. Length of fore wing 7.0 mm, antennal ring yellowish,

basal 0.7 of Ist tergite blackish brown, length of body 7.3 mm, antennal segments

45, length of malar space 0.3 times basal width of mandible, length of Ist tergite 1.9

times its apical width, and sculpture and shape of claws as in holotype.

Homolobus (Oulophus) armatus spec. nov.

(figs. 646—660)

Holotype, 9, length of body 8.9, of fore wing 10.3 mm.

Head. — Antennal segments 48, 3rd segment 1.3 times 4th segment, without

ridge, length of 3rd and 4th segments 4.0 and 3.0 times their width, respectively,

length of both penultimate segments 2.0 and 1.7 times their width; length of 4th

segment of labial palp 2.8 times 3rd segment; length of maxillary palp 1.6 times

height of head; dorsal length of eye 1.8 times temple; temple rather directly

narrowed posteriad (fig. 653); POL : @ ocellus : OOL= 5: 6: 5; frons smooth and

rather flat; vertex rather flat, almost smooth; face mainly flat, medially weakly

convex, indistinctly punctulate, near antennal sockets weakly rugulose (fig. 652);

clypeus convex, punctulate; apical margin of clypeus thin, almost straight

medially, not differentiated; length of malar space 0.7 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

largely coarsely rugose-striate, dorsally narrowly smooth (fig. 647); epicnemial

area anteriorly almost smooth, posteriorly rugose; precoxal suture widely rugose-

reticulate; rest of mesopleuron punctulate; metapleural flange large, rounded and

rather slender apically (fig. 647); metapleuron punctulate, with some carinae

ventrally; notauli narrowly crenulate anteriorly, more widely so posteriorly (fig.

660); mesoscutal lobes indistinctly punctulate; surface of propodeum smooth,

except for some rugae and an irregular transverse carina, with a short medial

carina, areola absent; posterior part of propodeum somewhat separated from

antero-dorsal part (fig. 647).

Wings. — Fore wing: r : 3-SR : SR1= 8: 16: 57; SRI weakly curved (fig. 649);

cu-a slightly inclivous, antefurcal; 2-M + CUI : 1+2-CUl= 1: 16; 2-SR : 3-SR : r-

m= 11: 16 : 7; 2A distinctly developed and sclerotized (fig. 649); area basally of

344 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

2A sparsely setose (fig. 648). Hind wing: r absent; 2-SC+R transverse; SC+RI

weakly curved (fig. 655); basal third of SR weakly pigmented, not sclerotized (fig.

649).

Legs. — Hind coxa punctulate, except for some striae dorso-apically; tarsal

claws with a comparatively large subapical tooth (figs. 656, 657); length of femur,

tibia and basitarsus of hind leg 6.4, 12.4, and 10.2 times their width, respectively;

length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface smooth,

but posterior half rugulose (fig. 658); dorsal carinae of Ist tergite absent; length of

ovipositor sheath 0.77 times fore wing.

Colour. — Brownish-yellow; stemmaticum yellowish; hind tarsus not or weakly ‘

contrasting with its tibia.

Holotype in CNC, Ottawa: “Capulin Nat. Mon., 6 mi. SW. Folsom, N. Mex.,

56—6000 (ft), 15 mi. S. Sierra Vista, Huachuca Mts., Ariz., Sternitzky, 23.X.67”

(CNC); 12 9, topotypic with holotype (CNC, RMNH); 5 9, “Ute Park, N. Mex., 3

mi. SW., 7300’ (ft), 14.IX.1968, D. F. Hardwick” (CNC, RMNH); 1 9, “Ramsey

Cyn., 6000’ (ft), 15 mi. S. Sierra Vista, Huachuca Mts., Ariz., Sternitzky, 19.X.67”

(CNC); 2 9, “Portal, Ariz., 5 mi. SW., 5400’ (ft), 3.X.1969, D. F. Hardwick” (CNC,

RMNH); 1 9, “Mex., Chis., 9600 ft, Zontehuitz, nr. S. Crist., 25 June 1969, W. R.

4.X.1969, D. F. Hardwick” (CNC); 1 9, “Ozumbilla, Hidalgo, Mex., 10-30-57, R.

& K. Dreisbach” (MSU); 1 9, “Arizona: Cochise Co., Southwestern Res. Sta., 5

mi. W. Portal, 30.IX.1966, 5400 ft, P. H. Arnaud, Jr.” (CAS). Furthermore I

examined 3 ¢ from Ute Park, which probably belong to armatus, but are excluded

from the type-series because they are not distinguishable with certainty from

antefurcalis and mesoxiphius.

Variation: length of fore wing 8.2—10.3 mm; length of body 7.6—8.9 mm;

antennal segments 49— 50; length of 4th segment of labial palp 2.2—2.8 times 3rd

segment; length of ovipositor sheath 0.68—0.79 times fore wing; length of Ist

tergite 2.6—2.7 times its apical width; vein cu-a of fore wing antefurcal,

exceptionally interstitial.

Homolobus (Oulophus) obscurus spec. nov.

(figs. 667—679, 703)

Holotype, 9, length of body 7.4, of fore wing 8.4 mm.

Head. — Antennal segments 53, 3rd segment 1.1 times 4th segment, without

ridge, length of 3rd and 4th segments 3.5 and 3.1 times their width, respectively,

length of both penultimate segments 2.8 and 3.2 times their width; length of 4th

segment of labial palp 3.9 times 3rd segment; length of maxillary palp 1.9 times

height of head; dorsal length of eye 4.2 times temple; temple narrowed into a line

posteriad (fig. 677); POL : @ ocellus : OOL= 7: 11 : 10; frons medially smooth,

laterally rugulose, rather flat; vertex flat, coriaceous; face weakly convex,

punctate, weakly transversely rugose dorsally (fig. 675); clypeus weakly convex,

VAN ACHTERBERG: Revision Zelinae auct. 345

punctate and somewhat coriaceous; apical margin of clypeus thin, almost straight

medially, and not differentiated (fig. 675); length of malar space 0.7 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.2 times its height; side of pronotum

dorsally punctulate, medially and posteriorly crenulate and ventrally mainly finely

rugose (fig. 667); epicnemial more or less rugose; precoxal suture dorsally

crenulate, and its anterior half widely rugose (fig. 667); rest of mesopleuron finely

punctate; metapleural flange large, lamelliform, wide and rounded apically (fig.

667); metapleuron punctulate, ventrally rugose-reticulate; notauli rather coarsely

crenulate (fig. 674); mesoscutal lobes remotely punctulate; surface of propodeum

mainly finely and densely reticulate-rugose, only anteriorly and posteriorly partly

smooth (fig. 667), with a rather long medial carina anteriorly, and areola absent,

posterior part of propodeum not separated from antero-dorsal part (fig. 667).

Wings. — Fore wing: r : 3-SR : SR1= 8: 13 : 44; SRI curved (fig. 670); cu-a

inclivous, interstitial; 2-SR : 3-SR : r-m= 11: 13: 8; 2A basally rather long and

slender, sclerotized (fig. 671); area basally of 2A evenly setose. Hind wing: r

absent; 2-SC + R transverse; SC + RI evenly curved (fig. 669); basal third of SR

almost straight, unsclerotized (fig. 670).

Legs. — Hind coxa remotely and finely punctate, with some striae dorso-

apically (fig. 703); tarsal claws with a well-developed, sharp subapical tooth (figs.

678, 679), setose; length of femur, tibia and basitarsus of hind leg 6.8, 10.5, and 8.8

times their width, respectively; length of spurs of hind tibia 0.8 and 0.6 times

basitarsus.

Metasoma. — Length of Ist tergite 3.4 times its apical width, its surface behind

the spiracles mainly rugose (fig. 676); dorsal carinae of Ist tergite absent; length of

ovipositor sheath 0.08 times fore wing.

Colour. — Brownish-yellow; head (except for major part of mandibles), basal

half of antenna (except annellus and apex of scapus) and most wing veins, dark

brown; hind tarsus (but telotarsus yellowish) and ovipositor sheath, yellowish-

white; palpi rather light yellowish; wing membrane slightly yellowish.

Holotype in TC, Ann Arbor: “Nova Teutonia, Santa Catarina, June '53, Braz.,

IV.30.1948; 1 9 (RMNH), topotypic, June 1953. Variation: Length of fore wing

7.4—8.0 mm; antennal segments 53—54; length of Ist tergite 3.4—3.5 times its

apical width; vein cu-a of fore wing interstitial or postfurcal, 1-CUI : 2-CUl= 5:

53; length of malar space 0.7 times basal width of mandible; metasoma somewhat

infuscated in allotype.

Homolobus (Oulophus) antefurcalis spec. nov.

(figs. 664— 666, 680—690)

Holotype, 9, length of body 9.6, of fore wing 10.3 mm.

Head. — Antennal segments 54, 3rd segment 1.2 times 4th segment, without

ridge, length of 3rd and 4th segments 3.9 and 3.2 times their width, respectively,

length of both penultimate segments 2.0 and 2.3 times their width; length of 4th

segment of labial palp 2.3 times 3rd segment; length of maxillary palp 1.6 times

346 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

height of head; dorsal length of eye 1.7 times temple; temple roundly narrowed

posteriad (fig. 686); POL : g ocellus : OOL= 12: 12: 11; frons flat, smooth; vertex

mainly flat, smooth; face rather flat, indistinctly rugulose-punctulate; clypeus

convex, punctulate; apical margin of clypeus thin, almost straight medially, not

differentiated (fig. 687); length of malar space 0.6 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

crenulate medio-anteriorly, ventrally crenulae connected with long rugae,

posteriorly rugose, and dorsally smooth (fig. 680); epicnemial area mainly smooth,

posteriorly indistinctly rugose; precoxal suture largely reticulate-rugose,

posteriorly mainly smooth (fig. 680); metapleural flange lamelliform, large,

rounded apically; metapleuron mainly smooth, ventrally with some carinae;

notauli narrowly and indistinctly crenulate (fig. 666); mesoscutal lobes indistinctly

punctulate; surface of propodeum largely smooth, with several vermiform,

irregular and remote carinae, with a long irregular medial carina, without well-

developed costulae and areola; posterior part of propodeum not separated from

antero-dorsal part (fig. 680).

Wings. — Fore wing: r : 3-SR : SR1= 17: 29: 110; SRI weakly curved; cu-a

inclivous, antefurcal; 2-M +CUI : 1+2-CU1= 1: 17; 2-SR : 3-SR : r-m= 25: 29:

15; 2A well developed, rather long and sclerotized basally (fig. 682); area basally of

2A sparsely setose (fig. 681). Hind wing: r absent; 2-SC+R transverse; SC+RI

curved (fig. 683); basal third of SR weakly curved and unsclerotized (fig. 682).

Legs. — Hind coxa punctulate, with some weak striae dorso-apically (fig. 664);

tarsal claws with a well-developed, subapical, lamelliform tooth (figs. 685, 690),

setose; length of femur, tibia and basitarsus of hind leg 7.4, 10.9, and 10.4 times

their width, respectively; length of spurs of hind tibia 0.6 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 3.7 times its apical width, its surface smooth,

except for some rugulosity laterally (fig. 665); dorsal carinae of Ist tergite absent,

except for a short basal remnant; length of ovipositor sheath 0.15 times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; most wing veins

infuscated; hind tarsus whitish-yellow.

Holotype in CNC, Ottawa: “Cimarron Canyon, 7900 ft, Sangre de Cristo Mts.,

9, “Ute park, N. Mex., 3 mi. SW. 7300’ (ft), 14.1X.1968, D. F. Hardwick”

Mason” (CNC); 1 9, id., 18 July 1964 (CNC). Variation: Length of fore wing

9.9—10.8 mm; antennal segments 54—56; length of vein 3-SR of fore wing

1.6—1.8 times vein r; length of Ist tergite 3.5—3.7 times its apical width; length of

ovipositor sheath 0.12—0.15 times fore wing; sometimes surroundings of

stemmaticum infuscated.

Homolobus (Oulophus) mesoxiphius spec. nov.

(figs. 691—702, 704, 705)

Holotype, 9, length of body 8.5, of fore wing 8.9 mm.

Head. — Antennal segments 50, 3rd segment 1.2 times 4th segment, without

ridge, length of 3rd and 4th segments 3.6 and 3.1 times their width, respectively,

VAN ACHTERBERG: Revision Zelinae auct. 347

length of both penultimate segments 1.9 times their width; length of 4th segment of

labial palp 2.6 times 3rd segment; length of maxillary palp 1.5 times height of head;

dorsal length of eye 2.4 times temple; temple rather roundly narrowed posteriad

(fig. 697); POL : @ ocellus : OOL= 11: 11 : 11; frons smooth, slightly concave;

vertex flat, smooth; face flat, punctulate, medially and dorsally weakly aciculate

(fig. 695); clypeus convex, punctulate; apical margin of clypeus straight medially

and well differentiated from clypeus (fig. 695); length of malar space 0.5 times

basal width of mandible.

Mesosoma. — Length of mesosoma 1.2 times its height; side of pronotum

dorsally smooth; medially coarsely crenulate, ventrally striate-rugose, and

posteriorly rugose (fig. 694); epicnemial area almost smooth, somewhat rugose

(fig. 694); precoxal suture reticulate-rugose, only posteriorly mainly smooth; rest

of mesopleuron punctulate; metapleural flange large, sublamelliform, rather thick,

rounded apically; metapleuron smooth, except for ventral and anterior carinae;

notauli anteriorly mainly smooth, posterior third crenulate (fig. 704); mesoscutal

lobes weakly punctulate; surface of propodeum rather coarsely and remotely

reticulate-rugose medially and laterally, in between mainly smooth, medial carina

and areola absent; posterior part of propodeum weakly separated from antero-

dorsal part of propodeum (fig. 694).

Wings. — Fore wing: r: 3-SR : SR1= 11: 32: 92; SRI almost straight (fig. 698);

cu-a inclivous, antefurcal; 2-M +CUI : 1+2-CUl= 1: 14; 2-SR : 3-SR : r-m= 26:

32 : 14; 2A shortly sclerotized basally (fig. 698); area basally of 2A remotely setose

(fig. 693). Hind wing: r absent; 2-SC+R transverse; SC+RI curved (fig. 692);

basal third of SR weakly curved, unsclerotized (fig. 698).

Legs. — Hind coxa punctulate, dorso-anteriorly weakly coriaceous and dorso-

posteriorly with some short striae (fig. 705); tarsal claws with a well-developed,

lamelliform subapical tooth (figs. 696, 699), indistinctly yellowish pectinate

basally, except inner hind claw; length of femur, tibia and basitarsus of hind leg

6.7, 11.8 and 9.0 times their width, respectively; length of spurs of hind tibia 0.6

and 0.4 times basitarsus.

Metasoma. — Length of Ist tergite 2.8 times its apical width, its surface mainly

smooth, posterior third rugulose (fig. 702); dorsal carinae of Ist tergite absent;

length of ovipositor sheath 0.34 times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; hind tarsus (except

telotarsus) and ovipositor sheath, whitish-yellow; wing membrane hyaline.

Holotype in CNC, Ottawa: “Ramsey Cyn., 5000’ (ft), 15 mi. S. Sierra Vista,

(6), viii.1968 (1), 18.ix.1967 (1), and 29.x.1967 (1) (CNC, RMNH). Variation:

Length of fore wing 9.0— 10.0 mm; antennal segments 49—S2; length of ovipositor

sheath 0.29—0.36 times fore wing; length of vein 3-SR of fore wing 2.2—3.7 times

vein r; vein cu-a of fore wing antefurcal, but sometimes not distinctly so.

Homolobus (Oulophus) macropterus spec. nov.

(figs. 714—728)

Holotype, 9, length of body 8.9, of fore wing 11.5 mm.

348 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Head. — Antennal segments 43, 3rd segment 1.1 times 4th segment, with a

rather weakly-developed ridge (fig. 720), length of 3rd and 4th segments 4.2 and

4.0 times their width, respectively, length of both penultimate segments 2.6 and 2.7

times their width; length of 4th segment of labial palp 3.9 times 3rd segment;

length of maxillary palp 1.7 times height of head; dorsal length of eye 2.0. times

temple; temple roundly narrowed posteriad (fig. 722); POL : @ ocellus : OOL= 8:

9: 16; frons medially somewhat concave, with some striae behind antennal sockets

(fig. 722); vertex rather flat and smooth; face mainly flat, medio-dorsally with a

tubercle, punctulate, and dorsally somewhat rugulose (fig. 721); clypeus convex,

punctulate; apical margin of clypeus almost straight medially, thin, and well

differentiated from clypeus; length of malar space 0.8 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

crenulate medio-anteriorly, medially punctulate, and posteriorly crenulate-rugose

(fig. 714); epicnemial area punctulate, posteriorly indistinctly rugulose; precoxal

suture only anteriorly weakly rugulose, punctulate; rest of mesopleuron

punctulate; metapleural flange long, lamelliform, very wide, rounded anteriorly

(fig. 714); metapleuron reticulate-rugose ventrally, rest punctulate; notauli

indistinctly crenulate anteriorly, more distinctly crenulate posteriorly (fig. 728);

mesoscutal lobes punctulate; surface of propodeum smooth, except for some

crenulae posteriorly and an interrupted lamelliform lateral carina (fig. 726), medial

carina mainly and areola completely absent; posterior part of propodeum not

separated from antero-dorsal part (fig. 714).

Wings. — Fore wing: r : 3-SR : SR1= 6: 14: 65; SRI straight; cu-a inclivous,

postfurcal; 1-CUI : 2-CUl= 2: 23; 2-SR : 3-SR : r-m= 17: 14: 7; 2A shortly

sclerotized basally (fig. 719); area basally of 2A bare. Hind wing: r absent; 2-SC+R

transverse; SC +RI rather straight and long (fig. 725); basal third of SR straight

and unsclerotized (fig. 716).

Legs. — Hind coxa punctulate, with some apico-dorsal striae (fig. 727); tarsal

claws with a well-developed subapical sharp tooth (figs. 723, 724), yellowish

pectinate basally; length of femur, tibia and basitarsus of hind leg 7.6, 12.4, and

10.4 times their width, respectively; length of spurs of hind tibia 0.5 and 0.4 times

basitarsus.

Metasoma. — Length of Ist tergite 2.9 times its apical width, its surface smooth

(fig. 726); dorsal carinae of Ist tergite mainly absent; length of ovipositor sheath

0.07 times fore wing.

Colour. — Brownish-yellow; stemmaticum blackish; hind tarsus yellowish-

white; all tibiae, fore and middle tarsi and tegulae, rather light yellowish.

Type in TC, Ann Arbor: “10 m W. Silvia, Cauca, Colombia, VII.5.(19)70, 10,000’

(ft), H. & A. Howden”.

Homolobus (Oulophus) rectinervis spec. nov.

(figs. 734—747)

Holotype, 9, length of body 7.2, of fore wing 7.6 mm.

Head. — Antennal segments 44, 3rd segment 1.1 times 4th segment, without

VAN ACHTERBERG: Revision Zelinae auct. 349

ridge, length of 3rd and 4th segments 3.8 and 3.6 times their width, respectively,

length of both penultimate segments 1.7 and 2.0 times their width; length of 4th

segment of labial palp 2.8 times 3rd segment; length of maxillary palp 1.3 times

height of head; dorsal length of eye 2.0 times temple; temple rather roundly

narrowed posteriad (fig. 741); POL : & ocellus : OOL= 4: 4: 5; frons smooth,

weakly concave medially; vertex smooth, rather flat; face flat, punctulate laterally,

more punctate medially; clypeus convex, punctulate; apical margin of clypeus

scarcely differentiated from clypeus, thin, and straight medially (fig. 744); length

of malar space 0.8 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

medially crenulate and posteriorly rugose, rest mainly smooth (fig. 734);

epicnemial area smooth, except for some punctulation; precoxal suture and rest of

mesopleuron punctulate (fig. 734); metapleural flange lamelliform, large, rather

rounded apically; notauli narrowly crenulate (fig. 746); mesoscutal lobes mainly

smooth; surface of propodeum mainly smooth, medio-anteriorly with an irregular

medial carina and medio-posteriorly rugose, without areola; posterior part of

propodeum not separated from antero-dorsal part (fig. 734).

Wings. — Fore wing: r: 3-SR: SR1= 4: 8: 45; SRI weakly sinuate (fig. 737); cu-

a almost straight, postfurcal; 1-CU1 : 2-CU1= 5: 33; 2-SR : 3-SR : r-m= 12: 8: 7;

2A shortly sclerotized basally (fig. 738); area basally of 2A bare. Hind wing: r

absent; 2-SC+R transverse; SC+R1 almost straight (fig. 739); basal third of SR

almost straight and unsclerotized (fig. 737).

Legs. — Hind coxa rugulose dorso-anteriorly (fig. 745); tarsal claws setose, with

a small, slender subapical tooth (figs. 736, 740); length of femur, tibia and

basitarsus of hind leg 7.3, 11.5, and 9.6 times their width, respectively; length of

spurs of hind tibia 0.4 and 0.5 times basitarsus.

Metasoma. — Length of Ist tergite 2.4 times its apical width, its surface

posteriorly remotely and weakly rugose, laterally and basally mainly smooth (fig.

747); dorsal carinae of Ist tergite distinctly developed in basal third of tergite;

length of ovipositor sheath 0.14 times fore wing.

Colour. — Brownish-yellow; stemmaticum slightly infuscated.

Holotype in CNC, Ottawa: ‘‘Fundo Malcho, Cord. Parral, Chile, L. E. Pena,

Chile, 4.1II.62, L. E. Pena” (RMNH); 1 g (NR), “Ria Aysen (=Aisén, South

Chile)”, “P. Dusén”, “Riksmuseum Stockholm”; 1 & (TC), “Marga Marga River,

III.14— 15.64, Chile, Luis E. Pefia”, allotype; 1 9, “Bosque de los Conservadores

Graneros, 1100 m, O'Higgins, Chile, 1—4.111.62, Pefia” (CNC); | g, “Pichinahuel,

Cord., Nahuelbuta, Arauco, Chile, 20—28.1.1959, L. Pena” (CNC); 2 g, “El

Coigo, Curico, Chile, 1/7.11.1961” (CNC); I &, id., 1.1961 (CNC); 1 9, “Chile,

Cubillo, C. Curico, Curico, 24/26.1.1961, L. E. Pefia” (CNC); 1 9, “Pichinahuel,

“Tregualemu, Maule-Nuble, Chile, 7.XII.1953, L. Pefia” (CNC); 1 g, Penalolén,

Santiago, Chile, X.1953, L. E. Pena” (RMNH); 1 9, “Enco, Chile, Valdivia,

2.111.1955, L. E. Pefia” (RMNH); 1 9, “Las Nieves, 15.1X.47, Chile, L. E. Pena”

(CNC); 1 9, “Laguna Amarga, Magallanes, XII.12.60, Chile, T. Cehalovick”

350 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

(TC); 1 9, “Santiago Prov., Maipa, Chile, XII.28.66, Lionel Stange” (TC); 1 &,

“Pto. Aqua, L. Traful, Neuquen, Argentina, January 30, 1968, L. & J. Stange”,

(TC).

Variation: Length of body 6.8—7.8, of fore wing 7.1—8.4 mm; antennal

segments 42—44; length of malar space 0.6—0.8 times basal width of mandible;

length of fore wing 1.1—1.2 times body; length of ovipositor sheath 0.13—0.14

Key to the Palaearctic species of the genus Homolobus

|. Claws simple, without any protuberance (fig. 160); hind tibial spurs of &

rounded and pigmented apically, without a sharp, hyaline apex (fig. ne

(subgenus Apatia); avn dede ent DERE ee LA

— Claws with at least a minute subapical tooth (fig. 643) or with a lamella ce

629); hind tibial spurs of Z sharp and hyaline apically ............ 4

2. Length of outer aspect of 4th segment of labial palp 1.6—2.8 times its medium-

sized 3Ird segment (fig 309) Fr. ne rl ey ae Ra RA 3

— Length of outer aspect of 4th segment of labial palp 4—5 times its tiny 3rd

sevment (Fig TOA) jar ee ESTEN ER AR NRE truncator (Say) (p. 285)

3. Frontal aspect of head comparatively transverse (fig. 301), upper condyli of

mandibles of 9 close to lower level of eyes; length of malar space 0.3—0.7

times basal width of mandible; claws distinctly yellowish pectinate basally (fig.

DIA RES SE an ophioninus (Vachal) (p. 298)

— Frontal aspect of head more trapezoid, longer (fig. 311), upper condyli of

mandibles of 9 distinctly below lower level of eyes; length of malar space

0.6—1.1 times basal width of mandible; claws not or weakly pectinate basally

(figs 31353804) Er Ar EE truncatoides spec. nov. (p. 300)

A Vein-lA +2Asstraight (figs:396,419) I a Wa aoe Nee 6

— Vein 1A +2A curved (fig. 343) (subgenus Chartolobus) ............ 5

5. Antennal ridge at inner side of 4th—6th segments straight (fig. 348); vein 2A of

fore wing slender compared with its surrounding veins (figs. 343, 353)

CRATERE N Ss En infumator (Lyle) (p. 305)

— Antennal ridge of 4th—6th segments undulate (fig. 366); vein 2A of fore wing

widened basally, if compared with surrounding veins (fig. 369) ..........

oh Garhi SU SG gh MA ETE AE III undulatus spec. nov. (p. 309)

6. Vein SR of hind wing strongly curved and sclerotized basally (fig. 507); area

basally of vein 2A of fore wing remotely setose (fig. 496) (subgenus Phylacter)

ah DELI TEE RE EEN Ee 7

— Vein SR of hind wing straight or weakly curved and unsclerotized basally (figs.

539, 649); if, exceptionally, extensively sclerotized, then area basally of vein

2A of forewing mainly barei(fig. 630) RR 9

7. Claws bifurcate, subapical tooth large and in 9 truncate apically (figs. 488,

491); length of ovipositor sheath 0.12—0.14 times fore wing ............

de dale DIE a REEN bifurcatus spec. nov. (p. 322)

— Claws with a subapically sharp and tooth-shaped ventral lamella (figs. 498,

VAN ACHTERBERG: Revision Zelinae auct. 351

510); length of ovipositor sheath 0.17—0.25 times fore wing ......... 8

. Vein 2-SC +R of hind wing transverse, longer than wide (fig. 495) or quadrate;

hind tarsus more yellowish basally than medially, its 2nd—4th segments

whitish and contrasting with hind tibia ..... annulicornis (Nees) (p. 324)

Vein 2-SC+R of hind wing vertical, wider than long (fig. 507); hind tarsus

equally whitish yellow, only weakly contrasting with hind tibia .........

MI ee LY I ae AR meridionalis spec. nov. (p. 326)

. Inner aspect of 3rd—6th antennal segments of 9 with a longitudinal ridge (fig.

468); inner hind claw of 9 with a concavity ventro-subbasally (fig. 476), not

equal to its outer claw (fig. 475) (subgenus Homolobus) ............ 10

Inner aspect of 3rd—6th antennal segments of 9 without a ridge, or

exceptionally with a faintly developed ridge; inner hind claw of 9 convex or

straight ventro-subbasally (fig. 534), (sub)equal to its outer claw (fig. 536)

(eubeesnustOnlophus)user ow Shem a kono in Ai Fee 12

. Tarsal claws with a submedial lamella (figs. 390, 392); lamellae of middle and

hind claws of 4 with a 2nd lamella situated at the Ist lamella (figs. 393, 394)

sn En ee BIR NS a simplex (Watanabe) (p. 313)

Tarsal claws with a subapical tooth (fig. 443); claws of & without lamellae 11

. Ovipositor sheath short (fig. 459), 0.09—0.12 times fore wing; propodeum

without an areola, mainly smooth, except for some rugae (fig. 459);

mesopleunonsmooth van te ..… discolor (Wesmael) (p. 319)

Ovipositor sheath rather long (fig. 467), 0.36—0.39 times fore wing;

propodeum with a suboval areola, its surroundings usually rugose (fig. 467);

mesopleurompunctulate STI we dauricus Shestakov (p. 320)

. Malar space comparatively long (figs. 525, 532), its length 1.0—1.3 times basal

width of mandible; hind coxa more or less coarsely sculptured (figs. 523, 527);

both penultimate segments of antenna of 9 rather stout, their length 1.2—1.6

Limes their IK (IES Sete, 331) Aa. man en ae 13

Malar space comparatively short (figs. 549, 554), its length 0.3—0.8 times basal

width of mandible; hind coxa at most punctulate (fig. 553); both penultimate

segments of antenna of 9 more slender (fig. 640), their length 1.7—2.9 times

rele WYLIE vasten Re AP Ne RR RE E 14

. Subapical tooth of tarsal claws comparatively stout, rather blunt and subequal

to its apical tooth, resulting in sub-bifurcate claws (figs. 522, 524); body mainly

black; medially propodeum coarsely reticulate; vein r of hind wing absent (fig.

