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EN Ti LN \ I Ade . Dent UT RE: | ge Ti a pa di E —— i DE e Pe al n TL) | J À dr È, FA (EP | i uw; 7 j 2 MR: 4 Le PA Vi ae da 7} a LA ) a x rede By ont Eu x Le 22, apen N, 4 . A 4 1 + 1 ) EM ) Qi l'AS PORN Fae NRA TOR TE MR REN DEEL 126 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING Tijdschrift voor Entomologie, deel 126, 1983 NEDERLANDSE ENTOMOLOGISCHE VERENIGING BESTUUR (BOARD) Voorzitten (Chairman EER eeeeae rere C. A. W. Jeekel Wice-voorzitter (Vice President) Peene L. H. M. Blommers Secretarisi(SECrEta gE R. de Jong AGGRESSORI RI Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden 2311 PL lePenninemeesten(liieasurenl) Pe L. P.S. van der Geest Address ee Doornenburg 9, Landsmeer 1211 GP 2elPenmnemeesten(lineasunenil) gr ae PRE P. Oosterbroek Addresse Re APRES SR TROT AR E Ce ES, OR Baanstraat 2, Edam 1135 CB Bibliorheeanisi(leibrarıan) Pen W. N. Ellis Plantage Middenlaan 64, Amsterdam 1018 DH MM i Ans A RE B. van Aartsen TIJDSCHRIFT VOOR ENTOMOLOGIE Nedactrei(Editonralloard) peer P.J. van Helsdingen, R. de Jong, J. Krikken, M. A. Lieftinck, C. van Achterberg, S. A. Ulenberg Address RETRO ERE Rijksmuseum van Natuurlijke Historie, Raamweg 2, Leiden 2311 PL The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. Subscription rate: D.Fl. 245, — per year. Issues 1—6 appeared on 20.V.1983 Issues 7—12 appeared on 15.X11.1983 ISSN 0040-7496 INHOUD VAN DEEL 126 Achterberg, C. van. — Revisionary notes on the genera Dapsilarthra auct. and Mesocrina Koessten(EelyinenoprerasBraconidaevAlysimae) REG ERO SEE eee Achterberg, C. van. — Revisionary notes on the subfamily Gnaptodontinae, with description of eleveninewaspeciesi(Eiymenopteray Braconidae) en Achterberg, C. van. — Six new genera of Braconinae from the Afrotropical Region (Hymeno- BEeraNBracomdac RER RE RE AE BI a Et Belle, J. — A review of the genus Zonophora Selys (Odonata, Gomphidae) … Belle, J. — On the species of the polygonus group of Progomphus with a description of a new speciest(OdonataGom pda) RE Lot CO TR Brock, Th. C. M. & G. van der Velde. — An autecological study on Hydromyza livens (Fabricius) (Diptera, Scatomyzidae), a fly associated with nymphaeid vegetation COMANDI ER SIA O Brussaard, L. — Reproductive behaviour and development of the dung beetle Typhaeus typhoeus Colon Cote es OR OMR RE alee eae Lae Duijm, M. & L. Oudman. — Interspecific mating in Ephippiger (Orthoptera, Tettigonioidea) . Duijm, M., L. Oudman & B. G. Veldstra. — Copulation in Ephippiger (Orthoptera, Tettigonioi- ONE en Ml e Evenhuis, H. H. & H. J. Vlug. — The hymenopterous parasites of leaf-feeding apple tortricids (L'epidoprerasdliontrie:dae)inttheiNerherland SERRA O NO Jong, R. de. — Annotated list of the Hesperiidae (Lepidoptera) of Surinam, with descriptions of DEDEN SOI TI LI LI Lieftinck, M. A. — Notes on the nomenclature and synonymy of Old World Melectine and An- thophorine bees (Hymenoptera, Anthophoridae) M Oudman, L., see Duijm, M. Oudman, L., see Duijm, M., e.a. Velde, G. van der, see Brock, Th. C. M. Veldstra, B. G., see Duijm, M., e.a. Vlug, H. J., see Evenhuis, H. H. 203 233 269 i i La 1 i ni ti } udite NET, Kae AAN wl 4 pA dn RA { ire re u N n N (eo | / A DI 5 f L ven end meds! eme B ht TONE ROEDEN Re COLLI Hi ah wen MAT tà ve cem ad ma ur | ì i 4 : 2 i i i 7 \ EX Ve n ” 4 Vi an, De Ades TT h BRUT Pen a lan {ua LIA î gi 4 Al } } i we pi Ÿ Ur KEINE IT Y ; j N \ N \ ’ gote Il mh “Ri vee EAT ATA n i i WE ar no vi Art en & QUE i Betty ut Le de at n \ Num. 4 HN A 4 [ 1 | hei: È È N i} | eN È if 7 I "x \ i Nile i j (©) DEEI 126 | AFLEVERING 1—2 1983 VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD C. van ACHTERBERG. — Revisionary notes on the genera Dapsilarthra auct. and Mesocrina Foerster (Hymenoptera, Braconidae, Alysiinae), pp. 1—24, figs. 1—108. C. van ACHTERBERG. — Revisionary notes on the subfamily Gnaptodontinae, with description of eleven new species (Hymenoptera, Braconidae), pp. 25—57, figs. 1—131. Tijdschrift voor Entomologie, deel 126, afl. 1—2 Gepubliceerd 20-V-1983 REVISIONARY NOTES ON THE GENERA DAPSILARTHRA AUCT. AND MESOCRINA FOERSTER (HYMENOPTERA, BRACONIDAE, ALYSIINAE) by C. VAN ACHTERBERG Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands ABSTRACT The described species of the genera Dapsilarthra auct. and Mesocrina Foerster are re- vised, keyed and partly illustrated. Dapsilarthra auct. is divided into two genera, Adelurola Strand and Dapsilarthra Foerster and a subgeneric division of Dapsilarthra is proposed. A neotype is selected for Mesocrina indagatrix Foerster, and lectotypes for Adelura gahani Baume-Pluvinel, Alysia api Curtis, A. florimela Haliday, and Heterolexis subtilis Foerster. New combinations are Prorima thienemanni (Bischoff), Adelurola amplidens (Fischer), A. florimela (Haliday), Dapsilarthra subtilis (Foerster), D. indagatrix (Foerster), D. dalhou- siensis (Sharma), and D. tirolensis (Königsmann). New synonyms of Dapsilarthra are Het- erolexis Foerster, Mesocrina Foerster, Pseudomesocrina Königsmann, and Paraorthostigma Königsmann. Opisendea Foerster is excluded from the synonymy with Dapsilarthra. Opı- sendea tenuicornis Foerster, 1862, is a new synonym of Pentapleura angustata (Haliday, 1838). Neocarpa Fischer is a new junior synonym of Adelurola and Phaenocarpa multiarti- culata Marshall, 1898, is a new junior synonym of Adelurola florimela (Haliday, 1838). Dapsilarthra nowakowskii Königsmann, 1959, is synonymized with D. gahanı (Baume- Pluvinel, 1914), Orthostigma americana Brues, 1907, with Dapsilarthra apu (Curtis, 1826), Dapsilarthra testacea Griffiths, 1968, with Dapsilarthra subtilis (Foerster, 1862), Mesocrina venatrix Marshall, 1895, with Dapsilarthra indagatrix (Foerster, 1862), and Dapsilarthra fuscula Griffiths, 1968, with D. rufiventris (Nees, 1814). Additionally a new species, Dapst- larthra carpathica spec. nov., is described from Romania, and a new genus, Prorima gen. nov., is erected to include the Oriental species Mesocrina thienemanni Bischoff, 1932. INTRODUCTION The capture of a new species of Dapsilarthra during my holidays in Romania is the main rea- son to publish some of the numerous synony- mies, which I had already established during my research for a generic revision of the Braconi- dae. Adelurola, previously included in Dapsı- larthra, stands clearly apart from the rest of the species by virtue of the shape of its mandible and its enlarged 4th antennal segment. The re- maining species may be grouped in three subge- nera of Foerster, viz., Heterolexis, Mesocrina, and Dapsilarthra. The scanty information on their biology indicates that the groups treated in this paper are endoparasites of mining dipterous larvae, belonging to the families Anthomyiidae, Agromyzidae, Tephritidae, Psilidae, and Scato- phagidae. Species marked with an asterisk are new to the Dutch fauna. For the terminology used in this paper, see Van Achterberg (1979: 242— 249). KEY TO THE SPECIES OF THE GENERA Dapsilarthra AND Mesocrina AUCT. 1. Antescutal depression distinct (figs. 95, 96); vein M + CU of hind wing shorter than vein 1-M (fig. 97); claws very slender, sickle-shaped (fig. 101); propodeum com- pletely carinated and antero-dorsal part of propodeum differentiated and about as long as posterior part of propodeum (figs. 95, 99); lower outer orbits of eyes depressed (ie 98)1(cenust2rommanove) ME BEM AMAMI Eeen thienemanni (Bischoff) — Antescutal depression (virtually) absent (figs. 15, 26, 72); vein M + CU of hind wing longer than vein 1-M (figs. 6, 70); claws less slender (figs. 10, 22, 32, 47); pro- podeum not carinated; antero-dorsal part of propodeum not differentiated (fig. 1) or much shorter than posterior part (fig. 26); lower outer orbits of eyes flat or convex esere) a A: 2 2. Mandible with a ventral (4th) lamelliform TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 protuberance (figs. 12, 13, 42); 4th antennal segment longer than 3rd segment or sube- qual (figs. 1, 2); pterostigma wide and ellip- tical (figs. 6, 14, 40); vein r of fore wing emerging (sub)medially from pterostigma (fig. 6); precoxal sulcus sculptured (fig. u (genus Adelurola Strand) Mandible without ventral lamelliform I tuberance (figs. 19, 27, 29, 46); 4th antennal segment shorter than 3rd segment (fig. 43), or subequal (fig. 72); pterostigma, vein r of fore wing and precoxal sulcus variable; (ge- nus Dapsilarthra Foester) .............. 5 . Vein 1-R1 (metacarp) of fore wing almost straight; marginal cell of fore wing compar- atively short (fig. 40) amplidens (Fischer) Vein 1-R1 of fore wing distinctly curved; marginal cell of fore wing comparatively long (ISRO) EE 4 . Head distinctly transverse; dorsal length of eye 0.8—1.1 times dorsal length of temple (fig. 9) Rd ANS florimela (Haliday) Head (sub)cubical; dorsal length of eye 0.6—0.7 times dorsal length of temple asiatica Telenga . Vein CU1b and apical half of vein 2-1A of fore wing absent, resulting in an open 1st subdiscal cell; vein CUla of fore wing at same level as vein 2-CU1 (fig. 17); vein m- cu of fore wing antefurcal (fig. 17) or sub- interstitial; (subgenus Heterolexis Foerster) SR SAS AI RARE ER PR 6 Vein CU1b and apical half of vein 2-1A of fore wing present, resulting in a closed 1st subdiscal cell (figs. 29, 58); vein CUla of fore wing situated below level of vein 2- CUI (figs. 58, 80); vein m-cu of fore wing variable. grams hotes Miele 10 . Middle lobe of mesoscutum evenly setose; medial groove of mesoscutum more or less developed posteriorly; antennal segments 45— 52; parasite of Tephritidae dictynna (Marshall) Middle lobe of mesoscutum largely gla- brous; medial groove of mesoscutum absent (fig. 24); antennal segments 25—48; para- SITESIOL Np COMIZI Ca CIRO RO 7 . Scutellar sulcus smooth (fig. 24); notauli ab- sent; precoxal sulcus crenulate (fig. 15); an- tennal segments 25—29 gahani (Baume-Pluvinel) Scutellar sulcus more or less crenulate; no- tauli and precoxal sulcus variable; antennal 8. 10. ale segments 27—48 Eyes rather converging below, the maxi- mum width of head 2.2—2.7 times the min- imum width of face; antennal segments 27—33; precoxal sulcus smooth; notauli not impressed in disk of mesoscutum; Ist tergite dark and strongly contrasting with yellowish 2nd tergite.... levisulca Griffiths Eyes scarcely converging below, the maxi- mum width of head 1.8—2.1 times the min- imum width of face; antennal segments 31—48, if 33 or less then 1st tergite not dis- tinctly contrasting with 2nd tergite; precox- al sulcus usually sculptured; notauli vari- ables ais. Li 22 oc o c 9 . The anteriorly impressed part of notauli longer, reaching the disk of mesoscutum; antennal segments 34— 48; 1st metasomal tergite dark brown or black, distinctly con- trasting with middle of 2nd tergite......... balteata (Thomson) Notauli short, not or slightly impressed in disk of mesoscutum; antennal segments 31—33; 1st metasomal tergite yellowish or reddish, not distinctly contrasting with middleof Znditereiter en subtilis (Foerster) Vein 3-CUI of fore wing longer than vein CUIb (figs. 28, 35, 51); pterostigma largely elliptical; vein r emerging submedially from elliptical part of pterostigma (figs. 28, 45); vein m-cu of hind wing present (figs. 28, 45); metasoma of ® of type- species strong- ly compressed, of 6 depressed; vein m-cu of fore wing shortly postfurcal (fig. 45); (subgenus Mesocrina Foerster)......... 11 Vein 3-CU1 of fore wing shorter than vein CU1b (figs. 57, 58, 80) or equal (fig. 89); pterostigma (sub)linear, with its sides (sub)parallel (figs. 58, 82); vein r emerging between base and middle of pterostigma (figs. 70, 82); vein m-cu of hind wing absent (fig. 70); metasoma of both sexes largely depressed; vein m-cu of fore wing variable (subgenus Dapsilarthra Foerster) ...... 12 Precoxal sulcus present (fig. 31); posterior part of propodeum subvertical, steep, and distinctly differentiated from antero-dorsal part of propodeum (fig. 26); dorsope com- paratively small (fig. 37); sides of 1st tergite diverging apically . .. dalhousiensis (Sharma) Precoxal sulcus absent (fig. 43); posterior part of propodeum continuous with antero- dorsal part, slightly oblique (fig. 43); dor- sope large (fig. 50); sides of 1st tergite sub- 12. 19. 14. 15. 16. VAN ACHTERBERG: Dapsilarthra and Mesocrina 3 parallel indagatrıx (Foerster) Vein r of fore wing (very) short, shorter than width of pterostigma (figs. 82, 90) or subequal (fig. 38); pterostigma not reaching beyond middle of marginal cell of fore wing, rather stout (figs. 38, 82, 90); vein m- cu of fore wing shortly antefurcal (fig. 38) Soo ess A EDI IE 13 Vein r of fore wing comparatively long, longer than width of pterostigma (figs. 58, 70); pterostigma reaching beyond middle of marginal cell of fore wing, and slender (figs. 58, 70); vein m-cu of fore wing shortly postfurcal (fig. 70) or subinterstitial .... 15 Antennal segments 40-41; vein 3-CU1 of fore wing subhorizontal (fig. 89); antenna about twice as long as body net EE REN isabella (Haliday) Antennal segments 25—33; vein 3-CU1 of fore wing oblique or subvertical (figs. 38, 92); antenna shorter than twice length of EO. 2.000 et eT 14 Antenna short, about 1.3 times length of body; marginal cell of fore wing distinctly removed from wing apex (fig. 38); hind fe- muspondiebiadarkibrown! AES. MS debet SA tirolensis (Königsmann) Antenna medium-sized, about 1.7 times length of body; marginal cell of fore wing comparatively close to wing apex (fig. 90); hind femur and tibia brownish-yellow; (nominate form has hind coxa and 2nd ter- gite yellowish, which are largely dark brown or blackish in forma fuscula Grif- NN) AEEA rufiventris (Nees) Notauli distinctly impressed posteriorly; precoxal sulcus extensively sculptured sylvia (Haliday) Posterior half of notauli absent (fig. 78); precoxal sulcus smooth or narrowly and su- perficially sculptured (figs. 56, 72)...... 16 Sides of mandible parallel (figs. 61, 62); laterope absent (fig. 56); 3rd and 4th anten- nal segments yellowish; vein 1-R1 (meta- carp) of fore wing less curved (fig. 58); ven- tral half of side of pronotum largely smooth (fig. 56); length of fore wing ca. 2.8 mm .... carpathica spec. nov. Mandible widened dorsally (figs. 68, 69), because of the protruding Ist tooth; laterope present (fig. 72); 3rd and 4th an- tennal segments dark brown; vein 1-R1 of fore wing more curved (fig. 70); ventral half of side of pronotum distinctly sculptured (fig. 72); length of fore wing 3.5—4.8 mm … PRE ORI, en EE apu (Curtis) Genus Prorima nov. Type-species: Mesocrina thienemanni Bi- schoff, 1932. Etymology: From “pro” (Latin for “in front of”, forward”) and “rima” (Latin for “cleft”) because of the antescutal depression on the an- terior part of the mesosoma. Diagnosis. — Fourth antennal segment slightly shorter than 3rd segment (including an- nellus); eyes glabrous, not emarginate (fig. 104); lower outer orbits of eyes depressed (fig. 98); mandible slender, parallelsided, with 3 small teeth, 2nd tooth somewhat longer than 1st tooth, without ventral protruding tooth or la- mella, and with carina to 3rd tooth (fig. 98); an- tescutal depression distinct (figs. 95, 96); pro- nope medium-sized (fig. 99); medio-posterior groove of mesoscutum present (fig. 99); precox- al sulcus present, superficially crenulate (fig. 95); metanotum blunt dorsally (fig. 95); propo- deum completely areolated (fig. 99), its antero- dorsal part differentiated and about as long as posterior part of propodeum (fig. 95); pterostig- ma elliptical; vein r emerges somewhat behind middle of pterostigma (fig. 97); vein CU1b of fore wing present, somewhat longer than vein 3-CUI (fig. 105); 1st subdiscal cell of fore wing closed; vein m-cu of fore wing distinctly post- furcal (fig. 97); vein M + CU of hind wing shorter than vein 1-M (fig. 97); vein m-cu of hind wing faintly indicated; marginal cell of hind wing medium-sized (fig. 97); claws very slender, sickle-shaped (fig. 101); dorsope large (fig. 102); 2nd tergite smooth; ovipositor sheath (fig. 106) longer than apical height of metasoma. Distribution. — Contains only the Oriental type-species. Prorima thienemanni (Bischoff) comb. nov. (figs. 95— 108) Mesocrina thienemanni Bischoff, 1932: 742— 744, fig. 5. Shenefelt, 1974: 996. Holotype, ®, length of head and mesosoma combined 1.5 mm, of fore wing 2.8 mm. Head. — Antenna missing, except for scapus and pedicellus; an earlier examination proved that the length of the 3rd segment is slightly longer than 4th segment; length of maxillary palp about 0.7 times height of head; dorsal length of eye 2.0 times temple (fig. 103); POL: 4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 & ocellus : OOL = 5 : 3 : 9; frons flat, smooth; vertex smooth, with deep medio-longitudinal suture (fig. 103); face smooth, rather flat; ante- rior tentorial pits medium-sized, far removed from eyes (fig. 104); clypeus rather convex, smooth, its apical margin not differentiated; length of malar space 0.3 times basal width of mandible; medial length of mandible 1.3 times its basal width. Mesosoma. — Length of mesosoma 1.8 times its height; side of pronotum smooth, except for the transverse crenulate groove (fig. 95); pre- coxal sulcus narrowly impressed, superficially crenulate anteriorly, but posteriorly absent (fig. 95); pleural suture smooth dorsally, narrowly crenulate ventrally; episternal scrobe large, deep, connected to pleural suture (fig. 95); metapleural flange absent; metapleuron largely smooth, but ventrally rugose; only anterior third of notauli impressed and crenulate, rest absent; medial suture long, reaching midpoint of mesoscutum (fig. 99); mesoscutum largely glabrous; scutellar sulcus wide, deep, without distinct longitudinal carinae; scutellum com- pletely smooth, rather flat; metanotum with medial carina anteriorly; medial carina of pro- podeum long (fig. 99); posterior part of propo- deum with a narrow areola; propodeal spiracle small, round and far from anterior margin of propodeum (fig. 95). Wings. — Fore wing: vein 1-SR rather long (19,197) ES SRIISRAE 42:519 SD SRI aM slightly sinuate; SR1 straight; cu-a short, post- kurcal ECU EC HIER MIS eRsubdiscal cell about 5 times its maximum width (fig. 105); basal half of M + CUI not or indistinctly scle- rotized; 2-SR : 3-SR : r-m = 12: 19 : 6; m-cu strongly converging to 1-M (fig. 97). Hind wing: cu-a short, rather reclivous; M + CU : 1- M = 10 : 12; marginal cell absent apically. Legs. — Hind coxa smooth; only claws of middle leg present, long setose, with minute lobe (fig. 101); length of femur and tibia of hind leg 5.2 and 8.6 times their width, respectively; 3rd and 4th tarsal segments of middle leg sub- quadrate (fig. 101). Metasoma. — Length of 1st tergite 1.7 times its apical width, its surface largely smooth, but between dorsal carinae rugose (fig. 102), basally concave, rest strongly convex; dorsal carinae of Ist tergite present in basal % of tergite; spira- cles of 1st tergite just in front of middle of ter- gite, protruding (fig. 102); glymma wide ante- riorly; laterope rather small, shallow (fig. 95); dorsope deep, diameter of dorsope equal to about half their distance apart (fig. 102); ovipo- sitor slender; length of ovipositor sheath 0.46 times fore wing; hypopygium sharp apically, large (fig. 108). Colour. — Dark brown; legs, scapus, pedi- cellus, palpi, tegulae, and metasoma ventrally, yellowish; pterostigma brown; wing membrane slightly infuscated. Holotype in Zoologisches Museum, Berlin: “West-Java, Tjibodas, 14.7.29, Thienemann S.”, “Mesocrina thienemanni Bisch., Typ., det Bis- choff”, “Typus”, “Präp. 18.2.55/ 1 2”. Metaso- ma largely on separate slide. Notes. — In existing keys this species runs to Pseudomesocrina Königsmann (= Mesocrina Foerster), however, it certainly does not belong in this genus. In particular the long vein 1-M of hind wing, the depression of the lower outer or- bits, the long vein CU1b of fore wing, the long anterodorsal part of the propodeum, the pres- ence of the antescutal depression and the shape of the claws indicate that this species is more closely related to Phaenocarpa. However, M. thienemanni differs from Phaenocarpa by the postfurcal vein m-cu of fore wing, the short 4th antennal segment (but some spp. of Phaenocar- pa have similar antennae), the presence of an an- tescutal depression, and of a depression at the lower outer orbits of the eyes (the latter is sometimes shallowly indicated in Phaenocarpa), preventing its inclusion in Phaenocarpa. The new genus also differs from Dinotrema, mainly by the wide pterostigma, the antescutal depres- sion, the large claws, and the antero-dorsal part of the propodeum being subequal to its posteri- or part. Genus Adelurola Strand stat. nov. Adelura Foerster, 1862: 267 (nec Bonaparte, 1854). Shenefelt, 1974: 986. Adelurola Strand, 1928: 51 (nom. nov. for Adelura Foerster). Shenefelt, 1974: 986—987. Neocarpa Fischer, 1966: 185. Shenefelt, Syn. nov. 1974: 987 Type-species: Alysia florimela Haliday, 1838. Diagnosis. — Fourth antennal segment lon- ger than 3rd segment (figs. 1, 2) or subequal; apical antennal segment without spine; lower outer orbits flat or convex (fig. 12); eyes gla- brous, not emarginate; mandible strongly widened dorsally, with a ventral (4th) lamelli- form protuberance (figs. 12, 42); pronope small and shallow or absent (fig. 4); antescutal depres- sion absent; medio-posterior groove of mesos- cutum present (fig. 4); precoxal sulcus sculp- VAN ACHTERBERG: Dapsilarthra and Mesocrina 5 tured (fig. 1); metanotum blunt dorsally (fig. 1); antero-dorsal part of propodeum not differ- entiated from posterior part of propodeum (fig. 1); pterostigma wide and largely elliptical (figs. 6, 14); vein M + CUI of fore wing largely un- sclerotized; vein r of fore wing emerging (sub)medially from pterostigma (fig. 6); vein CU1b of fore wing present, longer than vein 3- CUI (fig. 8); 1st subdiscal cell of fore wing closed; vein m-cu of fore wing slightly postfur- cal (figs. 6, 14); veinM + CU of hind wing lon- ger than vein 1-M (fig. 6); vein m-cu of hind wing absent; marginal cell of hind wing medi- um-sized basally (fig. 6); claws moderately slen- der (fig. 10), not sickle-shaped; dorsope large (fig. 7); 2nd tergite smooth; setae of metasoma in subapical rows; ovipositor sheath shorter than apical height of metasoma (fig. 1). Distribution. — Contains 3 Palaearctic spe- cies. Adelurola amplidens (Fischer) comb. nov. (figs. 40—42) Neocarpa amplidens Fischer, 1966: 185, figs. 9—11. Dapsilarthra amplidens; Shenefelt, 1974: 987. Only known from 4 d from Iraq, of which the holotype should be housed in the Zoolo- gische Sammlung des Bayerischen Staates in Munchen, where it could not be traced. Owing to the kindness of Dr. Fischer I was able to ex- amine two topotypic paratypes upon which the insertion in the key is based. This species is very close to A. florimela; the difference of the short marginal cell of fore wing is not significant be- cause this is also common in males of florimela (fig. 14). Adelurola florimela (Haliday) comb. nov. (figs. 1—14) Alysia florimela Haliday, 1838: 239. Dapsilarthra florimela; Shenefelt, 1974: 988—989. Phaenocarpa multiarticulata Marshall, 1898: 245. Syn. nov. Dapsilarthra multiarticulata; Shenefelt, 1974: 989. Dapsilarthra pentapleuroides Fischer, 1971: 85, figs. 25— 27 (unavailable name, used in synonymy with D. multiarticulata). Redescribed after a female from the Nether- lands (Linschoterbos), compared with the lecto- type, length of body 3.3 mm, of fore wing 3.7 mm. Head. — Antennal segments 43, long setose, length of 3rd segment 0.9 times 4th segment (fig. 2), length of 3rd and 4th segments 3.4 and 3.6 times their width, respectively; penultimate segment of antenna 2.6 times its width (fig. 5), and apical segment without spine; length of maxillary palp 1.1 times height of head; dorsal length of eye 1.1 times temple; POL : © ocel- lus: OOL = 5:3:12; frons slightly convex, medially somewhat rugose, laterally partly punctulate, rest smooth (fig. 9); vertex smooth, with medial groove; face medially convex, somewhat punctulate and microsculptured, shi- ny; anterior tentorial pits large, far removed from eyes (fig. 3); clypeus strongly convex, punctate; apical margin of clypeus straight, nar- row, thin and differentiated; malar space almost absent (fig. 12); 2nd tooth of mandible longer and sharper than both lateral, more lobe-shaped teeth (fig. 13). Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum rugose ventrally, rest smooth; epicnemial area rugose; precoxal sulcus only medially developed (fig. 2), deep, rugose; rest of mesopleuron smooth; pleural sulcus crenulate ventrally, almost smooth dor- sally (fig. 1); episternal scrobe deep, elliptical; metapleural flange absent; metapleuron largely coarsely reticulate-rugose; notauli only ante- riorly impressed, microsculptured, rest absent; mesoscutal lobes glabrous medially; medial groove of mesoscutum rather long (fig. 4); scu- tellar sulcus deep, with one longitudinal carina; scutellum punctulate; metanotum with rather long medial carina (fig. 4); surface of propo- deum coarsely reticulate-rugose, no medial cari- na or areola; propodeal spiracle small, round, submedially. Wings. — Fore wing; 1-SR rather long (fig. 6); r : 3-SR : SR1 = 4:18:42; 1-SR + M and SR1 slightly curved; cu-a medium-sized, post- UC CUIR CURE 252 SRE ESR r-m = 13 : 18 : 7; m-cu parallel to 1-M (fig. 6). Hind wing: cu-a rather short (fig. 6); marginal cell narrowed apically. Legs. — Hind coxa smooth; tarsal claws slen- der, setose (fig. 10); length of femur, tibia, and basitarsus of hind leg 4.1, 9.5, and 7.0 times their width, respectively; length of hind tibial spurs 0.3 and 0.4 times hind basitarsus. Metasoma. — Length of Ist tergite 1.8 times its apical width, its surface behind the spiracles coarsely rugose (fig. 7), medio-basally concave, rest convex; dorsal carinae of 1st tergite present almost to apex of tergite, united at about mid- segment (fig. 7); glymma rather wide, without laterope (fig. 1); dorsope deep, large, their diameter about equal to their distance apart; 6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 ovipositor straight; length of ovipositor sheath 0.06 times fore wing; hypopygium large (fig. 1), truncate apically. Colour. — Black; scapus, annellus, palpi, veins of hind wing mainly, tegulae, and legs, light yellowish; mandibles, antenna (except for scapus and annellus), pterostigma, most of fore wing veins, and metasoma behind 1st tergite, dark brown; apical % of hind tibia and base of hind tarsus darkened. Redescribed after a specimen in the Rijksmu- seum van Natuurlijke Historie, Leiden: “Lin- schoterbos, 8.x.1966, v. Ooststroom”, “com- pared & conspecific with lectotype of Alysia florimela Haliday, C. v. Achterberg, 1979”. The type-series in the Haliday Collection (Dublin) consists of 1 9 and 1 g, both with the label “British Haliday, 20.2.82/Box 10 AWS.” Origi- nally I was inclined to consider only the © an original type-specimen, because Haliday did not state he had a male. However, Stelfox in a copy of his notes (discovered by Dr. C. O’Riordan and a partial copy kindly supplied by Dr. J. P. O'Connor) stated under Alysia florimela Hal- iday, that Haliday did include the male: “In de- scribing the male of this species as unknown Marshall (Trans. ent. Soc. London, 1895, p. 365) fell into error. When describing species of which he had only one sex Haliday always be- gan his diagnosis with the words mas or fem. as the case might be: — e.g., “Sp. 43. Punctigera. Al. & c. fem. fusco-castanea ...” or again “Sp. 48. Perdita Al & c. mas nigra...” On the other hand, when he had both sexes before him, he only mentioned the sexes when referring to some character which belonged to that sex alone: in the present case “fem. terebra subexerta” and “fem. corpore duplo longiores articulis 50.” That he had both sexes of the pre- sent species is manifest, for box 10 I found a d & a 2 beside each other (on 26.1.1935).” The indirect evidence put forward by Stelfox in his notes makes it possible to accept both specimens from box 10 as types; I prefer to des- ignate here the © as lectotype (and not the d as Stelfox did without publishing it). The 9 lecto- type is in a fairly good condition, the antenna has 50 segments (as stated by Haliday) and the dorsal length of the eye is 1.1 times the dorsal length of the temple. The lectotype stems from England (London). In the Haliday Collection there are two specimens (1 ® (“Clifden, July 8, [18]39”) and 1 g, which do not belong to the type series. Additionally 9 © and 5 d were ex- amined from the Netherlands (Waarder; Aspe- ren; Udenhout), West Germany (Taufkirchen bei München), and Bulgaria (Rhodopi Moun- tains, Thagortsharn; id., N. Zdraves). The spec- imen from Den Haag reported by Snellen van Vollenhoven (1873: 195) belongs to Phaenocar- pa. Variation. — Antennal segments 43—54; length of 3rd segment 0.9—1.0 times 4th seg- ment; dorsal length of eye 0.8—1.1 times dorsal length of temple; vein 2-SR of one fore wing of specimen from N. Zdraves partly absent; pterostigma of male enlarged (fig. 14); metaso- ma of some specimens more or less yellowish banded. Note. — The proposed synonymy is based on the redescription by Fischer (1971) and the vari- ation found in the series collected in the Nether- lands. A. florimela is an easily recognizable spe- cies, with is not uncommon in Waarder (Neth- erlands). The enlargement of the pterostigma in the male misled Marshall, so that he described the male as a separate species (multiarticulata). A. florimela has been reared from Pegomya ni- gritarsis Zetterstedt (Anthomyiidae) and Acidia cognata (Wiedemann) (Tephritidae). Adelurola asiatica Telenga Adelurola asiatica Telenga, 1935: 186. Dapsilarthra asiatica; Shenefelt, 1974: 987. The type from Turkestan (USSR) is destroyed and additional specimens of this species have not been recorded. Genus Dapsilarthra Foerster Gnamptodon Haliday, 1833: 265 (suppression re- quested to ICZN). Dapsilarthra Foerster, 1862: 267. Shenefelt, 1974: 986—991. Marsh, 1979: 222. Heterolexis Foerster, 1862: 268; Shenefelt, 1974: 992. Syn. nov. Grammospila Foerster, 1862: 269; Shenefelt, 1974: 987. Mesocrina Foerster, 1862: 266; Shenefelt, 1974: 996. Syn. nov. Pseudomesocrina Kénigsmann, 1959b: 611; Shenefelt, 1974: 1018. Syn. nov. Paraorthostigma Köningsmann, 1972: 25—26, 1 fig. Syn. nov. Diagnosis. — Antennal segments 25—52; 4th antennal segment shorter than 3rd segment (fig. 43) or subequal (fig. 72); apical segment of an- tenna without spine; eyes glabrous, not emar- ginate: lower outer orbits of eyes flat or convex (fig. 29); mandible without ventral lamelliform protuberance (figs. 29, 46), robust (fig. 29) or VAN ACHTERBERG: Dapsilarthra and Mesocrina 7 slender (fig. 25); pronope usually absent (fig. 49), seldom present (fig. 24); antescutal depres- sion absent; medio-posterior groove of meso- scutum present (fig. 49) or absent (fig. 24); metanotum blunt dorsally (fig. 15) or slightly protruding (fig. 26); propodeum not carinated, its antero-dorsal part not differentiated from posterior part (fig. 72) or much shorter than posterior part (fig. 26); vein 1-SR of fore wing medium-sized (fig. 58), exceptionally short (fig. 28); 1st subdiscal cell of fore wing parallel- sided; vein M + CUI of fore wing largely un- sclerotized; vein M + CU of hind wing longer than vein 1-M; marginal cell of hind wing nar- rowed apically or subparallel; claws moderately slender, not sickle-shaped (figs. 22, 32); dorsope rather small (fig. 18) to large (fig. 50); laterope absent or shallow; 2nd tergite smooth; setae of metasoma in subapical rows; length of oviposi- tor sheath subequal to apical depth of metasoma (fig. 54) or shorter (fig. 72). Distribution. — All three subgenera occur in the Palaearctic region (including the Himalayan area), the subgenus Dapsilarthra Foerster also occurs in the Nearctic region (partly intro- duced). Subgenus Heterolexis Foerster stat. nov. Heterolexis Foerster, 1862: 268. Shenefelt, 1974: 992. Grammospila Foerster, 1862: 269. Shenefelt, 1974: 987. Syn. nov. Diagnosis. — Antennal segments 25—52; medio-posterior groove of mesoscutum absent (fig. 24), only present in dictynna; precoxal sul- cus impressed, smooth or crenulate (fig. 15); vein CU1b and apical half of vein 2-1A of fore wing absent, resulting in an open 1st subdiscal cell (fig. 17); vein CUla of fore wing at same level as vein 2-CU1; vein r of fore wing arising before middle of pterostigma; vein m-cu of fore wing antefurcal (fig. 17) or subintersitial; pterostigma linear (fig. 17); dorsope rather small (fig. 18). Distribution. — Contains five Palaearctic species; one species is introduced in North America (Hendrickson & Barth, 1979). Dapsilarthra (Heterolexis) dictynna (Marshall) Adelura dictynna Marshall, 1895a: 422—423, fig. 6. Dapsilarthra dictynna; Königsmann, 1972: 22. Shene- felt, 1974: 988. A scarcely collected but widespread species; not yet collected in the Netherlands. Examined: 1 2 (Bulgaria, Rhodopi Mountains, Pamporo- va) and 1 gd (Switzerland, Aeschi-Ried, 1000 m). Parasite of Pycnoglossa flavipennis (Fallén) (Tephritidae) in Pteridium aquilinum L. Dapsilarthra (Heterolexis) gahani (Baume- Pluvinel) (figs. 15—25) Adelura gahani Baume-Pluvinel, 1915: 47, 11 figs. Dapsilarthra gahani; Shenefelt, 1974: 989. Dapsilarthra nowakowskii Königsmann, 1959a: 591, _ figs. Shenefelt, 1974: 989. Syn. nov. Lectotype, 9, length of body 1.5 mm, of fore wing 2.1 mm. Head. — Antennal segments 29, length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th segments 4.5 and 4.0 times their width, respectively; penultimate segment of antenna 3.0 times its width (fig. 20); length of maxillary palp 0.6 times height of head; dorsal length of eye 1.3 times temple (fig. 21); POL : © ocellus : OOL = 14 : 5 : 18; frons flat and smooth; ver- tex smooth, with medial groove; face slightly convex, smooth; anterior tentorial pits very large, wider than long and almost reaching eyes (fig. 16); clypeus convex and smooth, its ventral margin not differentiated, straight medially; eye nearly touching base of mandible (fig. 15); man- dible rather slender, not widened apically (fig. 25), with three sharp teeth. Mesosoma. — Length of mesosoma 0.9 times its height; pronope distinct (fig. 24); pronotal side largely smooth, except for some rugae (fig. 15); precoxal sulcus rather shallow and rugu- lose; rest of mesopleuron smooth; pleural sulcus finely crenulate; episternal scrobe narrow (fig. 15); metapleural flange absent; metapleuron largely smooth; notauli absent on dorsal disc of mesoscutum (fig. 24), faintly impressed in front of disc (fig. 15); mesoscutum rather convex, largely glabrous and smooth; medio-posterior groove of mesoscutum absent; scutellar sulcus deep, with no carinae (fig. 24); scutellum rather flat and smooth; surface of propodeum largely smooth, except for some rugulosity, without medial carina or areola; posterior pat of propo- deum not differentiated from antero-dorsal part; propodeal spiracle small and in front ot middle of propodeum. Wings. — Fore wing: r : 3-SR : SR1 = 8: 24 : 92; SRI sinuate (fig. 17); 1-CU1 : 2-CU1 = SE AEN 2E SRE NSS RET EMISE RA E Hindwing: cu-a reclivous; marginal cell (sub)parallel-sided apically. 8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 Legs. — Hind coxa smooth; tarsal claws min- ute, slender (fig. 22); length of femur, tibia and basitarsus of hind leg 5.2, 7.8, and 4.5 times their width, respectively; inner hind spur about Va times hind basitarsus, outer spur not visible (at 80 X). Metasoma. — Length of 1st tergite 2.0 times its apical width, its surface rugulose, medially convex, basally and latero-posteriorly flat (fig. 18); dorsal carinae of 1st tergite present in basal half of 1st tergite; laterope absent; length of ovi- positor sheath 0.05 times fore wing, somewhat widened medially and upcurved (fig. 15); hypo- pygium large, truncate apically. Colour. — Dark brown (blackish according to the original description); base of 2nd tergite yellowish-brown; scapus and pedicellus ven- trally, 3rd and 4th antennal segments, mandi- bles, palpi, and legs, yellowish; wings hyaline; pterostigma light brown. Lectotype in U.S. National Museum, Wash- ington: “Ex Phytomyza sp. on Columbine”, “Lucerne, Switzerland”, “G. de Baume-Pluvi- nell Col.”, “PHO 14, La Baume. 3 Rd’Ulm Pa- ris”, “Adulura gahani B.-P., det. Gahan Ce- vest(?)/ Basis of original identification by A.B.G.” This ® is here designated as lectotype. One paralectotype, 9, topotypic, in same col- lection; it has the ovipositor sheath retracted, not protruding; antennal segments 29; length of fore wing 2.2 mm, of body 1.2 mm. Note. — Parasite of Phytomyza minuscula Goureau and P. aquilegiae Hardy (Agromyzi- dae); recorded from Poland and Switzerland. Dapsilarthra (Heterolexis) levisulca Griffiths Dapsilarthra levisulca Griffiths, 1968a: 10. Shenefelt, 1974: 989. Only known from the type-series from Yu- goslavia, partly reared from Phytomyza bellidi- na Hering (Agromyzidae). * Dapsilarthra (Heterolexis) balteata (Thomson) Alysıa balteata Thomson, 1895: 2288. Dapsilarthra balteata; Königsmann, 1972: 21. Shene- felt, 1974: 988. Hendrickson & Barth, 1979: 168, 1170), This common species has been reared from many leaf-mining species, belonging to the gen- era Agromyza, Cerodonta, Phytomyza, and Liriomyza (Agromyzidae). The record from Pegomya (Anthomyiidae) needs to be confirm- ed. Specimens examined: 91 © and 3 d from the Netherlands (Waarder, Driebergen, Wijster, Lienden, Ouddorp, Weesp, Kadier en Keer). Variation: antennal segments 39—48; precoxal sulcus smooth or slightly sculptured; 1st tergite black, dark brown or brownish; head behind eyes parallel-sided to strongly ane posteri- orly; 2nd and 3rd tergites of metasoma fre- quently medially brownish-yellow, but colour of metasoma behind 1st tergite varies from largely piceous and dark brown to light yellow- ish and brown. * Dapsilarthra (Heterolexis) subtilis (Foerster) comb. nov. Heterolexis subtilis Foerster, 1862: 268. Shenefelt, 1974: 992. Dapsilarthra testacea Griffiths, 1968a: 9—10; Königs- mann, 1972: 22. Shenefelt, 1974: 991. Syn. nov. In the Foerster Collection (Zoologisches Mu- seum, Berlin) there are two males on one pin (‘25/2127 “Aachen, Juni, Erst swiss Frst”), of which the d nearest to the pin is here designated as lectotype. Both are typical speci- mens of Dapsilarthra testacea. The 1st and 2nd segments of the metasoma are yellowish; the lectotype has 31, and the paralectotype 32 an- tennal segments. This species is seldom collected; reported from Germany and Austria. In the Leiden col- lection are specimens from the Netherlands (Ede and Putten). Subgenus Mesocrina Foerster stat. nov. Mesocrina Foerster, 1862: 266. Shenefelt, 1974: 996. Pseudomesocrina Königsmann, 1959b: 611. Shenefelt, 1974: 1018. Syn. nov. Diagnosis. — Antennal segments about 38 (only known for type-species); medio-posterior groove of mesoscutum present (fig. 49); precox- al sulcus absent (fig. 43) or impressed (fig. 26); vein CU1b and apical half of vein 2-1A of fore wing present, resulting in a closed 1st subdiscal cell (fig. 28); vein 3-CU1 of fore wing longer than vein CU1b (fig. 51); pterostigma largely el- liptical (figs. 28, 45); vein r of fore wing emerg- ing submedially from elliptical part of pterostig- ma; vein m-cu of hind wing present (fig. 28); vein m-cu of fore wing shortly postfurcal (fig. 45); dorsope small (fig. 37) or large (fig. 50). Distribution. — Contains one West Palaearc- tic species and one species from the Himalayan area. VAN ACHTERBERG: Dapsilarthra and Mesocrina 9 * Dapsilarthra (Mesocrina) indagatrix (Foerster) comb. nov. (figs. 43—55) Mesocrina indagatrix Foerster, 1862: 266. Shenefelt, 1974: 996. Mesocrina venatrix Marshall, 1895a: 429— 430, fig. Shenefelt, 1974: 1018— 1019. Syn. nov. The interpretation of this obviously scarcely collected species has been hindered by the loss of the type (Königsmann, 1959b: 611), but in my opinion the fairly complete original descrip- tion by Foerster enables a correct identification of the species to be made. This interpretation is close to that of Marshall (1895a: 430) and dis- agrees with the statements made by Königs- mann (1959b: 610). Kônigsmann’s argument that Foerster was unlikely to have overlooked the strongly compressed metasoma of the ® is not conclusive, because Foerster’s specimen may have been a d, which has a normally de- pressed metasoma. Because no closely related species are known, there is no reason to doubt the synonymy of venatrix and indagatrix. The holotype of D. veratrix (Marshall) has been ex- amined; it is a ® with the typical compressed metasoma. To fix the type-species of the genus Mesocrina Foerster I designate here the d de- scribed and figured below as neotype of Meso- crina indagatrix Foerster, 1862. Neotype, d, length of body 3,5 mm, of fore wing 3,9 mm. Head. — Antennal segments 21 (apical seg- ments missing), length of 3rd segment 1.7 times 4th segment, length of 3rd and 4th segments 4.5 and 2.4 times their width, respectively; penulti- mate segment of antenna of neotype missing, but in 2 from Ede (Netherlands), length 1.7 times its width (fig. 55); length of maxillary palp 1.2 times height of head; dorsal length of eye subequal to dorsal length of temple (fig. 53); POL : © ocellus : OOL = 12 : 7 : 20; frons smooth, vertex smooth and with medial suture: face rather flat and with some punctulation (fig. 44); anterior tentorial pits large and far removed from eyes, distance to eyes 1.5 times maximal diameter of pit; clypeus strongly convex, smooth, its apical margin narrowly differ- entiated and straight medially; malar space short, eye almost touching base of mandible (fig. 46); mandible slightly widened dorsally, 2nd tooth sharp and longer than both lateral, more obtuse teeth (figs. 46, 48). Mesosoma. — Length of mesosoma 1.4 times its height; pronope absent (fig. 49); side of pro- notum with a crenulate medial groove, rest smooth (fig. 43); precoxal sulcus absent; meso- pleuron smooth; pleural sulcus narrowly crenu- late ventrally, dorsally largely smooth (fig. 43); metapleural flange absent; metapleuron smooth, but crenulate rugose-reticulate; notauli only an- teriorly impressed and crenulate (fig. 49); meso- scutum largely glabrous, only anteriorly densely setose; medio-posterior groove long and nar- row, rather shallow (fig. 49); scutellar sulcus with one weak longitudinal carina; scutellum sparsely punctulate; side of scutellum smooth; surface of propodeum anteriorly and medio- posteriorly finely rugulose, rest mainly smooth, medial carina and areola absent; posterior part of propodeum not differentiated from antero- dorsal part; propodeal spiracle rather small, round, somewhat protruding and in front of middle of propodeum. Wings. — Fore wing: r much shorter than width of pterostigma (fig. 45); r : 3-SR : SR1 = 3:21:41; SR1 straight; 1-CU1 : 2-CU1 = GMA SREBESRE Em 37222039: lstasub- discal cell somewhat widened apically. Hind wing: cu-a straight; marginal cell absent apical- ly. Legs. — Hind coxa punctulate; tarsal claws rather slender (fig. 47); length of femur, tibia and basitarsus of hind leg 4.8, 8.2, and 6.2 times their width, respectively; length of hind tibial spurs 0.3 and 0.4 times hind basitarsus. Metasoma. — Length of 1st tergite 2.0 times its apical width, its surface basally smooth, its posterior half rugose (fig. 50), basally concave, medially convex; dorsal carina of 1st tergite pre- sent in basal 0.9 of tergite, but apically superfi- cial; dorsope large (fig. 50); hypopygium of neotype rather short, truncate apically; 2 from Ede has length of ovipositor sheath 0.13 times fore wing, ovipositor slender, with no distinct teeth or nodus, only slightly widened subapi- cally (fig. 54); hypopygium medium-sized and subtruncate apically; metasoma of neotype d depressed (fig. 43), of 2 strongly compressed (fig. 54). Colour. — Black; metasoma behind 1st ter- gite, wing veins, and tegulae, brownish-yellow; hind tarsus and apex of hind tibia somewhat in- fuscated; wing membrane subhyaline. Neotype in the Zoologische Staatssammlung, München: “Harthausen b. München, A. 27.9.68, Haeselb.”. Additional specimens exam- ined: 1 9, Museum Budapest, holotype of Mesocrina venatrix Marshall (“venatrix M., Coll. Marshall”, “Pseudomesocrina venatrix 10 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 (Marsh), det. Königsmann”) from England, which is conspecific with the neotype of inda- gatrix. Length of fore wing 3.7 mm, and length of ovipositors sheath about 0.12 times fore wing. First metasomal tergite missing (figured by Königsmann), rest of metasoma on separate slide, and antenna incomplete. 1 ©, Rijksmu- seum van Natuurlijke Historie, Leiden: Nether- lands (“Ede, trap [= Malaise-trap], 22—28.1x, Van Rossem”). Length of fore wing 4.2 mm, of body 4.3 mm; metanotum rather coarsely sculp- tured; length of 1st tergite 1.9 times its apical width. Parasite of Pegomya and Amaurosoma spp. (Anthomyiidae and Scatophagidae, re- spectively). Dapsilarthra (Mesocrina) dalhousiensis (Sharma) comb. nov. (figs. 26—37) Acrobela dalhousiensis Sharma, 1978: 127—128, figs. 46. Holotype, d, length of body 5.4 mm, of fore wing 4.7 mm. Head. — Remaining antennal segments 11, apical segments missing, scapus compressed; length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 3.0 and 2.6 times their width, respectively; length of maxillary palp 1.2 times height of head; dorsal length of eye 1.2 times temple; temple smooth and subpa- rallel-sided (fig. 31); POL : © ocellus : OOL = 8:4:14; frons flat and smooth; vertex with medial groove; face unevenly convex, largely smooth, with some punctulation (fig. 30); ante- rior tentorial pits large, far removed from eyes (fig. 30); clypeus convex, somewhat punctulate, rugulose near dorsal margin, its apical margin not distinctly differentiated medially, convex; malar space absent, eye touching base of mandi- ble (fig. 27); mandible widened dorso-apically, finely rugose medially, 3rd tooth large and lobe- shaped; ventral margin of mandible with small incision (fig. 29) (but without ventral protrud- ing lamella); 2nd tooth of mandible large, longer than lateral teeth, acute apically (figs. 27, 29). Mesosoma. — Length of mesosoma 1.2 times its height; pronope absent; side of pronotum medially crenulate, dorsally largely smooth, ventrally microsculptured (fig. 26); epicnemial area ventrally rugose (fig. 26); precoxal sulcus present in anterior 0.7 of mesopleuron, densely reticulate-rugose; rest of mesopleuron smooth; pleural sui: us rather shallow and narrow, finely crenulate (tig. 26); episternal scrobe large; metapleural flange small, obtuse apically; metapleuron finely rugose; notauli only ante- riorly impressed, finely crenulate (fig. 34); mesoscutal lobes smooth, medially glabrous; medio-posterior groove of mesoscutum slender, also anteriorly with a shallow groove (fig. 34); scutellar sulcus deep, long, with 8 longitudinal carinae; scutellum smooth; sides of scutellum rugose; metanotum somewhat protruding me- dially (fig. 26); posterior part of propodeum dif- ferentiated and much longer than antero-dorsal part (fig. 26); surface of propodeum densely ru- gose anteriorly, with some short apical carinae and indistinct sculpture posteriorly, medial cari- na and areola absent; propodeal spiracle rather small, round, and in front of middle of propo- deum. Wings. — Fore wing: r subequal to width of pterostigma (fig. 28); r : 3-SR : SR1 = 8: 28: 61; SR1 straight; 1-CU1 : 2-CU1 = 2: 20; 2- SR: 3-SR : r-m= 18: 28: 13; 3-SR of left wing with stub (fig. 36), indistinct in right wing (fig. 28). Hind wing: cu-a straight; marginal cell narrowed apically. Legs. — Hind coxa smooth; tarsal claws rath- er slender (fig. 32); length of femur, tibia and basitarsus of hind leg 3.9, 8.9, and 6.4 times their width, respectively; length of hind tibial spurs 0.3 times hind basitarsus, subequal. Metasoma. — Length of 1st tergite equal to its apical width, its surface largely striate (fig. 37), flat basally, and medially slightly convex; dorsal carinae distinct in front of spiracles, short (fig. 37); dorsope small; metasoma depressed; hypopygium medium-sized and truncate apical- ly. Colour. — Black; mesonotum, pronotum (largely), mesosternum and mandible, brown- ish-red; legs (except for middle and hind tarsi, apex of hind tibia and femur), palpi, tegulae, scapus, and pedicellus, yellowish; pterostigma, most wing veins, metasoma (dorsally rather blackish), fore and middle tarsi (except telotar- si), apices of hind femur and tibia partly, dark brown; wing membrane slightly infuscate. Holotype (and only known specimen) in Gupta Collection, Delhi: “India: H.P. Dalhou- sie, 2132 m, 22.1x.1971, Sykh. Dev. No. JD 158”, “Holotype Acrobela dalhousiensis V. Sharma, 1975”. Biology unknown. Subgenus Dapsilarthra Foerster Dapsilarthra Foerster, 1862: 267. Shenefelt, 1974: 986—991. Marsh, 1979: 222. Paraorthostigma Königsmann, 1972: 25—26, 1 fig. Syn. nov. VAN ACHTERBERG: Dapsilarthra and Mesocrina 11 Diagnosis. — Antennal segments 25—41: medio-posterior groove of mesoscutum present (fig. 67); precoxal sulcus present (fig. 81) or al- most absent (fig. 72); vein CU1b and apical half of vein 2-1A of fore wing present, resulting in a closed 1st subdiscal cell (fig. 58); vein 3-CU1 of fore wing shorter than vein CU1b (fig. 57) or equal (fig. 89); pterostigma (sub)linear, with its sides (sub)parallel (figs. 58, 82); vein r of fore wing emerging between base and middle of pte- rostigma (fig. 70, 82); vein m-cu of hind wing absent (fig. 70); dorsope (rather) large (figs. 66, 79,91). Distribution. — Five Palaearctic species, two of which also occur in the Nearctic region. Dapsilarthra (Dapsilarthra) rufiventris (Nees) (figs. 90, 92) Bassus rufiventris Nees, 1814: 213. Dapsilarthra rufiventris; Königsmann, 1972: 22. Shenefelt, 1974: 989— 990. Marsh, 1979: 222. Alysia flaviventris Haliday, 1838: 240. Dapsilarthra flaviventris; Shenefelt, 1974: 990. Alysia graalicornis Thomson, 1895: 2283. Shenefelt, 1974: 990. Phaenocarpa ochrogaster Szépligeu, 1898: 393, 406. Shenefelt, 1974: 1012. Dapsilarthra fuscula Griffiths, 1968b: 65—66. Shene- felt, 1974: 989. Syn. nov. Because the differences between rufiventris and fuscula seem to be completely restricted to differences in colour and I have examined inter- mediately coloured specimens from Romania and Bulgaria, I consider fuscula to be only a melanistic form of rufiventris. Total specimens examined; 20 ® and 5 d (nominate form) and 3 © of f. fuscula Griffiths, all in the Rijksmuseum van Natuurlijke Historie, Leiden, except for the Bulgarian specimens from the Zaykov Collec- tion, Plovdiv. The specimens originate from the Netherlands (Waarder, Breukelen, Thorn (ex Trilobomyza flavifrons Meigen in Melandrium rubrum Garcke), Lienden (ex ? Napomyza glechomae Kaltenbach in Glechoma hederacea L.), Wijster), Switzerland (Schwyz, Brunnen), Romania (Sinaia, S. Carpathians, 1300-1400 m), and Bulgaria (Pamporovo, Rhodopi Mountains (f. fuscula); Velinograd, Rhodopi Mountains (nominate from)). Parasite of Liriomyza, Phyto- myza, and Trilobomyza spp. (Agromyzidae) in herbaceous plants. Dapsilarthra (Dapsilarthra) isabella (Haliday) (figs. 81—89, 93, 94) Alysia isabella Haliday, 1838: 240. Dapsilarthra isabella; Shenefelt, 1974: 989. Redescribed from a ® in the Marshall Collec- tion from Barnstaple, England. Precise length of body unknown, because part of the metasoma has been removed; length of fore wing 3.1 mm. Head. — Antenna about twice length of body, with 41 segments (fig. 81), length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 3.7 and 3.0 times their width, respectively; length ot penultimate antennal segment 2.5 times its width (fig. 83); length of maxillary palp equal to height of head; dorsal length of eye equal to dorsal length of temple (fig. 87); temples subparallel-sided; POL: © ocellus OOR A MEEO Mons latin smooth; vertex smooth, without medial groove; face rather convex, laterally coriaceous (fig. 85); anterior tentorial pits large, far removed from eyes (fig. 85); clypeus convex and smooth, its apical margin straight medially and with a nar- ‚row rim; malar space almost absent; mandible smooth, slightly widened apically (figs. 84, 88), its middle tooth acute, somewhat longer than lateral, more lobe-shaped teeth. Mesosoma. — Length of mesosoma 1.3 times its height; pronope absent; side of pronotum smooth, except for a crenulate medial groove (fig. 81); epicnemial area largely crenulate; pre- coxal sulcus only medially distinctly impressed and crenulate (fig. 81); rest of mesopleuron smooth; pleural sulcus finely crenulate; epister- nal scrobe medium-sized; metapleural flange medium-sized, rather acute apically; metapleu- ron largely smooth (only ventrally somewhat rugose), and with a pit close to the mesopleuron (fig. 81); notauli only anteriorly impressed and finely crenulate, but absent on dorsal part of mesocutum (figs. 81, 94); mesoscutum largely glabrous, smooth; medio-posterior pit of meso- scutum elliptical and rather deep (fig. 94); scu- tellar sulcus deep, finely crenulate; scutellum smooth; side of scutellum smooth; metanotum posteriorly rather bluntly protruding (fig. 81); surface of propodeum densely and finely ru- gose; medial carina and areola of propodeum absent; posterior part of propodeum not differ- entiated from antero-dorsal part; propodeal spi- racle round, rather small and protruding, in front of middle of propodeum. Wings. — Fore wing: r very short, shorter than width of pterostigma (fig. 82); pterostigma not reaching beyond middle of marginal cell and rather wide (fig. 82); m-cu shortly antefurcal; 3- CU1 subhorizontal (fig. 89): marginal cell al- most attaining wing apex; r: 3-SR : SRI = 2:26:58; SR1 sinuate; 1-CU1 :2-CU1 = 12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 1:11; CUlb subvertical and equal to 3-CUI (fig. 89); 2-SR : 3-SR : r-m = 12:26 : 6. Hind wing: cu-a straight; marginal cell absent apical- ly. Legs. — Hind coxa smooth; tarsal claws slen- der (fig. 86); length of femur, tibia and basitar- sus of hind leg 4.7, 9.2, and 4.7 times their with, respectively; length of hind tibial spurs 0.3 times hind basitarsus, subequal. Metasoma. — Length of 1st tergite 1.6 times its apical width, its surface coarsely punctate- rugose (fig. 91), antero-medially concave, and medially convex; dorsal carina of 1st tergite ab- sent; distance between dorsope far more than their width (fig. 91); ovipositor missing, but according to Marshall (1895b: 366) subexserted. Colour. — Black or blackish-brown; antenna dark brown, but basally partly yellowish; palpi, legs (except for the infuscated apical half of hind tibia and base of hind coxae), and tegulae, brownish-yellow; wing membrane hyaline; pte- rostigma, wing veins, and 2nd tergite, brown. Type-series (obviously consisting of one male received from Walker, from the surroundings of London) could not be found in the Haliday Collection (Dublin) and is probably lost. The redescription is based on the 2 in the Marshall Collection (British Museum, London): “Eng- land, ND, Barnstaple, Marshall Coll., B.M. NO HOP Segel Bal I BML 1250, Under Adelura isabella Hal”, “prep. DSN Alysia isabella Haliday has not been recog- nized since Marshall (1895a, 1895b). Because the redescribed © fully agrees with Haliday’s fairly complete description and Marshall is the first revisor, Marshall’s interpretation has to be accepted. The biology is unknown. Dapsilarthra (Dapsilarthra) sylvia (Haliday) Alysia sylvia Haliday, 1839: 25. Dapsilarthra sylvia; Königsmann, 1972: 22. Shenefelt, 1974: 990—991. This is a sparsely collected species, not yet found in the Netherlands. Parasite of Agromyza, Cerodontha, Phytomyza, and Trilo- bomyza spp. (Agromyzidae) in herbaceous plants. Dapsilarthra (Dapsilarthra) tirolensis (Königsmann) comb. nov. Paraorthostigma tirolense Königsmann, 1972: 26, 1 fig. Owing to the kindness of Dr. E. Haeselbarth (München) I was able to examine the holotype of Paraorthostigma tirolense (8, Haeselbarth Collection: “Unser Frau in Schnals, Südtirol, 1600 m, 14.7.66, Hbth”, “Holotypus”, “Pa- raorthostigma tirolense n.sp”). It belongs to the subgenus Dapsilarthra, is close to rufiventris, and differs mainly by the dark legs, somewhat longer vein r (fig. 38), short antenna (fig. 39) and shorter marginal cell of fore wing (fig. 38 versus fig. 90). Dapsilarthra (Dapsilarthra) carpathica spec. nov. (figs. 56—67) Holotype, ?, length of body 1.8 mm, of fore wing 2.8 mm. Head. — Antennal segments 34 (but apical segments missing), length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th seg- ments 4.7 and 4.2 times their width, respective- ly; scapus rather slender (fig. 65); length of maxillary palp 1.1 times height of head; dorsal length of eye 1.6 times temple; temple roundly narrowed posteriorly (fig. 63); POL : © ocel- lus: OOL = 7: 4: 12; frons smooth, with small medial pit; vertex smooth, with shallow groove (fig. 63); face smooth, rather flat; ante- rior tentorial pits large, less robust than in ap, oval, not well differentiated from face, distance to eye about % of maximum diameter of pit (fig. 59); clypeus convex and smooth, its apical margin slightly differentiated and convex me- dially (fig. 59); malar space absent; sides of mandible parallel, rather slender (figs. 61, 62), 2nd tooth acute, longer than both, more lobe- shaped lateral teeth, without distinct carinae. Mesosoma. — Length of mesosoma 1.2 times its height; pronope absent; side of pronotum largely smooth, only medially crenulate (fig. 56); precoxal sulcus rather slender, superficially sculptured, anteriorly and posteriorly absent (fig. 56); rest of mesopleuron smooth; pleural sulcus dorsally largely smooth, ventrally finely crenulate (fig. 56); episternal scrobe slender; metapleural flange medium-sized, directed for- wards; metapleuron ventrally rugulose, dorsally largely smooth; notauli anteriorly impressed and crenulate (fig. 67), but posterior half absent; mesoscutum largely smooth and glabrous; me- dio-posterior groove of mesoscutum medium- sized, deep; scutellar suture with one longitudi- nal carina; scutellum (including sides) smooth; metanotum posteriorly coriaceous and medial VAN ACHTERBERG: Dapsilarthra and Mesocrina 13 carina short (fig. 67); surface of propodeum re- ticulate-rugulose; medial carina and areola of propodeum absent; posterior part of propo- deum not differentiated from antero-dorsal part; propodeal spiracle rather small, round, in front of middle of propodeum. Wings. — Fore wing: r longer than width of pterostigma (fig. 58); pterostigma reaching be- yond middle of marginal cell and slender; m-cu shortly postfurcal (fig. 58); 3-CU1 oblique and shorter than CUI1b (fig. 57); 1-R1 less curved than in api (fig. 58 versus fig. 70); r:3- SR : SRI = 13 : 49 : 119; SRI almost straight; 1-CU1 :2-CU1 = 7:22; 2-SR : 3-SR : r-m = 17:49:16. Hind wing; cu-a almost straight; marginal cell parallel-sided apically. Legs. — Hind coxa smooth; tarsal claws rath- er slender (fig. 64); length of femur, tibia, and basitarsus of hind leg 6.4, 10.4 and 6.3 times hind basitarsus, respectively. Metasoma. — Length of 1st tergite 2.2 times its apical width, its surface rugulose, flat medio- basally, rest convex; dorsal carinae of 1st tergite present in front of spiracles; laterope absent; dorsope medium-sized (fig. 66); ovipositor straight, its apex somewhat curved downwards, without notch or nodus, with some ventral teeth; length of ovipositor sheath 0.07 times fore wing, sheath somewhat widened subapi- cally; hypopygium large, truncate apically (fig. 56). Colour. — Black; tegulae basally dark brown, apically yellowish; palpi whitish; face, antenna (except 4 basal segments) and meta- soma behind ist tergite, dark brown; wing veins, and pterostigma, brown; mandibles, la- brum, legs, and 4 basal segments of antenna, largely yellowish; apex of hind tibia and telotar- si somewhat infuscated; wing membrane hya- line. Holotype (and only known specimen) in the Rijksmuseum van Natuurlijke Historie, Leiden: “Museum Leiden, Romania, Sinaia, S. Carpathi- ans, 1300-1400 m, 8—10.viii.1978, C. van Achterberg”. Dapsilarthra (Dapsilarthra) apii (Curtis) (figs. 68—80) Alysia apu Curtis, 1826: 141, figs. Dapsilarthra api; Shenefelt, 1974: 987. Alysia laevipectus Thomson, 1895: 2288. Orthostigma americana Brues, 1907: 59—60. Syn. nov. Dapsilarthra americana; Fischer, 1973: 256—258, fig. 8. Redescribed after a female from Austria (Aschbach, Tirol), which is compared and con- specific with the lectotype of Alysia apu Curtis; length of body 3.6 mm, of fore wing 4.8 mm. Head. — Antennal segments 50, 3rd segment equal to 4th segment, length of 3rd and 4th seg- ments both 4.0 times their width; scapus very robust, compressed (fig. 75); penultimate seg- ment of antenna 3.3 times its width (fig. 71); length of maxillary palp 1.3 times height of head; dorsal length of eye 1.2 times temple (fig. 73); temples subparallel-sided, rounded posteri- orly; POL : @ ocellus; OOL = 6 : 3 : 12; frons smooth, except for a shallow medial impression (fig. 74); vertex smooth, with medial groove: face rather flat, largely shiny-coriaceous; ante- rior tentorial pits large, not well differentiated from face, distance to eye about equal to maxi- mum diameter of pit (fig. 74); clypeus convex, punctulate, its apical margin superficially differ- entiated, weakly convex (fig. 74); malar space absent; mandible robust, widened apically (figs. 68, 69), with 2nd tooth large and acute, longer than both large and more lobe-shaped lateral teeth. Mesosoma. — Length of mesosoma 1.2 times its height; pronope absent, except for a narrow slit-shaped depression (fig. 78); side of prono- tum dorsally smooth, its ventral half largely ru- gose (fig. 72); epicnemial area ventrally crenu- late, dorsally smooth; only anterior half of pre- coxal sulcus impressed, smooth (fig. 72); rest of mesopleuron smooth; pleural sulcus very finely crenulate; episternal scrobe linear (fig. 72); metapleural flange medium-sized, directed for- wards; metapleuron rugulose, but medially smooth; notauli anteriorly impressed (fig. 78), their posterior half absent; between notauli two rows of punctures, rest of mesoscutal lobes smooth; medio-posterior groove medium-sized, droplet-shaped (fig. 78); scutellar sulcus finely crenulate (but in lectotype of ap: and in speci- men from Sölden with one medial carina only); scutellum and its sides smooth; metanotum with weak superficial crest (fig. 78), slightly protrud- ing dorsally (fig. 72); surface of propodeum densely rugulose, with no carina; posterior part of propodeum not differentiated from antero- dorsal part (fig. 72), with a scarcely differ- entiated oval areola; propodeal spiracle rather small, round and in front of middle of propo- deum. Wings. — r longer than width of pterostigma (fig. 70); pterostigma reaching beyond middle 14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 of marginal cell, and slender; m-cu shortly post- furcal; marginal cell ends close to wing apex (fig. 70); 1-R1 strongly curved; r: SRI EPS RSS SR Curved IE CUIR NDE CURE CRC UbMlonsenelrannss GU CUI oblique (fig. 80); 2-SR : 3-SR : r-m = 10: 25 : 8. Hind wing: cu-a reclivous; marginal cell parallel-sided apically. Legs. — Hind coxa smooth; tarsal claws rath- er slender (fig. 76); length of femur, tibia, and basitarsus of hind leg 5.6, 10.8, and 6.2 times their width, respectively. Metasoma. — Length of 1st tergite 1.7 times its apical width, its surface rugulose, flat medio- basally, medially convex; dorsal carinae present in basal third, in front of spiracles (fig. 79); laterope deep and large (fig. 72); dorsope large, their distance apart much more than diameter of dorsope (fig. 79); ovipositor straight; ovipositor sheath 0.05 times fore wing, long setose (fig. 72); hypopygium large, truncate apically. Colour. — Black; antenna, 2nd tergite, cly- peus, hind tibia and tarsus largely, wing veins (largely), and tegulae, dark brown; pterostigma brown; palpi, labrum, and rest of legs, yellow- ish; wing membrane subhyaline. Redescribed ® in the Rijksmuseum van Na- tuurlijke Historie, be “Austria, Tirol, Aschbach, 1400 m, 16.viii.1975, C. J. Zwak- hals”. Lectotype of Alysia api Curtis, 1826, here designated: 9, Curtis Collection (Melbourne): 0, “an, Type”, “Type of Alysia api Curt., G. Nixon det. 1948” [unpublished]. The Berre of apu given in the original description is fairly correct, but the separately figured wing (fig. 9) obviously belongs to another species (vein 3-SR shorter than 2-SR!). Curtis had more than one specimen, because he used the pleural form (“For specimens of this insect and their history I am indebted to a lady... .”); the only remaining specimen has to be accepted as lecto- type. The holotype of Orthostigma americana Brues (Milwaukee Public Museum, Milwaukee) is a female of rather small size and with whitish palpi, not, however, essentially differing from api. It bears the following labels: “Milw., Co. Nm selen 1008, Mme, Orion doer Bos? “22062”. Additionally exam- ined 1 2 (Rijksmuseum van Natuurlijke Histo- rie, Leiden). “Austria, Tirol, Sölden, 1800 m, 17.viii.1975, C. J. Zwakhals”. It has vein SR1 of the fore wing less curved, while the palpi, sca- pus, pedicellus, and hind tibia (except apex) are 3-SR :, more reddish than ot the figured specimen. The specimen reported from Driebergen by Snellen van Vollenhoven (1873: 195) belongs to D. bal- teata (Thomson). Parasite of Philiphyllla heraclei L. (Tephriti- dae) and Psila rosae F. (Psilidae). Species excluded from Dapsilarthra Dapsilartha tenuicornis (Foerster) Opisendea tenuicornis Foerster, 1862: 266. Dapsilarthra tenuicornis; Shenefelt, 1974: 991. Pentapleura angustula Haliday, 1838: 229 Syn. nov. The lectotype (9, Zoologisches Museum, Berlin) of O. tenuicornis has the ventral protu- berance of the mandible comparatively weakly developed, and vein 3-SR of fore wing is somewhat longer than vein 2-SR; however, both features are not uncommon among Penta- pleura spp. Dapsilarthra sulcifera Papp Dapsilarthra sulafera Papp, 1967: 209, figs. Shenefelt, 1974: 990. Opius comatus Wesmael, 1835; Fischer, 1974: 48. As pointed out by Fischer (1974) this species was wrongly assigned to the Alysiinae. Dapsilarthra barthii (Brues) Asobara barthii Brues, 1907: 57—58. Dapsilarthra barthu; Fischer, 1973: 258—260, figs. 9—10. I have examined the lectotype selected by Fischer (9, Milwaukee Public Museum, Mil- waukee); it proved to be a Phaenocarpa species with stout antenna (however, 3rd antennal seg- ment distinctly shorter than 4th segment). Phaenocarpa barthu (Brues, 1907) is a new combination, except for the (accidental?) use of this combination in the text to figs. 9—10 in Fischer, 1973. ACKNOWLEDGEMENTS I wish to express my sincere thanks to the fol- lowing persons for the loan of (type-)specimens and/or gift of unidentified specimens: Dr. T. van Dijk (Wijster), Dr. M. Fischer (Wien), Mr. A. van Frankenhuyzen (Wageningen), Mr. M. J. Gijswijt (Ankeveen), Prof. Dr. V. K. Gupta (Gainesville), Dr. E. Haeselbarth (München), Mr. T. Huddleston (London), Mr. K. J. Huis- man (Melissant), + Dr. E. K6nigsmann (Berlin), Mr. B. J. Lempke (Amsterdam), Dr. P. M. VAN ACHTERBERG: Dapsilarthra and Mesocrina 15 Marsh (Washington), Dr. A. Neboiss (Mel- bourne), Dr. G. R. Noonan (Milwaukee), Dr. J. P. O’Connor (Dublin), + Dr. S. J. van Oost- stroom (Oegstgeest), Dr. J. Papp (Budapest), Mr. G. van Rossem (Ede), Prof. Dr. J. van der Vecht (Putten), Dr. A. Zaykov (Plovdiv), Drs. CM Zalkhals(Arkel)s and Drs) RAW. R. Zwart (Wageningen). REFERENCES Achterberg, C. van, 1979. A revision of the subfamily Zelinae auct. (Hym., Braconidae). — Tijdschr. Ent. 122: 241—479, figs. 1—900. Baume-Pluvinel, G. de la, 1915. Evolution et formes larvaires d’un braconide (Adelura gahani n.sp.), parasite interne de la larve d’un Phytomyzinae (diptère). — Archs Zool. exp. gén. 55: 47—59, 11 figs. Bischoff, H., 1932. Hymenoptera (excl. Formicidae und Cynipidae) der Deutschen Limnologischen Sunda-Expedition. — Arch. Hydrobiol., Suppl. 9: 738—746. Brues, C. T., 1907. Notes and descriptions of North American parasitic Hymenoptera. III. — Bull. Wis. nat. Hist. Soc. 5: 54—62. Curtis, J., 1826. British Entomology, pt. 3, Hymeno- ptera. London. Fischer, M., 1966. Studien über Alysiinae (Hym., Bra- conidae). — Annln naturh. Mus. Wien 69: 177— 205, figs. 1—24. ——, 1971. Untersuchungen über die europäischen Alysiini mit besonderer Berücksichtigung der Fauna Niederösterreichs (Hym., Braconidae). — Polskie Pismo ent. 41: 19—160, figs. 1—56. ——, 1973. Redeskriptionen von Alysiinen (Hym., Braconidae). — Annln naturh. Mus. Wien 77: 245—261, figs. 1—10. , 1974. Studien an Alysiinen-Typen (Hym., Bra- conidae, Alysiinae). — Z. Arb. ost. Ent. 25: 47— 51, figs. 1—3. Foerster, A., 1862. Synopsis der Familien und Gat- tungen der Braconen. — Verh. naturh. Ver. preuss. Rheinl. 19: 225—288. Griffiths, G. C. D., 1968a. The Alysiinae (Hym., Bra- conidae) parasites of the Agromyzidae (Diptera). V. The parasites of Liriomyza Mik and certain small genera of Phytomyzinae. — Beitr. Ent. 18: 5—62, figs. 171185. ——, 1968b. Id.VI. The parasites of Cerodontha Rondani s.l. — Beitr. Ent. 18: 63—152, figs. 186—209. Haliday, A. H., 1833. An essay on the classification of the parasitic Hymenoptera of Britain, which cor- respond with the Ichneumones minuti of Lin- naeus. — Ent. Mag. 1: 259—276. , 1838. Essay on the classification of parasitic Hymenoptera. — Ent. Mag. 5: 209—248. —, 1839. Hymenoptera Britannica: Alysia, p. 1— 28. — Ballière, London. Hendrickson, R. M., & S. E. Barth, 1979. Introduced parasites of Agromyza frontella (Rondani) in the U.S.A. —N.Y. ent. Soc. 87: 167—174. Königsmann, E., 1959a. Revision der palaarktischen Arten der Gattung Dapsilarthra. 1. Beitrag zur systematischen Bearbeitung der Alysiinae (Hym., Braconidae). — Beitr. Ent. 9: 580—608, figs. 1— 9, pls. 33—38. — — 1959b. Revision der paläarktischen Arten der Gattung Mesocrina. 2. Beitrag zur systematischen Bearbeitung der Alysiinae (Hym., Braconidae). — Beitr. Ent. 9: 609— 619, figs. 1—6. — 1972. Zur Kenntnis verschiedener Gattungen der Alysiinae nebst Beschreibung der neuen Gat- tung Paraorthostigria (Hym., Braconidae). — Dtsch. Ent. Z. (N.F.) 19: 21—30, 1 fig. Marsh, P. M., 1979. Braconidae, p. 144-313. In: K.V. Krombein et all. Catalog of Hymenoptera in America North of Mexico, 1: i-xvi, 1—1198. Marshall, T. A., 1895a. Species des Hyménoptères d’Europe & d’Algerie. V. Les Braconides, p. 401—480, pls. 12—14. ——, 1895b. A Monograph of British Braconidae. VI. — Trans. ent. Soc. London, p. 363—398, fıgs. 1212: ——, 1898. Species des Hyménoptères d'Europe & d’Algérie, V.bis. Les braconides (supplément), p. 145—288, pls. 7—12. Nees von Esenbeck, C. G. 1814. Ichneumonides adsciti, in genera et familias divisi. — Mag. Ges. Naturf. Fr. Berlin 6: 183—221. Papp, J., 1967. Ergebnisse der Zoologischen For- schungen von Dr. Z. Kaszab in der Mongolei. 101. Braconidae 1. — Acta Zool. Acad. sci. hung. 13: 191—226. Sharma, V., 1978. Taxonomic Studies on Indian Bra- conidae (Hym.). — Oriental Ins. 12: 123—132, figs. 1—14. Shenefelt, R. D., 1974. Hymenopterorum Catalogus (nov. ed.) Part 11, Braconidae 7: 937— 1113. Junk, ’s-Gravenhage. Snellen van Vollenhoven, S. C., 1873. Nieuwe naam- lijst van Nederlandse vliesvleugelige insecten (Hym.). — Tijdschr. Ent. 16: 147—220. Strand, E. 1928. Miscellanea nomenclatorica zoolog- ica et palaeontologica. — Arch. Naturgesch. 92 (A): 30—75. Szépligeu, G. V., Beiträge zur Kenntnis der ungari- schen Braconiden, 3. Theil. — Természetr. Füz. 21: 381— 408. Telenga, N. A., 1935. Beträge zur Kenntnis der Trib- us Alysiini (Braconidae, Hym.) aus der USSR. — Konowia 14: 186—190. Thomson, C. G. 1895. LII. Bidrag till Braconidernas kannedom. — Opusc. ent. 20: 2141—2339. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 1, 1983 16 15] UO ıy3ıs [ny ‘oygrpuew ‘Ey ‘41001 pig uo quais [my suraA Jo Jreiop ‘g ‘10adse Jesıop ‘9113101 18] ‘4 !sZurm © < 9 "X ELETTI OI GX 9:64 PX I ur-ajess Fp] “TT “9 ‘] Sum zo Fey jesıde ‘1 Sy1901 ‘01 “aoadse jesıop ‘pray ‘6 ‘Burm 2107 Jo 1ND-€ pur IND ojgipueur ‘ZI “ay purq 6 JL ‘aejo pury J9UUI “euuorue jo xode ‘ç ‘aoadse jesıop ‘wmouosow ‘y :199dse jeruoag ‘pray ‘ç !sauaudas jeuuaiue YI pue pie Jo [re19p ‘7 ‘109dse Jessie] ‘snugeH ‘I :(ropseem ‘Spuejsoion ‘9 wor GI ang) SOGIIOYISUIT] ‘ SPUEHOHAN ‘è ‘(AEPIEH) vprumso]f vIOANIAPY ‘1-1 "SSA 17 VAN ACHTERBERG: Dapsilarthra and Mesocrina "x VLG? “C7 ‘OZ “61 SX 91:97 “12 ‘81 “OLS x I UN-oJess :€7 ‘ZI “GT ‘001 187 vo aydıs [ny oyqipuew ‘oz foadse jesıop ‘Kero ‘7 ‘Bay pury ‘eq fav] appiu Jouur ‘77 “aoadse jes10p ‘pray ‘17 ‘euuaiue zo xade ‘07 ‘41001 pag uo ıydıs [ny ‘oyqrpuew ‘61 fa0adse jeszop ‘93193 as] ‘gy ‘s8urm ‘/y fioadse jesuorz ‘peoy ‘9] ‘aoodse jezorer ‘snargey ‘Gy ‘ad/10299] 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‘96 ‘edAojou (Hoyosig) Tuurwauargs VIO SOI —S6 ‘SIA REVISIONARY NOTES ON THE SUBFAMILY GNAPTODONTINAE, WITH DESCRIPTION OF ELEVEN NEW SPECIES (HYMENOPTERA, BRACONIDAE) by C. VAN ACHTERBERG Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands ABSTRACT The tribe Gnaptogastrini Tobias, 1976, is synonymized with the tribe Gnaptodontini Fischer, 1970, and the group is given subfamily rank. The genera Gnaptogaster Tobias, 1976, and Gnaptodon Haliday, 1837, are redescribed and illustrated. À neotype for Bracon pumilio Nees, 1834, (type-species of Gnaptodon) is selected, described and illustrated. A lectotype is designated for Diraphus pygmaeus Wesmael, 1838. Keys are given to the Pal- aearctic and Afrotropical species of Gnaptodon, of which eight new Palaearctic and two new Afrotropical species are described; additionally the first Oriental Graptodon species is described. Mesotages Foerster, 1862, is synonymized with Gnaptodon Haliday, 1837, and Mesotages decoris Foerster, 1862, is a new senior synonym of G. bachmaieri Fischer, 1957, and of G. klemensiewiczii (Niezabitowski, 1910). The ecology of the reared Palaearctic Gnaptodon species and the phylogenetic relationships of the Gnaptodontinae are discussed. INTRODUCTION The genus Gnaptodon contains some of the smallest Braconidae, usually scarcely longer than 1 mm and exclusively parasites of the min- ing caterpillars of Nepticulidae (Lepidoptera). Gnaptodon and its relatives are recognizable by the peculiar basal elevation of the 2nd tergite (figs. 8, 31). However, in a new species de- scribed in this paper both the elevation and the 2nd suture of the metasoma are absent (fig. 19). Owing to a fine collection of parasites of Nepti- culidae assembled by Dr. G. Bryan and Mr. E J. van Nieukerken (Vrije Universiteit, Am- sterdam) and the Gnaptodon species reared by Dr. J. Szôcs (Budapest) it has been possible to revise the Palaeartic species of Gnaptodon. The breviradialis group (only described species: bre- viradialis) could be sorted out, resulting in six new species. Tobias (1979: 240) on the basis of his material (which was not available for this study) con- cluded that all described Palaearctic forms actu- ally belong to one large polymorphic species with enormous variation: G. pumilio (Nees). I agree with Tobias that the degree of sculpture and the colour alone are inadequate and often very variable character-states. However, the wing venation, the setosity, the shape of the grooves of the metasoma and (to some extent) coloration appear to allow of the recognition of 25 several species, and a study ot the available reared series has confirmed the usefulness of these characters. Additionally the opportunity is taken to syn- onymize the tribe Gnaptogastrini Tobias, 1976, with the Gnaptodontinae, to redescribe the ge- nus Gnaptogaster Tobias, 1976, and to describe two new species of Gnaptodon from Africa (S. Africa and Somalia) and one new species from Thailand. In the cosmopolitan genus Gnapto- don 21 valid species have been described and in this paper 11 species are added. The generic combinations of the host species mentioned in this paper are on behalf of Mr. E. J. van Nieu- kerken (Amsterdam), who is currently studying the biosystematics of the Nepticulidae. If the genus is given in inverted commas then the ge- neric placement is uncertain. For the terminolo- gy used in this paper, see Van Achterberg (1979: 242246, figs. 1—8). I wish to thank the following persons, who provided unidentified material, types and/or in- formation: Dr. F. Bin (Perugia), Dr. G. Bryan (Amsterdam), Dr. P. Dessart (Brussels), Dr. E. Haeselbarth (München), Mr. T. Huddleston (London), Dr. E. Kierych (Warsaw), + Dr. E Königsmann (Berlin), Dr. J. Macek (Prague), Mr. E. J. van Nieukerken (Amsterdam), Mr. G. M. Nishida (Honolulu), Dr. J. Papp (Budapest), Dr. M. R. Shaw (Edinburgh), Mr. H. J. Vlug 26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 (Wageningen), Mrs. I. Wegener (Berlin), and Dr. A. Zaykov (Plovdiv). Dr. M. R. Shaw, Mr. T. Huddleston, and Mr. E. J. van Nieukerken provided useful critical notes on the text. Subfamily GNAPTODONTINAE Fischer stat. nov. Gnaptodonina Fischer, 1970: 85, 87. Gnaptodontini Fischer, 1972: 55—57. Gnaptogastrini Tobias, 1976a: 319. Syn. nov. Diagnosis. — Length of fore wing 1.2—1.9 mm.; antennal segments 14—26; maxillary palp with 5 or 6 segments; labial palp with 3 seg- ments; scapus truncate apically, stout (figs. 10, 22); hypostomal, occipital, and prepectal cari- nae usually absent, but occipital carina present in Liparophleps Enderlein and Gnaptodon novobrittanicus Fischer, and prepectal carina present in Liparophleps; eyes glabrous, slightly or not emarginate; antennal sockets closer to eyes than to each other (fig. 32); anterior tento- rial pits rather small and deep; clypeus (largely) smooth, rather convex; apical margin of clypeus slightly concave (fig. 14); hypoclypeal depres- sion present, transverse, but labrum (nearly) flat and glabrous; malar suture absent; length of malar space 1.0—1.5 times basal width of man- dible; mandible slender, twisted apically and 1st tooth much longer than 2nd; pronope, antescu- tal depression, lateral carina of mesoscutum, precoxal sulcus (except for a medial pit as in, Gnaptogaster, fig. 1) absent; posterior flange of propleuron virtually absent; notauli reduced posteriorly (figs. 9, 18); episternal scrobe nar- row (fig. 10); metapleural flange small or ab- sent; scutellum completely smooth; metanotum at most with a short antero-medial carina, pos- teriorly smooth and rather convex, not distinct- ly protruding (fig. 22); propodeum smooth, without any areolation or carinae; propodeal spiracle round, rather small and in front of mid- dle of propodeum; propodeal tubercle absent; 1st discal cell of fore wing petiolate; vein M + CUI of fore wing completely sclerotized; vein cu-a of both wings present, short (figs. 3, 24); veins CU1b and 2-1A of fore wing more or less reduced, resulting in a partly open 1st sub- discal cell (figs. 3, 24); parastigma small (fig. 13) or medium-sized (fig. 3); vein m-cu of fore wing antefurcal and parallel to vein 1-M; veins 2A and 2-R1 of fore wing absent; vein SR of hind wing largely absent apically (figs. 3, 24); vein M + CU of hind wing shorter than vein 1- M; length of vein 3-SR of fore wing 0.2—0.7 times vein 2-SR; tarsal claws slender apically, simple, without lobe or pectination (tig. 2); spurs of hind tibia subequal, their length 0.2— 0.3 times the basitarsus; fore tibia without spin- es or pegs; lst metasomal tergite sessile, rather flat medio-basally (at least in the Oriental and Palaearctic species); spiracles of 1st tergite sub- medial or behind middle (figs. 1, 8, 19, 31); laterope absent (fig. 1) or small and pit-shaped (fig. 22); 2nd tergite with a curved basal eleva- tion (figs. 8, 31), except in Gnaptodon apheles spec. nov. (fig. 19) which has only weak lateral traces; 2nd intersegmental suture of metasoma distinctly impressed, except in G. apheles spec. nov. (fig 19); spiracles situated in notum of 2nd and 3rd metasomal tergites; metasoma de- pressed; ovipositor almost straight (fig. 1, 22); length of ovipositor sheath in Palaearctic and Oriental species 0.03—0.12 times fore wing. Contains 4 genera: Gnaptodon Haliday, 1837 (cosmopolitan), Pseudognaptodon Fischer, 1965 (New World, possibly only a subgenus of Gnaptodon), Gnaptogaster Tobias, 1976 (Pal- aearctic), and (provisionally included) Lipa- rophleps Enderlein, 1920 (Neotropical). KEY TO GENERA OF THE SUBFAMILY GNAPTODONTINAE 1. Occipital and prepectal carinae present..... Liparophleps Enderlein (Synonym of Plesa- demon Fischer, 1961; may belong to the Doryctinae, but lacks the characteristic pegs of the fore tibia). — Occipital and prepectal carinae absent, at most with a faint trace of the occipital cari- Na Present i... a EE 2 2. Veinir-m\of fore wing absentee eee SEE Pseudognaptodon Fischer — Vein r-m of fore wing present (as an unscle- rotized, slightly pigmented vein, figs. 3, 13) ER EEE N eee ee 30.0 20 3 3. Dorsal carinae of 1st tergite absent (fig. 8); mesoscutal pit present (fig. 9); apex of an- tenna without short spine (fig. 4); pleural sulcus crenulate (fig. 1); maxillary palp with SISC SM CHIESE RANE Gnaptogaster Tobias — Dorsal carinae of Ist tergite present (figs. 19, 31); mesoscutal pit absent (fig. 18); apex of antenna with short spine (fig. 11); pleural sulcus smooth (fig. 10); maxillary palp with 6 segments (fig. 16) ... Gnaptodon Haliday Genus Gnaptogaster Tobias, 1976 Gnaptogaster Tobias, 1976a: 319. Type-species: Gnaptogaster mongolica Tobi- as, 1976. Gender: feminine. VAN ACHTERBERG: Gnaptodontinae 27 Diagnosis. — Length of fore wing ca. 1.8 mm, length of body ca. 2.1 mm; antennal seg- ments 18—20; apex of antenna without spine apically (fig. 4); maxillary palp with 5 segments; occipital carina completely absent; marginal cell of fore wing very narrow (fig. 3); apical margin of clypeus not differentiated from clypeus (fig. 6); mesopleuron smooth, except for the medial pit (fig. 1); mesoscutal pit present (fig. 9); no- tauli shallow, finely crenulate, but posteriorly absent (fig. 9); pleural sulcus crenulate (fig. 1); dorsal carinae of 1st metasomal tergite absent (fig. 8); basal elevation of 2nd tergite distinctly differentiated by the curved groove (fig. 8); length of ovipositor sheath ca. 0.06 times fore wing. Contains only the East Palaearctic type-spe- cies. Gnaptogaster mongolica Tobias, 1976 (figs. 1—9) Gnaptogaster mongolica Tobias, 1976a: figs. 2—5. 319—321, Paratype, 9, length of body 2.1 mm, of fore wing 1.8 mm. Head. — Antennal segments 19, 3rd segment 1.2 times 4th segment, length of 3rd and 4th segment 2.3 and 2.0 times their width, respecti- vely, penultimate segment 2.1 times its width; length of maxillary palp 0.5 times height of head; length of eye 2.6 times temple in dorsal view; POL: © ocellus: OOL = 10 : 5 : 8; frons concave behind antennal sockets, smooth; ver- tex rather flat and finely granulate near stemma- ticum; face convex and smooth; epistomal su- ture present, but medially shallow (fig. 6); length of malar space equal to basal width of mandible. Mesosoma. — Length of mesosoma 1.3 times its height; side of pronotum smooth, except for a narrow crenulate area posteriorly (fig. 1); metapleuron smooth; middle lobe of mesoscu- tum with weak medial suture (fig. 8); mesoscu- tum smooth; scutellar sulcus medium-sized, rather shallow, with 4 weak longitudinal cari- nae; scutellum rather convex, smooth; surface of propodeum smooth; medial carina absent. Wings. — Fore wing: r : 3-SR : SR1 = 4:5: SIE OIC 75772387 2-SR 7 3-SR eet m= 17:5: 14; m-cu far antefurcal (fig. 3). Hind wing: m-cu present as a weak, unpig- mented trace (fig. 3). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.3, 6.1 and 6.0 times their width, respectively (fig. 5). Metasoma. — Length of 1st tergite 0.6 times its apıcal width, its surface smooth, basally rath- er flat, medially convex, and laterally rather flat- tened (fig. 8); spiracles of 1st tergite after mid- dle of tergite and not protruding (fig. 8); glym- ma completely absent; 2nd tergite smooth and 2nd intersegmental suture deep and smooth; all metasomal setae widely spaced, scattered; 2nd and 3rd segments with a sharp lateral crease; length of ovipositor sheath 0.06 times fore wing. Colour. — Black; legs (but hind coxa black- ish basally and hind tarsus somewhat infus- cated), tegulae, palpi, mandibles, scapus and an- nellus mainly, and metasoma (but apically infus- cated), yellowish- brown; pterostigma dark brown, wing membrane hyaline. Holotype and paratypes in the Zoological Institute, Leningrad. The figured and redes- cribed paratype is topotypic with the holo- type: “Mongolia, South Gobijskij ajmak, 20 km. West of well of Barin-Bugatyn-Khuduk, Kerzhner, 25—27.vii.1969” (translated), “Para- typus Gramptogaster (sic!) mongolica Tobias”. Note. — Tobias erected a separate tribe for his Gnaptogaster, but this is evidently unneces- sary. It fits well into the Gnaptodontinae be- cause it shares several synapomorphous charac- ter-states with Gnaptodon. For example, the curved basal elevation of the 2nd tergite, the short 2nd submarginal cell of the fore wing, the absence of vein CU1b of the fore wing, the 3- segmented labial palp, the narrow and shallow hypoclypeal depression, and the (nearly) flat and glabrous labrum. Genus Gnaptodon Haliday, 1837 Gnaptodon Haliday, 1837: 220. Fischer, 1972: 569. Shenefelt, 1975: 1123. Tobias, 1976a: 315—318; 1976b: 22, 47. Marsh, 1979: 173. Diraphus Wesmael, 1838: 89. Fischer, 1972: 569. Shenefelt, 1975: 1123. Marsh, 1979: 173. Mesotages Foerster, 1862: 258. Syn. nov. Type-species: Bracon pumilio Nees, 1834. Gender: masculine. Diagnosis. — Length of fore wing and of body, both 1.0—1.9 mm; antennal segments 14—26; apex of antenna with short spine (fig. 11); maxillary palp with 6 segments (fig. 28); apical margin of clypeus narrowly differentiated from clypeus (fig. 14); mesopleuron completely 28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 smooth, exceptionally precoxal sulcus superfi- cially impressed; mesoscutal pit absent (fig. 18), exceptionally with a shallow impression; pleural sulcus smooth (fig. 10); notauli of Palaearctic and Oriental species smooth, and posteriorly (nearly) absent (figs. 18, 26); dorsal carinae of 1st tergite basally present (figs. 19, 31); length of ovipositor sheath of Palaearctic and Oriental species 0.03—0.12 times length of fore wing. Biology. — The larvae are obligatory para- sites of (mining) larvae of Nepticulidae (Lepido- ptera). Pupation takes place in the host cocoon, but endoparasitism remains to be proven (see chapter on phylogenetic relationships). Distribution: Palaearctic: 12 species; Nearc- tic: 7 species; Neotropical: 1 species; Afrotropi- cal: 3 species; Oriental: 1 species; Australian: 8 species. A. PALAEARCTIC REGION KEY TO THE PALAEARCTIC SPECIES OF THE GENUS GNAPTODON HALIDAY 1. Second intersegmental suture of metasoma deep medially (figs. 31, 43, 51); transverse elevation of 2nd tergite distinct medially (figs. 31, 63), exceptionally obsolete (ig. 102); de segment vari- able, if dark brown, then hind femur yel- lomashars. srl. hornet) a 2 — Second suture (almost) absent (fig. 19); transverse elevation of 2nd tergite absent medially (fig. 19); 3rd antennal segment dark brown; hind femur largely dark brown Meek ent eet apheles spec. nov. 2. Distance between wing apex and apex of marginal cell 1.5—3.6 times vein 1-R1 (figs. 57, 65, 74, 91); length of pterostigma 1.5— 3.0 times vein 1-R1; intermediate speci- mens have posterior margin of elevation of 2nd tergite weakly curved or straight me- dially (figs. 63, 80) and metasoma of ? comparatively robust (fig. 110) ......... 3 — Distance between wing apex and apex of marginal cell 0.2—1.4 times vein 1-R1 (figs. 20, 24), exceptionally up to 1.8 times (fig. 46); length of pterostigma 0.7—1.9 times vein 1-R1; intermediate specimens have posterior margin of elevation of 2nd tergite distinctly curved (figs. 51, 53) and metaso- ma of comparatively slender (fig. 53) .. 