516) or only present as a short remnant (fig. 519); pterostigma dark brown;

hind basitarsus black or brownish basally and at least apical half of basitarsus

KC VERUEX SINOOtH OF punetulate (fig. 520) EEE was oe

oe ora rien dt ne ee carbonator (Shestakov) (p. 330)

Subapical tooth of tarsal claws slender, sharp, much shorter than the apical

tooth (figs. 531, 536); body yellowish and/or brownish; propodeum, except for

the carinae, only indistinctly sculptured (fig. 537); vein r of hind wing present,

but anterior half usually not distinctly developed (fig. 539), exceptionally

mainly absent; whole of hind basitarsus and pterostigma yellowish; vertex

EORTATCOUSMER O98) A UNE SU AN Ar bohemani (Bengtsson) (p. 332)

352

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

. Only basal quarter of vein SR of hind wing pigmented, not equal to vein Ir-m

(fig. 555); precoxal suture mainly smooth (fig. 553); antenna equally coloured,

without a white or yellowish ring; vein r of hind wing present (fig. 607) .. 15

Basal quarter of vein SR of hind wing equally sclerotized as vein Ir-m of hind

wing (fig. 635) precoxal suture extensively sculptured (fig. 633); antenna with a

white or yellowish ring medially; vein r of hind wing absent (fig. 635)

TE RIO IE MAR ER ai ene nae ea annulatus spec. nov. (p. 342)

. Ovipositor sheath rather long (fig. 577), 0.25—0.52 times fore wing;

mesonotum mainly black; pterostigma dark brown medially ........ 16

Ovipositor sheath short (fig. 553), 0.08—0.12 times fore wing; mesonotum

mainly yellowish-brown; pterostigma yellowish flagitator (Curtis) (p. 334)

. Length of ovipositor sheath subequal to length of metasoma and 0.51—0.52

times fore wing (fig. 588); length of vein 3-SR of fore wing 1.7—2.0 times vein

r; face blackish; length of malar space 0.6—0.7 times basal width of mandible

en NRA Bi SR ROSIE EN ENG: nipponensis spec. nov. (p. 338)

Length of ovipositor sheath 0.25—0.26 times fore wing and much shorter than

metasoma (fig. 602); length of vein 3-SR of fore wing 1.0—1.3 times vein r;

face reddish-brown; length of malar space 0.4—0.5 times basal width of

Mandiblest:t ees Pe Samen Hr a nepalensis spec. nov. (p. 340)

Key to the Nearctic species of the genus Homolobus

. Tarsal claws with at least a minute subapical tooth (fig. 545), which is

sometimes lamelliform (fig. 350); hind tibial spurs of & sharp and hyaline

APICI nst ur 2.0 mer an cee niece EEE 2

Tarsal claws simple, without any protuberance (fig. 160); hind tibial spurs of ¢

rounded and pigmented apically (fig. 112) (subgenus Apatia) ...........

RO ch ite cette: Lace RAN truncator (Say) (p. 285)

Vein IA +2A of fore wing straight (fig. 542) (subgenus Oulophus) ...... 3

Vein IA +2A of fore wing curved (fig. 353) (subgenus Chartolobus) .......

LEA erin Ming ENE OS DE od 2 infumator (Lyle) (p. 305)

Vein r of hind wing present (figs. 543, 555), at least posteriorly, as a brownish

pigmented stripe; precoxal suture mainly smooth (figs. 541, 553) . ...... 4

Vein r of hind wing completely absent (fig. 649); precoxal suture extensively

sculptured, at least dorsallly(fig.64 7) RENNES 6

Pterostigma and parastigma of 9 bicolorous, yellowish and dark brown; hind

tarsus brownish-yellow; vein r of hind wing short and comparatively straight

(fig.540); vertex comaceousi(hie550)) aaa bicolor spec. nov. (p. 333)

yellow; vein r of hind wing comparatively long and strongly reclivous (fig.

355); vertex. smooth (ig. ISOLE a ee 5

Precoxal suture with some rugae antero-dorsally (fig. 553); subapical tooth of

claws comparatively slender and claw weakly concave ventro-medially (figs.

558, 560); vein cu-a of fore wing parallel to vein 3-CUI (fig. 555); palpi, fore

and middle legs, more or less whitish-yellow; costulae of propodeum at least

VAN ACHTERBERG: Revision Zelinae auct. 353

partivipresent (fig SS3)f FAN IN flagitator (Curtis) (p. 334)

Precoxal suture smooth antero-dorsally (fig. 564); claws with an apical sharp,

tooth-shaped ventral lamella and claws straight medio-ventrally (figs. 570,

571); vein cu-a of fore wing more inclivous than vein 3-CUI (fig. 567); palpi,

fore and middle legs, brownish-yellow; costulae of propodeum absent (fig.

DOSE RENNENS) vena te rase sali hd, à acares spec. nov. (p. 336)

Ovipositor sheath medium-sized (fig. 694) or rather short (fig. 680), 0.12—0.36

times fore wing; subapical tooth of claws medium-sized (fig. 678) or rather

large (fig. 699); hind tarsus more whitish-yellow, contrasting with its brownish

IDE. ore CORE RO O SI MEA A Ti

Ovipositor sheath long (fig. 646), 0.68—0.79 times fore wing; subapical tooth

of claws large (fig. 657); hind tarsus and its tibia almost equally coloured, not

omwcaklyicontrasting., io. nerina armatus spec. nov. (p. 343)

Ovipositor sheath rather short (fig. 680), 0.12—0.15 times fore wing; length of

vein 3-SR of fore wing less than twice length of vein r (fig. 682) ..........

ME KURT oi antefurcalis spec. nov. (p. 345)

Ovipositor sheath medium-sized (fig. 694), 0.29—0.36 times fore wing; length

of vein 3-SR of fore wing more than twice vein r (fig. 698) .............

LER] ER ois RS Bla, mesoxiphius spec. nov. (p. 346)

Key to the Neotropical species of the genus Homolobus

Tarsal claws with at least a minute subapical tooth (fig. 740), which is

sometimes lamelliform (fig. 350); hind tibial spurs of Z sharp apically, with a

[VALINE QOS BE ee oe EEE: 2

Tarsal claws simple, without any protuberance (fig. 160); hind tibial spurs of &

rounded and pigmented apically (fig. 112) (subgenus Apatia) ...........

ATA Be au reede truncator (Say) (p. 285)

Vein IA +2A of fore wing straight (fig. 738) (subgenus Oulophus) ...... 3

Vein 1A +2A of fore wing curved (fig. 353) (subgenus Chartolobus) .......

vee dei NEE infumator (Lyle) (p. 305)

Vein r of hind wing present (fig. 567), at least posteriorly, as a brownish

OMS MIS CES CHEN ei Mia D pee SA las igs ithe wees 3 Bees CRN staan ge ie NM 4

Neinmot hind wine. completelyabsent,(fig..670) 6 "cer Len 5

Ovipositor sheath short (fig. 564), 0.08—0.10 times fore wing; pterostigma

unicolorous, yellowish; vertex smooth (fig. 568) .. acares spec. nov. (p. 336)

Ovipositor sheath medium-sized (fig. 577), ca. 0.43 times fore wing; ptero-

stigma bicolorous, medially dark brown, basally and apically yellowish; vertex

POLOSTRAAT a fa occidentalis spec. nov. (p. 337)

Vein SC+RI of hind wing curved (fig. 669); area basally of vein 2A of fore

wing sparsely setose (fig. 671); precoxal suture extensively sculptured

anteriorly (fig. 667); head and antenna dark brown .................

RE EE ae obscurus spec. nov. (p. 344)

Vein SC+RI of hind wing straight (fig. 725); area basally of vein 2A of fore

wing bare (fig. 719); precoxal suture usually mainly smooth (fig. 714) .... 6

354

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Length of fore wing of 9 ca. 1.3 times length of body (compare fig. 716 with

fig. 714); inner aspect of 3rd—8th antennal segments with a rather weakly

developed ridge (fig. 720); subapical tooth of claws larger, comparatively stout

(fig. 723); length of ovipositor sheath ca. 0.07 times fore wing ...........

Li sss RIE N. macropterus spec. nov. (p. 347)

Length of fore wing of 9 1.0—1.1 times length of body (compare fig. 737 with

fig. 734); inner aspect of 3rd—8th antennal segments without ridge; subapical

tooth of claws comparatively small and slender (fig. 740); length of ovipositor

sheath 0.13—0.14 times fore wing ........ rectinervis spec. nov. (p. 348)

Key to the Afrotropical species of the genus Homolobus

Hind claws of 9 with at least a minute sharp subapical tooth (figs. 426, 443)

sometimes claws bifurcate (fig. 406); inner aspect of 3rd—6th antennal

segments of 9 with a ridge (fig. 424); inner hind claw of 9 weakly concave

ventro-subbasally (figs. 427, 439), not the same shape as outer hind claw (figs.

426, 443) (subgenus Homolobus) ...... atd aN nata er Re 2

Hind claws of 9 simple, without a sharp subapical tooth (fig. 252), at most with

a blunt scarcely visible, subapical prominence (figs. 153, 237); inner aspect of

3rd—6th antennal segments of 9 without a ridge; inner hind claw of 9 convex

or straight ventro-subbasally (fig. 314), of the same shape as outer claw (fig.

313) (subgenus:Apatia)c.. 208 LE ER eee ee 5

Claws bifurcate (fig. 406); 2nd tergite rugose (fig. 404); vertex punctate (fig.

409); vein SC + RI of hind wing short, Rl mainly absent, and hamuli separated

fromRl (fig? 412) u A are rugosus spec. nov. (p. 314)

Claws with a small subapical tooth (fig. 426); 2nd tergite smooth (fig. 428);

vertex at most punctulate (fig. 429); vein SC+RI of hind wing longer, RI

shortly developed, and hamuli situated:at Ris AOS 3

Second tergite whitish; subapical tooth of tarsal claws of 9 scarcely visible at

80x (fig. 443) or, if easily visible, then length of ovipositor sheath 0.14—0.17

times fore wing: an 0a Bk eaten RS Du a SA 4

Second tergite dark brown and partly reddish or yellowish-brown; subapical

ovipositor sheath 0.07—0.08 times fore wing ethiopicus spec. nov. (p. 318)

Length of ovipositor sheath 0.14—0.17 times fore wing, about as long as apical

height of metasoma, slender as ovipositor (fig. 416); subapical tooth of tarsal

Ve le oe Nt Tn RUE NE cingulatus (Granger) (p. 315)

Length of ovipositor sheath 0.06—0.09 times fore wing, distinctly shorter than

apical height of metasoma, rather stout as ovipositor (fig. 431); subapical tooth

of tarsal claws of 9 minute and scarcely visible at 80 x (figs. 439,443) .....

M ut a or inopinus spec. nov. (p. 316)

First tergite of metasoma black, strongly contrasting with the (at least in part,

laterally) whitish 2nd and 3rd tergites; vein SR1 of fore wing straight (fig. 147);

VAN ACHTERBERG: Revision Zelinae auct. 355

precoxal suture mainly smooth (fig. 144) ..... albipalpis (Granger) (p. 283)

Three basal tergites of metasoma brownish-yellow, if more or less dark brown,

then 2nd tergite yellowish, dark brown or blackish and less contrasting with

Iste tergite; vein SRI of fore wing more or less curved (figs. 184, 196), if

straight (figs. 206, 219), then precoxal suture extensively sculptured (figs. 204,

ORE Sen haul delicata SER teh ra SRE EME Mit, 6

Length of outer aspect of 4th segment of labial palp 3.0—5.0 times the small

3rd segment (figs. 200, 222, 235), if intermediate, then vein cu-a of fore wing

antefurcal (fig. 206) and/or apical half of metasoma mainly dark brown or

blaekish pad, Serie ACLI A AE EES NR dl

Length of outer aspect of 4th segment of labial palp 1.6—2.8 times the

medium-sized 3rd segment (figs. 246, 257, 265); vein cu-a of fore wing more or

less postfurcal (figs. 266, 329); metasoma mainly yellowish apically .... 10

Tarsal claws of 9 with a tiny prominence subapically (figs. 212, 225); veins

SRI of fore wing and SR of hind wing straight or nearly so (figs. 206, 234) . 8

Tarsal claws of @ without any prominence (figs. 202, 203); vein SRI of fore

wing curved (fig. 196) vein SR of hind wing sinuate (fig. 196) ............

ENE esterni ada cla wiicaeorac rufithorax (Granger) (p. 289)

Length of malar space 1.2— 1.6 times basal width of mandible (fig. 210); apical

half of metasoma blackish or dark brown ... maculatus spec. nov. (p. 291)

Length of malar space 0.7—1.0 times basal width of mandible (figs. 229, 238), if

intermediate, then apical half of metasoma yellowish .............. 9

Vein r of fore wing longer than 3-SR (fig. 219); subapical prominence of claws

of 9 very small, scarcely visible at 80x (figs. 225, 226); head, antenna and hind

leg, mainly dark brown; palpi, tegulae, fore and middle coxae, yellowish

WINGS) ARTE alternipes spec. nov. (p. 292)

Vein r of fore wing shorter than 3-SR (fig. 234), exceptionally of equal length;

subapical prominence of claws of 9 small, but at 80x easily visible (fig. 237);

head, antenna, hind leg, palpi, tegulae, fore and middle coxae, brownish

KEOMO AEN a RT HE AT priapus (Nixon) (p. 293)

. Vein SR of hind wing strongly sinuate (figs. 243, 258); marginal cell of hind

wing distinctly narrowed medially, in respect to its basal width (figs. 254);

middle lobe of mesoscutum finely and densely punctate or punctulate (figs.

250, 262); scapus more or less dark brown; vein SC+RI of hind wing

Comparatively shorti(figssi254; 239, m eeN, LENS, MATE PSE Ne, 11

Vein SR of hind wing weakly sinuate (fig. 266); marginal cell of hind wing not

or weakly constricted medially, in respect to its basal width (fig. 290); middle

lobe of mesoscutum smooth or weakly punctulate (fig. 300); scapus mainly

yellowish; vein SC + RI of hind wing somewhat longer (figs. 267, 307) .. 12

. Marginal ceil of hind wing constricted just after middle of the cell (fig. 243);

length of ovipositor sheath 0.24—0.26 times fore wing, the exserted ovipositor

longer than 1.5 times length of Ist tergite (fig. 240); propodeum and Ist tergite

irregularly sculptured (figs. 240,253) ...... lacteiceps spec. nov. (p. 294)

Marginal cell of hind wing constricted in front of middle of the cell (fig. 258);

length of ovipositor sheath ca. 0.14 times fore wing, the exserted ovipositor

-356

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

slightly longer than Ist tergite (fig. 280); propodeum and Ist tergite evenly,

finely and densely rugulose (figs. 142, 255) ... pulchricornis (Nixon) (p. 296)

. Vein 2-SC+R of hind wing transverse, longer than wide (fig. 290); length of

hind femur 5.6—7.2 times its maximum width, usually comparatively slender

(fig. 291), if intermediate, then upper condyli of mandibles rather far below

lower level of eyes or Ist tergite more slender, longer than 2.2 times its apical

width (fig. 311); lateral aspect of hind tibial spurs of & more or less truncate

apically (figs: 2965 297) STEN LR IP Er sade Ais are APA 13

Vein 2-SC +R of hind wing vertical or quadrate (fig. 267); length of hind femur

4.6-5.8 times its maximum width, comparatively stout (fig. 269); upper condyli

of mandibles comparatively close to lower level of eyes (fig. 270); Ist tergite

stout, its length 1.7—2.2 times its apical width (fig. 271); lateral aspect of hind

tibial spurs of ¢ sharp apically (figs. 272, 273) huddlestoni spec. nov. (p. 297)

. Frontal aspect of head comparatively long, trapezoidal (figs. 311, 330); upper

condyli of mandibles of 9 distinctly below lower level of eyes (figs. 311, 330);

length of malar space 0.8—1.1 times basal width of mandible, if exceptionally

shorter, then claws setose basally (figs. 313, 314) ................ 14

Frontal aspect of head comparatively short, transverse (fig. 301); upper

condyli of mandibles of 9 close to lower level of eyes (fig. 301); length of

malar space 0.3—0.7 times basal width of mandible; claws yellowish pectinate

basally (fie 294) ee m UIID ophioninus (Vachal) (p. 298)

. Vein SC+RI of hind wing somewhat curved and shorter (figs. 306, 307);

marginal cell of hind wing usually less widened apicad, its apical width

1.9—2.2 times its maximum basal width (fig. 306); length of fore wing 3.5—7.1

mm; claws only setose or indistinctly pectinate basally (figs. 313, 314);

ovipositor sheath in undistorted position rather wide apically (fig. 303)... ..

POEUN Oa EE MEN BE ACER ees EE truncatoides spec. nov. (p. 300)

Vein SC+RI of hind wing almost straight and somewhat longer (figs. 329,

337); marginal cell of hind wing more widened apicad, its apical width 2.4—2.6

times its maximum basal width (fig. 329); length of fore wing 7.0—9.5 mm;

claws distinctly pectinate basally (figs. 339, 340); ovipositor sheath slightly

moreslenden(fig 335) been eee pallidistigmus (Cameron) (p. 303)

Key to the Oriental and Australian species of the genus Homolobus (Himalayan

area included)

Claws simple, without any protuberance (fig. 295); hind tibial spurs of ¢ often

rounded and pigmented apically (fig. 112) (subgenus Apatia) ......... 2

Claws with at least a minute subapical tooth (fig. 638) or with a lamella (fig.

624); hind tibial spurs of 3 always sharp and hyaline apically ......... 6

Vein r of hind wing absent (fig. 130); hind tibial spurs of & rounded and

pigmented apically (ig 112) eee, eee ee 3

Vein r of hind wing present (fig. 122); hind tibial spurs of Z sharp and hyaline

apically 20} Lo enen E elagabalus (Nixon) (p. 280)

First tergite of metasoma black, strongly contrasting with the laterally whitish

VAN ACHTERBERG: Revision Zelinae auct. 397

2nd and 3rd tergite; anteriorly precoxal suture mainly smooth (fig. 128) ....

EE PART CEOS ET RATEN australiensis (Nixon) (p. 282)

Basal three tergites of metasoma equally brownish-yellow; anteriorly precoxal

sutunrerextensively sculptured (figs: 2872302) 72%. nme nn. 4

Length of outer aspect of 4th segment of labial palp 4—5 times its tiny 3rd

Stond OU) PRA da SS truncator (Say) (p. 285)

Length of outer aspect of 4th segment of labial palp 1.6—2.8 times its medium-

Srederdiseement (Mg SOD) Pee ee MI 4 5

Frontal aspect of head comparatively transverse (fig. 301); upper condyli of

mandibles of 9 close to lower level of eyes; length of malar space 0.3—0.7

times basal width of mandible; claws yellowish pectinate basally (figs. 294,

DOS) à 41010 EAA ENE: ophioninus (Vachal) (p. 298)

Frontal aspect of head comparatively long (fig. 311); upper condyli of

mandibles of 9 distinctly below lower level of eyes (fig. 311); length of malar

pectinate basally (figs. 313, 314) ........ truncatoides spec. nov. (p. 300)

Vein 1A+2A of fore wing curved (figs. 353, 369, 380) (subgenus Chartolo-

Di MEER GAS athe MA ta BONS ALT de AU AR AOR LC Eed 7,

Vein 1A +2A of fore wing straight (fig. 619) .................... 9

Vein 2A of fore wing widened, if compared with its surrounding veins (figs.

DOS D RRL DA Lab AE A RTRT TA RISATA e AAT EIA 8

Vein 2A of fore wing slender, if compared with its surrounding veins (fig. 353)

MP EN RE EN MIE) ERBE IR, gui infumator (Lyle) (p. 305)

Basal third of vein SR of hind wing weakly curved (figs. 367, 368); pterostigma

light brownish or yellowish brown; hind tarsus yellowish or whitish .......

Bee TO, WL Ed enr JTE. undulatus spec. nov. (p. 309)

Basal third of vein SR of hind wing almost straight (fig. 382); pterostigma and

findktarsus Dlackish#(s YY 10 Hu Ta See, nigritarsis spec. nov. (p. 310)

Vein SR of hind wing strongly curved and extensively sclerotized basally (fig.

484); area basally of vein 2A of fore wing remotely setose (fig. 485); claws

bifurcate, the subapical tooth large and in Q truncate apically (figs. 488, 491)

GubgenusPhylacter) ron nenn dir bifurcatus spec. nov. (p. 322)

Vein SR of hind wing straight or weakly curved, usually only pigmented

basally, if exceptionally extensively sclerotized, then area basally of vein 2A of

fore wing mainly bare (fig. 619); claws with lamella (fig. 624) or with a sharp

medium-sized or small subapical tooth (fig. 534) (subgenus Oulophus) .. 10

. Frontal aspect of head comparatively long (figs. 525, 535); length of malar

space 1.0—1.3 times basal width of mandible; hind coxa more or less coarsely

sculptured (figs. 513, 527); length of both penultimate segments 1.2—1.6 times

hote Ra SLOSS EE EN Il

Frontal aspect of head comparatively transverse (fig. 620); length of malar

space 0.3—0.8 times basal width of mandible; hind coxa at most punctulate

and somewhat rugose dorsally (fig. 616); both penultimate segments of

. Subapical tooth of tarsal claw of 9 comparatively stout, rather blunt and

358 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

subequal to the apical tooth, resulting in sub-bifurcate claws (figs. 522, 524);

body mainly black; medially propodeum mainly coarsely reticulate; vein r of

hind wing absent or at most an indistinctly developed remnant present (figs.

516, 519); pterostigma dark brown; basally hind basitarsus more or less

blackish or dark brown and at least its apical half white; vertex smooth or

punctulater(higo20) ns Sepals oes EE carbonator (Shestakov) (p. 330)

— Subapical tooth of tarsal claws of Q slender, sharp, much shorter than the

apical tooth (figs. 534, 536, 707); body brownish and/or yellowish; propodeum

(except for the carinae) indistinctly sculptured (fig. 537); vein r of hind wing

present, but anterior half usually indistinctly developed (fig. 529); excep-

tionally largely absent; pterostigma and whole hind basitarsus yellowish;

VEGECXICORACECOUS (NES) an bohemani (Bengtsson) (p. 332)

12. Vein r of hind wing present, at least as a brownish pigmented stripe (figs. 555,

607); basal quarter of vein SR of hind wing only pigmented, unsclerotized (fig.

S55) ney Ala ancl ell ie a pe ees Zara SIE 18

— Veinr of hind wing completely absent; basal quarter of vein SR of hind wing

equally sclerotized as vein 1-M of hind wing (fig. 618) ............. 14

13. Length of ovipositor sheath 0.08—0.12 times fore wing (fig. 553); apical two-

thirds of hind tibia and metasoma yellowish .... flagitator (Curtis) (p. 334)

— Length of ovipositor sheath 0.25—0.26 times fore wing (fig. 602); apical two-

thirds of hind tibia and mesosoma mainly blackish ..................

AT OT N BO ca de oe pen D nepalensis spec. nov. (p. 340)

14. Vein 2-SC +R of hind wing long, transverse (fig. 618); base of vein SR of hind

wing straight (fig. 631); tarsal claws with a ventral lamella (fig. 624); antenna,

hind tibia and body, yellowish. .......... crenulatus spec. nov. (p. 341)

— Vein 2-SC+R of hind wing short, vertical (fig. 635); base of vein SR of hind

wing weakly curved (fig. 641); tarsal claws with a subapical tooth (fig. 638);

antenna mainly dark brown, but with a white or yellowish ring medially; body

and apical 0.7 of hind tibia mainly brownish black .................

Subfamily EUPHORINAE Foerster

Foerster, 1862, Verh. naturh. Ver. preuss. Rheinl. 19: 228, 250.

Syn.: Perilitinae Foerster, 1862.

Diagnosis. — Antescutal depression and hypoclypeal depression absent; Ist

discal cell of fore wing (sub)petiolate, vein 1-SR present or nearly so; dorsope of

Ist tergite present or (less frequently) absent; Ist tergite of metasoma petiolate,

subsessile or sessile, more or less narrowed in front of spiracles, exceptionally

weakly narrowed behind spiracles; apical segment of antenna variable (figs. 767,

787, 847); occipital carina connected with the hypostomal carina above the

mandibular base; veins a and CUIb of fore wing absent (fig. 836), exceptionally a

short part of CUIb present (fig. 792); pronope of pronotum more or less developed

(fig. 874); metapleural flange variable; hypostomal and prepectal carina present;

lateral carina of scutellum absent; lateral carina of mesoscutum more or less

VAN ACHTERBERG: Revision Zelinae auct. 359

developed; vein m-cu of fore wing usually antefurcal or interstitial with 2-SR,

exceptionally postfurcal; subbasal cell of hind wing usually large; antennal

segments 15—50; lobes of mesoscutum equally convex; trochantelli simple,

without teeth; scapus (sub)truncate apically; vein 2A of fore wing usually absent or

shortly developed; Ist subdiscal cell of fore wing more or less open ventro-distally;

plical lobe of hind wing and laterope of Ist tergite variable; maxillary and labial

palpi with 6 and 3—4 segments, respectively; metasoma usually sparsely setose,

less frequently densely and evenly setose; occipital carina present, at least

laterally; postpectal carina absent or nearly so; hypopygium truncate apically,

large to medium-sized; ovipositor straight or curved ventrad, usually with a small

subapical notch.

Distribution. — Cosmopolitan. Contains three tribes: Meteorini Cresson, 1887;

Centistini Capek, 1970; Euphorini Foerster, 1862.

Tribus METEORINI Cresson

Cresson, 1887, Trans. Am. ent. Soc., Suppl.: 55, 60.

Syn.: Zelini Ashmead, 1900; Petiolarini Szepligeti, 1904; Zemiotini Van Achter-

berg, 1976.

Diagnosis. — Occipital carina complete; 3rd segment of labial palp reduced

(figs. 809, 821) or (virtually) absent (figs. 759, 800); anterior tentorial pits deep,

medium-sized (fig. 773) or large (fig. 781); mesopleuron more or less protruding

antero-dorsally (fig. 772); medial lobe of mesoscutum more or less rounded

anteriorly (figs. 755, 874); vein 2A of hind wing absent (fig. 388), or at most present

as a vague stripe (figs. 750, 784); vein 2-RI of fore wing absent (fig. 758), or

(exceptionally) well developed (fig. 836); Ist tergite of metasoma petiolate and its

spiracles situated submedially (figs. 754, 775, 852); vein r-m of fore wing present

(fig. 18); Ist tergite widened apicad (figs. 754, 852).

Distribution. — Cosmopolitan. Contains two described genera: Meteorus

Haliday, 1835, and Zele Curtis, 1832.

Genus Zele Curtis

Curtis, 1832, Br. Ent. 9: 415.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 49.

Shenefelt, 1970, id. 5(2): 220.

Capek, 1970, Can. Ent. 102: 848.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 254— 300, figs. 3, 6.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 222—224.

Capek, 1972, Ent. Problémy 10: 133, 134, 136, 138.

Mason, 1973, Proc. ent. Soc. Wash. 75: 213—215.

Jakimavicius, 1974, Tr. AN Lit. SSR B2(66): 97.

Van Achterberg, 1976b, Tijdschr. Ent. 119: 37, 50, figs. 107, 111.

Tobias, 1976, Opr. Fauna SSSR 110: 113, fig. 33: 14—16.

Type-species: Zele testaceator Curtis.

Synonyms: Zemiotes Foerster, 1862, Syn. nov.; Protelus Foerster, 1862, Syn. nov.;

360 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Meteorus auct. p.p.

Diagnosis. — Length of body 4.1—10.3, of fore wing 3.9—10.8 mm; antennal

segments 29—50; length of 3rd segment of antenna 0.9—1.2 times 4th segment;

length of maxillary palp 1.0—1.5 times height of head; length of malar space

0.1—0.7 times basal width of mandible; mandible with a pair of (more or less)

protruding thin lamelliform carinae (figs. 809, 858), more or less twisted apically;

ventral margin of clypeus rather wide, lamelliform, straight medially and well

differentiated from clypeus (figs. 765, 781); eyes bare, immarginate, more or less

converging ventrad (figs. 765, 793, 830, 862), larger in ¢ than in 9; epistomal

suture complete (fig. 803); propleural lamellae more or less developed (figs. 785,

796); notauli complete (figs. 790, 849); side of scutellum more or less rugose (figs.

755, 808, 811); scutellum sculptured medio-posteriorly (figs. 755, 785); episternal

scrobe medium-sized, elliptical (figs. 748, 778, 785); metapleural flange more or

less lamelliform (figs. 748, 796); propodeal spiracle small and round (figs. 748,

796); propodeal tubercles absent; propodeum with a more or less developed

anterior transverse carina (figs. 754, 762, 775, 801); vein m-cu of fore wing more or

less antefurcal (figs. 750, 758, 784); Ist discal cell of fore wing shortly petiolate

anteriorly (figs. 758, 768, 836); short remnant of vein 2A of fore wing present (figs.

814, 827, 846); marginal cell of hind wing widened apicad (figs. 784, 788); vein SRI

of fore wing straight; vein r of hind wing present (fig. 788) or absent (fig. 758);

tarsal claws with a large submedial lobe (figs. 752, 757, 791); length of hind femur

3.8—7.6 times its width; length of Ist tergite 1.6—4.1 times its apical width, its

dorsope more or less developed (figs. 762, 794); at least apical half of 3rd and

following tergites densely setose (figs. 783, 794); 2nd and following tergites

smooth, only in gracilis weakly coriaceous-punctulate (fig. 852); length of

ovipositor sheath 0.19—0.60 times fore wing, slender (fig. 843); length of hind

tibial spurs 0.3—0.4 times hind basitarsus, subequal.

Distribution. — Widespread, but absent in the Afrotropical and Australian

regions. The distribution is rather similar to that of the subgenus Oulophus of the

genus Homolobus and may be due to the same factors, e.g., a primarily Holarctic

centre of speciation, combined with a secondary centre in the Neotropical area.

Biology. — Endoparasites of larvae of the Geometridae Pyralidae, Noctuidae,

Lasiocampidae, Lymantriidae, Arctiidae, Limacodidae, and Saturniidae. Aberrant

records from Tortricidae, Momphidae, Douglasiidae, Yponomeutidae, Lyone-

tiidae, Gelechiidae, Conchylidae, Pterophoridae and Nymphalidae need to be

confirmed.

Note. — There is some confusion about the gender of the genus Zele Curtis.

Because it is frequently used in generic combinations (Austrozele, Palinzele,

Neozele, Xiphozele, all belonging to other subfamilies) certainty about its gender is

required. The name Zele is a fantasy-name which takes the gender expressly

attributed to it by its author. If no gender is assigned or implied, the name is to be

treated as masculine unless the ending is clearly a natural classical feminine or

neuter one when the gender is that appropriate to the ending (according to Article

30a(ii) of the International Code of Zoological Nomenclature). Curtis (1832: 415)

‘id not expressly attribute a gender to his new genus, while of the ten species

VAN ACHTERBERG: Revision Zelinae auct. 361

included, only two may imply a certain gender of Zele, viz., thoracicus, and

longicauda . Thus Curtis himself was uncertain about the gender, furthermore also

the ending is not a natural classical feminine or neuter one. Therefore the name

Zele (and the derived names as well) has to be treated as a masculine noun

according to Article 30a(ii) ofthe International Code.

Key to the species of the genus Zele

l. Precoxal suture smooth or only narrowly sculptured (figs. 748, 758, 766), if

intermediate, then length of ovipositor sheath ca. 0.2 times fore wing (fig. 756);

vein 1-M of hind wing 0.7—1.1 times vein cu-a (figs. 750, 758), exceptionally

somewhat shorter; Ist tergite usually comparatively stout (figs. 754, 762, 775);

Ueinicu-atefifiore Wing postiurcal (fig, 768) 0 CAR ee e nn CURE )

— Precoxal suture widely sculptured, at least anteriorly or medially (figs. 778,

785, 796), if intermediate, then vein 1-M of hind wing shorter than 0.7 times

vein cu-a (figs. 836, 855) or vein cu-a of fore wing antefurcal (fig. 798); Ist

tergite comparatively slender (figs. 824, 839, 852, 864) or length of ovipositor

sheathtear 0 3 times tore Wingi(fip: 833), 1 RER VE ee, ee oe kal ae 4

2. Length of ovipositor sheath 0.19—0.28 times fore wing (figs. 756, 767); hind

femur slender (figs. 761, 774), its length 5.2—6.4 times its width ....... 3

— Length of ovipositor sheath 0.38—0.39 times fore wing (fig. 748); hind femur

somewhat swollen (fig. 751), its length 4.4—5.l times its width; North

Paldicanclicmeree. seats A siete hin. bate a annulicrus (Thomson) (p. 363)

3. Body, hind tibia and its tarsus mainly dark reddish-brown; pterostigma of 9

more or less brown; mesopleuron somewhat more sculptured (fig. 756);

Palaearctic and North Nearctic ........... caligatus (Haliday) (p. 364)

— Body, pterostigma, and hind leg of 9, yellowish; mesopleuron comparatively

smooth (fig. 766); South Nearctic .......... levis (Muesebeck) (p. 365)

4. Scutellum protruding dorsad, with a tubercle (figs. 778, 785); laterope absent

(nes 88 18S)vertex punctate (lies. 779,789) 20... 2. n 5

— Scutellum at most rather strongly convex, without tubercle (figs. 796, 825);

laterope present, at least shallowly (figs. 825, 843); vertex smooth or

purciulate (VEN ide ee EER RA Re ne 6

5. Tubercle of scutellum rounded apically (figs. 778, 780); dorsope of Ist tergite

well developed (fig. 783); wing membrane hyaline; hind tibia brownish, only

with blackish setae; South Neotropical ...... punctatus spec. nov. (p. 367)

— Tubercle of scutellum sharp apically (figs. 785, 795); dorsope almost absent

(fig. 794); wing membrane light brownish; apical 0.7 of hind tibia (at least in ¢)

blackish; North Neotropical ........... tuberculifer spec. nov. (p. 368)

6. Length of ovipositor sheath 0.41—0.60 times fore wing, longer than 1.5 times

Ist tergite of metasoma (figs. 796, 815); vein cu-a of fore wing antefurcal (figs.

MOERS wexceptionally interstitial wears ee. ER fis ots ser 7

— Length of ovipositor sheath 0.19—0.33 (exceptionally 0.37) times fore wing,

shorter than 1.5 times length of Ist tergite (figs. 825, 834, 843, 853); vein cu-a of

fore wing postfurcal (figs. 827, 855), seldom (sub)interstitial (fig. 836), and

362

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

exceptionally shortly antefurcalis Presenter 9

Hind tarsus yellowish; eyes somewhat smaller, their dorsal length in 9

1.3—2.1 (in & 1.2—1.6) times length of temple (fig. 817); West and Middle

Palaearctie „na ee Rs ee chlorophthalmus (Spinola) (p. 370)

Hind tarsus whitish; eyes of 9 comparatively large, their dorsal length

2.4—3.2 (in ¢ 1.8—2.3) times length of temple (fig. 892); (niveitarsis Cresson

SI) ecn Le AO ORO er 8

Hind tibia and pterostigma mainly dark brown; East Palaearctic and

Oriental rer ie a toes as niveitarsis f. peronatus (Shestakov) (p. 370)

Hind tibia and pterostigma mainly brownish-yellow; Nearctic ..........

EE niveitarsis f. niveitarsis (Cresson) (p. 369)

Scutellum rather strongly convex (figs. 811, 825); surroundings of veins 1-M, 1-

CUI, and r of fore wing dark brown pigmented (fig. 827); basal two-thirds of

hind tibia, all trochanters and trochantelli, white; pterostigma of 9 dark

brown, with base and apex whitish; malar space of 9 very short, its length 0.1

times basal width of mandible (fig. 830); South Nearctic ..............

wi ith hee ln AERC te RT picinervis spec. nov. (p. 373)

Scutellum weakly convex (fig. 834); surroundings of veins 1-M, 1-CUI and r of

fore wing hyaline or faintly brownish; basal half of hind tibia, all trochanters

and trochantelli, yellowish or dark brown; pterostigma of Q uniformly

yellowish or dark brown; length of malar space of 9 0.3—0.4 times basal width

of mandible, somewhat longer (figs. 842,802). casei cee eee 10

. Hind femur stout, its length 3.8—4.4 times its maximum width (fig. 840); head

more transverse (fig. 838); vein 2-R1 of fore wing well-developed, somewhat

longer than vein r (fig. 836); Nearctic ... crassifemur (Muesebeck) (p. 374)

Hind femur slender, its length 5.1—7.6 times its maximum width (fig. 860);

head less transverse (figs. 850, 863); vein 2-RI of fore wing rather short, usually

shorter than veinir. (figs: 840,899) EEN 11

. Length of vein r of fore wing 1.1—1.2 times vein 3-SR (fig. 846); 2nd tergite

evenly setose and finely coriaceous-punctulate (fig. 852); base of Iste tergite

and ventral half of temple, yellowish white; length of ovipositor sheath ca. 0.37

times fore wing (fig. 843); South Palaearctic .... gracilis spec. nov. (p. 375)

Length of vein r of fore wing 0.3-0.6 times vein 3-SR (figs. 855, 868); at least

basal half of 2nd tergite bare and smooth (fig. 853); base of Ist tergite and

ventral half of temple, brownish-yellow or blackish; length of ovipositor

sheath 0.19—0.33 times fore wing (fig. 853); Holarctic; (albiditarsus Curtis

Sel.) sen deet rte E Ge ab ee Pe er 12

. Length of vein I-M of hind wing 0.9—1.3 times cu-a, subequal (fig. 868) .

Se Pade SARAI albiditarsus f. pallitarsis (Cresson) (p. 379)

Length of vein 1-M of hind wing 0.3—0.8 times vein cu-a, usually much shorter

than-cuza (fig. 853), a. eren A Ped Oa 13

. Middle of hind tarsus yellowish or infuscated, if intermediate, then similarly

coloured as middle of hind femur; length of fore wing usually less than 8

MD. pate er ae eee albiditarsus f. deceptor (Wesmael) (p. 377)

Middle of hind tarsus white or whitish-yellow, lighter coloured than middle of

VAN ACHTERBERG: Revision Zelinae auct. 363

hind femur; length of fore wing usually more than 8mm ..............

RL PAN ACNE re SUR albiditarsus f. albiditarsus Curtis (p. 380)

Zele annulicrus (Thomson) comb. nov.

(figs. 748— 755)

Thomson, 1895, Opusc. ent. 20: 2161 (as Meteorus).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 52, 53.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258.

Lectotype, 9, length of body 4.2, of fore wing 4.4 mm.

Head. — Remaining antennal segments 11, 3rd segment 0.9 times 4th segment,

length of 3rd and 4th segments 3.4 and 3.6 times their width, respectively; length of

maxillary palp 1.1 times height of head; dorsal length of eye 1.4 times temple;

temple weakly roundly narrowed posteriad (fig. 749); POL : @ ocellus: OOL = 14:

5:6; frons weakly concave, smooth; vertex convex, punctulate; face weakly

convex, indistinctly punctulate; clypeus strongly convex, punctate (fig. 753);

length of malar space 0.4 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

largely smooth, medially crenulate and ventro-anteriorly and posteriorly rugose

(fig. 748); epicnemial area crenulate anteriorly, dorsally rugose near posterior

subalar depression; precoxal suture narrowly crenulate medially (fig. 748);

metapleural flange medium-sized, apically lamelliform and rounded (fig. 748);

metapleuron almost smooth, ventrally rugose; notauli indistinctly crenulate,

narrow (fig. 755); mesoscutal lobes slightly punctulate; scutellum convex,

punctulate; surface of propodeum with a small areola medio-anteriorly and a well-

developed medial carina and costulae (fig. 754), its surroundings almost smooth;

posterior part of propodeum not separated from antero-dorsal part (fig. 748).

Wings. — Fore wing: r: 3-SR : SRI = 4: 7: 37; cu-a postfurcal; 1-CUI : 2-CUI

= 2: 15; 2-SR : 3-SR: r-m = 10:7: 7. Hind wing: r mainly absent (fig. 750); length

of 1-M 0.9 times cu-a.

Legs. — Hind coxa punctulate; hind femur somewhat curved (fig. 751); length

of femur, tibia, and basitarsus of hind leg 4.4, 10.2 and 8.6 times their width,

respectively.

Metasoma. — Length of Ist tergite 1.6 times its apical width, its surface

superficially and remotely striate (fig. 754); dorsal carinae present in front of

spiracles; dorsope and laterope large and deep; 2nd tergite smooth and bare;

length of ovipositor sheath 0.38 times fore wing.

Colour. — Dark brown; scapus, pedicellus, clypeus, labrum, mandibles mainly,

palpi, pterostigma, tegulae, postero-dorsal corner of pronotum, and legs,

yellowish; hind tibia basally whitish, its apical three-quarters and hind tarsus

darkened.

Lectotype in ZIL, Lund: ‘“Hbg”, “Meteorus annulicrus Th., Type, det. Fischer”,

‘1977, 39”. Paralectotypes: 4 specimens, of which | 9 and 2 Z were examined. |

31; length of fore wing 4.6—4.7 mm; length of Ist tergite 1.6—1.7 times its apical

364 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

width; length of hind femur 4.4—5.1 times its apical width; length of ovipositor

sheath 0.38—0.39 times fore wing; pterostigma of ¢ dark brown.

Zele caligatus (Haliday) comb. nov.

(figs. 756—765)

Haliday, 1835, Ent. Mag. 3: 25 (as Meteorus).

Ruthe, 1862, Berl. ent. Z. 6: 22, 23 (Meteorus neesii).

Ashmead, 1902, Proc. Wash. Acad. Sci. 4: 247 (Dyscoletes alaskensis). Syn. nov.

Fahringer, 1930, Ark. Zool. 21A: 8 (Meteorus caligatus var. sibiricus). Syn. nov.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 51, 55, 56.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 222.

Mason, 1973, Proc. ent. Soc. Wash. 75: 214.

Lectotype, 9, length of body 5.1, of fore wing 5.3 mm.

Head. — Antennal segments 35, 3rd segment equal to 4th segment, length of 3rd

and 4th segments both 3.2 times their width, length of both penultimate segments

1.3 and 1.6 times their width; length of maxillary palp 1.3 times height of head;

dorsal length of eye 2.0 times temple; temple roundly narrowed posteriad (fig.

760); POL : & ocellus : OOL = 7: 4: 5; frons smooth, but with some rugosity near

antennal sockets, almost flat; vertex convex, weakly punctulate; face rather flat,

punctate medially and near antennal sockets, laterally punctulate (fig. 765);

clypeus convex, punctate; length of malar space 0.4 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely punctulate, antero-medially and apically crenulate and indistinctly rugose

ventrally (fig. 756); epicnemial area mainly smooth, rugose postero-dorsally;

precoxal suture narrowly and irregular crenulate, anteriorly and posteriorly almost

smooth (fig. 756); metapleural flange medium-sized, narrowly lamelliform apically

(fig. 756); metapleuron reticulate, only dorsally almost smooth; notauli narrowly

crenulate (fig. 764); mesoscutal lobes indistinctly punctulate; scutellum rather

convex, weakly punctulate; surface of propodeum mainly smooth anteriorly,

except for a weakly developed transverse carina and a short part of the medial

carina (fig. 762), posteriorly rugose; posterior part of propodeum not separated

from its antero-dorsal part (fig. 756).

Wings. — Fore wing: r: 3-SR : SRI = 10: 24: 114; cu-a postfurcal; 1-CUI : 2-

CUI = 1:20; 2-SR : 3-SR : r-m = 19: 12: 12. Hind wing: r absent; length of 1-M

0.9 times cu-a.

Legs. — Hind coxa weakly punctulate; hind femur rather straight (fig. 761);

length of femur, tibia and basitarsus of hind leg 5.3, 10.2 and 7.7 times their width,

respectively.

Metasoma. — Length of Ist tergite 1.7 times its apical width, its surface

indistinctly rugulose, almost smooth (fig. 762); dorsal carinae of Ist tergite absent;

laterope and dorsope large and deep (fig. 756); 2nd tergite mainly bare and

smooth; length of ovipositor sheath 0.19 times fore wing.

Colour. — Dark reddish-brown; palpi, mandibles, clypeus ventrally, antenna

(except apically), tegulae, dorso-posterior corner of pronotum, legs (but hind tibia

VAN ACHTERBERG: Revision Zelinae auct. 365

and tarsus mainly infuscated), 2nd and 3rd tergites and their sternites, apex of

hypopygium, apex of ovipositor sheath, yellowish; pterostigma brown; hind tibia

basally with a whitish ring.

Lectotype in NMI, Dublin: “Jullymore”, “Ireland, Haliday, 20.2.82/Box 8,

A.W.S.”, “Meteorus caligatus Hal., det. Muesebeck”, “Meteorus caligatus Hal., 9,

Type!, AWS. 29.7.1948”. Lectotype of Perilitus neesii Ruthe, 1862, in BM,

London: 9, “Type H.T.”, “B.M. type Hym., 3.c.758”, “em. Type. Hym. Meteorus

neesii Ruthe, 1862”, ‘‘P. Neesii m.”, “P. neesii Rut.”, “59.101 Germany”, “Ruthe

Coll. 59.101”. Holotype of Meteorus sibiricus Fahringer, 1930, in NR, Stockholm:

o, “Kamtschatka, Malaise”, “1870”, “Meteorus 9 caligatus Hal. var. sibiricus m.”

(in Fahringer’s handwriting), “405, 77°, “Riksmuseum Stockholm”. Holotype of

Dyscoletes alaskensis Ashmead, 1902, in USNM, Washington: &, “Popoff Island,

Alaska, July 11”, “99”, “Harriman Expedition ’99, T. Kincaid, Collector”, “5,

Type, No. 5703, U.S.N.M.”, “Dyscoletes alaskensis Ashm. &” (in Ashmead’s

handwriting). These three holotypes were examined and proved to be rather

typical specimens of caligatus.

Additional specimens examined: 26 9 and 18 G. From the Palaearctic region:

Finland (Sääksmäki, Kivirikko; Kenru), Denmark (no locality), USSR (Irkutskaja

obl., S. Siberia), Japan (Mt. Arakura, 1300 m; Kamitakai, 800 m, both Nagano),

and Italy (Campi, Riva s. Garda, 1200 m) (CNC, HC, WHC, RMNH, UZM). From

the Nearctic region: Alaska (Haines), North West Territories (Norman Walls),

British Columbia (Woodfibre; Gison’s Landing; Triumph Bay; Coquitlan L.;

Seymour Cr.; Squamish; Canyon Cr.; Harrison L.; Hixon; Howe Sound), Alberta

(E. Jaspar Gate; Clearwater), Ontario (Stittsville), Quebec (Mt. Lyall), and

Newfoundland (St. Georges) (CNC, USNM, RMNM).

Variation: Length of fore wing 4.6—5.9 mm; antennal segments 35—37; length

of ovipositor sheath 0.19—0.28 times fore wing; length of malar space 0.4 times

basal width of mandible; length of hind femur 5.2—6.1 times its width; length of

Ist tergite 1.7—1.8 times its apical width; length of vein 1-M of hind wing 0.6—1.0

times vein cu-a of hind wing; cocoon whitish.

Known hosts of examined specimens belong all to the genus Eupithecia

(Geometridae, Lepidoptera): E. luteata Packard, E. palpata Packard, E. ?usurpata

Pears., E. satyrata (Hübner), E. filmata Pears., and E. indigata (Hübner).

Zele levis (Muesebeck) comb. nov.

(figs. 766-776)

Muesebeck, 1923, Proc. U.S. natn. Mus. 63: 11 (as Meteorus).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 76.

Holotype, 9, length of body 4.5, of fore wing 5.0 mm.

Head. — Antennal segments 29, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 5.0 and 4.2 times their width, respectively, length of both

penultimate segments 1.7 times their width (fig. 767); length of maxillary palp 1.3

times height of head; dorsal length of eye 2.1 times temple; temple roundly

narrowed posteriad (fig.770); POL : @ ocellus : OOL = 14: 9: 8; frons almost flat,

366 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

smooth; vertex weakly convex, smooth; face rather flat, somewhat weakly

punctulate; clypeus convex, almost smooth (fig. 773); length of malar space 0.4

times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

densely punctulate, with some crenulae medially and some striae apically (fig.

766); epicnemial area largely smooth, postero-dorsally punctate-rugulose;

precoxal suture smooth, except for some short crenulae medially (fig. 766);

metapleural flange rather large, lamelliform; metapleuron largely smooth,

ventrally punctulate; notauli indistinctly developed dorsally, almost smooth (but in

other specimens finely rugose); mesoscutal lobes faintly punctulate; scutellum

rather convex and punctulate; surface of propodeum finely rugose, only anteriorly

smooth, anterior transverse carina well developed, and posteriorly with a short

medial carina (fig. 775); posterior part of propodeum not separated from antero-

dorsal part of propodeum (fig. 766).

Wings. — Fore wing: r: 3-SR:SRI = 6: 11 : 64; cu-a postfurcal; 1-CU1: 2-CUI

= 3:32; 2-SR : 3-SR : r-m = 16: 11: 10. Hind wing: r present posteriorly (fig. 768);

length of 1-M 1.1 times cu-a.

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

6.4, 12.5, and 9.0 times their width, respectively.

Metasoma. — Length of Ist tergite 2.0 times its apical width, its surface striate

and basally rugose (fig. 775); dorsal carinae of Ist tergite weakly developed in front

of dorsope; dorsope and laterope deep and large (fig. 766, 775); 2nd tergite mainly

bare and smooth; length of ovipositor sheath 0.19 times fore wing.

Colour. — Brownish-yellow; palpi and base of hind tibia, whitish; eyes greenish

iridescent.

Holotype in CU, Ithaca: “Jemez Springs, IX.6-13, N.M., John Woodgate”,

““Meteorus levis Mues., Type, Det. Mues.”, “Holotype Cornell U., No. 616.1”.

Additional specimens of levis examined: 6 Q and 1 &. From Mexico (Dgo., 3 mi.

E. El Salto, 8500 ft; id., 10 mi. W. El Salto, 9000 ft; Chis., 9600 ft, Zontehuitz, nr. S.

Christ.), California (Berkeley), and Wyoming (5 mi. W. New Castle, 4200 ft).

Variation: Length of fore wing 5.0—5.9 mm; antennal segments 29—33; length of

hind femur 6.3—6.4 times its width; length of vein 1-M of hind wing 0.8—1.1 times

vein cu-a of hind wing; length of Iste tergite 1.7—2.2 times its apical width; length

of ovipositor sheath 0.19 times fore wing; & from Mexico has pterostigma,

metasoma basally and apically, propodeum, hind tibia (except its base) and hind

tarsus, more or less dark brown (CNC, USNM, RMNH).

which are intermediate between /evis and caligatus. The females have the shape of

the Ist tergite and face of levis, and the hind leg and metasoma yellowish brownish

(but Ist tergite infuscated). Head and mesosoma are mainly dark brown, the

pterostigma is rather infuscated and the precoxal suture is narrowly and rather

irregularly sculptured as in caligatus. The male is completely melanistic, with also

the legs more or less infuscated. Further collecting is needed to make a decision

about the synonymy of /evis with caligatus.

VAN ACHTERBERG: Revision Zelinae auct. 367

Zele punctatus spec. nov.

(figs. 777— 784, 813)

Holotype, 9, length of body 8.6, of fore wing 8.4 mm.

Head. — Antenna absent except for scapus and pedicellus, 3rd segment of

allotype 1.2 times 4th segment, and length of 3rd and 4th segment 3.9 and 3.2 times

their width, respectively; maxillary palp subequal to height of head; dorsal length

of eye 2.2 times temple; temple punctulate and roundly narrowed posteriad (fig.

779); POL : @ ocellus : OOL = 9: 6: 1; frons smooth, concave behind antennal

sockets; vertex rather flat, punctate (fig. 779); face rather flat, densely punctate

(fig. 781); clypeus strongly convex, coarsely punctate; length of malar space 0.3

times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely reticulate-rugose, dorsally punctate and antero-medially crenulate (fig.

778); epicnemial area rugose-reticulate; precoxal suture crenulate-rugose dorsally

(only rather smooth posteriorly) and coarsely reticulate-punctate ventrally (fig.

778), its surroundings punctate; metapleural flange large, lamelliform and rounded

apically; metapleuron coarsely reticulate; notauli crenulate (fig. 782); mesoscutal

lobes densely punctate (fig. 782); scutellum with rounded tubercle, punctate (fig.

778, 780); surface of propodeum coarsely reticulate, its medial carina rather

weakly developed and without an areola; posterior part of propodeum rather

separated from antero-dorsal part of propodeum (fig. 778).

Wings. — Fore wing: r: 3-SR: SRI = 10: 14: 89; cu-a postfurcal; 1-CUI : 2-

CUI = 2: 17; 2-SR : 3-SR: r-m = 10: 7: 8. Hind wing: r faintly developed; length

of 1-M 0.8 times cu-a.

Legs. — Hind coxa densely and coarsely punctate; length of femur, tibia and

basitarsus of hind leg 5.6, 12.1 and 12.0 times their width.

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface smooth

anteriorly, posterior half rugose (fig. 783); dorsal carinae of Ist tergite absent;

laterope absent (fig. 778); dorsope deep, medium-sized (fig. 783); 2nd tergite

mainly bare and smooth; length of ovipositor sheath 0.40 times fore wing.

Colour. — Brownish-yellow; hind tarsus (except the yellowish telotarsus) white;

apical three-quarters of hind tibia blackish setose; ovipositor sheath dark brown;

wing membrane hyaline; palpi somewhat infuscated.

Holotype in IML, Tucumän: “R. A. Tucuman, Aconguya, XI. (1)946, Coll. R.

Golbach”, “Inst. M. Lillo”, ‘“‘Zemiotes sp., Det. Muesebeck”. Paratypes: 3

specimens, | specimen without metasoma, topotypic with holotype (IML); 1 G

(allotype, IML), “R. A. Tucuman, Dpto. Tafi, 18.X11.50, Coll. Golbach”; 1 &

(RMNH), topotypic with allotype.

Note. The protuberant scutellum indicates a relationship with the South

Nearctic Zele picinervis spec. nov., but punctatus is more coarsely sculptured, has

the wing membrane hyaline, and the ovipositor is longer.

368 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Zele tuberculifer spec. nov.

(figs. 785— 795, 812)

Holotype, 3, length of body 9.0, of fore wing 7.8 mm.