9 3. Vein r of fore wing about as long as vein 3- SR or longer (fig. 91); antero-lateral grooves of 3rd tergite rather deep (excep- tionally obsolete) and (very) finely crenu- late (fig. 92); vertex (largely) smooth; an- tennalisezments; 1719 RARE MR I breviradialis Fischer Vein r of fore wing shorter than vein 3-SR (fig. 74, 84); antero-lateral grooves of 3rd tergite absent or shallow and smooth (figs. 72, 80); vertex often coriaceous; antennal segments 8-23) e 4 . Marginal cell of fore wing narrow (fig. 57); distance between wing apex and apex of marginal cell about 3 times vein 1-R1 (fig. 57); metasoma completely — yellowish; length of vein SR1 of fore wing about 3 times vein 3-SR (fig. 57); episternal scrobe deep (fig. 55); ocelli small, POL 2.5—3 times diameter of posterior ocellus (fig. 56); medial length of 2nd tergite about 1.5 times medial length of 3rd tergite (fig. 63)........ TRL ao Fr Ae brevis spec. nov. Marginal cell of fore wing wider (figs. 65, 96); distance between wing apex and apex of marginal cell 1.5—2.2 times vein 1-R1; colour of metasoma variable; length of vein SR1 of fore wing 4.0—6.7 times vein 3-SR (figs. 74, 96); episternal scrobe rather shal- low or obsolete (fig. 94); ocelli larger, POL about twice diameter of posterior ocellus or shorter (figs. 93, 98); medial length of 2nd tergite less than 1.5 times medial length of Srditergite (1125280, 85) 5 . Vertex almost completely smooth and strongly shiny (fig. 71); 3rd tergite with shallow and smooth antero-lateral grooves (fig. 72); Ist and 2nd tergites, antero-lateral corners and hind tarsı (except telotarsus), ivory-whitish; 2nd suture of metasoma dis- tinctly crenulate medially (fig. 72); vein SRI Ofiore wing istrale Ae (ies o>) MEE lers O 0... vlugi spec. nov. Vertex coriaceous, moderately shiny (fig. 83); 3rd tergite with no antero-lateral grooves (figs. 80, 102), colour and vein SR1 variable; 2nd suture smooth or indistinctly Crenulate (es 30,102), Sr 6 . Long dense setose between antennal sock- ets (figs. 76, 77); medio-longitudinal groove of mesoscutum anteriorly as deep as notauli (fig. 81); medial length of 3rd tergite equal to medial length of 2nd tergite (fig. 80) or slightly less; 2nd tergite dark brown poste- Oy pilosus spec. nov. Sparsely setose between antennal sockets (fig. 83); medio-longitudinal groove ot mesoscutum anteriorly shallower than no- 10. VAN ACHTERBERG: Gnaptodontinae 29 tauli (figs. 82), 100); medial length of 3rd tergite variable (figs. 85, 110); 2nd tergite frequently completely yellowish ........ 7 . Hind coxa at least basally and hind femur dorso-apically infuscated; 2nd tergite pos- teriorly and hind tarsus dark brown; medial length of 3rd tergite of 9 1.1—1.3 times medial length of 2nd tergite (fig. 85); mar- ginal cell of fore wing comparatively nar- row (fig. 84), distance between wing apex and apex of marginal cell 1.9—2.4 times vein 1-R1; face (except medially) densely and transversely rugulose-coriaceous ...... Er in no nieukerkeni spec. nov. Hind coxa, hind femur apically, 2nd tergite posteriorly and hind tarsus (except telotar- sus), yellowish; medial length of 3rd tergite of 2 0.7—0.8 times medial length of 2nd tergite (figs. 102, 110); marginal cell of fore wing somewhat wider (figs. 96, 104), dis- tance between wing apex and apex of mar- ginal cell 1.6—1.9 times vein 1-R1; face coriaceous or smooth . Head and apical half of metasoma yellow- ish; 2nd suture of metasoma rather angular- ly bent (fig. 102); pterostigma and 3rd and 4th antennal segments dark brown; anten- nal segments of 2 about 21; scutellar sulcus distinct (fig. 100) ruficeps spec. nov. Head and apical half of metasoma black; 2nd suture of metasoma rather evenly curved (fig. 110); pterostigma light brown; 3rd and 4th antennal segments of 2 about 18; scutellar sulcus very shallow (fig. 107) .. erasmi spec. nov. . Distance between wing apex and apex of marginal cell of fore wing 0.2—0.5 times vein 1-R1 (fig. 24); vein SR1 of fore wing largely straight (fig. 24); area behind trans- verse elevation of 2nd tergite often more or lessisculpruredi(ie:91) ANA rom 10 Distance between wing apex and apex of marginal cell of fore wing 0.7—1.8 times vein 1-R1 (figs. 20, 46); vein SRI rather sin- uate (fig. 46); area behind elevation of 2nd tergite often largely smooth (figs. 51, 53) odin NERO EEL AN. 11 Length of vein 1-R1 of fore wing 2—4 times distance between wing apex and apex of marginal cell of fore wing (fig. 24); 3rd antennal segment yellowish, exceptionally dark brown; mainly parasite of Nepticuli- dae in trees pumilio (Nees) Length of vein 1-R1 of fore wing about 5 times distance between wing apex and apex of marginal cell of fore wing (fig. 96 in Fischer, 1966); 3rd antennal segment dark brown Care. Se nepalicus Fisher 11. Four basal segments of antenna of ® infus- cated or dark brown dorsally, exceptionally (largely) yellowish; medial length of basal elevation of 2nd tergite less than 0.6 times medial length of rest of tergite or, if longer, less narrowed laterally (cf. fig. 31); medio- longitudinal groove of mesoscutum often distinct; metasoma usually dark brown or black, if partly yellowish then elevation of 2nd tergite indistinct medially; antennal segments of 2 mostly 21; parasites of Nep- ticulidae in herbs and (low) shrubs ........ decoris (Foerster) — Four basal antennal segments of 2 yellow- ish; medial length of basal elevation of 2nd tergite 0.6—1.0 times medial length of rest of tergite and elevation distinctly narrowed laterally (figs. 51, 53); medio-longitudinal groove of mesoscutum usually obsolete (fig. 49); several tergites of metasoma yellowish; basal elevation of 2nd tergite distinct me- dially; antennal segments of 2 usually 19 or 20; parasites of Nepticulidae in trees and (IS georginae spec. nov. Note. — Of the 7 Nearctic species, 3 are close to Palaearctic spp.: Gnaptodon bicolor Fischer, 1965, is near erasmi, but bicolor has length of mesosoma about 1.5 times its height and distance between apex of fore wing to apex of marginal cell about 1.5 times vein 1-R1 (fig. 876 in Fischer, 1977). G. glaber Fischer, 1965, is close to georginae but the latter has infuscated hind tarsi, pterostigma dark brown, and face less sculptured. Finally G. nepticulae (Rohwer, 1915) (= G. pulchrigaster Fischer, 1965, accord- ing to Marsh, 1974) is close tot pumilio, but nepticulae differs by the reddish base of the metasoma and the deep, crenulate, antero- lateral grooves of 3rd tergite. The only Neo- tropical species described, G. novoteutonicus Fischer, 1977, is exceptional because of its long ovipositor. Gnaptodon apheles spec. nov. (figs. 10—19) Holotype, ?, length of body and of fore wing both 1.6 mm. Head. — Antennal segments 20, length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 3.5 and 2.8 times their width 30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 respectively, penultimate segment 1.7 times its width; length of maxillary palp 0.9 times height of head; length of eye 1.5 times temple in dorsal view; POL : 2 ocellus:OOL = 8:4:10; frons slightly convex, smooth; vertex convex, faintly coriaceous; face laterally coriaceous, shi- ny; length of malar space 1.5 times basal width of mandible. Mesosoma. — Length of mesosoma 1.5 times its height; middle lobe of mesoscutum slightly impressed (fig. 18); scutellar sulcus rather nar- row and shallow, and with one medial longi- tudinal carina besides the crenulae (fig. 18); episternal scrobe medium-sized (fig. 10). Wings. — Fore wing: r:3-SR:SR1 = 6 sil 2 62e TCM s2CUil = 18e ZAR 2 B= SR :r-m = 16:11 : 11; distance of wing apex to apex of marginal cell 0.5 times length of vein 1-R1; length of pterostigma 0.8 times length of vein 1-R1 (fig. 13); vein SR1 nearly straight (fig. 13). Legs. — Length of femur, tibia and basitarsus of hind leg 4.4, 8.5, and 7.5 times their width, respectively. Metasoma. — Length of 1st tergite equal to its apical width, its surface smooth, and dorsal carinae weakly developed in front of spiracles (fig. 19); 2nd tergite smooth, without distinct transverse elevation basally, except for some lateral traces (fig. 19); 2nd intersegmental suture absent; length of ovipositor sheath 0.06 times fore wing. Colour. — Black; antenna, palpi, coxae, tro- chanters, middle and hind femora, tegulae, pte- rostigma, wing veins, and metasoma ventrally, dark brown; rest of legs yellowish brown; wing membrane hyaline. Holotype in the Haeselbarth Collection, Munchen: “St. Peter/Ahrntal, Sudtirol, 1900 m, J/26.8.(19)67, Haeselbarth”. Paratype: 19 (Rijksmuseum van Natuurlijke Historie, Lei- den), topotypic, but from 2200 m. Antennal segments 20, length of fore wing 1.7 mm; length of ovipositor sheath 0.06 times fore wing; length of body 1.5 mm; other characters essen- tially as holotype. Note. — The wing venation of G. apheles is similar to the venation of G. pumilio (Nees), but pumilio differs (in addition to its sculptured 1st tergite, the distinct transverse groove of its 2nd tergite, and the deep 2nd suture of its metaso- ma) by its yellowish scapus, pedicellus, 3rd an- tennal segment, and its more extensive yellow- ish legs. Gnaptodon breviradialis Fischer (figs. 91—93) Gnaptodon breviradialis Fischer, 1959: 259; 1972: 571—572, fig. 435. Shenefelt, 1975: 1123. Tobias, 1976b: 48. Szócs, 1979: 200, 201. Variation. — Antennal segments 17—19; length of fore wing and of body, both 1.4—1.8 mm; 4 or 5 basal antennal segments yellowish; vein r of fore wing about as long as vein 3-SR of longer (fig. 91); length of pterostigma 1.6—3 times vein 1-R1 of fore wing; distance between wing apex and apex of marginal cell of fore wing 2.1—3.6 times vein 1-R1; 3rd tergite with usually finely crenulate antero-lateral grooves, but these sometimes shallow and smooth; ver- tex smooth. Specimens examined: Holotype, 2, also 7 9 and 7 d. The examined specimens originate from France, Hungary and Greece. Hosts. — Nepticulidae in the tree zone: in Loranthus europaeus Jacq. (in Greece col- lected from Castanea trees): ex Niepeltia loranthella (Klimesch); Greece, Evvoia, Dhirfis Oros, 2 km NE Steni, 500 m, Castanea-Abies forest; id., S slopes of Dhirfis Oros, 700—900 m; Hungary, Nor- mafa. in Ulmus spec.: ex Stigmella ulmivora (Fologne); France, Digne. in Prunus mahaleb L.: ex Ectoedemia mabalebella (Klimesch); Greece, Mt. Timfristos, Evritania, above Karpenission, Spartium shrub, 1200—1400 m. in P. cocomilia Ten.: ex Stigmella amygdali (Klimesch); Greece, 4 km NW Timfristós, (vill.), Fthiotis, 1400 m, clearings in Abies wood. Tobias (1976b: 48) reported breviradialis from the European part of the U.S.S.R. and gives Stigmella prunetorum (Stainton) as host. Gnaptodon brevis spec. nov. (figs. 55—63) Holotype, ©, length of body 1.4 mm, of fore wing 1.3 mm. Head. — Antennal segments 20, length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 3.7 and 3.0 times their width, respectively, the penultimate segment 2 times its width (fig. 61); length of maxillary palp 0.8 times height of head; length of eye 1.6 times VAN ACHTERBERG: Gnaptodontinae 31 temple in dorsal view (fig. 56); POL : © ocel- lus : OOL = 14: 5 : 19; frons rather flat and coriaceous; vertex distinctly coriaceous, face medially smooth, otherwise coriaceous (fig. 62); length of malar space 1.5 times basal width of mandible. Mesosoma. — Length of mesosoma 1.2 times its height; medio-longitudinal depression of mesoscutum absent (fig. 60); scutellar sulcus rather shallow and medium-sized, crenulate (fig. 60); episternal scrobe deep (fig. 55). Wings. — Fore wing:r:3-SR:SR1 = BEIGE MGE OU ED -E@ UF Sel 28; 02-SR:5- SR :r-m = 16:11:11; length of pterostigma 2.3 times vein 1-R1; distance between apex of fore wing and apex of marginal cell 3 times vein 1-RI (fig. 57); vein SRI straight. Legs. — Length of femur, tibia and basitarsus of hind leg 3.1, 8.5, and 5.3 times their width, respectively. Metasoma. — Length of 1st tergite 0.6 times its apical width, its surface superficially coria- ceous (fig. 63), and dorsal carinae present in basal half of tergite; 2nd tergite behind basal el- evation coriaceous, basal elevation smooth and distinctly transverse (fig. 63), its medial length 0.4 times medial length of rest of tergite; medial length of 2nd tergite 1.5 times 3rd tergite; 3rd and following tergites largely coriaceous, with no antero-lateral grooves; 2nd suture deep, curved and indistinctly crenulate; length of ovi- positor sheath 0.12 times fore wing. Colour. — Dark brown (including pterostig- ma and veins); antenna dark brown, but 3 basal segments ventrally yellowish; palpi, legs (but te- lotarsi infuscated), tegulae, and whole metaso- ma, yellowish; mesosoma laterally largely rath- er reddish; wing membrane slightly infuscated. Holotype in Museum Budapest: “Budaòrs, Törokugrato” (= Hungary), 1975.vii.16, Szöcs J., “Filipendula vulgaris”, “ex Nepticula hexa- petalae Szöcs”. Host: Ectoedemia hexapetalae (Szöcs) in Fil- pendula vulgaris Moench. Gnaptodon decoris (Foerster) comb. nov. (fig. 20) Mesotages decoris Foerster, 1862: 258. Gnamptodon bachmaieri Fischer, 1957: 41. Syn. nov. Gnaptodon bachmaieri; Fischer, 1972: 570—571. Shenefelt, 1975: 1123. Tobias, 1976b: 48. Szöcs, 1979: 200—201. Gnaptodon bachaieri (sic!); Fischer, 1980: 202. Gnamptodon klemensiewiczu Niezabitowski, 1910: 44. Shenefelt, 1975: 1123—1124. Syn. nov. Variation. — Antennal segments 20—23 (mostly 21); length of fore wing 1.2—1.7 mm, of body 1.1—1.7 mm; 5th—7th tergites of metasoma sometimes yellowish-brown, usually dark brown; length of pterostigma 1.0—1.6 (usually 1.1—1.2) times length of vein 1-R1; distance between apex of fore wing and apex of marginal cell 0.9—1.5 times vein 1-R1; 3rd an- tennal segment and base of metasoma dark brown, exceptionally yellowish; 2nd and 3rd tergites usually smooth, but sometimes superfi- cially sculptured behind the transverse elevation and the 2nd metasomal suture; vertex coria- ceous. Specimens examined: 22 ® and 29 6. The examined specimens originate from Great Britain (Wales), Netherlands (Wijster, dunes of Meijendel (near the Hague) and of Oostvoorne, Kunrade (Kunderberg), and Hil- versum (Spanderswoud)), West Germany, Italy, Austria, Hungary, Bulgaria and Greece. Hosts. — Nepticulidae in herbs and (low) shrubs: in Lotus corniculatus L.: ex Trifurcula cryptella (Stainton); Nether- lands, Kunrade, Kunderberg (2nd tergite dark brown and hind coxa infuscated). in Coronilla varia L.: from same host; Hungary, Svar. in Fragaria vesca L.: ex Stigmella fragariella (Heyd.) (on label, cf. aurella (F.)); Hungary, Budapest, Petnehazi- rét (apex of metasoma reddish, rest of meta- soma dark brown, hind coxa yellowish). in Sanguisorba officinalis L.: ex S. geminella (Frey) (on label, cf. poteri (Stainton)); Hungary, Föt, Mogyorodi p. (only 1st tergite and apex of metasoma dark brown, rest yellowish, hind coxa infuscated). in Rubus spec.: ex S. splendidissimella (H.-S.) or aurella (F.); Netherlands, Hilversum, Spanderswoud (2nd tergite yellowish). in Rosa pimpinellifolia L.: ex S. spec.; U.K., Wales, Pembrey, Carmar- then. in Agremonia agrimonoides (L.): ex Ectoedemia agrimoniae (Frey); Greece, Evvoia, Dhírfis Oros, S. slopes, Castanea- Abies-forest, 700—900 m (elevation of 2nd tergite largely absent, 2nd tergite and hind coxa yellowish). in Potentilla erecta L.: ex Stigmella poteru (Stainton); Hungary, Svar. 32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 in Potentilla spec: ex S. occultella (Hein.) (on label, cf. poteru (Stainton)); Austria, Schörfling, north of At- tersee; id., Hinter-Stoder. in Prunus spinosa L.: ex S. plagicolella (Stainton); Hungary, Vérz- teskozma (aberrant because of yellowish 3rd and 4th antennal segments; 2nd tergite dark brown). in Betula nana L.: ex S. betulicola nanivora Petersen; West Germany (types of G. bachmaieri (Fischer), examined: 2nd tergite dark brown, hind coxa yellowish. Notes. — The type of Gnaptodon klemensie- wiczu Niezabitowski, 1910, was not available for study (it is part of a private collection; according to Tobias (1979: 237) the type is lost). Judging from the original description (1st and 2nd tergites smooth and 3rd antennal segment blackish) it is most likely to be a junior syno- nym of G. decoris (Foerster). During my visits to the Berlin Museum I ex- amined the holotype of Mesotages decoris Foerster, 1982. It is a completely yellowish specimen, probably owing to ageing (as many other small specimens in the Foerster collection are bleached). The length of the pterostigma is 1.5 times vein 1-R1 (metacarp) and the distance between the apex of the fore wing and the apex of the marginal cell is 1.4 times vein 1-R1. The vertex is distinctly coriaceous and the 2nd su- ture of the metasoma smooth. The basal eleva- tion of the 2nd tergite is rather transverse. Foerster stated in the original description that the transverse elevation of the 2nd tergite is ab- sent, but the surface of the 2nd tergite of the metasoma 1s obscured by some glue and by part of the hind wing. If looked for carefully the typ- ical transverse elevation of the 2nd tergite can be seen. This explains how Foerster arrived at his incorrect statement in the original descrip- tion and Fischer’s description of this species as bachmaieri. The holotype of decoris in the Foerster Collection (Zoologisches Museum, Berlin) bears the following labels: “24/926”, “Rheinprovinz”, “Erst”, “decoris Frst”, “Gnap- todon bachmaieri Fi., det. Fischer”, “Glaube nicht dass dieser Ex. die Type von Mesotages decoris sein kann. Foerster schreibt ausdrück- lich dass Mesotages keine bogenformige gekrümmten Fürchen an den 2. Tergit hat, Fischer, 1963”. As pointed out above, I disagree with Fischer’s statement and synonymize Gnamptodon. bachmaieri Fischer, 1957, with Gnaptodon decoris (Foerster, 1862) comb. nov. There is a second specimen under decoris in the Foerster Collection, which does not belong to the type-series, and also has a reduced, easily overlooked transverse elevation on the 2nd ter- gite. Gnaptodon erasmi spec. nov. (figs. 103—111) Holotype, 9, length of body 1.6 mm, of fore wing 1.5 mm. Head. — Antennal segments 18, length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th segments 3.5 and 3.2 times their width, respectively, penultimate segment 1.9 times its width (fig. 105); length of maxillary palp 0.7 times height of head; length of eye twice temple in dorsal view (fig. 108); POL: © ocel- lus : OOL = 6 : 4 : 9; frons slightly convex, co- riaceous; vertex completely coriaceous; face largely smooth, only laterally coriaceous, rather flat (fig. 111); length of malar space 1.7 times basal width of mandible. Mesosoma. — Length of mesosoma 1.4 times its height; medio-longitudinal groove of mesos- cutum absent, represented only by a wide shal- low depression (fig. 107); scutellar sulcus very shallow, narrowly crenulate (fig. 107); epister- nal scrobe rather shallow (fig. 103). Wings. — Fore wing: r:3-SR:SR1 = Ja MO 46E CUP CUI = be Ske SR :r-m = 11:5:8; length of pterostigma twice length of vein 1-R1; distance between wing apex and apex of marginal cell 1.9 times vein 1-R1; vein SR1 slightly curved (fig. 104). Legs. — Length of femur, tibia and basitarsus of hind leg 3.2, 6.8 and 4.5 times their width, respectively. Metasoma. — Length of 1st tergite 0.9 times its apical width, its surface coriaceous, and pos- teriorly rugulose (fig. 110); dorsal carinae dis- tinct in basal 0.6 of 1st tergite; basal elevation of 2nd tergite distinct medio-posteriorly (fig. 110), its medial length 0.3 times rest of tergite; meta- soma behind elevation of 2nd tergite largely co- riaceous (fig. 103); 2nd suture deep, finely cre- nulate and rather evenly curved (fig. 110); medi- al length of 3rd tergite 0.7 times length of 2nd tergite; length of ovipositor sheath 0.07 times fore wing. Colour. — Black; 4 basal segments of anten- na (rest dark brown), palpi, tegulae and legs, brownish-yellow; 1st and 2nd tergite, and ante- ro-lateral corners of 3rd tergite, reddish-brown; pterostigma light brown. VAN ACHTERBERG: Gnaptodontinae 33 Holotype in Haeselbarth Collection: “Riva s. Garda (North Italy), 150 m, G/10.9.67, Hae- selb.”; (G = collected in vegetation of Quercus ilex-shrubwoodland, on lime, with very strong dominance of Q. ilex, mixed with Fraxinus or- nus). Paratype, 1 d (Rijksmuseum van Natuur- lijke Historie, Leiden): “Ectoedemia groschkei (Skala) in Vitex agnus-castus, VU no. 80675 KE”, “Ellas (Greece), Sept. 1980, S. B. J. Men- ken & E. J. v. Nieukerken”, “4 km SW Pa- padates (Préveza), cult. area, roadside, 100 m, 39.17N-20.46 E, 25.ix.1980, st. 53”; antennal segments 19, basal elevation of 2nd tergite obso- lete medially, otherwise as holotype. Host: “Ectoedemia” groschkei (Skala) in Vi- tex agnus-castus L. Note. It is a pleasure to me to dedicate this species to Dr. E. Haeselbarth (Minchen). Gnaptodon georginae spec. nov. (figs. 44— 54) Holotype, 6, length of body 1.5 mm, of fore wing 1.4 mm. Head. — Antennal segments 20, length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th segments 3 and 2.7 times their width, respectively, penultimate segment 1.8 times its width (fig. 48); length of maxillary palp 0.7 times height of head; length of eye twice temple in dorsal view; POL :@ ocellus : OOL 7:5:10; frons flat, largely coriaceous; vertex distinctly coriaceous (fig. 52); face superficially coriaceous-pimply (fig. 45); length of malar space 1.3 times basal width of mandible. Mesosoma. — Length of mesosoma 1.3 umes its height; medio-longitudinal groove obsolete (fig. 49); scutellar sulcus narrow, rather shallow (fig. 49); episternal scrobe indistinct (fig. 44). Wing. — Fore wing: r : 3-SR : SRI = 5:9: ÉDMÉCUIRPE CI 3235 2-SR SES Ren m = 24:9: 15; length of pterostigma 1.9 times vein 1-R1; distance between wing apex and apex of marginal cell 1.8 times vein 1-R1; vein SR1 slightly sinuate (fig. 46). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.4, 7.7 and 5 times their width, respectively. Metasoma. — Length of 1st tergite 0.8 times its apical width, its surface smooth; dorsal cari- nae distinct in basal 0.7 of 1st tergite (fig. 51); basal elevation of 2nd tergite distinctly nar- rowed laterally, medio-posteriorly distinctly curved (fig. 51), its medial length 0.8 times rest of tergite; area directly behind elevation of 2nd tergite, as rest of tergites basally, weakly coria- ceous (fig. 44); tergites otherwise smooth; 2nd suture rather deep, smooth; 3rd tergite with no antero-lateral grooves; medial length of 3rd ter- gite 0.9 times length of 3nd tergite (fig. 51). Colour. — Black; tegulae (largely), pterostig- ma, lst tergite medially, and antenna, dark brown; palpi, rest of Ist tergite, 2nd and 3rd tergites, antero-lateral corners of 3rd tergite, and legs (except the infuscated apex of hind tibia and the hind tarsus), brownish-yellow. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden: “ex Stigmella hybnerella (Hbn.), Crataegus monogyna, VU no. 80061 K1” “11 km ESE of Morris O.m., 26.iv.1980, Marsh & Pistacia carr.”, “Stat. 19”, “Algeria, E. v. Nieukerken, G. Bryan & P. Oosterbroek”. Raratypess oi) st ZARO (uscii dapest): “Tahi, Pilis hegys.” “1973.v.28, Szöcs J.”, “Nepticula malella Str. det. J. Szöcs”, “Mal- us sylvestris”; 2 2 (Shaw Collection, Edin- burgh): “Sulzchopf, BL., Switz(erland), LT/96, 470 m, H: Stigmella tityrella, mine coll. UOTE SEE STER hitebre dés (Zaykov Collection, Plovdiv): “22.9.1968, Ro- dopi, Parvenez, leg. A. Germanov”; 1 d (id): “21.5.1968, Plovdivsko Starosel, leg. A. Ger- manov”; 1 2 (Rijksmuseum van Natuurlijke Historie, Leiden): “Jambol, Ormana, 1981.v.27, leg. Zaykov”. The latter 3 specimens are from Bulgaria. Variation. — Antennal segments 19-21 (9) or 20—23 (4); 46 basal segments of 2 anten- na yellowish, penultimate segment twice its width (fig. 54), 6 has basal antennal segments infuscated or blackish; length of fore wing 1.4 mm (3 specimens); length of pterostigma 1.2— 1.9 times vein 1-R1; distance between apex of fore wing and apex of marginal cell 1.1—1.8 times vein 1-R1; medial length of basal eleva- tion of 2nd tergite 0.6-1.0 times rest of tergite; 2nd tergite often completely smooth (fig. 53); 2nd and 3rd tergites usually yellowish, if dark brown then apical half of metasoma yellowish (ex S. tityrella); length of ovipositor sheath 0.04 times fore wing. Hosts: Stigmella hybnerella (Hubner) in Cra- taegus monogyna L., S. malella (Stainton) in Malus sylvestris (L.), S. ruficapitella (Haworth) in Quercus, and S. tityrella (Stainton) in Fagus sylvatica L. It is a great pleasure to me to dedicate this species to Dr. Georgina Bryan (Amsterdam), who made an important collection of reared parasites of Nepticulidae available for study. 34 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 Gnaptodon nepalicus Fischer Fischer, 1966: 159—161, figs. 96—97. Shenefelt, 1975: 1124. Fischer considered G. nepalicus an Oriental species; however, I prefer to include it among the Palaearctic spp., because it was collected at 6800 ft. in the Himalayas. The fauna at such high altitudes is more closely correlated with the (South) Palaearctic region than with the Oriental region. As shown in the foregoing key nepalicus is close to G. pumilio and seems to dif- fer mainly in the colour of the antenna and in the wing venation. Gnaptodon nieukerkeni spec. nov. (figs. 82—90) Holotype, ©, length of body and of fore wing 1.4 mm. Head. — Antennal segments 18 (right anten- na) or 19 (left one), length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th seg- ments 3.2 and 3.0 times their width, respective- ly, penultimate segment 2.2 times its width (fig. 89); length of maxillary palp 0.7 times height of head; length of eye 2.1 times temple in dorsal view (fig. 83); POL :@ ocellus :OOL = 14 :7 :18; frons coriaceous, flat; vertex com- pletely coriaceous; face rather convex and obliquely, finely rugulose-coriaceous, medially narrowly smooth (fig. 90); length of malar space 1.7 times basal width of mandible. Mesosoma. — Length of mesosoma 1.4 times its height; medio-longitudinal groove obsolete (fig. 87); scutellar sulcus deep, rather narrow, and distinctly crenulate (fig. 87); episternal scrobe absent. Wings. — Fore wing: r : 3-SR :SR1=2:5: 24 M=CÜI 2=EUle= 177 10572-SRE3-SR Zr m = 8:5: 6; length of pterostigma 2.1 times vein 1-R1; distance between wing apex and apex of marginal cell 2.2 times vein 1-R1; vein SRI slightly sinuate (fig. 84). Legs. — Length of femur, tibia and basitarsus of hind leg 3.4, 7.3 and 5.0 times their width, respectively. Metasoma. — Length of 1st tergite 0.8 times its apical width, its surface smooth (fig. 85); dorsal carinae present in basal third of 1st ter- gite; basal elevation of 2nd tergite distinct, its medial length 0.4 times medial length of rest of tergite; 2nd tergite behind elevation and rest of metasoma largely coriaceous; 2nd suture deep and smooth; 3rd tergite with no antero-lateral grooves and its medial length 1.2 times medial length of 2nd tergite (fig. 85); length of oviposi- tor sheath 0.03 times fore wing. Colour. — Black; antenna (except yellowish annellus), tegulae, pterostigma and wing veins, dark brown; palpi, and legs, yellowish (but hind coxa dorsally largely, hind femur apico-dorsally and hind tarsus (other tarsi slightly) infuscated; 2nd tergite medially and antero-laterally brown, rest of tergite dark brown. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden: “Parnassós Oros, 2 km, W. Summit (Voiotia), rocks, dwarf shrub, 38.32 N- 22.35: E, 2000 my, 281x319 SOMS SN o} 80.685 KE, 21/24.x.80, in leaf-litter!”, “ex Stig- mella cf. rhamnophila (Amsel), Rhamnus saxa- tilis, VU no. 80.685 K.”, “Ellas, Greece, Sep- tember 1980, S. B. J. Menken, E. J. van Nieu- kerken”. Paratypes: 12 © and 2 d from same series (Vrije Universiteit, Amsterdam; Rijksmu- seum van Natuurlijke Historie, Leiden; Mu- seum Budapest; Collectie Zaykov, Plovdiv). Variation. — Antennal segments 17—20; length of fore wing 1.2—1.4 mm; vein SR1 of fore wing usually more curved than in fig. 84; length of pterostigma 1.8—2.3 times vein 1-R1; distance between apex of fore wing and apex of marginal cell 1.9—2.4 times vein 1-R1; medial length of 3rd tergite 1.1—1.3 (9) or 0.9 (d) times medial length of 2nd tergite; 2nd tergite often (rather dark) brownish medially. Host: Stigmella cf. rhamnophila (Amsel) in Rhamnus saxatilis Jacq. Gnaptodon pilosus spec. nov. (figs. 73— 81) Holotype, ®, length of body 1.4 mm, of fore wing 1.3 mm. Head. — Antennal segments 21, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 2.2 and 1.7 times their width, respectively, penultimate segment 1.4 times its width (fig. 75); length of maxillary palp 0.8 times height of head; length of eye 2.2 times temple in dorsal view (fig. 77); POL : © ocel- lus: OOL = 7:4:11; frons slightly concave behind antennal sockets, coriaceous; vertex dis- tinctly coriaceous (fig. 77); face rather convex and largely smooth, only microsculptured laterally (fig. 76); long and densely setose be- tween antennal sockets (figs. 76, 77; this setosi- ty is absent in all other spp. treated in this pa- per); length of malar space 1.1 times basal width of mandible. Mesosoma. — Length of mesosoma 1.1 times its height; medio-longitudinal groove rather VAN ACHTERBERG: Gnaptodontinae 35 deep, similar to notauli anteriorly (fig. 81); scu- tellar sulcus rather wide and deep, narrowly crenulate (fig. 81); episternal scrobe indistinct (fig. 73). Wings. — Fore wing: r:3-SR:SR1 = eo Glen te CUI 2 CUI Sat: 802 SRE SR :r-m = 20:9:11; length of pterostigma 1.8 times vein 1-R1; distance between wing apex and apex of marginal cell 1.8 times vein 1- R1; vein SR1 straight (fig. 74). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.4, 6.4 and 4 times their width, respectively. Metasoma. — Length of 1st tergite 0.8 times its apical width, its surface aciculate in posterior half, the rest smooth (fig. 80), dorsal carinae present in basal 0.7; basal elevation of 2nd ter- gite distinct (somewhat cariniform at posterior edge), its medio-longitudinal length half the length of rest of 2nd tergite; metasoma behind elevation of 2nd tergite largely coriaceous (fig. 80); medio-longitudinal length of 2nd tergite equal to length of 3rd tergite; 2nd suture finely crenulate and rather deep; length of ovipositor sheath 0.08 times fore wing. Colour. — Black; 4 basal segments of anten- na (but scapus and pedicellus dark dorsally), palpi, humeral plate (tegula dark brown), legs (except dark brown tarsi), metasoma ventrally, Ist tergite (medially infuscated) and anterior half of 2nd tergite, yellowish; rest of antenna medially brown, its apical half dark brown; pro- notal sides and mesopleuron with brownish stripe ventrally (absent in paratypes); apex of pterostigma whitish, rest dark brown. Holotype in Museum Budapest: “Budapest, Rupphegy”, “1974.vi.9, Szöcs J.”, “Quercus pu- bescens, semen”, “ex Nepticula sp., 1974.v11.8.” Paratypes, 1 © + 1 d:1 d (Museum Budapest): “Hungaria, Budaôrs. Naphegy”, “1977.v.26., Szöcs J.”, “Quercus pubescens”, “ex Nepticula eberhardi Joh., det. J. Szöcs”. As holotype, but metasoma slender, antennal segments 23, Ist tergite smooth, and pterostigma completely dark brown; 1 2 (Rijksmuseum van Natuurlijke Historie, Leiden): “Fót (= Hungary), 1960.ix.19”, “lg. Mihalyi”. Antennal segments 21; as holotype, but medial length of 2nd tergite 1.1 times 3rd tergite. Host: Stigmella eberhardi (Joh.) in Quercus pubescens Willd. Variation. — Antennal segments 21—23, length of fore wing 1.3—1.5 mm; length of pte- rostigma 1.5—2.1 times vein 1-R1; distance be- tween apex of fore wing and apex of marginal cell 1.6—2.1 times vein 1-R1. Note. G. pilosus resembles the Nearctic G. glaber Fischer, 1965, but glaber has the pteros- tigma yellowish, length of mesosoma about 1.5 times its height, 1st tergite longer than apical width, metasoma completely smooth and face distinctly coriaceous. Gnaptodon pumilio (Nees) (figs. 22—33) Bracon pumilio Nees, 1834: 90—91. Gnaptodon pumilio; Fischer, 1972: 572—574, figs. 436—437. Shenefelt, 1975: 1124. Tobias, 1976b: 48. Van Achterberg, 1976: 60, figs. 37, 38. Hud- dleston, 1978: 47. Fischer, 1980: 202. Gnamptodon pumilio; Shaw & Askew, 1976: 131. Diraphus pygmaeus Wesinael, 1838: 90, figs. 11, F. A neotype of Bracon pumilio Nees, 1834, is here selected, because the original type-series is lost, it is the type-species of the genus Grapto- don Haliday, 1837, and closely related species occur in the same faunal region. The hind coxae of the neotype are infuscated contrary to the original description, but Nees probably over- looked this feature. Description of neotype, ®, length of body and of fore wing 1.9 mm. Head. — Antennal segments 21, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 4.3 and 3.0 times their width, respectively, the penultimate segment 1.8 times its width; length of maxillary palp equal to height of head; eyes slightly emarginate (fig. 32); length of eye 2.6 times temple in dorsal view; POL: 2 ocellus:OOL = 12:7: 18; frons flat, coriaceous; vertex largely smooth, convex; face rather convex (more so than in decoris) and laterally distinctly coriaceous (fig. 32); length of malar space 1.4 times basal width of mandible. Mesosoma. — Length of mesosoma 1.5 times its height; middle lobe of mesoscutum antero- medially slightly depressed (fig. 26); scutellar sulcus narrow, merely crenulate (fig. 26); medi- al carina of propodeum absent. Wings. — Fore wing: r : 3-SR:SR1= 4:7: 552 TCUM ¢ DCI = 5e Bile ZON 8 Basis 3 Er m = 27: 14: 15; distance of wing apex to apex of marginal cell 0.3 times vein 1-R1; length of pterostigma 1.1 times vein 1-R1 (fig. 24); vein SR1 straight (fig. 24). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.8, 9.2 and 7.5 times their width, respectively. Metasoma. — Length of Ist tergite equal to 36 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 its apical width, its surface behind the spiracles finely rugulose and in front of the spiracles largely smooth (fig. 31); dorsal carinae of the 1st tergite present in basal half of tergite; posterior margin of transverse elevation of 2nd tergite straight medio-posteriorly, behind it rugulose; medial length of elevation of 2nd tergite 0.6 times medial length of rest of tergite; 2nd tergite with some punctures, its medial length 1.5 times medial length of 3rd tergite (fig. 31); 2nd inter- segmental suture medially deep and laterally obliterated, with a pair of obsolete, posteriorly diverging submedial grooves antero-laterally and a pair of obsolete grooves close to suture; ovipositor straight, without notch, and with a dorso-subapical nodus and some minute ventral teeth (fig. 30); length of ovipositor sheath 0.11 times fore wing. Colour. — Black; mouthparts (including pal- pi), 3 basal antennal segments, tegulae, vein C + SC + R of fore wing, legs (but hind coxa largely infuscated) and metasoma ventrally, more or less yellowish-brown; pterostigma and 4th an- tennal segment dark brown; metasoma behind Ist tergite dorsally blackish-brown; wing mem- brane hyaline. Neotype in Rijksmuseum van Natuurlijke Historie, Leiden: “Nederland, Wijster (Dr.), opposite Biol. Stat., 18—25.vi.1976, C. v. Ach- terberg”. Note. Gnaptodon pumilio (Nees) differs from decoris, in addition to the characters given in the key, by the distinctly setose back of the head (above the hypostomal carina) and the more convex face. The type-series of Diraphus pygmaeus Wes- mael, 1838, consists of one typical female speci- men of G. pumilio (Nees) with the 2nd and 3rd tergites brownish, one female with yellowish 2nd and 3rd tergites (“var. 1” of Wesmael) and one male (probably not conspecific). The first specimen mentioned is here designated as lecto- type. The lectotype has no distinct antero- lateral grooves at the 3rd tergite, the whole 1st tergite is dark brown, the 2nd metasomal suture is rather deep but smooth, the vertex is coria- ceous, the 2nd tergite is distinctly sculptured and the hind coxae are yellowish. Variation. — The variation within pumilio as interpreted in this paper is considerable. I have tried to split up the complex, but the results were unsatisfactory. The variation (especially of coloration) within a reared series is so extensive that any attempted division has been frustrated. Considering the available data, it is likely that the specialisation within the complex has not re- sulted (yet) in recognizable species. At least three forms can be discerned, of which form A seems to be ecologically differ- entiated from both other forms because it has been reared from Nepticulidae in herbs and bramble (Rubus spp.). The other forms have been reared from Nepticulidae in trees. I men- tion these three forms only to indicate the main lines of variations encountered. All characters mentioned are very variable, even within the same reared series. Form A (including pygmaeus var. 1 (Wes- mael)) has the 1st and 2nd tergites yellowish, with the middle of the 1st tergite usually dark brown, but sometimes the whole metasoma (ex- cept Ist tergite) is yellowish; vertex behind stemmaticum coriaceous; 2nd suture of metaso- ma shallow; antero-lateral grooves of 3rd tergite absent; hind coxa yellowish. Form B has basal half of metasoma blackish or yellowish; vertex behind stemmaticum often smooth; 2nd suture of metasoma deep and cre- nulate; antero-lateral grooves of 3rd tergite deep and often finely crenulate; exceptionally the 3rd tergite has a separate crenulate and curved croove, directly behind 2nd suture in stead of antero-lateral grooves (this indicates that these grooves may be the result of an amal- gamation of the groove and the 2nd suture in the middle of the tergite); hind coxa yellowish or infuscated. Even within series from the same host there is considerable variation. The typical form has the whole metasoma blackish-brown; vertex behind stemmaticum smooth; 2nd suture of metasoma deep and smooth (fig. 31); antero- lateral grooves of 3rd tergite incomplete, shal- low and smooth; hind coxa infuscated or yel- lowish. Intermediates to form B occur frequent- ly, e.g. in a series from North Italy (St. Peter, Ahrntal, 1270 m, Haeselbarth Collection). Specimens examined: 52 ® and 22 d (of which 7 © and 4 & belong to form A). Antennal segments 19—23 (of both sexes mostly 21 or 22); length of body 1.6—1.8 mm; length of fore wing 1.7—1.9 mm; 4th antennal segment yel- lowish, exceptionally dark brown; area behind transverse elevation of 2nd tergite smooth to rather extensively rugulose; apex of metasoma frequently reddish brown; length of pterostig- ma: vein 1-R1 of fore wing : distance between wing apex and apex of marginal cell of fore wing = 0.8—1.2 : 1 : 0.25—0.6. The examined specimens originate from Norway, Sweden, Great Britain (Scotland, England), Ireland, VAN ACHTERBERG: Gnaptodontinae 37 Netherlands (Bemelen (Bemelerberg); Cadier (Schiepersberg); Castricum (dunes); ’s Graven- land (Ankeveense Plassen); Grevenbicht; Hil- versum (heath Spanderswoud); Hulshorst (Leu- venumse Bos); Nederhorst den Berg (Spiegel- polder); Waarder; Wijster; Winterswijk), France, Italy, Austria, Hungary, Bulgaria and Greece. Hosts: Reared from Nepticulidae in trees (form B and typical form): in Betula verrucosa Ehrh.: (= pendula auct.) ex Stigmella luteella (Stainton); U.K., Dan- bury, and Debden, both Essex. in Betula spec.: ex S. betulicola (Stainton); U.K., Thorpe- ness, Suffolk. ex S. confusella (Wood); Norway, Ändalsnes, More & Romsd. ex S. continuella (Stainton): Netherlands, Hilversum, heath. ex S. lappomica (Wocke); Netherlands, dunes of Castricum; U.K., Delamere, Cheshire. in Salix caprea L.: ex S. salicis (Stainton); U.K., Benenden, Kent, (intermediate between form A and typical form, apex of metasoma partly yel- lowish). in Tilia spec.: ex S. tiliae (Frey); Hungary, Vásárosbée. ini Ulmus procera Salisbury: ex S. ulmivora (Fologne); U.K., Hallow, Worcestershire. in U. glabra Mill.: ex Nepticulidae; Hungary, Bátorliget. in Ostrya carpinifolia Scopoli: ex S. carpinella (Heinemann); Greece, Oiti Oros, Fthiotis, 4 km E, pavliani, 850 m. in Sorbus aucuparia L.: ex S. magdalenae (Klimesch); Norway, Storen, Sor-Trondelag, 60 m. ex S. sorbi (Stainton); U.K., Blackford Hill, Edinburgh. ex S. spec.; Ireland, Ballinahinch, Co. Gal- way. in Malus domestica L.: ex S. malella (Stainton); Netherlands, Gre- venbicht (one & has 2nd tergite yellowish, the other dark brown!). ex S. pomella (Vaughan); Netherlands, Win- terswijk. in Rhamnus catharticus L.: ex S. rbamnella (H.-S.); Hungary, Budaòrs. in Quercus robur L.: ex S. ruficapitella-group; Netherlands, Hulshorst, Leuvenumse Bos; id., Hilversum, heath; ıd., Bemelen, Bemelerberg. ex Ectoedemia caradjai (Hering); France, Digne. Form A: in Rubus fruticosus agg.: ex Ectoedemia erythrogenella (de Joannis); U.K., Benfleet, Essex. ex Stigmella splendidissimella (H.-S.) or au- rella (F.); Netherlands, Nederhorst den Berg, Spiegelpolder; id., ’s Graveland, An- keveense Plassen, (part of series has almost whole metasoma yellowish). in Rubus spec.: from same host(s); Netherlands, Hilversum. Spanderswoud, (whole metasoma (except middle of 1st tergite) yellowish). in Agrimonia eupatoria L.: ex S. aeneofasciella (H.-S.); Netherlands, Cadier, Schiepersberg (part of series has whole metasoma yellowish (@) or behind 2nd tergite blackish (d)); Austria, Schörfling, north of Attersee. Gnaptodon ruficeps spec. nov. (figs. 94— 102) Holotype, 9, length of body 1.6 mm, of fore wing 1.5 mm. Head. — Antennal segments 21, length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th segments 3.5 and 3 times their width, respectively, penultimate segment 1.8 times its width (fig. 95); length of maxillary palp 0.8 times height of head; length of eye 1.8 times temple in dorsal view (fig. 98); POL : © ocel- lus: OOL = 12 :7 : 18; frons slightly convex, coriaceous; vertex completely coriaceous (fig. 98); face rather convex and largely coriaceous (fig. 101); length of malar space 2.1 times basal width of mandible. Mesosoma. — Length of mesosoma 1.2 times its height; medio-longitudinal groove of mesos- cutum absent, represented by only a wide, shal- low depression (fig. 100); scutellar sulcus rather wide and deep (fig. 100); episternal scrobe shal- low. Wings. — Fore wing: r : 3-SR : SR1 = 3 :6: Ds CIM 2= GO — Ie EP Shere SNe met m = 10:6: 7; length of pterostigma 1.5 times vein 1-R1; distance between apex of fore wing and apex of marginal cell 1.6 times vein 1-R1; vein SR1 slightly curved (fig. 96). Legs. — Length of femur, tibia and basitarsus of hind leg 3.9, 9 and 6 times their width, re- spectively. 38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 Metasoma. — Length of 1st tergite 0.6 times its apical width, its surface superficially coria- ceous; dorsal carinae present in basal half of 1st tergite (fig. 102); basal elevation of 2nd tergite indistinct, its medial length 0.5 times rest of ter- gite; 2nd tergite behind elevation and rest of metasoma largely coriaceous (fig. 94); 2nd su- ture deep, smooth and distinctly bent (fig. 102); 3rd tergite with no antero-lateral grooves; me- dial length of 3rd tergite 0.8 times medial length of 2nd tergite (fig. 102); length of ovipositor sheath 0.08 times fore wing. Coulour. — Yellowish-brown; stemmaticum, antenna (but scapus largely yellowish), mesoso- ma (but propleuron, mesopleuron dorsally and metapleuron brownish), and pterostigma, dark brown; vertex (partly) and telotarsi, infuscated. Holotype in Museum Budapest: “Nagy- kovàcsi (= Hungary), 1967, vii.9, Szöcs J.”, “ex Nepticula dorycniella Suire”, “Dorycnium ger- manicum”. Host: Trifurcula dorycniella (Suire) in Doryc- nium germanicum (Grenli). Gnaptodon vlugi spec. nov. (figs. 64—72) Holotype, d, length of body 1.2 mm, of fore wing 1.4mm. Head. — Antennal segments 18, antenna widened apically (fig. 64), length of 3rd segment 1.1 times 4th segment, length of 3rd and 4th segments 3.5 and 3.2 times their width, respecti- vely, penultimate segment 2.2 times its width (fig. 66); length of maxillary palp 0.9 times height of head; length of eye 1.7 times temple in dorsal view (fig. 71); POL : © ocellus : OOL = 16 :8:19; frons almost smooth, slightly de- pressed behind antennal sockets; vertex largely smooth; face largely smooth, laterally indis- tinctly micro-sculptured, rather convex (fig. 68); length of malar space equal to basal width of mandible. Mesosoma. — Length of mesosoma 1.3 times its height; medio-longitudinal groove of mesos- cutum shallow but distinct (fig. 69); scutellar sulcus rather wide, distinctly crenulate (fig. 69); episternal scrobe deep (fig. 64). Wings. — Fore wing: r: 3-SR: SR1 = 6: 11:54; 1-CU1 : 2-CU1 = 3: 11; 2-SR : 3-SR : r-m = 24: 11: 16; length of pterostigma 1.9 times vein 1-R1; distance between apex of fore wing and apex of marginal cell 2.2 times vein 1- RI (fig. 65); vein SRI straight. Legs. — Length of femur, tibia and basitarsus of hind leg 3.6, 6.6 and 5.2 times their width, respectively. Metasoma. — Length of 1st tergite 0.9 times its apical width, its surface smooth, and dorsal carinae obsolescent (fig. 72); basal elevation of 2nd tergite weakly differentiated, strongly transverse, its medial length 0.4 times medial length of rest of tergite (fig. 72); 2nd tergite be- hind elevation coriaceous but posteriorly smooth; medial length of 2nd tergite 1.2 times medial length of 3rd tergite; 3rd tergite smooth and with shallow and smooth antero-lateral grooves (fig. 72); 2nd suture deep, medially dis- tinctly crenulate. Colour. — Black; palpi, legs (except infus- cated telotarsi), 5 basal segments of antenna, te- gulae, 1st and 2nd tergites, antero-lateral cor- ners of 3rd tergite and anterior half of metasoma ventrally, ivory-whitish; pterostigma brown; rest of antenna (dark) brown. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden: “Sweden, Hallandsl., Särö- Hamra, swept, 26.6.1977, leg. H. J. Vlug”. It is a pleasure to me to dedicate this species to its collector, Mr. H. J. Vlug (Scherpenzeel). THE ECOLOGY OF THE REARED PALAEARCTIC GNAPTODON SPECIES As shown in fig. 131, if reared more than once, the hosts of Gnaptodon species are usual- ly found on plants of several families. Obvious- ly the parasites do not select certain plant groups. If the host plants are arranged accord- ing to their morphology (trees, (low) shrubs, and (non-woody) herbs), however, the picture changes (fig. 130). The assignment of host plants of the Nepticulidae to a tree, shrub, or herb zone is, of course rough: the low shoots of a tree may be lower than a large herb, and the same applies for a dwarf shrub. Nevertheless, the species seem to select for a certain vegeta- tion zone; if more than one zone is included (e.g. pumilio, no. 1 in fig. 130) then the speci- mens in another zone are aberrant in colour and/ or morphology with respect to the nominate form (e.g. form A of pumilio in herbs) and may constitute already a good biological species or be in the process of active sympatric speciation. If two morphologically very close species oc- cur in the same zone (nos 3 and 5 in fig. 130) then one of the species is specialized: in this case no. 5 (pilosus) seems to be restricted to Nepticu- lidae in Quercus, while no. 3 (georginae) parasi- tizes Nepticulidae in other trees (and high VAN ACHTERBERG: Gnaptodontinae 39 shrubs). In the case of erasmi (no. 6) and nieu- kerkeni (no. 7), the latter seems to be restricted to dwarf shrubs among rocks at high altitude. G. erasmi is known from 100—150 m altitude and occurs in higher shrubwood. Summarizing, the selection of a vegetation zone is of major importance. Within one zone closely related species are ecologically separated because of their preference to à certain host- plant (pilosus in Quercus, georginae in other trees) or to a certain altitude (erasmi at low alti- tude, nieukerkeni at high altitude). Obviously a lot of research, especially cross-breeding experi- ments, still has to be done before definitive con- clusions can be drawn. B. AFROTROPICAL REGION KEY TO AFROTROPICAL SPECIES OF THE GENUS GNAPTODON HALIDAY 1. Second tergite of metasoma with distinct transverse elevation basally (figs. 119, 128); length of vein SR1 of fore wing 7—10.5 timessyemW3-SRA(ties. 114122) #67 ee 2 — Second tergite with no distinct transverse elevation basally; length of vein SR1 of fore wing about 3 times vein 3-SR ............. RR vader le unifossa Fischer 2. Distance between fore wing apex and apex of marginal cell about 0.7 times vein 1-R1 (metacarp); length of pterostigma about 1.1 times vein 1-R1; length of vein SRI of fore wing about 10 times vein 3-SR (fig. 122); Ist tergite largely aciculate-rugulose (fig. 128); antenna comparatively slender basally (aliene. ngi similis spec. nov. — Distance between fore wing apex and apex of marginal cell about 1.3 times vein 1-R1; length of pterostigma 1.5—1.9 times vein 1- R1; length of vein SRI of fore wing 7—8 times vein 3-SR (fig. 114); 1st tergite largely smooth (fig. 119); antenna rather robust rally (fino bini spec. nov. Gnaptodon bini spec. nov. (figs. 112—120) Holotype, ©, length of body, and of fore wing 1.1 mm. Head. — Antennal segments 14, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 2.5 and 2.0 times their width, respectively, penultimate SE 1.8 times its width; length of maxillary palp 0.8 times height of head: length of eye 1.9 times temple in dorsal views POW): Drocellus OOL =" 9: 4 M0; frons slightly impressed, coriaceous (figs. 116, 117); vertex coriaceous; face rather convex, laterally coriaceous, rest smooth (fig. 117); length of malar space 1.2 times basal width of mandible Mesosoma. — Length of mesosoma 1.4 times its height; medio-longitudinal groove of mesos- cutum absent; scutellar sulcus narrow (fig. 120); episternal scrobe obsolescent (fig. 112). Wings. — Fore wing: r : 3-SR : 3-R1 = 5: I0OGSMECUAE ME CUS EPEPESRERESRE r-m = 25: 10: 11; length of pterostigma 1.9 times vein 1-R1; distance between wing apex and apex of marginal cell 1.4 times vein 1-R1; vein SR1 slightly curved. Legs. — Length of femur, tibia and basitarsus of hind leg 3.2, 6 3 and 4 times their width, re- spectively. Metasoma. — Length of 1st tergite 0.7 times its apical width, its surface smooth; dorsal cari- nae of 1st tergite obsolescent (fig. 119); basal el- evation of 2nd tergite slightly developed medial- ly, rest of 2nd tergite and 3rd tergite coriaceous, rest of metasoma largely transversely micro-ac- iculate (fig. 112); medial length of basal eleva- tion of 2nd tergite 0.4 times rest of tergite; me- dial length of 2nd tergite 0.9 times length 3rd tergite (fig. 119); 2nd suture deep, smooth, 3rd tergite with no antero-lateral grooves; length of ovipositor sheath 0.08 times fore wing. Colour. — Blackish or dark brown; clypeus ventrally, mandibles, 4 basal segments of anten- na, tegula, legs (but fore and middle telotarsi and hind tarsus infuscated), and 3 basal seg- ments of metasoma, brownish-yellow; rest of metasoma dark brown; humeral plate whitish; pterostigma dark brown; palpi whitish. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden: “Somalia, Afgoi, iv.1977. Lower Shabelli Valley, F. Bin”. Additional material : 1 ® (non-type), in same institute, to- potypic, length of fore wing i mm, antennal segments 16, robust antenna as holotype (length of 3rd and 4th segments 2.5 and 2 times their width, respectively), length of pterostigma 1.5 times vein 1-R1, distance between apex of fore wing and apex of marginal cell 1.3 times vein 1- R1; hind tarsus and whole metasoma yellowish; head (except stemmaticum) browmish-yellow; 1st tergite with some micro-aciculae. Not la- belled as paratype because of aberrant colora- tion. Note. The combination of the short distance between the apex of the fore wing and the apex of the marginal cell, the short vein 1-R1, and the 40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 low number of antennal segments, differentiates it from the known Palaearctic species. The only Oriental species (described in this paper) differs e.g. by its slender pterostigma. I am pleased to dedicate this species to Dr. F. Bin (Perugia), who collected several interesting Braconidae in Somalia. Gnaptodon similis spec. nov (figs. 121— 129) Holotype, ©, length of body 1.2 mm, of fore wing 1.3 mm. Head. — Antennal segments 17, length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 3 and 2.5 times their width, respectively, penultimate segment 2.3 times its width (fig. 123); length of maxillary palp 0.8 times height of head; length of eye 1.1 times temple in dorsal view; POL : © ocellus : OOL = 6 :3 : 8; frons slightly impressed, coriaceous; vertex strongly shiny, coriaceous (fig. 124); face rugulo-coriaceous laterally, rest smooth, rather convex (fig. 125); length of malar space equal to basal width of mandible. Mesosoma. — Length of mesosoma 1.4 times its height; medio-longitudinal groove of mesos- cutum absent (fig. 129); scutellar sulcus narrow, merely finely crenulate (fig. 129); episternal scrobe small and shallow. Wings. — Fore wing: r : 3-SR : SR1 = 4 : 4 : AD CON CUNEMEAMEPESREM SRE m = 14 : 4 : 10; length of pterostigma 1.1 times vein 1-R1 (fig. 122); distance between wing apex and apex of marginal cell 0.7 times vein 1- R1; vein SRI straight (fig. 122). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.6, 6.2 and 4.5 times their width, respectively. Metasoma. — Length of 1st tergite equal to its apical width, its surface largely aciculate-ru- gulose (fig. 128); dorsal carinae distinct in basal half of 1st tergite; basal elevation of 2nd tergite distinct, posteriorly straight (fig. 128), its medial length 0.3 times length of rest of tergite; 2nd tergite medially and 3rd tergite basally coria- ceous, rest of metasoma smooth (fig. 121); 2nd suture deep and smooth; medial length of 2nd tergite 1.1 times length of 3rd tergite; length of ovipositor sheath 0.08 times fore wing. Colour. — Brownish-yellow; 6th—17th an- tennal segments, 3rd and 4th metasomal ter- gites, ovipositor sheath and stemmaticum, dark brown or blackish; mesosoma and rest of meta- soma, brown; pterostigma rather light brown. Holotype in British Museum (Natural Histo- ry), London: “Port St. John(s), Pondoland, April 5—30, 1923”, “S. Africa, R. E. Turner, Brit. Mus., 1923—286”. Note. Resembles the Palaearctic G. decoris, but similis has vein SR1 of fore wing straight, head and base of antenna yellowish, metasoma distinctly sculptured, and fewer antennal seg- ments (20—23 in decoris). C. ORIENTAL REGION Gnaptodon orientalis spec. nov. (figs. 34-43) Holotype, ®, length of body 1.0 mm, of fore wing 1.2 mm. Head. — Antennal segments 19, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 3.3 and 2.5 times their width, respectively; penultimate segment 2.3 times its width (fig. 41); length of maxillary palp 0.6 times height of head; length of eye 2.6 times temple in dorsal view; POL: © ocellus: OOL = 8: 4: 12; frons virtually flat and smooth; vertex convex and smooth; face rather convex and smooth; length of malar space 1.5 times basal width of mandible. Mesosoma. — Length of mesosoma 1.2 times its height; mesoscutal lobes evenly setose, with- out medial depression (fig. 42); scutellar sulcus narrow and virtually smooth (fig. 42). IR — Fore wing:r:3-SR : SRI = 3 : À : ; 1-CU1 : 2-CU1 = 3: 25; 2-SR: 3-SR: m = 11:4: 8; length of pterostigma 1.4 times vein 1-R1; length of distance between apex of wing and apex of marginal cell 0.8 times vein 1- R1 (fig. 37); pterostigma slender (fig. 37; all other spp. treated in this paper have the pteros- tigma more robust (figs. 13, 20, 24)); vein SR1 slightly curved (fig. 37). Legs. — Length of femur, tibia, and basitar- sus of hind leg 3.5, 8.0, and 5.5 times their width, respectively. Metasoma. — Length of Ist tergite equal to its apical width, its surface largely smooth, with dorsal carinae developed in basal two-thirds of tergite (fig. 43); curved transverse elevation of 2nd tergite distinct, in front smooth, behind it superficially coriaceous (fig. 43); 2nd interseg- mental suture of metasoma medially distinct, smooth, with no additional grooves; setae rath- er sparse and spread; ovipositor straight, with nodus subapically (fig. 34); length of ovipositor sheath 0.05 times fore wing. Colour. — Yellowish-brown; antenna (ex- cept scapus); stemmaticum, mesoscutum VAN ACHTERBERG: Gnaptodontinae 41 largely, 3rd-5th tergites medially, ovipositor sheath, pterosigma, and vein 1-R1, dark brown; rest of veins brown; scutellum, medial part of metanotum, propodeum, and 1st tergite basally, hind tarsus and telotarsi, infuscated; wing mem- brane hyaline. Holotype in Bernice P. Bishop Museum, Ho- nolulu: “Thailand: Trang Prov., Khaophappha Khaochang, 200 m., 11—15.1.1964”, “G. A. Samuelson, Malaise Trap, Bishop”. Note. — G. orientalis spec. nov. runs in my key to the Palaearctic species to G. decoris (Foerster); however, orientalis differs by the more slender pterostigma and longer marginal cell of the fore wing (fig. 37 versus fig. 20), the yellowish head and base of the metasoma, and by the (nearly) smooth face and 1st tergite (both more or less sculptured in decoris). Of the Afro- tropical spp., it most resembles G. similis spec. nov. from S. Africa; however, orientalis differs by the slender pterosigma and marginal cell of fore wing (fig. 37 versus fig. 122) and by the smooth Ist tergite (fig. 43 versus fig. 128). It runs in the key to the Australian species of Gnaptodon by Fischer (1978: 397—398) to G. novobrittanicus Fischer, 1971, because of the rather evenly setose mesoscutum. However, G. novobrittanicus is not closely related; it has a longer ovipositor sheath (longer than half length of metasoma), longer maxillary palp (equal to height of head), more antennal segments (26 in holotype), different colour (mesosoma black), and is twice as large. Additionally G. novobrit- tanicus has a weakly developed ventral part of the occipital carina and the propodeum has a medial carina. PHYLOGENETIC RELATIONSHIPS The Gnaptodontinae may be easily mistaken for a group of the Opiinae because of the ab- sence of the prepectal carina, the almost flat and glabrous labrum, and the presence of a shallow hypoclypeal depression. However, the wing ve- nation, the more or less developed transverse el- evation of the 2nd metasomal tergite, the pit in front of the propodeal spiracle, and the 3-seg- mented labial palp of the Gnaptodontinae would be aberrant within the Opiinae. Also the biology is aberrant; Opiinae are obligatory lar- val endoparasites of Diptera, while the Gnapto- dontinae (as far as known) are obligatory (?en- do)parasites of Nepticulidae (Lepidoptera). The cephalic structures of the final instar lar- va of Gnaptodon are fairly complete and include robust and toothed mandibles (fig. 43 in Capek, 1970). The mandibles of the larvae of Opiinae are slender and toothless. The toothed mandi- bles of the larvae of the Gnaptodontinae are pe- culiar (if it is a real endoparasite), because endo- parasites such as Opiinae, Alysiinae and the en- doparasitic groups, e.g. of the Rogadinae, have smooth mandibles. This may indicate that Gnaptodon actually is an ectoparasite, which glues the egg to the host’s intersegmental mem- brane (as in the ectoparasitic Rhysipolis; Dr. M. R. Shaw, in litt.). A really sound and accurate proof that Gnaptodon is an endoparasite has not yet been obtained; it is only deduced from circumstantial evidence (e.g. the host remains look as though an endoparasite has been there and the parasitized host is mobile and able to spin a cocoon when it leaves the mine). The del- icate cocoon of Gnaptodon is formed in the co- coon of the host. The cephalic structures of the larvae of some ectoparasitic Rogadinae are similar to those of the Gnaptodontinae. However, he similarity is based on symplesiomorphous character-states (e.g., the toothed mandibles). The cephalic structures of the larvae of Gnaptodontinae are completely different from the more or less re- duced cephalic structures of the larvae of Opii- nae. Despite this evidence Fischer (1970, 1972 & 1977) and Tobias (1976a, not 1976b) favour the inclusion of the Gnaptodontinae as a tribe in the Opiinae, because, for example, the trans- verse elevation of the 2nd tergite is considered to be too aberrant an evolutionary tendency to include it in the Rogadinae (Fischer, 1972: 56) as proposed by Capek (1970). Peculiarly, it is forgotten that this evolutionary tendency is also absent in the Opiinae. Fischer (1972: 56) provi- sionally retains his Gnaptodontini in the Opii- nae because of “konventionellen Gründen”. Within the Rogadinae the Gnaptodontinae would fit in near the Exothecini. The synapo- morphous character-states shared with the Exo- thecini are the absence of the prepectal carina and the precoxal sulcus, and the scarcely devel- oped laterope. Unfortunately these are all re- ductions and such “negative apomorphous character-states” are comparatively less reliable for a phylogenetic arrangement than “positive apomorphous character-states”, such as the pe- culiar curved transverse elevation of the 2nd tergite in the Gnaptodontinae. The only positive synapomorphy with the Exothecini is in the bi- ology and then only in being parasites of leaf- miners. The morphology of the Gnaptodontinae (as 42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 2, 1983 far as I have examined the group) seems too de- rived to give any firm clue about its sister- group. An exception may be the rather flat la- brum combined with the transverse, rather shal- low hypoclypeal depression (figs. 6, 14), which is aberrant for (almost all of) the Rogadinae and normal for a large part of the Opiinae. The only alternative to its inclusion in the Rogadinae or in the Opiinae seems to be to consider it an old archaic subfamily, and the presence of rather numerous species both in the Australian and Palaearctic regions supports this view. Thus the Gnaptodontinae are considered to have branched off somewhere from the stem of group B (Van Achterberg, 1976, fig. 123), which includes the Opiinae and Alysiinae. All of the group B are now exclusively parasites of Diptera and originally endoparasites of mining larvae of Diptera (as the majority of the Opiinae still are). Later they switched to larvae in (more or less decaying) fruits, followed by accepting hosts in decaying vegetable matter (e.g. dung), and finally in decaying animal matter (carrion). It is interesting to note that the Gnaptodontinae are (still) specialized (?ecto)parasites of mining larvae of an archaic group of Lepidoptera; this makes it likely that the Gnaptodontinae branched off before the Opiinae became special- ized on dipterous hosts (and if Graptodon is an ectoparasite, even before the development of endoparasitism), since the group from which the Gnaptodontinae, Opiinae and Alysiinae stem seems to consist of solitary ectoparasites of various orders, including Lepidoptera and Di- ptera, as still are found in the Rogadinae. Another problem is the position of the group B within the Braconidae. Tobias (1967, fig. 43) placed it as a sister-group of the Helconinae, while Capek (1970) and Van Achterberg (1976) considered the group a branch of the “cyclosto- me”-group, including the Rogadinae. The latter view has recently gained much support from the work of Edson & Vinson (1979) on the mor- phology of the venom apparatus of the female. The venom apparatus of the Opiinae and Alysu- nae is similar to that of the Rogadinae and dif- ferent from the venom apparatus of the Helco- ninae. Unfortunately the venom apparatus of the Gnaptodontinae remains still to be investi- gated; thus its similarity to that of the Opiinae remains to be proved. REFERENCES Achterberg, C. van, 1976. A preliminary key to the subfamilies of the Braconidae (Hym.). — Tijdschr. Ent. 119: 33—78, figs. 1—123. ——, 1979. A revision of the subfamily Zelinae auct. (Hymenoptera, Braconidae), — Tijdschr. Ent. 122: 241—479, figs. 1—900. Capek, M., 1970. A new classification of the Braconi- dae (Hym.) based on the cephalic structures of the final instar larva and biological evidence. — Can. Ent. 102: 846—875, figs. 1—58. Edson, K. M. & S. B. Vinson, 1979. À comparative morphology of the venom apparatus of female Braconids (Hym., Braconidae). — Can. Ent. 111: 1013—1024, figs. 1—2. Fischer, M., 1957. Zwei neue Parasiten aus der in den Blattern der Zwergbirke minierende Raupe von Stigmella nanivora Pet. (Hym., Braconidae). — NachrBl. bayer. Ent. 6: 41—43. — — 1959. Die europäische Opiinae (Hym., Braconi- dae). — Sb. ent. Odd. nár. Mus. Praze 33: 241— 263. ——, 1963. Die Opiinae der athiopische Region (Hym., Braconidae). — Beitr. Ent. 13: 662—747. ——, 1966. Revision der Indo-Australischen Opiinae (Hym., Braconidae). — Ser. ent. 1: 1—166, figs. 1—97. ——, 1970. Probleme der Systematik bei den Opiinae (Hym., Braconidae). — Z. ArbGem. ost. Ent. 22: 81—88, figs. 1—14. — —, 1972. Opiinae, Hym., Braconidae. — Tierreich 91: i-xii + 1—620, figs. 1—463. , 1977. Subfamilia Opiinae II, Amerika (neark- tische und neotropische Region). — Tierreich 96: i-xxvil + 1—1001, figs. 1—890. ——, 1978. Neue Opiinae (Hym., Braconidae) von der australischen Region, besonders aus Tasma- nien. — Polskie Pismo Ent. 48: 371—412, figs. 138% ——, 1980. Opiinen aus den österreichischen Al- penlandern und angrenzenden Gebieten, gesam- melt von Herrn Dr. E. Haeselbarth (München) (Insecta: Hym., Braconidae). — Ber. naturw-med. Ver. Innsbruck 67: 185—212, figs. 1—6. Foerster, A., 1862. Synopsis der Familien und Gat- tungen der Braconen. — Verh. naturf. Ver. preuss. Rheinl. 19: 224— 288. Haliday, A. H., 1837. Essay on parasitic Hymeno- ptera. — Ent. Mag. 4: 203 —221. Huddleston, T., 1978. Braconidae, p. 46—62. In: Kloet, G. S. & W. D. Hincks. A check list of Brit- ish Insects, Hymenoptera (ed. M. G. Fitton et all.). — Handbk. Ident. Br. Insects 11 (4) : i-ix + 1—159. Marsh, M. P., 1974. New combinations and new syn- onyms in North American Braconidae (Hym.). — Proc. ent. Soc. Wash. 76: 285—289. —, 1979. Braconidae, p. 144295. In: Krombein, K.V. et all. Catalog of Hymenoptera in America North of Mexico 1:1-xvi + 1—1198. Nees von Esenbeck, C. G., 1834. Hymenopterorum Ichneumonibus affinum monographiae, genera Europaea et species illustrantes 1: 320. — Cottae, Stuttgartiae et Tubingae. VAN ACHTERBERG: Gnaptodontinae Niezabitowski, E. L., 1910. Materyaly do Brako- nidów Polski, I. — Spraw. Kom fizyogr. Krajow. 44: 47 —106. Shaw, M. R., & R. R. Askew, 1976. Ichneumonoidea (Hym.) parasitic upon leaf-mining insects of the orders Lepidoptera, Hymenoptera and Coleo- ptera. — Ecol. Ent. 1: 127—133. Shenefelt, R. D., 1975. Hymenopterorum Catalogus (nov. ed.). Part 12, Braconidae 8: 1115—1262. Junk, ’s-Gravenhage. Szócs, J., 1979. Angaben zu den Parasiten der mini- renden Motten (Hym., Braconidae). — Folia ent. hung. (ser. nov.) 32: 199—206. Tobias, V. I., 1969. A review of the classification, phylogeny and evolution of the family Braconidae (Hym.). — Ent. Obozr. 46: 645—669, figs. 1—43. 43 English transl.: Ent. Rev. 46: 387—399. , 1976a. Gnaptogaster mongolica gen. et sp. n. from a new tribe of braconids (Hym., Braconidae, Opiinae) and some problems of the phylogeny and evolution of this family. — Nasekomye Mon- golu 4: 315—321, figs. 1—5 (Russian). ——, 1976b. Brakonidy Kavkaza (Hym., Braconi- dae). — Opred. Faune SSSR 110: 1—287, figs. 1— 67. , 1979. Sistematika, izmentsjivost 1 istorija v Pal- earktike roda Gnaptodon Hal. (Hym., Braconi- dae). — Verh. VII. int. Symp. Entomofaun. Mitte- leuropa (1977): 237—240, figs. 1—5. Wesmael, C., 1838. Monographie des Braconides de Belgique. — Mém. Acad. Brux. 11: 1—166, figs. 1—17 + AF. CHECKLIST OF NAMES USED IN THE GENUS GNAPTODON HALIDAY Name apheles spec. nov. bachmaieri (Fischer, 1957) bicolor Fischer, 1965 bini spec. nov. breviradialis Fischer, 1959 brevis spec. nov. catamaranensis Fischer, 1978 clarimacula Fischer, 1978 decoris (Foerster, 1862) dispar Fischer, 1978 erasmi spec. nov. georginae spec. nov. glaber Fischer, 1965 klemensiewiczii Niezabitowski, 1910 longicauda Fischer, 1965 nepalicus Fischer, 1966 nepticulae (Rohwer, 1915) nieukerkeni spec. nov. novobritannicus Fischer, 1978 novoteutonicus Fischer, 1967 orientalis spec. nov. pilosus spec. nov. pulchrigaster Fischer, 1965 pumilio (Nees, 1834) pygmaeus (Wesmael, 1838) recticarinatus Fischer, 1965 rotundincisus Fischer, 1978 ruficeps spec. nov. rugulosus Fischer, 1965 similis spec. nov. sinuatus Fischer, 1965 talumalausensis Fischer, 1978 tasmanicus Fischer, 1978 tricrenulatus Fischer, 1978 unifossa Fischer, 1963 vlugi spec. nov. Type-locality page Northiltalyats to 5. 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Diagram depicting the vegetation zone wherein the host occur versus the indices of distance between the apex of the fore wing and the apex of the marginal cell (“c” in fig. 96) and length of vein 1-R1 (“b” in fig. 96). 1, Gnaptodon pumilio (Nees); A, form A; 2, G. decoris (Foerster); 3, G. georginae spec. nov. (in trees except Quercus); 4, G. ruficeps spec. nov.; 5, G. pilosus spec. nov. (in Quercus); 6, G. erasmi spec. nov; 7, G. nieuker- keni spec. nov; 8, G. breviradialis Fischer; 9, G. brevis spec. nov; T, tree zone; S, shrub zone; H, herb zone. © Lv OSD Plant % = © a = Vv 9 5 È 3 . 5 oO & & Vv 5 (=! © (SI Fe} ny GO À kan ee ICE RS = 5 9 Ses E D S a à 5 Gnaptodon O. cd ge nee ns Pp an & E D & species pumilio decoris georginae ruficeps Di pilosus erasmi nieukerkeni breviradialis. di brevis Fig. 131. The occurrence of Gnaptodon species in plant families infested by Nepticu Sa En At i UPPER IQ TE REA ETS M iu Fra CEE KARO TE TT NE pa i Leo, BE i DETTE Kira) u RL | platani AT va tetes cadi tone oui ige ea hund ke wnd NOTE Ù 7 9 AT CRE _ I ê bi = i \ + ge: PE FA SI x 5 NP = Jae, OENE he 5 2 ’ ODEEL 126 AFLEVERING 3 | 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD TH. C. M. Brock and G. van DER VELDE. — An autecological study on Hydromyza livens (Fabricius) (Diptera, Scatomyzidae), a fly associated with nymphaeid vegetation dominated by Nuphar, pp. 59—90, figs. 1—25. Tijdschrift voor Entomologie, deel 126, afl. 3 Gepubliceerd 20-V-1983 i ie à D Ù ve er HA ni a ct A ki 5 4 fi : 3 CL DE a ? AN AUTECOLOGICAL STUDY ON HYDROMYZA LIVENS (FABRICIUS) (DIPTERA, SCATOMYZIDAE), A FLY ASSOCIATED WITH NYMPHAEID VEGETATION DOMINATED BY NUPHAR!) by Th. C. M. BROCK and G. VAN DER VELDE Laboratory of Aquatic Ecology, Catholic University of Nijmegen, The Netherlands ABSTRACT The life history and autecology of the fly Hydromyza livens (Fabricius) were investigated in detail in two Dutch waters. Special attention was paid to its effects on and interrelations with vegetation dominated by Nuphar lutea (L.) Sm. The food plants, geographical distribution, aspects of population dynamics and the morphology of H. livens are described and discussed. Investigations have been made on general behaviour, feeding behaviour, spatial occurrence, sexual behaviour, sex ratio, oviposition, mortality and predation of the adults. The function of the chorion of the egg is discussed. Some experiments have been carried out to study the development of the eggs in relation with humidity, oxygen and temperature. The process of hatching, feeding behaviour, mine patterns, the effect of temperature on feeding behaviour and the mortality of the larvae are described. Data are given of the position in the leaf stalks, emergence and hibernation of the pupae. The life history of H. livens is compared with that of H. confluens Loew, a species bound to Nuphar in North America. INTRODUCTION Nymphaeid vegetations, which are very com- mon and show an optimal development in The Netherlands, have been studied by us during several years. Nymphaeids are aquatic plants with floating leaves, which root in the bottom and have flowers above or floating on the water surface. Because of their structure they have an important function as a habitat for semi-aquatic insects. The species Hydromyza livens (Fabricius, 1794) (syn. Musca livens Fabricius, 1794, and Nupharia rivularis Robineau-Desvoidy, 1830) of the dipteran family Scatomyzidae (syn. Cor- dyluridae, Scatophagidae) is such an insect oc- curring abundantly on the nymphaeids studied, especially on Nuphar lutea (L.) Sm. The larvae of most species belonging to this family live in dung; only a few genera are known to mine in leaves or petioles of plants (Sack, 1937). According to Hering (1926) the larvae of most Hydromyza species mine the leaves and/or peti- oles of Liliaceae and Orchidaceae. 1) Contribution no. 19 of the Nymphaeid Project. 59 Hydromyza livens, however, can complete its whole life cycle on Nuphar lutea (Nymphaea- ceae), one of the nymphaeid species of which the structure and function is a subject of investi- gations in our laboratory. The larvae of H. livens mine the floating leaves of Nuphar. By their consumption of living plant tissue, the lar- vae are an important link in the grazer food chain and decomposition processes connected with Nuphar lutea. Because of our interest in these interrelations the life history and habits of H. livens were studied in detail and a review was made of the literature. Furthermore the results of our study could be compared with those of Welch (1914, 1917) who studied the North American species H. confluens Loew, 1863, of which the life his- tory and habits resemble those of H. livens in many ways. DESCRIPTION OF THE STUDY AREAS Most observations have been made in the Oude Waal near Nijmegen (municipality of Ub- bergen, Province of Gelderland); some addi- tional observations have been made in the Haarsteegse Wiel (municipality of Vlijmen, 60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 Province of Noord-Brabant). In both water bodies pure stands of Nuphar lutea occur. The waters differ, however, in their dimensions, wa- ter quality and bottom composition. The Oude Waal (fig. 1) is an old river branch cut off from the river Waal, where extensive nymphaeid vegetation dominated by Nuphar lutea, Nymphaea alba L. and Nymphoides pel- tata (Gmel.) O. Kuntze occur. It consists of a large shallow water body, 1 km in length and about 250 m wide, and three interconnected ponds (fig. 1, D, E and F). The depth of the large shallow water body varies from 0.5 to 1.5 m; in the dry summer of 1976 it became almost completely dry. About once every two years the area is flooded in winter or spring by water from the river Waal; summer high waters occur only occasionally. The depth of the three inter- connected ponds increases towards their centres to 2.5, 5.5 and 5.5 m, respectively (D, E and F). The bottom of the Oude Waal consists of a pat- tern of clay and sand, sometimes mixed, covered by a sapropelium layer of varying thickness. The Haarsteegse Wiel (fig. 1) originated from two, but not simultaneous, bursts in the dike of the river Meuse. This water body has a surface area of 17.9 ha (44.2 acres) and a maximum depth of 17 m. The nymphaeid vegetation dom- inated by Nuphar lutea and Nymphaea candida Presl, is almost completely restricted to the southern and western parts of the lake, where they are sheltered against wave- and wind ac- tion. The bottom of the Haarsteegse Wiel con- sists mainly of sand (wash-over deposits), but where nymphaeids grow there is a layer of sa- propelium, becoming thicker towards the litto- ral border. The hydrology of the lake is depen- dent on rain and ground water only; the water- level shows only small fluctuations. Apart from the two described areas of inves- tigation several waters in The Netherlands with vegetation dominated by Nuphar have been vis- ited and checked for floating leaves with mines of the larvae of A. livens. GEOGRAPHICAL DISTRIBUTION AND FOOD PLANTS The ecological distribution of H. livens is mainly governed by its affinity for the food plants on which the larvae feed and on which they can complete their life-cycle. In table 1 a literature review is given of the food plants of the larvae of A. livens. According to most authors the larvae mine in the leaves 200m Fig. 1. Left side: the location of the areas studied in The Netherlands (open dot: Haarsteegse Wiel, closed dot: Oude Waal). Right side: bathymetric map of pond F of the Oude Waal, showing the nymphaeid beds (dotted area), the situation of transects studied in the Nuphar zone and the isolated Nymphaea bed (indicated by an asterisk). BROCK & VAN DER VELDE: Hydromyza livens 61 Table 1. Review of the food plants of H. livens as mentioned in the literature. Taxon Nymphaeaceae Nymphaea Nymphaea alba L. Nuphar Nuphar lutea (L.) Sm. Author Collin, 1958 Wesenberg-Lund, 1943 De Meyere, 1940 Hackman, 1956; Vockeroth, 1978 De Meyere, 1895; Eberle, 1943; Gercke, 1877; Brauer, 1883; Hendel, 1928; Gaevskaya, 1966; Grünberg, 1910; Hering, 1926; Heslop-Harrison, 1955; Monchadskii, 1940; Sack, 1937; Seguy, 1934; Schütte, 1921; Van der Velde, 1978; Van der Velde et al., 1978 Nuphar pumila (Timm) DC. Nuphar X intermedia Ledeb. Caltha palustris L. and petioles of Nuphar lutea. This is confirmed in the present study; in all investigated Dutch waters with Nuphar stands floating leaves with mines of H. livens larvae were found. Heslop- Harrison (1955) mentioned H. vens larvae also from the leaves and petioles of Nuphar pumila (Timm) DC. and Nuphar X intermedia Ledeb. (= N.lutea x N.pumila). They were also found by us in floating leaves of N. X interme- dia. It can be concluded from these data that all European Nuphar species are adequate food plants for A. livens. In the botanical garden of Haren (State University of Groningen, The Netherlands) the second author of this paper observed floating leaves of the North-American Nuphar advena Ait. with mines of H. livens. The occurrence of H. livens in America has not been reported. In the literature there is some misunderstand- ing concerning the plant species on which H. livens can complete its life-cycle. De Meijere (1940) reported the occurrence of H. livens lar- vae in floating leaves of Nymphaea alba; in his paper, however, a drawing of a Nuphar leaf with mines of A. livens is given. In Wesenberg- Lund’s “Biologie der Süsswasserinsecten” (1943) an obscure picture of Hydromyza mines in a so-called Nymphaea leaf is shown. Eberle (1943), however, who transplanted Hydromyza larvae from Nuphar leaves to Nymphaea leaves, and also to other Nuphar leaves, concluded that the transplanted larvae fail to develop in floating leaves of Nymphaea. We confirmed this in the laboratory for transplanted larvae in leaves of Nymphaea alba and Nymphaea candida. We never observed Hydromyza larvae in Nym- phaea leaves. Heslop-Harrison, 1955 Heslop-Harrison, 1955 Gaevskaya, 1966; Monchadskii, 1940 According to Gaevskaya (1966), Monchadskii (1940) reported the occurrence of Hydromyza mines in leaves of Caltha palustris L.; this fact, however, is neither confirmed by other litera- ture sources nor observed by the authors of the present paper. Therefore, it can be concluded that in Europe H. livens is strictly bound to Nuphar species. In the Limnofauna Europaea, Vockeroth (1978) reported several distribution areas for H. livens (see fig. 2), viz.: (A) Areas with real records: the western secondary mountain chain (8), the central secondary mountain chain (9), the western low- lands (13), the central lowlands (14), Ireland (17) (Coon River near Partry, North of Lough Mask, 3 July 1982 on Nuphar lutea (L.) Sm. (G. van der Velde), the United Kingdom (18), Northern Sweden (22), Taiga (23), Uzbekistan, Siberia; (B) Areas in which H. livens is expected to occur: the Alps (4), the Carpathians (10), the Hungarian lowlands (11), the Baltic Province (15), the eastern lowlands (16); (C) An area in which A. livens does absolu- tely not occur: Iceland (19). The distribution of H. livens does not extend beyond the confined distribution of the genus Nuphar in Europe. In fig. 2 both the geograph- ical distribution of Nuphar (according to Meu- sel et al., 1965) and H. livens (according to Vockeroth, 1978) are compared. Both distribu- tion areas coincide nearly perfectly. When con- sidering fig. 2, the statement in the Limnofauna Europaea that H. livens does not occur in Ice- land (19) is quite obvious, because Nuphar does not occur there either. On account of the geo- 62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 Fig. 2. Map of the European distribution of both the genus Nuphar (dotted areas) (modified after Meusel et al., 1965) and H. livens (areas with numbers according to Limnofauna Europaea). graphical distribution of Nuphar, H. livens might also be expected to occur in other regions such as Italy, the Balkans and Norway. DEVELOPMENT IN TIME In order to obtain an idea of the population dynamics of H. livens the occurrence in time of adults and larvae was quantified in various ways. Both the larval and adult stage can be eas- ily recognized in the field. Adult H. wens can be found on the upper side of the nymphaeid floating leaves; the flies are not adapted to walk- ing over the water surface, they have to fly from leaf to leaf. To quantify adult H. livens in time the insect-lime method was used (adhesive or sticky trap). The mine tracks of the larvae of H. livens in the floating Nuphar leaves can be easily recog- nized and counted. To quantify the larvae and the number of affected floating leaves in time, the leaf-marking method was used. The quantitative data of adults and larvae are dealt with together in order to maintain to over- all picture of the various generations of H. livens per year. The insect-lime method. — Each week, from June until October, 1977, six undamaged float- ing leaves (five of Nuphar lutea and one of Fig. 3. Adhesive trap (for explanation, see text). BROCK & VAN DER VELDE: Hydromyza livens 63 Nymphaea alba) of the same size were gathered in the Oude Waal, pond F, and covered with in- sect-lime (Tangletrap or Stikem), which has no odour and consequently does not attract or re- pel the insects. These leaves, of which the peti- oles were removed, were attached onto six other floating leaves and then protected from preda- tion by birds by wire netting (with meshes of 3 cm) which was supported by cork floaters (fig. 3). The wire netting with cork floaters was attached with a string to the petiole of the sup- porting leaf. After 24 hours these leaves were taken to the laboratory in plastic bags, so that the catch remained as undamaged as possible. The fauna was washed from the floating leaves with refined petrol and then fixed in 70% alco- hol. The leaves were laid out in transects from the littoral border vegetation to the open water, so that the distribution of the animals over the en- tire nymphaeid zone could be studied (see sec- tion on spatial occurrence on the nymphaeids, further below). Leaf-marking method. — During two years (1976 and 1977) floating leaves and flowers of Nuphar lutea have been marked with 12 mm Rotex-tape on which a number was printed with a Universal 12 printer. One end of the tape was fixed around the petiole by means of a sta- ple and the other end was drawn through the leaf-sinus and then bent over the leaf surface like a flag so that it could be easily read (fig. 4). One plant forms too many leaves to follow the development in detail so that plots of one square metre were made with frames of PVC tubes. The perforated tubing was sunk into the water and held at approximately 15 cm depth by using a string-corner attachment to cork float- Fig. 4. A Nuphar leaf, marked with rotex tape (leaf- marking method). ers. Each frame was anchored (Van der Velde, 1980). Observations and marking of newly un- rolled leaves took place every week during the season. The marking of floating leaves made it possible to study the numbers of mines and the numbers of affected leaves per plot per time. Temperatures of air and water were read regu- larly. Results. — In fig. 5 the leaf-marking observa- tions for 1976 in the Oude Waal are presented; the summer of that year was characterized by a longlasting drought resulting in the emergence of parts of the Nuphar vegetation. The numbers of mines, the numbers of af- fected leaves, and their percentages of the total numbers of floating leaves per plot, show the same tendency (fig. 5). The number of mines and affected floating leaves of Nuphar per plot remained relatively low in the months of May, June and July, while at the end of August and in the beginning of September the numbers of lar- val H. livens clearly reached their maximum. Also the quantity of mines per floating leaf was at its maximum in this period (fig. 5). Figs. 6 and 7 present the data of 1977 in the Oude Waal (marking-method and insect-lime method) and the Haarsteegse Wiel (marking- method). The summer of 1977 was quite normal without droughts. At the end of May and the beginning of June small numbers of H. livens larvae occurred in both waters. The quantity of affected leaves reached a maximum in the Haarsteegse Wiel from the last week of July un- til the second half of September. The larval pop- ulation of H. livens in the Oude Waal was also large in this period with a small peak at the end of July and an absolute maximum in the last week of August. The data concerning the adult stage of H. livens, obtained with the insect-lime method, are summarized in table 2 and fig. 6. Among the total catch of approximately 23,158 specimens (of which 99.4 Hexapoda and 0.6% Arachnida) 1,055 specimens were adult H. livens, ie. 4.56% of the total catch (see for some other re- sults Van der Velde (1978), Van der Velde & Brock (1980) and Van der Velde (1980)). The highest numbers of adult H. livens were caught from 18 August until 8 September. Discussion. — With respect to the insect-lime method one should keep in mind that the result is not only dependent on the size of the A. li vens population, but also on changes in the ac- tivity pattern of adult H. livens as, for example, caused by weather conditions. This is also valid 64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 total number of floating leaves of Nuphar lutea (+) per m? and number of floating leaves with mines of Hydromyza livens (+) per m2 the percentage of leaves with mines of H. livens of total number of en Nuphar-leaves per m? [---) \/\ OUDE WAAL 1976 total number of mines of Hydromyza livens on Nuphar-leaves in a plot of Im? ( --~) A 59 Fand the number of mines observed for the 2 first time in that plot (-—° ) / / 40 ! OUDE WAAL 1976 / miele] lo olulolala wloala| |» S = = & #1 [rollen [un [on [no ala un | [on COUS o ~ n œ =] ~ co Kk EN œ = IS] in n n JAN. FEB. MARCH APRIL MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC. Fig. 5. Above: Total number of Nuphar leaves and absolute numbers and percentages of Nuphar leaves affected by H. livens per m?, during 1976, in the Oude Waal. Middle: Total number of Hydromyza mines in a 1 m? Nu- phar plot and the number of mines observed for the first time in that plot, during 1976. Below: The presence in time of all Nuphar leaves which were affected by larvae of H. livens in a plot of 1 m2. Each line represents the occurrence in time of an affected floating leaf. The numbers above these lines represent the numbers of mine tracks of H. livens in these leaves. BROCK & VAN DER VELDE: Hydromyza livens 65 Table 2. Number of adult H. livens caught with insect-lime on floating leaves from 16 June until 29 September, 1977. For the localities of the different leaves see fig. 1 and the text. Date 1977 Leaf Number of H. livens caught Total number % H. livens number 1 à à AE 6 Total ee de of total catch Broad Nuphar Narrow N. alba P 5 zone Nuphar zone 16-VI = == = = — = 1401 — 23-VI i — — —_— —Z — 1 715 0.14 30-VI ia ee — 3 1 7 1195 0.59 8-VII Ben me ale. 3er 2 34 2234 1.52 12-VII Te Taw 2 6 7 = 29 3168 0.92 21-VII Ze nz: 9 14 22 48 2287 2.10 28-VII Bre aaa 4: 6 — 1 26 1284 2.02 5-VIII 15, 9% 6 — 5 3 38 1580 2.41 12-VIII 7 17 20 — 7 10 61 1578 3.87 18-VIII 54 22 66 1520 20 197 1322 14.90 25-VIII 33 88 64 29 14 13 241 1303 18.50 1-IX 12 15 54 233 12 126 1421 8.87 8-IX 16 68 66 14 21 5 190 1876 10.13 15-IX dl 15 3. 