Head. — Antennal segments 41, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 4.2 and 3.6 times their width, respectively, length of both

penultimate segments 1.8 and 2.2 times their width; length of maxillary palp 1.1

times height of head; dorsal length of eye 2.2 times temple; temple roundly

narrowed posteriad, punctulate (fig. 789); POL : & ocellus : OOL = 19: 11:6;

frons deeply concave behind antennal sockets, mainly smooth; vertex punctate,

face rather flat, densely reticulate-rugose (fig. 793); clypeus strongly convex,

reticulate-rugose; length of malar space 0.5 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

densely punctate, medially coarsely crenulate and ventrally reticulate-rugose (fig.

785); epicnemial area anteriorly rather smooth, posteriorly rugose; precoxal

suture dorsally narrowly crenulate, medially and ventrally rather coarsely

punctate (fig. 785); metapleural flange large, wide, truncate and lamelliform

apically; metapleuron coarsely rugose-reticulate; notauli remotely and widely

crenulate (fig. 790); mesoscutal lobes densely punctulate; scutellum with a sharp

tubercle and punctate (figs. 785, 790, 795); surface of propodeum coarsely

reticulate, with a medial carina; posterior part of propodeum separated from

antero-dorsal part of propodeum (fig. 785).

Wings. — Fore wing: r : 3-SR : SRI = 13: 16: 94; cu-a postfurcal; 1-CUI : 2-

CUI = 1: 16; 2-SR : 3-SR : r-m = 9: 8: 7. Hind wing: r present as a brownish

stripe (fig. 788); length of 1-M 0.9 times cu-a.

Legs. — Hind coxa densely and coarsely punctate (fig. 785); length of femur,

tibia and basitarsus of hind leg 6.1, 11.6 and 10.0 times their width, respectively.

Metasoma. — Length of Ist tergite 2.8 times its apical width, its surface smooth

anteriorly, but posteriorly (behind the spiracles) rugulose (fig. 794); dorsal carinae

of Ist tergite absent; laterope absent; dorsope almost absent (fig. 799); 2nd tergite

mainly bare and smooth.

Colour. — Reddish-brown; pterostigma, wing veins, antenna (but scapus and

antenna medially more brownish), and posterior two-thirds of hind tibia, mainly

dark brown or blackish; hind tarsus (except telotarsus) yellowish-white; wing

membrane light brownish.

Holotype in RMNH, Leiden: “Museum Leiden, North Panama, 1050 m,

Fortuna, Chiriqui, 8°44’, 82°15’W, 19.X.1976, H. Wolda, at light”.

Zele niveitarsis (Cresson) comb. nov.

(figs. 796— 807)

Cresson, 1872, Can. Ent. 4: 81 (as Perilitus).

Shestakov, 1940, Ark. Zool. 32A: 16 (Meteorus peronatus), Syn. nov.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 82, 86.

Capek, 1970, Can. Ent. 102: 848.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 222.

Mason, 1973, Proc. ent. Soc. Wash. 75: 214.

VAN ACHTERBERG: Revision Zelinae auct. 369

Zele niveitarsis f. niveitarsis (Cresson)

(figs. 796—807)

Lectotype, 9, length of body 7.5, of fore wing 6.7 mm.

Head. — Antennal segments 40, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 3.6 and 3.4 times their width, respectively, length of both

penultimate segments 2.0 times their width; length of maxillary palp 1.2 times

height of head; dorsal length of eye 3.0 times temple; temple rather directly

narrowed posteriad (fig. 802); POL : @ ocellus : OOL = 8: 6: 2; frons concave,

with some striae anteriorly; vertex rather flat, punctulate; face flat, finely and

densely punctate (fig. 803); clypeus convex and remotely punctate; length of malar

space 0.2 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

largely reticulate-rugose, with some crenulae antero-medially, and dorsally

smooth (fig. 796); epicnemial area largely smooth, anteriorly crenulate; precoxal

suture densely punctate, dorsally narrowly crenulate (fig. 796); metapleural flange

large, rounded and narrowly lamelliform apically (fig. 796); metapleuron coarsely

reticulate; notauli finely crenulate (fig. 807); mesoscutal lobes finely and densely

punctulate; scutellum convex, finely punctulate; surface of propodeum with a long

medial carina anteriorly, and rest of surface remotely and coarsely reticulate and

areolate (fig. 801); posterior part of propodeum separated from antero-dorsal part

of propodeum (fig. 796).

Wings. — Fore wing: r: 3-SR : SRI = 4: 11:45; cu-a antefurcal; 2-M + CUI:

1+2-CUI = 1: 18; 2-SR : 3-SR : r-m = 13: 11: 7. Hind wing: r present, weakly

pigmented (fig. 798); length of 1-M 0.7 times cu-a.

Legs. — Hind coxa weakly punctate; length of femur, tibia and basitarsus of

hind leg 5.1, 11.8, and 12.2 times their width, respectively.

Metasoma. — Length of Ist tergite 2.1 times its apical width, its surface

transversely rugose basally, more irregularly and remotely rugose posteriorly (fig.

801); dorsal carinae present in front of laterope; laterope and dorsope deep and

large (fig. 796, 801); 2nd tergite only medially setose, smooth; length of ovipositor

sheath 0.43 times fore wing.

Colour. — Brownish-yellow; all tibiae basally and entire tarsi, white or nearly

so; telotarsi, hind tibia medially and apically, and antenna, somewhat darkened;

pterostigma and tegulae, light yellowish; stemmaticum and ovipositor sheath, dark

brown; tip of ovipositor sheath indistinctly yellowish; wings hyaline; eyes greenish

iridescent.

Lectotype in ANSP, Philadelphia: “Mass.”, “Type No. 1766”, “Perilitus

niveitarsis Cr./Perilitus albitarsis Cresson’’. Paralectotypes: 1 9 and 3 &, topotypic,

not examined. Additional specimens examined (15 9 and 8 g) from the Nearctic

region: Ontario (Blackbury; Ft. Francis), Quebec (Quebec; Tenaga); Nova Scotia

(Smith’s Cove), Massachusetts (Williamstown; Sterking), New Jersey

(Moorestown), Main (Bar Harbor; Augusto; Mt. Desert), New York (Orient;

Riverhead), Kansas (Riley Co.), Wisconsin (Gibson Lake, Polk Co.), and

Connecticut (Stafford Springs) (USNM, CNC, RMNH).

370 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Variation: Dorsal length of eye of @ 2.4—3.2 times temple (1.8—2.3 times in &);

length of ovipositor sheath 0.42—0.60 times fore wing, longer than 1.5 times Ist

tergite; length of vein 1-M of hind wing 0.7—0.8 times vein cu-a; sometimes head

posteriorly, stemmaticum and mesosoma anteriorly more or less dark brown and

hind tibia blackish apically; hind tibia of g and antenna mainly dark brown or

yellowish brown; pterostigma light brown; vein cu-a of fore wing antefurcal or

interstitial. The dense silken cocoon is whitish, spindle-shaped, in a darker and

larger lepidopterous cocoon.

Known hosts of examined specimens belong all to the Pyralidae (Lepidoptera):

Salebria virgatella (Clements) on Robinia, S. contatella Grote, Acrobasis

rubrifasciella Packard on Alnus; A. betulella Hulst, A. ostryella (?), A. sylviella (?), A.

comptomiella Hulst, and Meroptera pravella (Grote).

Zele niveitarsis f. peronatus (Shestakov)

Holotype, 9, length of fore wing 5.9 mm; dorsal length of eye 2.4 times temple;

vein cu-a of fore wing antefurcal; 2-M +CUI : 1+2-CUl = 5: 66; vein 1-M of hind

wing 0.4 times vein cu-a; antennal segments 39; length of ovipositor sheath 0.46

times fore wing and 1.7 times length of Ist tergite; pterostigma and body, dark

brown; Ist tergite basally, mesopleuron dorsally, face, clypeus, and basal half of

antenna, brownish; fore and middle legs, tegulae, and hind trochanters, yellowish,

apical three-quarters of hind tibia dark brown, its basal quarter yellowish white;

hind coxa and its femur, brown; hind tarsus mainly white, but basally dark brown.

Holotype in NR, Stockholm: ‘“‘Vladivost., Sedanka/10/8, 1930, Malaise”,

‘“‘Meteorus peronatus sp. n. typ., det. Shestakov”, ‘‘Holotype of peronatus

Shestakov, det. T. Huddleston, 1976”, ‘406, 77°, “Riksmuseum Stockholm”.

Additional specimens examined: | Q from Sumatra (N. Sumatra, Bivouac 3, Mt.

Bandahara, ca. 1810 m, 3°45’N, 95°45’E, 10—16.VII.1975, J. Krikken, no. 25

(RMNH, 1 9 (ex E. pylonitis (?)) and 1 g (ex D. abietella (?)) from Lower Topa (?)

(both USNM). Variation: Length of fore wing 5.9—8.1 mm; antennal segments

39—43; length of Ist tergite 2.3—2.8 times its apical width; length of ovipositor

sheath 0.46—0.50 times fore wing; dorsal length of eye 2.4—3.1 times temple (1.8

times in &).

Zele chlorophthalmus (Spinola) comb. nov.

(figs. 814—824)

Spinola, 1808, Insect. Liguriae 2: 133, 134 (as Bracon).

Nees, (1811) 1812, Mag. Ges. nat. Fr. Berl. 5: 21 (Bracon chrysophthalmus). Syn. nov.

Thunberg, 1822, Mem. Acad. sci. St. Petersbourg 8: 263 (/chneumon nudator). Syn. nov.

Costa, 1884, Rc. Accad. Sci. fis. mat., Napoli 22: 171 (Meteorus splendens). Syn. nov.

Thomson, 1895, Opusc. ent. 20: 2150 (Meteorus (Zemiotes) nigricollis). Syn. nov.

Wagner, 1928, Ver. naturw. Unterh. Hamb. 20: 7.

Cavro, 1954, Suppl. Bull. Soc. ent. N. Fr. 75: 109.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 58, 59, 81, 82.

Shenefelt, 1970, id. 5(2): 222.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258, 268— 270.

VAN ACHTERBERG: Revision Zelinae auct. 371

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 223.

Capek, 1972, Ent. Problemy 10: 134, 136.

Mason, 1973, Proc. ent. Soc. Wash. 75: 213—215.

Tobias, 1976, Opr. Fauna SSSR 110: 113, fig. 33: 14, 15.

Neotype, 9, length of body 7.4, of fore wing 6.4 mm.

Head. — Antennal segments 39, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 4.2 and 4.0 times their width, respectively, length of both

penultimate segments 2.0 and 2.3 times their width (fig. 819); length of maxillary

palp 1.2 times height of head; temple weakly roundly narrowed posteriad (fig.

817); dorsal length of eye 1.3 times temple (fig. 817); POL : g ocellus : OOL = 12:

9 : 6; frons mainly smooth, weakly concave; vertex convex, slightly punctulate;

face rather flat, punctulate; clypeus convex and punctate; length of malar space

0.1 times basal width of mandible, eyes almost touching mandibular condylus.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

punctulate, medially and ventrally reticulate-rugose (fig. 815); epicnemial area

crenulate anteriorly and posteriorly punctate-rugose; precoxal suture widely

reticulate-punctate, dorsally indistinctly crenulate (fig. 815); metapleural flange

rather large, lamelliform apically; metapleuron largely rugose-reticulate, dorsally

weakly sculptured notauli rather widely crenulate (fig. 818); mesoscutal lobes

punctulate; scutellum rather convex, punctulate; surface of propodeum rather

coarsely reticulate-rugose, only smooth anteriorly (fig. 824), with a medial carina;

posterior part of propodeum not separated from antero-dorsal part (fig. 815).

Wings. — Fore wing: r: 3-SR : SRI = 7: 15: 70; cu-a antefurcal; 2-M+CUl:

1+2-CUI = 3: 52; 2-SR : 3-SR : r-m = 24: 15 : 14. Hind wing: remnant of r

present (fig. 814); length of 1-M 0.7 times cu-a.

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

6.2, 12.5, and 12.0 times their width, respectively.

Metasoma. — Length of Ist tergite 2.4 times its apical width, its surface behind

dorsope rugulose-punctate (fig. 824); dorsal carinae short anteriorly, in front of

dorsope; laterope large and deep (fig. 815); dorsope medium-sized, deep (fig. 824);

2nd tergite mainly bare and smooth; length of ovipositor sheath 0.42 times fore

wing.

Colour. — Brownish-yellow; apical half of antenna and ovipositor sheath

(except the apex) dark brown; eyes greenish iridescent.

Neotype of Bracon chlorophthalmus Spinola to be deposited in the collection of

(Zemiotes) chrysophthalmus (Nees), det. C. v. Achterberg, 1973”.

According to Mr. T. Huddleston, who kindly examined the holotype of Meteorus

splendens Costa, 1884 (9, Museum of Naples), splendens agrees with my

interpretation of chlorophthalmus. Length of fore wing 4.0 mm, length of ovipositor

sheath 0.44 times fore wing and 1.9 times length of Ist tergite, propodeum postero-

dorsally and anterior half of Ist tergite darker than rest of body (blackish

according to Costa). The type bears two labels: ““Decimoputzu 4” and “Meteorus

splendens” (the latter in what is probably Costa’s handwriting). The type-locality is

situated at the South of Sardinia, nr. Cagliari.

372 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Mr. Huddleston also examined the holotype of /chneumon nudator Thunberg,

1822 (g, Thunberg Collection, Uppsala), which proved to have been correctly

synonymized with Bracon chrysophthalmus Nees in the past. The holotype of

Meteorus (Zemiotes) nigricollis Thomson, 1895 (9, ZIL, Lund: ‘“Degeberga”,

“nigricollis m.”, “holotype Meteorus (Zemiotes) nigricollis T., det. T. Huddleston,

1976”, “1977, 40”) agrees well with the neotype of chlorophthalmus. The eyes are

slightly less convex, length of eye 1.3 times temple, POL : @ ocellus : OOL = 16:9

: 9, frons striate anteriorly, length of fore wing 6.4 mm, length of ovipositor sheath

0.50 times fore wing, length of Ist tergite 2.2 times its apical width, mesosoma

mainly dark brown, middle of frons and vertex faintly infuscated.

The type of Bracon chrysophthalmus Nees, 1812, is lost; a neotype is selected

from the Wesmael Collection, because Wesmael is the first revisor of this species.

The neotype of chrysophthalmus (9, KBIN, Brussels: “Coll. Wesmael”, “1743”,

“Perilitus 3 9 chrysophthalmus N. v. Es., det. C. Wesmael”, “type”) agrees well

with the descriptions by Nees and Wesmael (1835: 24—26) and with the neotype of

chlorophthalmus: length of fore wing 6.0 mm, length of ovipositor sheath 0.49 times

fore wing, eyes rather flat, and length of malar space 0.3 times basal width of

mandible.

Mariehamn), Sweden (Degeberga), Denmark (Bornholm; Nordsjglland; Sj.

Jungshoved; Ebsjorg; Allnye; Adserbo; Stube in Sondbg), England (Dartmoor,

SD, Lustleigh; Hants., Hawkley Warren; Whetstone, Hertfordshire; South-

ampton; Berks., Windsor Forest; SR, Claygate; H., Bricket Wood), Netherlands

(Oostkapelle; Melissant; Oostvoorne; nr. Breda; Heerde (G.); Putten (G.);

Meijendel; Castelre; Udenhout; Savelsbos; Tilburg; Assel; Crailo; Venlo; Bergen

op Zoom; Gliphoeve; Tegelen; Zundert, De Krochten; Meinweg, Melick &

Herkenbosch; Texel, landside dunes nr. Fonteinsnol; St. Pietersberg (ex Crataegus

stem); Middelharnis; Naardermeer, Ouddorp; Wijster; Veenhuizen; Herpen;

Voorburg; Berghem), West Germany (Annatal nr. Honnef, Siebengeb.; nr.

Eichstadt); Czechoslovakia (B. Stiavnica), Austria (Piesting; Leitha Geb.;

Donnerskirchen; Innsbruck), Poland (Gdansk), USSR (Tsav, Armenia; MSSR,

Benderespsjij; Moldavia, C. Choresji, garden; Jonava, Dukstas), Bulgaria

(Rodopi, Nicoloro); France (Baton; Lille Nord; Tours; Paris), and Spain

(Santander, Potes; 25 km SW Salou) (RMNH, LH, CVR, BM, CNC, UZM, ITZ,

ZMH, ZMB, HC, USNM, ZIL, IZP). Variation: Length of fore wing 4.0—6.8 mm;

antennal segments 36—42; dorsal length of eye of 9 1.3—2.1 times temple

(1.2—1.6 times in £); length of Ist tergite 2.1—2.4 times its apical width; length of

ovipositor sheath 0.41—0.53 times fore wing; vein cu-a of fore wing antefurcal or

interstitial; body colour variable, varies from completely yellowish or brownish to

mainly dark brown of blackish. Cocoon whitish.

Notes. Known hosts of examined specimens belong to the Pyralidae

(Lepidoptera): Acrobasis consociella (Hübner) and R. formosa (?= Salebria formosa

Haworth) and to the Zygaenidae (Lepidoptera): Zygaena lonicerae (Esp.).

The application of the name chlorophthalmus in the Braconidae has led to a lot of

confusion and misinterpretation. The name was first used by Spinola in 1808, but

VAN ACHTERBERG: Revision Zelinae auct. 373

the type is lost according to information kindly provided by Prof. Dr. C. Vidano

and Dr. P. Passerin d’Entreves. Luckely the original description by Spinola is

comparatively clear: the petiolate metasoma, the long ovipositor (about as long as

metasoma), vein m-cu of fore wing about interstitial and the size (ca. 7 mm)

indicate its synonymy with Zele chrysophthalmus (Nees). Nees (1812: 21) clearly

made a mistake by attributing the name chrysophthalmus to Spinola, when no such

name was published by Spinola. Probably it was a miswriting of chlorophthalmus

but actually he created a new binomen. In 1834 (p. 35) he exacerbated the

situation by retaining his chrysophthalmus and misinterpreting chlorophthalmus for a

species of Homolobus which differs by e.g., a sessile metasoma. Finally the name

chlorophthalmus was misapplied by Haliday for what is now known as Homolobus

flagitator (Curtis).

Zele picinervis spec. nov.

(figs. 811, 825—833)

Holotype, 9, length of body and of fore wing both 8.9 mm.

Head. — Antennal segments 42, 3rd segment 1.2 times 4th segment, length of

3rd and 4th segments 3.7 and 3.1 times their width, respectively, length of both

penultimate segments 2.0 and 2.3 times their width (fig. 832); length of maxillary

palp 1.3 times height of head; dorsal length of eye 2.7 times temple; temple

roundly narrowed posteriad (fig. 831); POL : @ ocellus : OOL = 9: 6: 4; frons

weakly concave, mainly smooth; vertex weakly convex, punctulate; face flat,

indistinctly punctulate; clypeus strongly convex (fig. 825), punctulate; length of

malar space 0.1 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum

densely rugulose ventrally and posteriorly, crenulate medially, punctulate dorsally

(fig. 825); epicnemial area slightly punctulate-rugulose; precoxal suture widely

rugose-punctate, only dorsally narrowly crenulate (fig. 825); metapleural flange

large, lamelliform (fig. 825); metapleuron coarsely rugose-reticulate; notauli

anteriorly narrowly crenulate, posteriorly widely crenulate-rugose (fig. 811);

mesoscutal lobes densely punctulate; scutellum rather strongly convex (fig. 811,

825), without tubercle, mainly smooth; surface of propodeum coarsely carinate,

anteriorly with an almost straight transverse carina, medially with a long

longitudinal carina, enclosed areas weakly rugose; posterior part of propodeum

not separated from antero-dorsal part (fig. 825).

Wings. — Fore wing: r : 3-SR : SRI = 6: 15: Sl; cu-a postfurcal, somewhat

inclivous (fig. 827); 1-CUI : 2-CUI = 1 : 22; 2-SR : 3-SR : r-m = 13: 15: 9. Hind

wing: r mainly absent; Ir-m weakly curved (fig. 827); length of 1-M 0.8 times cu-a.

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

5.9, 11.8, and 9.0 times their width, respectively.

Metasoma. — Length of Ist tergite 2.6 times its apical width, its surface smooth,

except for some rugulosity behind spiracles (fig. 833); dorsal carinae of Ist tergite

absent; laterope and dorsope deep and large (fig. 825, 833); 2nd tergite evenly and

densely setose, bare; length of ovipositor sheath 0.24 times fore wing.

374 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Colour. — Brownish-yellow; all trochanters and trochantelli (but apex of hind

trochantellus dark brown), base of fore and middle tibiae and tarsi, apex of

parastigma and pterostigma, basal quarter of pterostigma, basal two-thirds of hind

tibia, its spurs and tarsus, white or nearly so; pterostigma medially, veins I-M, cu-

a, CUI, 1-SR+M, 2-SR and r of fore wing, and apical third of hind tibia, dark

brown; palpi and wing membrane, slightly infuscated, but surroundings of veins 1-

M, 1-CUI and r of fore wing dark brown pigmented (fig. 827); ovipositor sheath

dark brown, but apex narrowly yellowish.

Holotype in CNC, Ottawa: “Ramsey Cyn., 5000’(ft), 15 mi. S. Sierra Vista,

Huachuca Mts., Ariz., Sternitzky, VIII.1968’”, ““Zemiotes n. sp., W.R.M. Mason

72”. Paratypes: (19 and 1¢); 1 g (allotype, CNC), “Mex., Dgo., 24 mi. W. La

Kritsch”.

Variation: Length of fore wing 8.9—11.4 mm; length of ovipositor sheath

0.19—0.24 times fore wing: length of vein 1-M of hind wing 0.8—0.9 times cu-a;

length of Ist tergite 2.5—3.2 times its apical width; length of malar space 0.1(9)or

0.2 (4) times its apical width.

Zele crassifemur (Muesebeck) comb. nov.

(figs. 808, 809, 834— 842)

Muesebeck, 1939, Proc. ent. Soc. Wash. 41: 84 (as Meteorus).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 63.

Mason, 1973, Proc. ent. Soc. Wash. 75: 214.

Head. — Antennal segments 43, 3rd segment subequal to 4th segment, length of

3rd and 4th segments 3.0 and 3.1 times their width, respectively, length of both

penultimate segments 1.7 and 1.8 times their width; length of maxillary palp 1.1

times height of head; dorsal length of eye 1.8 times temple; temple rounded

posteriad, punctulate; dorsal aspect of head rather transverse (fig. 838); POL : 5

ocellus : OOL = 11 : 6 : 7; frons rather flat, rugulose medially and punctate

laterally; vertex rather flat, punctulate; face medially convex, densely punctate,

rather wide (fig. 842); clypeus convex, punctate; length of malar space 0.4 times

basal width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum

crenulate medially, rugulose ventrally and posteriorly; epicnemial area more or

less rugose (fig. 834); precoxal suture densely and narrowly rugose-crenulate

dorsally, widely and densely punctate medially and ventrally; metapleural flange

large, narrowly lamelliform apically; metapleuron almost smooth dorsally, rugose-

reticulate ventrally; notauli distinctly crenulate (fig. 808); mesoscutal lobes weakly

punctulate; scutellum weakly convex, punctulate; surface of propodeum

completely and rather coarsely reticulate, with a long medial carina; posterior part

of propodeum not separated from antero-dorsal part of propodeum (fig. 834).

Wings. — Fore wing: r: 3-SR: SRI = 11 : 21 : 91; cu-a postfurcal in right wing

(fig. 836), but subinterstitial in left wing; 1-CUI : 2-CUI of right wing = 2: 45; 2-

VAN ACHTERBERG: Revision Zelinae auct. 375

SR : 3-SR : r-m = 28: 21 : 21; vein 2-R1 well developed, longer than r. Hind wing: r

shortly developed posteriorly (fig. 836); length of 1-M 0.4 times cu-a.

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

4.2, 11.5, and 7.4 times their width, respectively.

Metasoma. — Length of Ist tergite 2.3 times its apical width, its surface largely

smooth, apical third superficially, longitudinally striate (fig. 839); dorsal carinae of

Ist tergite absent; laterope and dorsope deep and large (fig. 834, 839); 2nd tergite

smooth and mainly bare; length of ovipositor sheath 0.28 times fore wing.

Colour. — Brownish-yellow; ovipositor sheath (except apex) dark brown; apical

half of antenna, hind tibia dorso-apically, somewhat infuscated; all tarsi whitish;

base of hind tibia and tegulae, whitish-yellow; pterostigma yellowish.

Holotype in USNM, Washington: “Wellington Kans(as)”, “E. G. Kelly

Collector”, “Experiment 151539”, “Type 53036 U.S.N.M.”, “ Meteorus crassifemur

Mues., Type, Det. Muesebeck”. Two paratypes were examined: | 9 from Texas

and | 9 from Brookings, S. D., both in USNM. Additional specimens examined:

(4 9) from Illinois (Principia College, Elsah, Jersey Co.), North Carolina

(Highlands), South Dakota (Brookings, light trap), and Texas (Lost Pines Pk.,

Bastrop) (CNC, USNM, UZM).

Variation: Antennal segments 43—44; length of fore wing 8.8—10.1 times fore

wing: length of ovipositor sheath 0.25—0.29 times fore wing; length of hind femur

3.8—4.4 times its width; Ist tergite as well as length of vein 1-M of hind wing, as in

holotype.

Zele gracilis spec. nov.

(figs. 810, 843—852)

Holotype, 9, length of body 10.6, of fore wing 9.7 mm.

Head. — Antennal segments 49, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 4.4 and 4.2 times their width, respectively, length of both

penultimate segments 2.5 and 3.0 times their width; length of maxillary palp 1.5

times height of head; dorsal length of eye 2.8 times temple; temple directly

narrowed posteriad (fig. 850); POL : @ ocellus : OOL = 13: 11 : 4; frons mainly

smooth, concave behind antennal sockets; vertex punctulate, rather flat; face

rather flat, punctulate; clypeus convex, punctulate; length of malar space 0.4 times

basal width of mandible.

Mesosoma. — Length of mesosoma 1.6 times its height; side of pronotum

largely coarsely and densely reticulate-punctate, medially crenulate and dorsally

punctulate (fig. 843); epicnemial area coarsely punctate; precoxal suture reticulate

dorsally, densely ventrally (fig. 843); rest of mesopleuron more or less punctate;

metapleural flange large, lamelliform apically (fig. 843); metapleuron coarsely and

finely reticulate; notauli rather shallow and narrowly crenulate (fig. 849);

mesoscutal lobes densely punctulate; scutellum rather flat, punctulate; surface of

propodeum densely and coarsely rugose-reticulate, only anteriorly narrowly

almost smooth, medial carina only anteriorly present; posterior part of propodeum

not separated from antero-dorsal part (fig. 843).

376 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Wings. — Fore wing: r: 3-SR : SRI = 8: 7: 57; cu-a postfurcal; 1-CUI : 2-CUI

= 1 : 22; 2-SR : 3-SR : r-m = 15: 7: 10; 2-R1 absent. Hind wing: r weakly

developed; length of 1-M 0.8 times cu-a.

Legs. — Hind coxa densely and finely punctate; length of femur, tibia and

basitarsus of hind leg 7.6, 14.4, and 13.4 times their width, respectively. À

Metasoma. — Length of metasoma 4.1 times its apical width, its laterobasal half

smooth (fig. 843) and its surface smooth in front of spiracles, convergently striate

behind spiracles (fig. 852); dorsal carinae of Ist tergite absent; laterope and

dorsope deep and large (fig. 843, 852); 2nd tergite densely setose, shiny and weakly

coriaceous-punctulate (fig. 852); length of ovipositor sheath 0.37 times fore wing.

Colour. — Yellowish-brown; stemmaticum, head medio-posteriorly, mesosoma,

basal half of hind coxa, dark brown; apex of antenna infuscated; wing membrane

light brownish; pterostigma yellowish; palpi, lower half of temples, mandibles,

face, fore leg, tegulae, hind tarsus (except base and apex) and Ist tergite in front of

spiracles, more or less yellowish-white; pronotum anteriorly and posteriorly, and

propodeum ventrally, narrowly brownish.

Holotype in CNC, Ottawa: “Nepal, Ktmd., Pulchauki, 7300'(ft), 7—16.VIII.

1967, Mal. Tr., Can. Exp.”.

Zele albiditarsus Curtis

(figs. 853—876)

Curtis, 1832, Br. Ent. 9: 415—4, figs.

Curtis, 1832, Br. Ent. 9: 415—3, fig. (Zele testaceator). Syn. nov.

Nees, 1834, Hym. Ichn. affin. Mon. 1: 34 (Perilitus albitarsus).

Haliday, 1835, Ent. Mag. 3: 24 (Meteorus albitarsis).

Wesmael, 1835, Nouv. Mem. Acad. Brux. 9: 22 (Perilitus dispar).

Wesmael, 1835, id. 9: 26 (Perilitus deceptor). Syn. nov.

Curtis, 1837, Guide Br. Insects: 118 (Meteorus calcitrator).

Cresson, 1872, Can. Ent. 4: 81 (Perilitus pallitarsis). Syn. nov.

Thomson, 1895, Opusc. ent. 20: 2149 (Meteorus (Zemiotes) rufulus). Syn. nov.

Muesebeck, 1923, Proc. U.S. natn. Mus. 63: 13 (Meteorus maximus). Syn. nov.

Muesebeck, 1923, id. 63: 14 (Meteorus reticulatus). Syn. nov.

Wagner, 1928, Verh. Ver. naturw. Unterh. Hamb. 20: 8.

Fahringer, 1930, Ark. Zool. 21A: 8 (Meteorus romani). Syn. nov.

Fischer, 1957, Opusc. zool. 3: 3 (Meteorus (Zemiotes) separandus). Syn. nov.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 51—89.

Shenefelt, 1970, id. 5(2): 226.

Capek, 1970, Can. Ent. 102(7): 848.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258, 275—277, fig. 6.

Tobias, 1971, Tr. Vsesoyuzn. ent. Obshch. 54: 222—224.

Capek, 1972, Ent. Problemy 10: 133, 138.

Mason, 1973, Proc. ent. Soc. Wash. 75: 214.

Papp, 1973, Acta Mus. Mac. Sc. nat. 14: 3.

Jakimavicius, 1974, Tr. AN Lit. SSR B2(66): 97.

Gauld & Huddleston, 1976, Entomologist’s Gaz. 27: 43, fig. 19.

Van Achterberg, 1976b, Tijdschr. Ent. 119: figs. 107, 111.

Tobias, 1976, Opr. Fauna SSSR 110: 113, figs. 33: 16.

VAN ACHTERBERG: Revision Zelinae auct. 377

Zele albiditarsus f. deceptor (Wesmael) comb. nov.

(figs. 853— 864)

Lectotype, 9, length of body 7.5, of fore wing 6.8 mm.

Head. — Antennal segments 34, but apical segments missing, 3rd segment 1.1

times 4th segment, length of 3rd and 4th segments 3.6 and 3.2 times their width,

respectively; length of maxillary palp 1.5 times height of head; dorsal length of eye

2.2 times temple; temple roundly narrowed posteriad (fig. 863); POL : @ ocellus :

OOL = 10: 6: 2; frons smooth, weakly concave; vertex weakly convex,

punctulate; face rather flat, indistinctly punctulate; clypeus strongly convex,

weakly punctate (figs. 853, 862); length of malar space 0.3 times basal width of

mandible.

Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum

punctate-reticulate ventrally and posteriorly, crenulate medially, and mainly

smooth dorsally (fig. 853); epicnemial area rugose-reticulate dorsally, narrowly

crenulate anteriorly; precoxal suture crenulate dorsally, coarsely punctate-

reticulate ventrally (fig. 853); rest of mesopleuron punctulate; metapleural flange

large, lamelliform apically; metapleuron reticulate; notauli distinctly crenulate

(fig. 861); mesoscutal lobes punctulate; scutellum weakly convex and weakly

punctulate; surface of propodeum remotely and coarsely reticulate, medial carina

irregular, long (fig. 864); posterior surface of propodeum not separated from

antero-dorsal part (fig. 853).

Wings. — Fore wing: r: 3-SR : SRI = 5: 11: 42; cu-a straight, postfurcal; 1-CU1

:2-CUl = 2: 21; 2-SR : 3-SR: r-m = 14: 11 : 8; 2-R1 short (fig. 855). Hind wing: r

mainly absent; length of 1-M 0.6 times cu-a.

Legs. — Hind coxa punctulate; length of femur, tibia and basitarsus of hind leg

7.1, 13.8 and 12.2 times their width, respectively.

Metasoma. — Length of Ist tergite 2.5 times its apical width, its surface smooth

in front of spiracles, longitudinally striate behind spiracles, only medially almost

smooth (fig. 864); dorsal carinae of Ist tergite absent, except for a short basal

remnant; laterope and dorsope large and deep (figs. 853, 864); 2nd tergite mainly

bare and smooth; length of ovipositor sheath 0.27 times fore wing.

Colour. — Brownish-yellow; antenna apically and ovipositor sheath (except its

tip), somewhat darkened.

Lectotype in KBIN, Brussels: “Coll. Wesmael”, “1774”, “Perilitus deceptor

mihi, g 9, det. C. Wesmael”, “Type”. Lectotype here selected, and labelled

accordingly. There are two 9 paralectotypes, both heavily damaged and two,

rather dark Z paralectotypes.

The holotype of Meteorus reticulatus Muesebeck, 1923 (9, USNM, “Mt.

Wash’n”, “58”, “collection Ashmead”, ‘“Meteorus areolatus Ash., 9, type (MS-

name), “Type No. 24967 U.S.N.M.”, ““Meteorus reticulatus Mues., Type”) is a

typical deceptor.

The holotype of Meteorus separandus Fischer, 1957 (8, ZSB, “Holotypus”, “8/

440”, “Staatssamml. München, München, Pusing, 11.6.1884, leg. J. Kriech-

baumer”, ““Meteorus separandus n. sp. det. Fischer, Holotype’’) is only a melanistic

378 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

specimen of albiditarsus, of which the males are usually darker than the females.

Additional specimens examined: 180 9 and 93 Z. From the Nearctic region:

Alaska (Gulkana Gla., from snowfield (!)), Yukon Territory (Dawson, 1100 ft),

British Columbia (Andalis Cr.; Hazelton; St. Croix; 15 mi. Beatton R.; Green R.;

Lakevale, Kettlevalley; Lac le Jeune; Duck Range; Lower Nicola; Kimberley; Mt.

Thornhill, nr. Terrace, 700 ft; Green Inlet; Knouff L.; Jesmand; Field; mi. 103

Cariboe Hwy.; Bostock Cv.), Quebec (Parke Reserve, Kam. Co.; Covey Hill; Sept

Iles: Lac Mondor, Ste Flore; Sinclair Mills; Mara; Spiller Chl.), Ontario (Gogama;

Nakina; Glomcoe; Hearst; Port Arthur; Kapuskasang; Black Sturgeon Lake),

Alberta (Miette Springs Rd.; Poeahontas; Hargwen; Slave L.; Jasper),

Saskatchewan (Prince Albert), New Brunswick (Restigouche Co.), Nova Scotia

(Truemanville, Cumberland Co.), Wisconsin (Illakee, Vitsap Co.; Trout L.), South

Dakota (2 mi. S. Sylvan L., Black Hills), Virginia (Mt. Washington), Oregon

(Milton Freewater; Saddleback Mt., Lincoln Co.), Utah (Aspen Grove Camp, Mt.

Timpanogos, Utah Co., 6800 ft), Idaho (Cronwall), New Hampshire (Hanover;

Durham), and California (Cisco).

From the Palaearctic region: Finland (Tvarmine; Perna; Norv. b. Skiervö;

Lemland; Carelia or., Soutjärva), Sweden (Lapland; Höör, Skane), Denmark

(Ems; Sonderburg; Allerup; Sondbg; Dyrhavn), West Germany (Fuss Hohen

Acht, Eiffel; Jungfernhardt, Siebengeb.; Vorgebirge, Kottenforst; Fuss Lohrberg

im Siebengeb.; Mainz; Ennert nr. Beuel; Tiergarten N. Blankenheim, Eifel;

München, Pusing; Bramwald, Nd. Sachsen; Hedemünden; Wiesen, Spessart;

Tremalzo, Voralpen, 1300 m; Hochstadt, Obb.; Bergen, 600 m, Bayr. Alpen;

Grainbach, Obbay., 700 m; Reither Alm, 1100 m; Heidelberg; Ziegenhagen,

Hessen; Eberschütz, id.), England (Epping Forest, Essex; Spratton, Northants. ;

Isle of Rhum, Kimloch; Oxon; Goring Heath; Sherwood, NM), Scotland

(Aviemore; Invern., Tollochmoor; NS., Inchnadamph), Ireland (Killykeen),

Netherlands (Oostkapelle; Putten (G.); Woold; Overveen; Bentveld; Lienden;

Velzen; Wijster; Rijs (Fr.); Meinweg, nr. Herkenbosch; Nunspeet; Meijendel,

dunes; Rockanje, Stekelhoekduin; Oostvoorne; Drijber; Haamstede; Wapenveld;

Rijsbergen; Schayk; Waarder; Wageningen, Wageningse berg; Assel; Crailo;

Naardermeer; Muiderberg; Otterlo; Venlo; Heerde (G.); Asperen; Loenen;

Melissant), Belgium (Lk., Stavelot), France (La Bégude de Mazenc, Drome;

Saumane de Vaucluse), Switzerland (Wallis, Unterbach, 1500 m), Austria

(Hofgastein, Schossalm, 2000—2150 m; Semmeringgeb., Reichenau dist,

Salzburg, Judenbergalm; Rainberg, Steinbruch, Salzburg; Söllheim, Autobahn nr.

Salzburg; Innsbruck; Gneisermoor, Salzburg; Aschbach, 1400 m, Tirol),

Czechoslovakia (Prachatitz, Bohemia); USSR (Sveneioniy, Lit. SSR; Kazachstan,

Karagandinsk), Bulgaria (Rodopi, Velinograd), and Italy (Campi, Riva s. Garda,

800 m; id., 220 m) (CAS, ZMH, UZM, RMNH, ITZ, CNC, ZC, ZSB, HC, ZIL,

IZP).

Variation: Length of fore wing 4.5—8.3 mm; antennal segments 32—43; vein cu-

a of fore wing postfurcal or seldom interstitial; length of hind femur 5.1—6.4 times

its width; length of Ist tergite 2.1—2.5 times its apical width; length of ovipositor

sheath 0.26—0.32 times fore wing; length of vein 1-M of hind wing 0.5—0.8 times

VAN ACHTERBERG: Revision Zelinae auct. 379

vein cu-a; hind tarsus yellowish, brownish, or infuscated. Silken cocoon rather

tough, dense and whitish or brownish.

Known hosts of examined specimens belong to the Geometridae (Lepidoptera):

Semiothisa sexmaculata (Packard), S. granitata (Guenée), S. unipunctaria perplexa

(McDunnough), Rheumaptera hastata (L.), Enypia moilietti (2), Eupithecia

pseudotsugata MacK., Nyctobia nigroangulata Strecker, and Hydriomena furcata

(Thunberg); to the Noctuidae (Anarta myrtilli (L.)), Saturniidae (Antheraea

polyphemus Cramer on Quercus macrocarpa), and Tortricidae (Acleris variana

(Fernald)). The latter host especially needs to be confirmed.

Zele albiditarsus f. pallitarsis (Cresson) comb. nov.

(figs. 865—876)

Holotype, &, length of body 6.6, of fore wing 5.6 mm.

Head. — Antennal segments 43; length of maxillary palp 1.4 times height of

head; dorsal length of eye 1.3 times temple; temple roundly narrowed posteriad

(fig. 866); POL : @ ocellus : OOL = 9: 5: 7; length of malar space 0.7 times basal

width of mandible.

Mesosoma. — Length of mesosoma 1.4 times its height; sculpture of mesosoma

as in forma deceptor (fig. 865).

Wings. — Fore wing: r: 3-SR : SRI = 4: 10: 43; 1-CUI : 2-CUI = 1 : 22; 2-SR:

3-SR : r-m = 14: 10: 9. Hind wing: short remnant of r present; length of 1-M 0.95

times cu-a.

Legs. — Length of femur, tibia and basitarsus of hind leg 5.9, 12.2, and 8.8 times

their width, respectively.

Metasoma. — Length of Ist tergite 2.4 times its apical width, its surface smooth

in front of spiracles, superficially punctate-rugose posteriorly (fig. 876); 2nd tergite

mainly bare and smooth.

Colour. — Brownish-yellow; pterostigma brown; hind tibia infuscated apically;

palpi, two basal segments of hind tarsus, whitish, rest of hind tarsus mainly

yellowish.

Holotype in ANSP, Philadelphia: “N.J.”?, “Type No. 1767”, “Perilitus pallitarsis

Cress.”. Essentially as deceptor, but vein 1-M longer and hind tarsus more whitish

basally. Additional specimens examined: 24 9 and 21 g. From the Nearctic

region: Yukon Territory (Dawson), Newfoundland (South Branch), Michigan

(Ann Arbor; Lake Odessa), Ontario (Rondeau Park; St. Pelee; Florence; Aylmer

West), Quebec (Hull), Alberta (14 km N. Sundre), New York (Orient, L.I.; Ithaca),

Oregon (Bedford; Hinckley), Virginia (Rosslyn), Pennsylvania (Germantown;

Grantville, Dauphin Co.), Maryland (Cabin John), New Jersey (Moorestown),

Maine (Bar Harbor), and Mexico (Dgo., 30 mi. W. La Cuidad, 6500 ft; Sin., 4.5 mi.

W. EI Palmito, 6500 ft), and from the Palaearctic region: Denmark (Sondbg.)

(CNC, UZM, USNM, RMNH). One specimen was reared from Semiothisa

sexmaculata (Packard) (Geometridae).

Variation: Length of fore wing 4.6—6.2 mm; antennal segments 39—43; length

of vein 1-M of hind wing 0.9—1.3 times vein cu-a of hind wing: length of Ist tergite

380 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

2.2—2.4 times its apical width; length of ovipositor sheath 0.24—0.32 times fore

wing; length of hind femur 5.9—6.6 times its width; hind tibia, pterostigma and

antenna of male usually infuscated or dark brown.

Zele albiditarsus f. albiditarsus Curtis comb. nov.

Holotype, & (from Regent's Park, England (NMV)) not examined, but the

original description, especially the remark on the coloration and the figures given

by Curtis are clear enough to enable a correct identification of the species. The

albiditarsus; the following redescription is based on this lectotype of testaceator.

Length of body 8.8, of fore wing 8.9 mm.

Head. — Antennal segments 46, 3rd segment 1.1 times 4th segment, length of

3rd and 4th segments 3.4 and 3.0 times their width, respectively, length of both

penultimate segments 1.8 and 2.0 times their width; length of maxillary palp

subequal to height of head; dorsal length of eye 1.7 times temple; temple roundly

narrowed posteriad; POL : @ ocellus : OOL = 13: 10: 7; frons concave, somewhat

rugose near antennal sockets; vertex smooth and rather convex; length of malar

space 0.3 times basal width of mandible.

Mesosoma. — Length of mesosoma 1.2 times its height; sculpture of mesosoma

as in deceptor.

Wings. — Fore wing: r: 3-SR : SRI = 10: 26: 124; cu-a interstitial; 2-SR : 3-SR:

r-m = 31:26:21; 2-RI short. Hind wing: r present; 1-M much shorter than cu-a.

Legs. — Length of femur, tibia and basitarsus of hind leg 6.1, 11.0, and 9.3 times

their width, respectively.

Metasoma. — Length of Ist tergite 2.0 times its apical width, anterior half very

finely rugulose, basal half almost smooth; 2nd tergite mainly bare and smooth;

length of ovipositor sheath 0.25 times fore wing.

Colour. — Yellowish-brown; surroundings of ocelli and ovipositor sheath

(except tip), dark brown; pterostigma yellowish; hind tarsus yellowish-white.

Lectotype in NMV, Melbourne: “Coomb, 25 July” (old handwritten label, refers

3.albiditarsus 3, G. Nixon, det. 1948”. This specimen is selected as lectotype,

because Curtis mentioned a second locality (viz., Regent’s Park), and is labelled

accordingly.

The lectotype of Perilitus dispar Wesmael, 1835, is here selected, and labelled

accordingly; it is the only well-preserved female of the type-series of dispar: “Coll.

Wesmael”, “1742”, “Perilitus dispar mihi & 9, det. C. Wesmael”, “Type”. Four

additional paralectotypes examined: 1 9 and 34 with the same labels. Despite the

small differences given by Wesmael (which are mainly due to the less accurate

description of Nees) dispar is a junior synonym of Perilitus albitarsus Nees, 1834,

while both are junior synonyms of albiditarsus.

The holotype of Meteorus maximus Muesebeck, 1923 (9, USNM, “Coll”, “Am.

Ent. Soc. Collection”, ‘“Zemiotes coloradensis Ashm., 9” (MS-name), “Type No.

VAN ACHTERBERG: Revision Zelinae auct. 381

24966 U.S.N.M.”, ““Meteorus maximus Mues., Type”) is a typical albiditarsus.

The lectotype of Meteorus rufulus Thomson, 1895 (3, ZIL, “p”’, “1977, 37”) is

herewith selected, and labelled accordingly. Length of fore wing 7.5 mm; hind

tarsus largely whitish, basally and apically yellowish; vein cu-a of fore wing

interstitial; length of Ist tergite 2.1 times its apical width, and length of hind femur

6.7 times its width. There is one paralectotype (¢, ZIL, “rufulus”, “1977, 36”),

which is also not essentially different from albiditarsus.

The lectotype of Meteorus romani Fahringer, 1930 (9, NR, “Kamtschatka,

Malaise”, “1231”, “Type”, “Meteorus Romani n. sp. prope tabidus” (in Fahringer’s

handwriting), “Holotype of Meteorus (Zemiotes) romani Fahr., det. T. Huddleston,

1976”, ‘403, 77”. “Riksmuseum Stockholm”) is here selected, and labelled

accordingly. Body mainly dark brown, but eye margins mainly, antennal sockets,

clypeus, mandibles, and metasoma ventro-apically, brownish; palpi and apex of

ovipositor sheath, whitish-yellow; legs largely yellowish, coxae dark brown, hind

femur and tibia apically brownish tinged, and hind tarsus mainly whitish; vein r of

hind wing absent; length of fore wing 6.7 mm; dorsal length of eye 2.4 times

temple; length of vein 1-M of hind wing 0.3 times cu-a; length of ovipositor sheath

0.21 times fore wing. A melanistic specimen of albiditarsus, comparable with the

type of Meteorus separandus Fischer, but hind tarsus mainly whitish. There is one

paralectotype (&, NR, topotypic): length of fore wing 5.4 mm; hind tarsus whitish-

yellow; hind femur more dark brown than in female; dorsal length of eye 1.3 times

temple; length of vein 1-M of hind wing 0.85 times vein cu-a.

Alaska (Mile, Elliott Hwy), Yukon Territory (Dawson, 1000 ft), Manitoba (Lac du

Bonnet), Ontario (Pt. Pelee; Rondeau Park; Florence), New Brunswick (St.

Andrew), Newfoundland (South Branch), Michigan (Gull Lake Bio. Sta.,

Kalamazoo Co.), New York (Orient, L.I.; Green Co., 2500 ft; Greenport;

Cranberry L.), New Jersey (Ocean View, Cape May Co.), Maine (SW. Harbor),

Maryland (Patuxent Ref., Cabin John), Virginia (Mountain L.), North Carolina

(Highlands; Tryon), South Carolina (Clemson), Georgia (Raban Bald; Athens),

Pennsylvania (Roxborough), Oregon (Saddleback Mt., Lincoln Co.), California

(Camino), and Mexico (Dgo., 9000 ft, 10 mi. W. El Salto) (CNC, MSU, RMNH,

UCA, USNM).

From the Palaearctic region: Finland (Taivassalo; Eckerö; Jomaba), Sweden

(Sk., Dalby), Denmark (Kirksby; Adserbo; Roden-Skov, Lolland; Dyrehavn;

Wittenberge; Hus, Westgylland; Kége), West Germany (Dollendorfer Hardt im

Siebengeb.; Kiel; Lowenberg, Siebengeb.; Mayschoss, mittl Ahr, nr. Bonn;

Hirschwaihar im Kottenforst; Katzenlochsbachtal, nr. Bonn; Lohrberg im

Siebengeb.; Geisenheim; Adenau-Hohe, Acht, Eiffel; Steinbach-Talsperre, 7 km

S. Euskirchen, Rhld; Röndorfer Tal; Mühltal, nr. München; Hochstatt, nr.

Rosenheim: Mittenwald, Hasel-Lähne, ca. 1700—1900 m; Seierskopf, nr.

Mittenwald, ca. 1000 m; Riedbergstharte, ca. 1500 m; Titisee, Schwarzwald;

Spessart, nr. Lochmühle; Hedemünden; Obbayern, nr. Gauting), East Germany

(Thüringen; Berlin), USSR (K. Merija, Nida; Varenosz, Merkini, Susviesa;

Kazakhstan, Balkarija; Georgiansk, Absjaro-Imerinsk, 10 km Bachmaro),

382 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

England (Herts., Arkley), Netherlands (Putten (G.); Heerde (G.); Ulvenhout; Rijs

(Fr.); Muiderberg; Melissant; Oostvoorne; Driebergen; Breda; Ameland, | kmS.

Ballum: id., Nesserbosch, 1 km NE. Nes; Zundert, De Krochten; Amsterdamse

Duinwaterleiding, nr. Vogelenzang; Loenen (G.); Meinweg, nr. Herkenbosch;

Mon (Z.L.); Vijlen; St. Pietersberg; Bergen; Bergerbosch; Molenven (nr.

Oisterwijk?); Vaals; Den Haag; Delft; ’s-Graveland; Baarle-Nassau; Tegelen, De

Holtmühle; Noordwijkerhout, Nieuw Leeuwenhorst; Stein; Hilversum; Ame-

rongen; Nagele; Woold; Eerde; Garderen; Epen; Oldenzaal; Haarlemmerhout;

Bussum; Beetsterzwaag; Oegstgeest, Oud-Poelgeest; Soest; Waarder; Winters-

wijk; Schayk; Vessem; Oploo; Venlo; Crailo; Nunspeet; Weesp; Asperen; Naar-

dermeer; Assel; Drijber; Botshol; Bredevoort; Overveen; Ede (G.); De Steeg;

Winterswijk; Wijster; Weert; Drunen; Keperbosch (Z.L.); Bergen op Zoom;

Meijendel, dunes; Castelre), Belgium (Lk., Stavelot; Hautes Fagnes, Mt. Rigi, 670

m; Virton, 230 m), Austria (Leitha Geb., Donnerskirchen; Tirol, Aschbach, 1400

m; Piesting; Wien; Judenbergalm, Salzburg; id., Sam-Moos; id., Wallersee; id.,

Kasern; Riedegg nr. Gallneukirchen), Switzerland (Wallis), Italy (Monte Lessini,

Pr. Verona, S. Rocco, 300 m), France (Pyr. or., Perpignan), Poland (Wroclaw;

Gdansk), Czechoslovakia (B. Stiavnica, Slov.), China (Beh Luh Din, 30 mi N.

Chengtu, Szechuan; 30 mi. N. Tatsienlu, 12000 ft, Szechuan; Mt. Omei,

6000— 7500 ft, Szechuan), Japan (Nagano, 400 m; Mt. Takao, 600 m, Tokyo;

Kiyose, Tokyo; Nippara, Tokyo; Mt. Asama, Nagano), Nepal (Ktmd., Godavari,

6000 ft; nr. Ktmd., Gulubhanjyang, 7500—8500 ft, pastures; Ktmd., Pulchauki,

8000 ft; 11100 ft, 27°58’N, 85°00’E; 9900 ft, 28°00’N, 85°00’E), Birma (N.E. Birma,

Kambaiti, 2000 m), and India (Kalatop, 2438 m, H. P.; Dalhousie, 2133 m, H. P.;

Ahla, 2286 m, H. P.) (WHC, CNC, DZD, CVR, LH, RMNH, ITZ, ZMH, IZP,

ZMB, UZM, USNM, BM, HC, ZI).

Variation: Length of fore wing 4.3—11.1 mm; antennal segments 37—50; length

of vein 1-M of hind wing 0.3—0.8 times vein cu-a; length of hind femur 5.9—7.1

times its width; length of Ist tergite 1.6—2.8 times its apical width; length of

ovipositor sheath 0.19—0.33 times fore wing; hind tarsus white or whitish-yellow,

lighter coloured than middle of hind femur; vein r of hind wing more or less

developed or completely absent; specimens from China have mesoscutum and

mesopleuron punctate or punctulate, but vertex at most punctulate; male has

apical 0.7 of hind tibia more or less infuscated; melanistic specimens of both sexes

with dark brown pterostigma, infuscated legs and body or mainly blackish body

occur rather frequently; vein cu-a of fore wing postfurcal or interstitial,

exceptionally antefurcal; frons and Ist tergite sculptured or almost smooth;

temples of male sometimes somewhat swollen; tough silken cocoon rather greyish

or almost whitish.

Known hostst of examined specimens belong to the Geometridae (Rheumaptera

hastata (L.), Macaria notata (L.)), and Noctuidae (Hypena proboscidalis (L.), Zale

spec.).

Notes. There has been a great deal of confusion about the identity of testaceator,

the type-species of the genus Zele. Up till now it has been confused with

Homolobus (Phylacter) annulicornis (Nees). This is surprising because Curtis figured

VAN ACHTERBERG: Revision Zelinae auct. 383

the metasoma (fig. 415—416) with a clearly visible laterope, far removed from the

base of the tergite. This may have been overlooked because Curtis did not state

explicitly that the figure was made after testaceator. But he explicitly stated the

presence of the vein r of the hind wing for both albiditarsus and testaceator. This

vein is always absent in Homolobus annulicornis (Nees). In my opinion Curtis was

misled by the rather pronounced sexual dimorphism in albiditarsus; actually he

named the dark male albiditarsus and the yellowish female testaceator. Already

Reinhard (in Ruthe, 1862: 2) indicated the possible synonymy of testaceator and

albitarsus (Nees). Additionally Bengtsson (1918: 29—32) proved that testaceator

has to be a synonym of albiditarsus, without the examination of the types.

Curiously he was not followed by later authors; unfortunately Bengtsson made the

wrong choice for the replacement name of Zele auct., viz., Phylacter Reinhard,

1863, instead of Homolobus Foerster, 1862.

The variability in size and colour is very large in albiditarsus. This has led a large

number of entomologists to naming forms structurally not or only slightly different

from the nominate form. Actually on the basis of colour two main groups may be

formed. The first group with more or less whitish hind tarsus includes the

synonyms: testaceator, albitarsus, dispar, calcitrator, pallitarsis, rufulus, maximus,

and romani (the last mentioned being the melanistic form). The second has the

hind tarsus more or less brownish or yellowish and includes the synonyms deceptor,

reticulatus, and separandus (the last mentioned being the melanistic form of this

group). The body colour varies from completely yellowish to completely blackish;

the latter colour is very common in Nepal. The males are usually more infuscated

than the females. I have tried hard to split up the complex, but all efforts were

unsuccessful. For instance, the shape of the mesoscutum anteriorly, the length of

the fore wing, the number of antennal segments and the index of the length of vein

1-M and vein cu-a of hind wing could not be reliably applied. Also the hosts of the

various forms are, at least partly, the same; the difference in size of the hosts may

account for the more than 100% difference in size between small and large

specimens; smaller specimens have also smaller numbers of antennal segments.