3 5 44 762 5.77 22-IX A et — — — 13 584 2.23 29-IX — — — — — — — 448 — Total number per leaf 171 265 324 87 134 74 1055 23158 4.56 for the other insects, so that percentages of H. livens with respect to the total catch per date have been calculated (table 2). The absolute numbers and percentages showed the same trend, so that the catch gives a realistic picture of the adult population development in time. When considering figs. 5, 6 and 7 it is clear that the development of the population of both adult and larval A. wens is correlated with the development of the floating leaves of Nuphar lutea. The first mine tracks on the floating leaves can be recognized early in the growing season of Nuphar, and at the end both the total numbers of floating leaves per plot and the numbers of H. livens diminish with the same tendency. From the data obtained it can be concluded that H. livens has two or three generations per year in the studied waters. The third generation of the year before, which has hibernated in the pupal stage, occurs in May and June and the number of animals is small. The experiment with the insect-lime started too late to show this generation in the adult stage; adult flies, howev- er, were caught by us with a hand-net in this pe- riod. In the Oude Waal adults of H. livens were observed in 1975 from 5 June until 29 Septem- ber, and in 1977 from 23 May unul 22 Septem- ber. The first generation in the Oude Waal in 1977 was present as larvae in May and the first week of June; adults of this generation were present at the end of June and in July. The max- imum of adults was on 21 July. The second gen- eration was present as larvae in the second half of July and the beginning of August. The maxi- mum of adults was reached on 25 August. The third generation was present as larvae at the end of August and the first half of September. This generation hibernates as pupa and the adults of this generation were present in the next spring. The data from the Oude Waal in 1976 do not indicate so clear the occurrence of three genera- tions, that year, however, was characterized by a longlasting drought. This may have caused the establishment of only a minor second genera- tion. The population data from the Haarsteegse Wiel show two peaks (fig. 7), a minor one in the second half of May and the first weeks of June (first generation) and a large one in the second half of July, August, September and October. That the second peak is spread out over such a long period probably indicates that the second and third generation overlap, resulting in one broad peak only, as shown in fig. 7. It can be concluded from the numbers of af- fected leaves per plot that in the Oude Waal in 1977 the second larval generation is ca. 6 times larger than the first generation while the third 66 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 % 2 e——e water temperature 22 œ—0 alr temperature Re 20 9: he EN, e OUDE WAAL Sf À N LL 36 Nuphar lutea 32 OUDE WAAL 1977 1977 number of leaves with mines Hydromyza livens OUDE WAAL 1977 200 Hydromyza livens Adults pol OUDE WAAL 1977 Fig. 6. Course of water and air temperature, presence of floating leaves and flowers of Nuphar lutea in a plot of 1 m?, numbers of floating leaves affected by Hydromyza larvae in that plot and numbers of adult A. livens caught with insect-lime on the floating leaves during 1977 on the Oude Waal. Brock & VAN DER VELDE: Hydromyza livens 67 24 e—e water temperature 22 / \ 20 HAARSTEEGSE WIEL Jor Seen, 40 36 Nuphar lutea 32 HAARSTEEGSE WIEL J F M A M J 1977 number of leaves with mines 24 2 ; ydromyza livens 20 HAARSTEEGSE WIEL 1977 J F M A M J Fig. 7. Course of water temperature, presence of floating leaves and flowers of Nuphar lutea in a plot of 1 m? and numbers of floating leaves affected by H. livens in that plot during 1977 on the Haarsteegse Wiel. generation is ca. 8 times larger than the first one. In the Haarsteegse Wiel in 1977 both the sec- ond and third larval generations are ca. 7 times larger than the first one. The first, and under favourable conditions al- so the second, generation complete their life-cy- cle within the growing season of Nuphar while the pupae of the third (under bad environmental circumstances probably also those of the sec- ond) generation hibernate and form the parents of the first generation, the next year. That there are more generations per year is also supported by differences in behaviour and morphological characteristics of the larvae and pupae of the various generations, as will be dis- cussed later. Schütte (1921) reported that sum- mer and winter pupae are both present in the 68 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 beginning of August indicating that two genera- tons overlap each other (see section on the pu- pa). The emergence of adults from summer pu- pae occurs in the same growing season of the food plant while the winter pupae hibernate. Schütte (1921) also suggested the possibility of the existence of three generations. ADULTS Morphological description H. livens (fig. 8 and 9) is a fly of 7—9 mm length; the females (8—9 mm) are larger than the males (7—8 mm) (20 specimens of each sex were measured). The flies have a remarkably bright-yellow frons, chestnut-brown eyes and black antennae. The thorax and the abdomen are dusty blue-grey. The mesonotum has five more or less visible, grey-brown, longitudinal stripes. The legs are for the larger part red- Fig. 8. Dorsal view of adult H. wens (wings omitted). Fig. 9. Lateral view of adult H. livens (female). BROCK & VAN DER VELDE: Hydromyza livens 69 brown, with yellow-brown ginglymae; the fem- ora are partly blackish, tinged with a dusty blue-grey. The wings are of a translucent dusty- grey while the veins are slightly brownish. Veins R 4 + 5 and M are slightly convergent to- wards the apex; vein 1A does not reach the mar- gin of the wing. The relatively very short bris- tles on the body and the legs and the three very small proclimate frontal bristles and the single sternopleural bristle are characteristic for Hydromyza. For further descriptions, see Grinberg (1910), Séguy (1934), Sack (1937), Hackman (1956), and Collin (1958). General behaviour In the areas studied the flies usually occurred on the floating leaves of the nymphaeids Nym- phaea alba, Nymphaea candida, Nymphoides peltata, but preferably on those of Nuphar lu- tea. In spite of the fact that these insects have well-developed powers of flight they are not found at any great distance from the nym- phaeids. They are difficult to capture and move restlessly around over the floating leaves, alter- nately walking or flying. They fly only short Table 3. Temperature measurements in °C with a Wallacoy Universal Thermometer (GST 32; probes Ni-101x and Ni-106) in the flowers and on the floating leaves of Nuphar lutea and of the ambient water and air. FLOATING LEAVES Date (1977) Tw ue 8-VI 16.0 18.0 8-VI 16.5 18.5 23-VI — 22.6 6-VII 24.5 28.2 8-VII 23.2 DA 8-VII 23.4 26.1 8-VII 2055 25.5 7-VII — 24.2 7-VII — 22.0 29-VI 19.2 17.8 29-VI 19.1 17.5 29-VI 19.0 19.1 29-VI 11922 1725 FLOWERS Date (1977) Tw Ta 8-VI 16.0 18.0 23-VI — 22.6 23-VI _ 22.0 6-VII 24.5 28.2 8-VII ZON 24.1 8-VII 23.4 26.1 7-VII 20.0 23701 7-VII 22.8 20.0 7-VII 23.0 21.0 29-VI 19.0 19.1 29-VI 19.2 125 29-VI 17.2 17.8 29-VI 19.1 17/5) Tl TI-Tw TI-Ta weather type 19.2 3.2 1.2 sun 195 3.0 1.0 sun 25.4 = 2.8 sun 29.0 4.5 0.8 sun 25.0 1.8 0.9 bright sun 26.5 Sell 0.4 brightsun 25.0 1.5 —0.5 _ bright sun 24.0 — —0.2 heavily clouded 23.0 — 1.0 clouded 19.1 0.9 1.3 drizzle 192 0.1 LAN CAE 11929 0.9 0.8 rain 19.5 0.3 2.0 rain The Tf-Ta weather type 19.0 1.0 brightsun 26.9 4.3 sun 272 DA sun 33.0 2.8 bright sun 26.8 2.7 bright sun 28.0 1.9 bright sun 23.3 0.2 heavily clouded 212 12 heavily clouded 21.0 0.0 heavily clouded 19.0 = drizzle 18.6 Ll drizzle No 153 drizzle 17.4 =0.1 heavy rains Tw = water temperature in °C measured 3 cm below water surface. Ta = air temperature in °C measured 50 cm above water surface. Tl = temperature upper surface floating leaf measured near midrib. Tf = temperature of flower near stigma. 70 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 distances to reach other floating leaves or flow- ers in the neighbourhood; even when they are disturbed they make only short flights, seldom to open water or to the shore. Especially under sunny weather conditions the flies are very ac- tive on the floating leaves or in the flowers of Nuphar performing feeding, walking, flying, copulating, polishing, and sunning. The nymphaeid floating leaves and flowers grow in maximum exposure to sunlight; conse- quently they can reach somewhat higher tem- peratures in comparison with water and air, as is illustrated in table 3 for Nuphar lutea. Such high temperatures probably stimulate the activity of the flies. During bad weather conditions, e.g. heavy rain, they seek shelter under the aerial leaves of Nymphaea alba (leaves of this species often project entirely or partly above the water sur- face), in the flowers of Nuphar, or between the littoral helophytes, staying there more or less motionless. Feeding behaviour According to Eberle (1943) A. livens feeds in the adult stage on small insects occurring on the floating leaves of the nymphaeids, like Hydrel- lia (Ephydridae) and Chironomidae. These small Diptera are squashed by the proboscis of this predatory fly (fig. 10) and the body fluids are consumed. On the floating leaves nearly all insects which move are attacked by H. livens, but only small or soft skinned insects such as Nematocera are killed. In the laboratory ten H. livens specimens were kept in an aquarium with Nuphar leaves together with some insects, which also occur abundantly on the nymphaeids, viz., the chiro- nomid Cricotopus sylwestris (Fabricius) and the ephydrid flies Hydrellia griseola Fallén and Notiphila brunnipes Robineau-Desvoidy. The chironomids and the tiny Hydrellia were cap- tured and consumed by H. livens, the larger Notiphila (2.8— 4.5 mm) were attacked but not harmed, while dead, squashed Notiphila’s were consumed. In the areas investigated adult H. livens was regularly observed to occur in the flowers of Nuphar lutea, especially in those just opened, but never in the flowers of Nymphaea or Nym- phoides. The flowers of Nuphar possess nectaria situated on the outer side of the small petals; they produce nectar containing glucose and fructose in equal amounts as could be proved with enzymatic tests. The production of nectar occurs especially on the first day of flowering; the flower has then a strong brandy scent (Van der Velde & Brock, 1980). The flowers of Nu- phar also have many anthers, which produce abundant pollen. Eggs of the fly Notzphila brunnipes and specimens of the aphid Rhopalo- siphum nymphaeae (L.) are usually present in the Nuphar flowers (Van der Velde et al., 1978; Van der Velde & Brock, 1980). Because of the regular occurrence in Nuphar flowers the flies were suspected of obtaining food products from them. It was however difficult in the field to ob- serve the shy flies in the flowers without dis- turbing them. In the laboratory the ten Hydromyza speci- mens were offered food in the form of Notiphila eggs, the aphid Rhopalosiphum nymphaeae and moist sugar as a substitute for nectar. The flies were repeatedly observed licking the moist sug- ar. In most cases this behaviour continued too long to be interpreted as a mere random inspec- tion. Further some Notiphila eggs and the aphids were squashed by the proboscis of the flies and consumed. Probably adult H. livens feeds in the Nuphar flowers on nectar, Notiphi- la eggs and small insects such as aphids. To check whether adult A. livens also con- sumes pollen, the intestines of some flies cap- tured in Nuphar flowers were studied under the microscope; in the intestines diffuse material and micro-organisms could be recognized but no pollen grains. The adult flies are numerous and it can be as- sumed that they play a role as pollinators of Nu- phar. A regular flight pattern from flower to flower was never observed. However, by their regular occurrence in the flowers and by their activities, the flies certainly can cause pollina- tion. Spatial occurrence on the nymphaeids Method. — To study the distribution of adult H. livens over the nymphaeid zone, floating leaves with insect-lime (see insect-lime method) were laid in transects from the littoral border vegetation to the open water (fig. 1). Three Nuphar leaves were laid in the north- ern part of pond F; leaf 1 near the broad littoral border vegetation, consisting of Typha angusti- folia L., Rumex hydrolapathum Huds. and Scir- pus lacustris L., here the floating leaves of Nu- phar lutea covered up to 60%; leaf 2, 6 m from leaf 1, in the centre of the Nuphar zone (maxi- mum coverage 70%); leaf 3, 12 m from leaf 1, at the margin of the Nuphar zone with the open Brock & VAN DER VELDE: Hydromyza livens Fig. 10. Scanning electron microscope photograph of the head of H. livens. Note the proboscis by which small preys are squashed. Fig. 11. Scanning electron microscope photograph of hydrophobous hairs on the body of H. livens by which the fly avoids wetting. water, where Nuphar covered up to 60% of the surface. Two leaves were laid in a narrow Nuphar zone at the southern side of pond F; leaf 4 near Table 4. Absolute numbers and percentages, of adult H. livens and its preys caught on floating leaves with insect-lime. For the localities of the different leaves see fig. 1 and the text. Absolute numbers per leaf and per taxon Leaf number 1 2 3 4 5 6 total Hydromyza livens 171 265 324 87 134 74 1,055 Nematocera spp. 9503112882777, 734 895 599 6,138 Chironomidae spp. PRI OA) 2589) 663 705 463 5,267 Cricotopus spp. 83 288 1,790 437 358 142 3,098 Hydrellia spp. 346 293 137 180 154 317 1,427 broad Nuphar zone narrow N. alba Nuphar zone Percentages per leaf and per taxon Leaf number 1 2 3 4 5 6 total % Hydromyza livens 16.2 23 307 82 127 7.0 100 Nematocera spp. 5.8 13.3 44.6 12.0 ME 9.8 100 Chironomidae spp. be lien GOP Mo 194 8.8 100 Cricotopus spp. Dell 9975728 file Wl 4.6 100 Hydrellia spp. Die HOLD 9.6 126 108 222 100 broad Nuphar zone narrow N. alba Nuphar zone the littoral helophyte vegetation, which borders on grassland, consisting of a narrow zone of the same plant species mentioned for the northern side, but also with Mentha aquatica L. (maxi- 72 TiJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 mum coverage of Nuphar 50%) and leaf 5 at 3 m distance from leaf 4, at the margin of the Nu- phar zone with the open water (maximum cov- erage 40%). Leaf 6, a Nymphaea leaf, was laid in an iso- lated vegetation of Nymphaea alba, surrounded by open water and at 6 m distance from the shore at the western side of the pond (maximum coverage of Nymphaea 85%). Results. —;The general distribution of the flies within the nymphaeid zone can be read from table 2, by comparing the results from the different leaves. The flies appeared to have a distinctly higher abundance on the floating leaves bordering the open water (leaves 3 and 5); the lowest number of flies has been caught near the littoral helo- phyte vegetation (leaves 1 and 4). Adult H. livens also showed a higher abun- dance on the broader Nuphar zone (leaves 1, 2 and 3) in comparison with the narrow one (leaves 4 and 5). The lowest number of H. livens specimens was caught on the Nymphaea leaf (leat 6), which is comparable with the Nuphar leaves 3 and 5 because of their similar location near the open water. This distribution suggests a prefer- ence for Nuphar leaves. Discussion. — The higher abundance of adult H. livens on the nymphaeids bordering the open water may be explained by the fact that there is more food available for this predatory fly on floating leaves situated towards the open water. As mentioned before, particularly Ne- matocera and Hydrellia are food objects. With the insect-lime method the spatial dis- tribution of these predated insects could also be determined; the data of the most important preys are presented in table 4. When consid- ering the spatial distribution, the absolute num- bers and percentages of Nematocera and H. lı- vens specimens caught on the floating leaves show the same trend. By far the largest part of Nematocera specimens is formed by the genus Cricotopus, mostly C. trifasciatus (Meigen) and C. sylvestris (F.). The spatial distribution of Hy- drellia species does not show the same tendency as that of H. livens. The numbers of Hydrellia specimens on the floating leaves, however, are much smaller than those of the Nematocera (table 4). The higher abundance of adult H. livens on Nuphar leaves in comparison with those of Nymphaea may be explained by the facts that the quantity of the Nematocera is relatively low on Nymphaea (leaf 6, table 4) and that the fe- males of H. livens deposit their eggs on Nuphar leaves only. Sexual behaviour and sex ratio As already mentioned adult H. livens reso- lutely approaches all insects in motion on the floating leaves. This behaviour has the function to collect food but also to find a suitable part- ner. The fly can be observed regularly in copu- lation on the floating leaves of the nymphaeids mentioned and in the flowers of Nuphar. When two Hydromyza specimens meet, mostly they attempt to mate, resulting in copu- lation in the case of two adequate partners, or in a skirmish in the case of e.g. two males. A. li- vens mostly does not show an elaborate and dis- tinct mating behaviour. Sometimes the male, during the posturing phase of mating, toddles towards the female, while repeatedly and quick- ly spreading the wings sidewards. If the female accepts the invitation, she spreads the wings permitting the male to mount. It also has been observed that a male approached a female in a more aggressive way leading to copulation. During the insemination the females often con- tinue feeding, polishing and even walking. Males and females of H. livens can easily be separated from each other because they differ in size and in position of the genital aperture. The genital aperture is situated on the ventral side of the abdomen in the males and near the caudal tip of the abdomen in the females. Some samples from the insect-lime catch on the floating leaves were studied for calculating the sex ratio. The results are summarized in table 5. Usually more males than females were caught on the floating leaves; the percentage of males was 61.4. Maybe the males are more active, resulting in larger quantities of males in the catches with insect- lime. Another explanation might be a higher mortality of the females. Deposition of eggs As described by De Meijere (1940) and Eberle (1943), females of H. livens deposit eggs on the underside of the floating leaves of Nu- phar lutea. In the study areas normally no eggs were found under the floating leaves of other nymphaeids although Nuphar, Nymphaea and Nymphoides locally intermingle in the same beds. Apparently the female has the ability to recognize the food plant in the presence of oth- er nymphaeids. Only during the dry summer of 1976 eggs were found under the floating leaves BROCK & VAN DER VELDE: Hydromyza livens 73 of Nymphaea and Nymphoides as well, perhaps due to the fact that the oviposition sites were strongly diminished. Observations in the field suggest that the fe- males inspect the floating Nuphar leaves before oviposition; we often observed the females walking a number of times from the leaf margin to the midrib and back again. Probably the fly in this way inspects the condition of the leaf. When a floating leaf is affected, in whatever way (even by Hydromyza larvae) the leaf is re- jected and oviposition does not take place. From fig. 5 it can be concluded that most Nu- phar leaves become affected by larvae of A. li- vens when the leaves have just reached the wa- ter surface. Only at the end of the season, when the numbers of adult 7. Iivens are at their maxi- mum, also older undamaged leaves (fig. 5; leaves nr. 42, 48, 81 and 65) are affected by these larvae. From these data it can be con- cluded that oviposition preferably takes place on just unrolled Nuphar leaves. When the female has selected an adequate Nuphar leaf, she crawls over the leaf edge, goes into the water, and than proceeds upside down, onto the underside of the floating leaf where the egg is deposited. After approximately 20 sec- onds the female crawls via the leaf out of the water again and nearly immediately flies away. Experiments in which adults were submerged showed that under such conditions the flies rise to the surface if opportunity affords, because their specific gravity is less than water. They stay under water only by clinging to a sub- merged object (the floating leaf). When going under water a good supply of air clings around them like a coat. The fly has a dense body-cover of small hydrophobous hairs (fig. 11). When the flies come to the surface again, they immedi- ately lose the air coat and are apparently as dry as if they had never been under water. Because of the air coat, which they take down and the ease with which they resist wetting, oviposition can take place under water without difficulties. A female H. livens deposits only one egg at a time per floating leaf. Once it was observed that a fly went down under a floating leaf on which afterwards no egg was found; when examining the underside of the leaf it appeared that a lot ot periphyton occurred there, probably inhibiting Oviposition. To find out how many eggs one female of H. livens can deposit, a number of females were dissected and examined for eggs; 16 eggs were found at the most in the ovaries and oviducts, of which eight eggs were in a more or less mature state. The exact way of oviposition on the under- side of the floating leaves could not be observed in the field without disturbing the flies. As al- ready reported by Eberle (1943) (and this was also observed by us in the laboratory) the fe- males of H. livens, which were kept in aquaria, did not deposit their eggs under the floating leaves of Nuphar. Oviposition took place above the water surface on the upper side of the float- ing leaves and even against the glass walls of the aquaria. These eggs, which were also deposited singly, were consumed by the adults. Probably stress caused by the imprisonment changed the oviposition behaviour. The flies in the aquaria did not survive very long, only one or two days; they easily drowned. Probably the circumstances in the aquaria (e.g. high humidity of the air) dimin- ished their resistance in getting wet. Eberle (1943) pointed out that the position of the deposited eggs of H. livens under the float- ing leaves of Nuphar gives much information about the oviposition of the females when being under water. In the second half of August 1979, 20 more or less undamaged floating leaves of N. lutea were collected in the Oude Waal. The quantity of eggs per floating leaf varied from 0 to 15. Be- cause each female only deposits one egg under a particular leaf, more females of H. livens must be responsible for the number of eggs per leaf. The maximum number of eggs found by us un- der a Nuphar leaf was 29. Of 100 eggs deposited by H. livens on the collected Nuphar leaves the positions were recorded by measuring the dis- tance to the midrib and the petiole (attachment place to leaf blade). In fig. 12 the positions of these eggs are projected on a single floating leaf of Nuphar. The graph of fig. 13 shows the dis- tance between these eggs and the midrib of the Nuphar leaf measured along the longitudinal ax- is of the eggs. According to these data and those of Schutte (1921) and Eberle (1943) several regularities in the positions of the eggs appeared to exist, viz.: (a) The broad rostral end of the egg is situ- ated mostly towards the midrib. In dissected fe- males the more pointed caudal end of the egg lies more towards the oviduct, during oviposi- tion this end leaves the abdomen before the broader end. During oviposition, therefore, the female must be directed with her head towards the midrib. 74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 IN N ; Ex x SAS! DI 2 Fig. 12. The positions of 100 eggs of H. livens (collected from 20 leaves) projected on the underside of a single Nuphar leaf. (b) The longitudinal axis of the egg is mostly situated parallel to the lateral leaf nerves, mostly between two of them. (c) Most of the deposited eggs can be found near the basis of the midrib. (d) Most of the eggs are deposited approxi- mately 7—10 mm from the midrib (fig. 13). From these data it can be concluded that a fe- male of H. livens mostly crawls into the water via the edge of the broadest part of the floating leaf where the lateral nerves are dichotomously branched. When under water, clinging upside down to the leaf and covered with an air coat, the female crawls towards the midrib, guided by a lateral leaf nerve, which is translucent. Proba- bly the visual observation or touching of the midrib is a strong stimulus for oviposition be- cause the distances between the eggs and the midribs mostly were approximately the length of the flies. These conclusions agree with those of Eberle (1943). Spatial occurrence of egg deposition In order to find out where egg deposition mostly takes place within the nymphaeid zone, both in the Oude Waal and Haarsteegse Wiel, in 1977 regularly 15 floating Nuphar leaves were collected at random at three sites, viz., 15 near BROCK & VAN DER VELDE: Hydromyza livens 75 the littoral border vegetation, 15 in the middle of the Nuphar zone and 15 near the open water. Leaves were collected every month from May until September in the Oude Waal and every two weeks from June unul September in the Haarsteegse Wiel. The eggs and larvae occur- ring under and in the leaves were counted per site per date. In the Oude Waal in total 62 larvae and eggs were counted on the leaves bordering the litto- ral helophyte vegetation, 38 on leaves from the middle of the Nuphar zone and 65 on floating leaves from the location near the open water. In the Haarsteegse Wiel, in total 81 larvae and eggs were found on the leaves near the littoral border vegetation, 79 on leaves from the middle of the Nuphar zone and 94 on leaves bordering the open water. It can be concluded that oviposition in the Oude Waal takes place more often under leaves at the two edges of the Nuphar belt than under leaves in the middle of the Nuphar zone. The results from the Haarsteegse Wiel show this trend less clearly. From the data obtained with the insect-lime method it appeared that the adults were defi- nitely more abundant on the nymphaeids bor- dering the open water because of the higher abundance of food organisms there. The large number of eggs and larvae under and in floating leaves of the sites near the open water can be ex- plained in this way. The high frequency of egg deposition under floating leaves bordering the littoral helophytes may be explained by the fact that wind and wave action are less pronounced here. 9 15 012345 6 7 8 9 10111213 14 15 16 17 18 19 20>20 mm. DISTANCE Fig. 13. The distances of 100 eggs of H. livens to the midribs of the floating leaves on which they were de- posited, measured along the longitudinal axis of the eggs (see also fig. 12). Mortality and predation As may be concluded from fig. 6, the num- bers of adult H. Iivens decrease at the end of the growing season of Nuphar. The mortality of adults may have several reasons, e.g. fallıng temperatures, diminishing of the floating leaf area (figs. 5, 6 and 7) and lack of food (table 2). Exhaustion of females and risks under water during oviposition, such as predation by fish, might also cause mortality; the sex ratio data mostly showed a majority of males (table 5). Af- ter heavy rain the population size of adult H. /i- vens seems to be diminished. Although the flies have hydrophobous hairs, they easily drown when their skins become wet. Probably a long- lasting contact with water or even very humid conditions (aquaria) diminish their resistance against getting wet. Predation on adult H. livens has never been observed by us. However, Odo- nata, swifts (Apus apus (L.)), swallows (Hirun- do rustica L., Delichon urbica (L.)), terns (Ster- na hirundo L., Chlidonias niger (L.)) and the Green frog (Rana esculenta (L.)) prey on the in- sects from the floating leaves and thus can occa- sionally catch H. livens. THE EGG Morphological description The eggs of A. livens have a length of 1.9— 2.0 mm and a width of 0.5—0.6 mm. Freshly laid eggs are yellow-white in colour, a few hours before hatching they become darker. The eggs have the appearance of a boat (fig. 14a), with differently shaped ends: the caudal end is bluntly pointed, while the broader rostral end is more rounded and contains the micropyle. A straight, acute carina extends longitudinally from end to end and runs medially along the bottom of a rather deep, broad, longitudinal fossa (fig. 14b), dividing it into two similar parts. This divided fossa is laterally bordered by two other longitudinal carinae, which run al- most parallel to the median carina, converging and uniting with the ends of the egg. The eggs are always deposited singly under the floating leaves of Nuphar and are firmly fixed to the substrate so that it is nearly impossible to sepa- rate them from a floating leaf without damage to the chorion or the leaf tissue. Apparently a small amount of sealing fluid accompanies the egg at oviposition causing the firm attachment to the floating leaf. Eggs have been studied by means of scanning 76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 Table 5. Sex distribution and the percentage of males in samples of H. livens caught with insect-lime. Date (1977) Number of Males Females percentage specimens of males examined 30-VI 2 1 1 50.0 12-VII 13 11 2 84.6 21-VII 19 12 7 63.2 28-VII 7 3 4 2 5-VIII 25 13 12 52.0 12-VIII 24 6 18 2570 25-VIII 125 82 43 65.6 1-IX 91 63 28 69.2 8-IX 5 0 5 0 Total 311 191 120 61.4 electron microscopy; in this way the external and internal appearance of the chorion could be studied (fig. 14). The external surface of the chorion, especially near the carinae, appears more or less reticulate, being composed of poly- gonal units (fig. 14b). The reticulation is elon- gated in the direction of the long axis of the egg and within the netlike structures many pores can be recognized (fig. 14b and 14c). To study the internal structure of the chorion the eggs were sliced with a razorblade and ex- amined with the scanning electron microscope (fig. 14d, e and f). The chorion appeared to pos- sess cavities, which are apparently in contact with the pores in the external walls. The cho- rion is distinctly thicker at the carinae and the depression between them than on the lateral sides, as shown in fig. 14d. The function of the chorion The chorion protects the eggs against certain predators but also can function as a plastron. According to Hinton (1967, 1971) the terrestrial eggs of the majority of insects have structures in the chorion which have the function to hold air to maintain respiration on becoming wet, e.g. by a rain shower; the chorion of many aquatic and semi-aquatic eggs is solid. It appears from the scanning electron micro- graphs (fig. 14) that the chorion of the aquatic egg of H. livens contains cavities. These cavities possibly hold an air layer which functions as a plastron. Hinton (1960) has shown that a sys- tem of hydrofuge structures in the chorion can form the architecture for a permanent, un- shrinkable, physical gill, a plastron. An egg with a plastron can remain immersed indefinitely and obtains the oxygen it requires from the ambient water, provided that the water is well-aerated. Normally the eggs of H. livens are deposited under the floating leaves of Nuphar so during their development into larvae they are immersed in the extreme upper layer of water. In natural habitats, such as the Oude Waal, the water un- der the floating leaves is well-aerated. On 25 August 1977 the amount of oxygen in the water of the Oude Waal was measured with an oxygen meter (Y.S.I. model 151A). Directly under the floating Nuphar leaves the measured oxygen saturation percentage was 78%. At depths of 50, 100 and 150 cm (near the bottom) the oxy- gen saturation percentages diminished and were 75, 70 and 45% respectively. Eggs with a plastron are able to survive under the floating leaves because of the well-aerated conditions. In the laboratory, eggs of A. livens normally developed into larvae when floating leaves with eggs were kept floating in aquaria and also, when they were kept under very humid condi- tons outside the water. The eggs, however, soon desiccated when floating leaves with eggs were kept in a dry environment outside the wa- ter. Also when floating leaves with eggs were kept on the bottom of an aquarium, 20 cm be- low the water surface in non-aerated water, the eggs had not yet hatched when the leaves had already decayed; probably the eggs did not de- velop because of depletion of oxygen. It can be concluded that the eggs develop in well-aerated water or in very humid air; the chorion, howev- er, is not adapted to prevent desiccation and probably forms the architecture for a physical gill or plastron. Fig. 14. Scanning electron microscope photographs of the egg stage of H. livens: (a) overall view, note the mi- cropyle at the frontal end; (b) detail of the chorion near the median carina; (c) detail of the chorion showing polygonal units and pores; (d) transversal section, showing the internal structure of the chorion near a carina; (e) detail of the internal structure of the chorion, showing many pores and cavities; (f) detail of the innerside of the chorion also showing pores. 78 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 DAYS 15 10 5 0 8 12 16 20 24 °c TEMP Fig. 15. Duration of the development of A. livens eggs into larvae in relation with temperature. Development in relation to temperature Eberle (1943) already discussed the fact that the development of eggs into larvae is depen- dent on temperature. In spring, when the water is relatively cold, the duration of the devel- opment can last out up to two weeks, while in summer, when the water is warmer, the larvae hatch after 3 to 4 days after the time of egg de- position (Eberle, 1943). It is difficult to study the development of the eggs in relation to temperature under controlled circumstances because it is difficult to obtain enough eggs of the same age which are freshly deposited by the females of H. livens. As al- ready discussed one can only sporadically ob- serve the oviposition in natural habitats, while the flies in captivity do not deposit their eggs in the normal way under the floating leaves. In or- der to obtain eggs which were more or less of the same age, 25 undamaged, young leaves of Nuphar, under which no eggs were deposited, were marked in the Oude Waal at the end of August 1979. After 24 hours these leaves were collected and 19 eggs of H. livens were obtained from them. The differences in age of these eggs could be 24 hours at most; the temperature of the Oude Waal water was at that time 18 °C. In the laboratory the floating leaves with eggs were kept in aquaria and placed in different cli- mate chambers which had constant tempera- tures of 8, 12, 16, 20 and 24 °C. Temperatures within this range had also been measured in the Oude Waal (fig. 6). In the climate chamber of 8 °C three eggs were placed, in the other cham- Fig. 16. Photograph of a H. livens larva sampled from a leaf blade. bers four each. By daily observations the devel- opment of the eggs into larvae was followed (see Ho) The data thus obtained confirm that the de- velopment of the eggs is indeed dependent on temperature and that the development goes faster at temperatures which are normally mea- sured in summer in comparison with colder cir- cumstances in spring and autumn. Below a cer- tain temperature, between 8 and 12 °C, the eggs do not develop at all into larvae. The large dif- ferences in developmental rate of the eggs, espe- cially in the climate chambers of 12°C and 16 °C, are probably caused by the way the eggs were obtained, viz. at a temperature of 18 °C. THE LARVA Morphological description The larva of H. livens (fig. 16) is yellow- white in colour with the exception of the black cephalopharynx skeleton (fig. 18), the brown, kidney-shaped, frontal stigmata and the dark terminal stigmata (fig. 17). The anterior stigmata consist of short pro- cesses, each with several buds; the total number of buds on one stigma being about 200 (De Meijere, 1895). According to Schütte (1921) there is no difference between the number of stigmatic buds of larvae and pupae of the differ- ent generations. In the summer larvae (in our case the first and second generation) the diame- ters of these buds are smaller, however, than in the winter larvae (in our case the third genera- tion). The anterior stigmata are not yet visible on freshly hatched larvae. In the translucent body the long, green intes- tine, the main tracheae and the very well devel- Brock & van DER VELDE: Hydromyza livens 79 Fig. 17. Terminal stigmata (spiracular disc) of a H. Iivens larva. Fig. 18. Cephalopharynx skeleton of a larva of H. livens. oped salivary glands are visible. The digestion tract of the full-grown larva is at most 8.5 cm long (about eight times the total length of the larva) (Schütte, 1921). The body is cylindrical and elongate, and consists of 12 more or less visible segments. Some intersegmental grooves are very distinct. The anterior end of the body is moderately rounded with the exception of the acute, emer- gent teeth of the cephalopharynx skeleton. Pos- teriorly, the body is bluntly tapering to the more or less rounded or acute chitinous projec- tions of the terminal stigmata (fig. 17). The integument is covered with pointed, chitinous projections which are directed back- wards. These projecuons are rather uniform in size and shape over the body except on the ante- rior margin of the first thoracic segment, where they are larger and more distinct. On the younger larvae, bristle-like projections can be recognized above the mouth-hooks. In fig. 19, scanning electron microscope photographs of heads of larvae of H. livens are presented. The larva of fig. 19a was collected from a floating leaf of Nuphar lutea and was 2—3 days of age. The larva of fig. 19b was collected from a peu- ole of a floating Nuphar leaf and is a few days older. According to Schütte (1921) the larvae of the summer generation measure 9.5 mm in length at a width of 2.5 mm, while the length and width of larvae of the last generation measure 11 and 3 Fig. 19. Scanning electron microscope photographs of the heads of two H. livens larvae, sampled from a leaf blade (a) and from a petiole (b). 80 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 mm, respectively. According to Eberle (1943) In the Oude Waal full-grown larvae of the | full-grown larvae of the first generation mea- last generation were 10.5— 13.0 mm in length, | sure 9 mm in length, while the mature larvae of witha width of 1.5—2.5 mm. the last generation can reach a length of 11 mm. According to Schütte (1921) there are four TT: | ) p IT Ht A ©: (VI) [7 CT Lo US Fig. 20. Daily length increase of mine tracks of three H. livens larvae in Nuphar leaves at a temperature of 20 °C (A), 16 °C (B) and 12 °C (C), respectively. Each number corresponds with a day. Transversal section of a part of a Nuphar leaf with a mine track (arrow) of a just hatched larva (D) and an older larva (E) in the leaf blade. Brock & VAN DER VELDE: Hydromyza livens 81 mine tracks of H. livens larvae. instars; this author also gives an extensive anatomical description of the larva and its way of nutrition. The process of hatching from the egg Shortly before the time of hatching, the egg becomes darker in colour, indicating that the larva within the egg is fully developed. The place of emergence of the larva through the egg shell is very constant. With the aid of relatively large, acute, chitinous mouthparts the larva makes an exit through the egg shell on the side next to the leaf. The position of the exit hole is determined by the position of the larva in the egg. So the exit is made near the blunt, broad, rostral end of the egg, which is mostly directed towards the midrib of the floating leaf. Via this hole the larva bores directly through the lower epidermis into the mesophyll of the floating Nuphar leaf. After the process of hatching the empty egg shell remains sticking to the leaf, closing off the exit, so that water is not able to enter the mine gallery in the leaf. General feeding behaviour In the investigated areas the larvae of H. h- vens feed on the floating leaf and petiole tissue of Nuphar lutea. During the early period of feeding only the spongy parenchyma of the floating leaf is consumed, while in a later stage, when the larva has increased in volume, also the palissade parenchyma of the mesophyll is eaten (fig. 20d and 20e). While eating through the floating Nuphar leat the larva produces a mine gallery in it, and af- terwards also in the petiole where the larva eats itself spirally downwards (fig. 21 and fig. 22c). With the aid of the large acute mouth-hooks Fig. 22. (A) Just hatched larva of H. Iivens in its mine track and the empty egg shell. (B) Larval consumption by day (white) and by night (black and marked with a) in a Nuphar leaf. (C) Mine tracks of H. livens in a pet- iole of a Nuphar leaf. (Modified after Eberle, 1943.) the larva loosens the plant cells while making mowing movements with the head. According to Schütte (1921) the larvae of H. livens secrete digestion enzymes which dissolve the plant cells before they are ingested. Consequently one can find no recognizable fragments in the intestines of the larvae. In the floating leaves and petioles of Nuphar many sharp, hair-like branched trichosclereids occur which consist of calcium oxalate. The se- creted digestion enzymes also dissolve these sharp trichosclereids; the intestines of the larvae are not harmed by them in this way. The calci- um oxalate is transformed into calcium carbon- ate in the Malpighian organs (Schütte, 1921). Eberle (1943) already demonstrated that the larva of H. livens feeds by day and by night, as is illustrated in fig. 22b. The larval consumption and consequently the length of the mine is somewhat larger by day than by night, probably due to differences in temperature between day and night. 82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 Regularities in larval feeding and mine patterns According to Schütte (1921) the larvae are negatively phototactic, which may help and guide their position in the leaf tissue. According to Eberle (1943) and to our own observations the way of larval feeding shows regularities, VIZ.: (a) The larva, which has recently left the egg, mostly creates a mine gallery that makes a sharp bend of approximately 180° (fig. 22a). (b) After that the larva mostly mines in the direction of the leaf margin, more or less paral- lel to the lateral leaf nerves. When eating to- wards the leaf edge, the thickness of the meso- phyll decreases while the volume of the larva in- Fig. 23. Various types of mine tracks of larvae of H. livens in leaves of Nuphar lutea. The arrows show the loca- tions of dead larvae. BROCK & VAN DER VELDE: Hydromyza livens 83 creases; this results in a more or less sharp gradual bend of the mine gallery parallel to the leaf margin (fig. 23a—e). (c) After this the feeding behaviour of the larva suddenly changes. This change is corre- lated with the first moulting period of the larva. After having cast the old skin, the larva mines more or less at right angles to the previously created part of the mine gallery in the direction of the midrib or petiole of the floating leaf (fig. 23a—e). (d) On reaching the petiole the larva mines spirally downwards where ultimately the full grown larva pupates. These regularities in larval feeding behaviour cause regularities in the shape of the mine galle- ries and consequently one can distinguish cer- tain mine types, in the leaf blades, viz.: (1) The T-shaped mine gallery (fig. 23a, b and c). This type is created by larvae which after the first moulting period crawl backwards in the older part of the mine gallery before starting to eat in the direction of the midrib or petiole. (2) The arch-like mine gallery (fig. 23d and e). In this type, the larva does not crawl back- wards after moulting. (3) The ae or irregularly shaped mine gallery (fig. 23f, g, h, 1). This type of mine gal- lery is the result of disturbances in the mining behaviour of the larva, or it is caused by the damaged state of the leaf blade. Although the eggs of H. livens are mostly deposited under young and undamaged floating leaves, these leaves might be damaged before the eggs have hatched. This damage on the leaves of Nuphar lutea might, for instance, be caused by the feed- ing by the Coot (Fulica atra L.), the gastropod Lymnaea stagnalis (L.), the beetles Donacıa crassipes F. and Pyrrhalta nymphaeae (L.), the caterpillar Nymphula nymphaeata (L.), but above all by other H. livens larvae. When the floating leaf is not damaged too much, the younger larva has the ability to avoid the older affected leaf part (fig. 20a). When the larval numbers are at their maxi- mum, there is a great chance that a larva of H. livens crosses an older mine gallery. In that case the larva sometimes crosses the mine under a right angle but mostly the larva makes use of the older gallery for a while (fig. 23g). This can result in creeping into wrong directions of the larva, and, consequently, in an irregular shape of the mine gallery. It also happens that a larva mines in the wrong direction after the first moulting (fig. 23h, 1) also resulting in an irregu- lar shape of the mine gallery in the leaf blade. In order to get an idea of the occurrence of the different types of mine galleries, 34 floating leaves were collected in the Oude Waal on 11 September 1979. In total, 207 mine galleries could be recognized in these leaves. Of the min- es 60.5% could be classified within the T- shaped type, 30.5% in the arch-like type, and 9.0% in the cross-like or irregular shaped type. Feeding behaviour and temperature In the laboratory the larval consumption in floating leaf blades was followed in different cli- mate chambers. In fig. 22b the larva has reached the petiole on the 4th day. This also happened with a larva in the laboratory in the climate chamber of 24 °C. In the climate chamber of 20 °C a larva reached the petiole on the 6th day, while at constant temperatures of 16 °C and 12 °C this happened on the 11th an 18th day after the moment of hatching from the egg, respectively. The daily consumption of the three larvae in the climate chambers of 20°C, 16°C and 12°C was fol- lowed in detail (fig. 20 and table 6). Although the consuming activity of only a few larvae was followed under controlled cir- Table 6. Daily length increase of mine tracks in mm of three H. livens larvae in leaf blades of Nuphar at temperatures of 20, 16 and 12 °C, respectively. Number of days 20 °C 16 °C PRG mm mm mm 1 12 8 7 2 16 10 8 3 23 11 7 4 8 9 6 5 32 13 7 6 34 7 6 7 — 2 7 8 — 5 4 9 = 17 5 10 — 15 6 11 — 7 4 12 — — 1 13 — = 1 14 — = 5 15 = = 8 16 = = 11 (7 — = 10 18 — — 5 Total length of mine track 125 104 108 mm Mean length per day 20.8 95 6.0 mm 84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 cumstances one can draw some general conclu- sions from the data thus obtained. The con- suming activity is temperature dependent and apparently the development rate is faster under warmer conditions during the summer, than in spring and autumn when colder air and water temperatures occur. The daily consumption of the larvae is not constant at a constant temperature. Generally the larvae consume more when they become older, only during the moulting period the lar- val consumption is very low (fig. 20, table 6). Also the time needed for moulting seems to be temperature dependent and consequently takes a longer time under colder circumstances. Differences in consumption and feeding behaviour between the generations According to Schütte (1921) and Hendel (1928) the length of the mine galleries in the petioles of the summer and winter pupae is ca. 10 cm and 20—25 cm respectively. In order to get an impression of the larval consumption of plant tissue of the various generations both in the second half of July and in September, 1977, a number of floating leaves and leaf stalks of Nuphar were gathered in the Oude Waal. The lengths of 25 mine tracks in the leaf blades and another 25 in the leaf stalks were measured (ta- ble 7). The measurements of July, 1977, correspond with the second generation of H. livens. This generation, as well as the first one, completes its life cycle within one growing season of Nuphar. The lengths of the mine galleries measured in September correspond with the third genera- tion, which generation hibernates as pupa. Three stages of the mine track have been dis- tinguished in the floating leaf and petiole, viz.: (1) mine stage 1; this stage corresponds with Table 7. Lengths of 25 mine tracks of H. livens larvae in leaf blades and petioles of Nuphar, in July and September, 1977. For explanation of the mine stages, see the text. July, 1977 n Mine stage 1 Mine stage 2 Total length Mine stage 3 inmm inmm in leaf blade in petiole (mm) (mm) 1 37 27 64 37 2 34 84 118 36 3 25 76 101 29 4 33 33 66 48 5 34 35 69 38 6 24 36 60 32 7 52 51 103 29 8 38 59 97 39 9 31 36 67 38 10 30 116 146 29 11 39 104 143 54 12 53 65 118 40 13 38 48 86 30 14 41 55 96 84 15 40 63 103 58 16 54 79 133 41 17 41 50 91 40 18 48 35 83 42 19 57 68 125 71 20 60 207 267 49 21 49 61 110 41 22 62 75 137 44 23 40 55 95 43 24 41 56 97 35 25 38 39 Zi 39 mean length 41.6 64.5 106.1 42.6 SD + 10.3 + 36.9 + 41.8 as 12,6) (Continued on page 85) BROCK & VAN DER VELDE: Hydromyza livens 85 the consumed length of the mine gallery before the first larval moulting period in the leaf blade; (2) mine stage 2; this stage corresponds with the consumed length of the mine gallery in the leaf blade after the moulting of the larva; (3) mine stage 3; the length of the mine gal- lery in the petiole. The results of the measurements are summa- rized in table 7. The differences in the mean length of the mine galleries in the leaf blade be- tween the second and the third larval generation of H. livens are not large. The differences in mean length of the mine galleries in the petioles, however, are very conspicuous between these generations with mean lengths of 42.6 + 12.9 and 95.1 + 28.3 mm, respectively. Therefore, the total length of the mine and consequently the consumption of plant tissue of the genera- tion that completes its life cycle within the growing season of Nuphar is smaller than that of the one which hibernates as pupa during the next winter. Apparently the generation that hi- (Continuation from page 84) September, 1977 bernates needs more energy to survive this cold period. Also the feeding behaviour of the first and second generation differs from the third one, in the Oude Waal. Schütte (1921) and Eberle (1943) already recorded that the fullgrown lar- vae of the first generation construct a provision for the emergence of the adults in the form of a window. In constructing this window the full- grown larva eats towards the exterior of the pet- iole until it reaches the epidermis. Here it re- moves all surrounding tissue from a more or less circular area with a diameter of 2—3 mm, which is destined to be the window and is composed of the epidermis only. The window is only large enough to allow the passage of the emergent adult. In the Oude Waal the fullgrown larvae of both the first and second generation construct such a provision. According to Eberle (1943) and according to our own observations the fullgrown larvae of the last generation (in our case the third one) do n Mine stage 1 Mine stage 2 Total length Mine stage 3 in mm in mm in leaf blade in peuole (mm) (mm) 1 50 64 114 90 2 59 78 137 88 3 41 90 131 75 4 42 55 97 80 5 41 62 103 73 6 55 75 130 93 7 37 42 79 101 8 35 120 155 80 9 48 61 109 80 10 42 50 92 86 11 51 66 117 102 12 40 155 195 94 13 45 52 97 76 14 572 55 il 90 15 50 VI 127 78 16 Si 60 111 72 17 47 71 118 116 18 59 82 141 75 19 35 121 156 124 20 60 75 135 74 21 34 41 75 80 22 46 35 81 88 23 39 74 113 197 24 35 61 96 150 25 47 73 120 116 mean length 45.8 71.8 117.6 95.1 SD + 8.1 + 26.9 ae 27.0 + 28.3 86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 not construct windows for emergence. When they have pupated they stay as long as possible in the petioles, in the broadened end part of the mine gallery. Mortality Predators of the larvae of H. livens were nev- er observed. The floating leaves and petioles of Nuphar not only provide the larvae with food and oxygen, but also with protection against possible predators. As already discussed a damaged state of the floating leaf can cause irregularities in the min- ing behaviour of the larvae, resulting in their death (fig. 23g). Also when the larvae mine in wrong directions they die before pupation (fig. 23g, h). When a particular floating leaf is affected by various larvae of A. livens, the leaf becomes too much damaged causing a faster rate of decomposition. The larvae, which have not reached the petiole in time drown, die be- cause of lack of fresh plant tissue, or become in- fected by fungi. When too many larvae reach a particular petiole and consume too much tissue the leaf stalk easily breaks off or decomposes too fast. Then only the larvae which arrived first in the petioles, and which have eaten themselves a longer way downwards survive. An important factor of mortality is the intra- specific competition especially at the end of the season when the number of larvae is maximal and more larvae mine in the same floating leaf. In the last week of August, 1977, 25 floating leaves of Nuphar which were affected by A. li- vens larvae were gathered in the Oude Waal; the mean number of mine tracks per floating leaf was four. At the end of September, 1977, 25 leaf stalks of affected leaves were gathered and in them 1—3 pupae were found with a mean number per leaf stalk of two. So in 1977 about 50% of the larvae of the last generation did pu- pate, while apparently the other 50% died be- cause of competition, especially with the older larvae. On 11 September, 1979, the mean num- ber of mine galleries per floating leaf was 6 (maximum 15) (34 affected leaves from the Oude Waal were examined). On 15 October, 1979, 20 leaf stalks were gathered and in them 32 pupae were found. The maximum number of pupae per petiole was three, while the mean number per leaf stalk was 1.6. Therefore, in 1979 less than 1/3 of the larval population of the third generation in the Oude Waal pupated. It can be concluded that the petiole offers room to 1—3 pupae only. Schutte (1921) re- corded that the mine tracks of the larvae in the triangular petioles are mostly situated in their edges, using the space optimally in this way. Pupation According to Schutte (1921) the metamor- phosis of summer larvae into summer pupae takes only a short time; transitional stages were never found. The metamorphosis of larvae into winter pupae, however, takes a period of at least 8—10 days. The process of metamorphosis starts after the 4th moulting period. Just before pupation the larvae excrete calcium carbonate via their diges- tive tracks and originating from the Malpighian organs. A clod of calcium carbonate can be found near the caudal parts of the pupae in the petioles (fig. 24). During the pupation of the winter generation the skin becomes ca. 9 times thicker by the for- | mation of chitinous layers; furthermore calcium _ and silicium are chemically bound to chitine and thus incorporated in the pupal skin (Schütte, | 1921). THE PUPA Morphological description The pupae which emerge within the growing season of Nuphar, according to Schütte (1921) and Eberle (1943) summer pupae (in the Oude Waal and Haarsteegse Wiel the first and second generation), measure 7.0—8.3 mm in length nn li i| | III [lil m) | | I il N | | Fig. 24. Positions of a summer and a winter pupa in the petioles of Nuphar leaves (Eberle, 1943). BROCK & VAN DER VELDE: Hydromyza livens 87 with a width of 2.6—3.3 mm (n = 10). Their pupal skin is rather soft, yellow-brown in col- our and more or less translucent so that the de- veloping adult is to be seen. The cylindrical body mostly consists of 12 clearly visible seg- ments, while a large amount of minor chitinous projections on the pupal skin gives it a faintly, transversally striated appearance. The kidney- shaped frontal stigmata and the acute projec- tions of the terminal stigmata are brown-black in colour and thus can be easily recognized. The pupae of the third generation, according to Schütte (1921) and Eberle (1943) winter pu- pae, which hibernate during the winter, measure 6.2—7.2 mm in length at a width of 3.4—4.1 mm, so they are shorter but more robust than those of the first and second generations fig. 24). The pupal skin is stout, yellow-brown to brown-black in colour and not translucent. The thick-set body, on which the 12 segments are not all clearly visible, has on its dorsal side a more or less visible yellow-brown line running from the kidney-shaped anterior stigmata to the large, dark-brown, acute projections of the ter- minal stigmata. The skin is also faintly transver- sally striated (fig. 25). In the summer pupae the thoracic stigmata are in connection with the anterior stigmata of the pupal skin. This is not the case in the winter pupae; here the anterior stigmata of the pupal skin are in connection with a well-developed air layer, ca. two times as large in the winter pupae as in the summer pupae (Schütte, 1921). Position of the pupae in the leaf stalks Eberle (1943) already described that the pu- pae of the first generation have an other posi- tion in the petiole than the pupae which have to hibernate (fig. 24). The pupa which emerges within the growing season of Nuphar (the first and second generation in our case) is situated in a small room in the leaf stalk with its rostral end below and its caudal end above. This rostral end faces the window, the provision for the emer- gence of the adult, while the acute projections of the terminal stigmata are stuck into the living tissue of the leaf stalk. The room in which the pupa matures is closed off from the remainder of the mine gallery by a plug of loosened plant cells. The pupa of the last generation (in our case the third one) is situated in a larger room in the petiole with its rostral end upward and its cau- dal end downward, so the other way round, when compared with the other generations. The acute projections of the caudal stigmata are also stuck into the leaf stalk tissue. The broad room in the petiole in which the pupa occurs is often closed off by a small plug of loosened cells. As already mentioned this third generation has not made a window for emergence. Emergence and hibernation According to Schutte (1921) the pupal stage of the summer generation lasts 2—3 weeks. At the time of emergence the adult, originating from a pupa of the first or second generation, crawls out of the pupal skin and out of the leaf stalk via the window by means of its ptilinum. The new adult rises towards the water surface because its specific gravity is less than that of water. The third generation hibernates as pupa and stays in the leaf stalk as long as possible until the petiole tissue has completely decomposed. Dur- ing the decomposition of the leaf stalk, the pupa together with parts of the petiole, can sink to- wards the bottom of the pool, but when the pet- iole has decayed too much the pupa rises to the water surface. The pupal skin is partly filled with gas so that the pupa remains floating. In the Oude Waal floating pupae could not be found in autumn or the first part of the winter. In spring however a few of them were found be- tween material washed ashore. Apparently the decomposition rate of the leaf stalks is not very fast; the pupae can stay a fairly long time in the petioles before they rise to the surface or wash ashore. During the hibernation period a lot of pupae die; the spring generation is only very small, while in autumn most petioles of Nuphar con- tain pupae of H. livens. The solid pupal skin of the winter pupae has probably the function of protection against ice, predators or desiccation. SUMMARY AND DISCUSSION There are several relationships between the fly H. livens and the vegetation dominated by Nuphar lutea. . Adult H. livens visit the flowers of Nuphar for several reasons, viz.: (a) collecting food in the form of nectar, fur- thermore preying in the flowers on soft skinned insects or on Notiphila brunnipes eggs; (b) protection and shelter against bad weath- er conditions, e.g. heavy rain; (c) resting site; the temperature in the flow- ers is higher than those of the surroundings dur- ing sunny weather conditions (table 3), which 88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 Fig. 25. Scanning electron microscope photographs of the skin of a summer (a) and a winter pupa (b). probably is favourable for the flies (faster diges- tion and warming up of flight muscles); (d) copulation site. By their regular occurrence in the flowers and by their activities, the flies can cause pollination. In the areas investigated flowering of Nuphar took place from mid-June to mid-August so that predominantly the second generation of H. livens could visit Nuphar flowers. Adult H. livens visit the nymphaeid floating leaves for: (a) food; the fly preys here on Chironomidae and Hydrellia while the water drops occurring on the floating leaves are used for drinking; (b) copulation site; (c) egg deposition; oviposition takes place under the floating leaves of Nuphar; (d) resting site, e.g. for sunning; the upper surface of the floating leaves can reach tempera- tures higher than those of the ambient air and water (table 3); (e) shelter against bad weather conditions under aerial leaves of Nuphar and Nymphaea or leaf margins which have been curled up and dried out. Because of their many activities on floating leaves, adults of H. livens might function as vec- tors of spores of parasitic fungi, which occur on the floating leaves such as Glomerella cingulata (Ston.) Spauld. et Schrenk. From mid-August till October decomposition of the nymphaeids exceeds production; at this time also the popu- lation of adult A. livens is at its maximum. The eggs are deposited upon the underside of the floating leaf blades. An advantage of this kind of oviposition might be protection against desiccation and against predators such as adults of the own species or parasites. The larvae and pupae of H. livens are bound to the floating leaves and petioles of Nuphar. The leaf blades and petioles function as: (a) food; the larvae consume mesophyll and petiole tissue, creating mine galleries in this way; (b) protection; the larvae and pupae are pro- tected against possible predators in these mine galleries; (c) oxygen supply; the larvae obtain oxygen from the intercellular cavities; the pupae thrust their caudal stigmata into the petiole tissue, ob- taining oxygen in this way. The consumption of living tissue by H. livens larvae initiates a decomposition process in the floating leaves of Nuphar. The tissue around the mine galleries very soon decays because of in- fection by fungi and bacteria. The comsumption and damage caused by H. livens in this way can amount to 5— 10% of a floating leaf blade (Van der Velde, 1978), when the larval numbers have reached their maximum. Parts of the leaf blades break off easily by wind and wave action on places where mine tracks of H. livens occur. Also the petioles break off easily on places where the larvae have pupated. In 1977, 15.3% of the floating leaves in the Haarsteegse Wiel and 11.7% of the leaves BROCK & VAN DER VELDE: Hydromyza livens Table 8. Comparison of the most important features of the life histories of A. livens and H. confluens. Food plant(s) Geographical distribution Number of generations Adults Food - on leaves - in flowers Pollinator Spatial occurrence Copulation site Oviposition site Hydromyza livens Hydromyza confluens Nuphar lutea (L.) Sm. Nuphar advena Ait. Nuphar pumila (Timm) DC. Nuphar X intermedia Ledeb. (Nuphar advena Ait.) Europe and Asia North America 2—3 2% Chironomidae, Hydrellia nectar, soft-skinned insects, Notiphila eggs + more abundant on floating leaves bordering the open water flowers and floating leaves underside of leaf-blades of floating leaves dead bodies of Chironomus ssp. pollen ?, liquid substances + more abundant on floating leaves bordering the open water flowers and floating leaves submerged petioles of floating leaves 89 Deposition of eggs singly singly Air coat when going under water + + Eggs Plastron +? ? Larvae Food leaf- and petiole tissue petiole tissue Type of damage mine gallery interior excavation of petiole Construction of window for + + emergence Pupation site in petiole in petiole Pupae Occurrence of summer pupae + > Occurrence of winter pupae + ? in the Oude Waal separated and floated away, a process in which A. livens plays an important role. These free floating parts of leaf blades and leaf stalks can have an important function else- where for other organisms for instance for the hydrozoan Cordylophora caspia (Pallas) (Roos, 1979). From all these data it can be concluded, that the fly H. livens is an important element in the nymphaeid ecosystem dominated by Nuphar. It appears from this study on H. livens and that of Welch (1914, 1917) on the North-Amer- ican species Hydromyza confluens that the life- histories of both species in many ways resemble each other. The most important features of both H. livens and H. confluens are summarized in table 8. Both flies have an important function in nym- phaeid vegetation, H. confluens in stands domi- nated by Nuphar advena and H. livens in simi- lar habitats dominated by European Nuphar taxa. Furthermore, it is interesting that A. li- vens has the ability to affect the natural food plant of H. confluens. It is not known whether the larvae of H. confluens can also complete their life cycle on European Nuphar taxa. It should be mentioned that Beal (1956) considers Nuphar lutea (L.) Sm., Nuphar pumila (Timm) DC. and Nuphar advena Ait. to be subspecies of one species, viz. Nuphar lutea. Berg (1950) reports a normal development of a H. confluens pupa in the silty, humous bottom of a stream with respirating plates embedded in a root of Potamogeton alpinus Balb. ACKNOWLEDGEMENTS We are much indebted to Prof. dr. C. den Hartog and to Mr. J. Krikken for critical re- marks, to Dr. P. J. van Helsdingen and Dr. W. J. Kabos for the help with the identification of the flies, and to Mr. A. W. Dicke for making the SEM photographs and to the Illustration Service of the University for preparing drawings. 90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 3, 1983 REFERENCES Beal, E. O., 1956. Taxonomic revision of the genus Nuphar Sm. of North America and Europe. — J. Elisha Mitchell Sci. Soc. 72: 317—346. Berg, C. O., 1950. Hydrellia (Ephydridae) and some other acalyptrate Diptera reared from Potamoge- ton. — Ann. Ent. Soc. America 43: 374398. Brauer, F., 1883. Die Zweiflügler des Kaiserlichen Museums zu Wien III. Systematische Studien auf Grundlage der Dipteren-Larven nebst einer Zusammenstellung von Beispielen aus der Litera- tur über dieselben und Beschreibung neuer For- men. — Denkschr. mat.-naturwiss. Cl. K. Akad. Wiss. 47: 1—100, 5 pls. Collin, J. E., 1958. A short synopsis of the British Scatophagidae (Diptera). — Trans. Soc. British Ent. 13: 37—56. Eberle, G., 1943. Die Mummelfliege (Hydromyza livens Fall.). — Natur und Volk, Frankfurt am Main 73: 293—306. Gaevskaya, N. S., 1966. The role of higher aquatic plants in the nutrition of the animals in fresh-wa- ter basins. Vol. 1: 1—232. — National Lending Library for Science and Technology. Boston Spa, Yorkshire, England. Gercke, G., 1877. Ueber die Metamorphose der Hy- dromyza livens Fall. — Verh. Ver. naturwiss. Un- -terh. Hamburg (1879) 4: 229—234. Grünberg, K., 1910. Diptera, Zweiflügler. Heft 2a. In: A. Brauer (ed.), Die Süsswasserfauna Deutsch- lands, eine Excursionsfauna: 1—312. — Gustav Fischer, Jena. Hackman, W., 1956. The Scatophagidae (Dipt.) of Eastern Fennoscandia. — Fauna Fenn. 2: 1—67. Hendel, F., 1928. Blattminenkunde Europas. I. Die Dipterenminen 2 (Bog. 5—7, pl. 3—5). Hering, E. M., 1926. Die Okologie der blattminieren- den Insectenlarven. — Zoologische Bausteine, Ausschnitte aus dem Gesamtgebiet der Zoologie. Berlin 1 (2): 1—253. Heslop-Harrison, Y., 1955. Nuphar Sm. Biological flora of the British Isles. — J. Ecol. 43: 342—364. Hinton, H. E., 1960. How some insects, especially the egg stages avoid drowning when it rains. — Proc. S. Lond. ent. nat. Hist. Soc. 1960: 138—154. , 1967. The respiratory systems of the egg-shell of the common housefly. — J. Insect Physiol. 13: 647—651. , 1971. Polyphyletic evolution of respiratory sys- tems of egg shells, with a discussion of structure and density-independent and density dependent selective pressures. In: V. H. Heywood (ed), Scanning electron microscopy. Systematic and evolutionary applications. The Systematics Asso- ciation Special Volume 4: 1—331 (pp. 17—36). — Academic Press. De Meijere, J. H. C., 1895. Ueber zusammengesetzte Stigmen bei Dipterenlarven, nebst einem Beitrag zur Metamorphose von Hydromyza livens. — Tijdschr. Ent. 38: 65— 100. ——, 1940. Hydromyza livens Fall. en Notiphila brunnipes Rob.-Desv., twee Dipteren, levenswijze verband houdt met Nymphaea alba L. — Ent. Ber. 10: 220—222. Meusel, H., E. Jäger & E. Weinert, 1965. Verglei- chende Chorologie der zentraleuropäischen Flora. Vol. 1: Text: 1—583. Vol. 2: Karten: 1—258. — Gustav Fischer Verlag. Jena. Monchadskii, A. S., 1940 (according to Gaevskaya, 1966). (Diptera excluding Tendipedidae and He- leidae). In: V. I. Zhadin (ed.). Freshwater life in the USSR 1: 233—263. — Moscow Acad. USSR. Roos, P. J., 1979. Two-stage life cycle of a Cordylo- phora population in the Netherlands. — Hydro- biologia 62: 231—239. Sack, P., 1937. Cordyluridae. 62a. In: E. Lindner (ed.). Die Fliegen der palearktischen Region 7 (1): 1—103. — Stuttgart. Séguy, E., 1934. Diptères (Brachycères) (Muscidae, Acalypterae et Scatophagidae). — Faune de France 28: 1—832 + pls. 1—27. Paris. Schütte, L., 1921. Die Metamorphose von Hydromy- za livens. Inaug. Diss., Greifswald: 1—48. — E. Hartmann, Greifswald. Van der Velde, G., 1978. Structure and function of a nymphaeid-dominated system. — Proc. EWRS 5th Symp. on Aquatic Weeds (Amsterdam) 1978: 127—133. ——, 1980. Studies in nymphaeid-dominated systems with special emphasis on those dominated by Nymphoides peltata (Gmel.) O. Kuntze (Menyan- thaceae). Thesis Nijmegen, 163 pp. —— & Th. C. M. Brock, 1980. The life history and habits of Notiphila brunnipes Robineau-Desvoidy (Diptera, Ephydridae), an autecological study on a fly associated with nymphaeid vegetations. — Tijdschr. Ent. 123: 105—127. , Th. C. M. Brock, M. Heine & P. M. P. M. Pee- ters, 1978. Flowers of Dutch Nymphaeaceae as a habitat for insects. — Acta Bot. Neerl. 27: 429 — 430. Vockeroth, J. R., 1978. Scatomyzidae. In: J. Illies (ed.), Limnofauna Europaea. Eine Zusammenstel- lung aller die europäischen Binnengewässer be- wohnenden mehrzelligen Tierarten mit Angaben über ihre Verbreitung und Okologie. (2nd revised ed.): 1—532 (p. 489). — Gustav Fischer Verlag, Stuttgart, New York. Swets and Zeitlinger, Amsterdam. Welch, P. S., 1914. Observations on the life history and habits of Hydromyza confluens Loew (Di- ptera). — Ann. Ent. Soc. America 7: 135—147. —, 1917. Further studies on Hydromyza confluens Loew, (Diptera). — Ann. Ent. Soc. America 10: 35—45. Wesenberg-Lund, C., 1943. Biologie der Süsswasser- insecten: 1—682. — Gyldendalske Boghandel Nordish Forlag, Kopenhagen und Verlag J. Springer, Berlin, Wien. wier | En LI Ni) BRA ED. Cet ar Vel db a ” Mn A Le Ws NE M } i \ iy gm ) DEEL 126 AFLEVERING 4—5 1983 Pipes rr RAPT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD M. Duijm, L. Oupman and B. G. VELDSTRA. — Copulation in Ephippiger (Orthoptera, Tettigonioidea), pp. 91—96, figs. 1—4. M. Duijm and L. Oupman. — Interspecific mating in Ephippiger (Orthoptera, Tet- tigonioidea), pp. 97—108, figs. 1—6. Tijdschrift voor Entomologie, deel 126, afl. 4—5 Gepubliceerd 20-V-1983 oe i # ARR ME as COPULATION IN EPHIPPIGER (ORTHOPTERA, TETTIGONIOIDEA) by M. DUIJM, L. OUDMAN and B. G. VELDSTRA Dept. of Zoology, State University Groningen, P.O. Box 14, Haren, The Netherlands ABSTRACT Observations have been made in a number of Ephippiger species from Southern France and the role various reproductive structures play during the copulatory process. INTRODUCTION A number of species belonging to the genus Ephippiger occur in Southern France and in the adjacent parts of Spain and Italy. In some cases these species vicariate geographically, but there are also several instances of overlapping distri- bution of two or more species. The question has to be raised whether all taxa distinguished as species (e.g. Chopard, 1951; Harz, 1969) are re- productively isolated. In this paper we will in- vestigate whether the interspecific differences in the form of cerci, epiproct, uullators and sub- genital plate could function as mechanical bar- riers possibly operating against interspecific matings. In another, the next, paper the likeli- hood of interspecific mating is dealt with. MATERIAL AND METHODS Animals of six species were collected during field trips in August and September, 1977, 1979 and 1980. They were measured and taken to the laboratory where observations were made dur- ing the second half of September and during October. In November most animals died. De- tailed observations of copulations were made with a stereomicroscope (magnifications 6.5— 16x). Some copulations were documented by photography or videorecording. The species studied and their collecting sta- tions are the following. Ephippiger ephippiger (Fieb., 1784): Niaux (Ariège) 1977, 1979; Mezel (Ht. Provence) 1979, 1980; Causse de Larzac (Aveyron) 1979; Plan d’Aups (Var) 1979, 1980. Ephippiger cunii (Bol., 1877): Mont Louis 1) Nadig (1980) considers E. bormansi a subspecies of E. terrestris, but the conventional classification is retained here for the time being. 91 (By Orient.) 219772 Orient.) 1979. Ephippiger cruciger (Fieb., 1853): Neffiès (Hérault) 1979; Leucate (Aude) 1979. Ephippiger provincialis (Yers., 1854): Plan d’Aups (Var) 1979, 1980. Ephippiger terrestris (Yers., 1854)!): Mézel and several other places in the vicinity of Digne (Ht. Provence) 1979, 1980, 1981; Fayence (Var) 1980; Col de Braus (Alp. Mar.) 1980; Tende (Alp. Mar.) 1980. Ephippiger bormansi (Br., 1882)!): Col de Tende (Alp. Mar.) 1980; Vallone del Arma (Pie- mont) 1981. Some pairs were killed during or just after copulation and preserved to study the position of the genital organs and their elements with re- spect to each other as well as the position of the spermatophore. 1979; Canigou (Pyr. THE COPULATION Description of the behaviour The general course of copulation in Ephippig- er conforms to that in other Tettigonioids, as described by Gerhard (1913, 1914). A short de- scription will be given here emphasizing the de- tails relevant to the questions raised. A normal copulation will take about half an hour. Ephippiger males stridulate during part of the day and night, while sitting on a plant or bush, the height of which generally does not ex- ceed two metres. A sexually motivated female will walk straight to a stridulating male (Busnel & Dumortier, 1954; Duijm & Van Oyen, 1948). On encountering the male the female stops. Both male and female touch each other with the antennae. The male may tremble with his body several times (Busnel et al., 1955). After some time the male will place the extremity of his ab- 92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 4 (1983) tem Fig. 1. A pair of Ephippiger ephippiger just before the cercal clamp: ce, cercus, ep, epiproct, gr, copulatory groove, ov, ovipositor, pa, paraproct, ph, phallus with: // lateral lobes, dl, dorsal lobes and vl, ventral lobes; pl, pleural region, sa, saddle-like part of sper- matophore, sp, subgenital plate, st, stylus, sx, sperma- tophylax, s6, sixth sternite, t9, ninth tergite, t10, tenth tergite. domen in front of the female’s head; the female will then touch the male’s somewhat ventrally curved abdomen with her maxillary palps. Mov- ing forward slowly, continually touching, she finally mounts the male’s back. When the fe- male has entirely mounted, the male bends his abdomen dorsally, extends the lobes of the phallus and turns his subgenital plate down- ward. The titillators are now visible between the phallus lobes. The cerci are extended and spread in a laterodorsal direction. The female is now sitting on the back of the male with her head above his pronotum and the median planes of both partners have to: coincide at this point. In the female the caudal part of the ventral abdom- inal wall is contracted, the ovipositor making an angle of 20—45° with its longitudinal axis (fig. 1). Only in this position the female’s subgenital plate is situated between the male’s cerci in such a way that the copulation may proceed. The male circles with its cerci caudally and medially. Suddenly the inner teeth of the cerci get a grip on the female’s subgenital plate and at the same time the cerci clamp the female tightly. Clamp- ing does not occur randomly, but only on two small grooves — the copulatory grooves — which are easy to distinguish by their dark col- our, owing to extra sclerotisation. By adducting the cerci the male pulls the female firmly against its body. In this position the epiproct pushes dorsocaudally against the female’s subgenital plate and provides one of the factors for deter- mining the position of the male abdomen with respect to the female’s. At the same time, push- ing the epiproct against the female subgenital plate results in the lifting of the caudal part of the subgenital plate from the female gonotrema. By this action the gonotrema will be opened up and exposed. Immediately after being clamped, the female brings the ovipositor downward very rapidly to such a position that it fits in the emar- gination of the caudal rim of the subgenital plate and contacts the styli (fig. 2). After some sec- onds the ovipositor returns to its normal posi- tion. The function of this action is not clear. Perhaps in this way a final check is made to de- termine whether the partners are in the correct position to enable the next step in the copula- tion procedure, viz., the rotation. In this posi- tion the male will extend and contract the phal- lus alternately. The female now makes a few steps forward, the male meanwhile somersault- ing until he gets hold of the female’s ovipositor with his fore and middle legs (fig. 3). In doing so, the male is rotated about an axis through the inner cercal teeth. This rotation is only possible when the cerci are tightly clamped. The position of the male is now entirely fixed with respect to the female and evidently in the only one in which the spermatophore can be deposited. When extended the phallus now reaches the ba- sis of the ovipositor. The phallus makes pulsat- Fig. 2. The position of male and female E. ephippiger immediately after the cercal clamp, the female having lowered the ovipositor to contact the male’s subgeni- tal plate and styli. See fig. 1 for explanations. Duijm, OUDMAN & VELDSTRA: Copulation in Ephippiger 93 2 Fig. 3. The relative position of male and female after the rotation in Ephippiger. See fig. 1 for explanations. ing movements during which its dorsal lobes, together with the apical parts of the uullators, will glide over the basis of the ovipositor. After a number of very intensive pulsating move- ments the apical parts of the titillators will be put into the female’s gonotrema at the basis of the ovipositor. Thereafter, the pulsating movements of the phallus will continue for a while but less intensively: meanwhile the uulla- tors in the gonotrema are moved backward and foreward. They are dorsocaudally directed and their apices reach the roof of the genital cham- ber, where the female’s genital lobes are situ- ated. Possibly these genital lobes are expanded during this phase. After some time the pulsating movements stop, the phallus being in a maxi- mally extended position. The phallus lobes part and the spermatophore is extruded (fig. 4a). The globular caudal part of the spermatophore bears dorsally a saddle-like structure. With the aid of the phallus the male pushes the spermatophore with the saddle against the ventral valves of the ovipositor (fig. 4b). The saddle is sticky so that the spermatophore adheres to the ovipositor for some time. The phallus is then contracted, especially its dorsal lobes (fig. 4c). In this phase the stalk of the spermatophore is inserted into the gono- trema. At the same time the female will make sculling movements with the ovipositor by moving the right and left valves with respect to each other rostrally and caudally, respectively. Apparently the female is playing an active role while receiving the spermatophore. At the end of this phase the extremity of the stalk of the spermatophore is firmly held by the genital lobes in the genital chamber. After some seconds follows the deposition of the spermatophylax, a rather voluminous gelati- nous mass (fig. 4d). Meanwhile the male with- draws the phallus from the spermatophore. The titillators have been withdrawn from the gono- trema and are visible again. The female will now make some strides, the cerci will loosen and the partners part. The male’s phallus is contracted and the subgenital plate closed. The female will soon begin to eat parts of the spermatophylax and ultimately, after an hour or so, also the spermatophore. In the meantime sperm from the spermatophore has had ample opportunity to enter the female’s spermatheca. The openings of the sperm canals in the spermatophore-stalk are situated just opposite the entrance to the spermatheca. Experiments The role of the inner cercal teeth was verified by carefully cutting them away. This was done in males of E. ephippiger and E. terrestris. Five copulation attempts by these experimental males were observed, all showing the same course. After mounting, the female performed downward movements with the ovipositor without however reaching the male’s subgenital plate. Although the male tried to grasp the fe- male with his cerci, he never succeeded. It is concluded that the inner cercal teeth are essen- tial for providing a hold on the female. Further- more, these experiments showed that the fully deflected position of the ovipositor will only be realised as a reaction to the accomplishment of the cercal clamp. With regard to the role of the titillators the following morphological information is rele- vant. A male has two symmetrical titillators. Each titillator consists of a basal part, on which two muscles insert, and an apical part, which may protrude from the phallus lobes. The latter enters the female gonotrema during copulation. One muscle inserts on the lateral part of the titillator basis and originates from the apodeme of the 10th tergite. This muscle is probably ho- mologous with the ventral retractor muscle (Snodgrass, 1940). The other muscle inserts on the medial part of the titillator basis and has its origin on the subgenital plate. This muscle is the homologue of Snodgrass’s dorsal retractor mus- 94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 4 (1983) 9 Fig. 4. Production of the spermatophore in Ephippiger: (a) the globular part of the spermatophore becomes vis- ible; (b) the saddle-like part of the spermatophore is being pressed against the ovipositor; (c) the dorsal lobes of the phallus are being contracted; (d) the spermatophylax appears. See fig. 1 for explanations. cle. The following movements have been ob- served: (a) dorsal deflection, (b) lateral deflec- tion. In (a) and (b) the apices of both titillators stay close together pointing in the same direc- tion, whereas when the phallic lobes are maxi- mally extended, the apices may diverge. To ob- tain some insight into the way the titillators function during copulation the following exper- iments were performed. In eight males of E. ephippiger the apical parts of the titillators were cut off. Both titillators where shortened to the same extent, but the length of the cut parts dif- fered. In the extreme case the entire denticu- lated part was removed. The operation did not alter sexual motivation or copulatory behav- iour. One copulation was observed in each male. In no case were the remnants of the titilla- tors inserted into the female’s gonotrema, inde- pendent of the length of the remaining part. Probably the cut ends of the titillators cannot readily enter the gonotrema because they lack the rounded apex which is characteristic of the intact titillator. Notwithstanding the lack of in- sertion, a spermatophore was deposited in all cases. In half of these cases the stalk was brought into the normal position within the gonotremal space. In the remaining cases this did not occur and the female then lost the sper- matophore immediately after the copulation. From these observations it follows that the in- sertion of the titillators is not a prerequisite for completion of the copulation. Evidently the ac- tion of the male’s phallic lobes together with the action of the female’s genital lobes may result in normal reception of the spermatophore stalk. The number of failures was nevertheless high; Dumm, OupMan & VELDSTRA: Copulation in Ephippiger 95 this points to the role of the titillators in provid- ing additional reliability. In our opinion the titillators do not play a role in opening up the female gonotrema nor in delivering a stimulus to the female, which is essential for receiving the spermatophore. Insertion of the titillators into the gonotrema probably orientates the position of the gonotrema just opposite the male’s gono- phore. In the end phase of copulation, the titil- lators are situated against the rostral wall of the gonotrema in such a way that the stalk of the spermatophore may glide along the smooth ventral sides of the titillators and in this way may be led to the gonotrema, as by a shoe-horn. Because the curved denticles on the apical parts of the titillators are situated on their dorsolat- eral sides, they will therefore grip the rostral wall of the gonotrema in the end phase of the copulation. Without this gripping, however, successful insertion of the spermatophore still remains possible. In two other males of E. ephippiger a part of only one titillator was cut off, the other being left intact. Insertion ap- peared to be normal in these males during copu- lation. In one copulation the stalk of the sper- matophore was inserted quite normally, in the other case the spermatophore was lost by the fe- male. In the latter case, orientation of the male gonopore with respect to the female’s gono- trema was in our view not precise enough to en- sure reception of the stalk by the female’s geni- tal lobes. Discussion The copulatory behaviour in Ephippiger con- sists of two closely interlocked complementary series of movements by male and female. Both female and male copulatory apparatuses func- tion as mechanical systems of considerable com- plexity. For their successful interaction not only has highly standardised behaviour to be per- tormed, but also the form and dimensions of the various parts in male and female must conform to a high extent. In the course of the copulation a consecutive series of mechanical conditions has to be sausfied after the female has mounted: (1) do the median planes of both partners co- incide? (2) is the female’s subgenital plate situated between the male’s cerci? (3) can the inner cercal teeth get a hold in the copulatory grooves of the female’s subgenital plate? (4) does the ventrally deflected ovipositor fit the emarginated rim of the male’s subgenital plate so that their gonopores are just in front of each other? (5) do the phallic lobes with the titillators reach the female’s gonotrema? Only when all conditions are fulfilled is the cercal clamp carried out correctly and rotation ensues. After this, spermatophore and sperma- tophylax will be produced and the spermato- phore stalk will be inserted into the female’s gonotrema when: (6) the titillators have been introduced in the gonotrema. Form, position and size of the various parts of the genital apparatus have to correspond; it is analogous to a lock and key system. However, this system does not appear to be very rigorous- ly determined: interspecific copulations may oc- cur occasionally under laboratory conditions. Moreover, not every morphological detail ap- pears to beiof decisivel importance as is indicated by the experiments with cut titillators. In the following paper, the differences be- tween species with regard to their genital appa- ratus and copulatory behaviour will be consid- ered together with the possibility of interspecif- ic matings. SUMMARY The course of copulation in the genus Ephip- piger has been described emphasising the suc- cession of mechanical conditions that must be satisfied for the copulation to be completed. The male cerci play a central role; they have to grasp into the copulatory grooves on the fe- male’s subgenital plate with their inner teeth. Only after having successfully clamped the fe- male does the male rotate about the axis through the inner cercal teeth and continues copulation. The titillators appear to be used as shoe-horns for guiding the spermatophore stalk when it is being introduced into the female gonotrema. ACKNOWLEDGEMENTS Our thanks are due to the students who as- sisted in collecting the insects and in all other connected activities: Geert Damsma, Gertjaap van Klinken, Willy Koolstra, Bert Lotz, An- neke Oerlemans, Gerben Poortenga, Gerard van der Veen and especially Wybren Landman. We are grateful to Dr. G. Thomas for critically reading the manuscript and for correcting the English. Furthermore we thank Mrs. J. Poel- stra-Hiddinga for typing the manuscript and to Mr. D. Visser for making the drawings. 