However, large difference in size is a not uncommon phenomenon among the

parasitic Hymenoptera if various host-species are attacked which differ

considerably in size.

EXCLUDED SPECIES

Austrozele assamensis (Cameron) comb. nov.

Cameron, 1910, Tijdschr. Ent. 53: 53 (as Zele).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 153.

This species will be treated in a proposed revision of the Macrocentrinae.

Macrocentrus bengtssoni (Fahringer) comb. nov.

Fahringer, 1930, Ark. Zool. 21A(8): 5 (as Phylacter).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 221.

384 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

The illustrated redescription will be published in a revision of the Macro-

centrinae.

Meteorus brunnipes Ruthe

Ruthe, 1862, Berl. ent. Z. 6: 37.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 55.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 275.

Fischer (1970: 275) has given brunnipes as a synonym of his Meteorus deceptor

(Wesmael) without giving a justification of this synonymy. Fortunately Mr. T.

Huddleston (London), who examined the lectotype, kindly informed me that this

species is a true Meteorus and is not related to deceptor.

Meteorus dubius Ruthe

Ruthe, 1862, Berl. ent. Z. 6: 27, 28.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 65.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258.

Fischer (1970: 258), in his key to the Meteorus species, has given dubius as a

synonym of Zele caligatus (Haliday), without any additional information. Again

Mr. T. Huddleston was kind enough to inform me (after examination of the

holotype) that dubius is not closely related to caligatus, and has to be referred to

the genus Meteorus s.s.

Macrocentrus dubius (Wesmael) comb. nov.

Wesmael, 1835, Nouv. Mém. Acad. Brux. 9: 168 (as Eubadizon).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 223.

Traditionally this species is considered to be related to Charmon extensor (L.),

but examination of the holotype revealed its true nature. The illustrated

redescription will be published in a revision of the Macrocentrinae.

Austrozele filicornis (Cameron) comb. nov.

Cameron, 1903, J. Straits Brch R. Asiat. Soc. 39: 128 (as Zele).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 153.

See note under Austrozele assamensis (Cameron).

Eubazus longicaudus (Curtis) comb. nov.

Curtis, 1832, Br. Ent. 9: 415—10 (as Zele longicauda).

Shenefelt, 1970, Hym. Cat. 5(2): 224.

The examination of the holotype (NMV, Melbourne, which is not a lectotype as

suggested by Shenefelt (1970: 224), because there is only one type-specimen in the

VAN ACHTERBERG: Revision Zelinae auct. 385

Curtis Collection and Curtis did not indicate that he had more than one specimen

at hand) reveals that it belongs to the genus Eubazus Nees (Helconinae,

Brachistini). However, there it becomes the senior homonym of Eubazus

longicaudis (Ratzeburg, 1844). Therefore, I have to rename E. longicaudis

(Ratzeburg, 1844). I call it E. denticulatus nom. nov., because of its small clypeal

tooth.

Austrozele maculiceps (Cameron) comb. nov.

Cameron, 1912, Annls Soc. ent. Belg. 56: 372 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 224.

Examination of the holotype (MAC, Tervuren) shows its relationship to the

Macrocentrinae and particularly the genus Austrozele Roman. It will be dealt with

in a revision of the Macrocentrinae.

Hymenochaonia melanonotus (Cameron) comb. nov.

Cameron, 1911, Timehri 1: 317 (as Zele).

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 225.

Examination of the holotype (BM, London) reveals that it belongs to the genus

Hymenochaonia Dalla Torre and it will be dealt with in a revision of the subfamily

Macrocentrinae.

Phylax nigricornis Walker

Walker, 1871, List Hym. Egypt. Arab.: 5.

Shenefelt, 1970, Hym. Cat. (nov. ed.) 5(2): 225.

Repeatedly Mr. T. Huddleston has searched, without success, for the type of

nigricornis, while it should be in BM, London. Therefore the type is considered to

be lost. Unfortunately the original description is too vague to be certain about its

identity. The colour of the hind leg and antenna exclude it from the known

Afrotropical and South Palaearctic species of Homolobus. If the note about the

“thick” antenna is taken not too literally and if Austrozele longipes (Holmgren)

occurs in Eritrea (where Phylax nigricornis was captured), it may well be this

species which belongs to the Macrocentrinae. Until more is known about the

fauna of Eritrea, the synonymy of nigricornis with longipes remains uncertain. The

interpretation by Szepligeti of nigricornis is incorrect and refers to several species

ofthe subgenus Apatia of the genus Homolobus.

Meteorus pallidus (Nees)

Nees, (1811)1812, Mag. Ges. nat. Fr. Berl. 5: 22 (as Bracon).

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 84.

Fischer, 1970, Wiss. Arbeiten Bgld. 44: 258.

386 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Fischer (1970: 258) has added M. pallidus (Nees) to his Zemiotes section, in this

paper treated as the genus Zele Curtis. But Nees stated explicitly that the Ist

tergite of metasoma has the dorsope absent (‘“abdominus segmentum petiolare

elongato-obconicum, punctulatum, nec sulcatum;….”), while all Palaearctic

species of Zele have large and easily visible dorsope (or sulcate petiolar segment

in the words of Nees). Another indication of the misinterpretation of pallidus is the

sculpture of the Ist tergite; in pallidus sensu Fischer it is striate or rugose, while

Nees calls it punctulate. Therefore I exclude pallidus from Zele Curtis and include

it in Meteorus Haliday s.s. This leaves pallidus sensu Fischer without a name, but in

my opinion it is only a rather robust form of Zele albiditarsus Curtis.

Zele somaliensis Szépligeti

Szépligeti, 1914, Mitt. zool. Mus. Berl. 7: 223.

Shenefelt, 1969, Hym. Cat. (nov. ed.) 4(1): 170.

Examination of the holotype (ZMB, Berlin) reveals its relationship to a new

genus near Austrozele in the Macrocentrinae. The illustrated redescription will be

published in a revision of the Macrocentrinae.

ACKNOWLEDGEMENTS

To the following persons I am much indebted for the loan of types, for providing

valuable information and/or for gifts of unidentified specimens (the abbreviations

used for the collections are given in brackets): Mr. B. van Aartsen, 't Harde; Dr. P.

Alayo D., Habana (PAC); Dr. P. H. Arnaud, California Academy of Sciences, San

Francisco (CAS); Dr. J. G. Betrem, Deventer; Dr. G. W. Byers, Snow

Entomological Museum, University of Kansas, Lawrence (SEM); Mrs. J. Cardale,

CSIRO, Canberra City (CSIRO); Mr. P. A. Clancey, Durban Museum and Art

Gallery, Durban; Dr. J. P. O'Connor, National Museum of Ireland (NMI); Dr. R.

Danielsson, Zoological Institute, Lund (ZIL); Dr. J. Decelle, Musée Royal de

l'Afrique Centrale, Tervuren (MAC); Dr. P. Dessart, Koninklijk Belgisch Instituut

voor Natuurwetenschappen, Brussel (KBIN); Mr. E. H. Diller, Zoologische

Sammlung des Bayerischen Staates, München (ZSB); Dr. T. van Dijk, Wijster;

Mrs. M. Favreau, American Museum of Natural History, New York (AMNH),

Prof. Dr. R. Fischer, Michigan State University, East Lansing (MSU); Dr. I. D.

Gauld, Commonwealth Institute of Entomology, London; Mr. C. Gielis,

Vlaardingen; Mr. M. J. Gijswijt, Ankeveen; Dr. H. N. Greenbaum, Gainesville,

Florida; Prof. Dr. W. Hackman, Zological Museum of the University of Helsinki,

Helsinki (ZMH); Dr. E. Haeselbarth, Institut für angewandte Zoologie, München

(HC); Dr. K. J. Hedqvist, Naturhistoriska Riksmuseet, Stockholm (NR); Dr. W.

Hellén, Helsinki (WHC); Mr. T. Huddleston, British Museum (Natural History),

London (BM); Mr. K. J. Huisman, Melissant; Prof. Dr. V. K. Gupta, Dept. of

Zoology, Delhi (DZD); Dr. A. Jakimaviëius, Institute of Zoology and

Parasitology, Akademia NAUK Lit. SSSR, Vilnius (IZP); Mlle Dr. S. Kelner-

Pillault & Mr. B. Sigwalt, Muséum National d’Histoire Naturelle, Paris (MNHN);

VAN ACHTERBERG: Revision Zelinae auct. 387

Dr. E. Kierych, Instytut zoologii PAN, Warszawa (PAN); Dr. E. Königsmann,

Zoologisches Museum der Humboldt-Universität zu Berlin, Berlin (ZMB); Mr. D.

van der Laan, Oostvoorne; Br. V. Lefeber. Maastricht; Dr. P. M. Marsh,

Systematic Entomology Lab., USDA, U.S. National Museum, Washington

(USNM); Dr. W. R. M. Mason, Biosystematics Research Institute, Agriculture

Canada, Research Branch, Ottawa (CNC); Prof. Dr. G. Morge & Dr. J. Oehlke,

Institut für Pflanzenschutzforschung Kleinmachnow, Eberswalde (IPK); Mr. T.

Munk, Vejle; Dr. A. Neboiss, National Museum of Victoria, Melbourne (NMV);

Mr. G. M. Nishida, Bernice P. Bishop Museum, Honolulu (BPBM); Dr. D. Otte,

Academy of Natural Sciences of Philadelphia, Philadelphia (ANSP); Mr. P.

Oosterbroek, Instituut voor Taxonomische Zoologie, Amsterdam (ITZ); Dr. S. J.

van Ooststroom, Oegstgeest; Dr. J. Papp, Zoological Dept. of the Hungarian

Natural History Museum, Budapest (TMA); Prof. Dr. L. L. Pechuman, Cornell

University, Ithaca (CU); Dr. E. C. Pelham-Clinton, Royal Scottish Museum,

Edinburgh (RSM); Dr. B. Petersen, Universitetets Zoologiske Museum, Kob

enhavn (UZM); Mr. G. van Rossem, Plantenziektekundige Dienst, Wageningen

(CVR); Mr. J. Scoble, Transvaal Museum, Pretoria (TMP); Dr. M. Sheehan,

University of Connecticut, Storrs (UC); Drs. M. Suwa & C. Watanabe,

Entomological Institute, Sapporo (EI); Mr. H. Teunissen, Oss; Dr. V. I. Tobias,

Zoological Institute, Akademia NAUK SSSR, Leningrad (ZI); Dr. H. K. Townes,

American Entomological Institute, Ann Arbor (TC); Prof. Dr. J. van der Vecht,

Putten; Prof. Dr. C. Vidano & Dr. P. Passerin d’Entréves, Museo ed Istituto di

Zoologia sistematica, Torino (MIZ); Prof. Dr. A. Willink, Instituto Fundacion

Miguel Lillo, Tucuman (IML); Dr. R. Wharton, Somerset, California; Dr. H.

Wolda, Smithsonian Tropical Research Institute, Balboa; Mr. J. B. Wolschrijn,

Heerde; Mr. A. P. M. van der Zon, Jakarta; Mr. C. J. Zwakhals, Arkel; Mr. K. W.

R. Zwart, Landbouw Hogeschool, Wageningen (LH); (RMNH) = Ryksmuseum

van Natuurlijke Historie, Leiden. Additionally I wish to thank Mr. T. Huddleston

(London) for his invaluable help, Mr. C. van Heijningen and Mr. J. Sloos for their

useful technical assistance.

LITERATURE

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VAN ACHTERBERG: Revision Zelinae auct. 389

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390 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

INDEX OF NAMES USED IN THE GENERA Charmon, Exasticolus, Homolobus, Zele AND

Name

acares spec. nov.

aestivalis Snellen van Vollen-

hoven, 1858

alaskensis Ashmead, 1902

albiditarsus Curtis, 1832

albipalpis Granger, 1949

albitarsis Haliday, 1835

albitarsus Nees, 1834

alternipes spec. nov.

annulatus spec. nov.

annulicornis Nees, 1834

annulicrus Thomson, 1895

antefurcalis spec. nov.

armatus spec. nov.

assamensis Cameron, 1910

atrator Curtis, 1832

atriceps Riley in Riley & Howard, 1890 Nom. nud.

australiensis Nixon, 1938

basalis Provancher, 1888

bengtssoni Fahringer, 1930

bicolor spec. nov.

bifurcatus spec. nov.

bohemani Bengtsson, 1918

brevicauda Hellén, 1958

brevinervis spec. nov.

brunnipes Ruthe, 1862

calcarator Wesmael, 1835

calcitrator Curtis, 1837

caligatus Haliday, 1835

carbonator Shestakov, 1940

chlorophthalmus Spinola, 1808

chrysophthalmus Nees, 1812

cinctus Provancher, 1880

cingulatus Granger, 1949

crassicalcaratus Viereck, 1905

crassifemur Muesebeck, 1939

crenulatus spec. nov.

cruentatus Haliday, 1833

curtis Provancher, 1886

dauricus Shestakov, 1940

deceptor Wesmael, 1835

discolor Wesmael, 1835

dispar Wesmael, 1835

dubius Wesmael, 1835

dubius Ruthe, 1862

elagabalus Nixon, 1938

THEIR SYNONYMS

Original Correct Type-locality (country) Page

genus genus and location

Homolobus Homolobus Panama, RMNH........ ee BRO

Phylax Homolobus Netherlands, lost .......... 319

Dyscoletes Zele Alaska, USNM .......... 364

Zele Zele England, NMV .......... 376

Zele Homolobus Malagasy, MNHN ....... 283

Meteorus Zele Ireland 2 NMIE fea: sree 376

Perilitus Zele Germany. 2lost va... 376

Homolobus Homolobus Kenya, MNHN .......... 292

Homolobus Homolobus India DZDs Fr ra 342

Rogas Homolobus Germany, lost

(neotype: KBIN) ......... 324

Meteorus Zele Sweden ZI zes 363

Homolobus Homolobus New Mexico, CNC ....... 345

Homolobus Homolobus New Mexico, CNC ....... 343

Zele Austrozele IndiaBME ree 383

Zele Meteorus England, NMV

Zele Homolobus Australia, BM ........... 282

Zele Hormius California, PC

Phylacter Macrocentrus USSIRANIRA SE 383

Homolobus Homolobus Mexico) CNG ee e 333

Homolobes Homolobus Burma Nike ee. eee ae 322

Phylacter Homolobus Sweden, NR............. 332

Eubadizon Charmon Finland WH@). E eee 268

Charmon Charmon New Guinea, RMNH ..... 267

Meteorus Meteorus Germany B MER. 384

Phylax Homolobus Belgium, KBIN .......... 285

Nom. nov. for albitarsis Haliday, 1835 ..................... 376

Meteorus Zele Ireland, NMI ........... 364

Zele Homolobus WSSIRMINIR aA NU 330

Bracon Zele Italy, lost

(neotype: RMNH) ....... 370

Bracon Zele Germany, lost

(neotype: KBIN) ......... 370

Phylax Bracon PARE

Zele Homolobus Malagasy, MNHN ....... 315

Zele Homolobus Kansas, SEM ............ 285

Meteorus Zele Kansas, USNM .......... 374

Homolobus Homolobus North Borneo, BM ....... 341

Charmon Charmon Treland NM IE EP PEER 268

Phylax Bracon Quebec, PC

Homolobus Homolobus USSR: NRC aan 320

Perilitus Zele Belgium, KBIN .......... 376

Phylax Homolobus Belgium, KBIN .......... 319

Perilitus Zele Belgium, KBIN .......... 376

Eubadizon Macrocentrus Belgium, KBIN .......... 384

Meteorus Meteorus Germany BM E 384

Zele Homolobus India. BM et Sys 280

ephippium Curtis, 1832

ethiopicus spec. nov.

extensor Linnaeus, 1758

filicornis Cameron, 1903

flagitator Curtis, 1837

(nom. nov. for chlorophthalmus Haliday, 1836, nec Spinola, 1808)

fulvifrons Curtis, 1832

fuscicornis Cameron, 1887

fuscitarsis Bengtsson, 1918

geminator Lyle, 1914

gracilis Provancher, 1880

gracilis Provancher, 1886

gracilis spec. nov.

huddlestoni spec. nov.

infumator Lyle, 1914

inopina spec. nov.

inopinus spec. nov.

japonica Watanabe, 1932

lacteiceps spec. nov.

levis Muesebeck, 1923

longicauda Curtis, 1832

luteus Cameron, 1911

macropterus spec. nov.

maculatus spec. nov.

maculiceps Cameron, 1912

maximus Muesebeck, 1923

melanonotus Cameron, 1911

melleus Cresson, 1872

meridionalis spec. nov.

mesoxiphius spec. nov.

neesii Ruthe, 1862

nepalensis spec. nov.

niger Provancher, 1885

nigriceps Riley & Howard, 1890

nigriceps Enderlein, 1920

nigricollis Thomson, 1895

nigricornis Walker, 1871

nigritarsis spec. nov.

nipponensis spec. nov.

niveitarsis Cresson, 1872

nudator Thunberg, 1822

obscurus spec. nov.

occidentalis spec. nov.

ochraceator Curtis, 1832

ophioninus Vachal, 1907

pacificus Provancher, 1885

pallidistigmus Cameron, 1911

pallidus Nees, 1812

pallitarsis Cresson, 1872

palliventris Provancher, 1880

pectoralis Curtis, 1832

pectoralis Nees, 1834

peronatus Shestakov, 1940

VAN ACHTERBERG: Revision Zelinae auct. 391

Zele Meteorus England, NMV

Homolobus Homolobus ManzaniayCNG@ eee. ase 318

Ichneumon Charmon Sweden, BM ............ 265

Zele Austrozele Borneo, BIMi ee). N. 384

Helcon Homolobus Irelands lostamn avec 334

Nom. nudum

Zele Exasticolus Guatemala,BM ......... 273

Phylacter Homolobus Sweden, ZE Aaa 285

Nom. nov. for chlorophthalmus Haliday, 1836, nec Spinola, 1808 334

Eubadizon Charmon POuebec IPC sew eae 265

Phylax Bracon Quebec, PC

Zele Zele Nepal.ENEr cit eh 375

Homolobus Homolobus TanzaniawBiMer ee) eer. 297

Zele Homolobus England,BM ............ 305

Charmontia Charmontia Chile ENG E CAT RE Ae 263

Homolobus Homolobus Malagasy, MNHN ....... 316

Zele Homolobus Japan, El Sa 305

Homolobus Homolobus Uganda, TORRES Aa 294

Meteorus Zele New Mexico, CU ........ 365

Zele Eubazus England, NMV .......... 384

Cyclocormus Charmon S'AfrCa MP nr 268

Homolobus Homolobus Colombia MER ET 347

Homolobus Homolobus WeandaiGN@ eee Fr 29]

Zele Austrozele Zane MUNG ees cano nododo 385

Meteorus Zele Colorado, USNM ........ 376

Zele Hymenochaonia Guyana, BM ......:..... 385

Phylax Homolobus Texas TANS PAN pn 285

Homolobus Homolobus Marocco lo NEE 326

Homolobus Homolobus Arizonas@GN@ ane a 346

Meteorus Zele Germany, BM... 364

Homolobus Homolobus NepaliGNG@ aeons 340

Phylax Doryctes British Columbia, PC

Nom. nudum

. Zele Exasticolus Miexicon RAIN Tee 275

Meteorus Zele Sweden; ZIE eren 370

Phylax excluded) EthiopiaNlost nnn 385

Homolobus Homolobus New Guinea, RMNH ..... 310

Homolobus Homolobus Japan El tre 338

Perilitus Zele Massachusetts, ANSP .... 368

Ichneumon Zele Sweden, Uppsala ......... 370

Homolobus Homolobus Brazile IC: ea 344

Homolobus Homolobus Bolivia MCR, ene 337

Nom. nudum

Meteorus Homolobus New Caledonia, MNHN .. 298

Phylax Doryctes British Columbia, PC

Macrocentrus Homolobus SWA fin Cami MIPA 303

Bracon Meteorus Germany, lost ........... 385

Perilitus Zele New Jersey, ANSP ....... 376

Phylax Bracon 2, PC

Nom. nudum

Eubadizon Charmon Germany, lost

(neotype: KBIN) ......... 268

Meteorus Zele USSRENR=-* earn 368

392 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

picinervis spec. nov.

pleuralis Cresson, 1872

priapus Nixon, 1938

pulchricornis Nixon, 1938

punctatus spec. nov.

rectinervis Spec. nov.

reticulatus Muesebeck, 1923

romani Fahringer, 1930

rosenbergi Viereck, 1911

rufipes Provancher, 1880

rufithorax Granger, 1949

rufulus Thomson, 1895

rugosus spec. nov.

separandus Fischer, 1957

sibiricus Fahringer, 1930

simillima Enderlein, 1920

simplex Watanabe, 1932

somaliensis Szepligeti

splendens Costa, 1884

terminalis Ashmead, 1889

testaceator Curtis, 1832

thoracicus Curtis, 1832

truncatoides spec. nov.

truncator Say, 1828

tuberculatus spec. nov.

tuberculifer spec. nov.

undulatus spec. nov.

unicolor Enderlein, 1920

wesmaeli Bengtsson, 1918

Zele

Eubadizon

Zele

Homolobus

Meteorus

Zele

Phylax

Zele

Meteorus

Homolobus

Meteorus

Apatia

Zele

Meteorus

Zele

Homolobus

Bracon

Exasticolus

Zele

Homolobus

Zele

Phylacter

Zele

Charmon

Homolobus

Zele

Homolobus

Zele

Exasticolus

Doryctes

Homolobus

Zele

Homolobus

Zele

Homolobus

Macrocentrinae

Zele

Macrocentrus

Zele

Meteorus

Homolobus

Exasticolus

Zele

Homolobus

Arizona, CNC ...........

Missouri, ANSP .........

SeAfncasBiM ayer

SAT ca IBM RENE

Argentine, IML..........

Chile {CNG UM ri o

New Hampshire, USNM ..

USSRe NRG Kerr

Peru USNIMUR ie werner

VARE

Malagasy, MNHN .......

Sweden, ZEN za.

Malagasy, MNHN .......

Germany,ZSB ..........

USSR,.NR tte ien

Costa Rica, PAN.........

JapanygEli senteren

Somalia, ZMB............

Italy, Naples:

Missouri, USNM

England, NMV ..........

England, NMV

Egypt ZM Bina

Indiana, lost

(neotype: RMHN) .......

Brazil ECS rer

Panama, RMNH..........

New Guinea, RMNH .....

Costa Rica, PAN.........

Belgium, KBIN ..........

393

VAN ACHTERBERG: Revision Zelinae auct.

x LITE OE STX S'7 :L7 ‘ST ET

‘x OS ZT X | ‘auij-a|eos :67 ‘97 ‘17 ‘07 ‘199dse jesıop ‘S2)18197 PIE-IS] pue wnapodoud ‘je ‘39adse [es10p ‘unjouosaw ‘Of :39] PUIY

‘67 199dse [esıop ‘peau ‘gz :19adse je UOoly

‘

peoy

‘

LZ ‘s8uim ‘97 ‘euuaque Jo xade ‘sq :euuajue ‘97

‘

‘Mmejo pury IQUUI “EZ

‘

‘joadse [eJuo1J

‘snadA]o Jo jeep ‘77 ‘1oysodiAo ‘17 :199dse jeraye] ‘snyıgey ‘07 ‘119901 ‘EPEUEI ‘Ò (snaeuurT) 4osuaixa UOWADY) ‘1€—07 ‘S34

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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XBL Pp TD X GE ‘Ob 6E: X GT JE SE: X I ‘OUIJ-O[BIS ‘gE

‘LE ‘pE—ZE odse jesiop ‘peau ‘pp :joadse [esiop ‘wnjouosaw ‘ep ‘39adse jejuou] ‘peau ‘7p joodse [essOp ‘9}1319} IS] ‘[p ‘Mejo puiy

Jano ‘op ‘snad£jo Jo xade Jo [re1ap ‘6€ ‘891 pury ‘ge :s8urm ‘LE ‘Burm 3105 JO WZ + WI SUISA Jo [rezop ‘gE ‘Burm 3105 JO INIT] pur

B-ND UI9A JO [IeJap ‘CE !euuajue ‘pe -1ONSOdIAO ‘ge ‘39adse jessye] ‘snyrgey ‘ze ‘odzojoy “Aou ‘dads 51442014249 UOUADYI ‘pp—ZE ‘SF 4

395

VAN ACHTERBERG: Revision Zelinae auct.

x O'S 266 ‘ES ‘ZS X 07.8695 ‘PS ‘OS ‘BH: X I “OUN-2]B9S :1S ‘Lp ch ‘1oxsodiao Jo xade ‘66 ‘joodse

[esiop ‘s91319) plg—js] pue wnapodoud ‘gs ‘idjed jo preep ‘Lg ‘399dse Jessop wnyouosaw ‘9g ‘snadA[9 Jo [revop ‘sg ‘399dse [eJUO1Y

‘peau ‘ps ‘euuaque jo xade ‘es ‘mejo pury Jauur ‘76 ‘82 pury ‘IS ‘yoadse [es1op ‘peau ‘OS ‘s8uim ‘6p ‘309dse je1ore) “uoanajdosauı jo

Sp ‘adAjoou ‘(s99N) S17DA099d uomany) ‘65 — St 'S3lA

‘

wed 10119]UP JO [reyop ‘gp ‘BUUIUE ‘Ly ‘ıoyısodıao ‘Op ‘399dse [1978] 'snyrgey

: x 0°S ‘69 ‘89 99: X OT IL ‘OL LI ‘59 T9: X | ‘aulf-2/80s

:99 ‘€9 ‘19 ‘09 ‘39adse [esıop ‘peay ‘I, :399dse jesıop ‘2118197 18] pue wnapodold ‘oy ‘Med pury 1ouur ‘69 !1oyısodıao Jo xade ‘gg

‘joodse [esıop ‘wnjouosaw ‘/9 :35] pury ‘99 ‘19adse [elaye] ‘uoJnajdosaw Jo ed 10L1aJue Jo jrezop ‘ç9 ‘snadAjo jo uIBIeW jeorde ‘pg

‘s8uim ‘£9 ‘joodse |e UO. ‘peay ‘79 ‘euuaque ‘[9 ‘joodse jeısye] ‘snygey ‘09 ‘adAzojoy (uaa H) snproomasg UOWIDY) ‘| L—09 'S31A

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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397

VAN ACHTERBERG: Revision Zelinae auct.

x C1 :6L'LLTPL: x | 'SUIJ-A[EIS :8L

‘SL “EL ‘ZL ‘Oodse jesıop ‘s9ui319) puz—3s] pue wnapodod

‘

6L :32] pury

gL ‘199adse [e]uo1] ‘peay ‘14 ‘399dse [esıop ‘peay ‘9/ !sduım

‘cy tyoodse jesiop “wnjouosauı ‘py !1oyısodıao ‘gz ‘399dse [el9]e] ‘smigey ‘7. ‘edAjojou ‘(uoISWIED) snainj UOUDY) ‘6L_TL "sag

: X O'S ‘88 °L8:X OT :16—68 ‘98 ‘98 ‘78 “18: X | ‘aulj-a|eos

CQ ‘€8 ‘O8 ‘399dse [e3uo1jJ ‘peau ‘16 ‘joodse [esıop ‘S2)1819} PIE—IS] ‘06 ‘yoadse [e19]g|-0SIOp ‘EX09 a[ppiu JO [IBJ2P ‘68 ‘MEIO PUIY

Ja]no ‘gg ‘me[9 pury Jauul ‘7g {199dse [esıop ‘peau ‘98 :SBUIM ‘Cg ‘FUIM 2105 JO WZ PUB VZ + VI SUISA Jo [IBJ9P ‘pg -euusyue ‘gg “ıdıed

‘zg ‘Burm pury JO [A + DS UIAA JO [IBJ2P ‘18 ‘joodse jesse] ‘snyigey ‘0g ‘adAjojoy “Aou ‘dads snippnsaagnı snjoa1spx7 ‘16—08 ‘SALA

E | |

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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399

VAN ACHTERBERG: Revision Zelinae auct.

x OZ :L6—S6 : X | ‘aur-21808 !y6— 76 ‘ded ‘16 :192dse jezuoIy ‘peau ‘96 ‘399d

-se [ESIOP ‘peau ‘66 ‘s3uim ‘pe ‘euuarue ‘6 ‘joodse [elsye] ‘snyiqey

‘

76 ‘adAjojoy ‘(uoroweg) 51U40019SNf SNIOINSDXxT ‘LE—T6 "SEA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

400

x OT POL “TOL: X | ‘ouI-A[BIS :001 “66: X O'S ‘EOI ‘TOI ‘86 ‘199dst Jessop ‘s911319) PIE —IS] ‘p0I ‘EUU9J

-ue Jo xade ‘¢g] ‘mejo pury 197n0 ‘7Q] :19>dse [es10p “wnjouosaw ‘[Q] ‘euuaque ‘OOI :3>] pury ‘66 :19>dse [EISJE-OSIOP ‘eIqn pury Jo

apis 1auur Jo xade ‘g6 "edjjesng ‘niog wol} à 1917 EO] PUB 001 ‘86 Iq ‘adAjojoy ‘(UOJaWIeD) 514402198nf SNJOINSDXY "POL —86 ‘SBIA

US (AIT

LOL

401

VAN ACHTERBERG: Revision Zelinae auct.

‘

x 8 TELL ‘Ol

‘x Tp 601 ‘901 SX 1 ‘eurj-ofeIs :gol ‘LOI ‘SOI ‘399dse [esıop ‘9313103 IS] ‘[ | ‘399dse [equo] ‘peau ‘OI | :euusjue Jo xade ‘601

-ue ‘801 ‘STuIM ‘LOI

‘MBO ojpprw 197n0 ‘90] ‘399dse je197e] ‘snjiqeu ‘SOI ‘adAjojou ‘(ura]19puq) sda21481u snjoonsoxX ‘| || —

‘euu9)]

I ‘STA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

402

‘ XBL LIT OTT HIT

x | ‘QUIj-a|eos :gII “ELL: X Tp SIL ‘ZII ‘399dse jessop “wnjouosaw ‘6] | ‘Baj pury ‘gj ‘idjed ‘7 jj ‘399dse jesıop ‘peau ‘o] | ‘adA}o]

-oy “(urajdapug) sdasıadıu snjoonspxd “61 1—91 1 ‘S314 "Med pury 1a3no ‘|| ‘399dse [esıop ‘3113197 IS] ‘pI] ‘Baj pui ‘ey ‘309dse |e19]

-e[ ‘sinds jerqn pury Jo jrevop ‘711 ‘(24403991 Jo E11 nq) P ‘ad(10399|esed (urajzopug) 40/001un (vuvdy) sngojowoy "SII—ZII 'S31A

403

VAN ACHTERBERG: Revision Zelinae auct.

x OT:LZITSTI:X O'S

PU ETI EX | Sauly-ayeos :771—071 ‘399dse jesıop ‘peau ‘L71 ‘Joadse jequody ‘peou ‘97 | ‘idjed ‘¢Z] ‘Melo ojpprw 1auul “p7Z] :ME9 910)

xarno ‘EZ ‘s3uim ‘77] ‘euuoue ‘I7] ‘399dse jeraye] ‘snyqey ‘071 ‘9d10]0Y ‘(UOXIN) s1/09v3D]2 (puvdy) snqojowoy ‘LTIT0ZI ‘S314

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

404

X OS EEL: X OT WEITET LEL: X I ‘aUI|-9|29S :OE 1 —8T1 ‘199dSt jessop ‘pray ‘p£] ‘Meo pury Jauul ‘ee | Joodse eyo ‘pray ‘ZE |

‘idjed ‘161 ‘s8urm ‘oe | ‘euuorue ‘671 ‘399dse jesaye] ‘snyrqey ‘gz] ‘adA10]0Y ‘(UOXIN) sisuaijpaisno (vijpdy) snqojowoH ‘y£ 1871 ‘S314

405

VAN ACHTERBERG: Revision Zelinae auct.

x 6O:1P1 GEL: X ST EPL Tri Obl BEI: X I OUI-[BIS :LET * X OT :9EI ‘SEI ‘199dse jesiop ‘pray ‘Ep] ‘399d

-SB [esJOp ‘2}1819] JS] ‘Zp ‘891 puly ‘[p] “adAjojoy ‘(UOXIN) siusoolayajnd (vijpdy) snqgojowoy ‘Ep{—| P| SA ‘joodse [esıop ‘0118197

IS] ‘OP ‘321 pury ‘6£I ‘309dse jesiop ‘peau ‘g£] ‘adA10[oy ‘(UOXIN) sndviad (puvdy) snqojowoy ‘Op{—8E] ‘SF14 39] pury ‘LEL ‘399d

-SB [esJop "wnJouosau ‘ge | ‘399dse jesJop ‘s1819) PIE—)S] ‘CEI “AdAJO[OY (UOXIN) sısuaıjp4ısno (DiJDdp) snqojowoy *LEI—SEI ‘SLA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

406

È / x O'S :ESI

ZS1 (691 © X OT LST ST PSI IST OSI ‘SPI ‘91 © X 1 ‘OUI-ATBIS :SSI ‘pl “Spl ‘ppl “Wedse jesıop ‘wnyouosau ‘/61 Joodse [esJop

‘sa1319) PIE—S] ‘951 139] pury ‘SI !rdjed ‘por !mejo pury Jarno ‘EST ‘Melo pury Jouur ‘76] ‘joodse jeyuouy ‘peau ‘]6] ‘joodse [esiop

‘pray ‘osi ‘euuajue Jo xade ‘gp] ‘3uim 210} JO WZ PUB VZ + VI SUISA JO [reJop ‘py ‘STUIM ‘/p] ‘FUIM PUY JO JU + DS UIA Jo [I19p

‘gp| ‘euuaque ‘sp :joodse [e19]e] ‘snyiqeu ‘pp ‘euejewouey ‘Asedejew ‘è ‘(198u819) sidjpdigjp (puody) sngojowog *LS1-p+1 "SHL

407

VAN ACHTERBERG: Revision Zelinae auct.

x 0°7:S9I EI: X O'S

79] (091 : x 1 ‘our-areos :191 ‘651 ‘851 ‘199dse yeiuoyy ‘peoy ‘91 ‘idjed ‘pgr yoodse [esiop ‘peau “¢9] ‘Euuaque JO xade ‘791 ‘SSUIM

‘19

] :Mej9 pury 19jno

‘

09] :euusyue ‘6SI

‘

‘yoadse [esse] ‘snyiqey ‘gc] ‘adAjoou (Aeg) 4orvauna (Dıypdy) sngojowoy *S91—

BSI ‘S8IA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

408

: x 0°£ ELE: X I ‘9UI-A|eos : [LL ‘691 “LOT: X OT :TLI ‘OLI ‘891 ‘991

‘idjed ‘LI ‘399dse Jessop ‘wnyouosaw ‘7/1 ‘821 pury ‘141 ‘9d10799] ‘(uoss13ua g) 17avusam (snqojorpyD ) sngojowog "ELI—ILI ‘S314

“joadse jes10p ‘wnjgouosaw ‘O1 ‘33| pury ‘691 ‘9d410192] ‘([aewsa MA) 4070951p (sngojowoH) sngojowor ‘OLI ‘691 ‘S314 "192dse [esıop

“wnjouosau ‘89] ‘Baj puiy ‘19] ‘39adse jesıop ‘say1819) piç-1S] ‘991 ‘adAjoou (Kes) 40100una1 (viipdy) sngojowog ‘89{|—99| ‘S314

CLI

feti

STE an

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Mens SSS Eee

gut EED

Q NS

ELL

409

VAN ACHTERBERG: Revision Zelinae auct.

x O'S :8LI

‘pL | 39] ajppru ‘adAjojou ‘(uoraweD) snwänsıpıjod (pıypdy) snqojowoy

‘LL1 ‘814 199dse [es10p wnyouosau ‘[g] isFurm Og] ‘39adse [eruo ‘peau ‘641 -euusJur JO xade ‘g/ | joodse Jessop ‘peau ‘9/ | :(jueu

(X99191A) snivaDojvaissp40 (vuvdy) sngojowog “181 —8L1 ‘9LI—vLI “SBA

-W91) Ba] pury ‘SLI ‘399dse [e19]E] ‘snjiqeu ‘p_| ‘edAojoq

€

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

410

x O'S :L81:% O'T 061 881 “981 ‘S81 ix I ‘auIj-a|eos :pg81—281I

‘yoodse jejuoiy ‘peoy ‘061 :199dse [esıop ‘peau ‘681 ‘joodse [esıop ‘3113137 IS] ‘gg] ‘Med 9104 ‘;g] ‘399dse [eSIOP “wnjouosaw ‘9g |

‘Idjed ‘sg :s8urm ‘pg ‘euuaque ‘eg | ‘zoodse [esaye] ‘snyrgey ‘7g] '>dA10399] ‘(ulajiopugq) 40/001un (ouvdy) snqojowop] ‘061—Z81 “S314

411

VAN ACHTERBERG: Revision Zelinae auct.

x OT ‘107 —L6I ‘P61 : X O'S :EOT “ZOT “S61 ‘T61 : X [ ‘SUIJ-[EIS :961 ‘E61 “161 MEID

pury Ja3no ‘Eq ‘me[o pury Jauur ‘707 ‘399dse [esıop ‘wnjouosow ‘107 ‘tdred ‘007 ‘19adse [esıop ‘peau ‘661 ‘192dse jeiuouj ‘peoy

‘861 ‘Burm 2105 JO VZ + WI UISA JO [rerop ‘L61 !STUIM ‘961 ‘Me[o 2107 JOUUT ‘661 ‘FUIM PUIY JO YS PUL [4 + IS SUISA JO [FEISP ‘pol

‘

‘euuaque ‘£6] ‘euuaque Jo xade ‘76] ‘399dse [eıaye] ‘snyqey ‘[6] 24410393] ‘(193ueIO) x0404Y11fn4 (brandy) snqojowoH ‘£OT— 161 ‘SSL

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

412

= x O'S :TIT TIT: X OT :SIT_EIZ ‘017 ‘807 “LOZ ‘SOT: X 1 ‘SUIJ-A[LIS :607

‘907 ‘907 ‘19adse [esıop ‘913193 18] ‘617 ‘joodse |esiop "wnJouossw ‘p|7 ‘399adse [esıop ‘peau ‘EZ ‘meo PUIU Jouur ‘zig ‘Euuaque

Jo xade ‘||7 ‘joodse [e]uo1j ‘peau ‘017 :euusjur ‘607 :Buim 3105 JO W7 + WI UISA JO [IEJSP ‘807 ‘Burm PUIU JO YS Pue [Y + DS SUI9A

JO [ré19p ‘LOC ‘S8uIM ‘907 :ıdjed ‘607 :199dse [eısye] ‘sniqey ‘907 ‘edAojou “Aou ‘dads snivjnonw (DIIDdp) snqojowoy ‘S1Z—t07 ‘S314

413

VAN ACHTERBERG: Revision Zelinae auct.

‘

‘euusjue Jo xade ‘g17

x O'S ‘977 STT SIT

‘

x O'€ :0E7:% OT (677 L77 WTL TUT 077

‘x | ‘QUT]-97BOS :£77 ‘617 “LIZ ‘917 ‘)99dse 1ouur ‘sruowSos [jeuusjue Yp pue PIE ‘OET ‘Joodse |e}uo1} ‘peau

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px ‘Boy pury ‘£77 ‘dyed ‘777 {Burm 3107 JO WZ + WI UI2A Jo [erop ‘177 ‘Burm PuIY JO YS PUL | Y + OG SUISA JO [FEISP ‘077 :sdum ‘617

‘

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

414

x O'S “LEZ ‘EET: X OT HET “BET “SET TET: X I “OUT]-2]B9S :9€7 ‘PET ‘1€7 “JOodse Jessop ‘wNn}

-ouosau ‘6£7 ‘joadse [e]uo1j ‘peay ‘867 ‘mejo pury Jeuur ‘/ç7 !euusjue ‘gez ‘idjed ‘seg ‘s3uim ‘pez ‘euusque Jo xade ‘EEZ ‘SUIM puy

JO HS Pue [Y+OS SUI9A JO |re19p ‘7EZ ‘1oodse [eıaye] ‘snyqey ‘17 ‘adAj0[oy ‘(uoxIN) sndviad (pipdy) snqojowoy "KET—IET 'S31A

415

VAN ACHTERBERG: Revision Zelinae auct.

pury J9UUI ‘767 !Mejo pury 193no ‘157

X O'S :TS7 IST PPT: X OT DST ‘EST OST ‘617 "LPT SVT

Opz ‘3uim pury Jo YS PUB [Y+9S SUISA Jo [Iejop ‘pq ‘399dse jesiop ‘2118197 IS] ‘EST :MPIO

210} JO VZ + VI UIAA Jo [erop ‘pc ‘idjed ‘pz ‘joodse jejuou ‘peau ‘647 !euusjue Jo xade ‘ppz ‘s8uim ‘pq ‘joodse [es10p ‘peau ‘7197

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-se [Joye] ‘sinds jerqty puly ‘957 ‘yoadse [elsye] 'snyigey ‘SZ ‘ad4Jo[ou ‘(UOXIN) s1440214Y2jnd (puody) sngopowog ‘T97—<SST ‘S814

417

VAN ACHTERBERG: Revision Zelinae auct.

~A]9 JO [IEIOP ‘LL7 ‘Bui 2105 JO VZ + VI UISA Jo [erop ‘9/7 :399dse jesiop ‘pray ‘9/7 !19adse jesiop ‘wmouosaw ‘FLz !199dse [esıop

‘P Jo sands jeiqn pury ‘er ‘oadse [essai] ‘© Jo sands jerqn pur ‘7/7 :39adse jesiop ‘ay1819) IS] ‘1/7 Hoedse jezuoay ‘pray ‘OL ‘321

PUIY ‘697 ‘MEjo pury Jajno ‘997 ‘Burm pury JO YS PUB | H + IS SUISA Jo [reJap ‘197 :sFurm ‘997 ‘idjed ‘997 ‘euuaque Jo xade ‘p97 :19ad

“SB [eJaje] ‘Snjiqeu ‘E97 ‘adAJOIIE JO LLT PUB ELT ‘TLT ‘997 INQ ‘adAjojou “Aou ‘dads 1v01sa|ppny (puody) snqopowoH ‘LLT—€97 ‘S314

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x OT :187 ‘6L7 ‘aUIJ-2/89S X 0°7 :987 ‘SLT ‘)9adse [esiop ‘wnjouossw ‘987 ‘Boy pury ‘687 ‘edAjojoy “Aou ‘dads snivs4nfig (4212D]

(UOXIN) snjvqvp8vje (pijvdy) snq

€87 (314 euuajue Jo xade ‘787 ‘SUIM 3105 JO W7+ VI

‘Zum PUIY JO YS PUB [Y+DS SUISA Jo [rerop ‘6/77 ‘2dAjo/je ‘(UOXIN) SIu1os1ayaınd

-Ayd ) sngojowog ‘987 ‘$87 “S314 ‘Burm pury JO no-w pue W-J] SUIDA Jo [IeJap ‘È ‘epuejy ‘sauiddi|iyg ‘

-0JOMOH ‘987 ‘814 35] pury ‘odAjojoy ‘’aou ‘dads sninjnovui (vijpdy) sngojowog ‘

UI9A JO [eJaP ‘187 ‘399dse Jeısye] ‘BUIOSeIOU ‘087

(pıpdy) snqojowoy ‘787—6LT ‘S314 ‘399dse |ejuo1j ‘peau ‘

ihr;

WE, À

419

VAN ACHTERBERG: Revision Zelinae auct.

x O'S S67 E67 : X OT :10E967 ‘767 ‘067 ‘887:

‘joodse jejuol ‘peau ‘I0E ‘399dse [esıop ‘wnjouosau ‘00€ :399dse [es10p ‘peau

x | ‘OUT]-3]B9S :167 ‘687 ‘L87

667 ‘39adse [esıop ‘2118191 Is] ‘867

‘padse [eıyusA

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(Teyse A) snuiuorydo (

L87

pijodp) sngojowog

L87 “S34

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421

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425

VAN ACHTERBERG: Revision Zelinae auct.

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3rd—7th antennal segments, inner aspect; 367, detail

Homolobus (Chartolobus) undulatus spec. nov., holotype. 366,

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VAN ACHTERBERG: Revision Zelinae auct.

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VAN ACHTERBERG: Revision Zelinae auct.

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-2p ‘619 ‘s3uim ‘819 ‘euuaque ‘/ [9 ‘Joodse [19e] ‘snjiqeuy ‘919 ‘adAjojou ‘AOU ‘sads sn70/nu249 (snydojno ) sngojowoy ‘779—919 ‘S314

O x | 9UI[-ABIS :0€9 : X O'T ITE9 EI “LZ9 ST : X O'S :6T9 ‘879 ‘979 ‘ETI ‘109dse jes

-Jop wnjouosaw ‘zeg : Burm pury JO YS PUR | H + IS SUI2A JO [IEJ>P ‘| ¢9 89] PUIU ‘OE9 ‘me pury Jouur ‘679 ‘Joodse JauuI Yuawdas

jeuusyur pig ‘879 ‘uoIna|dosaw jo Jed 10119]UE Jo [IEJ>P ‘179 ‘399dse Jesıop ‘9118191 Is] pue wnapodoid ‘979 ‘zoadse Jessop ‘peau

‘679 :MBID pury Jarno ‘79 ‘399dse 1ouur ‘sinds jerqn 3105 ‘€79 ‘adAjojou ‘AOU ‘dads sn70/nuaso (snydojno) sngojowog ‘"TEI—ETI ‘S314

929

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129 zo

Seg

x 0°S :EP9 ‘0p9 '8E9 X 0'T :Sp9 ‘tro ‘Tr9 ‘199 ‘6£9 ‘OEI “PED: x | ‘OUT]-9]B9S :LE9 “CEO ‘EE ‘399dse |E1UO1] ‘pray ‘CHO

‘joadse Jessop ‘say81a) PIE—S] ‘ppg ‘Meo pury Jarno ‘eyg ‘joodse [esıop ‘unjouosawu ‘zpg ‘FUIM PUIY JO YS Pue [Y + IS SUI9A JO

[rejop “| p9 !euusjue Jo xade ‘Opg ‘joodse [es10p ‘peau ‘6£9 ‘Mej9 pury JOUUT ‘g£9 !euusjue ‘/£9 ‘Burm 310} JO WZ PUB VZ + VI SuI2A JO

[re}op ‘9£9 !sduım ‘seg !ıdjed ‘peg ‘joodse [e1a7e] 'snyigey ‘çç9 ‘adAjojou “Aou ‘oads snzppnuuw (snydojno ) snqojowoH ‘Sp9—££9 ‘s814

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x O'S :LS9 ‘959 PS: X 0°7.:859 ‘S59 "ESITIS9 ‘899: X | ‘SUIJ-A[BIS :059 ‘619 ‘Lp9

D ‘99 ‘39adse [esıop ‘2118191 IS] ‘869 ‘MB PUY J9UUI ‘/69 ‘ME[9 PUTY Jarno ‘969 ‘Burm pury JO YS pue [ H + DS JO [IP12P ‘959 ‘euu UE

Jo xade ‘pcg ‘joodse [esıop ‘peau ‘E59 ‘1oadse [equouy ‘peau ‘759 ‘idjed ‘169 !euusyue ‘059 ‘sdUIM ‘6p9 :BUIM 310] JO WZ pue VZ + VI

SUI9A JO [IEJ>P ‘gp9 :399dse [|e19]e] 'snyrgey ‘/p9 :1oyısodıao ‘9p9 ‘adAjojou ‘AOU ‘dads snıpwan (snydopng ) sngojowoy ‘g59—9p9 “S314

VAN ACHTERBERG: Revision Zelinae auct.

x 0'7 :999 'S99 ‘E99 ‘199 ‘099: x | ‘aurj-a]eos :ÿ99 ‘799 ‘659 ‘19adse [esIop ‘Wnjouosau ‘999 :Joadse [esıop

‘s911319) puz pue Is] ‘<99 3a] pury ‘ÿ99 ‘adAjojoy “AOU ‘dads sijvoanforuw (snydojno ) snqojowoH ‘999 — 99 ‘S314 ‘19adse [es10p ‘wm

-ouosau ‘€99 32] pury ‘799 ‘zoadse [esiop ‘sd}1319} pug pue Js] ‘wnapodoid ‘199 ‘ad4jojoy ‘’aou ‘dads sauvan (snydojno ) snqojowop

‘£99— 199 'sd1 4 ‘JOodse jesıop ‘wunjouosaw ‘099 ‘82 pury ‘659 ‘adAjojoy ‘AOU ‘dads snJDUAD (snydojno ) sngojowoy ‘099 ‘659 ‘S3I4

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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VAN ACHTERBERG: Revision Zelinae auct.

x 0°S :6L9 ‘8L9 TL9 : X OT :LLITELI IL9I ‘699 : X | ‘9UIJ-A[ESS (049 ‘899 ‘L99 “MEI pury JAUUI ‘6/9

‘me[9 pury Jarno ‘g/9 joodse jesiop ‘peau ‘1/9 ‘399dse jesıop ‘2118191 IS] ‘9/9 :19>2dse jezuouy ‘peau ‘¢/9 !199dse [esıop "wnJouossw

‘pL9 :ıdjed ‘¢79 ‘euuaque Jo xade ‘7/9 ‘Burm 3105 JO YZ pue VZ + WI SUISA JO [IBJ2P ‘149 ‘s3uIM ‘OL9 ‘Burm PUIU JO YS PUB [U +IS

SUISA JO [erop ‘699 !euuajue ‘899 ‘Joodse je1arel 'snyigey ‘199 ‘adAjojoy ‘AOU ‘dads snunosgo (snydojno ) sngojowoy *6L9—L99 ‘SBA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x 0'S 069 ‘689 “S89: x 0°

T:L89 ‘989 ‘p89

‘€89 ‘189 : X | AUIJ-A[EIS :889 ‘789 ‘089 ‘MEJo pury Jono ‘069 “eUUG UR Jo xade ‘689 ‘euuajue ‘889 ‘39adse |e]uo1]} ‘peau ‘/g9 ‘100d

“SB [BSIOP ‘peau ‘989 :ME9 pury Jouui ‘989 !ıdjed ‘pgg ‘8urm pury Jo YS

PUB WZ + VI SUI9A JO [IeJap ‘[89 ‘joodse [U19Je] ‘snziqey ‘089 ‘adAjojoy ‘

PUB | Y + IS SUIAA JO [EJ>P ‘Egg !sduım ‘789 :BuIM 210] JO WZ

‘AOU “dads sijpounfo1uv (snydojno ) snqojowoy ‘069—089 ‘814

VAN ACHTERBERG: Revision Zelinae auct.

x O'S :00L ‘669 ‘969 : X 0°7 :TOL ‘IOL ‘L69L "S69 "E69 ‘769: X | ‘auIj-a]eos :869

‘p69 ‘169 ‘Joodse jesJop ‘3113137 38] ‘ZOL ‘died ‘107 !euusjue Jo xade ‘QOL :MEJ9 pury JOUUT ‘669 ‘SBUIM ‘869 :199dse [es10p ‘pray ‘/69

‘Mea pury Jajno ‘969 !199dse [eJuo1) ‘peau ‘<69 ‘joodse [8191] 'snyıgey

PUIY JO YS Pue JU + IS SUSA JO [II2P ‘769 iguu UE

‘

169 ‘edMojoy ©

‘769 ‘Burm pury JO WZ pue WZ + VI SUIAA Jo [IE]9p ‘£69 !sduım

nou ‘oads sniydixosau (snydoyng ) sngojowog ‘TOL—169 ‘S34

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

x O'S ‘1IL—60L "LOL: X OT EIL TIL “POL! x I

‘aUIj-ajeos :g0L ‘90L ‘SOL “EOL ‘399dse jeısye] ‘sinds JeIqn pury ‘ey, ‘309dse [etage] ‘ands puiy Jajno jo xade Jo [IP ‘OIL ‘Jooyseq

“S2OSBIA| “BILIJV'S ‘P (UOXIN) sndbiud (pivdy) sngojowog “EIL ‘OIL ‘S314 ‘39adse [erayej ‘Inds pury 1o3no Jo xade ‘syje [elskıyJ ‘ues

TYAN “VSN ‘© (KeS) 4osuna (vijpdy) sngojowop ‘| | L ‘314 ‘19adse [e193E] ‘sinds jergn pury ‘71 ‘ands pury 193n0 Jo xade jo [Ie]

-2P ‘602 ‘eqwede N ‘vluezuey ‘© (uorawe) snwänsıpıjjod (puvdy) sngojowo ‘TIL ‘60L ‘831-4 ‘821 pury ‘adAjojoy ““Aou ‘dads snipynu

-up (snydojno ) snqopowoy ‘gQL ‘314 ‘ME|o pury 193n0 ‘SH uoewny ‘eipur ‘è ‘(UOSs}B3Uag) 1042409 (snydojng ) sngojowog ‘LOL ‘Bid

“Baj pury ‘adAjojo ‘“Aou ‘dads 40/0019 (snydojno ) snqojowop ‘90L ‘314 ‘391 pury “SOL !12adse jesiop ‘wunjouosawu ‘pol ‘2d4joçou “AOU

‘aads smydixosou (snydojnQ) snqojowoH ‘SOL ‘+OL “83814 ‘891 pury ‘adAyojoy “Aou ‘99ds snanasqo (snydojno ) snqojowoy ‘COL ‘314

VAN ACHTERBERG: Revision Zelinae auct.

x O'S WZL ETL OCL: X OT

TTL ITL “OIL “LIL SIL: X | Naun-opeos “BIL ‘OTL “PIL Mejo pury 1970 ‘p7/ ‘Meo pury Jouur ‘eq, ‘oodse jesıop ‘peau ‘77 ‘1924

-se [eJuO1J ‘peau ‘]7/ ‘Joodse Jauul Yuawdas [euusjue PIE ‘OZL SWZ PUB VZ + VI SUIOA JO [rerop ‘614 euusjue ‘gj, !euusjue Jo xade

‘LIL ‘sduim ‘914 ‘idjed ‘sy, ‘y9odse [esse] ‘snyigey ‘pi, ‘adAjojou “Aou ‘oads sn4o1do4ovw (snydopng) snqojowoH ‘pZL—VIL ‘SA

= DIL

1979

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL.