96 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 4 (1983) LITERATURE Busnel, R. G., & B. Dumortier, 1954. Observations sur le comportement acoustico-sexuel de la fe- melle d’Ephippiger bitterensis. — C.R. Séanc. Soc. Biol. 148: 1589— 1592. Busnel, R. G., P. Pasquinelly & B. Dumortier, 1955. La trémulation du corps et la transmission aux supports des vibrations en résultants comme moyen d’information à courte portée des Ephip- pigères male et femelle. — Bull. Soc. Zool. France 80:18—22. Chopard, L., 1951. Orthopteroides. Faune de France 56: 1—359. — Lechevalier, Paris. Duijm, M., & T. van Oijen, 1948. Het sjirpen van de sabelsprinkhaan. — Lev. Nat. 51: 81—87. Gerhardt, U., 1913. Copulation und Spermatophoren von Grylliden und Locustiden. — Zool. Jahrb. 35: 415—532. , 1914. Idem, II. — Zool. Jahrb., Syst. 37: 1—64. Harz, K., 1969. Die Orthopteren Europas: 1—749. — Junk, The Hague. Nadig, A., 1980. Ephippiger terrestris (Yersin) und E. bormansi (Brunner von Wattenwyl) (Orthoptera): Unterarten einer polytypischen Art. Beschreibung einer dritten Unterart: E. terrestris caprai ssp. n. aus den Ligurischen Alpen. — Rev. Suisse Zool. 87:473—512. Snodgrass, A. E., 1940. Male genitalia of Ortho- pteroid insects. — Smiths. misc. Coll. 96: 1—102. INTERSPECIFIC MATING IN EPHIPPIGER (ORTHOPTERA, TETTIGONIOIDEA) by M. DUIJM and L. OUDMAN Dept. of Zoology, State University of Groningen, P.O. Box 14, Haren, The Netherlands ABSTRACT À number of (sub)species of Ephippiger were compared with respect to the possibility of interspecific mating, differences in the morphology of genital parts and geographical distri- bution. A tentative conclusion on the taxonomic status of the (sub)species studied is pre- sented. INTRODUCTION In Southern France a number of species of the genus Ephippiger occur, which are mainly en- demic. Their morphological characters are rath- er variable and show extensive overlapping. Therefore, the question may be raised whether this situation is at least partly due to interspecif- ic interbreeding. Nadig (1980) postulates an ex- tensive hybridisation zone in the French Alps between Ephippiger terrestris and E. bormansi, which he considers subspecies. To look further into this matter we have, in the preceding paper (Duijm, Oudman & Veldstra, 1983), analysed the copulatory behaviour in Ephippiger in gen- eral, emphasising those morphological and posi- tional details that could be decisive in determin- ing whether successful copulation between two partners will be possible. In this paper data are given with regard to the possibility of the oc- currence of interspecific matings under labora- tory conditions. Moreover, relations between these observations and the morphological dif- ferences in the copulatory apparatus are given. MATERIAL AND METHODS Specimens of eight (sub)species of Ephippiger were collected during field trips in August and September, 1977, 1979, 1980 and 1981. They were measured and taken to the laboratory where observations were made during the sec- ond half of September and October. Animals were kept individually in small cages and fed a diversity of wild plants (e.g. Stellaria media 1) Nadig (1980) regards E. bormansi as a subspecies of E. terrestris; the conventional classification is re- tained here for the time being. 97 (L.), Calystegia sepium (L.), Aegopodium poda- graria (L.), Taraxacum spec.) and occasionally a meal-worm. The cages were regularly sprayed with water and females could deposit their eggs in small boxes with moist turf. For observation a male and a female were placed together on a branch. During the copula- tions details were observed with a stereomicros- cope (magnification 6.5—16X). Some copula- tions were documented by photography or by videorecording. The species studied and their collection stations are: (France, unless stated otherwise) Ephippiger ephippiger vitium Serv., 1931: Causse de Larzac (Aveyron), 1979; Niaux (Ariège) 1977, 1979; Mézel (Ht. Provence) 1979, 1980; Plan d’Aups (Var) 1979, 1980; Ephippiger ephippiger vicheti Harz, 1966: Miglieglia (Ticino, Switzerl.) 1980, 1981; Nag- gio (Lago di Como, Italy) 1981; Ephippiger cunu Bol., 1877: Mont Louis (Pyr. Orient.) 1977, 1979; Canigou (Pyr. Orient.) 1979; Ephippiger cruciger (Fieber, 1853): Neffiès (Hérault) 1979; Leucate (Aude) 1979; Ephippiger provincialis (Yers., 1854): Plan d’Aups (Var) 1979, 1980, 1981; Ephippiger terrestris (Yers., 1854)!): Mézel and several other places in the vicinity of Digne (Ht. Provence) 1979, 1980; Collobrières (Var) 1980; Fayence (Var), 1980; Col de Braus (Alp. Mar.) 1980; Gréolières (Alp. Mar.) 1981; Col de Brouis (Alp. Mar.) 1981; Col de Castillon (Alp. Mar.) 1981; Tende (Alp. Mar.) 1980; Cle. Scra- vaion and Mt. Carmo (Ligur. Alps, Italy 1981; Ephippiger bormansi (Br. v. W., 1882)1): Col de Tende (Alp. Mar.) 1980, 1981; Vallone del Arma (Piemont, Italy) 1981; Naggio (Lago di 98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) Como, Italy) 1981; Miglieglia Ticino, Switzerl.) 1981. THE RESULTS OF ATTEMPTS TO INTERBREED IN THE LABORATORY We shall confine our attention to successful copulations. In this paper we speak of a success- ful copulation when the copulatory process takes the normal course and the spermatophore stalk is correctly inserted into the female’s gonotrema. No observations were made on sub- sequent fertilisation nor on the development of the eggs. The crosses attempted are dealt with below. (1) Ephippiger ephippiger Not all pairs of E. ephippiger formed by bringing together males and females of different populations copulated successfully. In all three cases observed of crossing a female of the sub- species E. e. vicheti with a male of the subspe- cies E. e. vitium (from different sources) the male’s attempt to clasp the female failed; appar- ently the male’s abdomen was too broad and therefore the distance between the cerci too big for getting hold in the female’s copulatory grooves. No data are available at the moment on the reverse combination (E. e. vicheti d x E. e. vitium ©). Among the populations studied of the sub- species E. e. vitium there is one that differs con- siderably from the others. This is the population from Niaux, where the animals are rather big. Their titillators are shorter than in the other populations of E. e. vitium and as seen in lateral view, they are more curved. Moreover the api- cal part, bearing teeth, is longer. There are also differences in song. Because of this heterogenei- ty the observations with specimens of E. e. vicheti and E. e. vitium from Niaux are left out of consideration when dealing with crosses be- tween E. ephippiger and other Ephippiger spe- cies. (2) Ephippiger terrestris Ten copulations were observed between twenty individuals from different sources. In all cases copulations were performed, not- withstanding the different sources and body sizes. The samples included rather small animals (forma minor, collected 1200 m above sea level) and rather big animals (nominate form, col- lected at 300 m), the difference in body length amounting to 15—25%. Also specimens from Tende, probably belonging to the subspecies E. terrestris caprai Nadig (1980) were included. Later observations showed the possibility of successful copulations between E. terrestris ter- restris and E. terrestris caprai (from Cle. Scra- vaion and Mt. Carmo in the Ligurian Alps). (3) Ephippiger bormansi One copulation between a pair of Ephippiger bormansi (from Col de Tende) was observed. As was expected the copulation ended success- fully. (4) Crosses between E. terrestris and E. bor- mansi On three occasions an E. terrestris 9 was placed together with an E. bormansi & (from Col de Tende). In all cases copulation was suc- cessful. The same applies to the reverse combi- nation: four successful copulations were ob- served. Later on, successful copulations were observed between E. bormansi (from Vallone del Arma and from Naggio) with E. terrestris caprai (both from Tende and from Mt. Carmo) and with E. terrestris terrestris. (5) Crosses between E. terrestris and other Ephippiger species (excluding E. bormansi) Six females of E. terrestris were placed with an E. ephippiger vitium male and two females of E. terrestris with a male of E. cuni. In none of these cases did a successful copulation ensue. Generally the female mounted and brought the ovipositor downward. The male tried to clamp the female with his cerci but did not succeed in getting hold on the female’s subgenital plate. The same result was obtained when placing a fe- male E. bormansi with an E. ephippiger male. On two occasions an E. terrestris male was placed with an E. ephippiger female. In both the male clamped the female with the cerci, but on the wrong place, i.e. on the soft rostral parts of the seventh sternum and not on the sclerotised copulatory grooves. In one of these cases the male pierced the female sternal wall whereupon the female left the male. In the other case, the copulatory rotation was effected and the male extruded the phallus. During pulsating movements of the phallus the uullators glided over the open gonotrema but they were not in- serted. A spermatophore was deposited but was lost by the female immediately after the copula- tion; obviously the spermatophore stalk had not been inserted into the female’s gonotrema. This failure was probably due firstly to the male go- nopore not being placed precisely opposite the female’s gonotrema and secondly to the uulla- tors not being able to conduct the sperma- tophore stalk. Comparable results were obtained in combi- nations (two cases) consisting of a male E. bor- Duijm & Oupman: Interspecific mating in Ephippiger 99 mansi and a female E. ephippiger. (6) Crosses between E. ephippiger and E. cunu As might be expected successful copulations between E. cunii males and females were readily obtained. On three occasions a male E. cunii was con- fronted with a female E. ephippiger: in one case the cercal clamp did not succeed; in the second case the cercal clamp held only for a short time; in the third case the cercal clamp held only after five failures. In the latter rotation followed and the titillators were inserted. After that the copu- lation attempt was terminated because the fe- male fled. On two occasions a male E. ephippiger was placed with a female E. cuni. Neither male suc- ceeded in getting a hold on the female’s copula- tory grooves and copulation attempts were ter- minated. (7) Crosses between E. ephippiger and E. cruciger One observation is available on a pairing be- tween a male E. cruciger with a female E. ephip- piger. The male did not succeed in clasping the female definitively; several times the cercal teeth entered the copulatory grooves but time and again they sprang loose with a conspicuous click. The female’s abdomen appeared to be too narrow for the male. Similar results were ob- tained in two comparable cases with female E. ephippiger from Niaux. No data are available on the reverse combination owing to a lack of E. cruciger females. (8) Crossings between £. cruciger and E. cunii. We have one observation of a fully successful copulation between a male E. cunzi and a female E. cruciger. INTERSPECIFIC DIFFERENCES IN THE MORPHOLOGY OF THE COPULATORY APPARATUS In order to elucidate the factors that may pre- vent successful copulation between a variety of Ephippiger species, a study was made of mor- phological characters of those genital parts, that could be relevant to the question raised. (1) The cerci (fig. 1) Notwithstanding the rather great variation in cercal form three groups of species may be dis- tinguished with regard to the general form of the cerci and the positioning of their inner teeth, mne): (1) Ephippiger provincialis: cercus short and stout with strongly curved inner tooth and also a strongly curved terminal tooth; (2) Ephippiger terrestris and E. bormansi with cerci that are more or less cylindrical in general outline, not conical as in the next group (Nadig, 1960). There is no terminal tooth. The inner cercal teeth are situated just near the cer- cal apex; in the living animal these teeth lay well behind the hind rim of the epiproct (Nadig, 1960). In preserved specimens this relative posi- tion may be altered; (3) Ephippiger ephippiger, E. cruciger and E. cunu: the cerci are more conical than cylin- drical. The inner teeth are not situated near the cercal apex but half or three quarters way along the cercus. In the living male the inner teeth are situated adjacent to the sides of the epiproct. (u) The epiproct (fig. 1) In group 3 the epiproct is entirely sclerotised; the connection with the 10th tergite, however, is not sclerotised and therefore the epiproct is movable with regard to the 10th tergite. On the other hand in group 2 the epiproct itself is not entirely sclerotised: neither its caudal edges nor its lateral rims are hardened. The sclerotised part of the epiproct is, in these species, not movable with respect to the 10th tergite. (iii) The titillators (fig. 2) In group 2 the apical parts of the titillators bear one row of denticles, whereas in group 3 there are often more rows of denticles. The bas- al edge of the titillators is broadened in group 2, but not so in group 3. Seen from the side the ap- ical parts of the titillators are distinctly curved in group 3 and rather straight in group 2. In group 1 the basal parts are strongly upcurved with a broadened end part. (iv) The seventh sternite and subgenital plate in the female (figs. 3 and 4) In group 1 nearly the entire subgenital plate is sclerotised, but in group 2 and 3 the seventh sternite and the subgenital plate of the female are only partially sclerotised. The two groups differ clearly with regard to the form of the scle- rotised parts. Indications of such differences are present in the systematic descriptions (e.g. Harz, 1969). A more detailed description is given here. In group 3 the subgenital plate is not as exten- sively sclerotised as in group 2. The caudal rim is sclerotised and generally there are two sepa- rate shield-like sclerotised parts on either side of the midline. (These shields are lacking in E. ephippiger vicheti). Moreover, the rostrolateral parts bearing the copulatory grooves are sclero- tised. These copulatory grooves are not situated 100 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) t10 E.provincialis E.ephippiger vitium Tal E.ephippiger vicheti od | E.terrestris E. bormansi E.cruciger E.terrestris caprai E.cunii BE ei cm Fig. 1. Hind parts of Ephippiger males, dorsal aspect: ce, cercus, ep, epiproct, gr, copulatory groove, ov, ovipo- sitor, pa, paraproct, ph, phallus with: //, lateral lobes, dl, dorsal lobes and v/, ventral lobes; pl, pleural region, sa, saddle-like part of spermatophore, sf, spermatophore, sp, subgenital plate, st, stylus, sx, spermatophylax, 56, sixth sternite, s7, seventh sternite, t9, ninth tergite, #70, tenth tergite. in an extreme lateral position as in group 2, but significant difference in the distance between more medially. The lateral limit of a groove is the copulatory grooves in group 2 and group 3. rather gradual, but the medial rim is very steep, On the average, this distance amounts to 10 to owing to a very distinct ridge, that is strongly 11% of the body length (range 9—12%) in sclerotised. group 3 and 14 to 16% (range 13—17%) in Owing to the difference in position there isa group2. Duijm & OUDMAN: Interspecific mating in Ephippiger 101 N E.provincialis Vi E.terrestris (Fayence) E. bormansi (Col de Tende) By E.terrestris caprai (cle. scravaion) apical part o E.ephippiger vitium (Couvertoirade ) N E.ephippiger vicheti E.cunii (Mont Louis) E.cruciger (Neffies ) | 1mm Fig. 2. Right uullators, dorsal view (left), right view (right). INTERSPECIFIC DIFFERENCES IN BEHAVIOUR (1) Trembling The trembling behaviour in Ephippiger is de- scribed by Busnel et al. (1955). The animal makes rapid up and down movements with the body, by flexing and extending the legs alterna- tely while keeping the tarsi to the substrate. The male often trembles after having contact with a female; also the female may tremble in the neighbourhood of a male, but this is seen less frequently. Males may tremble also towards other males. Trembling has been observed regu- 102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) E.bormansi E.terrestris Fig. 3. Hind parts of Ephippiger females, group 2, left view (left), ventral view (right); see fig. 1 for explana- tions. larly in E. ephippiger and also in Ephippiger cunii, E. cruciger and E. provincialis. It has nev- er been observed during copulation preliminar- ies in E. bormansi or in E. terrestris. (u) The course of the copulation A general description of the course of copula- tion in Ephippiger is given in the previous paper (Duijm, Oudman & Veldstra, 1983). In this pa- per we restrict ourselves to pointing to an ob- vious difference in the course of copulation be- tween group 2 and group 3. Before the cerci clasp the female’s subgenital plate, the partners have to position themselves very accurately with respect to each other. The female contracts the caudal part of the sternal wall and directs the ovipositor downward to- gether with the subgenital plate. In this position the male epiproct is directed caudally against the female’s subgenital plate. The female’s ven- tral wall (including the medial parts of the 6th and 7th sternum and the rostral part of the sub- genital plate) is pushed tightly against the male’s tergum including the dorsal side of the epiproct. In group 3, the caudal rim of the epiproct is held in place by the rostral part of the subgenital plate, but in group 2 by the caudal part of the subgenital plate, which is that part behind the two shield-like sclerotised parts. In both cases the distance between the copulatory grooves is determined by the connecting sclerotised parts, which are formed mainly by the 7th sternum in group 3 but by the subgenital plate in group 2 (fig. 5). THE GEOGRAPHICAL DISTRIBUTION OF THE EPHIPPIGER SPECIES To judge the likelihood of interspecific mat- ings occurring in the field and to consider the taxonomic status of the taxa dealt with, data on their geographical distributions are essential. Not all data from the literature are usable on ac- count of the confusion often met with regarding the exact distinction between the species. Our interpretation of the pattern of distribution is given in fig. 6 for the (sub)species studied. For the distribution of E. terrestris and E. bormansı we have mainly followed Nadig (1980) with the addition of a few of our own data. For the dis- tribution of E. ephippiger we used the data from Chopard (1951), Harz (1969), Dreux (1962) and Voisin (1979), with some additions and modifi- cations based on our own observations. The dis- tribution of E. cruciger and E. cunüù is mainly based on Chopard (1951) and Busnel (1963), with some additions, as is the distribution of E. provincialis. Duijm & OUDMAN: Interspecific mating in Ephippiger 103 E.cunii Fig. 4. Hind parts of Ephippiger females, group 3, left view (left), ventral view (right); see fig. 1 for explana- tions. The first question that comes up now is least, are likely to meet each other in such a situ- whether there are species with overlapping ation that interspecific matings could occur. For areas and if so, whether members of two or both of these field investigations are necessary. more species will indeed meet each other or, at From the distributional data available at the 104 2 d - / E. ephippiger t10 1cm TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) ©) E.terrestris 2 E. terrestris Fig. 5. Comparison of the relative position of male and female Ephippiger after cercal clamping; a: E. ephippiger vitium, b: E. terrestris; al, bl: left view; a2, b2: tentative diagrammatical median section. See fig. 1 for key expla- nations, sc, shield-like sclerotised area on the subgenital plate in E. ephippiger vitium. Highly sclerotised parts in the female indicated by thick lines. moment it appears that there are only two en- tirely vicarious species!) that vicariate not only mutually, but also with respect to all other spe- cies: E. cruciger (Chopard’s record of its occur- rence in Cavalaire (Var) is discarded) and E. cunu. Nevertheless, before the existence of con- tact or overlap zones may be completely pre- cluded, further field investigations are neces- sary. E. ephippiger vitium and E. ephippiger vicheti are considered to vicariate and E. ephippiger vitium appears to vicariate with both E. cruciger and E. cunii. On the other hand overlapping areas are pre- sent between: (a) E. ephippiger vicheti and E. bormansi: 1) According to Dr. G. Kruseman (pers. comm.), Chopard’s (1951) record of E. perforatus for France is in error. Italian-Swiss lake district; (b) E. ephippiger vitium and E. provincialis: Plan d’Aups; (c) E. ephippiger vitium and E. terrestris: neighbourhood of Mézel and Digne, Gorges du Verdon; (d) E.terrestris and E. provincialis: Collo- brières ; (e) E.terrestris and E. bormansi: Col de Tende and, according to Nadig (1980), in a rath- er broad hybridisation zone in the south-eastern part of the French Alps; possibly E. terrestris caprat is also involved. These cases of overlapping distribution are dealt with below in more detail. ad (a). According to Nadig’s extensive inves- tigations (1968) E. ephippiger vicheti, in the northern part of its area (Swiss), lives both at lower and higher altitudes up to 1650 m above sea level. In the southern part (Swiss-Italian lake 105 Duijm & OUDMAN: Interspecific mating in Ephippiger (08.9IavN) suoz uolzesipiugAy SE ISUBWHOG Sljsaue} ‘3 ||| || JOuIW S11}S2118} "3 aided suysaue} 3 I° SUJSOU9]} (3 === sijelouiaoid ‘3 und" 49619N19 ‘3 NOYoIA Jebiddiyde 7 — -— wnıya Jebiddiyde "7 — — NOTNOL dali xnv1a4og ‘21nejouau -ou (0861) sSipeN Surmorjoj ‘Yan apeop soroods(qns) 4237441947 ayı jo uonnqinsip jesıydeı3093 ayı Suimoys dew jeuorsiaoïg ‘9 ‘314 106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) district), however, where E. bormansi also oc- curs, a distinct niche separation is present, E. bormansi occurring above about 1000 up to 1900 m and E. e. vicheti occurring at lower alti- tude. Nevertheless, according to Nadig (1968) both species may live in an intermediate zone (950—1100 m). In September 1981 we found these two species living together above Naggio (Lago di Como) at 750 m and at the foot of the Mt. Lema between 920 and 1100 m. The two species differ considerably in song character and in behaviour. According to our observations in this region E. e. vicheti lives much less on ex- posed shrubs, but more in sheltered places and generally rather high up in trees and bushes, whereas E. bormansi occurs more in the open and on lower plants and shrubs. The differences in song will be dealt with in a separate paper. Specimens of both species may meet each other, but there is no indication of hybridisation in the field. This agrees with the laboratory observa- tions reported above (p.o.). ad (b). In the neighbourhood of Plan d’Aups and Mazaugues (Var) E. provinciaals and E. ephippiger vitium do occur together not only in the same geographical area, but also, to a certain extent, in the same habitat. In the field we had the impression that there is some difference in microhabitat. E. provincialis lives preferably on dwarf-shrubs, not more than 50 cm above the ground, whereas E. ephippiger mainly lives in the higher parts of bushes and trees, with pref- erence on oaks, blackberry, etc. As might be ex- pected from the considerable morphological dif- ferences between these species no hybridisation occurs. ad (c). As is indicated by Dreux (1962) E. ephippiger occurs in the French Alps generally at lower altitudes up to 1500 m (maximally 2000 m). We found E. terrestris and E. ephippiger oc- curring together at several places in the Alpes- de-Haute-Provence. In the neighbourhood of Mézel and Digne E. ephippiger occurred up to 1100 m and E. terrestris from 800 m upwards. In this area E. ephippiger lived mainly on bushes and high plants, whereas E. terrestris lived chiefly on low herbs and in the meadows. On the slope leading to the Gol de la Cine (coming from Digne), E. ephippiger was found at 770 m and E. terrestris at 870 m. In this case no real contact zone could be established. Near the Gorges du Verdon both species were found occurring in close proximity at 750 m as well as at an altitude of 850 m. In this case, too, E. ephippiger showed at preference for blackberry bushes along the road, unlike E. ter- restris. No indication of possible hybridisation was encountered in the field, which is in accordance with the results of the laboratory observations. ad (d). Near Collobrières (Var) at an altitude of about 130 m, E. terrestris lived on bushes, vine and low herbs, whereas E. provincialis was found preferentially in vineyards. ad (e). In the neighbourhood of Tende E. ter- restris occurs rather commonly at altitudes be- tween 700 and 1000 m. This population belongs probably to the subspecies E. terrestris caprai (Nadig, 1980). When climbing to the Col de Tende, we found E. bormansi between 1200 and 1750 m, but at lower altitudes individuals were found with intermediate morphological charac- ters (especially with respect to the form of cer- cus and epiproct). Therefore, hybridisation is likely to occur in this zone. Discussion The six species of Ephippiger studied have been arranged in three groups according to the morphology of the cerci and other genital parts. Also described is a difference between group 2 and group 3 in the copulatory positions, that re- sults from these morphological differences. Moreover, the premating “trembling” behav- iour has not been observed in group 2. The data presented above on the occurrence of successful interspecific copulations under laboratory (no choice) conditions agree with the division into three groups: between these groups mechanical barriers to interbreeding prevail, whereas within the groups interbreed- ing is not excluded in a number of combina- tions. Tentative inferences with regard to the grade of relationship may be drawn from the above. Ephippiger provincialis (group 1) forms the only example among the species studied, of completed speciation. In this regard it is inter- esting that this species is distributed on a very limited area without apparent barriers. All the other species studied appear to be in interme- diate stages of evolutionary divergence. The members of group 2, E. terrestris and E. bor- mansi, are evidently highly related taxa. No re- productive barrier whatsoever appears to exist between them. Nadig (1980) considers E. bor- mansi to be a subspecies of E. terrestris and de- scribed three vicarious subspecies, viz., E. ter- restris terrestris, E. t. bormansi and E. t. caprai. Our data do not contradict his opinion. Nadig Duijm & OUDMAN: Interspecific mating in Ephippiger 107 (1980), furthermore, describes an extensive hy- bridisation zone in the south western part of the French Alps, which he considered to result from secondary intergradation. Nadig’s hybri- disation zone extends in the east-west direction from the French-Italian border to Grasse, that is, over a range of more than 50 km. Genetical information should be collected to verify this hypothesis. More morphometric data are being collected. Research on the allozymic variation in the various populations studied is in progress. It is hoped that the independent elaboration of “genetic distances” will elucidate the phyloge- netic history. This applies also to the position of the species of the third group: E. ephippiger which is widely spread over Europe with a number of vicarious subspecies. The other spe- cies in this group are endemic ones with re- stricted areas, one (E. cruciger) being a lowland species, the other one (E. cumii) a mountain spe- cies. With regard to the systematic status of the three species within this group, Voisin (1979: 127) stated that it is really impossible to distin- guish E. cruciger, E. cunii and E. ephippiger and he cites Delmas (in litt.) that transition zones exist where hybridisation might occur. Accord- ing to Voisin (1979: 127) many species and sub- species have been previously described, which are not more than local forms. Therefore, Voi- sin tends to lump the three species together into one very polymorphic species, E. ephippiger. Our data do not support this proposition: mat- ing barriers do appear to exist between the three species (and also between £. e. vitium and E. e. vicheti), although these are generally not absolute. Probably this situation could best be charac- terised by regarding group 3 as a superspecies: a monophyletic group of essentially allopatric species that are too different to include in a sin- gle species (Mayr, 1970). Speciation appears to have proceeded furthest in those two taxa (E. cunu and E. cruciger) that are adapted to habitats that differ rather largely from the gen- eral habitat E. ephippiger inhabits. Probably all members of the third group have the status of semispecies, because their process of speciation is not fully completed. Gene exchange is still possible among them but this is usually prevent- ed by geographic isolation (Mayr, 1970). SUMMARY In a number of (sub)species of Ephippiger from southern France, Switzerland and north- ern Italy, observations have been made on the occurrence of interspecific matings in no choice combinations. Secondly, (sub)species differ- ences in the morphology of genital parts (cerci, epiproct, titillators, female subgenital plate) and in behaviour have been described. Thirdly, the geographical distribution is dealt with, specially emphasising the available data on partially over- lapping distribution. An interpretation of the data is presented and three groups are distinguished, between which no mating is possible: group 1: with E. provincialis, forming the on- ly example among the species studied of com- pleted speciation; group 2: including the species E. terrestris with (in accordance with Nadig, 1980) three subspecies, E. t. terrestris, E.t. bormansi and E. t. caprai; no mating barriers between these subspecies appear to exist; group 3: forming something like a superspe- cies, including E. ephippiger vitium, E. ephip- piger vicheti, E. cunii and E. cruciger; some mating barriers appear to exist between these species and subspecies, but these are not gener- ally absolute. ACKNOWLEDGEMENTS We are grateful to Dr. A. Nadig, who gener- ously supplied us with information about col- lection stations. We thank the students, who as- sisted in capturing the specimens and helped in a variety of activities: Geert Damsma, Arthur Decae, Henk Hiddema, Gertjaap van Klinken, Willy Koolstra, Bert Lotz, Lieke Tromp en es- pecially Wybren Landman. We wish to thank Dr. G. Thomas for critically reading the manu- script and for correcting the English. Finally we thank Mrs. J. Poelstra-Hiddinga for typing the manuscript and Mr. D. Visser for executing the figures. LITERATURE Busnel, M.-C., 1963. Charactérisation acoustique des populations d’Ephippiger écologiquement voi- sines. — Ann. Epiphyties 14: 25—34. Busnel, R. G., P. Pasquinelly & B. Dumortier, 1955. La trémulation du corps et la transmission aux supports des vibrations en résultant comme moyen d’information à courte portée des Ephip- pigères mâle et femelle. — Bull. Soc. Zool. France 80:18—22. Chopard, L., 1951. Orthopteroides. Faune de France 56: 1—359. — Lechevalier, Paris. Dreux, Ph., 1962. Recherches écologiques et biogéo- graphiques sur les Orthoptères des Alpes fran- çaises, 765 pp. — Thèse, Paris. 108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 5 (1983) Duijm, M., L. Oudman & B. G. Veldstra, 1983. Cop- ulation in Ephippiger (Insecta, Orthoptera, Tetti- gonoidea). — Tijdschr. Ent. 126: 91—96. Harz, K., 1969. Die Orthopteren Europas: 1—749. — Junk, The Hague. Mayr, E., 1970. Populations, species and evolution: 1—453. — Harvard Univ. Press, Cambridge. Nadig, A., 1960. Beiträge zur Kenntnis der Ortho- pteren der Schweiz und angrenzender Gebiete: I, Neubeschreibung von Ephippiger bormansı Br. v. W.— Mitt. Schweiz. ent. Ges. 33: 27—46. ——, 1968. Ueber die Bedeutung des Massifs de Re- fuge am südlichen Alpenrand (dargelegt am Bei- spiel einiger Orthopterenarten). — Mitt. Schweiz. ent. Ges. 41:341—358. , 1980. Ephippiger terrestris (Yersin) und E. bor- mansi (Brunner von Wattenwyl): Unterarten einer polytypischen Art. Beschreibung einer dritten Unterart: E. terrestris caprai ssp. nov. aus den Li- gurischen Alpen. — Rev. Suisse Zool. 87: 473— 512. Voisin, J. F., 1979. Autécologie et biogeographie des Orthoptères du Massif Central: 1—354. — Thèse, Paris. L i d'a Ù 7 ue MM hi al on a is > ji fata “An i i MAE ef DEEL 126 AFLEVERING 6 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING ï Er {IIS INHOUD H. H. EveNKuIs and H. J. VLuc. — The hymenopterous parasites of leaf-feeding apple tortricids (Lepidoptera, Tortricidae) in The Netherlands, pp. 109-135, figs. 1—103. Tijdschrift voor Entomologie, deel 126, afl. 6 Gepubliceerd 20-V-1983 THE HYMENOPTEROUS PARASITES OF LEAF-FEEDING APPLE TORTRICIDS (LEPIDOPTERA, TORTRICIDAE) IN THE NETHERLANDS by H. H. EVENHUIS and H. J. VLUG Research Institute for Plant Protection (I.P.O.), Wageningen, The Netherlands ABSTRACT Fifty-six Hymenopterous parasites, reared from apple leafroller caterpillars and pupae in the Netherlands, are listed. Brief descriptions and a key are given to separate related para- site species. The great difficulties encountered in discriminating between narrowly related, insufficiently described species are discussed. These difficulties in most cases make it im- possible to obtain reliable species names. Data about host specificity are given because iden- tification of host caterpillars and pupae now became possible. Knowledge of host specificity is considered an essential and often indispensable aid in species discrimination. INTRODUCTION In the Netherlands leafroller caterpillars be- long to the most important apple pests. In par- ticular Adoxophyes orana is harmful, though caterpillars of Archips rosana, A. podana, Pan- demis heparana, P. cerasana, Spilonota ocellana and Hedya nubiferana may also cause consid- erable damage to the fruits. The role that natural enemies play in control- ling apple leafrollers is appreciated in quite dif- ferent ways. Whereas Janssen (1958) does not consider the parasites of Adoxophyes orana es- sentially important in Western Germany, Auersch (1960) states that they contribute to a considerable reduction of its host numbers in the same region. In connection with investigations on inte- grated control in apple growing, it was neces- sary to study the parasites of apple leafrollers. This means in the first place an inventory of the several parasites, together with investigations on the relationships with their hosts, and esti- mates of their importance as mortality factors of the leafrollers. A number of papers concerning the parasite fauna of apple leafrollers in Europe have been published, viz., by Lehmann (1969), Papp & Reichart (1973), Carl (1974), Charles (1974), Evenhuis (1974a, b) and Miczulski & Koslinska (1976) for the D.D.R., Hungary, Switzerland, France, the Netherlands and Poland, respective- ly. These regions have many species in com- mon, but also show differences. This may be largely caused by different identifications for | one and the same species. 109 There is an urgent need for a reliable species identification of both parasites and hosts. It is a prerequisite to study their interrelationships and to compare the results with other regions. We will discuss these matters in the next chapters. METHODS Leafroller caterpillars and pupae were col- lected from apple in various orchards in the Netherlands from 1970 onwards. Each speci- men was separately kept in a glass vial under outdoor conditions and the caterpillars were provided with an artificial diet as recommended by Ankersmit (1968). They were inspected three times a week. The emerging adult insects were either leaf- roller moths or Hymenopterous or Dipterous parasites. The Hymenoptera were glued on a triangular card board. The host caterpillar re- mains with the parasite cocoon were glued on a separate card board on the same pin. We took care to show the inner sides of the caterpillar mandibles, as these present important characters (Evenhuis & Vlug, 1972). Egg parasites are ex- cluded from this study. IDENTIFICATION OF THE HOST Though identifying adult Dutch Tortricidae does not give too much trouble, thanks to the works of Hannemann (1961), Bentinck & Diakonoff (1968) and Bradley et al. (1973), identification of the caterpillars and pupae is much more of a problem. The older living instars of the apple leafroller caterpillars may be identified with the key of De Jong & Vlug 110 TijpscHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) (1974), based on superficial characters, particu- larly colours. Swatschek (1958) in his keys of last-instar caterpillars of Central European leaf- rollers mainly used the position of the chetae on thorax and abdomen as differentiating charac- ters (chaetotaxy). However, after a parasite has emerged and the host caterpillar has died, the less sclerotized parts of the body appear too much shrivelled and discoloured to make a reli- able identification possible. We have been looking for characters that would better suit our particular needs. We con- sider the shape of the teeth of the retinaculum in the last three instar caterpillars of Tortricidae Tortricinae important in this respect (Evenhuis & Vlug, 1972). This character, however, seems to hold only for the identification of the genera. Thus genera with only one species occurring on apple, e.g., the most important Adoxophyes ora- na, do not give particular difficulties. In genera with two or more species on apple, these species may be often distinguished by colour differ- ences in the more sclerotized parts as head and pronotum. The caterpillars of the three species of Tortricidae Olethreutinae occurring on apple, viz., Spilonota ocellana, Rhopobota naevana and Hedya nubiferana, are in the older instars, even in a shrivelled condition, generally sufficiently characteristic to be recognized. Though we were not able to distinguish be- tween the leafroller species in the first and sec- ond larval instars, this is hardly necessary as, ull yet, we only reared parasites from the older instars. As to the retinacular teeth we may state now that this character has turned out to be more useful than we suggested in our 1972 publica- tion. Abrasion of the teeth seems only an excep- tion. A number of parasites emerged from leafrol- ler pupae. Identification of pupae gives less problems than that of the caterpillars. A key for the Dutch species has been published (Even- huis, De Jong & Vlug, 1973). IDENTIFICATION OF PARASITIC HYMENOPTERA The present paper is an attempt to name the Hymenoptera species that we reared from apple leafrollers. When possible, specialists were con- sulted; for the rest we tried to find out names ourselves. These names, of course, remain doubtful as long as the groups to which the spe- cies in question belong, have not yet adequately been revised. Descriptions are given in which attention is paid to the characters we consider important for species discrimination, especially in those spe- cies about the identity of which we are in doubt. We tried to find new morphological characters. When closely related species for comparison were lacking, we assumed that certain morpho- logical structures might represent good charac- ters. We hope then that during a later revision of the taxonomical groups in question the iden- tity of those species might be established. For the several species a discussion on host specificity and the period of emergence of adults _ according to our investigations are given. In ta- ble 1 data about host ranges are represented. The drawings were made by the junior au- thor. Family ICHNEUMONIDAE In Europe this may be the largest insect fami- ly. Perkins (1959) estimates the number of Brit- ish species at 2000. The number of Dutch spe- cies is estimated to be of the same order of mag- nitude (Van Achterberg, 1982). In the sequence of subfamilies, tribus and genera, as well as for the taxonomic terminolo- gy, we follow Townes (1969— 1971), with some necessary alterations in the names. Subfamily PIMPLINAE (= Ephialtinae, sensu Townes, 1969) Scambus brevicornis (Gravenhorst) (figs. 1—4) Colour characters. — Female. Head black, except for maxillar and labial palps which are yellowish; an- tenna dark brown. Thorax black, tegula yellowish white, coxae and legs light brown, middle tibia sometimes with subproximal dark ring or spot; hind tibia whitish, subproximally and distally with dark ring; hind tarsus whiush, with distally darkened seg- ments. Abdomen black. Male. Differs from female in having scape and pedi- cel whitish yellow below. Fore and middle coxae whitish yellow, rest of fore and middle legs yellow, the latter without dark tibial ring or spot; hind coxa for the most part black, hind trochanter yellow. The smaller males are on the whole somewhat lighter. Morphological characters. — Female. Clypeus membranaceous and excavated (fig. 1). Prothorax smooth, shiny; mesoscutum and scutellum finely punctate and pilose; remainder of thorax more strong- ly pilose; propodeum with two incomplete longitudi- nal carinae, sparsely pilose in front outside the cari- nae, laterally longer and more densely pilose. Fore wing with small, rhombic areolet; hind wing with nervellus broken beneath the middle (fig. 2). First ter- gite with two distally converging carinae, area be- EvenHuIs & VLUG: Parasites of apple tortricids 111 tween them smooth in front, shiny, remainder rough- ly punctate and somewhat transversely wrinkled; re- maining tergites rather strongly punctate, except for the narrow hind parts of segments 2, 3, 4, and 5, which are finely striated transversely (fig. 3); oviposi- tor two times the length of hind tibia, lanceolate and serrate ventrally near tip (fig. 4). Male. Smaller and narrower than female. Carinae of propodeum sometimes extending to hind border. Narrow hind parts of tergites smoother, almost with- out transverse wrinkles. Length female: 5—7 mm, ovipositor: 3—4 mm, length male: 3.5—5.5 mm. The genus Scambus has not been revised in a modern way and this makes species identifica- tion doubtful. We use the name brevicornis be- cause its description, together with Schmiede- knecht’s interpretation (1914), apparently is closest to it. How intricate matters are, may be inferred from Schmiedeknecht’s remark (1914): “Die Gruppe der P. [Pimpla, in which Scambus at that time was incorporated] brevicornis ist die schwierigste der ganzen Gattung. Wir stos- sen hier auf eine ganze Reihe in Habitus und in der Färbung verschiedener Formen, deren Ab- grenzung als Art oder Varietät unübersteigliche Schwierigkeiten bereitet. Ich glaube nicht ein- mal, dass sorgfältige Zuchtversuche viel Klärung beiträgen würden, denn wollten wir die aus den einzelnen Wirten gezüchteten Formen als besondere Arten auffassen, wo wäre dann das Ende”. We quote Schmiedeknecht’s opinion especial- ly because of his misuse of the species concept. A species has to be considered a reproductive community. Only this species concept may be handled in ecological investigations, like that of apple leafroller parasites. As time-consuming cross breeding experiments cannot be easily performed, taxonomists have to rely upon mor- phological and colour characters from which they usually do not know the variation limits. In contrast to Schmiedeknecht’s view, we think that rearing parasites from well-known hosts is always a great help in fixing this variability. Oehlke (1965) reports Scambus brevicornis to be reared from the sawfly Neodiprion sertifer; he further mentions the species as a primary parasite of several Microlepidoptera, Diptera and of Anthonomus pomorum (Coleoptera). If these records are indeed reliable, Scambus brevicornis might be considered pantophagous. “Scambus brevicornis” in our investigations was reared in large numbers as a solitary ecto- parasite of second generation Adoxophyes orana caterpillars in well-kept orchards. However, the first instar parasite larva seems to live endopara- sitically; the larval moult may be seen protrud- ing from the caterpillar skin. It was observed to- gether with the gregarious ectoparasitic larvae of Colpoclypeus florus, which is much more common. The parasite larvae feed on dead cat- erpillars, just like those of C. florus. We do not know if any ecological relation between the two parasite species exists. Adults: 24 July—28 September. A single specimen appeared on 24 April next spring. Itoplectis alternans (Gravenhorst) (figs. 5, 6) Colour characters. — Female and male. Black. An- tenna brownish beneath, maxillar and labial palps light brown. Legs predominantly brown, coxae black, hind tibia light and dark brown with a white subbasal ring, tarsal segments of hind leg dark brown, proxi- mally white except for the short fourth one; all claws dark. Morphological characters. — Female and male. Body for the greater part pubescent. Antenna long, extending far beyond middle of abdomen; inner mar- gin of eye distinctly excavated above the middle, face finely and widely punctate, with short, thin hairs (fig. 5). Mesoscutum rather finely punctate, covered with short, fine hairs, distance between punctures much larger than their diameters; propodeum with two incomplete longitudinal carinae, shiny, bare, ex- cept for the pubescent outer front corners. Fore wing with small, rhombic areolet; hind wing with nervellus broken considerably above the middle (fig. 6). Ter- gites roughly punctate, first smooth between the two lateral carinae in front; tergites 6 and especially 7 more finely punctate than the preceding ones. Length female: 4.5—7.5 mm, ovipositor: 1—1.5 mm, length male: 2.5—8 mm. We reared this species as a primary parasite from the pupae of several leafroller species and as a secondary parasite from cocoons of other leafroller parasites. It is known as a primary parasite of the pupae of Lepidoptera belonging to several families and also as a hyperparasite of Lepidoptera through other primary Hymeno- pterous parasites. Perkins (1957) mentions sea- sonal dimorphism in the male that we could not confirm. Adults: 15 August—16 September. Itoplectis maculator (Fabricius) (fig. 7) Morphological and colour characters. — This spe- cies resembles the former in many respects. Differ- ences are given by Perkins (1941). The hairs on head and thorax are longer and the punctation is stronger. This is especially shown on the face (fig. 7) and the IZ TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) middle of the mesoscutum; on the latter the diameter of the punctures is about equal to the distance be- tween these punctures, whereas in the preceding spe- cies this distance is much larger. There are also a num- ber of colour differences, the hind borders of the mid- dle tergites and in the female also the lateral borders being brownish. Length female: 5.5—9 mm, ovipositor: mm, length male: 5.5—7.5 mm. 1.5—2.5 Though we reared this species in somewhat smaller numbers than the preceding one, ıt does not seem to be a rare parasite of apple leafroller pupae. We reared it from Pandemis heparana, Adoxophyes orana, Archips rosana and Hedya nubiferana. We also reared it from a parasite co- coon, probably Meteorus ictericus, on apple; the leafroller remains were lost, however. Adults: 21 August—1 October. One speci- men appeared on 18 July. Apechthis spp. (figs. 8, 9) We reared only a few specimens, belonging to three species of this genus, from apple leafroller pupae. Just like in the related genus /toplectis, the inner margin of the eye is distinctly exca- vated (fig. 8). However, in contrast with this ge- nus, the head shows distinct yellow markings and the ovipositor is bent downwards (fig. 9). With the keys of Perkins (1941) and Kaspa- ryan (1973) the species could be identified as Apechthis compunctor (L.), A. quadridentatus (Thomson) (= A. resinator Thunberg), and A. rufatus (Gmelin). Our rearings yielded one male of A. compunctor from Hedya nubiferana (collected as pupa on 24 June), one female of A. quadridentatus from Pandemis heparana (adult on 3 August), and two males of A. rufatus from Archips rosana and Ptycholoma lecheana, respectively (adults on 29 and 30 June). Considering the small numbers reared, it is of no use going into further details. Subfamily TRYPHONINAE Phytodietus segmentator (Gravenhorst) (figs. 10, 11) Colour characters. — Female. Black, with small, yellow markings above, especially on scutellum, post- scutellum and propodeum. Antenna brownish, above proximally black. Legs predominantly rufous. Hind borders of tergites to a varying extent yellowish. Male. Yellow, above for the most part black, with rather extensive yellow markings. Antenna and legs as in female. Hind borders of tergites yellowish. Morphological characters. — Female and male. See figures of fore tarsal claw (fig. 10) and of wings (fig. 11). Length female: 4.5—7 mm, ovipositor: mm, length male: 5.5—7.5 mm. 15225 We reared only a few specimens, from a num- ber of localities, exclusively from larvae of Archips, both Archips podana and A. rosana. This species is recorded as a well-known para- | site of Tortrix viridana. Horstmann (1971) | reared it from Archips xylosteana, À. rosana, Eudemis porphyrana and in autumn from Ancy- lis mitterbachiana on oaks. Bogenschütz (1965) considers it an oligophagous parasite of Tortri- cidae. Horstmann regards it to be plurivoltine. Adults: 29 June—5 July. Subfamily CRYPTINAE (= Gelinae, sensu Townes, 1969) Members of this in taxonomic respect often so difficult subfamily were reared from apple leafrollers as hyperparasites in only very small numbers. They do not seem to play an impor- tant role in the natural control of apple leafrol- lers. Therefore we will only mention them with- out going into details. We reared four specimens of Gelis sp., three from cocoons of Porizontinae, two of these from Archips rosana, and one probably from | Spilonota ocellana as indirect hosts; one speci- men was reared from a pupa of Adoxophyes orana. Furthermore, one specimen of Acrolyta sp. was obtained from Spilonota ocellana | through some unidentified Braconid as a direct host, and a few specimens of Lysibia nana (Gravenhorst) from cocoons of the gregarious Apanteles ater, Pandemis heparana being the in- | direct host. The identifications were made by Drs. K. W. R. Zwart and Mr. G. van Rossem. Subfamil The Glyptini, represented here by the genera | Teleutaea, Apophua, and Glypta, are character- ized by the presence of oblique furrows on the tergites 2, 3, and 4 (cf. figs. 14 and 15). We reared five species, viz., Teleutaea striata | (Gravenhorst), two Apophua and two Glypta | species. Teleutaea striata, the only species that | was reared in considerable numbers from apple | leafrollers, is easily known by its characteristic | black and yellow markings. | Identification of Apophua and Glypta species | provide considerable difficulties. There exists a | modern revision by Aubert (1978), but identifi- cation with his key is a precarious business y BANCHINAE Evenuuis & Vrug: Parasites of apple tortricids 113 Figs. 14. Scambus brevicornis: 1, head anteriorly; 2, wings; 3, gaster; 4, tip of ovipositor. Figs. 5—6. Itoplectis alternans: 5, head anteriorly; 6, wings. Fig. 7. Itoplectis maculator, head anteriorly. Figs. 8—9. Apechthis qua- dridentatus: 8, head anteriorly; 9, tip of ovipositor. Figs. 10—11. Phytodietus segmentator: 10, claw; 11, wings. Figs. 12—14. Teleutaea striata: 12, head anteriorly; 13, thorax dorsally; 14, first four tergites. Fig. 15. Glypta varicoxa, gaster dorsally. 114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) which, in our species, was not always conclu- sive. We think that head, pronotum and prepectal carina show sufficient differential species char- acters. These will be given in the following de- scriptions. Teleutaea striata (Gravenhorst) (figs. 12—14, 89) Colour characters. — Female and male. Preponder- antly black. Clypeus, cheek beneath and mouthparts, except for the tips, yellow; antenna brown yellow. Pronotum along front margin partially and along hind margin entirely yellow. Tegula and small adjoining part of mesopleurum yellow, just like a large spot in front of middle coxa. Scutellum, postscutellum and posterior part of propodeum before the apical trans- verse carina yellow; coxae and legs yellow and yel- lowish red. Hind margins of the tergites yellowish. The extension of the yellow markings may vary con- siderably. Morphological characters. — Female and male. Clypeus incised; a small unpaired tubercle before the insertion of the antennae (fig. 12). Parapsidal furrows distinct in front part of mesoscutum (fig. 13). Apical transverse carina of propodeum distinct, the other ca- rinae absent. Abdomen, see fig. 14; second, third, and fourth tergites, especially in the male, sometimes with a longitudinal middle carina in the first third. Length female: 7.5—12 mm, ovipositor 5—7.5 mm, length male: 8—12 mm. This species, easily recognized by its colour pattern, was reared in large numbers from apple leafroller caterpillars, mainly from Adoxophyes orana, but also in small numbers from Pandemis heparana and Ptycholoma lecheana, however not from other caterpillars. There appear to be two generations which coincide with those of Adoxophyes orana, perhaps the reason that it appears especially adapted to this host. Adults: 1 May—28 August. À single speci- men emerged on 6 April. Apophua cicatricosa (Ratzeburg) (figs. 16—19, 99) Colour characters. — Female and male. Black. Cly- peus and mouth-parts yellow; antenna brownish. Up- per embossed rim and hind corner of pronotum yel- low; tegula, tip of scutellum and hind border of post- scutellum yellow; legs yellowish brown, hind femur distally and hind tibia and tarsus darker except in the female, where the hind femur is whitish and both sub- proximally and distally darker. One male had no yel- low markings on the thorax, except for the hind cor- ner of pronotum and the tegula. Morphological characters. — Female and male. Clypeus broadly rounded, without incision in the middle (fig. 16); base of mandible 1.6 as long as the shortest distance between mandible and eye; genal ca- rina not strongly sinuated (fig. 17). Pronotal collar separated from the rest of the pronotum by a deep groove, the collar covering the propleurum entirely, thus not visible from above; epomia long, appearing as a sharp edge, reaching to near upper edge of prono- tum (fig. 18). Propodeal carinae complete and strong, more or less variable. Spur of front tibia distinctly ex- ceeding the middle of the first tarsal segment (fig. 19). Length female: 10 mm, ovipositor: 5.5 mm (as long as hind tibia), length male: 10 mm. We reared seven specimens, six from Pande- mis heparana and one from P. cerasana. Adults: June, July. Apophua sp. This species, which we can not identify, re- sembles À. cicatricosa in many respects. Our single specimen, a female, differs from it by be- ing somewhat larger, 11.5 mm, ovipositor 7 mm (two times length of hind tibia), having the cari- nae on the propodeum more oblique, and only the anterior part of the median longitudinal ca- rinae and the costula being distinctly defined. It was reared from Archips podana. Glypta nigrina (Desvignes) (figs. 20—23) Colour characters. — Female. Black. Clypeus with indistinct yellow markings; antenna brown. Tegula yellow; hind tibia without distinct bands, inner side whitish. Male. As female, however clypeus yellow, its front border brownish. Morphological characters. — Female and male. Clypeus narrow, rather convex, with small incision (fig. 20); base of mandible as long as shortest distance between mandible and eye; genal carina strongly sinu- ate (fig. 21). Pronotal characters similar as in A. cica- tricosa, but less distinctly defined; collar narrower, showing lower part of propleurum, visible from above (fig. 22); pronotum rather uniformly scattered and coarsely punctate; mesopleural fovea absent. Spur of fore tibia not quite reaching middle of first tarsal seg- ment (fig. 23). Length female: 7—8 mm, ovipositor: 4—5 mm, length male: 8 mm. This species seems quite similar to Glypta varicoxa, but may easily be distinguished by the structures of clypeus, genal carina and prono- tum. Considering the length of the front tibial spur in this species, the separation between Apophua with the spur reaching to middle of first tarsal segment or beyond and Glypta, the spur not reaching to the middle, seems hardly justified. We only reared two specimens from Adoxo- EvenHuis & VLUG: Parasites of apple tortricids 115 I) (i | = “Fl FEN, T 31 ==> 32 Figs. 16-19. Apophua cicatricosa: 16, clypeus with mandibles; 17, head laterally, showing genal carina; 18, pro- | notum laterally; 19, tibial spur of fore leg. Figs. 20—23. Glypta nigrina: 20, clypeus with mandibles; 21, head laterally, showing genal carina; 22, pronotum laterally; 23, tibial spur of fore leg. Figs. 24—28. Glypta varicoxa: 24, clypeus with mandibles; 25, head laterally, showing genal carina; 26, pronotum laterally; 27, tibial spur of fore leg; 28, tip of ovipositor. Figs. 29—30. Diadegma praerogator: 29, pronotum laterally; 30, first tergite laterally. Figs. 31—32. Tranosema arenicola: 31, pronotum laterally; 32, first tergite laterally. 116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) phyes orana and two from Ptycholoma lechea- na. Glypta varicoxa Thomson (figs. 15, 24— 28, 90) Colour characters. — Female and male. Black. Cly- peus dark brownish, palps yellow, antenna brownish. Hind corner of pronotum yellow; tegula yellow; legs yellow and reddish yellow, hind tibia with subbasal and apical dark ring, hind tarsus with dark rings. Morphological characters. — Female and male. Clypeus broadly rounded, more or less depressed in the middle, without median incision (fig. 24); base of mandible 1.5 as long as shortest distance between mandible and eye; genal carina only very shallowly sinuate (fig. 25). Pronotal collar parallel to front mar- gin as in Apophua cicatricosa, but less distinct and not especially deepened between collar and front margin; propleurum visible, epomia short (fig. 26); pronotum more or less uniformly punctate, without smooth or even weakly punctate middle part; spur of front tibia not reaching to middle of the first tarsal segment (fig. 27). Tergites 2, 3, and 4 with V-shaped impres- sions as in other Glyptini (fig. 15); ovipositor with ap- ical, dorsal notch (fig. 28). Length female: 6—7 mm, ovipositor: 4.5—5 mm, length male: 5.5—7 mm. As far as apple leafrollers are concerned, this parasite seems specialized on Spilonota ocellana. Adults: 28 June—17 July. Though the characters mentioned by Aubert (1978) do not exactly fit our species, we provi- sionally maintain the name; it was mentioned earlier under the same name by Evenhuis (1974b). Perhaps G. pedata Devignes should be better. Both Charles (1974) and Couturier (1973) report to have reared a Glypta species from Spilonota ocellana on apple in France, which they name G. pedata and G. (?) fracti- gena, respectively. There can be little doubt that they refer to the same species as ours. G. fracti- gena is considered by Aubert a synonym of G. nigrina Desvignes. Thus either the species mentioned as G. nigrina in this paper, or the species stated under that name by Couturier, or perhaps both, might be misidentified. Further taxonomical investigations could elucidate these questions. Lissonota complicator Aubert This species was mentioned earlier as Lisso- nota errabunda Holmgren (Evenhuis, 1974b). According to Aubert’s key (1978) it fits better into Lissonota complicator. Colour characters. — Female. Black. Antenna dark brown, mouth-parts yellow. Fore and hind margins and hind corners of pronotum yellowish brown. Legs red yellow. Male. Conspicuously lighter coloured than female. Black. Antenna dark brown, scape and pedicel yellow below; a yellow spot between eye and outer ocellus on each side of frons; face and mouth-parts yellow, with black middle stripe extending from insertion of antennae downwards, not reaching front margin of clypeus. Front margin of pronotum lighter and with yellow spot on the outer hind corners; mesoscutum broad yellow on the front margin on each side, tegula yellow; legs red brown, fore and middle coxae and trochantera whitish yellow, hind coxa darker, hind tibia distally and first three segments of hind tarsus darkened. Hind borders of proximal tergites lighter. Morphological characters. — As Aubert (1978) gives predominantly colour characters and as we have no specimens of related species to compare, we should refrain from giving morphological characters. Length female: 4.5—6.5 mm, ovipositor: 5 mm, length male: 4—5.5 mm. We reared this parasite in rather large num- bers from caterpillars of Archips podana on apple. It might be a rather strictly specialized parasite of this leafroller. However, we saw Swiss specimens reared from a caterpillar of Pandemis cerasana by Dr. A. Schmid, Nyon, Switzerland. It is a gregarious larval endoparasite with varying ratios of females and males per host. From one single host caterpillar 3 to 8 parasites emerged with an average of 6. Adults: 11 June —15 July. Subfamily PORIZONTINAE Diadegma praerogator (Linnaeus), (Diadegma interrupta (Holmgren) (Horstmann, 1973)) (figs. 29, 30) Colour characters. — Female and male. Black. Mouth-parts yellow. Tegula yellow; fore and middle coxae often partly yellow; fore and middle trochan- tera yellow; femora light brown, hind ubia yellowish white, with broad basal and apical dark rings, all tarsi light brown, distally darker, last tarsal segment espe- cially dark. Morphological characters. — See Tranosema areni- cola. Length female D. praerogator: 4—6.5 mm, oviposi- tor: 1—1.5 mm, length male: 3.5—5.5 mm. This species is one of the most common para- sites of apple leafrollers. We reared it from cat- erpillars of several leafroller species (cf. table 1). Till yet we did not rear it from caterpillars of other families of Lepidoptera on apple, repre- sentatives of which, however, were only col- lected in relatively small numbers. Neither does EvenHuis & Vrug: Parasites of apple tortricids 117 Herting (1965) mention it from Geometridae and Noctuidae on apple. Thus we consider Diadegma praerogator as a parasite, specialized on Tortricidae. Townes (1969) referred Diadegma interrupta to the genus Tranosema Förster. Horstmann in his type revision of the European Diadegma species, placed D. interrupta in the subgenus Nythobia Förster, which contains the extremely uniform majority of the Diadegma species. The latter of course seriously interferes with species identification. Often males cannot be idenufied at all, because important characters, as eventual notches in the hind borders of the posterior sternites and the length and shape of the ovipo- sitor, refer to the female sex only. Adults: 13 May—7 October. We reared some more Diadegma species from apple leafrollers, but only in very small numbers. They were idenufied by Dr. K. Horstmann as D. apostata (Gravenhorst), and as D. fenestralis (Holmgren). Three further specimens could not be identified. Tranosema arenicola (Thomson) (figs. 31, 32) Colour characters. — Female. As in Diadegma praerogator, however legs entirely light brown, but fore and middle coxae darker brown or even partly black, hind tarsus darkened. Male. As female, but fore and middle coxae light brown. Morphological characters. — The morphological characters may be treated together with those of the other Porizontini. As their general appearance is rath- er alike, it seems useful to compare some of their dif- ferentiating characters here. In the three Diadegma species mentioned above, the epomia is lacking (fig. 29), whereas this is short but evident in Trano- sema arenicola (fig. 31). The lateral carinae on the first abdominal segment, running from its hindborder for- ward, are evident in the three Diadegma species and pass distinctly beneath the spiracles (fig. 30); in Tranosema arenicola they stop distinctly quite near the spiracle (fig. 32). As two of the Diadegma species that we reared are not important as apple leafroller parasites at all, it will be enough to refer to the publi- cation of Horstmann (1969) for the differentiating characters. Horstmann (1973) discussed the similarity of Diadegma praerogator and Tranosema arenicola. Length female 7. arenicola: 5.5—6.5 mm, oviposi- tor: 1.5—2 mm, length male: 4.5—5.5 mm. Tranosema arenicola was quite often reared from several leafroller species, however, far the most from Archips rosana. It is a solitary, larval endoparasite. Adults: 14 June—26 July. A single specimen hibernated; the adult appeared on 7 May. Campoplex spp. Campoplex is a very large genus, like so many genera of Porizontinae. It gives tremendous dif- ficulties in species discrimination. No modern revision of the genus exists. Therefore, in most cases reliable identification is impossible. Be- cause our material from apple leafrollers is only scanty, we do not want to go into details. We obtained five specimens from Pandemis cerasana, Adoxophyes orana and Acleris rhom- bana, which were idenufied by Dr. K. Horst- mann as possibly Campoplex difformis Graven- horst. Six further specimens, no doubt conspeci- fic and of which five were reared from Spilonota ocellana, could not be idenufied, just like 13 specimens, probably belonging to at least three species, reared from several leafroller species. Subfamily MESOCHORINAE According to Townes (1971) the species be- longing to this subfamily are secondary para- sites through Ichneumonidae, Braconidae, Tachinidae (Diptera) and probably other fami- lies as direct hosts. He does not give informa- tion on indirect hosts. It is not excluded that some species might be primary parasites. We reared only four specimens, a male and a female of Mesochorus silvarum Curtis and two males of Stictopisthus lineatus Thomson (det. Prof. Dr. W. Schwenke). The two specimens of the former species were reared from the cater- pillars of Pandemis cerasana and Adoxophyes orana, respectively. The two specimens of the latter species were reared from a cocoon of probably Meteorus ictericus (Braconidae) with- out caterpillar host remains, and from a caterpil- lar of Adoxophyes orana, respectively. Subfamily METOPIINAE We reared only a few specimens of this subfa- mily, belonging to four species. They can easily be recognized, especially by their smooth body and the conspicuously short and stout legs (fig. 96); they are all larval-pupal parasites. Triclistus Förster The western palaearctic species have been dealt with by Aeschlimann (1973). Our material contains two species, that were idenufied by Dr. Aeschlimann as Triclistus pallipes Holmgren and Triclistus globulipes (Desvignes). 118 Triclistus pallipes Holmgren (figs. 33, 34, 96) Ten specimens of this species, all females, were reared exclusively from Rhopobota nae- vana. Aeschlimann (1973) mentions Strophedra weirana (Lepidoptera, Tortricidae) as a host, which species is associated with Fagus and Car- pinus. Adults: 16 May—15 June. Triclistus globulipes (Desvignes) (figs. 35—37) Morphological characters. — Figures 33—36 show differences in the form of the head and in the pattern of pubescence on the gaster in both Triclistus species. Fig. 37 shows the propodeum of 7. globulipes with the longitudinal carinae. Of this species we reared two females, Archips podana being the host, and one male from Archips xylosteana. Aeschlimann (1973) mentions five more Tortricid host species. Adults: 7 July—9 August. Exochus Gravenhorst We reared two species, each represented by one specimen, from pupae of Clepsis spectrana and Acleris rhombana. As there is no modern revision of the genus, it is not possible to name the two species. Exochus sp. 1 (fig. 38) Colour characters. — Black. Head and mouth- parts, except ocellar area, bright yellow; scape black and yellow, rest of antenna brown, underside lighter. Thorax conspicuously marked with yellow; legs bright yellow, hind femur basally and apically black. Morphological characters. — See Exochus sp. 2. Fig. 38 shows the propodeum with peculiarly shaped longitudinal carinae. This species was reared from Acleris rhom- bana. Adult: 7 July. Exochus sp. 2 Colour characters. — Black. Antenna apically brownish, coxae and trochantera dark brown, remain- der of legs lighter. Morphological characters. — Differs from the for- mer species in having basal vein less curved and hav- ing complete costulae. This species was reared from Clepsis spec- trana. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) Subfamily ANOMALONINAE The species of this subfamily are larval-pupal parasites. We reared only six specimens, be- longing to two species. Two specimens of Habronyx (Camposcopus) canaliculatus (Ratze- burg) were reared from Archips rosana. Four specimens, belonging to Agrypon anxium (Wes- mael), were reared from Acleris rhombana (three specimens) and from Spilonota ocellana (one specimen). The subfamily was recently treated by Gauld & Mitchell (1977). Adults of Habronyx canaliculatus: 7 July—19 July. Adults of Agrypon anxium: 9 June—24 July. Family BRACONIDAE Reliable identification of the species of Braco- nidae that parasitize apple leafrollers seems only exceptional. The difficulties will be discussed in the special cases. As to the division into subfa- milies we follow Van Achterberg (1976). Subfamily MICROGASTRINAE Apanteles Forster Two species of this very large genus, viz., A. ater (Ratzeburg) and A. xanthostigma (Hal- iday), may be considered the most important Apanteles species associated with apple leafrol- lers by far (Evenhuis & Vlug, 1973). Nixon (1965, 1973) gave a division in groups; in sever- al publications he dealt with the species in detail (Nixon, 1972, 1973, 1974, 1976). Several groups | of Nixon are treated as independent genera by Mason (1981). We reared seven species from apple leafrol- lers. They may be distinguished especially by characters of the pronotum (figs. 39—45), of colouration of the pterostigma (figs. 46—52), of hind wing venation and of propodeum and ter- gite of the first gastral segment (figs. 53— 59). Apanteles ater (Ratzeburg) (figs. 39, 47, 53, 60) Colour characters. — Female and male. Preponder- antly black. Mouth-parts, distal part of fore femur, fore tibia, fore tarsus, base of middle tibia, middle tar- sus and all tibial spurs lighter. Wing veins for the greater part pale, however costal vein and pterostigma | distinctly pigmented; the parts of 2r proximally of Rs and short distal part of M less distinctly pigmented; pterostigma brownish yellow in the center, margins darker (fig. 47). Morphological characters. — Female and male. Propodeum with two distinct carinae, converging in EvEnHuIs & VLUG: Parasites of apple tortricids Pal EN 2 o ñ = pi » 0° ' 5 v en def, nn MAIN alt sN RICO / NN AN “li, AO \ N ON PA i 1, MA UT) ‘ U 1 N 1 ¥ à \ N 8 Wry \ di 33 if | \ EAN il da N I o Sn \u 1 vay i Woy A ‘ 1 M Ihn Nil Un rn lu \ u ui Le TN SON 1 TIVO A ' ET dado 34 ai if (1 WI U in 1 Il i QUI ini AS u “ul z_ meal Kann ai. TEE TT EIT OS 119 Figs. 33— 34. Triclistus pallipes: 33, head laterally; 34, gaster dorsally. Figs. 35—37. Triclistus globulipes: 35, head laterally; 36, abdomen dorsally; 37, propodeum. Fig. 38. Exochus sp. 1: propodeum. Figs. 39-45. Micro- gastrinae spp., pronotum laterally: 39, Apanteles ater; 40, A. xanthostigma; 41, Dolichogenidea sicaria; 42, D. laevigata; 43, D. corvina; 44, D. dilecta; 45, D. longicaudus. 120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) front and thus forming a V, a litte coarse, but smooth and shiny between and just outside the keels. First tergite twice as long as broad, the sides rather strongly converging distally (fig. 53). Ovipositor valves about as long as hind tibiae. Length female: 2—2.5 mm, ovipositor: 0.5—1 mm, length male: 2—2.5 mm. Nixon (1976) states that this species is poorly characterized as a whole, but that it is easily rec- ognizable in the female by the presence of a fine, curved spine on the ventral side of the last tarsal segment of the front leg (fig. 60). This species has been treated by Wilkinson (1945) in such an accurate way, that we have no doubt about its identity. This author also studied Ratzeburg’s type material. The species was originally described as Microgaster carbon- arius Ratzeburg, 1848, but later on renamed Microgaster ater Ratzeburg, 1852, as the former name appeared to be preoccupied by Microgas- ter carbonarius Wesmael, 1837. Ratzeburg (cf. Evenhuis & Vlug, 1973) states about the host: “Von mir aus versponnenen Ap- felblüthen erzogen, wahrscheinlich aus der bru- mata, die diese versponnen hatte”. However, the host may as well have been some species of Tortricidae instead of Operophtera brumata. Wilkinson states the parasite to have been reared from Cacoecia (= Archips) podana, Hyponomeuta malinella (= Yponomeuta mali- nellus), Tortrix (= Lozotaenia) forsterana, No- tocelia udmanniana, and a specimen from a leaf- roller of Ribes nigrum. From this enumeration only Yponomeuta malinellus does not belong to the Tortricidae. It does not seem improbable that the mention of this host is due to a mistake, as Yponomeuta caterpillars live gregariously in large nests and some Tortricid host caterpillar may have been mixed with them. According to our opinion Apanteles ater must be considered a specialized parasite of the caterpillars of various Tortricidae. However, the species seems to show some preference for the larger caterpillars. We reared rather many specimens, almost exclusively from Archips podana and Pandemis heparana, but only a few from Adoxophyes orana, Ptycholoma lecheana, and Acleris spp. This is the only gregarious species within the genus that we reared from leafrollers associated with apple. Like other Apanteles species it is a larval endoparasite. Adults: 25 May-18 September. Apanteles xanthostigma (Haliday) (figs. 40, 46, 54) This species is very similar to the preceding one, both in colour and in morphological re- spect. Colour characters. — Female and male. The species differs from A. ater by the centre of the pterostigma being more yellowish instead of white (fig. 46) and in the female sex by the legs being more extensively light-coloured. Morphological characters. — Female and male. The most striking difference with the preceding species is, according to our opinion, the structure of the propo- deum. This contains a longitudinal trough in the mid- dle, which is at least distinct in its hind part and there bordered by short longitudinal carinae (fig. 54). The propodeum, on the whole, is also coarser. The female does not have a curved spine on the last tarsal segment of the fore leg. Length female: 2.5—3 mm, ovipositor: 0.5—1 mm, length male: 2.5—3 mm. The species has been dealt with by Nixon (1976), who mentions a number of hosts, be- longing to several families of Microlepidoptera. Telenga (1955) also enumerates the hosts, be- longing to various Lepidopterous families. Zwölfer (1962) characterizes its host preference as follows: „Die von uns seit einigen Jahren un- tersuchte, recht polyphage Braconide Apanteles . xanthostigma Hal. beschränkt ihr Suchgebiet - praktisch auf die Strauch- und Baumschicht, greift hier aber eine grössere Zahl von Wirten, meist Wickler und verwandte Kleinschmetter- lingsgruppen an”. We reared a large number of specimens from several leafroller species of apple (cf. table 1), but we have no experience with the hosts of other Lepidopterous families. If indeed only one and the same species is involved, it might be considered a polyphagous parasite of Lepido- ptera. Apanteles xanthostigma is a solitary en- doparasite. Adults: 13 May—8 September. Besides the two species mentioned, we reared some more Microgastrinae, however in quite low numbers: Dolichogenidea corvina (Reinhard) from Nixon’s “metacarpalis” group; Dolichogenidea laevigata (Ratzeburg), D. dilec- ta (Haliday), D. sicaria (Marshall) and D. longi- caudus (Wesmael) from Nixon’s “laevigatus” group. As these species have been dealt with thoroughly in taxonomical respect by Nixon EveNHuis & VLUG: Parasites of apple tortricids 121 & Figs. 46—60. Microgastrinae spp. 46, Apanteles xanthostigma, wings. 47—52, pterostigma: 47, Apanteles ater; 48, Dolichogenidea sicaria; 49, D. laevigata; 50, D. corvina; 51, D. dilecta; 52, D. longicaudus. Figs. 5359. Propodeum and first tergites: 53, Apanteles ater; 54, A. xanthostigma; 55, Dolichogenidea sicaria; 56, D. laevi- gata; 57, D. corvina; 58, D. dilecta; 59, D. longicandus. Fig. 60. Apanteles ater, last tarsal segment of fore leg of female ventrally. 122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) (1972, 1973) and as they apparently do not play any important role in the natural control of apple leafrollers, it does not seem necessary to treat them here in detail. Our material is far too poor to allow any con- clusion as to their host specialization. In figs. 39— 60, morphological details of the sever- al Apanteles species are given, which may be sufficient to separate them. Twelve further specimens of the subfamily Microgastrinae, all belonging to the genus Lis- sogaster Bengtsson (= Microgaster auct.), were reared. They belong to at least four species and apparently are not important as parasites of apple leafrollers. Nixon (1965) states: “Taxonomically, Micro- gaster is in a state of confusion and with regard to European species, there has been no im- provement on Thomson’s treatment of the ge- nus”. Thomson’s work appeared between 1869 and 1897! Subfamily AGATHIDINAE We reared only two species of this subfamily from apple leafrollers. Both species seem, at least within the host range that we investigated, restricted to Spilonota ocellana and are rather common solitary endoparasites. The subfamily may be easily recognized by the very narrow ra- dial cell in the fore wing (fig. 64). Notauli deep in both species. Agathis dimidiator (Nees) (figs. 61—63) Colour characters. — Female. Black. Mouth-parts yellow. Legs reddish yellow, often with somewhat lighter trochantera; hind coxa black or largely black; hind tibia in the distal two third more or less black, for the rest whitish, lighter as the remainder of the legs; hind tarsi and ultimate ends of fore and middle tarsi dark. Male. As female, but second gaster segment con- spicuously light. Morphological characters. — Female. The most salient characters of this species are, in comparison with the next one, the head elongated downwards slightly (fig. 61) and the first and generally the second tergite being partly rather coarsely striated longitudi- nally (fig. 62). Male. As female, but striae on second tergite largely varying and in some cases even lacking (fig. 63). Length female: 4—4.5 mm, ovipositor: 3.5 mm, length male: 4—4.5 mm. The characters given here, borrowed from Telenga (1955) who placed the species in the ge- nus Microdus Nees, are rather doubtful. A revi- sion of this genus and allied genera seems ur- gent. Unul so far we shall use the species name dimidiator, quoted from Telenga (1955). As stated above we reared this species only, but rather commonly, from Spilonota ocellana. Telenga (1955) mentions Archips rosana, Croe- sia bergmanniana and Epinotia tetraquetrana as hosts, all Tortricids. Adults: 4 July—13 August. Agathis rufipes (Nees) (Braunsia rufipes (Nees) (in Telenga, 1955) (figs. 64— 66) Colour characters. — Female and male. Black. Mouth-parts, tegula and legs reddish yellow, except for the extreme distal end of the hind tibia and the greater part of the hind tarsus; tibial spurs all more or less whitish. Morphological characters. — Female and male. As in the former species first radial and discoidal cells fused (fig. 64). Distal part of first tergite with fine lon- gitudinal striation; base smooth, in contrast to the for- mer species, where it is wrinkled all over its surface (figs. 65, 66); on each side a conspicuous, dorsolateral carina that does not reach the hind border, these cari- nae more lateral and thus inconspicuous in the former species. There are transverse grooves in the middle of each of the fused second and third tergites, so that it seems if one long tergite after the first tergite were di- vided into four sections by three transverse grooves; striation of first tergite continuing onto the first two sections. Length female: 4—5 mm, ovipositor: 3.5 mm, length male: 4—5 mm. This species superficially resembles the for- mer in many respects, may, however, easily be distinguished from it by the colour of the hind legs and by the sculpture of the gaster, as men- tioned. We reared this species only from Spilonota ocellana, but rather commonly. Telenga (1955) mentions quite a lot of host species, which may at least be partly erroneous, especially where non-Lepidopterous hosts are involved. Betz & Schwerdtfeger (1970) mention one specimen that was reared from Teleia luculella (Lepido- ptera, Gelechiidae) on oak. Adults: 29 June—30 July. A single specimen appeared on 8 June. Subfamily CHELONINAE From this subfamily we reared only two Ascogaster species. Though both species super- ficially resemble each other very much, they may be easily distinguished. Just like other EvenHuis & VLUG: Parasites of apple tortricids 123 members of the subfamily, as fas ar known, they are solitary egg-larval endoparasites. Ascogaster quadridentata Wesmael (figs. 67—69) Colour characters. — Female and male. Black. Scape for the greater part, rest of the antenna down- side, trochantera, fore femur distally, and fore tibia reddish yellow; tibial spurs whitish yellow; middle leg in male often a little lighter, at least tibia proximally. Morphological characters. — Female and male. Head with coarse sculpture, clypeus produced in a small angle (fig. 67). Thorax and abdomen with coarse sculpture, coarser than head, thorax coarsest (fig. 68). Horizontal front part and perpendicular hind part of propodeum distinctly separated; the boundary line with four strong teeth, from which the inner are a little smaller than the outer (fig. 69). Length female and male: 3.5—4 mm. The female lays the egg in the host egg, and the adult parasite emerges from the full-grown or almost full-grown caterpillar (Cox, 1932). Ascogaster quadridentata seems specialized on Tortricidae Olethreutinae; the females of these moths lay their eggs separately on the host plants (Evenhuis, 1969, 1974). We reared this species commonly from Spilonota ocellana and Hedya nubiferana. It is a well-known parasite of Laspeyresia pomonella. Adults: 19 June—24 July. Ascogaster rufidens Wesmael (figs. 70—72) Colour characters. — Female and male. Black. Mandible, palps, antenna basally downside, tegula, trochantera, femora and tibiae reddish yellow; middle of the tibiae and femora of the middle and hind legs a little darker; tibial spurs whitish yellow. Morphological characters. — Female and male. Sculpture somewhat less coarse than in the preceding species. There are three small, reddish teeth on the lower border of the clypeus (fig. 70). Wesmael (1835) (see Fahringer, 1934) mentions this explicitly. Length female and male: 3.5—4.5 mm. These characters, combined with others men- tioned by Wesmael in his detailed description, are sufficient to recognize the species. Charac- ters of mesoscutum and propodeum, to be com- pared with those of A. quadridentata, are shown in figs. 71 and 72, respectively. This species resembles the preceding one, but may be distinguished at once by the colour of the legs — preponderantly black in A. quadri- dentata and reddish yellow in A. rufidens — and by the structures of the propodeum. In contrast to the former species, its hosts seem restricted to the Tortricidae Tortricinae, which lay their eggs in batches. We reared many specimens, especially from Pandemis cerasana and P. heparana, but also often from Adoxo- phyes orana. Matthey (1967) reared the species from Pandemis cerasana, P.corylana and Batodes angustiorana on oak. Adults: 25 May—21 July. Subfamily EUPHORINAE Meteorus ictericus (Nees) (figs. 73— 75, 95) Colour characters. — Female. Reddish yellow. An- tenna and ocellar space darker. Often the whole tho- rax is entirely dark or even black, except for the great- er part of the prothorax. In some specimens only the propodeum is dark and often the distal ends of the tib- iae and the tarsi. Often the first tergites and the mid- dle part of the last tergites are black, or the abdomen is almost completely black except for the fused second and third tergites. Male. As female, but often a little lighter. Morphological characters. — Female and male. In figs. 73, 74 and 75, thorax and the first three tergites are shown, which might be of interest in the separa- tion of this species. The propodeum is more or less ir- regularly reticulated and shows some variability; in specimens reared from Adoxophyes orana it seems coarser than in other specimens. The first gastral seg- ment shows longitudinal wrinkles and two dorsal pits (dorsope: van Achterberg, 1974). Length female: 3.5—5 mm, ovipositor: 2 mm, length male: 3.5—4.5 mm. According to Fischer (1970) a critical revision of the species of Meteorus belongs to the most difficult questions in the taxonomy of Braconi- dae. The genus as such may be easily recog- nized, but separation of the many species is of- ten extremely difficult. In Fisher’s key our spe- cies runs to Meteorus ictericus. A critical revision of the Palaearctic species is given by Huddleston (1980). This parasite is one of the most common spe- cies that we reared from several apple leafroller species (cf. table 1). Adults: 2 May—28 August. A single speci- men appeared on 6 April. Subfamily ROGADINAE Oncophanes minutus (Wesmael) (figs. 76—78) Colour characters. — Female. Body dark or even black. Basal part of antenna brown, mouth-parts, in- cluding palps, yellowish. Legs and ovipositor yellow. Pronotum partly, tegula and extreme superior part of 124 TiJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) Figs. 61—63. Agathis dimidiator: 61, head anteriorly; 62, gaster female dorsally; 63, abdomen male dorsally. Figs. 64—66. Agathis rufipes: 64, wings; 65, gaster female dorsally; 66, gaster male dorsally. Figs. 67—69. Asco- gaster quadridentata: 67, front edge of clypeus; 68, thorax dorsally; 69, propodeum. Figs. 70—72. Ascogaster rufidens: 70, front edge of clypeus; 71, thorax dorsally; 72, propodeum. EvenHuis & VLUG: Parasites of apple tortricids 125 Figs. 73—75. Meteorus ictericus: 73, thorax dorsally; 74, propodeum; 75, first three tergites. Figs. 76—78. On- cophanes minutus: 76, thorax dorsally; 77, propodeum; 78, first three tergites. Figs. 79—80. Bracon sp.: 79, tho- rax dorsally; 80, propodeum. Figs. 81—87. Macrocentrus linearis: 81, head anteriorly; 82, mandible; 83, thorax dorsally; 84, wings; 85, tarsal claw; 86, propodeum; 87, first three tergites. 126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) mesopleurum reddish brown. Second tergite for the greater part yellowish. Male. As female, but second tergite not as yellow. Morphological characters. — Female and male. The exact difference between Oncophanes minutus and O. laevigatus (Ratzeburg) (= O. lanceolator (Nees)) is not clear from Fahringer’s revision (1930). This 1s the last revision of the genus anyway! Occipital carina above fine, but distinct. Mesopleurum polished, but extreme superior part coarse. Mesoscutum sparsely covered with fine long hairs, parapsidal furrows evi- dent (fig. 76). Propodeum with delimited areas (fig. 77). First tergite striated longitudinally, in front with two distally converging carinae, space between carinae smooth. Second and third tergites only sepa- rated by a shallow, in the middle almost faded trans- verse groove, second tergite longitudinally striated. The extension of this striation shows a rather large variability (fig. 78). Length female: 2—2.5 mm, ovipositor: 0.5 mm, length male: 2 mm. Oncophanes minutus is a gregarious ectopara- site. We reared it almost exclusively from Adox- ophyes orana and only from a few localities. It must be considered an occasional parasite of apple leafrollers, without economic importance. Adults: 6 August—26 August. Subfamily HOMOLOBINAE Charmon cruentatus Haliday We reared only one specimen from a caterpil- lar of Spilonota ocellana. It was identified by Dr. C. van Achterberg, Leiden. Subfamily BRACONINAE Bracon sp. (figs. 79, 80) Among Braconidae Bracon is one of the largest genera. It is considered one of the most difficult genera of the Hymenoptera. À modern revision is lacking and sufficient reliable species characters are not known at the moment. There are many morphological differences between this species and the preceding one. We shall de- scribe some that are most conspicuous, hoping that they may be an indication for the idenuty of the species, after the genus, eventually the group to which it belongs, has been revised. Colour characters. — Female. Body dark, locally lighter, especially thorax and gaster downside. Morphological characters. — Female. Mesoscutum smooth, with percurrent, converging notauli (fig. 79), which do not meet distally. Notauli shallow, with long setae, orientated mainly in two directions, in- wards and outwards. Hind part of mesoscutum and also scutellum densely pilose. Propodeum smooth, without areas (fig. 80). First tergite with two deep, distally converging grooves that enclose a triangular field. Length female: 2.5 mm. Of this species we only reared one female from Adoxophyes orana, which emerged on 12 August. Bracon obscurator Nees This species differs from the former by its smaller size and by the mesoscutum being much less pilose, the hairs on its hind part almost lack- ing. It was, provisionally, identified by Dr. C. van Achterberg. Length male: 1.5—2 mm. We reared two males from one caterpillar of Spilonota ocellana. Subfamily MACROCENTRINAE In this subfamily three Macrocentrus species were reared from apple leafrollers, mainly M. linearis (Nees), and a few specimens of the re- lated M. pallipes (Nees) and one specimen of M. thoracicus (Nees). A modern revision of Macrocentrus is that of Eady & Clark (1964). Macrocentrus linearis (Nees) (figs. 81—87) Colour characters. — Female and male. Preponder- antly light brownish, however variable. Interocellar space and a point on both sides of the insertion of the wings very dark to black. Upper side of thorax and gaster often dark in varying tinge and extension. Pterostigma dark centrally, with lighter margins (fig. 84). Morphological characters. — Female and male. As morphological differences with related species have been dealt with by Eady & Clark (1964), we refer to that publication. We may add that the mesopleurum in this species is rather uniformly and widely punctate at its centre and not more densely in the furrow, which is only slightly pronounced; prepectal carina strongly curved (in M. pallipes the mesopleurum is ir- regularly punctate, and especially in the shallow, however pronounced furrow above the hind coxae densely punctate; prepectal carina straight). Figs. 81—87 show head, mandible, thorax above, wings, tarsal claw, propodeum, and first three ter- gites, respectively. Length female: 4—5 mm, ovipositor: 6 mm, length male: 4—5 mm. We reared this parasite often from Pandemis cerasana, P. heparana and Archips podana, thus from the larger leafroller caterpillars. After Ea- dy & Clark (1964) the parasite has also been EvenHuIs & VLUG: Parasites of apple tortricids 127 reared from other Microlepidoptera than Tor- tricidae. It is a gregarious larval endoparasite, appar- ently showing polyembryony. In most cases the progeny of one single host consists of either males or females exclusively. However, in a number of cases we reared both sexes from one single host, probably indicating that then at least two eggs were laid in a single host, giving rise to a female and a male progeny respectively. Adults: 26 June—11 August. Macrocentrus pallipes (Nees) This species is very similar to the preceding one. Differentiating characters have been given by Eady & Clark (1964); a few supplementary characters were mentioned under M. linearis. We reared 18 female specimens from one sin- gle caterpillar of Hedya nubiferana. One specimen of Macrocentrus thoracicus (Nees) was reared from a caterpillar of Spilono- ta ocellana. It was, just like M. linearis and M. pallipes, identified by Dr. C. van Achter- berg, Leiden. Superfamily CHALCIDOIDEA This superfamily of parasitic Hymenoptera has found a number of prolific workers during the last decennia. Especially from the groups that interest us as parasites of apple leafrollers, there exist some modern taxonomic revisions. Only one species, Colpoclypeus florus, is of real importance in respect to apple leafrollers; the others were only reared in small numbers. Thus we will not treat them in detail and, as far as their identity is involved, refer to the revi- sions. Family PTEROMALIDAE Habrocytus chrysos (Walker) (fig. 101) This species has been dealt with by Graham (1969). We reared only three specimens as hy- perparasites of the caterpillars of Adoxophyes orana through Scambus brevicornis, Onco- phanes minutus and an unidentified Ichneumo- nid, respectively. Graham mentions some more hosts, so that H. chrysos must be considered a widely specialized hyperparasite of various Lepidoptera, through various species of parasit- ic Hymenoptera as direct hosts. Adults: 28 September. Habrocytus semotus (Walker) We reared this species three times from apple leafrollers, namely once as a hyperparasite of Archips podana through Scambus brevicornis and two times from pupae of Adoxophyes ora- na, possibly as hyperparasites. Graham (1969) mentions the parasite both as a primary and as a secondary parasite of various Lepidoptera, but also from certain Coleoptera and in one case from Hymenoptera Diprioni- dae. Adults: 25 August—25 September. Dibrachys cavus (Walker) (figs. 97, 102) According to Graham (1969) the species of Dibrachys have not yet been fully sorted out. On the authority of Peck (1963) he mentions Dibrachys cavus to be a very polyphagous para- site, usually hyperparasitic. We reared only two small specimens from co- coons of Apanteles ater, parasitizing a caterpil- lar of Pandemis sp. and two specimens from a cocoon of Diadegma sp., Archips podana being the host. Adults: 6 July Family EULOPHIDAE Pnigalio pectinicornis (Linnaeus) We reared only two specimens of this species, the hosts being caterpillars of Adoxophyes ora- na and Rhopobota naevana, respectively. Askew (1968) mentions it as a parasite of Microlepidoptera, especially of Lithocolletis (= Phyllonorycter), which are leafminers. Adults: 4 September. Sympiesis xanthostoma (Nees) We reared two specimens, both from caterpil- lars of Adoxophyes orana. Askew (1968) men- tions it a quite common parasite, especially of Tortricidae and Gracilariidae. Adults: 30 August and 4 October. Colpoclypeus florus (Walker) (figs. 98, 103) This is no doubt the most important parasite of Adoxophyes orana in apple orchards. We reared it in large numbers in some orchards dur- ing late summer and early autumn. There seems to be a serious discrepancy between the pheno- logy of the parasite and of Adoxophyes orana, as we were not able to recover the parasite before 128 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) 102 103 Figs. 88—95. Fore wing: 88, Lysibia nana; 89, Teleutaea striata; 90, Glypta varicoxa; 91, Mesochorus sp.; 92, Habronyx canaliculatus; 93, Agrypon anxium; 94, Lissogaster sp.; 95, Meteorus ictericus. Fig. 96. Triclistus pal- lipes, fore leg. Figs. 97—98. Fore tibia: 97, Dibrachys cavus; 98, Colpoclypeus florus. Fig. 99. Apophua cicatrico- sa, propodeum. Fig. 100. Diadegma sp, first tergite. Figs. 101—102. Fore wing: 101. Habrocytus chrysos; 102, Dibrachys cavus. Fig. 103. Colpoclypeus florus, front margin of clypeus. EvenHuIs & VLUG: Parasites of apple tortricids July (Evenhuis, 1974b). During and after this month its population increases rapidly. The dis- crepancy makes the feasibility of the parasite in integrated control doubtful. In spite of efforts in later years, we could not find any alternative leafroller host on which it could maintain itself and reproduce, neither in apple orchards, nor outside on other foodplants than apple. Thus the problem remains unsolved. More knowl- edge of the bionomics and ecological particu- larities of the parasite seems a prerequisite in us- ing it in integrated control in apple growing. Adults: From the end of July until October. It is interesting that in Swiss material, kindly sent to us by Dr. A. Schmid, Nyon, Switzer- land, four additional species of parasitic Hymenoptera, reared from apple leafrollers, but only in small numbers, are represented: Sympie- sis acalle (Walker) from Adoxophyes orana, Dicladocerus westwoodu Westwood from Ar- chips sp., Euderus albitarsis Zetterstedt, and Go- niozus claripennis Förster, both from Adoxo- phyes orana. The three former species are Chal- cidoidea Eulophidae, the latter belongs to the Bethylidae. KEY TO THE ADULT PARASITES OF APPLE LEAFROLLERS IN THE NETHERLANDS The characters used in this key are simple. Differences between the taxa only refer to spe- cies that are dealt with in the body of the paper. Where species identification is very doubtful, the key generally does not run to species level, especially when the species are no important apple leafroller parasites. 1. Fore wings with longitudinal veins and cross veins, delimiting several cells (cf. figs. 88—95); (Ichneumonoidea) ........ 2 — Venation of fore wing strongly reduced (figs. 101, 102). Small species with metallic green or blue body; (Chalcidoidea) .… 44 2. Second recurrent vein present (figs. 88—93: srv); first cubital cell and discoidal cell fused (figs. 88—93). Second and third gas- ter segments separated (cf. figs. 3, 14, 15, 34, 36); (Ichneumonidae) — Second recurrent vein lacking; first cubital cell (Cu) and discoidal cell (Di) separated by a vein (figs. 94, 95). Second and third gaster segments fused (cf. figs. 62, 63, 65, 66, 75, 78, 87); suture, however, generally visible; (Braconidae) 3. Gaster depressed, petiole broadly attached . Propodeal to propodeum (cf. figs. 14, 15).......... 4 Gaster compressed, petiole proximally slen- den(etsnes 2082/1100). 2727.00 18 . Tergites 2, 3 and 4 with oblique furrows (cf. fips SIMA) oretta: mes dato 5 Tergites 2, 3 and 4 without furrows (cf. ies 3004 Ba Len tel 9 . Areolet present (fig. 89: ar). Species gay- coloured black and yellow ............... Teleutaea striata Areolet absent (fig. 90). Yellow markings IEssfconspleuous eee 6 . Frontal tibial spur reaching to middle of first tarsal segment or beyond (fig. 19) ... 7 Frontal tibial spur not reaching to middle of first tarsal segment (figs. 23, 27). Genal ca- rina not strongly sinuated (fig. 25)......... Glypta varicoxa . Genal carina strongly sinuated (fig. 21); clypeus with small incision (fig. 20)........ Glypta nigrina Genal carina moderately sinuated (fig. 17); clypeus without incision (fig. 16)........ 8 carinae strongly developed essen reger Apophua cicatricosa Propodeal carınae more or less obliterated .. Apophua sp. . Clypeus and face conspicuously strongly convex (figs. 33, 35). Legs, especially femo- ra, conspicuously stout (fig. 96)........ 10 Clypeus and face not convex. Legs rather Slender nr eee 12 Antennal sockets separated by a high lamel- la, the latter with a deep median groove dorsally, just below ocellus ........... fil Antennal sockets not separated by a lamella ostsee nc eye owe tie. Exochus spp. . Tergites almost completely pubescent (CRT) en Triclistus globulipes Tergites almost completely glabrous. (fig. EN III SA Triclistus pallipes . Inner orbits at the level of the antennal sockets deeply incised (figs. 5, 7, 8) ..... 13 Inner orbits not or weakly incised (fig. 1) seats on cine te veen ode ees Seay 15 . Head of male with yellow markings. Ovi- positor curved downwards (fig. 9)......... pressato Apechthis spp. Head of male black. Ovipositor straight Bead i 14 . Head beneath antennae and centre of meso- scutum finely punctate (fig. 5). Tergites wholly black.......... Itoplectis alternans Head beneath antennae and centre of meso- scutum roughly punctate (fig. 7). Hind bor- 130 119. 16. 117% 18. NO). 20. Zale 26. ePAtcolet presente ee . Second tergite about as long as wide TIJDSCHRIFT VOOR ENTOMOLOGIE, denoitersmeslbrommish au ae to SAR AL Itoplectis maculator Clypeus incised (fig. 1). Tergites roughly punctate except for narrow hind strip which is finely transversely striated (fig. 3) BCM IGT Bruch) Scambus brevicornis Clypeus not incised. Tergites not roughly punctate 16 Areolet more or less regularly pentagonal, sometimes outer crossvein lacking (fig. 88) Cryptinae Areolet not pentagonal Claws pectinate (fig. 10). Scutellum, post- scutellum and propodeum with yellow markings Phytodietus segmentator Claws simple. Scutellum, postscutellum and propodeum entirely black................ RIDA Ne ek Lissonota complicator Areolet large, rhombical (fig. 91).......... Mesochorus sp. MORE AR ER 19 Body conspicuously slender. First gaster segment long and narrow, but not petio- lated anteriorly, following segments strong- Hcompressed IR 20 Body not conspicuously slender. First gas- ter segment petiolated anteriorly (fig. 100), following segments compressed........ Di Postnervulus intercepted near or below middle (fig. 92: arrow) Habronyx canaliculatus Postnervulus intercepted distinctly above middle (fig. 93: arrow) ... Agrypon anxium Pronotum moderately to strongly striated; central part of propodeum excavated and transversely striated...... Campoplex spp. Pronotum rather smooth or punctate; cen- tral part of propodeum flattened or a little CONVEX MANNEN Ne orn RG 22 VERROEN it 23 Hind tibia whitish, with dark ring proxi- mallygandidiscally PEER PRE E 24 Tranosema arenicola Areolet absent Diadegma apostata Diadegma praerogator Second tergite much longer than wide... ... Diadegma fenestralis . Gaster segments fused into a carapace with- out sutures, oval, roughly sculptured ... 26 Only second and third gaster segments fused, sutures visible (figs. 62, 63, 65, 66, ISMEA REA At ay Clypeus with two small incisions (fig. 70). 33% 36. DEEL 126, AFL. 6 (1983) Hind border of propodeum with two lateral spinal processes only (fig. 72). Legs largely rufous Ascogaster rufidens Clypeus entire (fig. 67). Hind border of propodeum except for lateral spines also with two medial spinal processes (fig. 69)... Ascogaster quadridentata . Radial cell distinctly defined and narrow RR... 28 Radialicellmotnarro were 29 . First and second tergites longitudinally striated, in the female also third tergite par- tially striated (figs. 65, 66).. Agathis rufipes Only first tergite longitudinally striated (figs. 62, 63) Agathis dimidiator . Some veins, e.g. radial vein, pale and hardly visible, distinctly different from the other veins (fig. 46) 30 At least radial vein well-developed . Second cubital cell present, small (fig. 94)... Lissogaster spp. Second cubital cell absent (fig. 46) 31 . Pterostigma hyaline (figs. 46, 47), borders slightly darkenedia=) Sea 32 Pterostigma dark, sometimes with light bas- al spot (figs. 48—52) 33 . Pronotal groove deep and crenulated over its whole length (fig. 39); propodeum with two carinae, constituting a V, space be- tween rather smooth (fig. 53). First tergite narrowed apically Apanteles ater Pronotal groove superficial, slightly crenu- lated in caudal half (fig. 40); propodeum with median, longitudinal, rather roughly: sculptured trough (fig. 54). Side borders of first tergite more or less parallel........... Apanteles xanthostigma light proximal spot Pterostigma with (figs. 49, 51) Pterostigma without light proximal spot (figs. 48, 50, 52) 35 . Propodeum strongly punctate reticulate (fig. 58) Dolichogenidea dilecta Propodeum smooth (fig. 56) Dolichogenidea laevigata . Middle hind part of propodeum excavated, crenulated in its caudal end (figs. 57, 59).... wdn EN SE SEE 36 Propodeum almost entirely smooth, with a few superficial wrinkles in the middle hind part (fig. 55) Dolichogenidea sicaria Rim near hind side of pronotum distinct (fig. 45). Pterostigma entirely dark (fig. 52) hg Dolichogenidea longicaudus Pronotal rim only superficial (fig. 43). EvenHuIs & VLUG: Parasites of apple tortricids 131 Centre of pterostigma somewhat lighter (fe 20) ORARIE Dolichogenidea corvina 37. Second tergite at least as long as wide ... 38 — Second tergite distinctly wider than long vesten 41 38. Mesoscutum divided into three vaulted lobes, the middle the highest (fig. 83) ... 39 — Mesoscutum more or less flat, not divided Ene ete. Charmon cruentatus 39. Pterostigma almost colourless (fig. 84) ..... a on. Macrocentrus thoracicus — Pterostigma dark, proximally somewhat ele bone REEN 40 40. Body predominantly rufous .............. ec... Macrocentrus linearis — Body predominantly black............... Ja à ce en. eee Ag Macrocentrus pallipes 41. First gastral segment narrow proximally, with two deep pits (dorsope) in front of the middlefio 75)... Meteorus ictericus — First gastral segment broadly inserted to propodeum, without deep dorsope (fig. 78) benson TE 42 42. Propodeum with dilimited areas, separated byairesularicarinae; (fig. 77) dolo. Hiko au... Oncophanes minutus — Propodeum without delimited areas (fig. BOMEN ele cody ent wo cts 43 43. Mesoscutum densely pilose (fig. 79) ....... Bracon sp. — Mesoscutum less pilose, hind part almost Darren. sori... Bracon obscurator 45. Postmarginal vein as long as stigmal vein (HSM Raw) ee Dibrachys cavus — Postmarginal vein longer than stigmal vein (GE TONE ON ee Habrocytus spp. 46. Anterior margin of clypeus medially nar- rowly incised (fig. 103) Colpoclypeus florus es @lypeusnorineised. i... 47 DA ropodeumiwaithtearinaer. avonden. OER Aa: Pnigalio pectinicornis — Propodeum without carinae .............. Sympiesis xanthostoma Discussion Among the Hymenopterous parasites, asso- ciated with apple leafrollers, some species were reared commonly and in rather large numbers, whereas most of them were only obtained occa- sionally. Many of these latter are no rare species whatever, but do not really belong to the apple orchard community (table ID) In provisional investigations parasites were also reared by collecting leafroller caterpillars on other foodplants and in different habitats. Many of these parasites belong to other species than those from apple orchards or to the same species but in entirely different ratios. The same feature holds also for some leafroller species, e.g., Adoxophyes orana was only found occa- sionally outside apple orchards. If these investi- gations might prove to have a general bearing, it could mean that leafroller caterpillars in habitats adjoining apple orchards, only play a minor role as a reservoir for apple leafroller parasites. It must be born in mind that there are several types of apple orchards, with different ecologi- cal conditions to satisfy the needs of the several parasite species. As an example of the special needs of a leafroller parasite might be men- tioned Zwölfer’s remark concerning Apanteles xanthostigma, the female of which only looks for its hosts in the shrubs and the lowermost parts of the trees (Zwölfer, 1962). Such ecologi- cal preferences might at least partly explain the scarcity in apple orchards of so many parasite species that are quite common elsewhere. It is obvious that numerous investigations are still necessary for obtaining a more clear insight in the intricate interrelationships between the different leafroller species on apple and their parasites. By rearing parasites from known hosts, large series of parasites belonging to one species, may be obtained. Rearing not only gives information about host specificity of the parasite, but also may be of interest in under- standing species variation. It does not seem superfluous to emphasize the tremendous importance of a sound taxo- nomical base in distinguishing the species. From our investigations it may be concluded that for most groups of Hymenopterous leafroller para- sites this base is not present and this might ap- ply as well to the group of parasitic Hymeno- ptera as a whole. Parasitic Hymenoptera constitute an insect group very rich in species. We estimate that Dutch species far outnumber 7000, the total number of insect species in the Netherlands probably not exceeding 25,000. We do not ven- ture to give figures for larger areas, but we sup- pose that parasitic Hymenoptera are extraordi- nary numerous throughout the world. Large genera with quite similar species are no excep- 182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 6 (1983) Table 1. Enumeration of parasitic Hymenoptera with the numbers that were reared from their apple leafroller hosts. Acleris spp./Croesia holmiana Hedya nubiferana Host unknown Ptycholoma lecheana Rhopobota naevana Pandemis cerasana Pandemis heparana Archips rosana Archips xylosteana Clepsis spectrana Adoxophyes orana Spilonota ocellana Scambus brevicornis Itoplectis alternans Itoplectis maculator Apechthis compunctor Apechthis quadridentatus 1 Apechthis rufatus Phytodietus segmentator 5 Gelis spp. 2 1 1 Acrolyta sp. 1 Lysibia nana 3 Teleutaea striata lh 3 65 7 2 Apophua cicatricosa 1 6 Apophua sp. 1 Glypta nigrina Pr Glypta varicoxa 20 Lissonota complicator 22 Diadegma praerogator . DAF AERO À mM 9 2 9 Diadegma apostata 1 1 Diadegma fenestralis 1 Tranosema arenicola te 0m 225" 2 1 3 2 Campoplex difformis 1 1 3 Campoplex sp. 5 1 Campoplex spp. 2 4 DA EDEN 33 Mesochorus silvarum 1 1 Stictopisthus lineatus 1 1 Triclistus pallipes 10 Triclistus globulipes Die Exochus sp. 1 1 Exochus sp. 2 1 Habronyx canaliculatus 2 Agrypon anxium Apanteles ater 24 43 WB QU Apanteles xanthostigma 26 8 38 2 14 2 7 Dolichogenidea laevigata 1 Dolichogenidea dilecta 1 Dolichogenidea longicaudus 1 2 Dolichogenidea corvina 1 1 1 Dolichogenidea sicaria Lissogaster spp. MG 1 2 2 Agathis dimidiator 2/7 1 Agathis rufipes 30 2 Ascogaster quadridentata 24 5085 3 2 ron Archips podana no pi N oo Won N n mo es m N ar an Na Ww Ce 00 CS — — an — a Ascogaster rufidens 13 28 Meteorus ictericus Waor WARY. 4 24 19 1 1 Oncophanes minutus 1 10 Bracon sp. 1 Bracon obscurator 1 Macrocentrus linearis 13 113 19 1 Macrocentrus pallipes 1 Macrocentrus thoracicus 1 Charmon cruentatus 1 Habrocytus chrysos 3 Habrocytus semotus 1 i 2 Dibrachys cavus 1 1 1 Pnigalio pectinicornis 1 1 Sympiesis xanthostoma 2 Colpoclypeus florus 3 248 3 EvenHuis & VLUG: Parasites of apple tortricids 133 tion in this group. Thus it is hardly surprising that identification meets with very large diffi- culties. Yet, exact species discrimination seems essential, as very similar species show consid- erable differences in biological and ecological respects. Descriptions of species by the earlier taxo- nomists are generally too poor to allow of any conclusion about their identity. They often suit more than one species. Later authors, who tried to interpret them in their own way, added new species, just as ill-described. In this way taxo- nomic literature has been overburdened with “species” from which no one can establish their identity to-day. In the past a number of taxonomic works have been written, dealing with larger groups of parasitic Hymenoptera and based mainly on a compilation of these insufficiently described species, which were often not fully understood by the compilators themselves. Some examples of these works are Schmiedeknecht’s Opuscula Ichneumonologica (1902—1936), Fahringer’s Opuscula Braconologica (1928—1937), Kief- fer’s work in “Das Tierreich” on Proctotrupoi- dea (1914—1926) and that of Dalla Torre & Kieffer on Cynipoidea (1910), also in “Das Tierreich”. It is not our intention to criticize these works. They present excellent surveys of the knowledge of the several species at that time, and any specialist who occupies himself with these groups now, has to consult them. However, they do not satisfy the needs of a modern entomologist who wants to know the reliable names and taxonomical status of the species which he encounters in his ecological in- vestigations. Unreliable species names of parasi- tic Hymenoptera show up in reviewing jour- nals, e.g., The Review of Applied Entomology, and from these they are compiled in compre- hensive works like Thompson’s host-parasite lists. In view of the difficulties discussed above, it cannot be expected that the gap in our taxo- nomic knowledge of the species of parasitic Hymenoptera will be filled up within a reasona- ble time. Fortunately in the last decades an in- creasing interest in the taxonomy of parasitic Hymenoptera has arisen, and many groups have now been or will soon be revised. Ecologists have to rely on identifications by specialists. Only these specialists are thoroughly acquainted with the taxonomic difficulties and their nomenclatorial consequences. Only a spe- cialist in a certain not too large taxonomic group can really judge in how far his identifica- tions are reliable. It might be desirable that a specialist, in iden- tifying material on behalf of ecological investi- gations, should add some more information than merely the name. The ecologist should also ‚be informed about the reliability of the species name, which can only be judged by the special- ist. ACKNOWLEDGEMENTS We wish to thank the following specialists in the various taxonomic groups of parasitic Hymenoptera: Dr. Ing. C. van Achterberg, the Netherlands, Dr. J. P. Aeschlimann, France, Mr. M. J. Gijswijt, the Netherlands, Dr. K. Horstmann, Bundesrepublik Deutschland, Dr. W. R. M. Mason, Canada, Dr. G. E. J. Nixon, United Kingdom, Mr. G. van Rossem, the Netherlands, Prof. Dr. W. Schwenke, Bundes- republik Deutschland, and Drs. K. W. R. Zwart, the Netherlands. Without their help with the identifications and their advice we could not have completed these investigations. Thanks are also due to Prof. Dr. J. T. Wiebes for critically reading the manuscript, Dr. P. Gruys, Drs. D. J. de Jong and Ir. P. M. J. Ra- makers, all from the Netherlands, and Dr. A. Schmid, Switzerland, for providing material, to Ir. A. B. A. M. Baudoin and Ir. E. Neering, both students of the Agricultural University at Wageningen at that time, who investigated the parasites of leafrollers on several foodplants in habitats outside apple orchards, and to Mr. C. van der Horst, IPO, Wageningen, for revising the English text. REFERENCES Achterberg, C. van, 1974. The features of the petiolar segment in some Braconidae. — Ent. Ber., Amst. 34:213— 214. ——, 1976. A preliminary key to the subfamilies of the Braconidae (Hymenoptera). — Tijdschr. Ent. 119: 33—78. | ——, 1982. Familietabel van de Hymenoptera in Noordwest Europa. — Wet. Meded. K. Ned. na- tuurh. Veren. 152: 1—50. Aeschlimann, J. P., 1973. Révision des espèces ouest- paléarctiques du genre Tviclistus Foerster (Hymenoptera: Ichneumonidae). — Mitt. schweiz. ent. Ges. 46: 219— 252. Ankersmit, G. W., 1968. 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Ent. 50: 93— 98. pathogens re f . un: ER rar î gn Fu “ i War De { AN pu Pi Les george pd went ï Ds SIENTE bali i i 4 ‘ wins À 2 Wes 4 wiet, rer } ke à A Ar y Ra N | u t È ii Zr a } fi ; i FO | RARE AU dt CL EE APN { dandy È pa shot à Te i ni Ù a t À hu hi it . i n 4 y \ \ À 7 4 u NU f f ‘ Pe hi 4 i aye RON ! ’ au j t CAES EME A f p i ; N 4 S \ ‘ ere : | it 1 di Ikea \ ( Î { 1 . ) I vid ‘ Ù 4 4 : t [i sint va, 92 AS DEEL 126 AFLEVERING 7—8 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING MUS: COMP. ZOOL LIBRARY ANO 0 1984 HARVARD UNIVERSITY INHOUD J. BELLE. — On the species of the polygonus group of Progomphus with a descrip- tion of a new species (Odonata, Gomphidae), pp. 137—144, figs. 1-23. J. BELLE. — A review of the genus Zonophora Selys (Odonata, Gomphidae), pp. 145—173, figs. 1—76. | Tijdschrift voor Entomologie, deel 126, afl. 7—8 Gepubliceerd 15-X11-1983 i re, De AE NE à CAR SLA RO EMO CRE WE) UT 2 ‘ I { à "A { HAT î 4 y ri “5 t be Malt en \ I Li N | > : » PA ON THE SPECIES OF THE POLYGONUS GROUP OF PROGOMPHUS WITH A DESCRIPTION OF A NEW SPECIES (ODONATA, GOMPHIDAE) by JEAN BELLE Onder de Beumkes 35, Velp, The Netherlands ABSTRACT P. occidentalis spec. nov. (& holotype: San Antonio, Venezuela?) is described and eluci- dated by figures. The right pair of wings of P. polygonus Selys is depicted. A distribution map of P. phyllochromus Ris is given. The male of P. abbreviatus Belle is redescribed and its female is described for the first time. The four species mentioned here are forming an infra- generic group. INTRODUCTION Since my revision of the genus Progomphus Selys in 1973, five new species have been de- scribed, two from Brazil (P. victor St. Quenun, 1973, and P. perithemoides Belle, 1980), one from Paraguay (P. flinti Belle, 1975), one from the USA (P. belle: Knopf & Tennessen, 1980) and one from Venezuela (P. racenisi De Mar- mels, 1983). In the present account a new spe- cies, P. occidentalis, is described from continen- tal South America. The new descriptions show that the genus Progomphus is very rich in spe- cies, and that we still have not come to the end of our discoveries. Together with P. polygonus Selys, P. phyl- lochromus Ris and P. abbreviatus Belle, the new species mentioned above belongs to an infrage- neric group, the polygonus group. The members of this group correspond in the occurrence of an extra cubito-anal cross-vein in the wings (occa- sionally present in P. abbreviatus), the more or less broken distal side of the fore wing discoidal triangle, the medially excised posterior margin of the occipital plate, the wedge-shaped first pale antehumeral stripes, and the lack of a sec- ond pale antehumeral stripe immediately in front of the humeral sutures, while the abdomen of the females is about as long as the hind wings. The new material of the polygonus group en- ables me to publish additional data on its known members. A figure of the right pair of wings of the female lectotype of P. polygonus is given. A distribution map of P. phyllochromus has been made with the aid of old and new records. The male of P. abbreviatus, known only from a sin- 137 gle male in poor condition, is redescribed from fully mature specimens; and the female of this species is described for the first time. The material on which the present paper is based belongs to the institutions and personal collections mentioned below; the names are preceded by the abbreviations used throughout the text and followed by the names of the per- sons from whom material was received on loan or as a gift. These persons are most gratefully acknowledged here. ANSP — Academy of Natural Sciences of Philadelphia, Philadelphia; Dr. Dave Rentz. — Florida State Collection of Ar- thropods, Gainesville; Prof. Dr. Minter J. Westfall, Jr. Instituto Miguel Lillo, San Miguel de Tucumán; Dr. A. Willink. Institut Royal des Sciences Natu- relles de Belgique, Bruxelles; Dr. Georges Demoulin. Instituto de Zoología Agrícola, Maracay; Dr. F. Fernández Yépez and Mr. Jorge de Marmels. Rijksmuseum van Natuurlijke Historie, Leiden; Dr. P. H. van Doesburg. Muséum National d'Histoire Na- turelle, Paris; Dr. Jean Legrand. Natur-Museum Senckenberg, Frankfurt am Main; Dr. Heinz Schröder. FSC SMF 138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 7 (1983) Progomphus polygonus Selys, 1878 (fig. 23) This species was described after two very ten- eral females from Merida, Venezuela. No addi- tional material is available. In 1973 a lectotype was designated by me. In the present paper a depiction of the right pair of wings of the lecto- type is published. P. polygonus is the largest representative of the polygonus group. The reticulation of its wings is denser than that of the other members. The trigonal interspace in the fore wings of the lectotype starts with three rows of cells from the triangle outwards, three cells long. The dis- tance between the nodus and the pterostigma is a little more than 2.75 times the length of the pterostigma. The abdomen of the female is rela- tively shorter than that of the females of the other members of the group. But the two fe- males of P. polygonus are very teneral and fully mature females may have a relatively longer ab- domen. Progomphus phyllochromus Ris, 1918 (figs. 1—4, 6) Material. — Argentina: Catamarca, Concep- Con 151960 MINS MN SES C Salt Or En 24 km of Agua Blancos), 9.v.1969, 1 4, IML, all A. Willink leg. — Colombia: 1 2 (ex coll. R. Mar- tin), MP. — Venezuela: Aragua, Cumbre de Choroní, 13.vin.1957, 1 © (author’s collection), 2, J, IkAceas legs Miane, IPerane, 26.x1.1961, 1 d, Bordon leg., IZM. This mountain species is chiefly Andean in its distribution and it occurs from the north coast of Venezuela to Catamarca in Argentina (dis- tance between the two localities almost 6000 km). It has been recorded from Argentina, Bo- livia, Colombia, Ecuador, Peru and Venezuela. In drawing up the distribution map (fig. 6) the present localities as well as those published ear- lier (Ris, 1918; Belle, 1973) were used. The male from Salta is small and its wings have a less dense reticulation. The mea- surements are: total length, 35.5 mm; abdomen, 26 mm; hind wing, 23 mm; costal edge of pte- rostigma in fore wing, 3.3 mm. The fore wings have an open subtriangle, a single row of cells in the anal field, and no extra cubito-anal cross- vein. The frons is higher than is normally the case, and there is a weak development of a fron- tal ridge. The two wedge-shaped, (first) pale an- tehumeral stripes are rather narrow and they are almost parallel-sided on the dorsal half. The female from Colombia has no extra cubi- to-anal cross-vein in the wings. The larva of Progomphus, described by Needham (1941) as No. 14, most likely belongs to P. phyllochromus. The venational characters found in the pair of wings of this larva are in agreement with those of the adults of P. phyl- lochromus, and its place of capture (Estado Mi- randa) lies within the range of the species. Needham’s larva No. 14 differs from that of P. abbreviatus described by De Marmels (1981b) in having the front margin of the middle lobe of. the labium armed with a single median tooth instead of one submedian pair of such teeth (Needham, 1941: 240). Progomphus abbreviatus Belle, 1973 (figs. 7—17) Material. — Colombia: Dept. Magdalena, Sierra San Lorenzo (4500-5600 ft), Hacienda Cincinatti, 20.vu.1920, 1 d, ANSP; Prima, 1.vi.1965, 1 d, J. & B. Bechyne leg, IZM; Sier- ra de Perija (1350), Mission Finca Marganta (6 km SW of Socorpa, rocky stream, half shaded), 7—8.viu.1968, 1 3d, Borys Malkin leg., ML. — Ecuador: 1 &, MP. — Venezuela: Miranda, Se- bucán, 16.x.1955, 3 d (author’s collection), 3 & all Klisans leg.; Miranda, El Marqués, Quebrada Pasaquire, Avila (1000 m), 10.x.1980, 1 2, J. de Marmels leg., IZM. This species was described after a single male from Colombia. The type is in poor condition, very teneral and flattened for its entire length. Due to this bad condition some features are misinterpreted or insufficiently described in the original description. Therefore descriptive notes of the present fully mature males are given be- low. The female is also described. Male. — Total length, 39—41 mm; abdomen, 29.5—30.5 (incl. app.); hind wing, 23—25.5 mm; costal edge of pterostigma in fore wing, JOS) Maan, Labrum with grey-green band along free bor- der, the width of the band very narrow in one male but very broad in another male. Antecly- peus, lateral sides of postclypeus and bases of mandibles externally grey-green. Frons low, without anterior ridge, its superior surface largely grey-green but dark brown at base. Ver- tex and occipital plate dark brown. Postocellar ridges well-developed. Mid-dorsal width of oc- Berre: The polygonus group of Progomphus cipital plate a quarter the length of frontal mar- gin of occipital plate. Posterior margin of occip- ital plate concave and with a shallow median ex- cision which is very small and V-shaped in some males. Crest of occiput with very long brown hairs. Rear of head brown but tempora with a pale green spot and a (lower) pale green band. Labrum and adjacent mouth parts pale green. Prothorax dark brown, the hind lobe blackish brown. Dark colour of pterothorax blackish brown. Pale antehumeral stripes grey-green, very broad in male from Ecuador. Three pale lateral stripes of pterothorax olive-green. Legs brown but inner sides of femora green. Third tarsi three-quarters the length of third tibia. Abdomen dark brown, becoming blackish brown on apical segments. Sides of 1 entirely green. Sides of 2 largely green. Genital hamules and hood of penial peduncle brown. Sides of 3 with a long, triangular, green basal spot. Seg- ments 2 to 7 with a pale mid-dorsal line which is rather wide and green on 2, narrow but green at base and becoming yellow at apex on 3, very fine and yellow on 4 to 7. Dorsum of 7 yellow on basal third. Caudal appendages black but up- per surface of superiors pale on apical two- fifths. Inner process of branches of inferior ap- pendage variable, generally reduced to a low hump but in some males entirely lacking; length of branch beyond this inner process (if avail- able) also variable. Venation of wings dark brown but costa with a very fine and inconspicuous yellow line. Pte- rostigma yellowish brown. Two cubito-anal cross-veins in addition to inner side of sub- triangle in seven fore wings; other wings with- out extra cubito-anal cross-veins. Discoidal tri- angle open in one fore wing, two-celled in other wings. Subtriangle three-celled in two fore wings and two hind wings, two-celled in other wings. Supratriangle (once) crossed in two hind wings, open in other wings. Trigonal inter- spaces in fore wings starting with two rows of cells from triangle outwards, those in hind wings starting with a row of two (one male) or three (other males) cells against triangle fol- lowed by two rows of cells. Anal field in five fore wings one cell wide, in other fore wings two cells wide. Second anal interspace in three hind wings starting with a single large cell against anal vein, in other hind wings with two cells against anal vein. Female (hitherto unknown; degutted and treated with aceton). — Total length, 37.5 mm; 139 abdomen, 28 mm (incl. app.); hind wing, 26.5 mm; greatest width of hind wing, 8.2 mm; cos- tal edge of pterostigma in fore wing, 3.7 mm. Similar to male regarding stature and colour design but abdomen stouter. Grey-green ante- rior band on superior surface of frons narrower than in male, in dorsal view occupying one- third of superior surface of frons. Pale markings on pterothorax greenish white, on abdomen yellowish white. Second pale antehumeral stripes somewhat narrower than in male. Pale markings on abdomen as follows: A fine mid- dorsal line on 2 to 8; lateral stripes becoming successively shorter on 4 to 6, reaching to apex of segment on 2 and 3, and to a point half way the segment on 6, the stripes being interrupted by the supplementary transverse carinae; dor- sum of 7 pale from base to supplementary trans- verse carinae. Abdominal segments 7, 8, 9, and 10 approximately in ratio 46 : 26 : 19 : 10, with the caudal appendages (stylets) 15 on the same scale. Vulvar lamina one-third the length of ninth sternum, its posterior margin medially ex- cised U-shaped for three-quarters the length of vulvar lamina. Caudal appendages white at tips and spine-like pointed. Basal subcostal cross-vein present. No extra cubito-anal cross-vein. Anal field in fore wings two cells wide. Nodal index 14 : 17—18 : 12/ 13: 13—14: 13. Second primary antenodal cross-vein the sixth in right fore wing, the fifth in other wings. Intermedian cross-veins 6—6/ 4—4 in fore and hind wings, respectively. Su- pratriangles open. Subtriangle in right fore wing two-celled, in other wings three-celled. Discoi- dal triangles two-celled. Trigonal interspaces starting with two (right fore wing) or three (other wings) cells against triangle followed by two rows of cells. Hind wings with five paranal cells, four (left) and five (right) postanal cells, and four to five rows of cells posterior to Cu2. The female of P. abbreviatus resembles that of P. phyllochromus but the latter has the frons distinctly angled, the occipital plate much broader (mid-dorsal width two-fifths the length of frontal margin), and the fore wings generally with extra cubito-anal cross-veins. Mr. De Marmels wrote me that several more females were seen and photographed at the place where the female described was captured (cf. De Marmels, 1981a: 40). The following ve- national characters could be determined from the colour slides: 9 (1—15.v111.1979), number of antenodal cross-veins in left fore wing 16 and 140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 7 (1983) Figs. 1-4. Progomphus phyllochromus Ris, & holotype: 1, tenth abdominal segment and caudal appendages, dorsal view; 2, the same, left lateral view; 3, the same, ventral view; 4, diagram of thoracic colour pattern. Fig. 5. Progomphus occidentalis spec. nov, 3 holotype: Diagram of thoracic colour pattern. in left hind wing 12, discoidal triangles and sub- triangles in left wings two-celled; ® (vili-ix. 1979), nodal index of left pair of wings 11 : 14/ 12 : 11. A female collected by him at the same place on 25.1x.1981 has the following features: total length, 37.5 mm; abdomen, 27.5 mm; hind wing, 25 mm; costal edge of pterostigma in fore wing, 3.8 mm. Sternum of abdominal segment 10 not shorter than sternum of abdominal seg- ment 9. All triangles and subtriangles two- celled. Trigonal interspace starting with two cells against triangle in fore wings, with three cells in hind wings. Nodal index 11 : 16—17 : 12/11 : 12-12 : 12. Second primary antenodal cross-vein the fifth in all wings (all data commu- nicated by Mr. De Marmels). The larva of P. abbreviatus was described by De Marmels (1981b). Progomphus occidentalis spec. nov. (figs. 5, 18—22) Material. — Venezuela (?): San Antonio, 1 d (holotype), MP. Male (abdomen broken between segments 5 and 6). — Total length, 43 mm; abdomen, 33.5 mm; hind wing, 26.5 mm; greatest width of hind wing, 8 mm; costal edge of pterostigma of fore wing, 3.6 mm. Face brown. Frons low and slightly angled, its superior surface with a broad, leaden-grey anterior band. Vertex dark brown, the posterior ridge of lateral ocelli swollen and provided with long brown hairs. Occipital plate dark brown, its posterior margin concave in middle portion, provided with long brown hairs. Rear of head brown above. Tempora with a brown-yellow BELLE: The polygonus group of Progomphus 141 + Fig. 6. Distribution of Progomphus phyllochromus Ris. spot. Lower parts of rear of head brown-yel- low. Labium and adjacent mouth-parts pale brown. Prothorax dark brown. Pterothorax dark brown with green stripes, its colour design shaped as shown in diagram fig. 5. Legs brown but inner side of first femora green. Third tibia three-quarters the length of third femur. Lamina tibialis of first tibiae one- fifth the tibial length. Wings clear. Venation brown but frontal side of costa with a fine yellow line. Pterostigma brownish yellow, surmounting 5'2—6 cells. Distance between nodus and pterostigma 22 times the length of pterostigma. Basal subcostal cross-vein present. Nodal index 11: 15— 16 : 10/11 : 11—12: 11. Second primary ante- nodal cross-vein the fifth. Intermedian cross- veins 6—5/5—5 in fore and hind wings, re- spectively. All supratriangles open. All subtri- 142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 7 (1983) 13 15 16 Figs. 7—17. Progomphus abbreviatus Belle: 7, tenth abdominal segment and caudal appendages of male, dorsal view; 8, the same, left lateral view; 9, the same, ventral view; 10, occipital plate of female; 11, vulvar lamina; 12, accessory genitalia, right lateral view; 13, frons of male, dorsal view; 14, ventral view of right branch of inferior caudal appendage, male from Miranda; 15, the same, another male from Miranda; 16, the same, male from Sierra San Lorenzo; 17, accessory genitalia, ventral view. angles and discoidal triangles two-celled. There is an extra cubito-anal cross-vein in each wing. Trigonal interspace in fore wings starting with two rows of cells from triangle outwards (that of left fore wing with an extra initial cell next hind angle of triangle) eight cells long, in ante- rior row, that in hind wings starting with three rows of cells against triangle followed by two rows of cells two (right) and three (left) cells long, in anterior row. Anal field in right fore wing one cell wide, that in left fore wing with one of the cells doubled. Hind wings with five (right) and six (left) paranal cells, four rows of cells behind Cu2 and a three-celled anal trian- gle. Abdomen predominantly dark brown. Mid- dorsum of segments 2 to 7 with a yellow line which is rather broad on segment 2 but which becomes very fine on segments 5 to 7. Sides of segment 2 with yellow marking behind and be- low auricle. Lower parts of sides of segments 3 to 6 with yellow marking from base to apex of segment. Basal third portion of segment 7 yel- low. Caudal appendages shaped as shown in ac- companying figures, blackish brown, the supe- riors becoming pale on apical half. The unique (pinned) specimen of P. occiden- talis was found in the collection of René Martin at Paris. Attached to the pin is an old label “Pro- gomphus polygonus Selys S. Antonio” but there is no label referring to the country in which the locality San Antonio lies. However, from its relationship with the other members of the polygonus group we can say that the specimen hails from western continental South America, hence the specific name occidentalis. But René Martin referred the male to P. polygonus, a spe- cies known only from Venezuela. I therefore suspect that the specimen is from the west of BELLE: The polygonus group of Progomphus 143 Figs. 18—22. Progomphus occidentalis spec. nov., 6 holotype: 18, tenth abdominal segment and caudal append- ages, dorsal view; 19, the same, left lateral view; 20, the same, ventral view; 21, occipital plate; 22, accessory genitalia, ventral view. Fig. 23. Progomphus polygonus Selys, 2 lectotype: right pair of wings. this country (State of Táchira) where a village San Antonio lies. P. occidentalis is intermediate between P. phyllochromus and P. abbreviatus but it is more closely related to the first than to the second species. The males of P. occidentalis and P. phyl- lochromus are distinguished by the following features: (1) apex of branches (part beyond an- teapical inner process) of inferior caudal ap- pendage longer in P. occidentalis than in P. phyllochromus; (2) inner process of branches of inferior caudal appendage acute and ending in two minute teeth in P. occidentalis, truncated and furnished with a marginal row of minute teeth in P. phyllochromus; (3) lateral outer side of superior caudal appendages between basal di- latation and proximal denticle of inferior carina not emarginated in P. occidentalis, widely emar- ginated in P. phyllochromus; (4) up of posterior hamule of accessory genitalia very acutely pointed and strongly recurved in P. occidentalis, and longer than that in P. phyllochromus. REFERENCES Belle, J., 1973. A revision of the New World genus Progomphus Selys, 1854 (Anisoptera: Gomphi- dae). — Odonatologica 2: 191—308. , 1975. Two new gomphine species from Para- guay (Anisoptera: Gomphidae). — Odonatologica 4:129= 135; 144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 7 (1983) , 1980. Two new gomphoidine species from Bra- zil (Anisoptera: Gomphidae). — Odonatologica 9:173—180. De Marmels, J., 1981a. Aeshna rufipes Ris in Venezu- ela, with a description of the male (Aniso- ptera: Aeshnidae). — Odonatologica 10: 39—42. ——, 1981b. The larva of Progomphus abbreviatus Belle, 1973 from Venezuela (Anisoptera: Gom- phidae). — Odonatologica 10: 147—149. ——, 1983. The Odonata of the region of Mount Auyantepui and the Sierra de Lema, in Venezue- lan Guyana. 3. Additions to the families Gomphi- dae, Aeshnidae and Corduliidae, with description of Progomphus racenisi spec. nov. — Odonato- logica 12: 5—13. Knopf, K. W. & K. J. Tennessen, 1980. A new species of Progomphus Selys, 1854 from North America (Anisoptera: Gomphidae). — Odonatologica 9: 247—252. Needham, J. G., 1941. Life history studies on Pro- gomphus and its nearest allies (Odonata: Aeshni- dae). — Trans. Am. ent. Soc. 67: 221—245; pl. 20. Ris, F., 1918. Libellen (Odonata) aus der Region der amerikanischen Kordilleren von Costarica bis Catamarca. — Arch. Naturg. 82 (9): 1—197; 2 tabs. Selys Longchamps, E. de, 1878. Quatriemes additions au synopsis des Gomphines. — Bull. Acad. r. Belg, (2) 46: 408471, 658698 (3—106 sep.). St. Quentin, D. 1973. Die Gomphidenfauna Südame- rikas (Ordn.: Odonata). — Annln naturh. Mus. Wien 77: 335—363. A REVIEW OF THE GENUS ZONOPHORA SELYS (ODONATA, GOMPHIDAE) by TEAN BELLE Onder de Beumkes 35, Velp, The Netherlands ABSTRACT The genus Zonophora Selys is revised. A number of problems in the classification of its members are solved and elucidated by figures. A new subfamily, the Zonophorinae, is in- troduced. Z. diversa spec. nov. (6 holotype: Nova Teutonia, Santa Catarina, Brazil), Z. campanulata annulata subspec. nov. (& holotype, ® allotype: Jataí, Goiás, Brazil) and Z. nobilis spec. nov. (6 holotype: Taracua, Amazonas, Brazil) are described. Z. machadoi St. Quentin and Z. obscura Belle are considered to be subspecies of Z. campanulata (Burmeis- ter) and Z. solitaria Rácenis, respectively. The females of these subspecies are also de- scribed. A neotype for the type-species of the genus and lectotypes for Z. supratriangularis Schmidt and Z. wucherpfennigi Schmidt are selected. Illustrations of some structures of the accessory genitalia of Z. surinamensis Needham are given. A key to the species and subspe- cies is constructed. New distributional records and distribution maps are added. CONTENTS PAGE Imre erde rd oee gelo. 145 Material and acknowledgements.............. 146 HOOD TONAR EVSA BDA. un... xi uae eye un 146 Disposition of species and infrageneric groups .. 148 SNSEEMAHLGSECHON NN en ns ene de ee 149 ISENALOREEN SUD SPECIES Anne ee ae 149 Wireattmentonthe species’ to. een 150 c. campanulata (Burmeister) ............... 151 Conachado!St Quentine itse ett 152 ciuamulatasubspec'inovi: i. vino edo: 152 HD ASP ECAMOVE REA). ns Hire dde 153 ©, CADDIE SON RN N ea: 154 GEspeetabi il CAMPION. 21. „u. me rien 155 & (ange Sec SSSR 155 EBSD Sets Ree MER 156 DG LAD CIMICISCAMIA MERE ee 156 SumnamenssiNeedhamt eni MN CT 156 nicola ANacenist tE Men. DAAMEN Jed. Bees 157 STIME ee a 157 FORTS ROUE CORRE cae tee ER eee okee 158 supratriangularis Schmidt ................. 158 ZODINSSBEEANOVERL AAR lee ie ahi 159 The species in the mimeographed list of Rácenis 159 Habitat and behaviour in the field ............ 160 iScoctapiic distribution, e enne 160 LOREM CES AEP E A IO ct 161 INTRODUCTION The genus Zonophora was created for a cer- tain Neotropical gomphid by Selys in 1854. Into this genus, he placed Diastatomma campanula- ta, described on the basis of a single male from Brazil by Burmeister (1839). No further addi- tions were made to the genus for fifteen years, when Selys (1869) published descriptions of two new species (batesi and calippus). Fifty-one years later, Campion (1920) described spectabi- lis. In 1941, Schmidt described two species (supratriangularis and wucherpfennigi) and one subspecies (klugi as subspecies of calippus). Needham (1944) described one species (surina- mensis), St. Quentin (1973) one species (macha- doi), Racenis (1970) one species (solitaria), and Belle (1976) two species (regalis and obscura). One species referred to Zonophora (Z. rokitanskyi St. Quentin, 1973) does not fall within the limits of this genus and was assigned to Diaphlebia Selys, 1854 (Belle, 1977a). Here I report on material of two new species (diversa and nobilis) and one new subspecies (campanu- lata annulata), bringing the total number ot taxa to fifteen. Schmidt (1941) treated spectabilis and klugı as subspecies but Belle (1963) elevated them to the rank of species. A restudy of these representa- tives, however, has revealed that there is little reason for retaining this status and that Schmidt’s classification should be followed. St. Quentin (1973) considered wucherpfennigi as a subspecies of batesi but here wucherpfennigi is treated as species, following again Schmidt. On the contrary, St. Quentin’s machadoi and Belle's obscura are considered subspecies of 145 146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) campanulata and solitaria, respectively. It is clear that in the interests of a stable nomencla- ture a neotype designation for the type-species of the genus is required since the holotype of Burmeister’s campanulata must be considered lost. The knowledge of the proventriculus of larval Zonophora has led to the introduction of the subfamily Zonophorinae, to contain the genus Zonophora. The diverse specific characters show that the members of the genus Zonophora can be conveniently divided into five groups. The critical structures on which the present classification is based are elucidated by original camera lucida drawings. Distributional maps tor the groups are added and a key to the species and subspecies is constructed. I take the oppor- tunity to publish illustrations of some important structures of the accessory genitalia of Z. suri- namensis. MATERIAL AND ACKNOWLEDGEMENTS The material on which the present review is based belongs to the institutions and personal collections mentioned below; the names are preceded by the abbreviations used throughout the text of this paper. AC — Author’s collection; BM — British Museum (Natural Histo- ry), London; Dr. Peter H. Ward; CC — Collection Cook, Center; Mr. Carl Cook; CJ — Collection Jurzitza, Karlsruhe; Prof. Dr. Gerhard Jurzitza; CM — Collection Machado, Belo Hori- zonte; Prof. Dr. Angelo B. M. Machado CU — Cornell University, Ithaca; Dr. L. L. Pechuman; IRSN — Institut Royal des Sciences Natu- relles de Belgique, Bruxelles; Dr. Georges Demoulin; IZM — Instituto de Zoologia Agricola, Nara cavi AND EA crnandez Wepez: ML — Rijksmuseum van Natuurlijke Historie, Leiden; Dr. P. H. van Doesburg; MP — Muséum National d'Histoire Na- turelle, Paris; Dr. Jean Legrand; UMMZ — Museum of Zoology, University of Michigan, Ann Arbor; Mrs. Leonora K. Gloyd and Dr. T. E. Moore; SMF — Natur-Museum Senckenberg, Frankfurt-am-Main; Dr. Heinz Schröder. For the loan of material I am also indebted to the following persons: Mr. Jorge de Marmels (Caracas) and Dr. Janis Rácenis (f) (Caracas). Zonophora Selys, 1854 Zonophora Selys, 1854: 80 (61 sep.); 1873: 517 (73 sep.). Selys & Hagen, 1858: 493—494 (233— 234 sep.). Kirby, 1890: 75. Karsch, 1890: 371. Car- pentier, 1897: 453. Navás, 1916: 28. Campion, 1920: 139—140. Williamson, 1920: 9, 11—12. Kennedy, 1928: 370. Tillyard & Fraser, 1938— 1940: 372. Byers, 1939: 21. Fraser, 1940: 549; 1957: 93. Needham, 1940: 388— 389; 1944: 172— 173, 215, 218, 221—222. Schmidt, 1941: 76—78. Belle, 1963: 60, 62—63; 1976: 197, 200, 206; 1979: 46: St. Quentin, 1967505 01078 8855 337—338, 341, 361. Paulson, 1977: 179. Kiauta, 1979: 268. Davies, 1981: 33. Type species. — The type-species of the ge- nus Zonophora is Diastatomma campanulata Burmeister, 1839, as designated by Kirby (1890). Position. — The systematic position of Zono- phora, based on a certain combination of vena- tional characters, has been changed several times. Williamson (1920) placed Zonophora in his “Zonophora series”, a series which only contained this genus. Zonophora was character- ized by the discoidal triangles and subtriangles of the fore and hind wings being similar, the dis- coidal triangles long and once crossed, the sub- triangles and supratriangles all free, absence of trigonal supplements and sectors of Rs and M4, and by the nearly right proximal angle at the point of separation of Al from Cu2 + Al and not as obtuse as the distal angle at the same point. However, the supratriangles are normally crossed in two species and incidentally crossed in some other species. In the classification as proposed by Tillyard and Fraser (1938— 1940), Zonophora becomes one of the members of the Ictinogomphinae, distinguished by their posi- tion of crossed discoidal, supra- and subtrian- gles. However, Zonophora has the subtriangles never crossed and the supratriangles of most species generally not crossed. In the revised re- classification, Fraser (1957) transferred Zono- phora into the Gomphoidinae, which were sep- arated from the other genera by the presence of crossed discoidal, supra- and subtriangles, and in having only a single cross-vein (Ac) in the cu- bital space. Zonophora, however, has uncrossed subtriangles and generally uncrossed supra- BELLE: Zonophora 147 triangles while in some species there are nor- mally two or three cubito-anal cross-veins in addition to the inner side of the subtriangle (Belle, 1963, tabs. 3, 4). Belle (1979), finally, placed Zonophora in the Epigomphinae, which were characterized by the uncrossed subtrian- gles in the fore wing, the number of intermedian cross-veins in the hind wing being two or more, and the absence of a trigonal supplement. How- ever, the aberrant proventricular armature of larval Zonophora (cf. below) shows the isolated position of this genus in the Gomphidae and justifies the erection of a sixth subfamily, the Zonophorinae, to contain Zonophora. The pro- ventricular character together with the vena- tional character for the hind wings cited above (proximal angle at point of separation of Al from Cu2+A1 nearly a right angle, and not as obtuse as distal angle at same point) separate Zonophora from the rest of the Epigomphinae in which it was lastly placed. Immature stages. — In 1944, Needham gave a description of a larval cast-off skin, which he re- ferred to Zonophora by a process of exclusion. In 1966, Belle published photographs of the ex- uviae of Z. batesi and Z. calippus, while he gave a brief description of the exuviae of the latter species. The reference of this exuvial material to these two species was based on the difference in size and on the fact that the two species were the only regional ones of the genus occurring in the well-explored surroundings of the airport Zanderij of Surinam. Some half-grown larvae of Zonophora were also secured by sifting mud or silt removed from the bottom and sloping banks of the creeks crossing the gallery forests in the surroundings of Zanderij. The gizzard of these larvae was recently examined by me. The result was most surprising as the proventriculus ex- hibited an unexpected specialized armature, es- sentially different from that hitherto found in the Gomphidae. In figs. 1—5 the complete proventriculus of a half-grown larva of Z. batesi is shown. Con- trary to what is normally found in the Gomphi- dae, there are two pairs of dissimilar, elevated, sclerotized areas (dental folds) in juxtaposition, one dorsal pair consisting of pyramid-shaped folds and one ventral pair consisting of molar- shaped folds. Each of the folds has a specialized dentition of large, posteriorly directed teeth and, on the posterior end, numerous minute posteriorly directed teeth. In shape, the dental folds of Zonophora much resemble those of Aphylla (cf. Beatty, 1956), but the occurrence of numerous minute teeth on these folds seems to be confined to Zonophora (and some Aeshnidae and Lestidae, cf. Bullens, 1966). In the Gomphidae, this aberrant proven- tricular armature apparently can be employed as a taxonomic character. Characterization. — The genus Zonophora is characterized by the following combination of features: In both sexes: (1) Numerous hind wings. (2) No trigonal supplements and sectors of Rs and M4. (3) Vein Al in hind wings well-developed, the proximal angle at the point of separation of A1 from fused portion Al + Cu? a right angle or nearly so, the distal angle at same point ob- tuse. (4) Discoidal triangles and subtriangles of fore wings more or less similar to those of hind wings; the discoidal triangles elongated in axis of wing and generally once crossed (occasional- ly two cross-veins); the subtriangles uncrossed. (5) Pterostigma in hind wings somewhat lon- ger than that in fore wings. (6) Meso- and meta-paraptera conspicuously produced in an acute point. (7) Vertex with conical process behind each lateral ocellus. (8) Occiput angled at rear. (9) Ventral tergal margins of abdominal seg- ments 8 and 9 not expanded. In the male: (10) Anal triangle in hind wings not reaching backward to anal angle of wing. (11) Superior caudal appendages twice to three times as long as abdominal segment 10 and incurved at tip; inferior caudal appendage about as long as abdominal segment 10 and with two branches widely spread apart. In the female: (12) Vulvar lamina reaching from a point one-third the way along ninth sternum to apex of ninth sternum; its posterior margin medially excised. In the larva: (13) No dorsal hooks on abdomen. (14) Abdomen with lateral spines only on ab- dominal segments 8 and 9. (15) Caudal appendages about three-and- one-half times as long as abdominal segment 10. (16) Lateral lobes of labium with incurving teeth along inner margin and with a strong end hook. intermedian cross-veins in 148 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) (17) Gizzard with two pairs of differentiated dental folds bearing undifferentiated large teeth and numerous minute teeth. DISPOSITION OF SPECIES AND INFRAGENERIC GROUPS (figs. 6— 30) There are excellent characters available for se- parating the species and subspecies of the genus Zonophora into five groups. The members of each group exhibit a striking resemblance in the shape of some structures although some of the statements concerning the shape of these struc- tures will have to be taken with due allowance for individual variation. (1) The campanulata group, comprising Z. campanulata campanulata (Burmeister) with its subspecies Z. campanulata machadoi St. Quen- tin and Z. campanulata annulata subspec. nov, and Z. diversa spec. nov. The distal segment of the penis has a pair of short and inconspicuous tusk-like spines. The anterior hamules of the male have a well-devel- oped, finger-like, internal lobe. The vulvar lami- na is more or less parallel-sided and long, reach- ing to the apex of the ninth sternum; the tip of the vulvar lamina exhibits much variation, the divisions being sometimes widely spread out (cf. Schmidt, 1941). The wings have no basal subcostal cross-vein and no extra cubito-anal cross-vein. There is, normally, no traversing cross-vein in the supra- triangles (cf. Belle, 1963, Verification tables). The hind wings have sometimes an open anal loop, i.e. open at the proximal side (cf. Schmidt, 1941), but generally an ill-defined anal loop of three to five cells. There are no distinct morphological differ- ences between Z. campanulata and its subspe- cies. The distinction is based only on colour dif- ferences. Z. c. campanulata is the darkest of the three. The pale markings on the sides of the pterothorax are grey-green. The abdomen is predominantly dark brown. The tergite of the segments 3 to 8 is densely set with minute black denticles, in particular on the basal part of the segments. The two subspecies Z. c. machadoı and Z. c. annulata have distinct greenish yellow and yellow markings which are extensive and very striking on the abdomen. The abdominal segments 3 to 8 of the two subspecies are scanti- ly set with minute black denticles. Z. diversa does exhibit great colour and mor- phological differences with Z. campanulata and its subspecies. The superior lobe of the anterior hamules of the male is vestigial in diversa, well- developed in campanulata. (2) The calippus group, comprising Z. calip- pus calippus Selys, Z. calippus spectabilis Camp- ion and Z. calippus klugi Schmidt. The distal segment of the penis has a pair of short flagella. The anterior hamules of the male are much smaller than those of the campanulata group; the internal lobe is small and the superi- or lobe is long and slender. The vulvar lamina is smaller and more slender than that of the cam- panulata group, and it exhibits but little varia- tion. The wings have, normally, a basal subcostal cross-vein and generally extra cubito-anal cross-veins. The anal loop in the hind wings is generally open at the proximal side. The supra- triangles often have a traversing cross-vein, in the females more frequently so than in the males. The three members of the group are subspeci- fically distinct only. They can be discerned on the ground of their coloration. Slight differences are found in the configuration of the anterior hamules of the males but no structural differ- ences are found in the shape of the vulvar lami- nae. Z. c. klugi is somewhat more robust than the other two. The superior lobe of the anterior hamules is very long and slightly upcurved at the tip. Z. c. klugi is also the darkest of the group. The pale frontal band on the superior surface of the frons is medially interrupted by a pitch black band that is broader in the female than in the male. Z. c. spectabilis is the palest of the group. It is distinguished from the other two by the pres- ence of large pale markings on the abdominal segments 8 to 10. The superior lobe of the ante- rior hamules is widened at the tip. (3) The batesi group, comprising Z. batesi Selys and Z. wucherpfennigi Schmidt. The distal segment of the penis has a pair of very long flagella. The anterior hamules of the male are as large as those ot the campanulata group; the superior lobe is short and stout; the inferior lobe is much larger than that of the campanulata group; the internal lobe is very small. The vulvar lamina is widest at base and roughly triangular in outline; it exhibits some variation, the divisions of the tip being often spread out. The wings have no basal subcostal cross-vein, no extra cubito-anal cross-vein, and no travers- ing cross-vein in the supratriangles. The anal BELLE: Zonophora 149 loop in the hind wings is open at the proximal side. The two species of the group differ struc- turally in the penile organ, the male caudal ap- pendages and vulvar laminae, but the differ- ences are slight and they can be shown more readily by figures than description (cf. Schmidt, 1941). The darkest of the two is wucherpfennigi. The seventh abdominal segment of the male is entirely black in this species; it is markedly yel- low-coloured at the base in batesi. The caudal appendages (stylets) of the female are largely blackish or brownish (paler in middle and black at base) in wucherpfennigi (cf. Schmidt, 1941: fig. 12f); they are largely pale-white (black at base) in batesi. (4) The supratriangularis group, comprising Z. supratriangularis Schmidt and Z. nobilis spec. nov. They differ structurally in the male caudal appendages and genital hamules. The distal seg- ment of the penis has a pair of very long flagella. The internal lobe of the anterior hamules of the male is small. The vulvar lamina is about half the length of the ninth sternum. The wings have no basal subcostal cross-vein but there are extra cubito-anal cross-veins. The supratriangles normally have a traversing cross- vein. The anal loop in the hind wings is open at the proximal side although not always very dis- tinct. The two species are the palest representatives of the genus. (5) The surinamensis group, comprising Z. surinamensis Needham, Z. solitaria solitaria Rá- cenis, Z. solitaria obscura Belle, and Z. regalis Belle. The anterior hamules of the male have no in- ternal lobe. The male superior caudal append- ages are not knotty at the internal margin. The vulvar lamina is about one-third as long as the ninth sternum. The wings have no basal subcostal cross-vein and generally no extra cubito-anal cross-vein. The supratriangles occasionally have a travers- ing cross-vein. The hind wings have a more or less distinct anal loop. Z. s. solitaria is the palest of the group having the lateral sides of the pterothorax largely palé and the dorsum of the eighth abdominal seg- ment with a well-developed pale basal spot. Z. s. obscura is the darkest of the group and likewise the darkest representative of the genus. It is dis- tinguished from all congeners by the lack of sec- ond pale antehumeral stripes (these stripes are partly developed in Z. s. solitaria). Z. regalis is the largest of the genus. SYSTEMATIC SECTION As considered in this study, the genus Zono- phora comprises 15 taxa of which three are de- scribed as new. The taxa are summarized in ta- ble 2, together with their type localities, type status and type locations. The males of all species and subspecies are known. The females of diversa, nobilis, regalis, solitaria solitaria, and surinamensis are yet un- known. Their discovery may corroborate the relationships hitherto found between some spe- cies. The recognition of the diverse species and subspecies is comparatively easy. Separate keys have been prepared for the identification of males and females. The key to the males has been based upon the structural characters dis- played by the anterior hamules, the penis, the superior caudal appendages, the reticulation of the wings, and some other details. The key to the females has been based on characters dis- played by the vulvar lamina and on some color- ation details. Key to the (sub)species Males 1. Basal subcostal cross-vein normally present (Calippuseroup) ger ER 2 — No basal subcostal cross-vein........... 4 2. Abdominal segments 8 to 10 with extensive yellow markings calippus spectabilis — Abdominal segments 8 to 10 without or with weakly developed yellow markings . 3 3. Interrupted median stripe of pale anterior band of frons well-developed and pitch black, its width normally one-third the mid-dorsal length of superior surface of OMS OT mor calippus klugı — Interrupted median stripe of pale anterior band of frons absent, ill-defined or very narrow, its width smaller than one-third the mid-dorsal length of superior surface of ONE sneer: are calippus calippus 4. Superior caudal appendages knotty near middle at internal margin; anterior hamules Withimtemallobeltie 0) PR 8 — Superior caudal appendages not knotty at internal margin; anterior hamules without internal lobe (suvimamensis group)....... 5 5. Lateral sides of pterothorax largely pale . MANNEN MATTE solitaria solitaria 150 10. IE IDE 113). 14. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) Lateral sides of pterothorax with well-de- velopecl care SWBDES s oo 0000000 aecce took 6 . Second pale antehumeral stripes undevel- Ope dees ae we teen n solitaria obscura Second pale antehumeral stripes well-devel- OPERARE HARE MICRA LI GI SEO ER 7 . Distal segment of penis with a pair ot very SONDE EEE A surinamensis Distal segment of penis with a pair of re- curved leaf-like plates regalis . Distal segment of penis with short tusk-like spines; internal lobe of anterior hamules- well-developed and finger-like (campanu- Aou) WAM OU INR TE LEN 9 Distal segment of penis with very long flag- ella; internal lobe of anterior hamules very small 12 . Distal halves of posterior hamules parallel; anterior hamules distinctly trilobate .... 10 Distal halves of posterior hamules diver- gent; anterior hamules not evidently trilo- bate due to an ill-defined superior lobe..... NA RTE AE Eee DRE Dot: sE diversa Terminal segments of abdomen without or with weakly developed yellow markings ... campanulata campanulata Terminal segments of abdomen with exten- sive wellom madsimes 5540000000000000c 11 First pale antehumeral stripes connected with pale collar and second pale antehu- meral stripes campanulata machadoi First pale antehumeral stripes neither con- nected with pale collar nor with second pale antehumeral stripes MER EIERN na campanulata annulata Supratriangles crossed (supratriangularıs SOUP A SIA, LORDO IAT 13 Supratriangles open (bates: group) 14 Superior caudal appendages almost three times as long as mid-dorsum of abdominal segment 10; internal tooth of each appen- dage at three-fifths the appendage’s length . . Ws EC ATL AL ea bettolo nobilis Superior caudal appendages twice as long as mid-dorsum of abdominal segment 10; in- ternal tooth of each appendage at half the appendage’s length supratriangularis Abdominal segment 7 with large, pale basal marking: rete ged or bode batesi Abdominal segment 7 entirely black wucherpfennigi Females . Basal subcostal cross-vein normally present (calippus group) > . Abdominal No basal subcostal cross-vein........... 4 Abdominal segments 8 to 10 with extensive yellow markings calippus spectabilis Abdominal segments 8 to 10 without or with weakly developed yellow markings . 3 . Pale frontal band on superior surface of frons widely interrupted in middle by pitch black calippus klugi Pale frontal band on superior surface of frons medially not or not distinctly inter- rupted by black calippus calippus . Vulvar lamina reaching to apex of ninth sternum Surinam 2:28, E, MP. This species was described by Burmeister (1839) after a single male from “Brazil”. This male must be considered to be identical with the darkest representative of the campanulata group since the statement “fusca, subtus dilu- tor; abdominis segmento septimo fascia basali pallida” in the original description evaluates the specimen. If all its abdominal segments has had extensive pale markings, they should undoubt- edly be mentioned in the description. The location of the type of D. campanulata 1s unknown. Calvert (1898) as well as Belle (1963) were unable to find it. Also further search made by myself was in vain. I am strongly inclined to believe that the original specimen is lost, possi- bly destroyed by a dermestid infestation (An- threnus). The species, however, has been very well described and figured in the Monographie des Gomphines (Selys & Hagen, 1858). In order to stabilize its identity a neotype is here desig- nated. Neotype designation of Diastatomma cam- panulata Burmeister. — The chosen neotype is a male from a remarkable uniform series placed under Zonophora campanulata (Burmeister) in the Selysian collection in Brussels. This series consists of the following specimens: (1) A male with the pin label “Cl. 2.” (2) A female with the pin labels “Cl. 3” and “Zonophora campanulata B ©”, this latter in Selys’ handwriting. (3) A male with the pin labels “122”, “122”, “Tijuca P.B.”, and “Zonophora campanulata B. 3”, this latter in Selys’ handwriting. (4) A male with the pin label “Tijuca P.B.” The abbreviation “P.B.” may refer to Paul de Borchgrave who has collected Odonata in Tiju- ca near Rio de Janeiro round-about 1860 (cf. Calvert, 1956: 88, 175—176). The abbreviation “Cl.” may refer to Dr. Clausen from whom Selys received the speci- mens mentioned in the Monographie des Gom- phines, p. 496 (236 sep.). I think, that the male which served for Ha- gen’s figures in this monograph belonged to the above mentioned series. This male should have borne at the pin the (missing) label “Cl. 1” and probably also Selys’s identification label of the Tijuca male. The specimens of Dr. Clausen are homotypes as may appear from Selys’s (1858) remark in parentheses: “C’est le type de M. Burmeister qui offre les nombres les plus grands”. The homotype male labelled “Cl. 2” is here designated as neotype of Diastatomma campanulata Burmeister, 1839. Z.c. campanulata exhibits some striking vari- ations. The hind wings of none of the present specimens have a distinct open anal loop, due to a rather well-developed vein A2. The hind wings of some specimens have more or less a distinct anal loop of 3 to 5 cells, with the vein A1 angled at the lower distal corner of the anal loop. The form of the vulvar lamina is also variable, in particular at the tip. The vulvar lamina of the present females approaches that depicted by St. Quentin (1973) but it differs considerably from that depicted by Schmidt (1941). The female of the neotype series has a vulvar lamina with a damaged tip. As this female was the only one available for Hagen, it apparently served for his figure in the Monographie des Gomphines (pl. 13, fig. 1t). Some data in Selys’s redescription of 1854 are incorrect. The measurement for the female ab- domen is 50 mm (incl. caud. app.) and not 58 mm (cf. Campion, 1920, foot-note on p. 137). Further, the number of given antenodal cross- veins in the fore wings of the male is 18 instead of 13 (cf. Selys, 1869). The nodal index of the neotype is 14 : 20— 20 : 13/13 : 145 : 14. Fraser (1940) published a figure of the penis 152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) of a Zonophora male, which he referred to Z. campanulata. But judging from the conforma- tion of the penile organ his male belonged to the batesi group. Remark. — Caution should be made regard- ing the locality labels of the specimens in the Martin collection (lodged in the Muséum Na- tional d'Histoire Naturelle, Paris) since several of these old labels prove to be unreliable. Some specimens of Z. c. campanulata have the pin la- bels “Surinam” and “Guyana” but these locali- Hesm May ab eMIMCOGKe ChsincelsnOr ml DMC: Geijskes nor I myself have encountered this species during our extensive explorations in Su- rinam and Guyana. On that account these re- cords have not been inserted in the distribution map. To further illustrate the unreliability of some locality labels of the Martin collection, in the boxes in which the specimens of Zonophora were stored, I found four males of Phyllogom- phoides pacificus (Selys, 1873) labelled “Minas Gerais” and “Bresil”, and a male of Aphylla car- aıba Selys, 1854, labelled “Bresil”. Zonophora campanulata machadoi St. Quentin, 1973 (figs. 21, 24, 35—36) Zonophora machadoi St. Quentin, 1973: 338—340; figs. 4a—b (6). Kiauta, 1981: 35, 60. Material. — Brazil: Minas Gerais, Belo Horizonte, 18 March 1979, 1 6, AC; Tabuoes, Ibirité (near Belo Horizonte), 22 March 1979, 1 &, CJ, both G. Jurzitza leg.; Ibirité, 21 March 1979, 1 2, A. B. M. Machado leg., CM. This subspecies was described by St. Quentin (1973) after a single male from Belo Horizonte, Brazil. The female is described below. Female (hitherto unknown; degutted and treated with acetone). — Total length, 64 mm; abdomen, 49 mm (including caudal appendages, 4.3 mm); hind wing, 45 mm; costal edge of pterostigma in fore wing, 6.0 mm, in hind wing, 6.5 mm. Coloration resembling that of male holotype. Face brown with conspicuous bright yellow band covering suture between postclypeus and vertical part of frons. Labium with pale cross- band in midfield. Anteclypeus becoming paler on outer edges. Superior surface of frons brown with oblong yellow spot on either lateral side. Colour pattern of prothorax and pterothorax very similar to that of male. Abdomen relatively shorter than that of male. Pale markings on sides of abdominal segments 8 and 9 much smaller than in male, and abdominal segment 10 entirely dark brown (fig. 36). Proportional length of abdominal segments 7, 8, 9, and 10 approximately as 45 : 30 : 20 : 10, with the caudal appendages 34 on same scale. Vulvar lamina shaped as shown in fig. 21. Pterostigma surmounting eight cells. No bas- al subcostal cross-vein. No extra cubito-anal cross-vein. No traversing cross-vein in supra- triangles. No anal loop in hind wings. Nodal in- dex 14 :20—22 : 13/13 :13—13 : 15. Second primary antenodal cross-vein the eighth in fore wings, the sixth in hind wings. Intermedian cross-veins 7—9/3—3 in fore and hind wings, respectively. Hind wings with six (left) and sev- en (right) paranal cells, six (left) and seven (right) postanal cells, and area posterior to Cu2 six to seven cells wide. Zonophora campanulata annulata subspec. nov. (figs. 22, 23, 37—38) Material. — Brazil: Gioäs, Jatai (Jatahy), 3 d holo- type and paratypes), 1 © (allotype), MP. This subspecies, though very similar in its morphology to Z. c. campanulata and Z. c. machadoi, is readily distinguished from both by the aberrant colour pattern of the pterothorax, which, in lateral view, shows a conspicuous pair of well-developed pale twin-stripes, the first pair being formed by the second pale antehu- meral stripe and the first pale lateral stripe, and the second pair by the second and third pale lateral stripes. The colour pattern of the dorsum of the pterothorax is most like that of Z. c. campanulata by the isolated pale antehumeral stripes and pale collar, while the abdomen greatly resembles that of Z. c. machadoi by the annulated appearance owing to the extensive pale markings at the base of the segments. According to some of the hitherto used char- acters, the colour differences mentioned above are important enough to consider the present examples from Jatai a distinct variety or subspe- cies. I prefer to regard them as a subspecies since they are forming a homogeneous series, and since they seem to be the only form occur- ring in that part of the distribution area. How- ever, as long as our knowledge is restricted to the scarcely available material it is difficult to indicate the value of the characters used for the subspecific distinction. The features often vary BELLE: Zonophora 153 in some widely ranging gomphid species to such a degree that it remains to be seen whether the status of the subspecies introduced will stand the test of future collecting. Male (holotype; abdomen broken on segment 6 and between segments 3 and 4; left superior caudal appendage broken off and lost). — Total length, 66 mm; abdomen, 50 mm (incl. caud. app., 4 mm); hind wing, 42 mm; costal edge of pterostigma in fore wing, 5.6 mm, in hind wing, 6.0 mm. Face yellowish green but brown along border of labrum and just below frontal ridge of frons. Superior surface of frons yellowish green on an- terior half, brown on basal half. Vertex black. Occipital plate and rear of occiput yellow- brown. Rear of head yellowish brown. Base of mandibles, genae, labium and adjacent mouth parts pale brown. Prothorax dark brown with greenish yellow markings. Median lobe with pale middorsal twin-spot and pale spot on each lateral side; posterior lobe with pair of pale oblong spots. Pterothorax dark brown to black, the pale markings greenish yellow, becoming yellow on metepimeron. Colour pattern of pterothorax shaped as shown in accompanying diagram. Middorsal carina and antealar sinus pale. Pale collar and first pale antehumeral stripe narrow. Second pale antehumeral stripe and pale lateral stripes rather broad. Femora reddish-brown but inner side of first pair green. Tibiae, tarsi, and claws dark brown. Lamina tibialis of first tibiae one-seventh the tibial length. Wings clear, its venation pale brown, becom- ing darker on posterior margin of wings. Pterostigma brown-yellow, surmounting 6— 6V cells. Brace vein present. No basal subcostal cross-vein. A single cubito-anal cross-vein in addition to inner side of subtriangle in all wings. Supratriangle in right hind wing with a travers- ing cross-vein, in other wings uncrossed. Sub- triangles open. Discoidal triangles two-celled. Nodal index 12 : 18—16 : 11/13 : 12-11 : 12. Second primary antenodal cross-vein the sev- enth in left fore wing, the sixth in other wings. Intermedian cross-veins 8—8/4—4 in fore and hind wings, respectively. Hind wings with open anal loop (open at proximal side), a four-celled anal triangle, five (right) and six (left) paranal cells, five (right) and six (left) postanal cells, and area posterior to Cu2 five cells wide. Abdomen dark brown to black, the pale markings yellow. Segment 1 pale on lower part of lateral sides and along dorsal posterior mar- gin. Segment 2 pale on lower parts of lateral sides, on auricles, and just above auricles. Seg- ments 3 to 8 with large pale basal markings which become successively larger on 3 to 8, laterally reaching to supplementary transverse carina on 3, and extending to beyond this carina on 4 to 8. Segment 9 with pale lateral markings extending along lower part of side to apex of segment. Segment 10 largely pale. Tubercles with externo-lateral pale spots. Inferior append- age black. Superior appendages pale yellow but extreme base brown. Female (allotype). — Total length, 69.5 mm; abdomen, 53 mm (incl. caud. app., 4 mm); hind wing, 46 mm; costal edge of pterostigma in fore wing, 6.4 mm, in hind wing, 6.6 mm. Very similar to male holotype in stature and coloration but abdomen stouter. Proportional length of abdominal segments 7, 8, 9, and 10 ap- proximately as 40 : 23 : 19 : 10, with the caudal appendages (stylets) 28 on the same scale. Pterostigma surmounting 5/—612 cells. No basal subcostal cross-vein. No extra cubito-anal cross-vein. No traversing cross-vein in supra- triangles. Second primary antenodal cross-vein the eighth in left hind wing and right fore wing, the seventh in other wings. Hind wings with a weak development of an anal loop of four cells. The male paratypes are very similar to the ho- lotype. All their wings have uncrossed sub- triangles and supratriangles. The triangles have one dividing cross-vein. The anal triangles are four-celled. The hind wings of one of the male paratypes have a weak development of an anal loop of four cells. Zonophora diversa spec. nov. (figs. 39—50) Material. — Argentina: Misiones, Dos de Mayo, January 1982, 2 d (paratypes), J. Foerster leg., AC and CJ. — Brazil: Santa Catarina, Nova Teutonia, 2 March 1942, 1 d (paratype, AC), 3 March 1942, 2 6 (paratypes), 4 March 1942, 1 d (paratype), UMMZ; same locality, January 1946, 1 d (holotype, CC), all Fritz Plaumann leg. — Paraguay: Alto Parana, Ta- queri (Tacuri), near Rio Nacunday, 20 February 1980, 1 d (paratype), H. Miers leg., CM. This species is closely related to the type-spe- cies of the genus, Z. c. campanulata, but it is rather easily distinguished in the male sex by the stouter superior caudal appendages, the ill-de- 154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) fined superior lobe of the anterior hamules, and the divergent distal halves of the posterior ham- ules. The female is unknown. Male (holotype). — Total length, 70 mm; ab- domen, 54 mm (including caud. app., 4 mm); hind wing, 47 mm; costal edge of pterostigma in fore wing, 5.0 mm, in hind wing, 5.4 mm. Face greenish-yellow but labrum black along free-border. Superior surface of frons black at base, greenish yellow on its three-fifths anterior portion, yellow on anterior ridge. Vertex black. Occipital plate yellowish-green, its posterior margin slightly and evenly convex, and heavily fringed with brown hairs. Prothorax dark brown, the middle lobe with reddish brown lateral markings and a more yel- lowish mid-dorsal twin-spot, the hind collar dark reddish brown. Pterothorax blackish brown with greenish yellow markings, its colour pattern shaped as shown in fig. 46. Femora blackish brown, save for a streak of yellowish brown on outer basal two-thirds portion of third femora. Tibiae and tarsi black. Claws dark reddish brown. Wings hyaline, its venation brown including frontal margin of costa. Pterostigma brown, surmounting 5/2—612 cells. Brace vein present. No basal subcostal cross-vein. Nodal index 14 : 18—18 : 14/13: 14—13 :15. Second prima- ry antenodal cross-vein the seventh in right hind wing, the eighth in other wings. Intermedian cross-veins 7—6/4—5 in fore and hind wings, respectively. No extra cubito-anal cross-vein. All subtriangles and supratriangles free from cross-veins. All discoidal triangles two-celled. Hind wings with an ill-defined anal loop of four (left) and five (right) cells, an anal triangle of six cells (two cells small), six paranal cells, seven (left) and six (right) postanal cells, and area pos- terior to Cu? six cells wide. Abdomen black with yellow markings, its colour pattern shaped as shown in fig. 43. Seg- ment 10 with brownish yellow mid-dorsal spot. Sterna of segments 6 to 8 brown to blackish brown but yellow on basal one-fifth; sternum of 8 with an extra, oblong yellow spot on either side half-way the sternum. Inferior caudal ap- pendage black. Superior caudal appendages dark brown. Caudal appendages and accessory geni- talia shaped as shown by the accompanying fig- ures. The following colour differences between the holotype and the paratypes from Nova Teuto- nia are noteworthy. One male has the black band along the free border of the labrum not de- veloped in the middle and at either side; two males have the black basal band on the superior surface of the frons medially interrupted by greenish yellow; in three males the black mark- ings on the dorsum of the pterothorax are re- duced; one male has the black antehumeral stripes almost lacking; one male has the pale markings of the prothorax yellow. The coloration of one of males from Argenti- na is very similar to that of the holotype. The other Argentine male has the black humeral and femoral stripes largely undeveloped. The male from Paraguay exhibits the most striking colour differences. Its labrum is entirely green. The black basal band on the superior sur- face of the frons is reduced to two narrow streaks at the level of the antennae. The vertex has a greenish yellow median stripe between the lateral ocelli. The occipital plate is yellow. The hind collar of the prothorax is entirely reddish brown. The pterothorax is almost entirely green; the dorsum is entirely green; the black humeral, interpleural and metapleural stripes are developed only near the subalar carinae. The terminal segments of the abdomen have exten- sive yellow markings; segments 5 and 6 are yel- low on the apical half; the yellow marking of 7 reaches to a point three-quarters the way along the segment on the sides; segment 8 is yellow on slightly more than the basal half; segment 9 is yellow on the basal half; segment 10 is yellow with black baso-lateral markings and a black hind margin, the tubercles are yellow. Zonophora calippus calippus Selys, 1869 (figs. 7, 10, 16, 73) Zonophora calippus Selys, 1869: 199—200 (36—37 sep.) (6, 2). Hagen, 1875: 54. Kirby, 1890: 75. Campion, 1920: 140. Schmidt, 1941: 76 —77, 80 — 81, 84. Needham, 1944: 171, 218—219, pl. 16, figs. 18a—f. Belle, 1963: 60, 62—63, 6569; 1966: 64, figs. 97—98, pl. 11b (exuviae); 1972: 217, 237—238, fig. 39. St. Quentin, 1973: 338 339. Kiauta, 1979: 267—269. Zonophora calippus calippus Schmidt, 1941: 8485, figs. 5d, 6a, 8a, 8f, 9a, 11a. Material. — Brazil: Pará, Santarém, Amazon River (Mahica), December 1920, 1 d, A. H. Fassl, leg., UMMZ, Pará, Fordlandia, February 1957, 1 9; Pará, Belem (Utinga Forest), February 1957, 2 6, all A.B.M. Machado leg.; Paraiba, Joao Pessoa (Bura- quinho Forest), May 1976, 1 d, Kesselring leg, CM. BELLE: Zonophora 155 — Guyana: Georgetown, March 1955, 1 d, M. Alva- renga leg., CM. This species was described by Selys (1869) af- ter a single pair from Santarém, Brazil. Need- ham (1944) recorded it from Surinam, Belle (1972) from Venezuela, and St. Quentin (1973) from Guyana. Some specimens of Zonophora already placed on record by St. Quentin (1973) were kindly sent to me for restudy by Prof. Ma- chado. The present male from Santarém has the pale frontal band on the superior surface of the frons narrowly interrupted in the middle by black; the pale band is normally not or not distinctly interrupted in the middle. The male from Parai- ba has the mid-lateral pale stripe on either side of the pterothorax almost absent. St. Quentin (1973) referred the female from Fordlandia (100 km south-west of Santarém) to the genus only, possibly due to the fact that in this specimen the wings have no basal subcostal cross-vein. The first and second pale antehumeral stripes are often confluent in Z. c. calippus. The occipi- tal ridge of the female has generally a pair of denticles which are often unsymmetrically situ- ated or partly developed. The larva of Z. c. calippus was described from Surinam by Belle (1966). Zonophora calippus spectabilis Campion, 1920 (fig. 12) Zonophora spectabilis Campion, 1920: 138—140; pl. 7, fig. 15 (8). Schmidt, 1941: 76-77, 80-81, 86. Belle, 1963: 61—62, 66—68; 1972: 237—238; figs. 36—38 (©). Kimmins, 1969: 298. St. Quentin, 1973: 338—339. Paulson, 1977: 176. Zonophora calippus spectabilis Schmidt, 1941: 86—88, 92; figs. 7c, 11c. Needham, 1944: 218. Material. — Bolivia: Santa Cruz, (San José de) Chiquitos, Roboré (300 m), 26 December 1953, 1 d, W. Forster leg., IZM. — Brazil: Sao Paulo, Itapeti- ninga, 10 February 1978, 1 d (very teneral), E. Dente leg., CM. Z. c. spectabilis was described by Campion (1920) after a single male from Sapucay, Para- guay. Belle (1972) recorded this subspecies from Bolivia. Paulson (1977) included Argentina in its range. I accordingly wrote to Dr. Paulson on May 24, 1980, about this record in his list and he kindly replied, under date of June 8, 1980, that he had wrongly entered on his cards Need- ham’s (1944) record that Campion described spectabilis from “Argentina” (instead of Para- guay). The presence of Z. c. spectabilis in Brazil was not known before. The yellow marking of the tenth abdominal segment of the male from Brazil is much more extended than as shown by Schmidt (1941). for a male from Paraguay. The male from Bolivia, on the contrary, has the yellow markings on the abdominal segments 8 to 10 smaller, and it lacks the lance-shaped yellow extension on the mid- dorsum of the segments 8 and 9. Zonophora calippus klugi Schmidt, 1941 (fig. 11) Zonophora calippus klugi Schmidt, 1941: 80—81, 85— 86; figs. 4a, 5a—b, 7a, 8g, 9b, 10a, 11b, 12a (ORD 058 238,251, Nacemıs, 1959: 492: Se. Quentin, 1973: 339. Zonophora klugi Belle, 1963: 60—63, 66—68; pls. 3— 4511972238: St. Quentin, 1973..338 339; Material. — Brazil: Amazonas, Tefe, January 1958, 1 d, Carvalho leg.; Acre, Feijó, no date, 1 d; Terri- tório de Rondônia, Pôrto Velho, 16 November 1962, 1 2, both Bokermann leg., CM; Territorio de Ron- dônia, Madeira-Mamoré River, near Jaci Paraná (km 87—91), 7 February 1922, 1 &, J. W. Strohm & J. H. Williamson leg.; Amazonas, Sao Paulo de Olivenca (Alto Rio Solimöes), April 1932, 1 d, F. Wucherpfen- nig leg., UMMZ. — Ecuador: Napo-Pastaza (Rio Na- po watershed), Río Ilapulin, Jatún Yacu (800 m), 13 March 1937, 1 2; same locality (700 m), 28 March 1937, 2 2, all Wm Clark-Macintyre leg., UMMZ. Z. c. klugi was described by Schmidt (1941) after several males and females from Mishiyacu, Peru. St. Quentin (1973) recorded it from Bra- zil. The subspecies is new to the fauna of Ecuador. Belle (1963) selected the male with the Senckenberg’s catalogue number 24000 as the lectotype. The triangular envelope with the male taken by Strohm & Williamson is labelled “State of Matto Grosso” instead of Territorio de Ron- dônia. The envelope has the field note: “Flew into wind-shield of motor-car (a fine wire screen which caught and held many kinds of in- sects)”. The inserted male has the pitch-black stripe which divides the pale frontal band on the superior surfacce of the frons about one-tenth the width of the frons. This stripe is generally one-fourth the width of the frons in the males ot this subspecies. The male has further three small yellow spots on either side of abdominal seg- ment 8. This segment is entirely black in the other examples of this subspecies. The present females from Ecuador are the 156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) largest specimens of the series; their mea- surements are; abdomen, 49 mm (incl. caud. app.); hind wing, 45—-46 mm. The male from Tefé was referred to Z. batesi by St. Quentin (1973). Zonophora batesi Selys, 1869 (figs. 1—5, 8, 13, 17, 74) Zonophora batesi Selys, 1869: 198—200 (35—37 sept.) (4). Hagen, 1875: 54. Kirby, 1890: 75. Campion, 1920: 137. Schmidt, 1941: 76, 80, 82, 84, 92— 94, 96; figs. 4c, 6d, 8d, Ie, 11f, 12d, 12g. Needham, 1944: 218—219; pl. 16, fig. 19a. Cal- vert, 1948: 61—62. Belle, 1963: 60—65, 67; 1966: 62—64; fig. 95— 96; pl. 11b (exuviae); 1972: 217, 236; 1977b: 292. Geijskes, 1971: 666. St. Quentin, 1973: 338—339. Zonophora batesi batesi St. Quentin, 1973: 338. Zonophora bodkini Campion, 1920: 136—138; pl. 7 figs. 10—14 (2). Schmidt, 1941: 76, 78, 92—94, 96. Needham, 1944: 218. Calvert, 1948: 62. Belle, 1963: 61, 65. Kimmins, 1969: 291. Geijskes, 1971: 666. Material. — Brazil: Amapá, Serra do Navio, Octo- ber 1957, 1 ®, Lenko leg.; Pará, Ananindeua, 7 Sep- tember 1964, 1 d, Souza leg., CM. — Guyana: Tuma- tumarı, 5 February 1912, 1 6,1 ©, L. A.