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x O'S :TEL:X PL 67L: X | ‘OUIJ-A[BIS :£EL “OEL

LUL: X OT EL STL ‘OL “STL 32] Pury ‘Egy ‘euuaque JO xade ‘7£/ :399dst [esıop ‘wnjouosauwu ‘TEL ‘adAjojoy “ou ‘ads snjpjnorog

“NI SNJOINSOXT “EEL EL "Bl 32] Puly ‘OEL :399dse [esıop ‘ond1a3 18] pue wnapodoid ‘67, ‘adAjojoy ‘’aou ‘dads snipynpun (sngoJoı

~ADYD) SNQOJOWOH ‘OEL ‘6TL ‘814 “adse jes1op ‘wunjouosauu ‘g7/ :33] pury ‘771 ‘yoadse [es1op ‘2}1819) 35] pue wnspodoud jo jjey [eo

-ıde ‘97, ‘Burm pury Jo YS pue [Y + DS SUISA JO [leap ‘CZL ‘edAjojou ‘’aou ‘dads snsa1dosonw (snydojng ) snqojowoy ‘87L— STL ‘SBI

VAN ACHTERBERG: Revision Zelinae auct.

x OT LPL ‘OPL ‘ppl ‘TL “IPL GEL BEL: X O'S EPL “OL “DEL: X I ‘9UIJ-A[BDS :Sp/ ‘LEL “SEL “PEL VOodse jesıop ‘2118197 JS]

‘ipl ‘oadse Jessop ‘umouosaw ‘gp, :39] pur ‘spl ‘oodse [eruo ‘peoy ‘pp, ‘euuaque Jo xade ‘ep/ !ıdjed ‘zp, :1oodse [es10p

‘Tp ‘MEjo pury 193no ‘Op BUIM pury JO YS PUR |Y + IS SUIAA JO [IIOP “GEL ‘FUIM 3105 JO WZ PUB WZ + VI SUI9A JO [1BI2P “BEL

‘

‘peau

‘SAUIM

‘LEL (MBI ‘9e, ‘euuaque ‘SEL ‘joodse [1918] 'snyigey ‘pEL ‘adAjojoy “Aou ‘oads s142U1924 (snydojng) snqojowoy ‘Lyl—vEL ‘SF4

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x OS ITSL!X OT SSL—ESL “6PL = X | '9UI-9[EIS

“ISL ‘OSL ‘Spl ‘199dse Jessop ‘wunjouosau *¢¢/ ‘9nF197 Js] pue wnapodoud ‘ps, ‘39adse jejuo1] ‘peay ‘ESL ‘MIS pury Jouul ‘76/ :32]

pu ‘162 ‘s8uim ‘OS ‘3oodse [esıop ‘pray ‘6p :19>dse Jesaye] ‘smugey ‘gp, ‘9d30)99] ‘(uoswoy J) snaoijnuuw ajaZ ‘SSL—8tL ‘S314

463

VAN ACHTERBERG: Revision Zelinae auct.

‘9, :yoodse [es10p “wnjouosaw ‘po, !euusjue ‘¢g/ ‘Joodse jesıop ‘an319) Js] pue wnapodoid ‘79, ‘Baj pury ‘194

‘peau ‘092 ‘idred ‘65, ‘s8urm ‘gg, :MeI9 puly Jajno ‘/6/ ‘joadse [ae] ‘snyqey ‘g¢/ ‘adAyojoy “(ayıny) ısoou 2/27 *

‘

‘joodse jesJop

S9L—ISL ‘SUA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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VIEL ELLOLL 220,592 LOE

x | ‘QUT]-9789S8 :pLL ‘69L '89L 99L ‘MEI9 PUIY Jouut “OLL ‘joodse [esıop ‘s9ni319) puz pue 15] ‘wnapododd ‘¢// :35] puly “pli :39adse

jequoyy ‘pray ‘ELL ‘oodse jessie] ‘uoina[dosau jo Wed 1011ajU8 JO [IEIOP "TLL ‘joodse jesıop ‘wnjouosaw ‘]// ‘39adse yesiop ‘peau

‘OLL “euuIue ‘69, SBUIM ‘g9/ !euuajur Jo xade ‘/9/ :399dse [eee] ‘smigey ‘99, “ad4jojou ‘(X99q9san]y) 51427 3/9Z ‘9LL—99L ‘STIA

46‘

VAN ACHTERBERG: Revision Zelinae auct.

X OTT OBL EX TI IL ‘6LL IX | DuNj-opeos :BLL ‘x 90 :LLL ‘Wadse [equo1] ‘peau ‘134 ‘399dse [eleve]

‘

wınj[a]n9s Jo [IeJ>p ‘087 ‘192dse

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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À x | ‘auij-a[eos ZL: X TI “EBL TEL

"SBUIM ‘pg/ ‘1oadse jesJop ‘saJ1819) PIE—S] ‘EBL ‘199dSe [eS10p ‘WnJouosau ‘Tg ‘adAyojoy ‘aou ‘dads snypjound 2127 ‘98L—T8L ‘STA

£8L

x 077 :S6L—T6L '06L 68L : *X O'S

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‘peau ‘C6L ‘8umm 910] JO Q[ AD UIA JO |IPJ2P ‘Z6L AME] pury JoUUT ‘164 ‘Joodse [esıop "wnJouosaw ‘O6L ‘19adse [esıop ‘pray ‘68L

‘sBuim ‘gg; ‘euuaque Jo xade ‘/g/ !euusjur ‘987 ‘joodse [else] ‘snziqey ‘Cg/ ‘adAjojou ‘’AOU ‘dads safıqnaaognı 21927 ‘S6L—SBL ‘SA

467

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Tong

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL.

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x 0°7 :€08—008 : X OS S08 ‘O8 “66L : X I ‘AUII-21B98 :B6L—96L ‘MEIO 2105 J9UUI ‘608

‘MBI PULY Jeuur ‘p0g ‘joodse jeJUO I ‘peau ‘€09 :19>dse [es10p ‘peau ‘708 ‘1oodse [esıop ‘911319) JS] pue wnapodoid ‘10g :ıdjed

‘puuajUue JO xode ‘664 !SFUIM ‘g6L ‘euuaqUue ‘L6L

‘

‘joodse jessie] ‘snjiqeu ‘96/ ‘ad303991

‘

(UOSSOID) sısapp1aalu 2/27 ‘

S08—96L

‘008

469

VAN ACHTERBERG: Revision Zelinae auct.

x Q'S E19 X | ‘Ul]-a]B9s :718 ‘O18: X OT 118 '608—L08 : X 11 :908 “MPLS PUIU IQUUI ‘adAyojoy “Aou ‘ads snjpjound ajaZ

‘€18 814 199] pury ‘adAjojoy “Aou ‘dads safijnosoqgni 2127 ‘718 ‘FIA ‘joadse [esıop "wnjouosau ‘adAjojoy ‘Aou ‘dads 5144241914 ajaZ

‘11g 314 35] pury ‘adf10[oy “Aou ‘dads syjov4s 2/2Z ‘018 314 ‘idjed ‘608 ‘1oadse [esıop ‘wnjouosaw ‘808 ‘edAyojoy “(Y9agasanyW)

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x OS “E78 ‘618: X OT PTS ITS 818-918 © X | ‘OUIJ-O[EIS :778 ‘078 ‘S18 ‘PIS Joodse Jessop ‘218197 JS] pue winapodo.d ‘978 :mejo

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118

VAN ACHTERBERG: Revision Zelinae auct.

‘678 LISST8

‘678

‘

ME]9 pury J9UUI ‘g7g ‘STUIAM

‘

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smiqeu ‘szg ‘adAjojou ‘“Aou ‘dads smaouroid ajaZ ‘CCR —ST8 ‘SBI

7,1979

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL.

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x O'7 ‘78 ‘6€8 “BEB: X O'S (018 ‘SES : X I ‘OUIJ-ABIS :[p8 ‘LES

‘ogg ‘peg joodse [e]uo1] ‘peau ‘748 ‘821 pury ‘| pg ‘mejo pury JOUUT ‘08 ‘39adse [esıop ‘911319 IS] ‘GEZ :199dse [esıop ‘peau ‘ggg ‘eu

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‘

VAN ACHTERBERG: Revision Zelinae auct.

x O'S :158 LWB: X OT :TS8 ‘058 ‘618 ‘Sp8*

‘

x | ‘auI[-a[eos :9pg ‘pps “Eps “JOodse jes1op 'saJ1d19) pug —JS] ‘758 MEI9 pui

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prs :joadse [else] 'snyrgey ‘Epg ‘adAjojoy “Aou ‘dads s1j10D48 aJaZ ‘7SR—E pg ‘SDA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

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x 07.998198 ‘65

8:% O'S ‘858 ‘968 PS8

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‘798 (399dse jesıop wnjouossw ‘]9g ‘821 pury ‘098 !ıdjed ‘668 :

pury J9UUI ‘968 :SBUIM ‘668

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475

VAN ACHTERBERG: Revision Zelinae auct.

x OS :1L8 ‘OLB : X 0°7 ‘9L8—€L8 ‘698

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‘6/8 ‘19adse [esıop ‘wm

+ IS SUIAA JO [IIOP ‘698

698 ‘adj0]oy ‘(uossa19) s1s4D11jJDd 2727 ‘9L8—S98 ‘SUA

476 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Figs. 877—880, inner aspect of 4th antennal segment. 877, 878, Homolobus (Chartolobus) infumator (Ly-

le), 9, Netherlands, Nunspeet: 879, 880, Homolobus (Apatia) truncator (Say), 9, U.S.A., Michigan, Ann

Arbor. 877: 1.1 x ; 878, 880: 2.5 x ; 879: scale-line, | x

VAN ACHTERBERG: Revision Zelinae auct. 477

Fig. 881, Homolobus (Apatia) truncator (Say), 9, U.S.A., Michigan, Ann Arbor, inner hind claw. Figs.

882—884, Exasticolus fuscicornis (Cameron), 9, Brazil, Nova Teutonia. 882, inner aspect of apex of

hind tibia; 883, 884: inner hind claw. 881: 2.1 x ; 882: scale-line, 1 x ; 883: 2.3 x ; 884: 4.6 x

478 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, AFL. 7, 1979

Figs. 885—888, inner hind claw of 9. 885—887, Homolobus (Homolobus) discolor (Wesmael), Nether-

lands, Wijster. 888, Homolobus (Chartolobus) infumator (Lyle), Netherlands, Nunspeet. 885: scale-line,

1 x ; 886: 2.5 x ; 887: 5.0 x ; 888: 2.3 x

VAN ACHTERBERG: Revision Zelinae auct.

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COLEOPTERA

armatus 49

castaneus 47 sqq

Cremastocheilus 47 sqq

stathamae 49

DIPTERA

Acutipula 113, 122

alpium 103

ambigua 115

americana 112

Apeilesis 118

Architipula 101

benesignata 103

Beringotipula 114, 122

Brachypremna 99, 102

Ceroctenia 118

cheethami 103

clara 101

Ctenophora 104, 111,

[112, 116, 118, 120, 121

Dolichopeza 102, 108,

[109, 112, 118, 119, 120

Dendrotipula 111, 115,

[122

Dicera 118

Dictenidia 112, 116, 118,

[121

Electrotipula 104

Emodotipula 122

Flabellifera 118

hoi 111

Leptina 118

Limnophila 99

Lindnerina 122

Lunatipula 105, 114, 115,

[122

Mediotipula 114, 122

Megistocera 99, 102, 108

Nephrotoma 92, 110,

[111, 115, 118, 120, 122

Nigrotipula 110, 112,

[118, 120

Nobilitipula 111

Nophretomodes 111

Odonatisca 114, 115, 122

odontostyla 103

Oreomyza 105, 114, 115,

[122

Oropeza 99, 120

Pachyrrhina 118

Pales 118

parva 116

Platytipula 113, 122

Plocimas 121

Plusiomyia 118

Prionocera 101, 110, 112,

[118, 120

Prionota 121

Pselliophora 120

Pseudolimnophila 99

Pterelachisus 105, 114,

[115, 122

quadristriata 92

rossica 116

rufina 103

Savtshenkia 103, 114,

[122

Schummelia 113, 122

simulans 103

Stygoropis 118

Tanyptera 111, 112, 116,

[118, 121

Tipula 92, 103, 110, 111,

[113, 114, 117, 120, 122

trifasciata 92, 115

Vestiplex 105, 114, 115,

[122

Xiphura 118

Yamatotipula 113, 122

HOMOPTERA

Opsius 188

stactogalus 188

HYMENOPTERA

*abbreviatus 131, 132, 143,

[148, 182

abdominator 253

*acares 329, 336, 353

aerofacies 132, 145, 152,

[183

aestivalis 319, 320

argentifrons 127, 133, 140,

[151, 192

alaskensis 364

albiditarsus 258, 259, 362,

[375

albifacies 130, 133, 145,

[152, 187

albipalpis 278, 283, 355

Aleiodes 2

*alphitopus 133, 140, 152,

[195

*alternipes 279, 292, 355

Amicrocentrum | sqq

angulicornis 131, 144, 147,

[157

annulicornis 258, 322, 324,

[351

annulicrus 361, 363

*annulatus 329, 342, 352,

[358

annulipes 127, 135, 136,

[146, 224

*antefurcalis 330, 345, 353

Apatia 246, 260, 276 sqq

*araguensis 134, 139, 150,

[221

arenivagus 127, 133, 144,

[152, 190

*armatus 327, 330, 343,

[353

assamensis 383

atra 127

atricornis 127, 133, 142,

[151, 194

australiensis 249, 278, 282,

[357

Austrozele 383, 384, 385

482 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, 1979

*ssp. ascillaris

[(P. pygmaeus) 130, 135,

[137, 149, 233

*basifuscus 131, 143, 145,

[148, 160

bengtssoni 383

*bicolor 329, 333, 352

*biformis 131, 148, 169

*bifurcatus 322, 350, 357

bohemani 328, 332, 351,

[358

brevicauda 268, 269

f. brevicaudus

[(C. cruentatus) 264, 269

*brevinervis 264, 267

brevipetiolatus 131, 143,

[144, 164

Brulleia 5, 29, 32

brunnipes 384, 385

burmensis 35, 36

calcarator 285, 288

calcitrator 376

caligatus 253, 258, 361,

[364

carbonator 328, 330, 351,

[358

*castaneipes 133, 151, 197

Charmon 246 sqq, 257,

[261, 263 sqq

Charmontia 246 sqq, 261,

[262

Chartolobus 246, 257, 260,

[304 sqq

chlorophthalmus 258, 285,

[289, 362, 370

chrysophthalmus 370

cingulatus 312, 315, 354

clavicornis 129, 134, 136,

[146, 206

*colonensis 133, 141, 198

compressiventris 32, 34,

[35

concolor 5, 10, 13

corneimacula 35, 36

crassicalcaratus 285, 289

crassifemur 362, 374

*crenulatus 329, 341, 358

cruentatus 257, 264, 268

Isqq

*ssp. cubanus (P. areni-

[vagus) 133, 144, 192

curvinervis 5, 7, 11, 14

*curvinervosa 38

Cyclocormus 268

dauricus 313, 320, 351

deceptor 376

f. deceptor (Z. albidi-

[tarsus) 259, 362, 375, 377

*denticollis 135, 138, 229

*depressus 132, 153, 181

discolor 257, 276, 313,

[319, 351

dispar 376, 380

*Distilirella 29, 34, 38

dubius 384

*duckei 134, 138, 212

elagabalus 249, 278, 280

[356

*emarginatus 132, 142, 185

*ethiopicus 312, 318, 354

*Eubadizon 254, 268

Eubazus 384

*evansi 132, 145, 153, 184

*Exastieolus 246 sqq, 257,

[260, 271

excavatus 2

*exilis 5, 11, 16

extensor 264, 265

*facialis 130, 135, 137, 149,

[234

ferruginea 252

filicornis 384

flagitator 258, 329, 334,

[352, 353, 358

flavipenne 5, 11, 17

Formica 47 sqq

*fritzi 133, 142, 150, 199

fuscicornis 271 sqq

fuscitarsis 285, 289

geminator 334

Gorytes 128

gracilis (Charmon) 265,

[266

*sracilis (Zele) 362, 375

Homolobus 246 sqq, 260,

[276 sqq, 311 sqq

*huddlestoni 290, 297, 356

Hymenochaonia 385

*incarinatus 134, 153, 222

infumator 249, 305, 350,

[352, 353, 357

*inopina (Charmontia)

[263

*inopinus (Homolobus)

[313, 316, 354

integra 47 sqq

f. japonica (H. testace-

[ator) 305, 307

joergenseni 128, 129, 135,

[136, 148, 235

*jugularis 132, 143, 178

*lacteiceps 280, 294, 355

levis 361, 365

littoralis 130, 132, 142,

[147, 171

longicaudus 384

longiventris 131, 136, 146,

[154

luteus 268

Macrocentrus 1, 7, 8, 383

*macropterus 330, 347, 354

*Macroxyela 252

*maculatus 279, 291, 355

maculiceps 385

marginatus 186

*marthae 134, 139, 153, 208

maximus 376, 380

melanonotus 385

melleus 285, 289

*meridionalis 322, 326, 351

medius 127, 133, 137, 150,

[200

*menkei 134, 139, 216

*mesoxiphius 330, 346, 353

*metanus 134, 139, 154,

[210

Meteorus 253, 261, 360,

[384, 385

mexicanus 49

Microtypus 251

Mimesa 127, 129, 173

minutus 131, 145, 162

Myrmecocystus 49

*nepalensis 329, 340, 352,

[358

nigriceps 271, 275

nigricollis 370

nigricornis 385

*nigritarsis 305, 310, 357

*nipponensis 329, 338, 352

*nitens 134, 139, 150, 210

niveitarsis 258, 362, 368,

[370

nudator 370

*obscurus (Homolobus)

[330, 344, 353

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, 1979

*obscurus (Pluto) 134, 140,

[214

*occidentalis 329, 337, 353

*occipitalis 134, 140, 217

ophioninus 249, 257, 278,

[280, 298, 350, 356, 357

Oulophus 246, 258, 261,

[327 sqq

pallidistigma (Pluto) 13i,

145, 147, 158

pallidistigmus

[(Homolobus) 300, 303, 356

pallidus 385

pallitarsis 376

f. pallitarsis

[(Z. albiditarsus) 362, 379

Parischnogaster 2

pauper 173

pectoralis 268, 269

f. peronatus

[(Z. niveitarsis) 362, 370

peronotus 368

Phylacter 246 sqq, 258,

[260, 276, 285, 321 sqq

Phylax 276, 285, 321, 385

*picinervis 362, 373

*Platyxanion 11

pleuralis 268, 269

Pluto 127 sqq

Polyergus 48

priapus 279, 293, 355

Protelus 359

Psenia 127 sqq

pulchricornis 280,

[296, 356

*punctatellus 130, 132, 142,

[152, 179

*punctatus 361, 376

pygmaeus 128, 135, 137,

[149, 230

*rectinervis 330, 348, 354

reticulatus 376

romani 376, 381

rosenbergi 273, 275

rotundus 131, 144, 165

*rufanalis 134, 140, 150,

[220

rufibasis 133, 141, 149,

[186

rufithorax 279, 289, 355

rufulus 376

*rugosus 312, 314, 354

*rugulosus 133, 151, 197

sayi 143, 153, 173

*scytinus 133, 141, 203

separandus 376

seyrigi 3, 5, 10, 11

var. sibiricus (Z. caligatus)

[364

simillima 277, 285, 289

simplex 312, 313, 351

*simplicicollis 134, 140, 215

smithii 128, 135, 136, 146,

[226

somaliensis 386

*spangleri 131, 146, 147,

[162

*spinicollis 134, 138, 217

splendens 370

*stenopygidialis 132, 141,

[176

*stramineipes 133, 140,

[151, 204

*strigellus 133, 141, 205

suffusus 131, 144, 146, 166

testaceator 305, 307, 359,

[376, 380

texanus 132, 142, 170

tibialis 145, 151, 188

townsendi 128, 134, 139,

[153, 207

*trilobatus 134, 138, 148,

[218

sp.aff. trilobatus 138, 220

*truncatoides 280, 300,

[350, 356, 357

truncator 257, 277 sqq,

[285, 350, 352, 353, 357

*tuberculatus 272

*tuberculifer 361, 368

undulatus 305, 309, 350,

[357

unicolor 285, 289

wesmaeli 305, 306

Xiphozele 29 sqq, 34, 36

Zele 29, 246 sqa, 258, 261,

[285, 359 sqq, 386

Zemiotes 246 sqq, 359

*zonatus 134, 139, 215

*ssp. zuliensis (P. medius)

[133, 137, 150, 202

LEPIDOPTERA

abietaria 258

Abraxas 259

483

absinthiata 258

aceraria 258, 259

aceris 258

Acleris 257, 259, 267, 270,

[379

Acrobasis 258, 370

acronycta 258

Adaina 259

adelphella 258

adusta 259

adustata 259

aemula 259

aeruginea 259

Aethes 259

Agonopterix 257, 306, 308

Agrotis 257

Alcis 257, 258, 306, 308,

[336

alniaria 258

alsines 259

Alsophila 257, 258

alstroemeriana 257, 307,

[308

Amathes 257, 286

Anarta 259, 379

Angerona 258

annulata 258, 336

Antheraea 259, 379

Anticlea 259

Apamea 258

Argyresthia 259

Aristotelia 259

armigera 257

Artaversala 64

apicitripunctana 257, 367

Apoda 259

arboreus 267

Archips 257, 258, 270

Arctornis 257

atomaria 257, 258, 308

atrupictella 257, 267

aurinia 258

avellana 259

badiata 259

badiocapitella 65, 68, 70

bajaria 257

ballovi 257

betulella 258, 370

bicornutus 65, 81

bidentata 258, 259

*bipinnatellus 65, 75

bitactata 257, 286

Blepharita 259

Boarmia 258

484

brassicae (Mamestra)

[258, 259

brizella 259

brockeella 259

bruceata 257, 267

brumata 259

Bupalus 257

Busseola 5

Cabera 258

caesiata 258, 259, 336

Campaea 256, 257, 258,

[336

canosaria 257, 308

canusella 257, 267

Caripeta 258, 336

Caryocolum 259

Catarhoe 259

cebraria 257

celtis 257

cervinalis 258

Chesias 259

Chloroclysta 259

Choristoneura 257, 267

citrata 258

Cnephalocrocis 258

Cnephasia 259

Coleophora 257, 267

Colotois 259

communana 259

comptoniella 258, 370

consociella 258

contatella 258, 370

continuaria 259

contortella 259

coronata 258

Cosmia 258

crocallis 258, 259

cruda 259

cuculata 259

Deileptenia 259

deodarae 257

Dichonia 259

Dioryctria 257, 267

distaleus 65, 66, 78

divisata 258, 336

Douglasia 259

Dryobotodes 259

Ectoedemia 63

Ectropis 257

elinguaria 258, 259

Ematurga 257, 258, 300

Enargia 258, 325

Eunomos 258

Entephria 258, 336

Enypia 259, 379

Epinotia 257, 259, 267

Erannis 257

eremita 259

Eucordylea 257, 267

Eucosma 257, 267

Eulophonotus 5, 16

Euphydryas 258, 259

Eupithecia 258, 336, 365,

[379

Euproctis 257

Eupsilia 258

Eurhodope 258

Eurrhypara 258

Evagora 257, 267

exempta 257

exigua 257

expallidata 258, 259

Falseuncaria 259

fasciata 257

fasciolaria 257

Fidonia 257

filmata 258, 365

fiscellaria 257, 308

flammea 258, 259

fluctuata 259

Fomoria 63, 82

forficalis 258

francillana 259

frugiperda 257

fumiferana 257, 267

furcata 259, 379

fusca 5

fuscana 257, 267

gamma 257

Glaucolepis 64

Gloveria 257

Gnorimoschema 257

goossensiata 258

gracilis 259

granitata 259, 379

Grapholitha 257, 270

griseana 259

grossulariata 259

harrisonata 258, 336

hastata 258, 259, 379,

[382

hastiana 259

Hypagyrtis 286

Hypena 259, 382

hypericella 85, 87

infuscana 257, 267

Heliothis 257

Hoffmannophila 257, 267

Hoplodrina 259

hostilis 258

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, 1979

indigata 258, 259, 267

interrogationis 259

Ipomorpha 259

Jaspidia 259

Hydriomena 259, 379

Hypagyrtis 257

koekeritziana 259

Lacanobia 259

lamda 258, 325

Lambdina 257, 308

Larentia 258

lariciata 259

Laspeyresia 259, 267

legatella 259

Leucoptera 259

Ligdia 259

limitata 258, 336

Lithocampa 258

Lithophane 258, 325

liturata 259

longipalpata 258, 336

luctuata 259

luteata 258, 365

Lycia 257, 286, 308

Lymantria 259

Macaria 259, 382

Malacosoma 259

Mamestra 258, 259

margaritata 256, 257

Margaritia 257, 258, 259

medinalis 258

Melanolophia 258, 336

Meroptera 258, 370

Metoponia 259

Microcalyptris 59 sqq

microdactyla 259

milleri 257, 267

miniosa 259

minutacinderella 257, 267

moillieti 259, 379

molesta 257, 267, 270

Mompha 259

monacha 259

murinana 257, 267

myrmeleon 5, 16

myrtilli 259, 379

Mythimna 258

nebulosa 259

Nephopterix 258

Nepticula 59

Nepytia 257, 308

neustria 259

nigricana 267

nigroangulata 258, 259,

[336, 379

notata 259, 382

nubilalis 259

Nyctobia 258, 259, 336,

[379

obeliscata 259

Obrussa 64

obsoleta 258

ocnerostomella 259

Odontopera 258, 259

oleracea 259

Oligoneura 64

olivaceae 258, 336

operculella 257

Operophthera 257, 259,

[267

Ophiusa 32, 36

Oporinia 336

ornithogalli 257

orthogonia 257, 286

Orthosia 257, 258, 259,

[325

Ostrinia 259

ostryella 258, 370

oxycoccana 257, 267

palpata 258, 365

Panolis 258, 259

pennaria 259

perlata 258, 336

Phlyctaenodes 258

Phycita 257, 258

piceaella 257, 267

pinatubana 257

piniarius 257

piniata 257, 286

placidata 258, 336

Plusia 257

Polia 259

Polyphaenis 258

polyphenus 259, 379

pomonaria 259

pomonella 259

populeti 258

Porosagrotis 257, 286

postalatratus 65, 77

pravella 258, 370

proboscidalis 259, 382

Prodenia 257

prunaria 258

pseudospretella 257, 267

pseudotsugata 259, 379

Ptelea 83

pteliaeella 84

pulchraria 336

punctulata 65, 66, 71

purpurata 259

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 122, 1979

pusaria 258

pygarga 259

quadripunctata 257

radicana 257, 267

ramosa 258

Recurvaria 257, 267

reniculella 257, 267

repandata 257, 258, 306,

[307, 308, 336

retusa 259

Rheumaptera 258, 259,

[379, 382

Rhyparia 259

ribeata 259

roborella 257, 258

rosaceana 257, 270

rosana 258

rubricosta 257

rubrifasciella 258, 370

ruficiliana 259

rufimitrana 258

ruralis 258

Salebria 258, 370

sanguiguttata 257

satyrata 258, 365

scirpi 60, 65, 66

scitella 259

segetum 257

selepa 257

Semasia 259

Semiothisa 257, 259, 286,

[379

sericata 258

sexmaculata 259

simillima 32, 36

smithii 257, 286

solandriana 259

somniaria 257, 308

sorditana 257

Spargania 259

Spilosoma 259

Spodoptera 257

stabilis 257, 258, 259

starki 257, 267

sticticalis 257, 258, 259

Stigmella 63

suasa 259

*suavella 258

subcaesiella 258

Sylepta 258

sylviella 258, 370

Syngrapha 259

tabulana 257, 267

tenuijuxtus 65, 66, 82

Thera 258, 259

thoracealbella 65, 67, 75

Tortrix 258, 259

transversa 258

trapezina 258

triangulum 258, 325

tricororella 259

trifoliata 83

tristicula 257, 286

truncata 259

tumidana 258

turbidalis 258

ulmifoliella 257, 267

unanimis 258

unicolor 258, 336

unipunctaria 259, 379

urticae (Spilosoma) 259

usurpata 258, 365

variana 257, 259, 267, 270,

[379

variata 258

venerabilis 257

virgatella 258, 370

viridana 258, 259

Xanthorhoe 259

Xestia 258, 325

ypsillon 258, 325

Zale 259

Zeiraphera 258, 259

zonaria 257, 286, 308

ORTHOPTERA

Opsius 130

stactogalus 130

PLANTAE

Alnus 370

Betula papyrifera 336

Ceanothus cuneatus 72

Hypericum prolificum 88

Quercus 32

Quercus macrocarpa 379

Rhamnus californica 72

Robinia 370

Salix 336

Scirpus olneyi 71

Scirpus paludosus 67

Sesuvium portolacastrum

[202

Tsuga canadensis 267

Vaccinium myrtillus 258,

[336

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