&E. B. Wil- liamson, and B. J. Rainey leg., UMMZ. — Surinam: Coropina Creek (upper part), Dauwdropkamp, 6 No- vember 1955, 1 2, J. Belle leg., ML. Z. batesi was described by Selys (1869) after a single male from Fonte Boa, Brazil. Campion (1920) recorded it (under Z. bodkini) from Guyana, Needham (1944) from Surinam and Geijskes (1971) from French Guyana. The larva of this species was described by Needham (1944) and Belle (1966). Some features of the female here recorded from Surinam were already published by Belle (1963) but its locality data were not yet put on record. Zonophora wucherpfennigi Schmidt, 1941 Zonophora wucherpfennigi Schmidt, 1941: 80, 84, 9496; figs. 3a, 4d, 5e, 8e, 11g, 12e—f (6, ©). Belle, 1963: 62—63, 69. St. Quentin, 1973: 338. Zonophora batesi wucherpfennigi St. Quentin, 1973: 338—339. Material. — Brazil: Pará, Itaituba, November 1961, IPB Eerrerales (No Const Quentin, 1972), CM. Lectotype designation of Zonophora wucherpfennigi Schmidt. — The type series of this Brazilian species belonged to the collection of Schmidt and consisted of two males from Manicoré and one female from Tefé. Schmidt did not designate a holotype so that a lectotype must be selected now. The penile organ of one of two males has been extruded and figured by Schmidt (his fig. 11g). The male which served for this figure is here designated as the lectotype of Zonophora wucherpfennigi Schmidt, 1941. This lectotype is in the collection ot Dr. S. Asa- hina, Tokyo, because Schmidt’s collection has been incorporated into it. The female of Zonophora wucherpfennigi in the collection formerly owned by Schmidt was tentatively referred to this species. The female from Itaituba, referred to Z. wucherpfennigi by St. Quentin (1973), was kindly lent to me for re- study by Prof. Machado. It fits fairly well the description and figures of the female from Tefé given by Schmidt (1941) except for the large pale basal marking of the seventh abdominal segment, which in the female from Itaituba is reduced to a pair of dorso-lateral spots. In the female of batesi this pale basal marking is mid- dorsally interrupted by a black line. In my opin- ion the females from Tefé and Itaituba are both conspecific with wucherpfennigi. Zonophora surinamensis Needham, 1944 (figs. 26—30, 76) Zonophora surinamensis Needham, 1944: 219—221; pl. 16, figs. 17a—b (6). Belle, 1963: 60, 62, 69; 1966: 62—63; figs. 93—94; pl. 11a; 1972: 238; 1976: 206. Racenis, 1970: 28. St. Quentin, 1973: 338. Z. surinamensis was described by Needham (1944) after a single male collected in Brazil at the Rio Mapaoni, a left-tributary of the Jari River, in the Serra de Tumucumaque. Belle (1972) recorded it from Surinam. The original description of this species was elucidated by depictions of the conventional dorsal and lateral views of the male caudal ap- pendages only. Belle (1966) published figures of the accessory genitalia of the male holotype in lateral and ventral views. The increasing num- ber of related species, however, makes it advis- able to pay more attention to some critical structures, hence the here offered original cam- era lucida drawings of the penis, penis guard, and anterior genital hamule of this species. A di- agram of the thoracic colour pattern is also added. The male of Z. surinamensis taken in Su- rinam (cf. Belle, 1972) has served to furnish the BELLE: Zonophora 11577 present illustrations. As clearly appears from the figures, the colour design of the pterothorax resembles greatly that of Z. regalis while the penis agrees with that of Z. campanulata in that the tip possesses a pair of very short spines. Contrary to the holotype, the male from Suri- nam has no extra cubito-anal cross-vein in the wings, whereas the anal loop in either hind wing consists of four cells only (five cells in the type). Zonophora solitaria solitaria Racenis, 1970 (figs. 51—54) Zonophora solitaria Racenis, 1970: 26—28; figs. 2—3 (6). De Marmels, 1980: 128; 1983: 8. This species was described by Rácenis (1970) after a single male from Guayaraca-Auyantepui, Venezuela. Mr. De Marmels, who investigated the male holotype of Z. solitaria Racenis in the Instituto de Zoologia Agricola in Maracay, generously provided me with valuable sketches of the markings of some important details (figs. 51— 54). He also compared the type with that of Z. obscura Belle and wrote from Egg in Zwitser- land, July 29, 1981: “It seems rather different from your obscura, at least at a first sight. The solitaria’s thorax sides are nearly uniformly green olivaceous, almost lacking black bands. On dorsum of (abdominal) segment 8 there is a big tripartite yellow basal spot. Pterostigma is darker brown. The anal appendages seem iden- tical, as well as the (accessory) genitalia al- though I did not extract the penis. Rácenis’s (or his assistant’s) figures are always very accu- rate”. Zonophora solitaria obscura Belle, 1976 (figs. 18, 55) Zonophora obscura Belle, 1976: 197, 203—206; figs. 16—24 (8). Kiauta, 1981: 35, 49. De Marmels, 1983: 8. Material. — Venezuela: Bolivar, at km 125 from El Dorado to Santa Elena (1100 m), 25 September 1967, 1 2, Rosales, Gelvez & Rodríguez leg., IZM. This subspecies was described by Belle (1976) after a single male from Sta. Elena, Venezuela. Mr. De Marmels, who compared the male holo- type of Z. obscura Belle with that of Z. solitaria Rácenis, did find structural differences neither in the caudal appendages nor in the accessory genitalia, although he did not extract the penis of solitaria. The colour differences between the two males, however, are great and they seem important enough to consider obscura a subspe- cies of solitaria, at least provisionally. The dis- tinction may be apparent from the following comparison: (1) In obscura, the second pale antehumeral stripes are not developed; in solitaria, they are developed on the lower (anterior) part of the thoracic dorsum. (2) In obscura, the dark lateral stripes of the pterothorax are well-developed, the first and second dark lateral stripes being entirely or largely confluent; in solitaria, the lateral sides of the pterothorax are almost uniformly pale, lack- ing distinct black bands. (3) in obscura, the abdominal segment 8 is en- tirely black; in solitaria, it has a large tripartite yellow basal spot. Differences are also found in the coloration of the face, the vertex and the occipital plate. If a comparison is made with the sketches of Mr. De Marmels (figs. 52, 54) the following differences are noteworthy: (4) In obscura, the vertical part of the frons and the postclypeus are black except for a pale cross-stripe covering the fronto-clypeal suture; in solitaria, they are largely pale. (5) In obscura, the vertex and occipital plate are entirely black; in solitaria, each of them has a pale central spot. Although obscura is a darker species than sol- itaria, the pterostigmata of obscura are lighter brown than those of solitaria. The female of Z. s. obscura is described be- low. Female (hitherto unknown; abdomen partly flattened; tips of fore wings broken off but not lost). — Total length, 63 mm; abdomen, 47 mm (incl. caud. app.); hind wing, 46 mm; costal edge of pterostigma in fore wings, 5.8 mm, in hind wings, 6.0 mm. Similar to male but labrum with an extra pale mid-basal spot, vertex with a pale spot between conical processes (tips of these processes yel- low), and pale metepisternal stripes weakly de- veloped near subalar ridge. Pale baso-lateral spots of abdominal segments 3 to 6 reduced, reaching to supplementary transverse carina on 3, being nearly absent on 6. Pale basal marking on dorsum of 7 well-developed. Occipital plate with several whimsical grooves in middle part, the crest densely set with long, stiff, black hairs (broken off in middle part of crest). Caudal ap- pendages (stylets) black-brown at bases, becom- ing paler (brown) towards tips. Abdominal seg- ments 7, 8, 9, and 10 approximately in ratio 158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) 37 : 24:17:10, with the caudal appendages (stylets) 19 on the same scale. Vulvar lamina one-third the length of ninth sternum, in ventral view triangular in outline, projecting in an oblique direction downwards and rearwards (possibly due to the fact that the abdomen is partly flattened), its posterior margin medially excised U-shaped for about two-thirds the length of vulvar lamina, the lobes triangular and acute. No basal subcostal cross-vein. Nodal index 16 :21—21 : 16/16 :17—17 : 16. Second prima- ry antenodal cross-vein the eighth in right hind wing, the seventh in other wings. Cell between first and second postnodal cross-veins divided by a cross-vein subparallel to costa. Intermedian cross-veins 10—9/4—5 in fore and hind wings, respectively. Supratriangle in left fore wing with one cross-vein, in other wings open. No extra cubito-anal cross-veins. Subtriangles open. Dis- coidal triangles two-celled. Hind wings without anal loop, with six paranal cells, six postanal cells, and area posterior to Cu2 five to six cells wide. Zonophora regalis Belle, 1976 Zonophora regalis Belle, 1976: 197, 200—203; figs. 5—15 (dg). Kıauta, 1981: 35, 49. This large species was described by Belle (1976) after three males from La Ceiba del Ven- tuari, Venezuela. The anal loop in the hind wings of the holo- type is two-celled owing to the fact that one of the dividing cross-veins is only partly devel- oped. The anal loop in the hind wings of the two paratypes consists of three cells. Mr. De Marmels kindly informed me, under date of February 8, 1982, that all the Odonata from the Racenis collection formerly in Caracas have been transferred to the Instituto de Zoolo- gia Agricola in Maracay. The holotype of Zono- phora regalis Belle, consequently, is now in that institution. Zonophora supratriangularis Schmidt, 1941 (figs. 14, 56, 75) Zonophora supratriangularis Schmidt, 1941: 77—78, 80—81, 83, 88: figs. 1, 2a, 4b, 6b, 7b, 8b, 9c, 10b—c, 11d, 12b (6, ©). Belle, 1963: 61—63, 65 (spuratriangularis is a misprint for supratriangula- ris), 69. Racenis, 1966: 8. St. Quentin, 1973: 338. Material. — Brazil: Amazonas, Manaus (Manáos), 17 July 1976, 1 2, collector unknown; same locality, 9 August 1977, 1 d, A. Castrilon leg., CM. — Venezu- ela: Territorio Federal Amazonas, San Fernando de Atabapo, 24 February 1957, 1 6, J. Racenis leg., IZM. Schmidt (1941) described this species after five males and three females from several locali- ties in Brazil. He again (see Z. wucherpfennigi, antea) did not designate a holotype. After Schmidt, in case of type designation in the fu- ture, preference should be given to the male from Borba since the pair from this locality has served him to furnish most of the figures of this species. Of the male from Borba illustrations of the wings, caudal appendages and accessory genitalia are given, but of the male from Tefé depictions of the whole insect (in side view) and of the basal abdominal segments (in dorsal view) are offered. In my opinion, there is no principle objection to select the male from Tefé as lecto- type. Lectotype designation of Zonophora supra- triangularis Schmidt. — There are five “coty- pes”, three males and two females, of this spe- cies among the material of the Ris collection in the Natur-Museum Senckenberg, Frankfurt- am-Main. Each of the specimens is stored in a triangular envelope with the printed label “det. Dr. Erich Schmidt”. The five envelopes possess the Senckenberg’s catalogue numbers 24025 till 24029. Of these syntypes, the male numbered 24026 is here designated as the lectotype of Zonophora supratriangularis Schmidt, 1941. The lectotype has the locality data: Brazil, Am- azonas, Tefé (Ega), IX.1921, A. H. Fassl leg. The specimen has served for the above men- tioned figures (Schmidt’s figures 4b and 7b) and it formerly has been lent to me for study (cf. Belle, 1963). Z. supratriangularis is new to the fauna of Venezuela. If compared with the Brazilean ex- amples of this species, the male from Venezuela is a darker specimen. The yellow markings along the ventral tergal margins of the abdomi- nal segments 4 to 9 are much reduced and very narrow; they are well-developed and broad in the other examples. The superior caudal ap- pendages are black for their proximal one-third; those of the male from Manaus are black at the extreme bases only. The caudal appendages (stylets) of the female from Manaus are entirely yellow. The present males have three cubito-anal cross-veins (in addition to the inner side of the subtriangle) in the left fore wing, two in the BELLE: Zonophora 159 other wings. The female has two in each of its wings. All wings have a traversing cross-vein in the supratriangle except for the left fore wing of the female which has none. Zonophora nobilis spec. nov. (figs. 57—66) Material. — Brazil: Amazonas, Taracuá (Uaupes River), 14 August 1964, 1 d (holotype), Machado & Pereira leg., CM. This species is most nearly allied to Z. supra- triangularis and it agrees with it in having the supratriangles traversed by a cross-vein. In the male sex it differs from supratriangularis in the following particulars: (1) Superior caudal appendages almost three times as long as mid-dorsum of tenth abdominal segment; in supratriangularis, twice. (2) Internal tooth of each superior caudal ap- pendage at three-fifths the appendage’s length; in supratriangularis, at half its length. (3) Inferior lobe of anterior hamules long and slender; short and broad in supratriangularis. (4) Inner margin of posterior hamules more strongly and more distally elbowed than in supratriangularis. The female is unknown. Male (holotype; much broken and missing some details such as labium, antennae, hairs of occipital crest, left posterior hamule, left superi- or caudal appendage). — Total length, 50 mm; abdomen, 40 mm (incl. caud. app., 3.1 mm); hind wing, 34 mm; costal edge of pterostigma in fore wing, 3.5 mm, in hind wing, 4.0 mm. Face black, the following yellow: A medially interrupted stripe parallel to free border of mi brum; a medially interrupted stripe on antecly- peus; and a complete cross-stripe on postcly- peus covering fronto-clypeal suture. Mandibles externally yellow, the tips black. Superior sur- face of frons black except for two widely sepa- rated anterior spots of yellow. Vertex black, the conical prominences low. Occipital plate black- ish brown, its posterior margin evenly concave. Rear of head black except for a brown median spot on occiput. Prothorax dark brown, with mid-dorsal twin- spot of yellow and a larger yellow spot at either side. Pterothorax dark brown with yellow stripes; its colour design shaped as shown in the diagram. Femora dark brown, the inner sides yellow. Tibiae, tarsi and claws blackish brown. Wings hyaline, the venation dark brown in- cluding frontal margin of costa. Pterostigma brown, surmounting 4V— 5% cells. Brace vein present. No basal subcostal cross-vein. Nodal index 12 : 19—19 : 10/12 : 14—14 : 12. Second primary antenodal cross-vein the eighth in fore wings, the seventh in hind wings. Intermedian cross-veins 8—7/5—4 in fore and hind wings, respectively. One extra cubito-anal cross-vein in each wing. Supratriangles and discoidal trian- gles two-celled. Subtriangles open. Hind wings with a three-celled anal triangle, six paranal cells, four postanal cells, and area posterior to Cu2 four to five cells wide. Anal loop in hind wings open but small and not distinct, A2 not angulated but curved. Abdomen black with yellow markings as fol- lows: Sides of 1 with large yellow marking; mid-dorsum of 2 with lanceolate yellow spot, the spot becoming wider at apex of segment, sides of 2 with large yellow marking interrupted by black behind auricle, margin of auricles black; sides of 3 with yellow marking bordering on whole ventral tergal margin, the marking be- ing enlarged just in front of supplementary transverse carina; sides of 4 to 6 with yellow marking reaching to supplementary transverse carina, the markings becoming smaller on 4, 5 and 6 squseessitnelyys basal half of 7 yellow, the yellow marking continued along ventral tergal margin, the supplementary transverse carinae black; sides of 8 with yellow basal marking, the marking continued along ventral tergal margin; segments 9 and 10 black. Superior caudal ap- pendages pale yellow, becoming dark brown at both ends. Inferior caudal appendage black. Genital hamules and vesicle black. Caudal ap- pendages and accessory genitalia shaped as shown in accompanying figures. THE SPECIES IN THE MIMEOGRAPHED LIST OF RACENIS Racenis, in his mimeographed list of 1966, re- corded four species of Zonophora from Venezu- ela. Three are indicated to be new and of these, two have been named (cruxenti and medinai). It is impossible to make out from the list which known species must be associated with these two names since Racenis did not discuss them. Mr. De Marmels wrote to me on July 29, 1981: “I have almost all of his (Racenis) handwritten manuscript notes, which later were published (or not), and also his collection catalogue. I can easily see what name he thought to put to which 160 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) Table 1. Distribution of Zonophora species and subspecies Species and subspecies a (oi TN > = È Eben Bil 5 5 = O 3 a E 3 2 S CNE el 5 a S IS = TD © = 5p = N v = N i E SS os 3 = © on = ao > © 5 a u uM 5 > © u ©) = S © 3 = Wes he ey n > batesi Selys x Rolex x calippus calippus Selys x x x calippus klugi Schmidt ou ex Ri calippus spectabilis Campion IES x campanulata annulata subspec. nov. x campanulata campanulata (Burmeister) x campanulata machadoi St. Quentin x diversa spec. nov. x x nobilis spec. nov. x regalis Belle x solitaria obscura Belle x solitaria solitaria Râcenis x supratriangularis Schmidt x x surinamensis Needham x x wucherpfennigi Schmidt species. .. medinai in the list became solitaria when published, and cruxenti in the collection catalogue became regalis Belle. He himself cor- rected the names in his catalogue.” The mimeographed list seems to have been composed for own use although copies ot it have been distributed among some workers. The list has been mentioned in a dedication arti- cle of Dr. Janis Rácenis by De Marmels (1980) but it is difficult to believe that it must be con- sidered an official publication. HABITAT AND BEHAVIOUR IN THE FIELD There is hardly any environmental and be- havioural information available about Zonopho- ra species. My experiences with regard to Zono- phora have been confined to the two species batesi and calippus calippus only. I found them at the small shady creeks of the gallery forests in the savannah zone of Surinam. They were never encountered near stagnant water and also never during my collecung trips along the large streams of Surinam. For notes on their behav- iour in the field reference is made to the paper by Belle (1963: 65, 67). GEOGRAPHIC DISTRIBUTION The genus Zonophora is limited in its geo- graphic distribution to the Neotropical Region with the greatest number of species in Brazil (table 1). The most favourable habitat is appar- ently the tropical rainforest. The type localities of the 15 species and sub- species are grouped as follows: Brazil 11, Vene- zuela 3, Paraguay and Peru each 1 (see table 2). The campanulata group (fig. 67). The range of this group covers the southern part of Brazil and Paraguay, and the north-eastern part of Ar- gentina. These parts do not fall within the area of the tropical rainforest. The calippus group (fig. 68). This group has the largest range of all Zonophora groups. Its range extends from the northern coast of the continent to the southern part of Paraguay. The ranges of the members of the calippus group do not seem to overlap each other. The batesi group (fig. 69). The two members of this group are primarily Amazonian in their distribution but Z. batesi also occurs along the northern coast of the continent. The supratriangularis group (fig. 70). This group is not merely Amazonian in its distribu- tion. Z. supratriangularis is also recorded from the Orinoco River in Venezuela (San Fernando de Atabapo). The surinamensis group (fig. 71). Till now, all members of this group have been collected in the northern coastal region of the continent but the range may be larger. In Rapoport’s paper of 1968, continental South America has been divided into two sub- regions, Guayano-Brasilena and Andino-Pata- gónica. The distribution of the genus Zonopho- ra lies within the sub-region Guayano-Brasile- BELLE: Zonophora na. Using the zoogeographic division of conti- nental South America after Sclater & Sclater (cf. Rapoport, 1968: 68) we can say that the mem- bers of the campanulata group occur in the Sud- brasileno region, the bates: group and calippus group mainly in the Amazónica region, and that both groups penetrate into the Colombiana re- gion as far as the northern coast (Venezuela and the Guyanas), while the latter group also pene- trates into the western part of the Sud-brasileno region. The supratriangularis group occurs mainly in the Amazonica region and the swrina- mensis group in the Colombiana region. The two cordilleras in the north-west of the conti- nent seem to form a natural barrier for the dis- tribution of Zonophora to the tropical rainforest of western Colombia and Central-America. REFERENCES Beatty, A. F., 1956. An inquiry into the significance of the larval proventriculus in the taxonomy of Odonata. — Proc. Xth. Intern. Congress of Ent. Montreal. I: 367—372. Belle, J., 1963. Dragon flies of the genus Zonophora with special reference to its Surinam representa- tives. — Stud. Fauna Suriname 5: 60—69; pls. 3— 4. ——, 1966. Additional notes on some dragon-flies of the genus Zonophora. — Stud. Fauna Suriname 8: 61—64; pl. 11. ——, 1972. Further studies on South American Gom- 161 phidae (Odonata). — Tijdschr. Ent. 115: 217— 240. , 1976. Three new gomphine species from Vene- zuela (Anisoptera: Gomphidae). — Odonatologi- ca 5: 197— 206. ‚ 1977a. Revisional notes on Diaphlebia Selys, 1854 (Anisoptera: Gomphidae). — Odonatologica 6: 111-117. ——, 1977b. Some gomphine material from Surinam, preserved in the Leyden Museum of Natural His- tory, with a note on the larva of Desmogomphus tigrivensis Williamson (Anisoptera: Gomphidae). — Odonatologica 6: 289-292, ——, 1979. An attempt at the subfamily classification of the Gomphidae, based on some new interpreta- tions of wing venation. — Odonatologica 8: 43— 46. Bullens, A. M., 1966. De fylogenie van de Odonata op basis van de larvale kauwmaagbetanding. — Natuurh. Maandbl. 55: 88— 95. Burmeister, F., 1839. Handbuch der Entomologie II (2), Berlin: 847— 862. Byers, C. F., 1939. A study of the dragonflies of the genus Progomphus (Gomphoides) with a descrip- tion of a new species. — Proc. Fla Acad. Sci. 4: 19—85. Calvert, P. P., 1898. Burmeister’s types of Odonata. — Trans. Am. ent. Soc. 25: 27—104; pl. 1. — — 1948. Odonata (dragonflies) of Kartabo, Bartica District, British Guiana. — Zoologica, N.Y. 33: 47—87; pls. 12. ——., 1956. The neotropical species of the “subgenus Aeschna” sensu Selys, 1883 (Odonata). — Mem. Am. ent. Soc. 15: 1—251. Table 2. Alphabetic list of the species and subspecies of Zonophora, with type localities, type status and type location. Species and subspecies Type locality Type (all males) holo lecto neo location batesi Selys Fonte Boa, Amazonas’), Brazil x IRSN calippus calippus Selys Santarém, Pará, Brazil x IRSN calippus klugi Schmidt Mishiyacu, Peru x SMF calippus spectabilis Campion Sapucay, Paraguay x BM campanulata annulata subspec. nov. Jatai, Gioás, Brazil x MP campanulata campanulata (Burmeister) “Brazil” lost x IRSN campanulata machadoi St. Quentin Serra do Cipo, Minas Gerais, Brazil x CM diversa spec. nov. Nova Teutonia, Santa Catarina, Brazil x CC?) nobilis spec. nov. Taracua, Amazonas, Brazil x CM regalis Belle La Ceiba del Ventuari, Venezuela x IZM solitaria obscura Belle Sta. Elena, Venezuela x IZM solitaria solitaria Racenis Guayaraca-Auyantepui, Venezuela x IZM supratriangularis Schmidt Tefé, Amazonas, Brazil x SMF surinamensis Needham Mapaoni, Território do Amapa, Brazil x CU wucherpfennigi Schmidt Manicoré, Amazonas, Brazil x @AS) 1) Because of practical reasons the states have been quoted following the locality names in Brazil. 2) Will be deposited in the National Museum of Natural History, Washington, D.C. 3) Collection Asahina, Tokyo. 162 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) 2 5 Figs. 1—5. Zonophora batesi Selys, larva: 1, proventriculus; 2, first (left) dental fold, seen in oblique direction from right; 3, second dental fold, the same; 4, third dental fold, the same; 5, fourth dental fold, seen in oblique direction from left. Campion, H., 1920. Some new or little known gom- phine dragonflies from South America. — Ann. Mag. nat. Hist. (9) 6: 130—141; pls. 6—7. Carpenter, G. H., 1897. The geographical distribution of dragonflies. — Scient. Proc. R. Dubl. Soc. 8: 439— 468; pl. 17. Davies, D. A. L., 1981. A synopsis of the extant gen- era of the Odonata. — Soc. int. odonatol. rapid Comm. 3: xiv + 59 pp. De Marmels, J., 1980. To Dr. Janis Racenis on his 65th birthday. — Odonatologica 9: 125—129. ——, 1983. The Odonata of the region of Mount Auyantepui and the Sierra de Lema, in Venezue- lan Guyana. 3. Additions to the families Gomphi- dae, Aeshnidae and Corduliidae, with description of Progomphus racenisi spec. nov. — Odonatolo- mea 123 Sl Dias dos Santos, N., 1970. Odonatas de Iiatiaia (Esta- do do Rio de Janeiro) da colecao Zikan, do Insti- tuto Oswaldo Cruz. — Atas. Soc. Biol. Rio de Janeiro 13 (5 & 6): 204. Fraser, F. C., 1940. A comparative study of the penes of the family Gomphidae (order Odonata). — Trans. R. ent. Soc. Lond. (A) 90: 541— 550; pls. 1—6. ——, 1957. A reclassification of the order Odonata. — R. zool. Soc. N.S.W., Sydney: 133 pp.; 1 pl. Geijskes, D. C., 1971. List of Odonata known from French Guyana, mainly based on a collection brought together by the Mission of the “Museum National d’Histoire Naturelle”, Paris, (1) (2). — Ann. Soc. ent. Fr. (N.S.) 7 (3): 655—677. Hagen, H. A., 1861. Synopsis of the Neuroptera of North America. — Smithsonian Inst., Washing- BELLE: Zonophora 163 nn 14 15 16 17 18 Figs. 6—8. Penes of Zonophora, right lateral view: 6, Z. campanulata campanulata (Burmeister); 7, Z. calıppus calippus Selys; 8, Z. batesi Selys. Figs. 9—14. Left anterior hamules of Zonophora, right lateral view, showing superior lobe (a), inferior lobe (b), and internal lobe (c): 9, Z. campanulata campanulata (Burmeister); 10, Z. calippus calippus Selys; 11, Z. calippus klugi Schmidt; 12, Z. calippus spectabilis Campion; 13, Z. batesi Selys; 14, Z. supratriangularis Schmidt (more enlarged). Figs. 15—18. Vulvar laminae of Zonophora, ventral view: 15, Z. campanulata campanulata (Burmeister); 16, Z. calippus calippus Selys; 17, Z. batesi Selys; 18, Z. solitaria obscu- ra Belle, first described female. 164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) Figs. 19—22. Vulvar laminae of Zonophora, ventral view, showing variability: 19 and 20, Z. campanulata cam- panulata (Burmeister); 21, Z. campanulata machadoi St. Quentin, first described female; 22, Z. campanulata annulata subspec. nov, female allotype. Figs. 23—25. Diagram of thoracic colour pattern of Zonophora: 23, Z. campanulata campanulata (Burmeister); 24, Z. campanulata machadoi St. Quentin, male holotype; 25, Z. cam- panulata annulata subspec. nov., male holotype. Figs. 26—30. Zonophora surinamensis Needham, male: 26, di- agram of thoracic colour pattern; 27, penis guard, frontal view; 28, the same, right lateral view; 29, penis, right lateral view; 30, left anterior hamule, right lateral view. BELLE: Zonophora 165 Figs. 31—34. Zonophora campanulata campanulata (Burmeister), male: 31, accessory genitalia (penis and penis guard not drawn), ventral view; 32, same, right lateral view; 33, tenth abdominal segment and caudal append- ages, left lateral view; 34, the same, dorsal view. Figs. 35—36. Zonophora campanulata machadoi St. Quentin, left lateral view of abdomen: 35, male holotype; 36, first described female. Figs. 37—38. Zonophora campanula- ta annulata subspec. nov.. left lateral view of abdomen: 37, male holotype; 38, female allotype. 166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) 47 48 49 50 Figs. 39—50. Zonophora diversa spec. nov., male holotype: 39, accessory genitalia (penis and penis guard not drawn), ventral view; 40, the same, right lateral view; 41, tenth abdominal segment and caudal appendages, left lateral view; 42, the same, dorsal view; 43, abdomen, left lateral view; 44, left anterior hamule, right lateral view; 45, the same, seen in oblique direction from right; 46, diagram of thoracic colour pattern; 47, penis guard, fron- tal view; 48, the same, right lateral view; 49, occipital plate; 50, penis, right lateral view. BELLE: Zonophora 167 DIS Figs. 51—54. Zonophora solitaria solitaria Rácenis, male holotype: 51, diagram of thoracic colour pattern; 52, face and frons, seen in oblique direction from front; 53, abdominal segments 7 and 8, dorsal view; 54, frons, vertex and occipital plate. (Figs. 51—54, after J. de Marmels, June 1981.) Fig. 55. Zonophora solitaria obscura Belle, first described female: Occipital plate. Fig. 56. Zonophora supratriangularis Schmidt: Left posterior ham- ule, ventral view. ton: XX + 347 pp. , 1875. Synopsis of the Odonata of America. — Proc. Boston Soc. nat. Hist. 18: 20—96. Karsch, F., 1890. Ueber Gomphiden. — Entom. Nachr. 16:370—382. Kennedy, C. H., 1928. Evolutionary level in relation to geographic, seasonal and diurnal distribution of insects. — Ecology 9 (4): 367—379. Kiauta, B., 1979. The karyotypes of some Anisoptera from Surinam. — Odonatologica 8: 267—283. , 1981. Annotated catalogue and bibliography of taxa introduced in Odonata trom 1971 to 1980. — Kimmins, D. E., 1969. A list of the type-specimens of Odonata in the British Museum (Natural Histo- ry), Part 2. — Bull. Brit. Mus. (Nat. Hist.) 23: 287—314. Kirby, W. F., 1890. A synonymic catalogue of Neu- roptera Odonata or dragonflies. — Gurney & Jackson, London: IX + 202 pp. Navas, L., 1914-1923. Neuröpteros colombianos. — Boll. Soc. Cienc. nat. (Bogota), 1914: 141148; 1916: 6—13, 21—29, 67—73, 105—110, 119— 121, 137—143, 150—154; 1919: 137—144; 1920: 169—176; 1922: 201—208, 233—240; 1923: 265—272, 279—280. Needham, J. G., 1940. Studies on Neotropical gom- phine dragonflies (Odonata). — Trans. Am. ent. Soc. 65: 363— 394; pls. 20—22. ——, 1944. Further studies on Neotropical gomphine dragonflies (Odonata). — Trans. Am. ent. Soc. 69: 171224; pls. 14—16. Paulson, D. R., 1977. Odonata. In: S. H. Hurlbert, Ed., Biota acuatica de Sudamérica Austral: 170— 184. San Diego St. Univ. Räcenis, J., 1959. Lista de los Odonata del Perú. — Acta biol. venez. 2: 335—367. —, 1966. Preliminary list of Venezuelan Odonata. Inst. Zool. trop., Univ. Centr. Venezuela, Cara- cas. , 1970. Los odonatos de la region del Auyantepui y de la Sierra de Lema, en la Guyana venezolana. 2. Las familias Gomphidae, Aeshnidae y Cordulu- dae. — Acta biol. venez. 7: 23—39. Rapoport, E. H., 1968. Algunos problemas bio- geográficos del nuevo mundo con especial referen- cia a la region neotropical. — Biol. Amer. aust. 4: 53—110. Schmidt, E., 1941. Revision der Gattung Zonophora Selys (Odonata Gomphidae neotrop.). — Dt. ent. Z., 1941: 76—96. —, 1952. Odonata nebst Bemerkungen über die Anomisna und Chalcopteryx des Amazonas-Ge- biets. — Beitr. Fauna Perus, Jena 3: 207—256, 4 pls. St. Quentin, D., 1938. Die Tibialleiste der Odonaten. — Zool. Anz. 121 (9/10): 225—239. — —, 1967. Die Gattung Gomphoides Selys (Ordnung Odonata) und ihre Verwandten in der neotropi- 168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) 63 Figs. 57—66. Zonophora nobilis spec. nov., male holotype: 57, apical segments of abdomen, right lateral view; 58, inferior caudal appendage, dorsal view; 59, penis guard, frontal view; 60, the same, right lateral view; 61, tenth abdominal segment and caudal appendages, dorsal view; 62, diagram of thoracic colour pattern; 63, penis, right lateral view; 64, left anterior hamule, right lateral view; 65, accessory genitalia, right lateral view; 66, the same, ventral view. schen Region. — Beitr. neotr. Fauna 5 (2): 132— 152. ——, 1973. Die Gomphidenfauna Südamerikas. — Annln naturh. Mus. Wien 7: 335—363. Selys Longchamps, E. de, 1854. Synopsis des Gom- phines. — Bull. Acad. r. Belg. 21 (2): 23—112 (3—93 sep.). ——, 1869. Secondes additions au synopsis des Gom- phines. — Bul.. Acad. r. Belg. (2) 28: 168—208 (5—45 sep.). ——, 1873. Appendices aux troisièmes additions et liste des Gomphines, décrites dans le synopsis et ses trois additions. — Bull. Acad. r. Belg. [2] 36: 492—531 (47—87 sep). Selys Longchamps, E. de & H. A. Hagen, 1858. Mo- nographie des Gomphines. — Mém. Soc. r. Sci. Liège 11: 257—720 (VIII + 460 pp. sep.); 23 pls; 5 tabs. Tillyard, R. J. & F. C. Fraser, 1938—1940. A reclassi- fication of the order Odonata. Based on some new interpretations of the venation of the dragonfly wing. — Aust. Zool. 9: 125—169; 195221; 359— 396. Williamson, E. B., 1920. A new gomphine genus from British Guiana with a note on the classification of the subfamily (order Odonata). — Occ. Pap. Mus. Zool. Univ. Mich. 80: 1—12; pl. 1. 169 VSUYAIQ VLVINNNV *9 IOCVHOYVN ‘9 VLVINNVANVO ‘9 BELLE: Zonophora “dnoa8 vivynurdwvs ayı jo uonnqinsiQ ‘9 “317 SITIAVIIUdIS IDATA SAddITVOI ‘dn013 snddijv> aya zo uonnqinsiq ‘89 ‘8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) 170 ISINNYAIUHHONM © Isaıva @ ‘dn033 159179 ayı Jo uonnquusigg ‘69 “BIJ ‘dnois szurpnäuviuvidns su Jo uonnquasig '0/ ‘814 sITIaon & SIHVINONVIULVHANS @ BELLE: Zonophora 171 SURINAMENSIS S. SOLITARIA S. OBSCURA REGALIS Fig. 71. Distribution of the surinamensıs group. 172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 8 (1983) AR =. = FATTER CA peas LQ ISES ORS IS OOS as. gen fen fante2 EEE Ts F TZ ELLE à ST CREER IO N LE RSR Ra TE NS EEE ELLE ER AE — ui Ty Dax: CT SE is eee > Di LJ PI TS SAN nor So Figs. 72—74. Right pair of wings of male: 72, Zonophora campanulata campanulata (Burmeister); 73, Zonopho- ra calippus calippus Selys; 74, Zonophora batesi Selys. BELLE: Zonophora Q IX KES Be ESS co dr TJ SEE > È LITI CARRE toe a a A Rd 8 sores PER ES OSS 173 Figs. 75—76. Right pair of wings of male: 75, Zonophora supratriangularis Schmidt; 76, Zonophora surinamen- sis Needman. an DEEL 126 AFLEVERING 9 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD C. VAN ACHTERBERG. — Six new genera of Braconinae from the Afrotropical Region (Hymenoptera, Braconidae), pp. 175— 202, figs. 1—147. Tijdschrift voor Entomologie, deel 126, afl. 9 Gepubliceerd son | NE ps 7 # SIX NEW GENERA OF BRACONINAE FROM THE AFROTROPICAL REGION (HYMENOPTERA, BRACONIDAE) by C. VAN ACHTERBERG Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands ABSTRACT The new tribe Adeshini is defined for the genera Adesha Cameron, 1912, and Adeshoides gen. nov. (type-species: À. asulcatus sp. nov. from Senegal). The genera with a facial protu- berance in the Braconinae (excluding the Atanycolus group with petiolate pedicellus) are keyed and (re)described. Three new genera from the Afrotropical region are described, viz., Malagopsis (type-species: Plaxopsis grandidieri Szépligeti, 1913), Deltaphyma (type-spe- cies: D. horstoki sp. nov. from South Africa), and Zanzopsis (type-species: Plaxopsis liogas- ter Szépligeti, 1913). Bracon nigriceps Brullé, 1846, is a new junior synonym of Lasiophorus lanceolator (Fabricius, 1804) and Lasiophorus seyrigi Granger, 1949, is synonymized with Malagopsis grandidieri (Szépligeti, 1913) comb. et syn. nov. The genus Plaxopsis Szépligeti, 1905, is removed from the synonymy with Lasiophorus Haliday, 1833, and a lectotype is designated for Lasiophorus seyrigi Granger, 1949, and Plesiobracon carinatus Cameron, 1903. Finally the Plesiobracon group is defined, the known genera are keyed and two new genera are described: Carinibracon (type-species: C. damielssoni sp. nov. from Senegal) and Kenema (type-species: K. quicker sp. nov. from Sierra Leone). Odontogaster Szépligeti, 1906, is a new junior synonym of Soter Saussure, 1892. In this paper some recently discovered new genera of the Braconinae from the Afrotropical region are described, illustrated and keyed. Among these are two new genera collected dur- ing the Gambia-Senegal Expedition of the Lund University in 1977, kindly sent on loan by Dr. R. Danielsson (Lund). Both new genera have a sistergroup in the Oriental region as far as can be deduced from our limited knowledge of the genera of Braconidae. Dr. D. Quicke (Notting- ham) kindly allowed me to describe a related new genus from Sierra Leone. During my visits to the Berlin Museum (1979, 1982) I found several genera from the Afrotrop- ical region which at present are included in the genus Plaxopsis. These genera are (re)described in this paper, together with a new genus from South Africa found in the collection of the Rijksmuseum van Natuurlijke Historie at Lei- den. For the terminology used in this paper, see Van Achterberg, 1979: 242—249. ADESHINI tribus nov. Diagnosis. — Contains the only known Bra- coninae with vein CUla of fore wing at same level as vein 2-CU1 (fig. 4) and vein CU1b of fore wing much longer than vein 3-CUI (a syn- apomorphy within the Braconinae); scapus 175 truncate apically (fig. 3); mesoscutum at least sparsely setose; vein r-m of hind wing very short (fig. 4); propodeum with complete longi- tudinal carina (fig. 11); 1st tergite movably con- nected to 2nd tergite; 2nd and 3rd metasomal tergites with no antero-lateral diverging grooves (fig. 1). Contains two genera: Adesha Cameron, 1912 (Oriental region) and Adeshoides gen. nov. (Af- rotropical region). The biology is unknown. KEY TO GENERA OF THE TRIBE A DESHINI NOV. 1. Mesoscutum with deep medio-posterior groove (fig. 146); laterope large and round (fig. 139); angle between veins 1-SR and C+SC+R of fore wing about 35° (fig. 140); head and mesosoma smooth; mesoscutum denselyasetose=e pre Adesha Cameron Mesoscutum without medio-posterior groove (fig. 11); laterope (virtually) absent (fig. 1); angle between veins 1-SR and C+SC+R of fore wing about 65° (fig. 7); head and mesosoma coriaceous; mesoscu- tum sparsely setose, mainly near the notauli Adeshoides gen. nov. Adeshoides gen. nov. Type-species: Adeshoides asulcatus sp. nov. 176 Etymology: Latin for resembling the genus Adesha. Gender: masculine. Diagnosis. — Head and mesosoma coria- ceous; scapus rather robust (fig. 3); apex of an- tenna with no spine (fig. 5); eyes glabrous and not emarginate (fig. 9); malar suture absent; mesoscutum sparsely setose (mainly near the notauli) and without medio-posterior depres- sion (fig. 11); pleural and mesosternal sutures smooth; antescutal depression virtually absent (fig. 1); metapleural flange absent; only anterior half of notauli impressed; scutellar sulcus nar- row (fig. 11); metanotum medially with weak longitudinal carina (fig. 11); propodeal spiracle round, situated submedially (fig. 1); angle be- tween veins 1-SR and C-SC+R of fore wing about 65° (fig. 7); vein 1-SR+M of fore wing straight; vein m-cu of fore wing far antefurcal and distinctly diverging from vein 1-M posteri- orly (fig. 4); vein 1-R1 of fore wing much lon- ger than pterostigma (fig. 4); hind wing setose basally; tarsal claws setose, with no lobe (fig. 6); ventral row of setae of hind tarsus indistinct; laterope (virtually) absent (fig. 1); dorso-lateral and dorsal (except basally: fig. 12) carinae of Ist tergite present; 2nd metasomal suture deep and smooth (fig. 12); 2nd tergite with no medio- basal area, with a weak medial carina and a pair of shallow parallel, sublateral grooves (fig. 12); 2nd-5th metasomal segments with no antero- lateral grooves and with distinct sharp lateral crease (fig. 1). Distribution. — Afrotropical: one species. Adeshoides asulcatus sp. nov. (figs. 1—12) Holotype, 6, length of body 2.5 mm, of fore wing 2.2 mm. Head. — Antennal segments 37, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 2.7 and 2.0 times their width, respectively; penultimate segment of antenna 2x longer than wide (fig. 5); length of maxillary palp 0.7 times height of head; length of eye in dorsal view 2.8 times temple (fig. 8); temples gradually narrowed posteriorly; POL : © ocel- lus : OOL = 5:4:10; frons weakly convex, coriaceous, with weak medial groove; face and clypeus rather convex and shiny coriaceous; ventral margin of clypeus not differentiated; length of malar space 0.8 times basal width of mandible. Mesosoma. — Length of mesosoma 1.4 times its height; episternal scrobe linear and rather shallow; mesoscutal lobes and scutellum rather TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 flat and coriaceous; posterior face of propo- deum rather differentiated and shorter than an- terior face. Wings. — Fore wing: r : 3-SR : SR1 = 5:13:35; veins M+CUT, EUR ES SR SM and 2-M distinctly wider than other veins; cu-a interstitials 2 SRE SRE rm AMIE: Legs. — Hind coxa smooth; length of femur, tibia and basitarsus of hind leg 4.6, 10.4, and 7 times their width, respectively; hind tibial spurs both 0.2 times hind basitarsus. Metasoma. — Length of 1st tergite 0.8 times its apical width, its surface largely rugulose-co- riaceous; dorsal carinae of 1st tergite nearly complete, but rather weak and not lamelliform (fig. 12); 2nd tergite coriaceous-rugulose; 3rd and 4th tergites shiny coriaceous; rest of meta- soma smooth and depressed; 4th and 5th ter- gites with smooth, deep anterior transverse groove (fig. 1). Colour. — Brownish-yellow; apical half of antenna and stemmaticum, dark brown; ptero- | stigma and veins (rather dark) brown; wing membrane slightly infuscated. Holotype in Entomological Museum, Lund: “Senegal, 3 km SSW Toubakouta, 10 km S Zi- guinchor, 4.111.1977, at light 19.00—22.00, Loc. No. 16, UTM 28PCJ 58 782”, “Lund Univ. Syst. Dept. Sweden, Gambia/Senegal, Febr.- March 1977, Cederholm, Danielsson, Larsson, Mireström, Norling, Samuelson”. Adesha Cameron, 1912 Cameron, 1912: 78; Shenefelt, 1978: 1430. Type-species: Adesha albolineata Cameron, 1912 (monotypic). Gender: feminine. Diagnosis. — Head and metasoma smooth; scapus rather robust (fig. 138); eyes glabrous and indistinctly emarginate (fig. 142); malar su- ture absent; mesoscutum densely setose and with deep medio-posterior groove (fig. 146); pleural and mesosternal sutures smooth; ante- scutal depression very narrow; metapleural flange small (fig. 139); anterior half of notauli | deep and smooth, rest shallow or absent (fig. 146); scutellar sulcus wide and distinctly crenulate (fig. 146); metanotum with distinct medio-longitudinal carina (fig. 146); propodeal spiracle rather small, round, and behind middle of propodeum (fig. 139); angle between veins 1- SR and C+SR+R of fore wing about 35° (fig. 140); vein 1-SR+M of fore wing straight; vein m-cu of fore wing moderately antefurcal VAN ACHTERBERG: Afrotropical Braconinae 177 and slightly diverging posteriorly from vein 1- M (fig. 141); tarsal claws setose, without lobe (fig. 143); ventral row of setae of hind tarsus distinct; laterope large and round (fig. 139); dorso-lateral carinae of 1st tergite distinct; dor- sal carinae on basal quarter of tergite, rest ab- sent (fig. 147); 2nd metasomal suture deep and crenulate (fig. 147); 2nd tergite with narrow tri- angular and smooth medio-basal area, bordered by crenulate grooves, laterally with a pair of posteriorly (weakly) converging depressions (fig. 147); 2nd-5th tergites with sharp lateral crease and without antero-lateral grooves (fig. 139); ovipositor normal, with neither no- dus nor teeth subapically. Distribution. — Oriental: one species. Adesha albolineata Cameron (figs. 138—147) Cameron, 1912: 78; Shenefelt, 1978: 1431. Holotype, 2, length of body 4.2 mm, of fore wing 3.8 mm. Head. — Antennal segments incomplete, 16 present, length of 3rd segment 1.5 times 4th segment, length of 3rd and 4th segments 3 and 2 times their width, respectively; length of maxil- lary palp 0.7 times height of head; length of eye in dorsal view 2.8 times temple; temples round- ly narrowed posteriorly (fig. 145); POL : © ocellus : OOL = 7 : 8 : 19; frons slightly con- vex, largely smooth and with medial groove (fig. 142); face weakly convex and smooth; cly- peus flat and smooth, its ventral margin not dif- ferentiated from clypeus, concave; length of malar space 0.8 times basal width of mandible. Mesosoma. — Length of mesosoma 1.7 times its height; side of pronotum smooth, except for the finely crenulate medial groove; mesopleu- ron smooth; episternal scrobe deep, medium- sized and round (fig. 139); mesoscutum rather convex, its middle lobe slightly impressed me- dially (fig. 146); scutellum weakly convex and somewhat pimply; propodeum largely smooth, with some short rugae near medial carina and its posterior surface not differentiated (fig. 146). Wings. — Fore wing: r : 3-SR : SRI = 5:10:38; veins of similar width; 1-CU1 : 2- ISO 172 SRE SRE Em AOR MORE Legs. — Hind coxa smooth and slender (fig. 139); length of femur, tibia and basitarsus of hind leg 4.1, 9.8, and 6.4 times their width, respectively; length of hind tibial spurs 0.2 and 0.3 times hind basitarsus. Metasoma. — Length of 1st tergite 0.8 times its apical width, its surface coarsely reticulate medially, rest smooth (fig. 147); 3rd tergite coarsely reticulate medially, as rest of metasoma (fig. 139); length of ovipositor sheath 0.20 times fore wing, widened apically; hypopygium medi- um-sized and truncate apically. Colour. — Brownish-yellow; antenna, dorsal part of head largely, meso- and metasoma dor- sally, mesosternum partly, ovipositor sheath, tarsi, patch on apex of hind tibia, hind femur dorsally, dark brown or black; metasoma laterally, and apex of Sth tergite yellowish; wing veins and pterostigma brown; wing membrane subhyaline. Holotype in British Museum (Natural Histo- ry), London: “Type”, “Kuching, J. H.” (= John Hewitt), “2.10”, “P. Cameron Coll, 1914- 110”, “Adesha albolineata Cam., Type, Bor- neo” (in Cameron’s handwriting). KEY TO THE GENERA OF THE BRACONINAE WITH FACIAL PROTUBERANCE (excluding the Atanyco- lus group) 1. Head with a protuberance anteriorly (figs. 13, 23, 35, 84); if rather cariniform and small, then in dorsal view (short) semi- circularly protruding; (some males of Magalopsis grandidieri (Szépligeu) have no protuberance but face is (narrowly) de- pressedimediallg RR 2 — Head without protuberance, at most cly- peus with a scarcely protruding, medially straight (in dorsal view), fine, dorsal carina; (face without medio-dorsal depression) .... Pedale da ei n other Braconinae 2. Pedicellus in dorsal view cylindrical, not narrowed basally; scapus rounded subba- sally (figs. 17, 32); inner aspect of scapus not excavated apically (fig. 38), except in Deltaphyma (fig. 56); length of fore tarsus lessichany IW7acinies to etiDIAEEPPREEERES 3 — Pedicellus in dorsal view petiolate, distinct- ly narrowed basally; scapus rather angular- ly protruding subbasally; inner aspect of scapus deeply excavated medio-apically; length of fore tarsus 1.7—2 times fore tibia be Ath 2 bosa. Atanycolus group 3. Middle of face with an irregularly incised, vertical lamella (fig. 13); antenna of 9 widened apically and with a blunt apex (figs. 13, 14); marginal cell of fore wing short, far removed from wing apex (fig. 15); vein 1-SR of fore wing short (fig. 15); tibial spurs dorsally glabrous and 178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 Mentrallasetose tre Victoroviella Tobias — Middle of face without vertical lamella (figs. 23, 58, 60); antenna of 9 not widened apically and with acute apex (figs. 24, 41); marginal cell of fore wing long, almost reaching wing apex (figs. 31, 70); vein 1-SR of fore wing medium-sized (figs. 31, 42, 59); tibial spurs evenly setose or nearly completely glabrous (at 50 x) .......... - 4. Face with protruding carına and/or lamella (figs. 35, 52, 77); clypeus with ventral cari- na (figs. 49, 77); mesoscutum largely gla- brous; hind spurs (partly) setose; medio- basal area of 2nd tergite robust (figs. 48, 65, 82); antero-lateral grooves of 3rd tergite (partly) crenulate (figs. 48, 93); (Afrotropi- ealfandisoudibalacaneus Peer — Face with robust horn above clypeus (figs. 23, 26 of type species, in other species spoon- shaped or tuberculiform); clypeus without distinct ventral carina (fig. 23); mesoscutum completely setose; hind spurs almost completely glabrous; medio-basal area of 2nd tergite very slender (fig. 34); antero-lateral grooves of 3rd _ tergite SMOOLHAUNEOTLO PCA) EE PAP EEE PEER Lasiophorus Haliday 5. Upper half of face protruding (figs. 58, 75) or slightly concave (fig. 87); posterior half of 4th and 5th tergites flat (figs. 57, 84); 1st tergite usually without dorsal carinae (figs. 65, 82), if present then not lamelli- form; vein Ir-m of hind wing straight (fig. 55); 2nd tergite with (sub)parallel de- pressions (figs. 65, 82, 93); vein 1-M of fore wing straight (figs. 55 A0 Os fee eliscal cell of fore wing less transverse (figs. 55, 91); lower facial protuberance of 2 dis- tinctly lamelliform or absent; (Continental Africa and South Palaearctical region) ... 6 — Upper half of face deeply concave (figs. 47, 49); posterior half of 4th and 5th tergites strongly convex (fig. 35); 1st tergite with lamelliform dorsal carinae (fig. 48); vein 1r- m of hind wing curved (fig. 39); 2nd tergite with (incomplete) diverging antero-lateral grooves (fig. 48); vein 1-M of fore wing somewhat bent posteriorly (fig. 42); 1st dis- cal cell of fore wing comparatively trans- verse (fig. 39); lower facial protuberance of ® sublamelliform and absent in d and ex- ceptionally in (small) 2 9 (Malagasy) ...... Malagopsis gen. nov. 6. Scapus truncate or at most moderately pro- truding ventrally, not beyond apex of pedi- cellus (figs. 69, 92); frons not or shallowly concave (figs. 74, 88); face with either a È | mella (figs. 84, 87) or a W-shaped protuber- | ance (figs. 75, 77); 2nd tergite with well-de- | fined basal area (figs. 82, 93); 2nd tergite | not projecting above 2nd suture (fig. 84); groove between eye and antennal socket rather shallow or absent (figs. 67, 84); | mesoscutum anteriorly at least as high as M the pronotum anteriorly (figs. 67, 84); scu- | tellum often with a small pit (fig. 76)..... 7 | — Scapus strongly protruding ventro-apically, | well beyond apex of pedicellus (fig. 54); | frons distinctly concave (fig. 62); face with | protruding lamella and with a triangular | protuberance above it (figs. 58, 60); tergite with ill-defined basal area (fig. 65); 2nd tergite medio-posteriorly projecting above 2nd suture (fig. 52); with a deep groove between eye and antennal socket (fig. 52); mesoscutum anteriorly lower than pronotum anteriorly (fig. 52); scutellum without smalll pit (fre N63) ARRE Deltaphyma gen. nov. 7. Face above clypeus with W-shaped protu- berance (figs. 75, 77); marginal cell of hind wing strongly narrowed apically (fig. 70) Zanzopsis gen. nov. — Face above clypeus without protuberance, only submedially with a more or less semi- circular (often minute) lamella (fig. 87); marginal cell of hind wing (sub)-parallel- sided or slightly narrowed apically (fig. 91) Plaxopsis Szépligeu Victoroviella Tobias, 1975 Tobias, 1975: 962—964. Type-species: Victoroviella deserticola Tobi- as, 1975. Diagnosis. — Scapus slightly longer ventrally than dorsally (fig. 13), curved, with a rounded anterior flange (figs. 13, 17) and subcylindrical; antenna of ® widened and rather compressed apically (fig. 13); apex of antenna blunt (fig. 14); face with longitudinal thin lamella, which has one deep medial and some smaller in- cisions (fig. 13), below this lamella a spoon- shaped, weakly concave protuberance (fig. 18); notauli absent; scutellar sulcus rather shallow and narrow; metanotum without carinae; vein 1-R1 (metacarp) of fore wing much shorter than pterostigma (fig. 15); fringe of wings very short; basal half of fore wing only sparsely se- tose; parastigma comparatively robust (fig. 15); 2nd # VAN ACHTERBERG: Afrotropical Braconinae 179 vein r-m of hind wing very short (fig. 15); mar- ginal cell of hind wing narrowed apically; tibial spurs dorsally glabrous and ventrally setose; tarsal claws large (fig. 20), setose, and with no lobe; dorsal and dorso-lateral carinae of 1st ter- gite absent; 2nd suture of metasoma deep, smooth and straight (fig. 21); 2nd and basal half of 3rd tergite with sharp lateral crease; 2nd and 3rd tergites with a pair of smooth, diverging grooves (fig. 21); ovipositor normal (fig. 19), with small dorsal nodus and ventral teeth. Distribution. — South Palaearctic: one spe- cies. The biology is unknown. Victoroviella deserticola Tobias (figs. 13—21) Tobias, 1975: 962—964, figs. 1—3. Paratype, ®, length of body 6.6 mm, of fore wing 5.0 mm. Head. — Antennal segments 34 (according to the original description 34—35), length of 3rd segment 1.6 times 4th segment, length of 3rd and 4th segment 2.1 and 1.3 times their width, respectively; length of penultimate segment 0.7 times its width (fig. 14); length of maxillary palp 0.6 times height of head; length of eye in dorsal view 1.3 times temple; temple subparallel (fig. 16) and long setose; POL : © ocellus : OOL = 3 : 12; frons flat and smooth; face with very long setae (fig. 13), remotely punctu- late and weakly convex; clypeus depressed ven- trally, its margin not differentiated; length of malar space 1.3 times basal width of mandible. Mesosoma. — Length of mesosoma 1.6 times its height; pronotal sides smooth (except for some punctures dorsally; fig. 13), and with a rather deep medial groove; mesopleuron smooth; episternal scrobe small, deep and round (fig. 13); mesoscutum and propodeum smooth; propodeal spiracle medium-sized, round, sub- medially situated (fig. 13). Wings. — r : 3-SR : SR1 = 3 : 11 : 16; 1- SR+M straight; cu-a subinterstitial (fig. 15); 2- SRB SR Siem — 16 22715; m-cu far ante- furcal and slightly converging to 1-M posteri- orly (fig. 15). Legs. — Hind coxa smooth; length of femur, tibia and basitarsus of hind leg 2.8, 7.1 and 5.2 times their width, respectively; length of hind tibial spurs 0.4 and 0.5 times hind basitarsus; hind tibia with much longer setosity dorsally than ventrally; fore tibia with a row of five short spines. Metasoma. — Length of Ist tergite 1.1 times its apical width, its surface smooth (fig. 21); glymma completely absent; 2nd and 3rd tergites smooth; 2nd tergite with a pair of short, smooth and converging grooves medio-anteriorly (fig. 21); rest of metasoma smooth and rather depressed; length of ovipositor sheath 1.19 times fore wing; hypopygium large and acutely protruding medio-apically (fig. 13). Colour. — Brownish-yellow; apex of mandi- bles, apex of antenna, wing veins partly, and pterostigma medio-anteriorly, dark brown; ovi- positor sheath black; pronotum dorsally, ante- rior subalar protuberance, middle of mesopleu- ron, mesonotum medially, scapus and pedicellus anteriorly, and face, largely whitish; wing mem- brane hyaline. Paratype in the Zoological Institute, Lenin- grad: “22/V. (19) 65), ksf. Kirpili, na strarinkus, T. Tokgaev”, “Paratypus Victoroviella deserti- cola Tobias”. Holotype in the Zoological Mu- seum at Moscow from Turkmenia (Repetek), not examined. Note. Not closely related to the other genera treated in this paper because of the peculiar shaped antenna, the longitudinal thin lamella of face, the shape of the scapus and the short mar- ginal cell of fore wing. Lasiophorus Haliday, 1833 Haliday, 1833: 213; Fahringer, 1930: 19 (key); Shene- felt, 1978: 1691—1694. Type-species: Bracon lanceolator Fabricius, 1804 (monotypy). Diagnosis. — Scapus very slender (fig. 23), widened apically and ventrally distinctly longer than dorsally (fig. 32); antennal sockets (toruli) protruding (figs. 23, 26); apex of antenna with short, not distinctly differentiated spine (fig. 35); face medially (just above clypeus) with a large, thick and rugose horn (figs. 23, 26, 27), or with spoon-shaped or tuberculiform protu- berance; clypeus with no ventral carina; notauli absent; mesoscutum completely setose; scutel- lar sulcus narrow and finely crenulate (fig. 28); metanotum with no medial carina; vein 1-R1 of fore wing much longer than pterostigma (fig. 31; as in the other genera); vein m-cu of fore wing shortly antefurcal, converging to vein 1-M posteriorly (fig. 31); vein 1-SR+M of fore wing straight, but basally weakly curved; vein r-m of hind wing long and straight (fig. 31); marginal cell of hind wing subparallel-sided api- cally (fig. 31); hind tibial spurs with indistinct 180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 and very short setosity, almost completely gla- brous (fig. 29); tarsal claws setose and with no lobe (fig. 33); hind tarsus with ventral row of setae; dorso-lateral carinae of 1st tergite distinct and complete; dorsal carinae of Ist tergite ab- sent (fig. 34); 2nd tergite anteriorly depressed and with a medial ridge instead of mediobasal area (fig. 34); antero-lateral grooves of 3rd and 4th tergites smooth and shallow, not reaching lateral margin of tergite (fig. 23); 2nd suture of metasoma distinct and smooth (fig. 34); no seg- ments (except Ist) with sharp lateral crease; ovi- positor normal (fig. 22), with small dorsal nodus and small ventral teeth. Distribution. — Neotropical: four species. The biology is unknown. Lasiophorus lanceolator (Fabricius) (figs. 22—34) Bracon lanceolator Fabricius, 1804: 106. Lasiophorus lanceolator; Shenefelt, 1978: 1692; Van Achterberg, 1982: 137 (lectotype designation). Bracon nigriceps Brullé, 1846, 334— 335. Syn. nov. Lasiophorus nigriceps; Shenefelt, 1978: 1692— 1693. Lectotype, ®, length of body 16.2 mm, length of fore wing 14.5 mm. Head. — Antennal segments 73, length of 3rd 1.7 times 4th segment, length of 3rd and 4th segments 2.2 and 1.3 times their width, respecti- vely; penultimate segment of antenna 1.2 times as long as wide; length of maxillary palp 0.9 times height of head; temples weakly narrowed posteriorly (fig. 27); POL : © of ocellus : OOL = 4:5:10; frons laterally flat, but medially narrowly depressed (fig. 27); face apart from the horn, rather concave, coriaceous, rather mat (fig. 26); length of malar space 0.9 times basal width of mandible. Mesosoma. — Length of mesosoma 1.7 times its height; pronotal sides smooth, except for some punctulation (fig. 23); meso- and meta- pleuron punctulate; episternal scrobe absent, except for an indistinct remnant; scutellar sulcus narrow and shallowly crenulate (fig. 28); meso- scutum and scutellum sparsely punctulate (fig. 28); surface of propodeum punctulate; pro- podeal spiracle elliptical, rather large and sıtu- ated behind middle of propodeum (fig. 23). Wings. — Fore wing: angle between 1-SR and C+SC+R about 35° (fig. 31); r : 3-SR : SRI = 8:43 : 58; cu-a postfurcal, long; 1-CUI1 : 2- CU = il PL PE SRE ORNE MIMO r-m sinuate (fig. 31). Legs. — Hind coxa punctulate; length of fe- mur, tibia, and basitarsus of hind leg 4.6, 12.9; and 8.6 times their width, respectively; length of hind tibial spurs 0.35 and 0.5 times hind ba- sitarsus; fore tibia with many spines (fig. 30). Metasoma. — Length of Ist tergite 1.7 times its apical width, its surface mainly smooth, punctulate medially and sublateral grooves cre- nulate anteriorly (fig. 34); glymma present, shallow (fig. 23); metasoma smooth behind the Ist tergite, rather compressed; length of ovipo- sitor sheath 2.08 times fore wing; hypopygium very large and truncate apically (fig. 23). Colour. — Yellowish-brown; head, antenna, propleura, coxae partly, hind leg (except tro- chanters partly, base and apex of femur, and basal half of tibia), 5th—7th tergites and ovipo- sitor sheath dark brown or blackish; palpi part- ly infuscated; wing membrane yellowish, but fore wing basally (shortly), apically and a band near pterostigma, dark brown (fig. 31); para- stigma and basal half of pterostigma, dar brown, but rest of pterostigma (light) brown. Lectotype in the Copenhagen Museum: “Type”, “Essequibo, Smidt, Mus: de Sehestedt, Bracon lanceolator Fabr.” (Essequibo is a river in Guyana). Paralectotypes: 2 ©, very similar to lectotype. In the Rijksmuseum van Natuurlijke Historie at Leiden are 11 © and 1 d. The male from Surinam (Avanavero, 25.v11.1973, D. C. Geijskes) has the facial horn about half the size of the ® and undivided apically, body more densely setose, and base of fore wing yellowish. The females originate from Trinidad (Tacariqua, 500—600 m, 10.ix.1953, G. F. Mees) and Suri- nam (Republiek, 23.xi.1963, D. C. Geijskes and 31.11.1963, P. H. van Doesburg; Kabalebo Riv- er, near Avanavero Falls, 14.1x.1965, and Brownsberg, 450—480 m, 16.xii.1973, both G. F. Mees; Mapane area, camp 8, 27 & 28.v.1963, and Zanderij, 21—25.vii.1963, both J. van der Vecht; Matta, 1.viii.1945; Wilhelminagebergte, 16.viii.1943; and Moengotapoe, 24.1x.1948, all D. C. Geijskes). Variation. — Length of fore wing 13.8—17.8 mm, of body 15.8—19.5 mm; length of oviposi- tor sheath 1.84—2.09 times fore wing; antennal segments 73—81. The known distribution 1s from Panama to Peru and Brazil. Note. The statement of Fabricius (1804: 106): “Aculeus excertus niger, longitudine corporis” leads to the misinterpretation of lanceolator by Fahringer. Actually the ovipositor is somewhat less than twice length of the body. A syntype of Bracon nigriceps Brullé, 1846, in the Paris Mu- VAN ACHTERBERG: Afrotropical Braconinae 181 seum has been examined. Another specimen may be present in the Spinola Collection (Turin) via Audinet-Serville. KEY TO SPECIES OF THE GENUS Lasiophorus HALIDAY (after Fahringer, 1930, modified) 1. Length of ovipositor sheath about twice length of fore wing; wings banded... 2 — Length of ovipositor sheath less than length of fore wing, about as long as metasoma; winesmmoticlearly banded tun 3 2. Middle of 1st—4th tergites black; (Guya- na); (dark form of lanceolator?) ........... ea en a. fortispinus Cameron — Middle of 1st—4th tergites yellowish; (Pan- amakosberuandWBrazil))....r.0:.0.20.0. 222. REED tan eren lanceolator (Fabricius) 3. Pterostigma dark brown or blackish; facial protuberance spoon-shaped, concave me- dially and truncate apically, regular; apex of metasoma blackish; (Ecuador) ............ PRU es ce aa à semirufus Fahringer — Pterostigma yellowish with its base and apex blackish; facial protuberance irregular tuberculiform; apex of metasoma yellow- ish; (Joinville Isl.) polaris Fahringer Malagopsis gen. nov. Type-species: Plaxopsis grandidieri Széplige- 913: Etymology: from the combination of “Mala- gasy” and “Plaxopsis” because this genus is close to Plaxopsis and is only known from Mala- gasy. Gender: masculine. Diagnosis. — Scapus medium-sized, ventrally strongly protruding (fig. 37) and inner aspect Li its apex not emarginate (fig. 38); apex of anten- na with short spine (fig. 41); face with a pair of large protuberances in front of antennal sockets (fig. 44); face very deeply concave medially, ventrally bordered by a sublamelliform semi- circular protuberance (absent in d and excep- tionally in small ©). Protuberance medially with a carina connected to a medial carina of the face (figs. 47, 49, 50); clypeus with a ventral carina; notauli shallowly impressed anteriorly (fig. 45); scutellar sulcus narrow and shallow; scutellum with no pit; metanotum with no medial carina; vein cu-a of fore wing slightly postfurcal; vein 1-M of fore wing somewhat bent posteriorly (fig. 42); vein m-cu of fore wing shortly ante- furcal, slightly converging to vein 1-M posteri- orly (fig. 39); vein 1-SR+M of fore wing bent basally; vein 1r-m of hind wing distinctly curved (fig. 39); marginal cell of hind wing par- allel-sided subapically; tarsal claws basally pec- tinate and with no lobe (fig. 46); hind tarsus with ventral row of setae; hind spurs setose; dorso-lateral carinae of 1st tergite lamelliform and complete (fig. 35); dorsal carinae of Ist ter- gite complete in posterior two-thirds of tergite, lamelliform and enclosing a long subpentagonal elevated area (fig. 48); 2nd tergite with smooth subtriangular medio-basal area, surrounded by sparsely crenulate grooves and with wide, pos- teriorly diverging antero-lateral grooves (fig. 48); 2nd suture obsolete laterally, rather deep and strongly crenulate medially (fig. 48); 3rd tergite with incomplete antero-lateral grooves; 2nd and 3rd tergites with sharp lateral crease; 4th and 5th tergites strongly convex (fig. 35); ovipositor with normal apex, with small nodus and teeth. Distribution. — Afrotropical (Malagasy): one species. Parasites of Anthribidae (Coleoptera). Malagopsis grandidieri (Szépligeu) comb. nov. (figs. 35—50) Plaxopsis grandidieri Szépligeti, 1913: 419. Lasiophorus grandidieri; Shenefelt, 1978: 1691. Lasiophorus seyrigi Granger, 1949: 34— 36, fig. 32; Shenefelt, 1978: 1694. Syn. nov. Holotype, 2, length of body 11.9 mm, of fore wing 10.5 mm. Head. — Antennal segments 64, length of 3rd segment 1.4 times 4th segment, length of 3rd and 4th segments 1.7 and 1.2 times their width, respectively; penultimate segment of antenna 1.3 times its width; length of maxillary palp 0.8 times height of head; groove between eye and antennal socket wide and rather deep (figs. 35, 47); eye slightly emarginate (fig. 47); length of eye in dorsal view 1.4 times temple; temples somewhat narrowed posteriorly (fig. 44); POL "O ocellus; = OOL = 4:3 285 trons, weakly, concave, with shallow medial groove (fig. 44); face convex laterally, coriaceous and depressed beside clypeus (fig. 47); length of malar space equal to basal width of mandible; malar suture distinct and narrow (fig. 35). Mesosoma. — Length of mesosoma 1.4 times its height; pronotal sides and mesopleuron smooth; episternal scrobe small, droplet-shaped (fig. 35); metapleuron punctulate; mesoscutum smooth and largely glabrous; surface of propo- deum punctulate laterally, rest smooth; propo- deal spiracle rather large, wide elliptical and sit- 182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 uated just behind middle of propodeum. Wings. — Fore wing: angle between veins 1- SR and C+SC+R of fore wing 58° (fig. 42); r : SOR 2 ON = 5223 2 De CUIR PE CUIR 18 B88 ZON 8 HSN Bien = iO) s 2s) IO) Legs. — Hind coxa smooth; length of femur, tibia and basitarsus of hind leg 2.9, 7.8 and 4.0 times their width, respectively; length of hind spurs 0.20 and 0.25 times hind basitarsus. Metasoma. — Length of Ist tergite 0.9 times its apical width, its surface largely smooth, me- dially with some wide punctures and weakly concave, and laterally with some coarse crenu- lae (fig. 48); glymma shallow and crenulate (fig. 35); metasoma behind 2nd tergite medially smooth, and 3rd—6th tergites punctate latero- posteriorly (fig. 35); length of ovipositor sheath 0.59 times fore wing; hypopygium large, and acute apically (fig. 35). Colour. — Yellowish-brown; head (except black middle of facial depression and of lamella, stemmaticum, and area behind it), and pronotal sides dorso-posteriorly, ivory; wing membrane brownish; pterostigma and veins rather dark brown. Holotype in Zoologisches Museum Berlin: “S. W. Madagascar, Tulear, Voeltzkow, S., 1.1904”, “Bracon grandidieri Sauss., det. Dr. Enderlein” (incorrect!), “Type”, “Plaxopsis grandidieri n. sp.” (in Szépligeti’s handwriting). Syntype-series of Malagopsis seyrigi (Granger, 1949) consists of 15 ® and 1 d. The 2 from Diego (“Madagascar, Diego, x1.29, A. Seyrig”) in the Museum National d'Histoire Naturelle at Paris is here designated as lectotype. Judging from this series of grandidieri there is consid- erable allometric variation in the size of the fa- cial lamella and flanges, and the punctation of the 5th and 6th tergites. The dark patch on the pronotum may be distinct but (especially larger) specimens have this patch absent or obsolescent. Variation. — Length of fore wing 7.9—10.5 m (6: 7.9 mm); length of body 8.8—11.9 mm (d: 8.8 mm); length of ovipositor sheath 0.59— 0.73 times fore wing; antennal segments 57—66 (d: 55); facial depression deep or moderately depressed (always distinct in 2, shallow in gd); facial protuberances large to nearly absent (ab- sent in d); 3rd sternite completely yellowish or with blackish patch, sometimes also 2nd and 4th sternites with such patch. Deltaphyma gen. nov. Type-species: Deltaphyma horstoki sp. nov. Etymology: from “delta” (Greek capital let- M ter, shaped like a triangle) and “phyma” (Greek | for “growth”) because of the peculiar triangular | protuberance of the face. Gender: neuter. | Diagnosis. — Scapus robust, ventrally much M longer than dorsally, surpassing pedicellus M (fig. 54) and inner aspect of scapus distinctly M emarginate (fig. 56); face convex, with spoon- M shaped lamella, with above it a triangular lamel- | la, with its corners (especially the top corner) M longer than its sides (figs. 58, 60); clypeus with- | out carinae; with deep groove from eye to an- | tennal socket (fig. 52); notauli complete, shal- M low (fig. 63); scutellar sulcus very shallow and | narrow, nearly smooth (fig. 63); scutellum with | no pit; metanotum without carina; vein cu-a of M fore wing subinterstitial; vein 1-M of fore wing | straight (fig. 55); vein m-cu of fore wing shortly | antefurcal, slightly converging to vein 1-M pos- | teriorly; vein 1-SR+M of fore wing evenly | curved (fig. 59); vein 1r-m of hind wing straight | (fig. 55); marginal cell of hind wing parallel- |! sided apically; tarsal claws yellowish pectinate | basally, without lobe (fig. 66); hind femur | densely setose dorsally, ventrally sparsely, | without distinct flange apically (fig. 64); hind | tarsus without distinct ventral row of setae; hind spurs setose; dorso-lateral carinae of 1st tergite complete and lamelliform behind the spi- | racle, absent in front of spiracle (figs. 52, 65); | dorsal carinae of 1st tergite completely absent | (fig. 65); 2nd tergite with droplet-shaped, coria- | ceous medio-basal area, ending posteriorly in # distinct keel, without distinct antero-lateral area | (fig. 65); 2nd tergite projecting over 2nd suture | (fig. 52); 2nd suture rather shallow and only | medially crenulate (fig. 65); 3rd and 4th tergites | with sharp lateral crease; 4th and 5th tergites rather flat (fig. 52); ovipositor normal, with small nodus and teeth (fig. 57). Distribution. — Afrotropical: The biology is unknown. one species. Deltaphyma horstoki sp. nov. (figs. 51—66) Holotype, ?, length of body 18 mm, of fore wing 14.5 mm. Head. — Antennal segments incomplete, 51, length of 3rd segment 1.7 times 4th segment, length of 3rd and 4th segments 2.2 and 1.3 times their width, respectively; length of maxillary palp equal to height of head; eyes slightly emar- ginate (fig. 61); length of eye in dorsal view 1.1 times temple; temple subparallel-sided behind eyes (fig. 62); POL : © ocellus : OOL = VAN ACHTERBERG: Afrotropical Braconinae 183 3:5:15; frons smooth, distinctly concave be- hind antennal sockets, medially with low eval- uation (fig. 62); face finely transversely striate, with some coarse striae; malar suture distinct; length of malar space 0.8 times basal width of mandible. Mesosoma. — Length of mesosoma 1.8 times its height; side of pronotum and mesopleuron smooth; episternal scrobe small and pit-shaped (fig. 52); metapleuron smooth; mesoscutal lobes largely glabrous and smooth; surface of propo- deum punctulate laterally, rest smooth; propo- deal spiracle medium-sized, rather elliptical and just behind middle of propodeum. Wings. — Fore wing: angle between 1-SR pade@+-S@+-R 55° (fie. 59); r : 3-SR : SR1 = ED SRE SR rm = 9): 22-710: Legs. — Hind coxa smooth; length of femur, tibia and basitarsus of hind leg 3.9, 9.9 and 6.6 times its width, respectively; length of spurs of hind tibia 0.3 and 0.4 times hind basitarsus. Metasoma. — Length of Ist tergite 1.2 times its apical width, its surface coriaceous at medial elevation, rest smooth; medial area of 2nd ter- gite surrounded by some rugae, rest smooth (fig. 65); 3rd and 4th tergites distinctly striate mediobasally, rest of metasoma smooth (fig. 52); length of ovipositor sheath 0.6 times fore wing; hypopygium large and acute apically. Colour. — Brownish-yellow; antenna, stem- maticum largely, metasoma behind Ist tergite (but medio-basal area of 2nd tergite yellowish), ovipositor sheath, middle telotarsus, hind tibia, spurs, and hind tarsus, blackish; basal third of wings, pterostigma (except its darkbrown apex), patch below pterostigma and near vein r-m of fore wing, more or less yellowish; rest of wings dark brown. Holotype in the Rijksmuseum van Natuur- lijke Historie at Leiden: “Horstok, Pr. b. Sp. (= Cape Town)”, “Museum Leiden, Bracon servil- le: Brullé” (incorrect!). Note. This species is named after its collector, Dr. H. B. van Horstok, a physician who col- lected for the Leiden Museum in South Africa during the period 1825—1835 (+ ca. 1838). Zanzopsis gen. nov. Type-species: Plaxopsis liogaster Szépligeu, 1915: Etymology: from a combination of “Zanzi- bar” and “Plaxopsis”, because the type species originates from Zanzibar and is closely related to the genus Plaxopsis. Gender: masculine. Diagnosis. — Scapus rather slender (fig. 67), ventrally slightly longer than dorsally (fig. 69) and its inner aspect not emarginate apically; apex of antenna without spine, acute and setose (fig. 81); face with large medial protuberance, surmounted by a W-shaped carina and next to protuberance somewhat depressed (figs. 75, 77); dorsal carina of clypeus nearly complete (fig. 75); clypeus ventrally cariniform (fig. 77); groove from eye to antennal sockets shallow and narrow (fig. 67); notauli anteriorly shallow- ly impressed, rest absent (fig. 76); scutellar sul- cus narrow and distinctly crenulate (fig. 76); scutellum with minute pit medio- -anteriorly; metanotum with very short medial carina (fig. 76); vein cu-a of fore wing postfurcal (fig. 70); vein 1-M of fore wing straight; vein m-cu of fore wing antefurcal, slightly converg- ing to vein 1-M posteriorly (fig. 70); vein 1- SR+M of fore wing distinctly bent subbasally (fig. 71); vein Ir-m of hind wing straight; mar- ginal cell of hind wing strongly narrowed api- cally (fig. 70); tarsal claws basally pectinate, without lobe (fig. 78); hind femur dorsally densely and ventrally sparsely setose, without apical flange (fig. 79); hind tarsus without ven- tral row of setae; hind spurs sparsely setose; dorso-lateral carinae of Ist tergite distinct be- hind spiracles, lamelliform, in front of spiracles absent (figs. 67, 82); dorsal carinae of Ist tergite absent, except for a weak basal elevation (fig. 82); 2nd tergite not projecting above 2nd suture (fig. 67), without antero-lateral grooves (only depressed sublaterally), and with smooth medio-basal triangular area surrounded by a crenulate depression (fig. 82); 2nd suture deep, completely and coarsely crenulate (fig. 82); 3rd and 4th tergites with crenulate antero-lateral grooves (fig.67); 2nd and 3rd tergites with sharp lateral crease; 4th and 5th tergites rather flat (fig. 67); ovipositor normal, with minute nodus and distinct teeth (fig. 73). Distribution. — Afrotropical and South Pal- aearctic (N. Africa): contains at least the follow- ing five species of which the type has been ex- amined: Zanzopsis liogaster (Szépligeti, 1913), Z. levis (Szépligeu, 1914), Z. pulchripennis (Szépligeu, 1911), Z. buettneri (Szépligeti, 1914), and Z. maculiceps (Szépligeti, 1914). The biology is unknown. Zanzopsis liogaster (Szepligeti) comb. nov. (figs. 67—82) Plaxopsis liogaster Szépligeu, 1913: 419. Lasiophorus hogaster; Shenefelt, 1978: 1692. 184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 Holotype, ®, length of body 15.3 mm, of fore wing 14.7 mm. Head. — Antennal segments 74, length of 3rd segment 1.5 times 4th segment, length of 3rd and 4th segments 1.5 and 1.0 times their width, respectively, penultimate segment 1.2 times its width (fig. 81); length of eye in dorsal view 1.3 times temple; temple parallel-sided (fig. 74); POL : @ ocellus : OOL = 4 : 5 : 14; frons flat, but shallowly concave near medial groove; face coriaceous and with some carinae (fig. 80); malar suture narrow; length of malar space 1.3 times basal width of mandible. Mesosoma. — Length of mesosoma 1.7 times its height; side of pronotum smooth and with medial groove; mesopleuron smooth, except for some punctulation ventrally; episternal scrobe shallow and narrow (fig. 67); metapleuron punctulate; mesoscutal lobes only setose near notauli, smooth, and covex; surface of propo- deum punctulate, but smooth medially; propo- deal spiracle nearly round, rather large and just behind middle of propodeum. Wings. — Fore wing: angle between 1-SR and C+SC+R of fore wing 58° (fig. 71); r : 3- SR ¢ SRI MOEI CU 2E CUI DEDTIDZSREIS-SREIT MORSE Legs. — Hind coxa smooth; length of femur, tibia and hind basitarsus of hind leg 3.4, 9.8, and 4.9 times their width, respectively; length of hind tibial spurs 0.3 and 0.4 times hind basitar- sus. Metasoma. — Length of 1st tergite 1.1 times its apical width, surface smooth, medially with a small, shallow depression with short carina (fig. 82); glymma narrow anteriorly (fig. 67); 2nd tergite smooth, except near basal area; 4th tergite crenulate anteriorly, rest of metasoma smooth (fig. 67); length of ovipositor sheath 0.78 times fore wing; hypopygium large, apical- ly acute (fig. 67). Colour. — Brownish-yellow; antenna, hind tibia and tarsus, and ovipositor sheath, blackish; metasoma and fore telotarsus, brown; middle telotarsus dark brown; basal third of wings, basal half of pterostigma, patch below pterostig- ma, area near vein r-m and below vein 2-SR+M of fore wing, yellowish; rest of pterostigma and wing membrane, dark brown. Holotype in Museum Berlin: “Sansibar, C. W. Schmidt”, “Type”, “Plaxopsis liogaster n. | sp.” (in Szépligeti’s handwriting). Plaxopsis Szépligeti, 1905 | Szépligeti, 1905: 1; Fahringer, 1931: 336 (key); | Shenefelt, 1978: 1691 (as synonym of Lasiopho- | rus). | Type-species: Plaxopsis sjoestedti Szépligeti, ; 1905 (monotypic). Gender: masculine. | Diagnosis. — Scapus medium-sized (fig. 84), | in dorsal view about twice its width, apically | truncate, not protruding ventrally (fig. 92) and | inner aspect of apex not emarginate; face with | pair of submedial depressions and near protu- | berance (rather) depressed (fig. 87); between | clypeus and face with a large, lamelliform, | spoonshaped protuberance, which is concave | ventrally (figs. 84, 87) and sometimes with horn | situated on it; clypeus with ventral carina; no | distinct groove between eye and antennal sock- | et; notauli shallow, but absent apically (fig. 89); | scutellar sulcus rather wide and coarsely crenu- | late (fig. 89); scutellum with pit medio-basally; | metanotum with short carina medially; vein cu- | a of fore wing slightly postfurcal (fig. 91); vein 1-M of fore wing straight; vein m-cu of fore | wing shortly antefurcal, distinctly converging to | vein 1-M posteriorly (fig. 91); vein 1-SR+M of | fore wing distinctly bent basally (fig. 91); vein i 1r-m of hind wing straight; marginal cell of hind | wing (sub)parallel-sided apically or slightly nar- | rowed, only medially distinctly narrowed | (fig. 91); tarsal claws setose (fig. 85); hind femur | without flange apically, dorsally distinctly and | ventrally sparsely setose (fig. 90); hind tarsus | with ventral row of setae; hind spurs normally | setose; dorso-lateral carinae complete (fig. 84); | dorsal carinae of 1st tergite absent (fig. 93); 2nd | tergite with smooth, subtriangular area medio- anteriorly, with pair of sublongitudinal grooves | laterally and with pair of small smooth triangu- | lar areas anteriorly (fig. 93); 2nd tergite not pro- | jecting above 2nd suture; 2nd suture deep and | coarsely crenulate; 3rd tergite with pair of near- ly complete, crenulate antero-lateral grooves (fig. 93); 4th and 5th tergites rather flat, with in- | complete antero-lateral grooves (fig. 84); ovipo- | sitor normal, with subapical nodus and with | small teeth ventrally. | Distribution. — Afrotropical and South Pal- | VAN ACHTERBERG: Afrotropical Braconinae 185 aearctic (N. Africa): moderately sized genus. The biology is unknown. Plaxopsis sjoestedti Szépligeti (figs. 83—93) Plaxopsis Sjöstedti Szépligeu, 1905: 1—3, fig. 1. Lasiophorus sjoestedti; Shenefelt, 1978: 1694. Holotype, 2, length of body 16.1 mm, of fore wing 16.2 mm. Head. — Antennal segments 64 (apical seg- ments missing), length of 3rd segment 1.5 times 4th segment, length of 3rd and 4th segments 1.6 and 1.1 times their width, respectively; length of maxillary palp equal to height of head; length of eye in dorsal view 1.5 times temple; POL : D ocellus: OOL = 3:3:11; frons weakly concave; face smooth; clypeus largely flat and smooth; malar suture shallow; length of malar space 0.9 times basal width of mandible. Mesosoma. — Length of mesosoma 1.7 times its height; side of pronotum and mesopleuron smooth; mesoscutal lobes largely glabrous, smooth and rather convex; surface of propo- deum smooth, posteriorly rather depressed (fig. 93); propodeal spiracle rather large, subel- liptical and at middle of propodeum (fig. 84). Wings. — Fore wing: angle between 1-SR An (SCORES (Ere 91); r : 3-SRe SRI = 9 : 50 : 51; 1-CU1 : 2-CU1 = 1 22 -SIRO 3 — SR : r-m = 24 : 50 : 22. Legs. — Hind coxa indistinctly punctulate; hind and fore claws missing; length of femur, tibia, and basitarsus of hind leg 4.3, 10.8, and 6.1 times their width, respectively; length of spurs of hind tibia 0.3 and 0.4 times hind ba- sitarsus. Metasoma. — Length of Ist tergite 1.7 times its apical width, smooth (fig. 93); glymma nar- row (fig. 84); 2nd tergite (except basal area) striate (fig. 93); rest of metasoma smooth; length of ovipositor sheath 0.84 times fore wing; hypopygium large and apically acute (fig. 84). Colour. — Dark reddish-brown; head, palpi, apex of antenna, prothorax, metasoma ventrally (but posteriorly brownish) and 9th tergite, yel- lowish; wing membrane dark brown, but fore wing with subapical yellowish, incomplete band and below pterostigma with a small light area (fig. 91). Holotype in the “Camerun”, “Sjöstedt”, n.g. n. sp.” Stockholm Museum: “Plaxopsis Sjöstedti (in Szepligeti’s handwriting). Plesiobracon GROUP The Plesiobracon group contains three genera known to me, which have the Ist tergite of the metasoma movably joined to the 2nd tergite, with strong and complete dorso-lateral carinae, the mesoscutum rather evenly setose and at the same time the propodeum with a strong and complete medio-longitudinal carina, the scapus is (sub)truncate apically and ovoid (figs. 97, 113), the clypeus has no dorsal carına, the angle between veins 1-SR and C+SC+R of fore wing more than 60°, vein cu-a of fore wing slightly postfurcal (figs. 98, 114), the laterope is absent, and the 2nd tergite has a pair of sublateral, con- verging or short longitudinal grooves (figs. 109, 121). The biology is unknown. KEY TO GENERA OF THE Plesiobracon GROUP 1. First tergite without dorsal carınae (fig. 137); posterior margin of hind wing (but sometimes hardly) concave subbasally and shorter setose than surrounding fringe (fig. 136); head strongly constricted behind eyes (fig. 132); vein cu-a of hind wing recli- vous (fig. 136); (Indo-Australian) ......... ee cnet. LER bed el Plesiobracon Cameron — First tergite with strong, united dorsal cari- nae (figs. 109,121); posterior margin of hind wing about straight subbasally and evenly setose (figs. 104, 114); head rounded posteriorly (fig. 107, 117); vein cu-a of hind wing (nearly) straight (figs. 98, 114); (Afro- tropical) 2. Medial length of 5th tergite ca. 0.8 times medial length of 4th tergite behind basal de- pression (fig. 95); vein 2-SR of fore wing straight (fig. 98); 5th tergite straight latero- apically (fig. 95); antescutal depression pre- sent (fig. 95); propleuron without posterior flange (fig. 95); 2nd tergite with small me- dio-basal area (fig. 109); mesoscutum punc- tulate (fig. 108); propodeal spiracle situated submedially at propodeum (fig. 95); 4th tergite more or less protruding medio-api- cally; 2nd submarginal cell of fore wing slender (fig. 98) ... Carinibracon gen. nov. — Medial length of 5th tergite (behind basal depression) 1.3—1.5 times medial length of 4th tergite (fig. 110); veins 2-SR of fore wing (rather) sinuate (fig. 114); 5th tergite oblique latero-apically (fig. 110); antescutal depression absent (fig. 110); propleuron with small posterior flange (fig. 110); 2nd 186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 tergite without medio-basal area (fig. 121); mesoscutum finely rugose (fig. 116); pro- podeal spiracle situated in front of middle of propodeum (fig. 110); 4th tergite straight medio-apically (fig. 122); 2nd submarginal cell of fore wing robust (fig. 114).......... SEE i az ari Kenema gen. nov. Plesiobracon Cameron, 1903 Cameron, 1903: 123; Shenefelt, 1978: 1717. Type-species: Plesiobracon carinatus Camer- on, 1903 (monotypic). Gender: masculine. Diagnosis. — Head strongly constricted be- hind eyes (fig. 132); apex of antenna with spine (fig. 130); ventral margin of clypeus cariniform and protruding (fig. 125); propleuron without posterior flange (fig. 123); anterior half of no- tauli distinct, smooth, rest absent (fig. 133); mesoscutum only laterally punctulate; scutellar sulcus deep, rather wide and distinctly crenulate (fig. 133); metanotum with some rugae medially (fig. 133); propodeal spiracle medium-sized, nearly round and behind middle of propodeum (fig. 123); vein 2-SR of fore wing straight; vein cu-a of hind wing reclivous (fig. 136); posterior margin of hind wing concave (but hardly ın vze- recki), more shortly setose than surrounding fringe (fig. 136); tarsal claws with large, round- ed submedial lobe (fig. 129); hind femur slen- der, its length about 4 times the maximum width (fig. 134); hind tibia with subapical trans- verse row of spiny setae (fig. 134); no ventral row of setae at hind tarsus; Ist tergite without dorsal carinae (fig. 137); 2nd tergite with a nar- row triangular area medio-basally (fig. 137); 2nd suture wide, straight and crenulate (fig. 137); 4th tergite protruding medio-posteri- orly (fig. 137); 2nd—5th tergites with sharp lateral crease; medial length of 5th tergite about 0.6 times medial length (behind basal depres- sion) of 4th tergite (fig. 123); 5th tergite nearly straight latero-apically (fig. 123); hypopygium large, acutely protruding beyond metasomal apex, medio-basally unsclerotized. Distribution. — Indo-Australian: two spe- cies. The 2nd species (besides the type-species from Borneo) is [phiaulax vierecki Strand, 1911 (= braconiformis Strand, 1911, nec Szépligeti, 1904) from New Guinea according to Dr. D. Quicke (in litt.). Plesiobracon carinatus Cameron, 1903 (figs. 123—137) Plesiobracon carinata Cameron, 1903: 123; Shenefelt, 1978: 1717. Lectotype, ®, length of body 5 mm, of fore wing 5.8 mm. Head. — Antennal segments 44, length of 3rd segment equal to 4th segment, length of 3rd and 4th segments both 1.9 times their width, penul- timate segment 2.3 times its width (fig. 130); length of maxillary palp 0.8 times height of head; eye glabrous, not emarginate; length of eye in dorsal view 3.5 times temple; POL : © ocellus : OOL = 7 : 7 : 20; frons flat, except for a medial groove (fig. 132); face indistinctly punctulate and rather flat; clypeus flat and smooth; length of malar space equal to basal width of mandible. Mesosoma. — Length of mesosoma 1.4 times its height; side of pronotum smooth, except for a crenulate groove medially; mesopleuron largely punctulate; episternal scrobe distinct and isolated; scutellum smooth; anterior half of propodeum punctulate, posteriorly smooth, long whitish setose. Wings. — Fore wing: angle between 1-SR and CHSCHR 816: BS REIS RUNNEN ERS 1-GUT : 2-CUL = 1 : 135 2 -SRAMESRE EME 10 811728 © Legs. — Hind coxa smooth; length of femur, tibia, and basitarsus of hind leg 4, 8.2, and 6.8 times their width; length of spurs of hind tibia 0.30 and 0.35 times hind basitarsus. Metasoma. — Length of 1st tergite 0.9 times its apical width, reticulate-rugose postero-me- dially, laterally with wide crenulate groove; spi- racles directed dorsally; 2nd—5th tergites fi- nally longitudinally rugose, 3rd—5th tergites with smooth apical rim (figs. 123, 137); 3rd ter- gite without lateral grooves; metasoma behind 5th tergite smooth; length of ovipositor sheath 1.32 times fore wing. Colour. — Blackish-brown; middle of frons, temples dorsally, and vertex, dark brown; rest of head, fore leg, and metasoma ventro-basally, yellowish-brown; pterostigma and parastigma, dark brown; middle femur, tibia and tarsus, brown; palpi light yellowish; wing membrane subhyaline. Lectotype in British Museum (Natural Histo- ry), London: “Type, SBMENIy peli 3.c.603”, “Plesiobracon carinata Cam., Type, Borneo” (in Cameron’s handwriting), “Kuch- ing, Feb. 3, 1902”, “Cameron Coll. 11903 121”. Lectotype here designated. In the Kuch- VAN ACHTERBERG: Afrotropical Braconinae 187 ing Museum I examined 2 ® in the collection under “carinata”: one specimen collected April 1, 1902, the 2nd specimen at May 24, 1903. Be- cause of the date of capture of the 2nd speci- men, this specimen is excluded from the type- series. The other ® is considered to be a para- lectotype. It differs from the lectotype by a nar- row antescutal depression. Carinibracon gen. nov. Type-species: Carinibracon danielssoni sp. nov. Etymology: from “carina” (Latin for “keel”) and “Bracon”, because it is closely related to the genus Bracon, but it possesses several carinae (propodeum, Ist and 2nd tergites) which are normally absent in Bracon. Gender: masculine. Diagnosis. — Head rounded posteriorly (fig. 107); apex of antenna without spine, setose (fig. 99); ventral carina of clypeus present, pro- truding (figs. 95, 103); propleuron without pos- terior flange (fig. 95); mesoscutum punctulate; notauli completely smooth (fig. 108); scutellar sulcus medium-sized, rather wide and crenulate (fig. 108); metanotum with short medial carina; propodeal spiracle round, near middle of propo- deum (fig. 95); vein 2-SR of fore wing straight; 2nd submarginal cell of fore wing slender (fig. 98); vein cu-a of hind wing straight (fig. 104); posterior margin of hind wing nearly straight basally and evenly setose (fig. 104); tar- sal claws with peculiar lamelliform lobe and with some pegs (fig. 100); hind femur robust, about 3.5 times its maximum width (fig. 102); hind tibia without subapical row of spiny setae (fig. 102); hind tarsus with distinct ventral row of setae; 1st tergite with dorsal carinae com- plete, meeting posteriorly, in front of strongly convex area (fig. 109); 2nd tergite with small subtriangular area medio-basally and with pair of weakly converging sublateral grooves (fig. 109); 2nd suture sinuate and crenulate (fig. 109); 4th tergite somewhat protruding me- dio-posteriorly; medial length of 5th tergite about 0.8 times medial length of 4th tergite be- hind basal depression (fig. 95); 5th tergite straight latero-apically (fig. 95); 2nd—6th ter- gites with sharp lateral crease; hypopygium large and acute apically (fig. 95). Distribution. — Afrotropical: one species. Carinibracon danielssoni sp. nov. (figs. 94— 109) Holotype, ®, length of body 4.7 mm, of fore wing 4.6 mm. Head. — Antennal segments 35, length of 3rd segment 1.3 times 4th segment, length of 3rd and 4th segments 2.3 and 1.8 times their width, respectively, penultimate segment 1.7 times its width (fig. 99); length of maxillary palp 0.7 times height of head; eye glabrous and slightly emarginate (fig. 103); length of eye in dorsal view 1.8 times temple; POL : © ocellus : OOL = 3:3:7; frons nearly flat, coriaceous and with short groove near anterior ocellus (fig. 107); face largely coriaceous, but smooth above clypeus; length of malar space 0.8 times basal width of mandible. Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum and mesopleuron smooth, but mesopleuron ventrally sparsely punctulate; episternal scrobe shallow (fig. 95); scutellum punctulate; surface of propodeum smooth and setose. Wings. — Fore wing: angle between 1-SR and C+SC+R 70° (fig. 101); r : 3-SR : SRI = 15 s 37 3 77/2 ICU 3 ZAC =S 2 3 25e DIN 3 D= SR :r-m = 28 : 37 : 17. Legs. — Hind coxa punctulate; length of fe- mur, tibia and basitarsus of hind leg 3.5, 7.5 and 5 times their width, respectively; length of spurs of hind tibia 0.35 and 0.4 times hind basitarsus. Metasoma. — Length of 1st tergite 0.8 times its apical width, largely smooth (fig. 109); 2nd tergite coarsely reticulate-rugose; 3rd—7th ter- gites finely rugulose, with narrow subapical transverse groove and posterior rim smooth (fig. 95); length of ovipositor sheath 0.8 times fore wing. Colour. — Brownish-yellow; antenna (of scapus and pedicellus only outer side), ptero- stigma, veins, and ovipositor sheath, dark brown; wing membrane brownish. Holotype in Entomological Museum, Lund: “Senegal, in forest, 1.5 km NE Djibélor, ca. 6.5 km SW Ziguinchor, 8.11.1977. At light, 19.00— 21805 Loe mor Zr WIM ZINC “Lund Univ, Syst. Dept. Sweden, Gambia/Se- negal Febr _— March 1977, Cederholm, Daniels- son, Larsson, Mireström, Norling, Samuels- son”. Note. — In existing keys the species runs to the genus Cratocnema, but Cratocnema differs e.g., by the presence of a dorsal carına at the clypeus. The descriptions of the Cratocnema spp. do not fit C. danzelssoni. It is a pleasure to me to dedicate this species to Mr. R. Danielsson (Lund), who has been very helpful in providing types and new taxa. 188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 9, 1983 Kenema gen. nov. Type-species: Kenema quickei sp. nov. Etymology: name based on the locality of the type. Gender: feminine. Diagnosis. — Head rounded posteriorly (fig. 117); apex of antenna with small spine (fig. 111); ventral carina of clypeus not differ- entiated (fig. 120); propleuron with small poste- rior flange (fig. 110); mesoscutum finely rugose (fig. 116); notauli only anteriorly distinct and finely crenulate (fig. 116), rest absent; scutellar sulcus narrow and finely crenulate (fig. 116); metanotum with complete longitudinal carina; propodeal spiracle small, round and in front of middle of propodeum (fig. 110); vein 2-SR of fore wing sinuate (fig. 114); 2nd submarginal cell of fore wing robust (fig. 114); vein cu-a of hind wing straight; posterior margin of hind wing straight and evenly setose (fig. 114); tarsal claws with small acute lobe (fig. 118); hind fe- mur moderately robust (fig. 119); hind tibia without row of spiny setae; hind tarsus with ventral row of setae; 1st tergite with dorsal cari- nae united medially, but medially rather weak (fig. 121); 2nd tergite without medial area and with short longitudinal grooves (fig. 121); 2nd suture distinctly sinuate and crenulate; 4th ter- gite straight medio-posteriorly (fig. 122); medi- al length of 5th tergite (behind basal, crenulate groove) 1.3—1.5 times medial length of 4th ter- gite (figs. 110, 122); 5th tergite oblique latero- apically (fig. 110); 2nd—5th tergites with sharp (cariniform) lateral crease. Distribution. — Afrotropical: two species. The 2nd species will be described in a forthcom- ing paper. Kenema quickei sp. nov. (figs. 110—122) Holotype, d, length of body 6 mm, of fore wing 4.5 mm. Head. — Antennal segments 52, length of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 1.7 and 1.4 times their width, respectively, penultimate segment 1.4 times its width (fig. 111); length of maxillary palp 0.6 times height of head; length of eye in dorsal view 2.6 times temple; temple ruguloso-coria- Geous ALOE IDKo ce USE OO frons slightly convex, laterally rugose, medially coriaceous, and with shallow medial depression (fig. 117); stemmaticum comparatively strongly protruding (fig. 120); face transversely rugose; length of malar space equal to basal width of mandible. Mesosoma. — Length of mesosoma 1.5 times its height; side of pronotum coriaceous, with some striae (fig. 110); mesopleuron coriaceous; episternal scrobe virtually absent; scutellum co- riaceous; propodeum densely rugulose, medio- laterally shortly lamelliform and angularly pro- truding (fig. 110). Wings. — Fore wing: angle between 1-SR and C+SC+R 61°; r-m distinctly sinuate and sclerotized (fig. 1114); 5 SES RS RUNE Ne 7 MIO 92 SRB SRE ten ORL Legs. — Hind coxa coriaceous; length of fe- mur, tibia, and basitarsus of hind leg 3.9, 8.2, and 5.2 times their width, respectively; length of spurs of hind tibia both 0.4 times hind ba- sitarsus. Metasoma. — Length of 1st tergite 0.4 times its apical width, basally smooth, behind dorsal carinae distinctly rugose; 2nd tergite rather coarsely reticulate; 3rd tergite reticulate-rugose; 4th and 5th tergites densely coriaceous-rugu- lose, with incomplete lateral grooves (fig. 110); 6th tergite shortly setose, exposed, smooth; 7th tergite glabrous; 2nd—5th tergites with acute lateral crease. Colour. — Brownish-yellowish; antenna (ex- cept annellus) black; wing membrane slightly infuscated (but more pigmented near parastig- ma; fig. 115); anterior margin of fore wing (ex- cept base) blackish, rest of pterostigma and most veins, brown. Holotype in Quicke Collection, Nottingham: “Kenema, Sierra Leone, Sept: 1981, Dt Quicke”. I am pleased to name this species after Dr. D. Quicke, who collected several interest- ing taxa in Africa. Note. This new genus resembles the genus Soter Saussure, 1892 (= Odontogaster Széplige- ti, 1906, syn. nov.), but Soter does not belong to the Plesiobracon group because it has the meso- scutum largely glabrous. Additionally it has the mesoscutum smooth, the 2nd submarginal cell is long, the 2nd tergite has converging grooves, the 5th tergite is serrate, and the spiracle of the propodeum is submedially situated. ACKNOWLEDGEMENTS I wish to express my gratitude to the follow- ing persons for the loan of types and of new taxa, and for assistance during visits: Mr. R. Danielsson (Lund), Dr. J. Decelle (Tervuren), Mr. A. A. Hamid (Kuching), Dr. K.-J. Hedq- vist (Stockholm), Mr. T. Huddleston (London), Mlle Dr. S. Kelner-Pillault (Paris), + Dr. E. VAN ACHTERBERG: Afrotropical Braconinae 189 Königsmann (Berlin), Dr. B. Petersen (Copen- hagen), Dr. D. Quicke (Nottingham), Mr. B. Sigwalt (Paris), Dr. V. I. Tobias (Leningrad), Mrs. I. Wegener (Berlin). The Uyttenboogaart-Eliasen Stichting finan- cially supported the visit to the Kuching Mu- seum (Serawak). REFERENCES Achterberg, C. van 1979. A revision of the subfamily Zelinae auct. (Hym., Braconidae). — Tijdschr. Ent. 122: 241—479, figs. 1—900. ——, 1982. Notes on some type-species described by Fabricius of the subfamilies Braconinae, Rogadi- nae, Microgastrinae and Agathidinae (Hym.: Bra- conidae). — Ent. Ber., Amst. 42: 133—139, figs. 1—9. Brullé, A., 1846. Hymenopteres. In: A. L. M. Lepele- tier de St. Fargeau, Histoire naturelle des insectes, pt. 4: 1689. Cameron, P., 1903. Descriptions of new genera and species of Hymenoptera taken by Mr. Robert Shelford at Serawak, Borneo. — J. Straits Br. Asıat. Soc. 39: 89— 181. — — 1912. Descriptions of new genera and species of parasitic Hymenoptera taken at Kuching, Sera- wak, Borneo, by Mr. John Hewitt, B. A. — Soc. ent. 27: 74—78. Fabricius, J. C., 1804. Systema Piezatorum, p. 1—439. — Reichard, Brunsvigae. Fahringer, J., 1930. Uber einige stidamerikanische Braconidengattungen. — Wien. ent. Ztg. 47: 19— dl, , 1931. Aethiopische Region. — Opuscula braco- nologica 5: 305—384. Granger, C., 1949. Braconides de Madagascar. — Mem. Inst. scient. Madagascar A: 1—428, figs. 1—426. Haliday, A. H., 1833. Essay on the classification of parasitic Hymenoptera. — Ent. Mag. 5: 209—248. Shenefelt, R. D., 1978. Braconidae, pt. 10. — Hym. Cat. (nov. ed.) 15: 1425—1872. Szépligeti, G. V., 1905. Beitrage zur Kenntniss der In- sektenfauna von Kamerun, 29, Braconidae. — Ark. Zool. 2 (14): 1—11. — — 1913. Braconidae von Madagascar und anderen Inseln Ostafrikas. In: A. Voeltzkov, Reise nach Ostafrika in den Jahren 1903—1905. — Wissen- schaftliche Ergebnisse 3: 419—423. Tobias, V. I., 1975. Novi tropitseskogo proischozjde- nya rod brakonid (Hym., Braconidae) v fauna srednej Azij. — Zool. Zhurnal 54: 962—964, figs. 1—3. 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AWE LIBRARY UNIVERSITY INHOUD L. BRUSSAARD. — Reproductive behaviour and development of the dung beetle Typhaeus typhoeus (Coleoptera, Geotrupidae), pp. 203—231, figs. 1—17. | Tijdschrift voor Entomologie, deel 126, afl. 10 Gepubliceerd 15-XII-1983 la REPRODUCTIVE BEHAVIOUR AND DEVELOPMENT OF DUNG BEETLE 7YPHAEUS TYPHOEUS (COLEOPTERA, THE Dept. of Animal Ecology and Dept. of Soil Science & Geology, Agricultural University, Wageningen, GEOTRUPIDAE) by LIJBERT BRUSSAARD The Netherlands ABSTRACT This paper is part of a study of the contribution of dung beetles to soil formation in sandy soils. Typhaeus typhoeus (Linnaeus) has been selected because it makes deep burrows and is locally abundant. The beetles are active from autumn until spring, reproduction takes place from February to April. Sex pheromones probably influence pair formation. The sexes co- operate in excavating a burrow (up to 0.7 m below surface) and in provisioning the burrow with dung as food for the larvae. Co-operation is reset by scraping each other across the thorax or elytra. Dung sausages, appr. 12.5 cm long and 15 mm in diameter, are manufac- tured above each other. Development is rapid at 13—17 °C. The life cycle is accelerated by a cold period in the third larval stage. These requirements are met by soil temperatures up to 15 °C in summer and down to 5 °C in winter. The life cycle lasts two years, but longer under certain conditions. Newly hatched beetles make their way to the surface through the soil, but do not follow the old shaft. Adults reproduce only once. Differential rate of com- pletion of the life cycle and occasional flying probably reduce the risk of local extinction. The study is thought to be relevant for behavioural ecology and soil science. CONTENTS oon En 203 Dung beetles and their association with soil .... 204 Choice of Typhaeus typhoeus as an object of EC rr AN Re 204 song biologe ee 206 DCO cb RENE M IE 206 WIZE OTOIS a odes E NEE LEE UA KE 207 Rieproductvebehaviouri wet. MR UNE. 209 DEelopmendmbedknt «Mon Ass mets de a 220 DScussionere te Werd. enen ilostennues db ui 226 Crdookss A re EE ERI 229 bichnowledeementst krts vet: 229 RESORT nde AS a 230 INTRODUCTION There were two main reasons for this study of dung beetles. First, ancient traces of former activity by small burrowing or crawling ani- mals, presumed to be dung beetles, can be found today in sandy soils. By studying the behaviour of dung beetles it should be possible to ascertain whether these are indeed the relics of dung beetle activity. Second, in areas where dung beetles are abun- dant nowadays, recent traces of their activity are present in the soil and this raises the ques- 203 tion of how much dung beetles contribute to soil formation today. These topics will be discussed in subsequent articles. The ancient traces (with an account of their age) will be described in a forthcoming pa- per and will be compared with the traces result- ing from recent dung beetle activity. It will be shown that dung beetles may indeed be respon- sible for the ancient traces. Knowledge about the reproductive behaviour of the dung beetles is required for a proper un- derstanding of their influence on soil morpholo- gy, and therefore this is discussed in the present paper. After a brief introduction to dung beetles as a group, the general biology of the species se- lected for study, i.e. Typhaeus typhoeus (Lin- naeus, 1758), is described and its geographical range is discussed. In order to quantify the beetles’ contribution to soil formation it will eventually be necessary to explain and predict their population dynam- ics. This considerable task was reduced to two basic investigations in the present study. The first pertains to the development of life stages during the season and to phenological phenom- ena that seem to be related to the persistence of 204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 the species in time and space. As this constitutes part of the biology of the species it has been in- cluded in the present paper. The second basic investigation pertains to the environmental con- ditions (including soil conditions) with which the beetles have to cope during the season of adult activity. These conditions may affect the beetles’ burrowing and reproduction. This will be treated in another paper. It will be shown that the number of dung targets and their distri- bution in the field is of paramount importance for the population ecology of T. typhoeus and its impact on soil. In its impact on soil Typhaeus typhoeus should serve as a model species for all Geotrupi- ni found in the temperate Holarctic and, to some extent, also for other dung beetle species of the paracoprid type in other parts of the world. Consequently, in this paper about 7. ty- phoeus, the emphasis will be on aspects of be- haviour that may help to explain the impact on soil formation caused by this kind of dung beetle. Aspects of the reproductive behaviour of T. typhoeus that have been published elsewhere will be re-described in terms of their relevance to soil formation. Furthermore, useful new in- formation about behaviour and the devel- opment of life stages will be presented. DUNG BEETLES AND THEIR ASSOCIATION WITH SOIL Representatives of several coleopteran fami- lies are regularly found inhabiting dung, e.g., Hydrophilidae and Histeridae. The term “dung beetle”, however, is usually restricted to a num- ber of species belonging to the superfamily Scarabaeoidea. Scarab beetles, whether dung beetles or not, show a close connection with soil. The larvae of most species live under- ground on a food-stock of dung or plant re- mains, prepared by their parents, or they show a free-living, root-sucking habit. Adults of most species forage above-ground on fungi, dead or- ganic matter or fresh leaves and burrow into the soil to prepare food-stocks for their larvae, to lay eggs or to hibernate or aestivate. Living on dung may have evolved from living on dead organic matter and fungi (Iablokoff- Khnzorian, 1977; Crowson, 1981). In the fami- ly Geotrupidae, to which Typhaeus typhoeus belongs, all these habits occur and this family has been placed near the base of the scarabaeoid evolutionary tree by the authors mentioned above. Dung-feeding beetles may have devel- oped the practice of digging into the soil as a re- | sult of searching for truffle-like fungi that | adopted a subterranean life history during times | of warming climate. The habit of making food- | stocks out of dead organic matter or dung | instead of searching for fungi is presumed to | have developed subsequently. | Dung beetles spread all over the world from | the Jurassic/Cretaceous periods onwards (Iab- | lokoff-Khnzorian, 1977; Crowson, 1981). The | Geotrupini tribe probably radiated out from the | area of the Tertiary Tethys Sea (Krikken, 1980) | and now shows a predominantly temperate, | Holarctic distribution. i In addition to the Geotrupidae, one other | family of scarab beetles is important for our | study, viz., the Scarabaeidae. This family not | only contains dung beetles (e.g. Scarabaeinae | and Aphodiinae), but also beetles of the cock- | chafer type (Melolonthinae), which, as adults, | live on fresh leaves and, as larvae, show a root- | sucking way of live. Cockchafers will be dis- | cussed in a subsequent paper. CHOICE OF TYPHAEUS TYPHOEUS AS AN OBJECT OF STUDY The impact of dung beetles on soil may be two-fold: enrichment with dung or plant re- | mains, and physical disturbance. On the basis of their impact on soil, dung beetles may be divid- ed into three ecological groups, as proposed by Bornemissza (1969) in a different context. First, the endocoprids, which pass their life cycle from egg to adult in the dung on the sur- face, or spend part of their life cycle a few centi- metres deep in the soil, e.g. Scarabaeidae-A pho- diinae. Second, the telecoprids, which make a ball out of the dung, roll it some distance away and bury it superficially as a food source for the larva, e.g. Scarabaeidae-Scarabaeinae. And, third, the paracoprids, which burrow a fairly deep shaft under or close to the dung patch; part of the burrow is filled with food for the lar- vae and part of it is back-filled with soil, e.g. Geotrupidae. Because of the depth of the shafts and the amount of dung transported below the ground, dung beetles of the paracoprid type, especially the larger species, may be considered to have the greatest impact on soil. In the temperate re- gions, paracoprid dung beetles of the Geotrupi- ni tribe are the most important in this respect, as has been shown by the studies of Fabre (+ 1910), Schreiner (1906), Spaney (1910), Von Lengerken (1954), Howden (1955, 1964, 1974), BRUSSAARD: Reproduction and development of Typhaeus typhoeus 205 Table 1. Depth of shafts in Geotrupidae. faunal species depth of source region shaft (cm) Palaearctic Geotrupes mutator* = 30 Teichert, 1955 G. spiniger* 25— 30 Lumaret, 1980 G. stercorarius* 50 Teichert, 1955 35— 60 Spaney, 1911 G. stercorosus* 35— 60 Spaney, 1910 G. vernalis* 12— 68 Teichert, 1959a Lethrus apterus 60—100 Frantsevich et al., 1977 75—100 Teichert, 1959b 50— 65 Schreiner, 1906 Typhaeus momus 10— 15 Baraud, 1977 T. typhoeus* 29—100 present study 60—100 (150) Kuyten, 1960 48—130 Teichert, 1959b 70—140 Spaney, 1910 = 150 Fabre, ca. 1910 T. hiostius 60—160 Crovetti, 1971 Nearctic Geotrupes egeriei 20— 75 (90) Howden, 1955 G. hornu 40— 75 id. Bolboceras farctum 57 id. Bradycinetulus ferrugineus 35—105 id. Mycotrupes retusus 45— 90 Olson, Hubbell & Howden, 1954 M. gaigei = 205 id. Peltotrupes youngi 140—270 Howden, 1952 * occurring in The Netherlands. Meichere 21955, 11956,..1957, 219592); Kuijten (1960), Crovetti (1971) and Klemperer (1978, 1979). There are differences between geotrupid species in, for example, geographical range, habitat, (use of) flight capability, reproductive season, depth of the shaft (table 1) and number of eggs laid (table 2). In northwest Europe, Typhaeus typhoeus (fig. 1) is one of the most obvious species to study. The beetles are locally abundant with a maximum of 1—2 pairs per m? and they make very deep burrows (tables 1 and 3). Occasional- ly I have found them as deep as 1 m and they may go even deeper, up to 1.50 m (Fabre, + 1910; Spaney, 1910; Teichert, 1959b; Kuijten, 1960). Moreover, they transport a fair amount of dung below the ground, as reflected by the number of dung sausages produced (= number of eggs laid; tables 2 and 4). 10 mm Fig. 1. Typhaeus typhoeus (Linnaeus, 1758). Left, male, and right, female. 206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 Table 2. Number of eggs laid per nest or per female (if reported) in Geotrupidae. number of eggs laid source per nest or per female species Geotrupes mutator* 4—16 per 9 Teichert, 1955 G. spiniger” 9—10 per ? Lumaret, 1980 G. stercorarius* 4—17 per 9 Teichert, 1955 (2) 3— 6 (8) Spaney, 1910 G. vernalis* 5—10 per 9 Teichert, 1959a Lethrus apterus 5— 7 Frantsevich et al., 1977 6—11 per 2 Schreiner, 1906 Typhaeus typhoeus* (1) 4—16 (21) per 2 present study 1— 6 per nest (field) id. = 10 per nest Palmer, 1978 = 15 per 2 Kuyten, 1960 3— 6 (8) Spaney, 1910 T. hiostius 2— 8 per nest (field) Crovetti, 1971 * occurring in The Netherlands. GENERAL BIOLOGY Nesting The reproductive behaviour consists of bur- rowing a branching shaft, provisioning the branches with food for the larvae and sealing the remaining burrow partly or completely with soil. Though the female, once fertilized, can perform the whole process on her own, the beetles normally operate in pairs. Some aspects of the reproductive behaviour have been out- lined previously by Fabre (+ 1910), Kuyten (1960) and Palmer (1978). Habitat The habitat is open to half open heathland, and the beetles are most abundant in bare areas surrounded by Nardus stricta, Cladonia spp. and Calluna vulgaris, and along paths. They al- sO Occur in open pine woods, predominantly along paths and in small clearings. A vital pre- requisite is the presence of dung. The beetles are found only on herbivore dung, mostly that of rabbits, as the rabbit is the most abundant her- bivore in the habitat of 7. typhoeus. They will also use dung from sheep, deer and roe. Seasonal and diurnal incidence From the second half of September onwards and throughout the winter, 7. typhoeus is active whenever the temperature is above zero and there is no snow. Intense activity occurs in Oc- tober and November, which is the main period of maturation feeding, and from February to April, which is the main period of reproduction. From May onwards the reproductive activity declines rapidly and from June to the latter half | of September there is no adult activity at the soil | surface (fig. 2). Contrary to the common as- | sumption that the beetles only reproduce after | the turn of the year (Fabre, + 1910; Main, 1917; Kuijten, 1960) I have established from field ob- | servations that pair formation, oviposition and provisioning with dung is not exceptional as early as the third week of October. On the oth- | er hand, unpaired beetles can be found in shal- low burrows as late as March, where they are apparently still involved in maturation feeding. | Although in overcast and humid weather ac- tivity on the surface by day is not exceptional, | T. typhoeus is usually active above-ground at dusk and at night. DISTRIBUTION The genus Typhaeus The genus Typhaeus Leach, 1815, contains six species of about the same size: 14—22 mm long and 8—11 mm wide. Five of these are re- stricted to the Mediterranean area: 7. hiostius (Gené, 1836), 7. momus (Olivier, 1789), 7. fos- sor (Waltl, 1838), 7. lateridens (Guérin, 1838) and 7. typhoeoides Fairmaire, 1852. The first of these is endemic to Sardinia. The sixth species, T. typhoeus has the largest geographical range: from Morocco to South Sweden, westwards to Ireland and eastwards to Poland (Horion, 1958; Lindroth, 1960) and Yugoslavia (Miksië, 1956), but it is absent in Hungary (pers. obs. and Endrödi, pers. comm., 1981). A preliminary map of the distribution in Europe is given in fig. 3. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 207 numbers so ee Fee in Po A months M | A Fig. 2. Number of Typhaeus typhoeus captured in standard pitfalls in various heathlands in the Dutch province of Drenthe during the years 1959—1967. (Courtesy of P. J. den Boer.) Typhaeus typhoeus in The Netherlands In The Netherlands, 7. typhoeus is found in sandy areas, even in isolated spots surrounded by peat, clay or loam soils (Gaasterland, Be- tuwe, South Limburg) but, remarkably, it is not present in the dunes along the west coast and on the Wadden islands in the north of the country. Fig. 4 is a preliminary map of the distribution of T. typhoeus in The Netherlands. Absence from the dune region In a preliminary experiment to ascertain the reasons for the absence of 7. typhoeus from the dunes, it was found that under laboratory con- ditions two pairs of beetles reproduced quite normally when supplied with pellets of rabbit dung from the Wadden island of Vlieland. The trial was stopped when the larvae were in their final instar. Under field conditions in enclosures in the dunes of North Holland near Castricum, beetles made normal burrows in which they provisioned dung for their offspring, whether supplied with pellets of rabbit dung from the dunes (two pairs) or from the inland (two pairs). When the burrows were excavated six months later, it appeared, however, that only one of the 22 dung sausages contained a live larva, whereas in most of the other cases the larva had died and in some cases the egg had evidently not hatched. This work needs to be continued to yield con- clusive results. METHODS To study the behaviour of 7. typhoeus in the laboratory, the beetles were kept in cages (1 m high and 0.60 m wide), similar to the one de- scribed by Main (1916/17): a wooden frame in which two windows (4 mm thick) were kept a distance of 15 mm (sometimes 12 mm) apart. The space between the windows was filled from above with tamped down portions of sand that came from a field at Wijster, in the Dutch prov- 208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 L Fig. 3. Geographical range of Typhaeus typhoeus. Grey area: after Pijpers (1981). Striped area: typhoeus pre- sent according to Horion (1958), Lindroth (1960) and Sinclair (1977). Barred area: typhoeus not occurring according to Endrôdi (pers. comm., 1981). Fig. 4. Distribution of Typhaeus typhoeus in The Netherlands. After Pijpers (1981). Each dot indicates at least one specimen. Stars constitute additional occurrences, assessed by present author. The dot near the west coast pertains to two specimens, captured around the turn of the century. ince of Drenthe, where T. typhoeus occurs nat- urally. The soil there is coversand to a depth of 1.40 m, in which a podzol has developed (Van Heuveln, 1965). The predominant particle size was 0.1—0.2 mm and particles larger than 0.6 mm were scarce. The organic matter content of the sub-soil was 0.6%. The sand was air-dried and subsequently moistened to a water content of 10% (by mass) to approximate field condi- tions in September at a depth of about 0.50 m below the surface. The tamping resulted in a bulk density of approximately 1.50 g/cm}, which was similar to the field situation at about 0.50 m below the soil surface. The beetles made their burrows in the sand between the windows. The light/dark regime in the rooms with the cages was the same as in na- ture. Daylight conditions were simulated with TL-33 tubes supplemented with normal bulbs. To prevent light affecting the beetles in the soil, the windows of the cages were covered with sheets of black plastic. A horizontal walking- surface (0.50 X 0.60 m2) on which dung could be offered was mounted on top of the glass cage and covered with 0.8 mm mesh wire-netting to prevent the beetles from flying away. Observa- tions were usually carried out under dim red light, after the plastic sheets had been removed. For the rearing 60 cages were used. To com- pare the laboratory results with the field situa- tion, five cages were dug into the soil in the field at Wijster. In addition, the experimental equip- ment in the field included five 1 X 1 m? and six 2 X 2 m? enclosures consisting of 0.50 m wide stainless steel plates which were inserted to a depth of 0.20 m in the soil. These enclosures were also spanned with the wire-netting. Almost all beetles used in the experiments were captured at the same site near Havelte in the province of Drenthe. Newly hatched adults can easily be collected in autumn from under the small hummocks of soil, where they have re- tired with some dung for their maturation feed- ing. Prior to experiments the beetles were kept in sand-filled plastic jars, 13 cm high and 10 cm in diameter, for a least six weeks at 5 °C. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 209 Further details about the experimental meth- ods will be given in the appropriate sections be- low. REPRODUCTIVE BEHAVIOUR Emergence and maturation feeding The first newly hatched adults of 7. typhoeus appear on the surface in the second half of Sep- tember, usually after heavy rain. They immedi- ately go in search of dung. As soon as a beetle has found a small collec- tion of dung it excavates a J-shaped feeding bur- row approximately 15—20 cm deep (in the case of females sometimes deeper) and 13—16 mm in diameter. The beetle carries a number of dung pellets (in the case of rabbit dung mostly 10— 20) down into its burrow and starts its matura- ton feeding. Given that reproduction was ob- served as early as the third week of October, the maturation feeding time in 7. typhoeus is proba- bly approximately four weeks, at a temperature of 13—16 °C. Flying To judge from the many beetles I found crawling around on the surface and the relative- ly few flying, it would seem that the beetles mostly move by walking and less so on the wing. Nonetheless, flying was observed in the field at a temperature of about 12 °C in the sec- ond week of October at dusk, in foggy weather with little wind. The beetles emerged from their burrows with their hind wings already unfolded and pumped up and flew off immediately. They flew low, zigzagging over the vegetation and the maximum distance I saw covered in one numbers flight did not exceed an estimated 50 m. At- tempts to fly were also observed at the same time of the day in the laboratory in the plastic jars in which the beetles were kept prior to ex- periments. Blut (1938) encountered 7. typhoeus flying at dusk in late May. Flying is possible even at very low temperatures, since in one of my laboratory experiments a female flew around in the walking area of a glass cage at 5°C, although she had been subjected to that temperature for over six weeks. To study flight movements in 7. typhoeus more closely, two window traps and a mist-net trap were placed in a study plot at Wijster, in the Dutch province of Drenthe. The window of the window trap measures 100 X 50 cm? and it catches beetles flying at a height of 150—200 cm above the ground. The mist-net trap measures 100 x 50 cm?, catching beetles flying 20—70 cm above the ground. After colliding with a trap the beetle falls down into a reservoir containing 4% formaline. The two window traps were in operation from 29 September 1978 and the mist-net trap from 8 November 1978 until the summer of 1980. In all, 19 beetles were trapped: eight males and eleven females (fig. 5). Fifteen beetles were captured in the single mist-net trap and only four in the two window traps, which confirms that the beetles fly low. Of the eleven females, ten were relatively unimpaired when the trap was inspected; two (captured on 27 September and 1 October, respectively) showed developing ovaries and had not yet mated; eight contained eggs and had mated, to judge from the presence of sperm cells. This shows that al- though no flying beetles were captured during peak reproduction in March, T. typhoeus can be months Fig. 5. Number of beetles trapped in flight at a study plot at Wijster (province of Drenthe, The Netherlands) during 1978/79 and 1979/80. 210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 i 1 cm i Fig. 6. Stance adopted by male Typhaeus typhoeus for supposed pheromone release during defecation. (Photo of beetle in nest entrance on walking surface of glass cage.) cumulative number of newly hatched beetles on the surface 24 20 16 SE TRO ZIA Zee EEE a 3 5 7 I DPOF Eise 17 18 September October date Fig. 7. Cumulative number of newly hatched male and female beetles appearing at the surface in September and October 1980 in a rearing trial started in late winter 1979. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 211 added to the list of species not obeying the oo- genesis-flight syndrome of Johnson (1969). The gut content of 14 of the trapped beetles was qualitatively estimated. In five beetles the gut was half filled or less, in nine the gut was more than half full. From these findings it can- not be concluded that a shortage of dung is the reason for flight. Settlement and pair formation Crovetti (1971) states that in Typhaeus hios- tius the male penetrates the feeding burrow of a female after maturation feeding. In 7. typhoeus I have observed behaviour that strongly sug- gests that pheromones may play a role, at least number of rabbit dung pellets taken down 20 in some stage of adult life, in pair formation: under laboratory conditions I have repeatedly observed that a male who has abandoned a nest with a female, then digs a shallow burrow near a food source, similar to the J-shaped feeding burrow. Next to this the male can be observed defecating in a characteristic stance, his body tilted at an angle of about 45° to the surface with his head above the entrance of the burrow and his abdomen lifted (fig. 6). This stance suggests that a pheromone is released with the excre- ment. Although the hypothesis of pheromone release needs experimental confirmation, it is significant that this stance was invariably adopted around the time that the light in the ex- 5 6 7 8 9 10 weeks after capture Fig. 8. Mean number of rabbit dung pellets carried down by beetles kept in plastic jars at 13—16 °C for ten weeks after their capture on 29 September and 4 October 1978. Vertical lines nd upper half of standard deviation. DR TijpscHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 perimental room was automatically switched on or off, i.e., around dawn or dusk. At those times the weather is usually favourable for odour communication because of high air humidity and moderate wind velocity. Crovetti’s (1971) observation that the male joins the female after maturation feeding and my observation that males appear to be trying to attract females after the pair has split up, sug- gests that the former behaviour changes to the latter in the course of the season (assuming the two Typhaeus spp. behave similarly). There is some evidence to support this. In the rearing trials I carried out, the females usually appeared on the surface and started maturation feeding before the males: the median of the total num- ber of females on the surface was reached eight days before that of males (fig. 7). Furthermore, in a cohort of beetles captured in the field, fe- males carried down the same quantity of dung for maturation feeding as did the males (fig. 8). If the duration of maturation feeding is the same for both sexes then it seems probable that fe- males mature sexually earlier than the males. For a female, the prerequisites for reproduction are a male and an adequate amount of dung, as a food source for the future larvae, and therefore her most profitable strategy seems to be not to go and search for one of the few males available for mating, but rather to settle near a spot rich in dung, make a shallow burrow and wait there for a male. This needs to be confirmed by addi- tional research. During the reproductive season the pattern changes: whenever a pair of beetles abandons a nest it is usually the male who leaves first, as will be reported in greater detail in another pa- per. Several days may pass before the female ap- pears outside the burrow. By the time the male leaves the burrow most females will be paired and involved in breeding, so that it is unprofit- able for him to search for one of the few bur- rows with an unpaired female. Instead, he seeks a spot with plenty of food, near which a new nest can be made, and tries to attract one of the females that will appear above-ground after abandoning a nest. Copulation On the first encounter, which usually takes place in a shallow feeding burrow, the male vig- orously sweeps his front tibiae across the fe- male’s thorax, the female turns around and then the male sweeps across her elytra while half mounted on her back. Finally, the female lifts her abdomen and copulation follows, lasting from 3 to 20 min. The female terminates the copulation by stepping forwards a few cm, turning around and pushing the male back. Burrowing Having paired and copulated the beetles make a nest that finally consists of a shaft that may or may not divide into tunnels, from which a num- ber of brood chambers branch off, provisioned with dung for the progeny and sealed by back- filling with soil (fig. 9). The female excavates by scraping the sand under her body with her front tibiae and then using her middle and hind legs to move it further back. As the sternites are dense- ly covered with backward-pointing hairs, the sand does not fall down past the beetle when she is in a vertical position. While excavating, the beetle intermittently turns around its length axis. Every time the beetle has excavated 0.5— 1.0 cm of the shaft she moves several mm back- wards, and by doing this tamps down the moist sand behind her into a plug. Then she turns around and pushes the plug into the shaft with her head and thorax, again intermittently turn- ing around her length axis. This turning enables her to apply force to a different point and thus facilitates the transport of the plug. The upper part of the shaft, within a depth of 30 cm from the surface, is made horizontal for some 10—15 cm (fig. 9). Here the female always leaves her sand plug before returning down the burrow to continue excavating. The male then burrows through the sand plug. Since the female has left the plug in a horizontal part of the passage, the sand does not fall down the shaft. Once past the small plug, the male turns around and shovels it to the exit, transporting it in the same way as the female. In this way a sand heap gradually ac- cumulates on the surface, finally achieving a height of some 5 cm and a diameter of 10—15 cm. In one of the glass cages the excavation of the shaft was closely monitored. Fig. 10 clearly shows that the beetles may continue to excavate for four days without pausing. On four successive days, during periods indi- cated in fig. 10, I recorded the intervals during which the female was involved in sand excava- tion and transport, respectively. On the first three days the duration of the periods of excava- tion was the same, on the fourth day the dura- tion was much longer, presumably indicating that the female was about to terminate the bur- depth(cm) | 4 10 16 22 28 34 40 46 52 58 64 70 76 82 88 94 100 BRUSSAARD: Reproduction and development of Typhaeus typhoeus 213 t t {—— path followed by emerging beetle Ì Ì Ì | <— tunnel edge of frame Fig. 9. Nest of Typhaeus typhoeus in one of the glass cages. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 214 ‚15/8 02°] Jo Auısuap xnq e YI pues ara pay advo sse]3 e ur snoogdKı snovgd(y zo mede Aq ©, ç Ie 10dsuen pue uoneaeoxa pueg ‘O1 “314 93 ep | 61 | 8l | EI | KAaenuer DI | ST (sanouy)owuL3 02 91 CI 8 yp wa OC OÙ gt 8 Pp ve OG OL el 8 v Pa 06 MEL 8 vb ve 0 EE EE e Aa | | Zi BuLmoyung JO uoL2JeuLW4g? zen 52 2 =o Ee A * Od eo À © LA ® * E ® . =. —06— i = — 019 y x a * CE oO a % // 5 D e Gala geo _* B SET © ei _* ® * a DI a Be: Bi 001 = * Zee sno zz a o A = -011- = == — A H4 _i __yAi — poruad bBuLp4o9au LZ 3 A è (T=U) €°62 eu) Ge ESS (£=U) G'°2+9°/T (ereU) ZA STESA (ULU) potdad .J4odsue4? JO UOLJPANP (d=) HTO TI (mel) Tie (GU) BIE (vI=u) 8°0+6°% (utw) potuad uotqeAe9X9 40 UOLZEANP BRUSSAARD: Reproduction and development of Typhaeus typhoeus 215 rowing. The duration of sand transport steadily increased to half an hour per plug. The cage had been filled and the sand tamped down under water. This resulted in a bulk density of 1.70 g/cm3, which is the maximum a beetle may gen- erally encounter in the field. The temperature in the room with the cage was 5 °C throughout the reproduction of the beetles. This approximates the soil temperature up to a depth of 1 m, mea- sured in the field at the beginning of March, fol- lowing the severe winter of 1978/79. This tem- perature was, therefore, near the lowest the beetles may encounter underground in the field in The Netherlands. Even so the beetles pro- gressed fairly rapidly (fig. 10). In a total of 60 cages filled with less densely packed sand (not exceeding 1.55 g/cm?) and at temperatures of 5— 17 °C the shaft was excavated within 1—3 days. The morphology of the shaft was very vari- able. However, a horizontal part was invariably present within a depth of 30 cm. During the process of excavation the female often made an- other horizontal or slightly sloping gallery where she left the sand plug while continuing deeper. The shaft terminated in an oblique or horizontal gallery 6—15 cm long: the future first brood chamber (fig. 9). The depth of the shaft varied considerably, as can be seen from table 3. The diameter of the shaft is determined by the size of the larger of the two beetles. In the field, casts of twelve shafts were made using liquid paraffin wax that solidified on cooling, enabling the diameter of the shafts to be measured. The mean diameters of all shafts were then averaged: the overall mean was 14.2 + 0.7 mm (S.E.). Oviposition and behaviour prior to and following it As soon as the female has terminated the bur- rowing she begins to make a small cavity (diameter about 0.5 cm) for the future egg by moving her head and fore legs in the blind end of the almost horizontal terminal part of the burrow, alternately scraping some sand away and pushing part of it back. When she has fin- ished she walks up and down the passage until the male arrives. The male joins the female, sometimes before she has finished the egg chamber, when he no longer encounters a sand plug to transport upwards. Then the beetles usually mate, as described before. Copulation at this stage, i.e. prior to oviposi- tion, is not obligatory but it is seldom omitted. After copulation the male starts scraping sand over a distance of about 10—15 cm from the bottom of the future first brood chamber. He may do this several times, so that a sand plug is formed. Then he turns around and transports the plug upwards. The female continues to pre- pare the egg chamber, breaking off to walk through the future brood chamber, with her ab- domen pulsating. These pulsating movements are probably connected with the transport of an egg into the oviduct. Finally she moves her ab- domen into the egg room and oviposits, making gentle pumping movements. Oviposition may last 10—25 minutes. After oviposition the fe- male shows the backward scraping behaviour, just like the male, over a distance of some 10— 15 cm, but in the opposite direction. Then she turns around and pushes the sand into the end of the burrow, thus sealing the egg chamber. The wall that thus separates the egg chamber from the future brood chamber becomes 10—15 mm thick (compare fig. 9). As a consequence of the scraping of sand by male and female the diameter of the brood chamber is slightly larger than that of the rest of the burrow: 15.0 + 0.7 mm (S.E.), n = 12. While the female is completing the egg cham- ber and preparing the brood chamber, the male is involved in widening the shallow horizontal part of the burrow, which is to become a store room for dung pellets. The diameter of the store room becomes 2—4 cm. Table 3. Depth of shaft in Typhaeus typhoeus under laboratory and field conditions. [Temperature in the lab 5° or 9 °C; bulk density in glass cages and casks about 1.55 g/cm} to match field conditions. Enclosures 1 X 1 m?; casks 0.5 x 0.5 x 1 m?. Ample supplies of dung were provided in all experiments. year experimental set-up n depth (cm) range (cm) 1979 glass cages lab (2-dim.) 12 67 + 22 29—100 1979 enclosures field (3-dim.) 5 SY ae 6 63— 80 1980 glass cages field (2-dim.) 5 68+ 9 52— 78 1980 casks lab (3-dim.) 4 58 + 13 46— 78 216 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 Dung provisioning When she has finished preparing the brood chamber the female walks up the burrow to meet the male in the store room or, alternative- ly, if he is ready first, the male walks down after enlarging the store room. When they meet, the female pushes the male upwards firmly and vig- orously sweeps his elytra with her fore legs. This continues, unul the male finally makes for the surface to fetch dung, the female waiting for him in the store room or in the brood chamber. When he has found a fecal pellet, the male usually takes it between his front tibiae and walks backwards with it to the nest entrance. Sometimes he holds the pellet between his man- dibles, but I have never observed the pellet be- ing carried on the horns as described by Fabre (+ 1910). The male drags the dung pellet into the direction of the nest entrance in a straight line, however tortuous his searching path may have been. If the dung pellet is found within about 40 cm from the nest entrance the male usually enters the hole at once. If the fecal pellet is found further away, however, the beetle drops it within 5— 10 cm from the entrance and then moves directly to the entrance, walking forwards, puts his head into the entrance for a few seconds, turns around, picks up the dung pellet and carries it down without further delay. Whenever the beetle misses the entrance he finds it after an area-restricted search. This be- haviour of searching for the entrance in the vi- cinity of the nest shows that the beetle is capa- ble of roughly estimating the distance from the place where the dung was found. During the procedure of dragging a dung pellet from a fair distance away, the behaviour of leaving it be- hind and walking forwards to the nest entrance may occur more than once. As soon as the male, dragging the dung pellet backwards into the nest, appears in front of the female below, she immediately begins to sweep his elytra. Then the male pushes the dung pellet underneath himself and walks up the shaft again to fetch more dung. The number of rabbit dung pellets dragged in successively by the male does not usually exceed 30; it depends on the ease with which he can find them and the distance to be covered. The time needed to collect them al- so varies, but seldom exceeds two hours. After this bout of dragging dung pellets, the male stays below-ground for some hours. The female takes a dung pellet from the store room and, holding it in her fore legs, lets herself fall down the shaft by drawing in her middle and hind legs close along the body. In the brood chamber she tears the pellet to pieces with the help of her mandibles and fore legs and then firmly presses the pieces into the blind end of the brood chamber with her head and thorax, intermittently turning around her length axis. This firm pressing causes a meniscate layering within the dung sausage that is going to fill the brood chamber. The female walks up the shaft to collect every dung pellet. Alternatively, the male may supply her with dung by carrying down a number of fecal pellets. Sometimes the male kicks the dung pellets out of the store room with his hind legs. As a consequence the lowest part of the burrow behind the female be- comes filled with fecal pellets. Often, the male walks down the shaft to the female. If the female progresses too slowly he may stimulate her and he often tries to copulate. When the female goes up to fetch more dung to provision the brood chamber and encounters the male before she reaches one of the pellets in the store room, she invariably stimulates him by sweeping his elytra. It thus appears that the co- operation of male and female in the stage of dung provisioning is often reset by interaction. The number of rabbit dung pellets processed per dung sausage varies between 30 and 65, av- eraging about 40. The number of dung sausages manufactured varies between 4 and 21, averag- ing about 10 (table 4). Sealing the dung sausage and excavating the next brood chamber When the dung sausage is finished the female Table 4. Number of eggs laid (= dung sausages manufactured) in Typhaeus typhoeus under laboratory and field conditions. [Experimental conditions as mentioned in table 3.] year experimental set-up n 1979 glass cages lab (2-dim.) 11 1979 enclosures field (3-dim.) 5 1980 glass cages field (2-dim.) 5 1980 casks lab (3-dim.) 4 number range JEANS 2 1 10.6 +49 6—19 VEND Aid WODael/ SZ BRUSSAARD: Reproduction and development of Typhaeus typhoeus DG Fig. 11a. Dung sausage being sealed over a predetermined stretch with sand from the shaft walls. Fig. Allo, Tine widening is plastered with sand from the new brood chamber. 218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 seals it with soil, using sand scraped from the wall of the shaft. Interestingly, this wall-scrap- ing starts some cm above the proximal end of the dung sausage and in this way the amount of the shaft to be filled with sand is determined (fig. 11a). Only half way or later in the stage of sealing is the next brood chamber excavated. The widened part of the shaft above the sand plug that seals the former dung sausage is inad- vertently plastered by the beetle with sand that it drops as it is carrying it upwards (fig. 11b). The next brood chamber is excavated above the former one (fig. 12). 29 glass cages 5 with only 1 tunnel 24 with >1 tunnel 23 with the deepest tunnel excavated first = with the deepest tunnel not excavated first Fig. 12. Brood chambers are excavated above each other, beginning with the deepest. Fig. 15 shows that the first tunnel excavated is usually the deepest. A tunnel is that part of the burrow from which one or more brood chambers branch off. This may be equivalent to a shaft, but more than one tunnel may be found branching off from the same shaft (fig. 9). A new tunnel is usually branched off when the latest brood chamber is relatively shallow. In the present study, the uppermost dung sausage in 22 cases with more than one tunnel was, on average, about 40 cm deep (table 5). While exca- vating the subsequent brood chambers the same behavioural sequence of transporting sand, cop- ulation, oviposition and dung provisioning 1s shown. As long as the male is present the female al- ways excavates a new brood chamber, at times preceded by a new tunnel, irrespective of the availability of dung or the number of eggs al- ready laid. If the male is no longer there, the be- haviour of the female varies. If there is still a supply of dung she may continue the whole process of excavating a brood chamber, egg-lay- ing, gathering dung and manufacturing dung sausages on her own. She may even excavate the next brood chamber with no male present and no dung around. If she has laid the next egg al- though there is no supply of dung, she may fill the newly excavated brood chamber with sand from the walls of the burrow, which shows that the behaviour after oviposition is fixed upon provisioning with whatever material there is around. Alternatively, the female may abandon the nest, invariably after finishing and sealing the last dung sausage, and continue reproduc- tion elsewhere. Behaviour of larvae and newly hatched adults As soon as the egg has hatched the larva makes its way through the 1—1.5 cm thick sand wall that separates it from the dung sausage and moves into the dung in a somersaulting motion by which it displaces material from in front of it to behind it. As a consequence, after the larva has passed through, a small wad of dung, several mm long and wide, remains at the distal end of the dung sausage. The larva eats its way through the dung sausage, back-filling the space behind it with its excrement (fig. 13), so that the cavity surrounding the larva becomes only 2—3 cm long. It may eat its way through the dung sau- sage several times. There are three larval stages. Finally the larva III moves out of the dung sau- sage at the distal end and makes a cavity at the site of the former egg chamber: the pupal cham- ber. As a consequence, the distal end of the dung sausage becomes filled with sand displaced by the larva. The pupal chamber is plastered with excrement by the larva, which finally lies on its back to pupate (fig. 14). After pupation the newly hatched adult often remains days or weeks in the pupal chamber be- fore going to the surface. In my rearing trials, 45 out of 51 emerged adults passed through the partly eaten dung sausage, which thus appeared to be the rule. The other 6 immediately bur- Table 5. Depth of uppermost dung sausage in shafts with more than one tunnel. [Experimental conditions as mentioned in table 3.] year experimental set-up n depth (cm) range (cm) 1979 glass cages lab (2-dim.) 10 36 + 10 20 —48 1979 enclosures field (Schim) a5 ADI 12 35 —54 1980 glass cages field (2-dim.) 3 38 + 10 25 —50 1980 casks lab (3-dim.) 4 BP ae 18 20.5—40.5 BRUSSAARD: Reproduction and development of Typhaeus typhoeus 219 ecm Ls Fig. 13. Larva of Typhaeus typhoeus eating its way from the distal end (right) to the proximal end (left) of a dung sausage and back-filling it with its own excrement. (Photo of dung sausage and larva in glass cage.) Fig. 14. Pupa of Typhaeus typhoeus in pupal chamber made outside the distal end of the dung sausage. The horn (right upper part of pupa) shows that this specimen will become a male. (Photo of pupa in glass cage.) rowed their own way upwards from the pupal chamber. At least 23 out of the 45 passing through the old dung sausage subsequently also burrowed their own way upwards. I believe this to be the normal behaviour because after pass- ing through the old dung sausage, the remaining 22 broke through the sealing sand and subse- quently followed the old shaft, but they were most probably forced to do so by the cages, so this should be regarded as abnormal behaviour (see the arrows in fig. 9). A beetle burrowing its way upwards scrapes. 220 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 80 tunnels 16 with 1 brood 64 with >1 brood chamber chamber 3 with the deepest brood chamber not excavated first 6 with next brood chamber not always above former one Fig. 15. The tunnel excavated first is usually the deepest. the sand away above itself, turns around and firmly presses it behind. Consequently, at the beginning of this behaviour the space in the old dung sausage or in the pupal chamber is filled with sand. Subsequently, a corridor about 8 cm long steadily extends upwards through the soil, the beetle scraping sand away above and pressing it beneath itself. An emerging beetle reaches the surface within a few hours, depend- ing on the depth from which it starts. On arrival at the surface the animal shows the behaviour as described in the section on emergence and mat- uration feeding. DEVELOPMENT Development of eggs and larvae To study the (rate of) development of the dif- ferent stages of 7. typhoeus, beetles were reared in cages as described in the section on methods. The results of the rearing trials, which lasted al- most three years, are given in tables 6 & 7, which cover the rearing period from winter 1979 to autumn 1980 and from autumn 1980 to autumn 1981, respectively. In the rearing trials the effect of administering cold winter periods was most noticeable. These cold periods were administered because soil temperature had been found to drop from 13 °C in August to 3 °C in February at a depth of 1 m in the field. At 0.20 m from the surface the fall in temperature was greater (from 16° to 1 °C). After presenting the results of the rearing tri- als in the laboratory, the development of larvae in the field will be described. To facilitate com- 61 with the deepest brood chamber excavated first 55 with next brood chamber always above former one prehension, the course of development is briefly outlined in fig. 16. Rearing trials (1979— 1980) The rearing trials were carried out in rooms with constant temperatures of 1°, 5°, 9°, 13° and 17°C, respectively. These temperatures were chosen because soil temperatures measured in 1979 in the study plot at Wijster (where 7. ty- phoeus occurs naturally) ranged from 3—10 °C in the reproductive period and increased to 13— 16 °C during summer, when the larvae develop. At all the temperatures they were subjected to, the beetles showed their reproductive behav- 1our. At 1 °C only very few eggs were laid, howev- er, and the dung sausages were abnormal, con- sisting partly or completely of whole dung pel- lets. The eggs did not hatch. Therefore the rear- ing trials carried out at 1°C will not be discussed further. At 5 °C reproductive behav- iour was normal, but the eggs did not hatch ei- ther, not even after 20 months. At a temperature at or exceeding 9 °C the eggs did hatch. The time eggs laid at 5 °C took to hatch at 9°, 13°, 17° and 20 °C was estimated (table 8). Though the number of observations is small in some groups (because this trial was not solely in- tended to study the hatching time of the eggs) it is quite clear that at 9 °C development is rela- tively slow. At the start of the rearing trials there were four cages at each temperature. However, three of the cages kept at 9 °C soon had to be discarded because in one cage the fe- male died without reproducing and although 221 Reproduction and development of Typhaeus typhoeus BRUSSAARD oY} AIUIY 1x91 995) Á[UO D, LINE CC «CHIC = DI *:1U911E91] PIO> E DAI9III JOU PIP EU sa3vo 211 wort po31owo LEU 9s0oU} put JUIUTEIII P[o9 12 SUIAI9III sa3svo 21 WOI po81ouo JEUI Sa]199q Jo JOQUINU au) U29MI9q J9U9T9FIP JULITFIUSIS e SEM AU LL SMOLT Do EI Put 6 oy us MY ger 241 ul yeorg (4 ‘99M [I ‘91 ‘18611 EZ 020861 IX S WOIJ À, g 1e poled sun jerorgnaw (¢ (È (i [ae sqerm SuIICAI wos} LIEP 01 ure11od ger 273 JO WONOG 20) IE STEIOT, "PIS 179] 941 UO UDAIS SIE [reds PI è O1 parsalqns 10U sıred UIOI} IVP ‘suwnjoo IUEA9[II 91 JO opis Bid ya UO paruasaid are 0861 Arenıgag 87 01 6/61 1240120 OE wor porzod pjoo v 01 poroolqns sıred wor Eq “MEIS OY] Joie SHIUOU QZ ‘aI ‘1861 Jaquiaidas ur paddorg “axor 998 “Med | 01 Bururjpoop uoos ‘sapoag jo sated + AJjenTuT “MEIS DY 191€ SYIUOW QZ ‘2'1 ‘0861 Joquiaaony UI poddorg ve 92 92 BL 08 0861 REIT vi 61 EE 2s 9€ SS SS 0861 AI LI 0861 AI LI vi LI ot gi vi 6261 Al è 6261 AI p 6/61 Al Vv 8€ LI 02 ee 111 62/82) °! | jon | 6261 III 62/82 6L6l III 62/82 6261 III 62/82 = u potdad 4aqULM FNOYZLMT potsad AaJULM UZLM 7 oft IVAYVI ‘same du auodojpip Ano} av sa8vo ssej8 ur suaogd (1 snavqdd 7 yo saved Sureoa JO synsoy ‘9 [GEL 222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 Table 7. Results of ongoing rearing trials in glass cages from November 1980 unul October 1981 in rooms kept at a temperature of 13° or 17 °C. winter o % periods ©, È offered au 5°C as indicated in Fig. 12 - + + + + = 7 7 6 7 Na = 20, DATES 18 3 15 (P << 0.005) five dung sausages were made in each of the other two cages they were very superficial and the larvae soon died, probably because the fluc- tuations in moisture were too great. On 30 October 1979, by which time virtually all the larvae had emerged, the cages were divid- ed into two groups, one group remaining at the temperature it had been subjected to so far, the second group receiving a five-month cold spell at 5 °C until 28 February 1980, after which the cages were once again subjected to their former temperatures. The single cage remaining at 9 °C was assigned by chance not to receive a cold spell. One year later, rearing at 9 °C was repeat- ed with one cage that was given a cold period (see table 6). The number of pupal chambers and the time that elapsed until they appeared can be read from table 6. At 9 °C the median number of pupal chambers was reached much later in the cage without a cold period than in the cages at 13 °C and 17 °C, whether or not the latter received the five-month cold spell. This j AU — SP} SU | — MI | SP y SU | — MI | Sa SU | = WI] SP | L979 1980 1981 1982 again indicates that development is very slow at DIG At 13°C and 17°C pupal chambers were made by the larvae of all groups. This proves that a cold period is not a prerequisite for mak- ing a pupal chamber. At 13 °C and 17 °C, in the groups receiving a cold period, the median number of pupal chambers was achieved after a similar period from the start of the trials: 55 and 52 weeks, respectively. The date by which half the beetles had emerged was also very similar for these two groups. Therefore, I felt justified in combining the data on the different stages in the cages kept at 13 °C and 17 °C (bottom of ta- ble 6). This resulted in a very significant differ- ence between the proportions of beetles emerg- ing from cages which had and those emerging from cages which had not been subjected to a cold period: only one beetle emerged from 19 pupal chambers in the latter, compared with twelve beetles out of 14 pupal chambers in the former (y2 = 211-855 PT 005) Ma iso adults eggs 7 larvae (pre)pupae adult progeny season year Fig. 16. Outline of development of life stages in Typhaeus typhoeus. At the bifurcations arrows indicate larvae changing into the pupal stage, while the continuous line indicates remaining in the larval stage. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 228 Table 8. Numbers of eggs hatching and median time that elapsed before hatching in a rearing trial in glass tubes. rature (°C) cold period, although not a _ prerequisite, certainly favours the completion of the life cycle. The single beetle that emerged from a cage without a cold spell appeared in the same period as those from the cages subjected to a cold interlude (table 6). Of the 13 beetles that emerged, six were fe- males and seven were males: a 1 : 1 sex ratio. From these results it can be inferred that soil temperatures in the field ranging from 3—10 °C in early spring to 13—16 °C in summer, favour reproduction and the development of the larvae. Two other experiments in cages were carried out in early spring 1979, one at 5 °C and one at 9°C. These will be reported in another paper. In June 1979, when reproduction in these ex- periments was over, the cages were taken to rooms in which constant temperatures of 13 °C or 17°C, respectively, were maintained. Thenceforth these cages were exposed to the same treatments as those in the above-men- tioned rearing trials, including the division into groups that did or did not receive the cold peri- od. Data on pupal chambers and the beetles that emerged are given in table 9. The results clearly confirm the conclusions already reached regard- ing sex ratio and effect of a cold spell on rearing results. Rearing trials continued (1980—1981) As mentioned before, in cages at 5 °C, the eggs had not hatched after 20 months. The two cages at 9 °C, one of which had received a cold winter period, were kept at 9°C until 20 months had elapsed from the start of the rearing trials. By that time they contained three and one live larvae in pupal chambers, respectively. It may thus be surmised that even at such a low temperature adult beetles might have emerged after a (new) cold period. Since the results of the 13 °C and 17 °C treat- ments of the three trials were similar and, more- over, the beetles emerged in the same period (ta- bles 6 & 9), those cages that apparently still con- tained life after the period of emergence of the beetles, were pooled and subsequently divided into four groups for continued rearing. Two of the groups were subjected to a cold period at 5°C from 5 November 1980 to 23 January 1981. This was done in such a way that after the rearing trials were completed all four combina- tions of 0 to 2 cold periods had been realized, as outlined in fig. 17. The results are given in table Te A beetle emerged from almost all pupal chambers in cages subjected to a cold period for the first time (23 beetles out of 27 pupal cham- bers). In cages that had already received a cold period during the 1979—1980 part of the rear- ing trials, only two pupal chambers remained in which the larva had not pupated. During the 1980—1981 part of the rearing trials this small number rose to six and, when subjected to a sec- ond cold period, a beetle emerged from five of Table 9. Rearing results from two additional experiments. Data on left of columns pertain to glass cages that had not received a cold period. Data on right of columns pertain to glass cages that did receive a cold period. additional experiment (13°C) additional experiment (17°C) 30 IX 1980 | 25 IX 1980 25 IK 1980 | DAR Wil 7 SP 4 SU TNA UA AUS sie 1979 1980 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 AU | WI, SP, SU | AU, season year Fig. 17. Cold spell at 5 °C (+) or no cold spell (—; constant temperature of 13 °C or 17 °C) administered in rearing trials during 1979—1981 and percentages of adults emerging. Percentages pertain to numbers mentioned in Table 6, columns 8 and 12, and Table 7, columns 4 and 5. these six pupal chambers. But in the cages not subjected to a second cold period no beetles emerged from the seven pupal chambers. Few beetles emerged from cages that received no cold treatments (five beetles from 14 pupal chambers). As in 1979-1980, the number of beetles that emerged after a cold period during the 1980—1981 part of the rearing trials differed very significantly from the number that emerged from cages not subjected to a cold pe- rod = 20.12: P << 0.005). Of the 33 beetles emerging from the cages in 1981, 18 were males and 15 females, which con- firms the 1 : 1 sex ratio found the previous year. All twelve larvae remaining after the 1980— 1981 part of the rearing trials were in their third instar (head capsules 4.6— 5.2 mm). Development of larvae in the field Evidently, the quantity of cold that larvae of a given generation experience in the field during winter will differ because of the differing depths at which eggs are laid in the soil and because some winters are more severe than others. In general, the larvae occurring closest to the sur- face will experience more cold in winter and more heat in summer, which may accelerate their development vis-à-vis that of deeper lar- vae. To test this, five cages were dug into the soil in the study plot at Wijster in November 1979, in such a way that their upper edges just reached the surface. At the beginning of March 1980 a pair of dung beetles and ample dung were introduced into each cage. The beetles were prevented from exchanging with the surrounding field popula- tion by wire-netting placed over the cages’ walking surface. The cages were dug out at the beginning of September 1981, by which time larvae that had completed their life cycle could be found as newly hatched but not yet emerged beetles; in addition, this was also the right time to check whether, as expected, some of the lar- vae had not yet completed their life cycle. The five cages proved to contain 39 dung sausages. In eleven of these a living larva was found, one of which was in a pupal chamber. All larvae were in the third instar (head capsules 4.5— 5.0 mm). In addition, one newly hatched beetle was found. Though this is minimal evidence, the tri- al does demonstrate that under field conditions larvae of the same generation may give rise to adult beetles in different years, assuming, of course, that some of the remaining larvae would complete their life cycle in the future, as oc- curred in the laboratory rearing trials. Interes- tingly, the cage with the adult beetle also con- tained a larva in the pupal chamber and some other larvae, still in dung, which shows that dif- ferent rates of development may occur in the offspring of one pair of beetles. The results of another field trial, executed in the same period, enable a judgement to be made about whether the poor survival obtained in the above-mentioned field trial (life in 12 out of 39 cases = 31%) was somehow induced by the glass cages. In this trial five pairs of beetles were released in enclosures measuring 2 X 2 m?, at the beginning of March 1980. Their nests were dug up in May and June 1981, i.e. about three months earlier than the cages. There proved to be 32 dung sausages, 14 of which (i.e. 44%) contained a living larva; three of them had al- ready made a pupal chamber. Assuming some mortality in the enclosures between May/June and September, the data suggest that there was BRUSSAARD: Reproduction and development of Typhaeus typhoeus 225 no difference in survival between the enclosures and the cages. Thus there is no evidence that the cages have a detrimental effect on survival. Conclusions on rearing The results from the rearing trials imply that the development of the larvae is very slow at 9°C as compared with that at 13° and 17°C. The latter temperatures closely approximate those in soil during summer. Some larvae pass through the life cycle at constant high tempera- ture (17 °C). But the proportion of larvae that develop into adults is greatly improved by a cold period in the third larval stage. This condi- tion matches to a varying degree the low soil temperature during winter. Depending on the experience of a cold winter period, larvae of the same generation and even from the same pair of beetles may give rise to adults in different years. So the life cycle may last three instead of two years and perhaps even longer. This variation in the duration of the life cycle enables genetic ex- change between reproductive pools to be passi- vely achieved in successive years. Longevity of adults Genetic exchange between reproductive pools in successive years might also be achieved if adult beetles reproduced more than once. This is unlikely to be the case in 7. typhoeus, since in all laboratory trials executed at a temperature above 5 °C and in all the field trials, the beetles died shortly after reproduction. In many insect species, the dissection of females starting re- peated reproduction reveals the presence of yel- low bodies (corpora lutea) at the terminal ends of the ovarioles; these are the remnants of the follicular epithelium, which degenerates after egg release (Krehan, 1970; Vlijm & Van Dijk, 1967). To investigate whether yellow bodies are formed and can be found in 7. typhoeus, adult females were collected in the course of their ac- tive period and subsequently dissected. More- over, females from several experiments that died after reproduction were dissected. Of 14 fe- males that died after reproduction, three proved to possess corpora lutea. This indicates that in T. typhoeus yellow bodies are not always formed to an extent that they can be recognized. Of 52 females, captured in the field in the course of the season, eight possessed distinct corpora lutea and three only a vague indication of these. These eight females were captured late in the season (May and June). Obviously, it was not known whether these females were on their way to aestivate or to die. Clearly, the absence of yellow bodies does not allow conclusions to be drawn about repeat- ed reproduction, and only if their presence is es- tablished at the onset of the reproductive period can these bodies be used as indicators of repeat- ed reproduction. Therefore, an experimental approach was fol- lowed. A trial to have beetles aestivated and let them reproduce in the following activity period was executed twice. From May 1979 onwards, 60 beetles, captured when newly emerged in October and November 1978, were kept in plastic jars (13 cm deep, 10 cm in diameter; one beetle per jar) containing moist sand at a tem- perature equal to that measured in the study plot at Wijster 20 cm below the surface. Natural daylength was applied. The beetles were regu- larly supplied with fresh rabbit dung. Of the 60 beetles, 25 had survived after reproduction (13 males and 12 females) before May at a constant temperature of 5 °C. The other 35 (23 males and 12 females) had been kept at 5 °C in the jars since their capture. By 2 August 1979 all the beetles had died except for two males (one male from each group). Shortly thereafter the tem- perature-regulating equipment in the laboratory broke down and the beetles died from heat. In 1980, 25 beetles that had not reproduced but that had been kept in plastic jars at 5 °C since their capture in autumn 1979 were used in a trial. No beetles that had already reproduced were available. In mid-March 1980 these 25 beetles were divided into four groups: staying at 5 °C (three males, one female); 9 °C (six males, three females); 13 °C (five males, one female); and 17 °C (five males, one female). By 1 July all the beetles kept at 9°, 13° and 17 °C had died; two males at 5 °C were still alive. These two males survived and were given the chance to pair with two fresh females in glass cages on 27/28 November 1980. One male died on 2 De- cember without having reproduced, the other died on 15 December after the female had laid a few eggs. From the results of the dissection and of the trials, it can be concluded that aestivation of adult beetles and repeated reproduction in T. typhoeus is very exceptional, if indeed it oc- curs. Most of the beetles die after the first re- productive season. Thus the contribution to ge- netic exchange between reproductive pools in successive years by means of survival of adults is virtually non-existent. DISCUSSION Influence of glass cages on behaviour and results of rearing Although the behaviour of the beetles seemed normal, the narrow space in which they were confined may have influenced their behaviour. Tables 3 & 4 show that there were no major dif- ferences in the depth of the shaft and number of dung sausages between beetles kept in the labo- ratory and those kept or found in the field. Moreover, the number of dung sausages pro- duced in the present trials was very comparable with figures reported in the literature (tables 2 & 4). As reported in the section on the devel- opment of eggs and larvae, the survival of larvae kept in glass cages dug into the soil and kept in enclosures in the field was similar. This con- firms that glass cages do not adversely affect the behaviour of the beetles Pheromones There are few data on settlement and pair for- mation in the literature. Halffter & Matthews (1966), in their extensive review of the biology of Scarabaeidae, assume that the sexes meet by chance near the dung patch. In addition to the case of Typhaeus hiostius mentioned earlier (Crovetti, 1971), the intrusion of a male into a female’s feeding burrow and joint feeding until sexual maturity, is also reported by Rommel (1961) in Copris hispanus and by Halffter & Lo- pez (1977) in Phanaeus daphnis. The hypothesis of pheromone communication in 7. typhoeus, tentatively suggested in the present paper, is supported by reports that in Scarabaeidae some dung patches are densely populated with beetles, whereas others are not, and that the densely populated dung patches often differ from each other in the taxon that is most repre- sented (e.g. Halffter & Matthews, 1966; Pascha- lidis, 1974). A similar clustered distribution is known in bark beetles (Scolytidae), and this is known to be caused by aggregation pheromones (Blight et al., 1980). By analogy, pheromones may be important in dung beetles, too. More- over, the stance adopted by the male 7. ty- phoeus during the supposed pheromone release is virtually the same as that described by Pas- chalidis (1974) in three scarabaeid Sisyphus spp. and by Tribe (1975) in Kbeper nigroaeneus. In the latter case pheromone release was proved. In Kheper the pheromone is released from a de- pression on either side of the first abdominal sternite, with the help of paraffin tubules as a TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 carrier. This is very different from the way the pheromone is supposed to be released by 7. ty- phoeus, i.e. along with the excrement. The latter mechanism is also known for other groups, e.g. in bark beetles, boll weevils and some Orthop- tera (Jacobson, 1972). If properly supported by additional research, pheromone release by T. typhoeus would constitute the first known example of pheromone communication in Geo- trupidae. The conformity in releasing stance and the difference in releasing mechanism point to a convergent development and may be added to arguments in favour of the modern view of clas- sifying scarabaeids and geotrupids in distinct families (Crowson, 1967, 1981) instead of in subfamilies (Halffter & Matthews, 1966). In all cases the pheromone release stance has so far only been described in male dung beetles. The possibility should not be excluded, howev- er, that after maturation feeding the male is also attracted to his first female by means of phero- mones. Fabre (+ 1910) had already noticed that at times two or three males of T. typhoeus can be found in a burrow with only one female. Tei- chert (1955) observed the same in G. mutator and from this he inferred that males were at- tracted by pheromones released by the female. This would certainly improve the female’s chances of mating. It has been stated by Halffter & Matthews (1966) that the similar size of the antennal clubs in almost all male and female dung beetle species is a sign of the unimpor- tance of pheromone communication in these beetles. But this argument does not hold if both sexes respond to pheromones. Burrowing One may wonder how much faster the female progresses in the burrowing stage thanks to help from the male. The gain is estimated by Teichert (1957) to be one-fourth to one-third of the excavation time. Co-operation between male and female in the burrowing phase is not so close in some species as it is in 7. typhoeus. It is marked in the rela- tively deep burrowing species Geotrupes verna- lis, Lethrus apterus and Typhaeus hiostius, whereas in the shallow burrowing species G. mutator, G. stercorarius and G. spiniger the male does little more than remove some soil from the entrance (Teichert, 1955, 1959a, 1959b; Crovetti, 1971). According to Klemperer (1979), in G. spinig- er the female stops excavating the shaft and the brood chamber as a response to their length. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 227 Behaviour preceding and following oviposition Oviposition in 7. typhoeus is almost always preceded by copulation. According to Weaver & Pratt (1977) repeated copulation has a stimu- lating influence on the reproductive effort in the cockroach Periplaneta americana. This is prob- ably why a female of 7. typhoeus that has lost the male continues reproduction much more slowly. During the backward scraping activity that follows oviposition, the beetle presses its head and thorax hard against the walls of the brood chamber, possibly to make the walls resist the pressure they will be subjected to when being provisioned with dung. Although he did not ob- serve the pressing behaviour, Fabre (+ 1910) noticed that the walls of the brood chamber . were smoother than those of the shaft. Dung provisioning One may wonder how the male orients him- self on the surface, when dragging a dung pellet to the nest entrance. Kuyten carried out some unpublished experiments in 1961/62 from which he concluded that the position of a light source provides the beetle with a directional cue. The same conclusion was independently reached by Frantsevich et al. (1977) after some very similar experiments with Lethrus apterus. But it has not yet been explained how typhoeus takes its bearings by night or on cloudy days, when most of the excursions for dung are made. Sealing and excavating brood chambers Klemperer (1979) states that in Geotrupes spiniger sealing a dung sausage is a side-effect of excavating the next brood chamber. In 7y- phaeus typhoeus, however, at the start of sealing the sand is scraped from the wall and firmly pressed into the shaft above the dung sausage. Moreover, the last dung sausage is invariably sealed; this is also evidence of a distinct behav- ioural phase. In the endemic Sardinian species Typhaeus hiostius, which closely resembles typhoeus both in its biology and in the reproductive season, the sequence of excavation of brood chambers is remarkably different. Contrary to typhoeus, hiostins constructs the uppermost dung sausage first, at about 60—70 cm below the surface, and every subsequent sausage is 10—15 cm deeper, the completed burrow generally reaching a depth of 1.30—1.60 m. As a result, under field conditions some weeks after the nest has been completed the uppermost dung sausage already contains a larva III, whereas the egg belonging to the lowest dung sausage has not yet hatched (Crovetti, 1971). This behavioural sequence may well be related to soil moisture conditions (Crovetti, pers. comm., 1981), the female begin- ning to lay eggs approximately at ground-water level and then deeper as the water table falls. The ultimate function of this sequence may be to ensure that eggs and larvae do not dry out during the hot summer. In T. typhoeus, however, the risk of drying out may be considerably lower. In this species, laying the first egg approximately at ground- water level and thenceforth more shallowly may ensure that the larvae and their dung sausages do not become waterlogged during the next winter. In this context it would be interesting to study the behavioural sequence in typhoeus in the southern part of its geographical range. Emergence through soil The way the beetle moves to the surface is seldom mentioned in the literature. Sano (1915/16) reports that Geotrupes stercorarius does not always use the original tube made by the parents, but makes its own route, as de- scribed for Typhaeus typhoeus in this paper. Crovetti (1971) reports that 7. hiostius breaks through the partly eaten dung sausage and the sand that seals it and follows the shaft made by the parents. The evidence available for 7. ty- phoeus (this paper) and for 7. hiostius, suggests that the newly hatched beetle starts moving up- wards at a place where the resistance from soil is least, i.e. through the remnants of the dung sau- sage. However, it seems to be luck, rather than strategy, whenever the beetle finds the open part of the parental shaft. Emergence through soil thus seems to be the rule. Behaviour as a reaction chain According to Klemperer (1979) the nesting behaviour of Geotrupes spiniger can be de- scribed as a reaction chain in which each action generates its own terminating stimulus and initi- ates the subsequent response. Although experi- mental analysis is beyond the scope of the pre- sent paper, the nesting behaviour of 7. typhoeus is in many respects very similar to that of G. spiniger. In addition, the way in which the stretch of the burrow that is to be filled with sealing sand is determined in 7. typhoeus (fig. 11) is an example of an action that gener- ates its own terminating stimulus. So it may well 228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 be that the behaviour of typhoeus can also be described as a reaction chain. The extent to which signalling between the sexes by means of pheromones, sweeping each other and stridula- tion alters this picture requires further research, however. The role of temperature in diapause and development Diapause is induced in the third larval instar and can be overcome by a cold winter period at 5 °C. Tauber & Tauber (1976) rightly point out that this does not prove that diapause develops under the influence of low temperature in na- ture, as in many instances diapause can artifi- cially be broken in several ways, including ma- nipulation of day-length and administering a high temperature. Yet it is difficult to imagine an environmental stimulus other than tempera- ture that would induce, maintain and develop the diapause in larval dung beetles, since they live in complete darkness and in well-drained soils in which, over the year, differences in moisture are rather unpredictable. On the other hand, although temperature is known to be the single environmental stimulus maintaining rhythms in a number of insects during the day (Saunders, 1976), total reliance on temperature as a diapause-regulating factor in the course of the year has, to date, only been reported in tropical insects (Tauber & Tauber, 1976). Regarding the role of temperature in larval development, the few larvae that deviated from the general pattern in the rearing trials (tables 6 & 7) are interesting. Some larvae did not com- plete their life cycle after a cold period. This suggests there may be considerable variation in the amount of cold (duration and/or degree) a larva needs before diapause is terminated. On the other hand, some larvae completed their life cycle without a cold interlude. As the latter were from the cages kept at 17 °C this suggests that a relatively high temperature can also over- come diapause. Raising the temperature from 15° to 21 °C has, for example, been shown to be sufficient to overcome diapause in rearing trials of Geotrupes spiniger (CSIRO, 1980: 66). This should be further studied in 7. typhoeus, since it may mean that in the southern part of the geo- graphical range a warm period in the first sum- mer (summer 1979 in fig. 16) may be sufficient to enable the species to pass through its life cy- cle within one year. This may help to explain why Fabre (+ 1910), working in southern France, found a one-year life cycle in his first rearing trial (though not in his second). The same phenomenon may occasionally occur in the northern part of the geographical range too, M e.g. in a hot summer, but as a rule in those re- gions development will continue after a cold pe- riod in winter (winter 1980 in fig. 16), which thus leads to a life cycle of a minimum of two years. Spaney’s suggestion (1910) (he worked in M Germany) that in most cases 7. typhoeus has al one-year life cycle, should be regarded with | suspicion. Main (1916/17), working in England, | found a two-year cycle. Though most Geotru- pidae have a one-year life cycle, some are | known to show a cycle of at least two years (Sa- M no, 1915/16; Main, 1916/17; Howden, 1955). Î Field results support the hypothesis that lar- 7 vae of the same generation develop at different | rates: the rearing trial in glass cages, dug into | the soil and started in March 1980, yielded one adult beetle and a number of larvae in their final instar in the autumn of 1981 (see section Devel- opment, p. 224). But as only one adult beetle emerged, the hypothesis that the rate of devel- | opment is related, by means of temperature, to the depth at which the larvae develop, could not be tested. Regarding the development of eggs laid be- fore the turn of the year, it is unlikely that these | eggs can hatch before winter, let alone reach the third larval stage, since soil temperature at 20 cm to 100 cm below the surface is uniformly + | 9°C by late October and subsequently de- creases. As a consequence, the eggs do not | hatch until early spring together with the eggs laid by that time, and the larvae do not experi- | ence a cold spell until the following winter. It may thus be inferred that a one-year life cycle | probably cannot be achieved in The Nether- lands, starting from these early eggs. This may not be true, however, in the southern part of the geographical range. Flying The Geotrupidae differ greatly in their use of flight. As early as 1910, Spaney remarked that Geotrupes stercorosus and Typhaeus typhoeus can rarely be observed flying, whereas G. ster- corarius regularly flies. He thought this was be- cause beetles of the first two species generally | find plenty of food around in the place where | they emerge and reproduce, whereas those of | the last need to move from one fresh dung | source to another. As regards the significance of | the flying behaviour in the life history of 7. ty- phoeus, three observations seem to be relevant. BRUSSAARD: Reproduction and development of Typhaeus typhoeus 229 | First, the behaviour is shown in the laboratory in the presence of an ample supply of dung. Sec- ond, most of the beetles captured flying in the field did not have an empty gut. And third, I have observed the flying behaviour in the field in places with adequate supplies of dung that are apparently easily accessible to the beetles. This shows that flying is not solely connected with a shortage of dung and it may well be that flying is mainly related with searching for a partner. At the end of the season, however, there may be less dung available because it has been used by the beetles and because the density of the rabbit population has decreased, and then the beetles may fly in search of new dung supplies. This as- pect needs to be studied further. Persistence of populations in space and time Clearly, much speculation remains about the cause and function of flying in adult 7. ty- phoeus. There is also speculation about the se- lective forces that have led to and maintain the pronounced differences, both within and be- tween pairs of reproducing beetles, in the depth at which the eggs are laid. Ignoring the causes, the different rates of development of larvae from one generation lead adult beetles to emerge in different years; this passively contrib- utes to a reduction of the risk of local popula- tion extinction in time. Similarly the dispersal of adults results in, although is not necessarily aimed at, reducing the risk of local population extinction in space (Den Boer, 1968). The extent to which these phenomena contribute to the persistence of 7. typhoeus populations within a certain area merits further study. OUTLOOK The present investigation of the biology of Typhaeus typhoeus will prove to be invaluable | for the follow-up studies. Among the most im- portant findings for the understanding of soil morphology and soil formation are: first, the observations of how, and to what extent, the burrows are excavated and, in particular, subse- quently back-filled with dung and soil; and, sec- ond, the way in which newly hatched adult beetles make their way to the surface. For the understanding of environmental fac- tors (including soil conditions) that play a role in the species’ burrowing behaviour and repro- ductive performance, results of rearing trials will prove indispensable. Apart from these aspects, which are closely related to the soil-oriented approach of the study, the present investigation reveals a num- ber of starting points that may also be worth de- veloping. The co-operation between male and female is so striking that further study is prom- ising both from an ecological point of view (what is the gain in time and energy for the fe- male and in net reproduction for both part- ners?) and, if carried out as a comparative study of related species, from an evolutionary point of view (how and when did co-operation devel- op?). In addition, the role of contests and the probable role of pheromones deserve further study, both in pair formation and in the exploi- tation of food resources. These aspects, phero- mones and co-operation (and concomitant in- teraction) between the sexes, may reveal new insights in future ethological analyses of the nesting behaviour of dung beetles. Finally, a long-term study may reveal how much the dif- ferential development time of larvae and the dis- persal of adults contribute to reducing the risk of local population extinction and to stabilizing beetle numbers. Such studies may reveal the adaptive value of numerous aspects of dung beetle behaviour, about which we can only speculate at present. ACKNOWLEDGEMENTS Thanks are due to Prof. Dr. L. J. Pons and Ir. K. J. Hoeksema for suggesting that dung beetles are important agents in reworking soil, thereby providing the stimulus for the present study. The stimulating discussions I had with Prof. Dr. He Klomp; Dr) Ps J. den Boers, Dry Ths) van Dijk, Dr. S. Slager, Mrs. C. C. M. Brussaard- Wüst, Ir. A. F. M. Olsthoorn, Mr. J. Krikken and Dr. C. A. D. de Kort were very valuable. Their constructive comments certainly ım- proved the manuscript. The students F. W. B. M. Loonen and J. G. A. Outhuis were of great help in observing beetle behaviour. The stu- dents W. J. F. Visser and A. F. M. Olsthoorn assisted in the fieldwork. I would also like to make special mention of the interesting dis- cussions I had with Mr. T. J. Palmer and Dr. H. G. Klemperer during my stay in England. This study would have been impossible, how- ever, without technical assistance from P. Looijen, H. Snellen, Ing. P. W. T. Huisman, Ing. T. H. P. van Huizen, H. J. Immink, J. Mo- lenaar, G. Buurman, R. de Fluiter, W. C. Th. Middelplaats and Z. van Druuten. Their help is gratefully acknowledged. Mrs. J. Burrough- Boenisch corrected the English. 230 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 10, 1983 REFERENCES Baraud, J., 1977. Coléoptères Scarabaeoidea. Faune de PEurope occidentale. — Publ. Nouv. Revue Ento- mol. 4: 1—352. Toulouse, Univ. P. Sabatier. Blight, M. C. J., C. J. King, L. J. 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J. van Helsdingen (eds.): Methods and results of Euro- pean Invertebrate Survey mapping schemes in The Netherlands. — Nieuwsbrief European Inverte- BRUSSAARD: Reproduction and development of Typhaeus typhoeus 231 brate Survey Nederland 10: 81—88. Rommel, E., 1961. Ernährungsbiologie und Brutpfle- geverhalten des Spanischen Mondhornkäfers Co- pris hispanus (L.) (Col., Scarab.). — Biol. Zen- tralbl. 80: 327—346. Sano, C., 1915/16. On the metamorphosis of Geo- trupes stercorarius L. — Proc. South Lond. Ento- mol. Soc. 1915/16: 25—28. Saunders, D. S., 1976. Insect clocks. — Oxford, Per- gamon. Schreiner, J., 1906. Die Lebensweise und Metamor- phose des Rebenschneiders oder grossköpfigen Zwiebelhornkäfers (Lethrus apterus Laxm.) (Col., Scar.). — Horae Soc. Entomol. Ross. 37: 197— 207: Sinclair, M., 1977. Typhaeus typhoeus (L.) (Col., Geo- trupidae) in Scotland. — Entomol. Monthly Mag. 113: 166. Spaney, A., 1910. Beiträge zur Biologie unserer ein- heimischen Rosskäfer. — Deutsche Entomol. Z. 1910: 625— 634. , A., 1911. Lebensweise und Brutpflege unserer einheimischen Rosskäfer. — Entomol. Rundschau 28: 15, 24, 26, 34. Tauber, M. J. & C. A. Tauber, 1976. Insect seasonali- ty: diapause maintenance, termination, and post- diapause development. — Ann. Rev. Entomol. 21: 81—107. Teichert, M., 1955. Biologie und Brutfürsorgemass- nahmen von Geotrupes mutator Marsh. und Geo- trupes stercorarius L. (Col. Scarab.). — Wiss. Z. Martin Luther-Univ. Halle-Wittenberg 5: 187— 218. , 1956. Nahrungsspeicherung von Geotrupes ver- nalis L. und Geotrupes stercorosus Scriba (Co- leopt., Scarab.). — Wiss. Z. Martin Luther-Univ. Halle-Wittenberg 5: 669 —672. — — 1957. Soziale Instinkthandlungen einheimischer Geotrupini bei der Anlage ihrer Brutbauten. — Wiss. Z. Martin Luther-Univ. Halle-Wittenberg 6: 1045— 1048. , 1959a. Zum Brutfürsorgeverhalten des Geo- trupes vernalis L. (Col, Scarab.). — Wiss. Z. Mar- tin Luther-Univ. Halle-Wittenberg 8: 873— 878. , 1959b. Die bodenbiologische Bedeutung der co- prohagen Lamellicornier. — Wiss. Z. Martin Lu- ther-Univ. Halle-Wittenberg 8: 879— 882. Tribe, G. D., 1975. Pheromone release by dung beetles (Coleoptera, Scarabaeidae). — S. Afr. J. Sci. 71:277—278. Vlijm, L. & Th. S. van Dijk, 1967. Ecological studies on carabid beetles. II. General pattern of popula- tion structure in Calathus melanocephalus (Linn.) at Schiermonnikoog. — Z. Morph. Okol. Tiere 58:396—404. Weaver, R. J. & G. E. Pratt, 1977. The effect ot en- forced virginity and subsequent mating on the ac- tivity of the corpus allatum of Periplaneta ameri- cana measured in vitro, as related to changes in the rate of ovarian maturation. — Physiol. Entomol. 2: 59—76. i 4 + quiet, 5 À i en deo Arre a a AR a mes f i LS as iù 3 ‘alco EL, ere stà Ya iva ca Tr : ; } 3 È x N NET LU È eg RUE i 5 4 54 ; à 1744 ried Kernen LE À | (el LATE Le i” ne its . N i 1 il U "4 N ot ee en A A ane tl he bé A"; ar val Plan LIENS ET an fi 2 \ yet CHAN oY « A Wk i t cu ne on Aah uen me PAPE hiken nlt i 5 i if if } Kirn I ki RAMIRO E TA f Ri à | r SIA i ad: org data ] Dos | vay î i that NÀ: ? Fa i} Ay \ À SE HEY, NA ET CET UE IE DEN Sr er i; | TO ni be RON 4 ì | TRL n M I % arn, ni MO Le AR i À FIA par j AU ni i AIT Len i ñ st A sr iR 5 ; ae u u 1 È i 1 Der i | A 4 en u Il 1 ir Ue A Ji ri i bi a I N i) AF \ À | SE | EREN = ble. QE 4 | i ¢ Milo 1 IL fi DEEL 126 ESE NV RUNG et 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING LIBRARY HARVARD UNIVERSITY SR MUS. COMP. ZOOL INHOUD R. DE JONG. — Annotated list of the Hesperiidae (Lepidoptera) of Surinam, with descriptions of new taxa, pp. 233—268, figs. 1—69. Tijdschrift voor Entomologie, deel 126, afl. 11 Gepubliceerd 15-XII-1983 ANNOTATED LIST OF THE HESPERIIDAE (LEPIDOPTERA) OF SURINAM, WITH DESCRIPTIONS OF NEW TAXA by R. DE JONG Rijksmuseum van Natuurlijke Historie, Leiden ABSTRACT A list is given of all known species of Hesperiidae of Surinam, based on the literature and examined material. Material not yet recorded in the literature is listed with locality data. Notes are given on many species, and the following taxa are described as new: E/bella bicuspis (spec. nov.), Urbanus ambiguus (spec. nov.), Bungalotis sipa (spec. nov.), Clito jon- kersi (spec. nov.), Cymaenes geijskesi (spec. nov.), Vehilius major (spec. nov.), Mnasitheus similis (spec. nov.), Cobalopsis dorpa (spec. nov.), Cobalopsis tanna (spec. nov.), Morys sub- grisea paradoxa (subspec. nov.), Eutychide subpunctata intermedia (subspec. nov), Phlebodes meest (spec. nov.), Penicula criska extrema (subspec. nov.), Polites vibicoides (spec. nov.), Panoquina panoquinoides minima (subspec. nov.), Cravera rara (gen. nov. and spec. nov.), Surina unica (gen. nov. and spec. nov.). INTRODUCTION Surinam has been popular with lepidopterists and naturalists in general since the early days ot nature exploration. The famous water colours of Surinam insects by Maria Sibylla Merian, who lived from 1647 until 1717, testify of this early interest, and the large number of Lepido- ptera from Surinam described by Cramer (1775—1780, continued by Stoll, 1780—1791) speaks volumes. A general impression of ento- mological research in Surinam up to 1950 was given by Geijskes (1951). As far as Hesperiidae | are concerned three works are important, viz., Cramer (just mentioned), Sepp (1829—1852, for publication dates, see De Jong, in press), and Môschler (1876, 1882). Cramer described 78 species of Hesperiidae from Surinam, Sepp 21 species, and Môschler listed 145 species. The work of Sepp is not outstanding because of the number of species described or the quality of the plates, but owing to the very detailed de- scriptions of the larvae, pupae and food plants, all described species having been bred. Shortly before Sepp started the publication of the work on Surinam Lepidoptera, Collin’s the- sis on the fauna of Surinam was published (1822, see Holthuis, 1958). This rare work is of little importance, since as far as the butterflies are concerned it is a not very accurate compila- tion of the species mentioned by Fabricius (1793) from Surinam. 235) After Môschler, Surinam butterflies are only incidentally mentioned in the literature. In 1951, however, Evans (1951—1955) started the publication of the catalogue of all American Hesperiidae in the British Museum (Natural History), London. That museum proved to have 194 species from Surinam. In the Netherlands the interest in South American Lepidoptera, already diminished in the beginning of the 19th century in a period of general impoverishment, seemed to have come to a standstill after the publication of Sepp’s work, at least if judged from the complete lack of publications on Surinam butterflies up to the present time. The very limited edition of the lat- ter work is significant in this respect. Although a scientific approach to entomology was gaining ground at the time in the Netherlands, most en- tomologists were interested in the Dutch fauna or the fauna of the Dutch East Indies. Never- theless, limited amounts of material from Suri- nam were still sent to the Rijksmuseum van Na- tuurlijke Historie, Leiden, e.g. by W. C. van Heurn and the Penard brothers (1908), while the druggist Bolten, who lived in Paramaribo from 1902—1910, presented his small collection of insects to the museum. More important material was assembled much later. Dr. D. C. Geijskes, entomologist of the Agricultural Ex- periment Station at Paramaribo, later biologist at the Department of Home Affairs and Direc- 234 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 tor of the Surinam Museum at the same place, lived in Surinam from 1938—1965. He collected all kinds of insects during his expeditions into the interior all over the country. His material, now in the Rijksmuseum van Natuurlijke His- torie, is not only important because of its quan- tity, but because of the outlying districts repre- sented. As Geijskes, like most other collectors in Surinam, was not particularly interested in Lepidoptera, the number of species in his material is limited, but as he also collected many small and dull-coloured species (also by running a Malaise trap) instead of restricting himself to the bigger and more brightly coloured ones as too often done, his material is quite interesting. By far the best collection of Lepidoptera from Surinam was made by E. H. Jonkers, who lived in Surinam from 1956-1975. As an amateur- 58 Sz F6 56 55 54 mG Zum DE Jong: Hesperiidae of Surinam 235 naturalist he collected mainly Lepidoptera. His collection, which came to the Rijksmuseum van Natuurlijke Historie in 1977 and 1980, con- tained a large number of species, most being represented by rather few specimens. He mainly collected in the vicinity of Paramaribo, but also made many trips into the interior of Surinam. It is remarkable that some species represented in the smaller collection of Geijskes, are absent from the Jonkers collection, probably due to a different way of collecting. Smaller amounts of Lepidoptera from Suri- nam are received irregularly by the Rijksmu- seum van Natuurlijke Historie from biologists working in Surinam for a longer or shorter peri- od. Among these I like to mention my colleague Dr. G. F. Mees, who visited Surinam several times as an ornithologist and found time to make some interesting collections of Lepido- ptera. The total number of species of Hesperiidae known (or at least recorded) from Surinam now amounts to 426. Eight of these are, however, highly unlikely and probably due to false locali- ty data or misidentifications. It seems to be a fair number, but as 126 species of Hesperiidae recorded from Guiana and/or French Guiana have not yet been found in Surinam, and the in- terior of the Guianas is still largely unexplored lepidopterologically, the total number of skip- pers occurring in Surinam must be far more than 500 and probably even exceeds 600. In spite of the apparent incompleteness of our knowledge of the Surinam Hesperiidae it seems worthwhile to publish a list of what is known at present, one hundred years after the last list Fig. 1. Localities of Hesperiidae in Surinam. Black dots represent more or less exactly recorded localities; cir- cles placed over mountains (hatched areas) or rivers indicate that the locality is not exactly known and could be anywhere in the mountains or along the rivers indicated. Afobaka 22 Gransoela 39 Oelemari 42 Alalapadoe 47 Groningen 69 Onverdacht 9 Alamandidon 44 Albina 33 Hebiwerie 54 Palaomeu 46 Avanavero Vallen 63 Jodensavanne 17 Paramaribo 1 Paranam 10 Parwabos 3 Bakhuis Gebergte 60 Batavia 71 Kabalebo 61 Belwaarde 5 Kabelstation 24 Benzdorp 40 Kabo 65 Berlijn 13 Kayser Gebergte 49 Bigiston 34 Koffiekamp 23 Blakawatra 16 Kroetoe 55 Blauwgrond 4 Kwatta 2 Boniface 1 Brokopondo 21 Lawa 41 Brownsberg 20 Leiding 1 Lelydorp 7 Caledonia 70 Lely Gebergte 37 Carolina Kreek 12 Leonsberg 4 Charlesburg 1 Linker Coppename 52 Coeroeni Eiland 50 Lucie Rivier 51 Copi 26 Coppename 59 Makambi Kreek 24 Corantijn Rivier 74 Manlobbi 38 Cultuurtuin 1 Maratakka 73 Ma Retraite 4 Domburg 8 Mariénburg 6 Matapi 64 Ephraims Zegen 1 Matta 14 Moengotapoe 30 Fallawatra Kreek 57 Moengo 31 Morico Kreek 28 Gansee 25 Goliath Kreek 66 Nassau Gebergte 35 Patamakka Rivier 32 Pepejoe 45 Phedra 19 Potribo 27 Powakka 15 Raleigh Vallen 57 Rama 18 Rechter Coppename 53 Republiek 11 Sara Kreek 25 Saramacca 67 Sipaliwini 48 Stoelmans Eiland 36 Temomairem 43 Tibiti 68 Vier Kinderen 1 Voltzberg 56 Wageningen 72 Wia-wia 29 Wonotobo 62 Zanderij 12 Zorg en Hoop 1 236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 (Möschler, 1882). It does not seem likely that the number will increase largely in the near fu- ture. Moreover, several species are new to sci- ence, and publication may be a stimulus to fur- ther research. In view of the incomplete knowledge of the species composition in Surinam and of the dis- tribution of the recorded species in Surinam as well as in South America in general, no attempt has been made to analyse the species as to possi- ble distributional types. At the moment the im- pression is, that by far most of the species re- corded from Surinam have a wide distribution in tropical South America. ACKNOWLEDGEMENTS Few people, if any, are equally well acquaint- ed with the insect world of Surinam as Dr. D. C. Geijskes and I am most grateful for his help in various ways. It is sad that Mr. E. H. Jonkers ( 1979), whose collection formed the immedi- ate cause and main basis for the present paper, did not live to see this paper published or even prepared. His enthousiastic tales and his wish to have his collection safely kept in the Rijksmu- seum van Natuurlijke Historie are gratefully re- membered and acknowledged. Dr. W. Diehl (Zoologische Staatssammlung, München), Mr. W. Hogenes (Instituut voor Taxonomische Zoologie, Amsterdam), and Mr. R. I. Vane- Wright (British Museum (Natural History), London), kindly permitted the author free ac- cess to the collections under their care. Mr. E. Neering (temporarily in Paramaribo) was help- ful in sending material from his own collection and the Nationale Zoologische Collectie (Para- maribo) on loan. LIST OF THE HESPERIIDAE OF SURINAM In the list below I have followed the sequence and nomenclature adopted by Evans (1951— 1955), unless recent literature or the material studied gave cause to deviation. The species name is followed by a reference to the literature records from Surinam if any (records by Evans always refer to the collection of the British Mu- seum (Natural History), London), and the number and localities of specimens from Suri- nam in the following institutions: Instituut voor Taxonomische Zoölogie, Zoölogisch Museum, Amsterdam (abbreviated: ITZ); Nationale Zoölogische Collectie, Universiteit van Suri- name, Paramaribo (NZC); Rijksmuseum van Natuurlijke Historie, Leiden (RMNH; by far the most extensive collection of Surinam Hespe- riidae), and Zoologische Staatssammlung, — München (ZSM; all Surinam material originat- | ing from Fruhstorfer, no precise locality data). | All localities are given on a map (fig. 1). Notes | and descriptions of new taxa are given after the | list. Pyrrhopyginae Pyrrhopyge phidias Linnaeus, 1758. — Evans (1951); Hewitson (1866); Williams & Bell | (1931). RMNH: 6 d, 2 2, Brownsberg, Par- | amaribo, Surinam, Vier Kinderen. ZSM: 3 | 6,1 ©, Surinam. See Note 1. | Pyrrhopyge aziza lexos Evans, RMNH: 3 d, Brownsberg. N Pyrrhopyge proculus cintra Evans, 1951. — | Evans (1951). RMNH: 2 6, 2 2, Domburg, | Lely Gebergte, Patamakka Rivier, Surinam. ZSM:1 4,1 2, Surinam. | Pyrrhopyge thericles ponicea Evans, 1951. — | Evans (1951). RMNH: 2 d, 1 2, Rama, Su- rinam, Zanderij. ZSM: 2 d, Surinam. Pyrrhopyge amythaon polka Evans, 1951. — | RMNH:1 4,1 ©, Kabalebo, Republiek. Pyrrhopyge sergius semana Evans, 1951. — Evans (1951). ITZ: 2 ®, Paramaribo, Suri- nam. RMNH: 2 6, 2 2, Bakhuisgebergte, Brownsberg, Republiek, Surinam. Pyrrhopyge charybdis semita Evans, 1951. — RMNH: 1 5, Surinam. See Note 2. Pyrrhopyge amyclas amyclas Cramer, 1779. — Cramer (1779); Evans (4950 MEabrems (1793, “Hesperia amiatus”); Möschler (1876); Williams & Bell (1931). ITZ: 3 d, 5 ?, Paramaribo, Surinam. RMNH: 13 6, 12 ?, Paramaribo, Surinam, Wageningen (at light), Zorg en Hoop. ZSM: 1 6, Surinam. Pyrrhopyge arinas arinas Cramer, 1779. — Cramer (1779). RMNH: 1 à, Surinam (ho- WI lotype). Pyrrhopyge creusae Bell, 1931. — RMNH: 1 6, Sipaliwini. Pyrrhopyge rubricollis Sepp, 1841. — Sepp (1841). RMNH: 1 ©, Surinam. Pyrrhopyge cometes staudingeri Plötz, 1879. — Cramer (1779). RMNH: 1 d, Surinam. See Note 3. Elbella intersecta intersecta Herrich-Schäffer, 1869. — RMNH: 1 9, Charlesburg. Elbella patrobas blanda Evans, 1951. — Evans (1951). RMNH: 1 4,1 2, Domburg. Elbella bicuspis spec. nov. — See Note 4. Elbella alburna Mabille, 1891. — RMNH: 1 6, Voltzberg. Nosphistia zonara Hewitson, 1866. — Môschler DE Jone: Hesperudae of Surinam 237 (1876); possibly confused with a species of Jemadia or Elbella. Jemadia hospita ulixes Plötz, 1879. — Evans (1951); Moschler (1882); Plotz (1879). ITZ: 1 (without abdomen), Surinam. RMNH: 1 9, Surinam. Jemadia menechmus Mabille, 1878. — Evans (1951). Jemadia hewitsonii hewitsonit Mabille, 1878. — RMNH: 2 6, 2 2, Brownsberg, Domburg, Surinam. Jemadia gnetus- Fabricius, 1781. — Cramer (1779, “Papilio vulcanus”); Evans (1951). RMNH: 1 2, Surinam. Jemadia fallax fallax Mabille, 1878. — RMNH: 4 gd, Coeroeni, Coppename, Zanderij. See Note 5. Amenis pionia ponina Herrich-Schaffer, 1869. — ITZ: 1 (without abdomen), Surinam. Mysoria barcastus barcastus Sepp, 1851. — Cramer (1775, “Papilio acastus”); Evans (1951); Moschler (1876, “Pyrrhopyge acas- tus”; 1882, “Pyrrhopyge barcastus” and “Pyrrhopyge acastus”); Sepp (1851); Wil- liams & Bell (1931). ITZ: 6 6, 3 2, Surinam. RMNH: 28 4, 11 2, Belwaarde, Marién- burg, Moengotapoe-Wiawia, Paramaribo, Surinam, Zorg en Hoop. ZSM: 2 à, Suri- nam. Mysoria thasus thasus Stoll, 1781. — Stoll (1781). RMNH: 2 6,1 ©, Surinam. Myscelus nobilis Cramer, 1779. — Cramer (1779); Evans (1951); Fabricius (1781, “Pa- pilio salus”). Myscelus assaricus assaricus Cramer, 1779. — Cramer (1779); Môschler (1876). RMNH: 2 9, Surinam. Passova glacia Evans, 1951. — Evans (1951). Passova passova stacer Evans, 1951. — Evans (25 ENNE SPS vr arn ZSNES Surinam. Aspıtha aspitha parıma Plötz, 1886. — Plötz (1886). Pyrginae Phocides palemon palemon Cramer, 1779. — @ramer (779). Evans = (19529 SPhocides polybius polybius Fabricius, 1793”); Möschler (1876). ITZ: 1 ®, Paramaribo. NZC: 1 ©, Paramaribo. RMNH: 1 6,9 ©, Boniface, Carolina Kreek, Paramaribo, Suri- nam (including holotype, d, of Papilio pale- mon Cramer). Phocides metrodorus Bell, 1932. — NZC: 1 9, Republiek. RMNH: 1 4, Domburg. Phocides pigmalion hewitsonius Mabille, 1883. — Môschler (1876, “Eryades pygmalion Cr.”; determination uncertain, as he gave as synonym “Gnetus Ltrll.”, which belongs to Jerada) NZ Gan Zor-MEnMECop. RMNH: 2 4,29, Paramaribo, Surinam. See Note 6. Phocides lincea lincea Herrich-Schäffer, 1869. — RMNH: 1 2, Charlesburg. Tarsoctenus papias Hewitson, 1857. — Williams & Bell (1931). Tarsoctenus corytus Cramer, 1777. — Cramer (1777); Stoll (1782, “Papilio pyramus”). RMNH: 2 2, Surinam (including holotype of Papilio corytus Cramer). ZSM: 1 6, 2 9, Surinam. Tarsoctenus praecia rufibasis Mabille, 1910. — RMNH:4 4,1 2, Carolina Kreek, Surinam, Zanderij. ZSM: 1 à, Surinam. Phanus vitreus Stoll, 1781. — Evans (1952); Möschler (1876, 1882); Stoll (1781); Wil- liams & Bell (1931). RMNH: 1 6, 3 ©, Av- anavero Vallen, Carolina Kreek, Voltzberg. ZSM: 1 d, Surinam. Phanus obscurior Kaye, 1924. — Evans (1952). Phanus marshalli Kirby, 1880. — Evans (1952). RMNH: 9 ®, Avanavero Vallen, Browns- berg, Carolina Kreek, Jodensavanne, Rama, Surinam. ZSM: 3 6,3 2, Surinam. Udranomia orcinus Felder, 1876. — Moschler (1876). Udranomia kikkawai Weeks, 1906. — ZSM: 1 2, Surinam. Drephalys phoenice Hewitson, 1867. — Möschler (1882). ZSM: 1 ©, Surinam. Drephalys dumeril Latreille, 1824. — Evans (1952); Möschler (1876). Drephalys oriander oriander Hewitson, 1867. — Evans (1952). RMNH: 1 ®, Goliath Kreek. Drephalys talbott Le Cerf, 1922. — Evans (1952). Drephalys opifex Evans, 1952. — Evans (1952). Drephalys alcmon Cramer, 1779. — Cramer (1779); Evans (1952); Möschler (1876). RMNH: 1 ©, Zanderij. ZSM: 1 6, 1 2, Su- rinam. Augiades crinisus Cramer, 1780. — Cramer (1780); Evans (1952); Möschler (1882); Wil- liams & Bell (1931). ITZ: 1 6, 4 ©, Parama- ribo, Surinam, Zanderij. NZC: 1 à, Parama- ribo. RMNH: 5 d, 2 2, Lelydorp, Moengo, Paramaribo, Rama, Zanderij. ZSM: 2 8, 3 2, Surinam. Hyalothyrus infernalis Moschler, infernalis 238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 1876. — Evans (1952); Möschler (1876). RMNH: 8 6,3 ®, Avanavero Vallen, Caro- lina Kreek, Fallawatra Kreek, Goliath Kreek, Hebiwerie, Lely Gebergte, Raleigh Vallen (Voltzberg), Rama, Sipaliwini. Hyalothyrus leucomelas Geyer, 1832. — Möschler (1876). RMNH: 1 6,1 9, Avana- vero Vallen. ZSM: 1 d, Surinam. Hyalothyrus mitocris Stoll, 1782. — Evans (1952); Möschler (1882); Stoll (1782); Wil- harms 6% Bell (OU), NZ MINS Surinam RMNH: 5 6, 3 2, Goliath Kreek, Rama, Zanderij. ZSM: 11 3, 4 2, Surinam. Hyalothyrus neleus pemphygargyra Mabille, 1888. — Evans (1952); Möschler (1882). RMNH: 1 d, Coppename (Kroetoe). Phareas coeleste Westwood, 1852. — Evans (1952); Williams & Bell (1931). RMNH: 2 3, Sipaliwini. ZSM: 2 2, Surinam. Entheus eumelus Cramer, 1777. — Cramer (1777); Williams & Bell (1931, “Entheus mina”). RMNH: 1 ©, Goliath Kreek. ZSM: 1 8, Surinam (lacks the costal fold). Entheus gentius Cramer, 1779. — Cramer (1779); Evans (1952); Moschler (1882); Wil- liams & Bell (1931). ITZ: 1 3, Surinam. RMNH: 3 6,9 ©, Avanavero Vallen, Blaka- watra, Goliath Kreek, Lely Gebergte, Phe- dra, Rama, Republiek, Zanderij. ZSM: 5 ó, 5 2, Surinam. See Note 7. Entheus priassus Linnaeus, 1758. — Evans (1952); Möschler (1876, “Entheus talaus L.” and “Entheus peleus L.”); Stoll (1782); Wil- liams & Bell (1931). ITZ: 3 6,1 2, Surinam. RMNH: 5 ®, Albina, Avanavero Vallen, Brownsberg, Surinam, Zanderij. ZSM: 10 à, 5 ©, Surinam. Entheus matho dius Mabille, 1897. — Evans (052) SRMNINIEER RC MR PRG bEls ation, Kroetoe, Ramam, Surinam, Zanderij. Cabirus procas Cramer, 1777. — Cramer (1777); Evans (1952); Stoll (1787, “Papilio julettus”). RMNH: 2 d, Copi, Lely Ge- bergte. Proteides mercurius mercurins Fabricius, 1787. — Cramer (1779, “Papilio idas”); Möschler (1882, “Thymele idas Cr.”); Sepp (1848, “Papilio idas Cramer”). ITZ: 2 3, Surinam. RMNH: 1 8, Oelemari. Epargyreus clarus Cramer, 1775. — Cramer (1775; described from Surinam, but locality undoubtedly false, the species is not known to occur south of Honduras); Evans (1952; locality not trusted by Evans). Epargyreus socus sinus Evans, 1952. — Evans (952); Epargyreus exadeus Cramer, 1779. — Cramer (1779); Möschler (1882). RMNH: 1 6, Suri- nam. ZSM: 1 d, Surinam. Polygonus leo Gmelin, 1790. — Evans (1952). ITZ:2 d, Surinam. Chioides catillus catillus Cramer, 1779. — Cramer (1779); Evans (1952); Môschler (1876); Sepp (1847, “Papilio longicauda”); Williams & Bell (1931): TTZ 772 maribo, Surinam. RMNH: 18 d, 11 2, Albi- na, Bigiston, Blauwgrond, Brokopondo, Coppename Rivier, Goliath Kreek, Lely- dorp, Ma Retraite, Onverdacht, Paramaribo, Tibiti. ZSM: 4 d,1 2, Surinam. Aguna asander asander Hewitson, 1867. — Evans (1952). Aguna claxon Evans, 1952. — RMNH: 1 d, Brownsberg. See Note 8. Aguna aurunce Hewitson, 1867. — RMNH: 1 3,1 2, Brokopondo, Carolina Kreek. Aguna coelus Stoll, 1781. — Evans (1952); Möschler (1876); Stoll (1781). RMNH: 2 9, Lely Gebergte, Rama. ZSM: 2 2, Surinam. Aguna ganna Môschler, 1878. — Evans (1952); Williams & Bell (1931). Typhedanus orion Cramer, 1779. — Cramer (1779); Evans (1952); Möschler (1882); Wil liams & Bell (1931). ITZ: 1 d, 2 2, Parama- ribo, Surinam. RMNH: 10 6, 4 ©, Carolina Kreek, Domburg, Lelydorp, Paramaribo. Typhedanus undulatus Hewitson, 1867. — Evans (1952); Môschler (1876). ITZ: 2 6,1 2, Paramaribo, Surinam. RMNH: 3 4,4 9, Ma Retraite, Paramaribo, Zorg en Hoop. ZSM: 1 4, Surinam. Typhedanus optica optica Evans, 1952. — RMNH: 1 6,2 2, Brokopondo. ZSM: 1 6, Surinam. Polythrix octomaculata octomaculata Sepp, 1844. — Môschler (1882); Sepp (1844). RMNH: 1 6,5 2, Paramaribo. ZSM: 1 d, Surinam. Polythrix asine Hewitson, 1867. — RMNH: 1 ® , Paramaribo. See Note 9. Polythrix minvanes Williams, 1926. — ZSM: 1 d, Surinam. Polythrix caunus Herrich-Schätfer, 1869. — Evans (1952). RMNH: 4 d, 2 2, Brokopon- do, Paramaribo, Rama, Republiek. See Note 10. Polythrix auginus Hewitson, 1867. — Evans (1952); Möschler (1876). ZSM: 1 d, Suri- nam. DE JONG: Hesperiidae of Surinam 239 Polythrix metallescens Mabille, 1888. — RMNH: 3 d, 1 2, Ma Retraite, Republiek, Surinam. Polythrix eudoxus Stoll, 1781.— Stoll (1781). | Chrysoplectrum pervivax Hübner, 1819. — Evans (1952); Hübner (1819); Kirby (1871, “Pamphila schelleri”); Möschler (1876, “Te- legonus schelleri Kirby”); Sepp (1832—1840, “Papilio pertinax Stoll”); Stoll (1790, “ Papi- lio pertinax”); Williams & Bell (1931). RMNH: 2 2, Leonsberg, Paramarıbo. Chrysoplectrum bahiana babiana Herrich- Schäffer, 1869. — Evans (1952). RMNH: 2 6, Paramaribo, Surinam. Chrysoplectrum otriades Hewitson, 1867. — RMNH: 2 dg, Brownsberg. Chrysoplectrum perniciosus perniciosus Herrich- Schaffer, 1869. — Evans (1952). RMNH: 1 3,2 2, Brownsberg, Paramaribo. Chrysoplectrum orpheus Plotz, 1882. — ZSM: 1 6, Surinam. Urbanus proteus proteus Linnaeus, 1758. — Evans (1952); Moschler (1876); Williams & Bell (1931). ITZ: 2 6, 7 2, Caledonia-Sara- macca, Paramaribo, Surinam. RMNH: 8 4, 9 2, Albina, Mariénburg, Paramaribo, Ra- leigh Vallen. ZSM: 1 6, Surinam. Urbanus viterboana alva Evans, 1952. — Evans (1952). NZC: 1 d, Charlesburg. RMNH: 1 3, 4 2, Brokopondo, Paramaribo, Sipaliwi- ni. Urbanus esmeraldus Butler, 1877. — Evans (1952). RMNH: 1 6, Surinam. Urbanus esma Evans, 1952. — RMNH: 2 &, Ma Retraite, Zanderij. See Note 11. Urbanus esta Evans, 1952. — RMNH: 1 a, Domburg (agrees with esta, but valves sym- metrical, both like Evans’ figure of right valve). ZSM: 1 2, Surinam. Urbanus acawoios Williams, 1926. — Evans (1952). ITZ: 1 6, Paramaribo. RMNH: 1 ©, Lelydorp. ZSM: 1 ©, Surinam. Urbanus dorantes dorantes Stoll, 1790. — Evans (1952); Möschler (1882); Stoll (1790); Wil- lams SorBEl AOS DI ZEN ROE Surinam. RMNH: 11 6, 10 2, Albina, Ma Retraite, Paramaribo. ZSM: 2 6,1 ©, Surinam. Urbanus teleius Hübner, 1821. — Evans (1952); Möschler (1876); Williams & Bell (1931, “Goniurus eurycles Latr.”). RMNH: 8 d, 6 2, Domburg, Onverdacht, Paramaribo, Zanderij-Saramacca. Urbanus tanna Evans, 1952. — Evans (1952). Urbanus cindra Evans, 1952. — RMNH: 1 6, Copi. Urbanus ambiguus spec. nov. — See Note 12. Urbanus zagorus Plötz, 1881. — RMNH: 1 d, Surinam. Unlikely locality; only known from S. Brazil and Argentina. Urbanus simplicius Stoll, 1790. — Evans (1952); Môschler (1876, 1882); Stoll (1790); Wil- liams & Bell (1931). ITZ: 7 8,4 2, Parama- ribo, Surinam. RMNH: 38 6, 17 2, Albina, Avanavero Vallen, Bigiston, Brokopondo, Brownsberg, Charlesburg, Domburg, Gro- ningen, Lelydorp, Ma Retraite, Paramaribo, Republiek, Zanderij, Zorg en Hoop. ZSM: 1 6, Surinam. Urbanus procne Plötz, 1881. — Evans (1952). ITZ: 4 d, 2 ®, Charlesburg, Paramaribo. NZC: 2 d, 2 2, Paramaribo. RMNH: 42 6, 17 2, Charlesburg, Cultuurtuin, Domburg, Kabelstation, Kwatta, Lelydorp, Ma Re- traite, Paramaribo, Republiek. Urbanus doryssus doryssus Swainson, 1831. — Evans (1952); Moschler (1876); Williams & Bellas) ZEN ESR MS urn an NZ d, Benzdorp. RMNH: 5 6, Brownsberg, Corantijn Rivier, Domburg, Paramaribo. ZSM:5 6,4 ®, Surinam. Urbanus albimargo takuta Evans, 1952. — Evans (1952); Williams & Bell (1931). RMNH: 2 ®, Avanavero Vallen, Browns- berg. ZSM: 1 à, Surinam. Urbanus virescens Mabille, 1877. — ZSM: 16, Surinam. Astraptes talus Cramer, 1777. — Cramer (1777); Môschler (1876); Sepp (1841, “Papi- how laadator) AZ 122027 Paramanıbo. RMNH: 4 6, 4 ©, Albina, Carolina Kreek, Leonsberg, Paramaribo, Surinam. ZSM: 1 ®, Surinam. Astraptes fulgerator fulgerator Walch, 1775. — Evans (1952); Möschler (1876); Sepp (1841, “Papilio fulminator”); Williams & Bell (1931). ITZ: 1 ©, Surinam. RMNH: 4 d, 7 2, Brownsberg, Domburg, Ephraims Zegen, Kabalebo/Corantijn Rivier, Lely Gebergte, Paramaribo, Surinam. ZSM: 3 6,5 ©, Suri- nam. Astraptes apastus apastus Cramer, 1777. — Cramer (1777); Evans (1952). ITZ: 1 8,1 2, Surinam. RMNH: 1 6,1 9, Domburg, Suri- nam. ZSM: 1 6, Surinam. Astraptes enotrus Stoll, 1781. — Stoll (1781). RMNH: 1 6, Rama. ZSM: 1 6, Surinam. Astraptes granadensis Möschler, 1878. — Evans (1952); Möschler (1882); Stoll (1780, “Papi- lio aulestes”). ITZ: 1 &, Surinam. RMNH: 1 6, Surinam. ZSM: 1 6, Surinam. 240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Astraptes narcosius Stoll, 1790. — Evans (1952, ssp. narcosius and ssp. aulina Evans, 1952); Stoll (1790). RMNH: 1 d, Surinam (ssp. narcosius). Astraptes alardus Stoll, 1790. — Evans (1952); Möschler (1876); Stoll (1790). ITZ: 3 ®, Domburg, Paramaribo, Surinam. RMNH: 6 3, 6 2, Charlesburg, Domburg, Paramari- bo. Astraptes alector hopfferi Plötz, 1882. — Möschler (1882, “Thymele hopfferi Plotz”). RMNH: 2 6, 1 2, Brownsberg, Lucie Riv- ıer, Rama. Astraptes cretatus Hayward, 1939. — RMNH: 1 8, 1 2, Carolina Kreek, Rama. See Note We Astraptes creteus creteus Cramer, 1780. — Cramer (1780); Evans (1952); Moschler (1876, “Telegonus parmenides Cramer”; 1882, previous record was based on misiden- tification, referred to “Thymele Hopfferi”); Stoll (1781, “Papilio parmenides”). ITZ: 1 2, Surinam. NZC: 2 6, Phedra. Astraptes latimargo Herrich-Schäffer, 1869. — Moschler (1882); Williams & Bell (1931). Astraptes chiriquensis oenander Hewitson, 1876. — ZSM: 1 6, Surinam. Astraptes anaphus anaphus Cramer, 1777. — Cramer (1777); Evans (1952); Sepp (1830, “Papilio leucogramma”). ITZ: 1 ®, Surinam. RMNH: 1 6, 3 ©, Brokopondo, Surinam (including d holotype, Surinam). Calliades zeutus Möschler, 1878. — ZSM: 1 9, Surinam. Autochton neis Geyer, 1832. — Evans (1952); Möschler (1876, “Cecropterus bocus Hopf- fer”); Williams & Bell (1931, “Cecropterus neis Geyer” and “Cecropterus bocus Plötz”). ITZ: 1 g, Surinam. RMNH: 3 d, Domburg, Surinam. ZSM: 1 ©, Surinam. Autochton longipennis Plôtz, 1882. — Williams & Bell (1931, “Cecropterus capys (Godman & Salvin”). ITZ: 1 2, Surinam. RMNH: 5 3, Brownsberg, Domburg, Sipaliwini, Suri- nam. ZSM: 1 d, Surinam. Autochton zarex Hubner, 1818. — Hubner (1818); Môschler (1876); Williams & Bell (1931, “Cecropterus aunus Fabricius”). RMNH: 9 d, 3 ®, Albina, Brownsberg, Carolina Kreek, Paramaribo, Surinam, Zan- derij. ZSM: 1 4,3 2, Surinam. Autochton bipunctatus Gmelin, 1790. — Moschler (1876). RMNH: 1 2, Albina. Autochton itylus Hubner, 1823. — Evans CO Brownsberg, Paramaribo. ZSM: 3 d, 2 9, Surinam. Thorybes daunus Cramer, 1777. — Cramer (1777); apparently wrong locality for this strictly North American species. Bungalotis erythus Cramer, 1775. — Cramer (1775). RMNH: 1 2, Kabalebo (at black light). Bungalotis diophorus Möschler, 1882. — Möschler (1882). Bungalotis midas Cramer, 1775. — Cramer (1775); Evans (1952); Fabricius (17819: Möschler (1876); Plötz (1882a). NZC: 1 9, Alalapadoe. RMNH: 2 d,2 2, Cultuurtuin, Paramaribo, Sarakreek. ZSM: 1 &, Surinam. Bungalotis astylos Cramer, 1780. — Cramer (1780); Möschler (1876). RMNH: 1 d,1 ?, Paloemeu, Surinam. ZSM: 1 d, Surinam. Bungalotis borax Evans, 1952. — RMNH: 1 d, 1 9, Nassau Gebergte, Saramacca Rivier. Bungalotis sipa spec. nov. — See Note 14. Bungalotis quadratum Sepp, 1845. — Evans (1952); Môschler (1876, “Telegonus annuli- cornis”); Sepp (1845). NZC: Paramaribo. RMNH: 1 6, 1 9, Paramaribo, Saramacca Rivier. See Note 15. Salatis salatis Stoll, 1782. — Evans (1952); Plötz (1882a); Stoll (1782). RMNH: 1 6, 2 9, Ephraims Zegen, Lely Gebergte, Onver- dacht. Salatis cebrenus (1777). Salatis fulvius Plôtz, 1882. — RMNH: 1 9, Wonotobo. Salatis flavomarginatus Sepp, 1851. — Sepp (1851). Sarmientoia eriopis Hewitson, 1867. — Evans (1952). RMNH:1 ©, Onverdacht. Dyscophellus euribates Stoll, 1782. — Evans (1952); Möschler (1876, “Telegonus hesus Westwood”); Stoll (1782). RMNH: 3 6, 1 2, Goliath Kreek (at light), Saramacca Riv- ier, Surinam. ZSM: 2 d, Surinam. Dyscophellus porcins Felder, 1862. — Evans (1952); Möschler (1882, “Thymele doriscus Hewitson”). Dyscophellus sebaldus Stoll, 1781. — Evans (1952); Stoll (1781). RMNH: 2 2, Sipalrwini (at light), Surinam. Dyscophellus erythras Mabille, 1888. — Evans (1952). Dyscophellus diaphorus Mabille & Boullet, 1912. — Mabille & Boullet (1912). Dyscophellus ramusis Stoll, 1781. — Evans (1952); Möschler (1876, 1882); Stoll (1781). Cramer, 1777. — Cramer DE JONG: Hesperiidae of Surinam 241 NZC: 1 9, Kabo. RMNH: 2 6,1 ©, Afoba- ka, Goliath Kreek, Surinam. Nascus phocus Cramer, 1777. — Cramer (1777); Moschler (1876, “Telegonus pherenice He- witson”); Sepp (1843, “Papilio decemmacu- lata”); Stoll (1782, “Papilio morpheus”). NZE: 1 ©, Paramaribo. RMNH: 2 6,1 ©, Domburg, Paramaribo, Surinam. Nascus solon Plötz, 1882. —RMNH: 1 ©, Suri- nam. Nascus broteas Cramer, 1780. — Cramer (1780). ZSM: 2 d, Surinam. Nascus paulliniae Sepp, 1842. — Evans (1952); Sepp (1842). RMNH: 1 6, 1 ©, Domburg, Paramaribo (at light). ZSM: 2 à, Surinam. Cephise cephise Herrich-Schäffer, 1869. — Evans (1952); Möschler (1876). Porphyrogenes passalus passalus Herrich- Schaffer, 1869.— RMNH: 1 6, Rama. Porphyrogenes probus Möschler, 1876. — Möschler (1876). Porphyrogenes zohra Möschler, 1878. — RMNH: 1 d, Coppename Rivier. Porphyrogenes pausias Hewitson, 1867. — RMNH:1 dg, Zanderij. ZSM: 1 d, Surinam. Ablepsis azines Hewitson, 1867. — Mabille & Boullet (1912, “Telemiades perseus”). RVING Zanderij. ZSM: 1 S, 109 Su- rinam. Orneatus aegiochus Hewitson, 1876. — Möschler (1882), probably confused with an Astraptes species, O. aegiochus being known from C. America only. Celaenorrbinus shema shema Hewitson, 1877. — Evans (1952); Môschler (1882, “Plesio- neura ochrogutta”). Celaenorrhinus similis bifurcus Bell, 1934. — Evans (1952). Celaenorrhinus spec. nov. — Will be described in a separate paper. Celaenorrhinus eligius eligius Stoll, 1782. — Evans (1952); Stoll (1782); Williams & Bell (1931). Celaenorrhinus syllins Felder, 1862. — Evans (1952). RMNH: 1 2, Brownsberg. Spathilepia clonius Cramer, 1775. — Evans (1953); Möschler (1876); Williams & Bell (1931). RMNH: 7 6, 3 ©, Brokopondo, Domburg, Paramaribo. Cogia hassan Butler, 1870. — Williams & Bell (1931, “Cogia freudiae”). RMNH: 13 d, 6 9, Temomairem, Zanderij. See Note 16. | Cogia calchas Herrich-Schaffer, 1869. — Möschler (1876). RMNH: 2 d, 1 ©, Broko- pondo, Rama. Telemiades vansa Evans, 1953. — ZSM: 6 @, Surinam. Telemiades squanda Evans, 1953. — RMNH: 1 ®, Brokopondo. Telemiades trenda Evans, 1953. — Evans (1953). Telemiades nicomedes Möschler, 1878. — Evans (1953). Telemiades epicalus Hübner, 1819. — Evans (1953). Telemiades penidas Hewitson, 1876. — Möschler (1876, “Telegonus mygdon”); Wil- liams & Bell (1931, “Telemiades ceramina Herrich-Schäffer”). RMNH: 1 6, 1 9, Bro- kopondo, Brownsberg. Telemiades avitus Stoll, 1781. — Stoll (1781); Williams & Bell (1931). Telemiades amphion amphion Hübner, 1826. — Môschler (1876, “Plesioneura compressa”); Williams & Bell (1931). ITZ: 1 2, Surinam. RMNH: 3 d, 2 ©, Domburg, Rama, Sipali- winl. Pyrdalus corbulo Stoll, 1781. — Môschler (1876); Stoll (1781). RMNH: 1 6, Surinam. ZSM:1 6,1 ®, Surinam. Ectomis cythna Hewitson, 1878. — Evans (1953). RMNH: 1 ©, Brokopondo (the first known female of this scarce species). Conognathus platon Felder, 1862. — RMNH: 1 3, Saramacca Rivier. Arteurotia tractipennis Butler & Druce, 1872. — Evans (1953). Eracon paulinus Stoll, 1782. — Stoll (1782). RMNH:1 ©, Lely Gebergte. Eracon onorbo Möschler, 1882. — Evans (1953); Möschler (1882). Spiomades artemides Stoll, 1782. — Evans (1953); Môschler (1876); Stoll (1782). RMNH: 1 2, Zanderij. ZSM: 2 g, Surinam. Spioniades libethra Hewitson, 1868. — Evans (1953). Polyctor polyctor Prittwitz, 1868. — RMNH: 1 d, Zanderij. ZSM: 1 ©, Surinam. Nisoniades bessus Môschler, 1876. — Evans (1953); Möschler (1876). RMNH: 4 6, Bro- kopondo, Paramaribo, Zanderij. Nisoniades laurentina Williams & Bell, 1939. — RMNH: 1 d, Brokopondo. Nisoniades rubescens Möschler, 1876. — Evans (1953); Möschler (1876); Williams & Bell (1931, “Pellicia bromias Godman & Salvin”). RMNH: 5 d, Domburg, Ma Retraite, Para- maribo. Nisoniades mimas Cramer, 1775. — Cramer (1775); Evans (1953); Sepp (1845, “Papilio 242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 bromius Stoll”); Stoll (1790, “Papilio bro- mius”). RMNH: 1 d, Sipaliwini. Nisoniades ephora Herrich-Schäfter, 1870. — Möschler (1882). Nisoniades rimana Bell, 1942. — RMNH: 1 dg, Brownsberg. ZSM: 1 d, Surinam. Nisoniades brunneata Williams & Bell, 1939. — Evans (1953). Nisoniades macarius Herrich-Schaffer, 1870. — Williams & Bell (1931). NZC: 1 à, Alaman- didon. Pachyneuria duidae Bell, 1932. — Evans (1953). Pellicia dimidiata Herrich-Schäffer, 1870. — Möschler (1876, “Pellicia didia”). Morvina fissimacula rema Evans, 1953. — ZSM: 1 ©, Surinam. Myrinia binoculus Möschler, 1876. — Môschler (1876). Cyclosemia herennius Stoll, 1782. — Evans (1953); Stoll (1782). Viola violella Mabille, 1897. — Evans (1953). RMNH:2 6,1 ©, Paramaribo. Plumbago plumbago Plôtz, 1884. — RMNH: 1 2, Paramaribo. Gorgythion begga pyralina Möschler, 1876. — Evans (1953); Môschler (1876, “Nisoniades plautia” and “Helias pyralina”); Williams & Bell (1931). NZE: 1 2, Potribo. RMNHE 1 d, Zanderij. ZSM: 1 ©, Surinam. Gorgythion beggina escalophoides Hayward, 1941. — RMNH: 6 d, 6 ©, Brokopondo, Domburg, Paramaribo. Gorgythion canda Evans, 1953. — ZSM: 1 2, Surinam. Identification must remain prelimi- nary, as long as female genitalia of Gorgy- thion species have not been studied. Ouleus matria dampa Evans, 1953. — ZSM: 1 2, Surinam. Ouleus fridericus fridericus Geyer, 1832. — Evans (1953); Môschler (1876); Williams & Bell (1931). RMNH: 8 6, 1 ©, Albina, Domburg, Ma Retraite, Paloemeu, Parama- ribo, Surinam. Zera tetrastigma Sepp, 1847. — Sepp (1847). RMNH:1 2, Blakawatra. Quadrus cerialis Stoll, 1782. — Evans (1953, “cerealis”, incorrect subsequent spelling); Möschler (1876); Sepp (1847); Stoll (1782); Williams & Bell (1931). ITZ: 2 8,2 9, Para- maribo, Surinam. RMNH: 8 6,6 ©, Avana- vero Vallen, Lely Gebergte, Ma Retraite, Paramaribo, Surinam. ZSM: 1 d, 3 ©, Suri- nam. Quadrus fanda Evans, 1953. — ZSM: 1 6, Suri- nam. Quadrus contubernalis contubernalis Mabille, 1883. — RMNH: 3 d, Blakawatra, Browns- berg. ZSM: 1 ©, Surinam. Quadrus deyrolli deyrolli Mabille, 1877. — Evans (1953). RMNH: 2 d, 1 2, Browns- berg, Copi, Zanderij. ZSM: 6 d, Surinam. Gindanes brebissoni Latreille, 1824. — RMNH: 1 2, Carolina Kreek. Comes closest to ssp. phagesia Hewitson, 1868. Pythonides jovianus jovianus Stoll, 1782. — Evans (1953); Möschler (1876); Stoll (1782); Williams & Bell (1931). ITZ: 1 &, 1 2 Suri- nam. RMNH: 6 d, 7 2, Albina, Avanavero Vallen, Brownsberg, Lely Gebergte, Nassau Gebergte, Rama, Surinam, Zanderij, Zande- rij-Matta. ZSM: 6 6, 4 9, Surinam. Pythonides lerina Hewitson, 1868. — Evans (1953); Môschler (1876); Williams & Bell (1931). RMNH: 3 d, Rama, Surinam, Zan- derij. ZSM: 6 6, 1 2, Surinam. Pythonides grandis assecla Mabille, 1883. — RMNH: 2 d, Koffiekamp, Rama. Pythonides herennius herennius Geyer, 1838. — ITZ: 1 d, Surinam. RMNH: 3 6, Surinam. Pythonides limaea limaea Hewitson, 1868. — Evans (1953); Möschler (1882). Sostrata festiva Erichson, 1848. — Evans (1953). ITZ21&, Sura mR MINE seni 2, Brokopondo, Surinam. ZSM: 1 6, Suri- nam. Sostrata bifasciata adamas Plötz, 1884. — Evans (1953). Paches exosa Butler, 1877. — Plötz (1884, “Py- thomides prudens”). Milanion hemes hemes Cramer, 1777. — Cram- er (1777); Evans (1953); Môschler (1876, 1882). ITZ: 1 6, Surinam. RMNH: 3 6, 3 2, Brokopondo, Brownsberg, Sipaliwini, Surinam, Zanderij. Milanion leucaspis Mabille, 1878. — ZSM: 1 6, Surinam. Milanion pilumnus albidior Mabille & Boullet, 19172 ZMS MS Suriname Paramimus scurra Hübner, 1809. — Evans (1953); Môschler (1882, “Pythonides leuco- desma Erichson”); Williams & Bell (1981). ITZ: 1 d, Surinam. RMNH: 5 6,2 ©, Albi- na, Blakawatra, Brownsberg, Surinam, Zan- derij. ZSM: 1 6,6 9, Surinam. Charidia lucaria lucaria Hewitson, 1868. — Evans (1953); Williams & Bell (1931). RMNH: 1 2, Brownsberg. ZSM: 1 2, Suri- nam. Mylon menippus Fabricius, 1776. — Cramer (1782, “Papilio melander”); Fabricius DE JONG: Hesperudae of Surinam 243 (1776); Möschler (1876, “Achlyodes melan- der Cramer”). ITZ: 1 d, Surinam. RMNH: 2 ©, Brownsberg. Mylon jason Ehrmann, 1907. — Evans (1953); Williams & Bell (1931). Carrhenes fuscescens conia Evans, 1953. — ZSM: 1 2, Surinam. Carrhenes canescens leada Butler, Evans (1953). Clito jonkersi spec. nov. — See Note 17. Clito clito Fabricius, 1787. — Möschler (1876). ZSM: 1 d, Surinam. Clito zenda Evans, 1953. — Evans (1953). Xenophanes tryxus Stoll, 1780. — Evans (1953); Möschler (1876); Stoll (1780); Williams & Bell (1931). ITZ: 2 g, Surinam. RMNH: 12 d, 5 2, Brokopondo, Brownsberg, Lely- dorp, Ma Retraite, Paramaribo, Sipaliwini, Surinam. ZSM: 2 ¢, Surinam. Antigonus erosus Hübner, 1812. — Evans (1953); Möschler (1876, “Antigonus wester- manni Latreille”; 1882). Timochreon doria Plötz, 1884. — RMNH: 1 ©, Rama. Sb == Anisochoria pedaliodina polysticta Mabille, | 1876. — Evans (1953). Achlyodes busirus busirus Stoll, 1782. — Möschler (1882); Stoll (1782). RMNH: 3 8, 1 2, Brownsberg, Surinam, Zanderij. Achlyodes thraso thraso Jung, 1792. — Evans (1953); Möschler (1882); Williams & Bell MN ZG MIN SC MCE Paramaribo: RMNH: 9 d, 6 ©, Brownsberg, Ma Re- traite, Paramaribo, Zanderij. ZSM: 1 6, Su- rinam. Anastrus sempiternus simplicior Möschler, 1876. — Möschler (1876). RMNH: 1 2, Ma Re- traite. Anastrus tolimus robigus Plötz, 1884. — Evans (1953). RMNH: 3 G, 4 2, Blakawatra, Bro- kopondo, Paramaribo, Patamakka Rivier, Surinam. Anastrus petins Môschler, 1876. — Evans (1953); Möschler (1876). ZSM: 5 d,2 2, Su- rinam. Anastrus obliqua Plôtz, 1884. — RMNH: 1 9, Zanderij. Anastrus obscurus narva Evans, 1953. — Evans (1953). ITZ: 3 8,2 9, Paramaribo, Surinam. RMNH: 3 6,2 ©, Brokopondo, Lelydorp, Matapı. ZSM: 1 6,2 9, Surinam. Ebrietas infanda Butler, 1876. — RMNH: 1 9, Brownsberg. Cycloglypha thrasibulus thrasibulus Fabricius, W723), —= levains (11055) IRVINE 1 dé 208 Brokopondo, Rama, Surinam. ZSM: 1 d Surinam. Cycloglypha caeruleonigra Mabille, 1904. — RMNH: 2 dg, Paramaribo, Sipaliwini. Cycloglypha enega Möschler, 1876. — Möschler (1876). RMNH: 2 4, 1 2, Broko- pondo, Paloemeu, Paramaribo. Helias phalaenoides phalaenoides Fabricius, 1807. — Evans (1953); Williams & Bell (19311) ZnS Sunmam RVING 3 9, Brownsberg, Coeroeni Eiland, Surinam. Chiomara asychis asychis Stoll, 1780. — Evans (1953); Möschler (1876), “Pythonides diluci- da” and “Achlyodes asychis Cramer”); Stoll (1780); Williams & Bell (1931). RMNH: 8 d, 5 2, Belwaarde, Kwatta, Ma Retraite, Mariënburg, Paramaribo, Surinam, Wia-wia. Chiomara mithrax Môschler, 1878. — Evans (1953). RMNH: 1 ©, Paramaribo. Chiomara punctum Mabille, 1878. — Evans (1953); Williams & Bell (1931). RMNH: 1 3,5 2, Copi, Ma Retraite, Zanderij. ZSM: 3 6,2 2, Surinam. Ephyriades arcas philemon Fabricius, 1775. — Moschler (1876, “Nisoniades otreus Cra- mer”); Stoll (1780, “Papilio otreus”, and “Papilio flyas”). RMNH: 1 6, Surinam. Pyrgus oileus orcus Stoll, 1780. — Evans (1953); Hubner (1809, “Papilio tartarus”); Moschler (1876, “Hesperia syrichtus Fabricius”); Stoll (1780); Williams & Bell (1931). ITZ: 6 d, 4 2, Paramaribo, Surinam. RMNH: 35 d, 21 2, Benzdorp, Brokopondo, Coeroeni Eiland (in Coeroeni Rivier), Domburg, Kabelsta- tion, Kwatta, Ma Retraite, Paramaribo, Par- anam, Stoelmans Eiland (in Marowijne). Brown & Heineman (1972: 389) considered orcus specifically distinct from ozleus, but ev- idence of real sympatry is not yet convinc- ing. Heliopetes arsalte arsalte Linnaeus, 1758. — Evans (1953); Möschler (1882); Williams & Bell (1931). ITZ: 3 d, 2 ®, Caledonia-Sara- macca, Paramaribo, South side Upper Sara- macca, Surinam. RMNH: 20 6,9 ©, Albina, Kayser Gebergte, Kwatta, Leiding, Lely- dorp, Maratakka, Paramaribo, Sipaliwini. AISNE DO PE Suriname Heliopetes leucola Hewitson, 1868. — Môschler (1882), probably confused this rare Brazilian species with the next species, which is com- mon in Surinam. Heliopetes alana Reakirt, 1868. — Evans 244 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 (1953). RMNH: 7 6, 2 2, Brokopondo, Goliath Kreek, Rama, Sipaliwini. ZSM: 1 9, Surinam. Hesperiinae Synapte silius Latreille, 1824. — RMNH: 1 4, Domburg. Lento ferrago Plötz, 1884. — RMNH: 1 9, Morico Kreek. Lento lento Mabille, 1878. — ITZ: 1 d, Repub- liek. Zariaspes mys Hübner, 1808. — Williams & Bell (1931). RMNH: 5 6, 5 ©, Domburg, Lelydorp, Ma Retraite, Surinam. Anthoptus epictetus Fabricius, 1793. — Evans (1955); Môschler (1882); Williams & Bell (1931). ITZ: 2 d, 1 ©, Leiding, Paramaribo. RMNH: 15 6, 1 ®, Brokopondo, Dom- burg, Goliath Kreek, Kayser Gebergte, Par- amaribo, Rama, Sipaliwini, Surinam. See Note 18. Corticea corticea corticea Plötz, 1883. — Evans (1955); Williams & Bell (1931, “Megistias noctis Plötz”). RMNH: 6 6,3 2, Domburg, Ma Retraite, Paramaribo, Surinam, Zanderij. Cantha celeus Mabille, 1891. — Williams & Bell (1931). Vinius tryhana Kaye, 1913. — NZC: 1 6, Pe- pejoe. RMNH: 3 d, Domburg, Ma Retraite, _ Paramaribo. See Note 19. Molo mango Guenée, 1865. — Evans (1955). RMNH: 3 d, Nassau Gebergte, Surinam. ZSM: 1 d,Surinam. Molo menta Evans, 1955. — Evans (1955). Apaustus gracilis smarti Evans, 1955. — Wil- liams & Bell (1931). RMNH: 4 6, 1 9, Av- anavero Vallen (Malaise trap), Raleigh Val- len. Apaustus menes Stoll, 1782. — Evans (1955); Möschler (1876); Sepp (1842); Williams & Bell (1931). RMNH: 3 6, Ma Retraite, Para- maribo. ZSM: 1 gd, Surinam. See Note 20. Callimormus radiola radiola Mabille, 1878. — Evans (1955); Williams & Bell (1931). RMNH: 3 d,1 2, Brownsberg, Domburg. Callimormus alsimo Môschler, 1882. — Möschler (1882). ZSM: 1 ®, Surinam. Callimormus juventus Scudder, 1872. — Evans (1955). RMNH: 1 6, 1 2, Kwatta, Lely- dorp. Callimormus corades Felder, 1862. — Williams & Bell (1931). RMNH: 4 d,1 2, Domburg, Lelydorp, Paramaribo, Zorg en Hoop. Callimormus saturnus Herrich-Schäffer, 1869. Ee (955), TZ: 2 2. Pammanbo. RMNH: 2 6, 1 2, Ma Retraite, Zorg en Hoop. Eutocus facilis Plötz, 1884. — Plotz (1884). Eutocus fabulinus Plotz, 1884. — Plotz (1884). Eutocus matildae vinda Evans, 1855. — RMNH: 3 6, Avanavero Vallen, Ma Re- traite, Surinam. Virga virginius Moschler, 1882. — Möschler (1882). Methionopsis ina Plötz, 1882. — Evans (1955). RMNH: 2 6, 3 ©, Domburg, Ma Retraite, Paramaribo. Sodalia sodalis Butler, 1877. — Evans (1955); Williams & Bell (1931, “Euroto saramacca”). RMNH: 3 6,3 ©, Ma Retraite, Paramaribo. Mnestheus servilius Moschler, 1882. — Moschler (1882). Artines aepitus Geyer, 1832. — Williams & Bell (1931, “Artines atizies Godman”). RMNH: 1 dg, 1 2, Goliath Kreek, Surinam. ZSM: 2 6,1 2, Surinam. Aecas aecas Stoll, 1781. — Stoll (1781); Wil- liams & Bell (1931). RMNH: 2 dg, 4 9, Domburg, Ma Retraite, Surinam, Zanderij. ZSM: 1 &, Surinam. Mnaseas bicolor Mabille, 1889. — Williams & Bell (1931). Thargella caura Plötz, 1882. — Möschler (1882); Plötz (1882b); Williams & Bell (1931, “Thargella fuliginosa Godman”). RMNH: 2 6, Domburg. Venas evans Butler, 1877. — RMNH: 1 4, Lely Gebergte. ZSM:9 6,2 9, Surinam. Venas caerulans Mabille, 1878. — ZSM: 1 d, Surinam. Monca telata Herrich-Schäffer, 1869 — Môschler (1882). Nastra chao Mabille, 1897. — Williams & Bell (1931, “Megistias huascari Lindsey”). It is not certain, if the name given by Lindsey is really a synonym of Mabille’s name. Nastra guianae Lindsey, 1925. — RMNH: 5 d, 4 ®, Charlesburg, Paramaribo, Zanderij, Zorg en Hoop. See Note 23. Cymaenes tripunctus theogenis Capronnier, 1874. — Evans (1955); Möschler (1882, “Pamphila ancus”); Williams & Bell (1931). RMNH: 2 d, 6 ®, Charlesburg, Kwatta, Lelydorp, Ma Retraite. Cymaenes geijskesi spec. nov. — See Note 21. Cymaenes tripunctata alumna Butler, 1877. — Evans (1955); Môschler (1882, “Pamphila obsoleta”); Williams & Bell (1931). NZC: 1 ©, Manlobbi. RMNH: 1 d, 2 2, Charles- burg, Domburg, Paramaribo. DE JONG: Hesperudae of Surinam Vehilius stictomenes stictomenes Butler, 1877. — Evans (1955); Môschler (1882, “Apaustus venosus Pittw.”); Williams & Bell (1931, “Vehilius venosus Plötz”). NZC: 1 6, Para- maribo. RMNH: 5 g, 7 2, Avanavero Val- len, Domburg, Lelydorp, Ma Retraite, Para- maribo, Zanderij. ZSM: 2 d, Surinam. Vehilius inca Scudder, 1872. — Evans (1955); Williams & Bell (1931, “Megistias labdacus Godman INZE: 71778, Paramaribo. RMNH: 7 6, 2 ©, Domburg, Kwatta, Lely- dorp, Ma Retraite, Paramaribo, Zorg en Hoop. Vehilius almoneus Schaus, 1902. — ITZ: 1 9, Paramaribo. RMNH: 5 6, 6 9, Charles- burg, Domburg, Kwatta, Paramaribo. Vehilius major spec. nov. — See note 22. Vehilius vetulus Mabille, 1878. — Evans (1955). RMNH: 2 d, Brownsberg, Lely Gebergte. Vehilius seriatus seriatus Mabille, 1891. — RMNH: 1 d, Coeroeni Eiland. (= Vehilius vetustus Mielke, 1968, nomen novum pro Cobalus vetulus Mabille, 1883). Mnasilus allubita Butler, 1877. — Williams & Bell (1931, “Mnasilus peniallatus God- man”). ITZ: 1 6, Paramaribo. RMNH:9 6, 4 ©, Charlesburg, Lelydorp, Ma Retraite, Paramaribo, Zanderij, Zorg en Hoop. ZSM: 1 4, Surinam. See Note 23. Mnasitheus _ simplicissima Herrich-Schäffer, 1870. — Williams & Bell (1931). Mnasitheus similis spec. nov. — See Note 24. Moeris remus Fabricius, 1798. — Evans (1955); Williams & Bell (1931). Parphorus storax Mabille, 1891. — Williams & Bell (1931). RMHN: 2 6, Brokopondo, La- wa. Parphorus decora Herrich-Schaffer, 1869. — Williams & Bell (1931), “Phlebodes fartuga Schaus”). RMNH: 2 6, Makambi-kreek, Ma Retraite. ZSM: 1 5, Surinam. Papias phainis Godman, 1900. — RMNH: 1 ?, Brownsberg. See Note 25. Papias phaeomelas Geyer, 1831. — Möschler (1882); Williams & Bell (1931). RMHN: 2 3,1 2, Brownsberg, Domburg. Papias subcostulata integra Mabille, 1891. — Evans (1955) NZ CIO RR Epubliek RMNH: 3 d, Brokopondo, Brownsberg, Domburg. Papias proximus Bell, 1934. — RMNH: 1 6 Sı- paliwini. Cobalopsis venias Bell, 1942. — RMNH: 1 d, Copi. 245 Cobalopsis dorpa spec. nov. — See Note 26. Cobalopsis tanna spec. nov. — See Note 27. Cobalopsis catocala Herrich-Schaffer, 1869. — Williams & Bell (1931). Lerema ancillaris Butler, 1877. — Plötz (1886, dentini RNN NC RC" Charlesburg, Domburg, Kwatta, Ma Re- traite, Paramaribo, Zanderij, Zorg en Hoop. Morys compta compta Butler, 1877. — Evans (1955); Plötz (1886, “Hesperia aethra”); Williams & Bell (1931, “Euroto compta” and “Euroto micythus Cramer”). Morys geisa geisa Möschler, 1978. — Evans (1955). RMNH: 4 G, 1 9, Lelydorp, Ma Re- traite, Paramaribo. Morys subgrisea paradoxa subspec. nov. — See Note 28. Vettius lafresnayei pica Herrich-Schäffer, 1869. — RMNH:2 4,1 2, Brownsberg, Sipaliwi- ni. Vettius richardi Weeks, 1906. — RMNH: 1 ó, 1 2, Berlijn, Brokopondo. Vettius triangularis Geyer, 1831. — Williams & Bell (1931). ZSM: 1 ©, Surinam. Vettius monacha Plötz, 1882. — Evans (1955); Möschler (1882). ZSM: 1 5,1 2, Surinam. Vettius phyllus phyllus Cramer, 1777. — Cram- er (1777); Evans (1955); Williams & Bell (1931, “Vettius laurea Hewitson”). RMNH: 16,1 2, Blakawatra. ZSM: 1 &, Surinam. Vettius marcus Fabricius, 1787. — Evans (1955); Williams & Bell (1931). Vettius fantasos fantasos Stoll, 1780. — Plotz (1882b, “Hesperia eucherus”); Sepp (1847); Stoll (17280), Nar ssc cll Osi RMNH:2 6,2 ©, Blakawatra, Domburg. Vettius artona Hewitson, 1868. — Evans (1955); Williams & Bell (1931). RMNH: 1 2, Zanderij-Phedra. Vettius yalta Evans, 1955. — RMNH: 2 ©, Co- eroeni Eiland, Paramaribo. See Note 29. Vettius tertianus Herrich-Schäffer, 1869. — Môschler (1882 “Pamphila warra” and “Pamphila zola”). Paracarystus hypargyra Herrich-Schäffer, 1869. — Evans (1955); Môschler (1876; 1882, re- cord of 1876 was false). RMNH: 3 d, Dom- burg, Paramaribo, Surinam. Paracarystus menestriesi rona Hewitson, 1866. — Evans (1955). RMNH: 2 6, 19, Browns- berg, Phedra. ZSM: 2 g, Surinam. Turesis lucas Frabricius, 1793. — Möschler (1876). RMNH: 3 6,1 ©, Domburg. Thoon dubia Bell, 1932. — RMNH: 2 6, 1 9, 246 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Brownsberg, Patamakka Rivier. Thoon taxes Godman, 1900. — RMNH: 1 a, Ma Retraite. Justinia phaetusa phaetusa Hewitson, 1866. — RMNH: 4 6, 1 ©, Lely Gebergte, Phedra, Rama. Justinia gava Evans, 1955. — Evans (1955). Justinia justinianus dappa Evans, 1955. — Wil- liams & Bell (1931, “Eutychide cingulicornis Herrich-Schäffer”). Eutychide complana Herrich-Schäffer, 1869. — Evans (1955); Williams & Bell (1931). RMNH: 2 2, Albina, Surinam. Eutychide subcordata subcordata Herrich- Schaffer, 1869. — Evans (1955). RMNH: 1 3,1 ®, Republiek, Sipaliwini. Eutychide subpunctata intermedia subspec. nov. — See Note 30. Onophas columbaria columbaria Schaffer, 1870. — Evans (1955). Naevolus orius orius Mabille, 1883. — RMNH: 2 8,2 2, Goliath Kreek, Paramaribo, Zan- derij. Enosis angularis Môschler, 1876. — Môschler (1876; 1882, “Proteides parvipuncta” and “Carystus infuscatus Plötz”). RMNH: 6 d, 3 2, Albina, Leonsberg, Paramaribo, Suri- nam. Vertica verticalis grandipuncta Mabille, 1883. — RMNH: 2 ©, Blakawatra, Ma Retraite. Ebusus ebusus Stoll, 1780. — Evans (1955); Môschler (1876, “Carystus psecas Cr.”); Stoll (1780; 1781, “Papilio psecas”). RMNH: INNER ma Suriname Zandeny#ZSMES d,2®,Surinam. Talides sergestus Cramer, 1775. — Cramer (1775); Môschler (1876); Williams & Bell (1931). RMNH: 2 6, 1 ®, Paramaribo, Zan- derij. Talides sinois sinois Hübner, 1819. — Evans (1955); Hübner (1819); Stoll (1781, “Papilio sinon”). RMNH: 2 6, 1 ®, Domburg, Para- maribo, Surinam. See Note 31. Talides alternata alternata Bell, 1941. — Evans (1955). RMNH: 2 gd, Paramaribo. See Note 31. Nyctus hiarbas Cramer, (1775). Carystus elvira Plötz, 1882. — RMNH: 1 9, Surinam. See Note 32. Carystus hocus Evans (1955). — Evans (1955). NZC: 1 d, Gransoela. Carystus jolus Stoll, 1782. — Evans (1955); Stoll (1782). RMNH: 1 ®, Saramacca Rivier. Herrich- 1775. — Cramer Carystus senex Plôtz, 1882. — ITZ: 1 6,1 ©, Paramaribo. RMNH: 1 ©, Paramaribo. Carystus junior Evans, 1955. — RMNH: 2 d, 3 ©, Charlesburg, Lelydorp, Mariënburg, Powakka, Surinam. Carystus phorcus phorcus Cramer, 1777. — Cramer (1777); Evans (1955); Möschler (1882, “Carystus claudianus Ltrll.” and “Proteides marpesia Hew.”). ITZ: 2 ©, Par- amaribo. RMNH: 3 ©, Paramaribo. ZSM: 1 2, Surinam. Telles arcalaus Stoll, 1782. — Sepp (1850, “Pa- pilio uraniae”); Stoll (1782). RMNH: 1 9, Surinam. Moeros moeros Moschler, 1876. — Evans (1955); Möschler (1876). RMNH: 1 6, 2 8, Paramaribo; agreeing with description and figures, but midtibiae spined (cf. Evans, 1955: 239). Cobalus virbius virbius Cramer, 1777. — Cramer (1777); Evans (1955); Möschler (1876); Williams & Bell (1931). ITZ: 1 9, Paramaribo. ZSM:5 6, Surinam. Cobalus calvina Hewitson, 1866. — RMNH: 1 3, Zanderij. ZSM: 9 d, Surinam. Dubiella fiscella Hewitson, 1877. — Williams & Bell (1931). Dubiella dubius Stoll, 1781. — Stoll (1781). RMNH:3 6,1 9, Moengotapoe-Wia-Wia. Tellona variegata Hewitson, 1870. — Bell (1931, “Thracides currani”). RMNH: 1 6,2 2, Brownsberg. Damas clavus Herrich-Schaffer, 1869. — Evans (1955); Möschler (1876, “Proteides cervus”). RMNH: 3 d, Patamakka Rivier, Zanderij. ZSM:4 6,1 9, Surinam. Orphe gerasa Hewitson, 1871. — RMNH: 1 6, Surinam. Carystoides basoches basoches Latreille, 1824. — Evans (1955); Môschler (1876, “Proteides brinoides” and “Caristus basochesit Ltrll.”); Williams & Bell (1931). ITZ: 1 ®, Surinam. RMNH: 4 dg, 2 ©, Paramaribo, Surinam, Zanderij. ZSM: 2 8, 1 2, Surinam (2 possi- bly type of Proteides brinoides Möschler). See Note 32. Carystoides noseda Hewitson, 1866. — Evans (1955). NZC:1 d, Lelydorp. See Note 33. Carystoides sicania orbius Godman, 1901. — Evans (1955). RMNH: 3 6, Coeroeni Ei- land, Linker Coppename Rivier, Patamakka. ZSM: 1 3, Surinam. Carystoides maroma Môschler, 1876. — Evans (1955); Môschler (1876); Plötz (1882b, DE JoNG: Hesperudae of Surinam 247 “Hesperia valentina”). Carystoides cathaea Hewitson, 1866. — Evans (1955). Perichares butus Môschler, 1876. — Môschler (1876). RMNH: 2 d, Domburg, Surinam. Perichares philetes philetes Gmelin, 1790. — Evans (1955); Môschler (1876), “Caristus corydon Fb.”); Williams & Bell (1931, “Peri- hares [sic!] coridon Fabricius”). RMNH: 4 3, 6 2, Domburg, Lelydorp, Paramaribo, Surinam. ZSM: 1 d, Surinam. Perichares lotus Butler, 1870. — Williams & Bell (1931). RMNH: 1 6, Domburg. Perichares deceptus Butler & Druce, 1872. — RMNH: 1 6, Rama. See Note 34. Orses cynisca Swainson, 1821. — RMNH: 1 6, Gansee. Lycas godarti boisduvalii Ehrmann, 1909. — Möschler (1876, “Proteides ceraca Hew.”). RMNH: 1 4, Domburg. Saturnus tiberius Möschler, 1882. Möschler (1882); Williams & Bell (1931, “Phlebodes tiberius Möschler” and “Phlebodes reticulata Plotz”). Phlebodes pertinax Stoll, 1781. — Evans (1955); Stoll (1781). RMNH: 1 d, Zanderij-Phedra. Phlebodes meesi spec. nov. — See Note 35. Joanna box: Evans, 1955. — RMNH: 1 8, Suri- nam. Quinta cannae Herrich-Schaffer, 1869. — Evans (1955); Môschler (1882, “Proteides osembo”). ITZ: 1 d, Paramaribo. RMNH: 3 d,4 2, Ma Retraite, Paramaribo. Cynea anthracinus luctatius Schaus, 1913. — RMNH:1 6 ,1 2, Brownsberg, Domburg. Cynea cyrus Plôtz, 1883. — RMNH: 2 d, Ra- ma. See Note 36. Cynea iquita Bell, 1941. — Evans (1955). Cynea corisana Môschler, 1882. — Môschler (1882). Cynea diluta Herrich-Schäffer, 1869. — Evans (1955); Môschler (1882, “Pamphila zeppa”); Plötz (1883, “Hesperia vellejus”); Williams & Bell (1931, “Cobalus zeppa Möschler”). ZR Sti Dam ERMNEIE STORE 2s Domburg, Ma Retraite, Paramaribo. Cynea corope Herrich-Schäffer, 1869. — Möschler (1882). Penicula bryanti Weeks, 1906. — Evans (1955). RMNH: 6 d, Domburg, Ma Retraite. Penicula criska extrema subspec. nov. — See Note 37. Decinea lucifer Hübner, 1831. — Hübner (1831). Conga chydaea Butler, 1870. — Williams & Bell (1931, “Prenes vala Mabille”). Hylephila phyleus phyleus Drury, 1773 (for cor- rect spelling, see Brown & Heineman, 1972: 408). — Williams & Bell (1931). ITZ: 6 6, 1 ®, Paramaribo. RMNH: 9 6, 4 2, Broko- pondo, Ma Retraite, Matapi, Paramaribo, Zanderij, Zorg en Hoop. Hesperia uncas Edwards, 1863. — Môschler (1882, 2 d from Paramaribo, identified as Hesperia uncas by Plôtz); identification or locality labels false, the species is North American. Polites vibex Geyer, 1832. — Williams & Bell (LOS) Zara abo and ery: NZC: 1 2, Paramaribo. RMNH: 8 6,2 9, Domburg, Paramaribo, Surinam. See Note 38. Polites vibicoides spec. nov. — See Note 39. Polites coras Cramer, 1775. — Cramer (1775); Williams & Bell (1931, “Catia otho Smith & Abbot”). A purely North American species. The locality of Cramer’s type must have been false. The observation of Williams & Bell must be based on a misidentification. Wallengrenia druryi curassavica Snellen, 1887. — Evans (1955); Möschler (1882). RMNH: 12 g, 15 ®, Albina, Domburg, Ma Retraite, Paramaribo, Raleigh Vallen, Rama, Surinam. Wallengrenia premnas Wallengren, 1860. — Evans (1955). Pompeius pompeius Latreille, 1824. — Evans (1955); Môschler (1882); Williams & Bell (1931, “Polites athenion Hübner”). ITZ: 15 3, 10 9, Charlesburg, Leiding, Paramaribo, Republiek, Zanderij. NZC: 1 ®, Paramari- bo. RMNH: 23 6, 12 2, Albina, Brokopon- do, Brownsberg, Charlesburg, Coeroeni Ei- land, Domburg, Lelydorp, Ma Retraite, Par- amaribo, Surinam. Choranthus vitellius Fabricius, 1793. — Wil- liams & Bell (1931). Mellana clavus Erichson, 1848. — Williams & Bell (1931, “Atrytone mella Godman” and “Atrytone barbara”). Mellana helva Möschler, 1876. — Möschler (1876). Mellana villa Evans, 1955. — RMNH: 1 d, 4 2, Brownsberg, Ma Retraite, Paramaribo, Surinam. Euphyes sirene kaye: Bell, 1931. — Evans (1955). RMNH:1 dg, Zanderij. Metron chrysogastra chrysogastra Butler, 1870. — RMNH: 1 dg, Sipaliwini. 248 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Metron fasciata Möschler, 1876. — Möschler (1876). RMNH: 3 d, Domburg, Ma Re- traite. Propertius phineus Cramer, 1777. — Cramer (1777); Evans (1955, “Propertius albistriga Tessmann”). RMNH: 1 d, Surinam, holo- type. See De Jong (1983). Calpodes ethlius Stoll, 1782. — Evans (1955); Stoll (1782). ITZ: 1 6, Paramaribo. RMNH: 5 6,6 ®, Ma Retraite, Paramaribo, Powak- ka, Zanderij. Panoquina panoquinoides minima subspec. nov. — See Note 40. Panoquina ocola Edwards, 1863. — Evans (1955); Môschler (1882, “Pamphila orty- gia”). ITZ: 11 d,8 ®, Charlesburg, Leiding, Paramaribo, Republiek. NZC: 1 4, Repub- liek. RMNH: 4 6, 2 2, Coppename Rivier, Ma Retraite, Morico Kreek, Paramaribo, Republiek. Panoquina sylvicola Herrich-Schätfer, 1865. — Evans (1955); Möschler (1876). RMNH: 4 3,5 2, Brownsberg, Carolina Kreek, Lely- dorp, Ma Retraite, Paramaribo. Panoquina fusina fusina Hewitson, 1868. — Evans (1955); Williams & Bell (1931). RMNH: 4 6, Charlesburg, Morico Kreek, Paramaribo. Panoquina evadnes Stoll, 1781. — Evans (1955); Plötz (1882, “Hesperia chlorus”). RMNH:5 6,4 ®, Paramaribo, Surinam. Nyctelius nyctelius nyctelius Latreille, 1824. — Evans (1955); Williams. & Bell (1931). RMNH: 1 d, 2 2, Lelydorp, Paramaribo, Surinam. Thespieus dalman Latreille, 1824. — Evans (1955). Vacerra bonfilius litana Hewitson, 1866. — Williams & Bell (1931). Oxynthes corusca Herrich-Schäffer, 1869. — ZSM: 1 2, Surinam. Niconiades yoka Evans, 1955. — RMNH: 1 d, 1 2, Morico Kreek, Patamakka Rivier. Niconiades nikko Hayward, 1948. — NZC: 1 6, Ma Retraite. RMNH: 5 6,1 ®, Ma Re- traite, Paramaribo. Niconiades caeso Mabille, 1891. — Williams & Bell (1931); possibly misidentification, southern species only known from Paraguay and S. Brazil. Aides epitus epitus Stoll, 1781. — Evans (1955); Stoll (1781). RMNH: 1 à, Zanderij. Aides brino Stoll, 1781. — Stoll (1781). Aides aegita Hewitson, 1866. — RMNH: 2 dg, 4 9, Paramaribo, Surinam. Aides ocrinus Plôtz, 1882. — Möschler (1882). Xeniades chalestra Hewitson, 1866. — RMNH: 1 4,2 2, Paramaribo, Surinam. Xeniades orchamus Cramer, 1777. — Cramer (1777). RMNH:2 d, 6 ©, Paramaribo, Suri- nam (including ® holotype, Surinam). Cravera rara gen. nov. et spec. nov. — See Note 41. Surina unica gen. nov. et spec. nov. — See Note 2 Saliana chiomara Hewitson, 1871. — Evans (1955). RMNH: 1 ©, Paramaribo. Saliana fischeri Latreille, 1824. — Evans (1955). RMNH:1 d, Surinam. Saliana nigel Evans, 1955. — Evans (1955). Saliana esperi Evans, 1955. — Evans (1955). ZSM: 4 d, Surinam. Saliana antoninus Latreille, 1824. — Evans (1955); Môschler (1876). ITZ: 1 ®, Parama- ribo. RMNH: 3 6, 3 ©, Brownsberg, Ma Retraite, Surinam, Zanderij. Saliana longirostris Sepp, [1840]. — Evans (1955); Sepp (1832-1840). RMNH: 3 4, 3 2, Albina, Domburg, Ma Retraite, Parama- ribo, Patamakka Rivier. ZSM: 1 d, 1 9, Su rinam. Saliana morsa Evans, 1955. — Evans (1955). RMNH: 1 d, Zanderij. Saliana salins Cramer, 1775. — Cramer (1775); Evans (1955); Möschler (1876, “Thracides telegonus Esper”); Williams & Bell (1931, “Thracides telegonus Esper”). ITZ: 1 ®, Paramaribo. NZC: 1 à, Republiek. ZSM: 4 d, Surinam. Thracides phidon Cramer, 1779. — Cramer (1779); Evans (1955); Möschler (1876); Sepp (1841, “Papilio pyrophoros”). RMNH: 2 6, Charlesburg, Paramaribo. Neoxeniades cincia Hewitson, 1871. — RMNH: 1 6,1 ®, Goliath Kreek. See Note 43. Neoxeniades braesia Hewitson, 1867. — RMNH: 1 ©, Goliath Kreek. See Note 44. Pyrrhopygopsis socrates orasus Druce, 1876. — RMNH:1 6,1 ®, Batavia, Surinam. NOTES ON THE HESPERIIDAE OF SURINAM 1. Pyrrhopyge phidias Linnaeus, 1758. | Evans (1951) recognized 10 subspecies. They | are, however, not geographically defined, sever- | al subspecies being sympatric. For Surinam, for | instance, the following “subspecies” are record- | ed: bixae Linnaeus, garata Hewitson, and | phidias Linnaeus. Among the specimens from | Surinam in RMNH there are, moreover, two | DE Jong: Hesperudae of Surinam 249 10 Figs. 2—4. Elbella species, inside of left valva; 2, E. bicuspis spec. nov., Paramaribo (Surinam); 3, E. umbrata acala, Muzo (Colombia); 4, E. umbrata umbrata, Rio Pancartambo (Peru). Figs. 5—8, Urbanus species, inside of left valva; 5, U. ambiguus spec. nov., Surinam; 6, U. teleius, Domburg (Surinam); 7, U. simplicius, Avanavero Vallen (Surinam); 8, U. zagorus, Surinam. Figs. 9—11, Bungalotis sipa spec. nov. Sipaliwini (Surinam); 9, inside of left valva; 10, dorsal view of tegumen and uncus; 11, lateral view of tegumen, uncus and gnathos. Figs. 12— 13, Clito jonkersi spec. nov, Domburg (Surinam); 12, inside of left valva; 13, lateral view of tegumen, uncus and gnathos. Figs. 2—11 drawn at same magnification, 12—13 at twice this magnification. 250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, males agreeing with “subspecies” zenodorus Godman & Salvin. Apparently the subspecific classification of this species does not correctly reflect the pattern of variation. In this case it is better to abandon the subspecies at all and to describe the variation in a more appropriate way. The possibility of more than one species being involved cannot be ruled out. 2. Pyrrhopyge charybdis Westwood, 1852. The single male from Surinam in RMNH has a forewing length of 26.4 mm, a dark indigo blue shine and veins not conspicuously black. It thus agrees with ssp. semita Evans, 1951, the southernmost of the two subspecies recognized by Evans for this essentially central and south- ern Brazilian species. A male in RMNH with identical genitalia and same colouring from Por- to Alegro, has a forewing length of 24 mm, and comes close to ssp. charybdis Westwood, as far as size is concerned. As the distribution areas of the two subspecies defined by Evans (1951) partly overlap, the recognition of two subspe- cies is doubtful. 3. Pyrrhopyge cometes Cramer, 1779. The single male from Surinam in RMNH has size and spots as in Peruvian specimens (ssp. staudingeri Plötz, 1879) and is unlike a male from French Guiana in RMNH (ssp. cometes) (cf. Evans, 1951: 35). Either the locality is false, or the variation is more complicated than sug- gested by the subspecific classification by Evans (1951), as is often the case in Pyrrhopyge spe- cies. 4. Elbella bicuspis spec. nov. External characters (figs. 49, 50). — Belong- ing to patrobas group of species (Evans, 1951: 42): Jemadia-like, forewing spot in space 3 free, spot in space 4 in line with those in 5—7. Fore- wing, no spot in space 9, a tiny one in space 10 far basad of the spot in space 8. Hyaline spot in space 1b narrow, pointed, not entirely reaching middle of space 1b. Blue subbasal bar on upper side at almost right angles to dorsum, not paral- lel with central hyaline band. Hindwing upper side, blue bands not extending beyond vein 7. Hindwing underside, subbasal blue band from space 8 across cell to vein 1b, directed to mid- way dorsum; central blue band as wide as dark area between it and subbasal band, directed to dorsum above tornus; inner discal band from vein 8 tapering to vein 3, narrow in spaces 3 and 4/5; outer discal band from vein 6 to vein 1b; 1983 = blue submarginal scaling inconspicuous, close to outer discal band; a narrow blue bar in space 7 between central and inner discal bands. Length of forewing, 24.2 mm. Male genitalia (figs. 2—4). — Very much like Elbella umbrata Mabille & Boullet, 1908, but cucullus slenderer, straighter, and with two dor- sal prongs instead of one (hence the name of the species). Identification. — As remarked by Evans (1951: 42) the species of the patrobas group can only be determined with certainty by genitalia examination. The present species, however, dif- fers from all other species of the group in the blue subbasal bands on the upper side of the forewing and underside of the hindwing being clearly less oblique. In all other species of the group the subbasal and central bands on the un- derside of the hindwing generally point to the tornus, while in the new species they point to the dorsum above the tornus. Material examined. — Holotype, 6, Parama- ribo, leg. E. H. Jonkers (RMNH). 5. Jemadia fallax fallax Mabille, 1878. The four males from Surinam in RMNH show some variation in the serration of the cu- cullus, and are strikingly different in the shape of the cucullus, which varies from semicircular to ovate. There is no reason to consider it other- wise than individual variation. 6. Phocides pigmalion hewitsonius Mabille, 1883. Hyaline spots of forewing hardly, if at all dif- ferent from those in males from Honduras and Colombia (ssp. pigmalion Cramer, 1779), but on upper side of hindwing blue postdiscal band reduced, especially in spaces 4/5 and 6. Therefore, the original figure of “Papilio pigma- lion” by Cramer (1779: fig. 245a) differs from specimens from Surinam, and as correctly sug- gested by Evans (1952: 13), Cramer’s figure must represent a specimen that did not come from Surinam. 7. Entheus gentius Cramer, 1779. In the females, there is much variation in the position of the hyaline spots of the forewing. The upper spot in space 1b and the spot in space 2 are always conjoined, but the position of the spot in space 2 varies relative to the cell spot from being completely in line to widely sepa- rate, so that the spot in space 2 is midway be- tween the spot in space 3 and the cell spot. In DE JONG: Hesperudae of Surinam 251 A | 16 | 20 SN > Figs. 14-17. Cymaenes geijskesi spec. nov., Zanderij (Surinam); 14, inside of left valva; 15, dorsal view of left valva; 16, dorsal view of tegumen and uncus; 17, lateral view of tegumen, uncus and gnathos. Fig. 18, Vehilius major spec. nov., Patamakka Rivier (Surinam), inside of left valva. Fig. 19, Vehilius vetula, Brownsberg (Suri- nam), inside of left valva. Figs. 20—22, Mnasitheus similis spec. nov., Surinam; 20, 6 forewing venation and brands (hatched); 21, dorsal view of tegumen and uncus; 22, inside of left valva. Fig. 23, Cobalopsis dorpa spec. nov., Paramaribo (Surinam), inside of left valva. Figs. 24—25, Cobalopsis tanna spec. nov., Domburg (Surinam); 24, inside of left valva; 25, dorsal view of tegumen and uncus. All parts of genitalia drawn at same magnification. addition there is much variation in the number and length of the processes on either side of the ostium bursae. The variation in the markings as well as in the genitalia seems to be continuous. 8. Aguna claxon Evans, 1952. The genitalia and external characters agree well with this species, but the spot in space 3 of the forewing is at least as far from the spot in space 2 as its own width and not overlapping as stated by Evans (1952: 60) in the original de- scription. 9. Polythrix asine Hewitson, 1867. The single female from Surinam in RMNH agrees well with this species, but on the upper side of the forewing the inner edge of the spot in space 2 is not in line with the inner edge of the cell spot, being at a right angle with vein 2. As the female genitalia of Polythrix species are practically unknown, the identification must re- main provisionally for the time being. 10. Polythrix caunus Herrich-Schäffer, 1869. The male genitalia agree completely with 252 TIJDscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, those of P. caunus, but contrary to what is stated by Evans (1952: 71), the spot in space 9 of the forewing is absent in two males and only a tiny dot in the other males. Moreover, on the forewing there is a well-developed spot in space 1b under the outer part of the spot in space 2 in both females, and a tiny dot in the same place in two males. Consequently, without the help of the genitalia it is impossible to idenufy these specimens with Evans’ key. 11. Urbanus esma Evans, 1952. The dorsal process of the cucullus is distinctly shorter and stouter than in U. esmeraldus But- ler, as stated by Evans (1952: 90), but the out- line of the valve seems to be similar in both spe- cies, contrary to Evans’ very schematic draw- ings. 12. Urbanus ambiguus spec. nov. External characters (figs. 51, 52). — Male, length of forewing 21 mm. No costal fold. Up- perside uniformly brown, including head, body and wing bases. Fringes brown, unchequered. Forewing with a narrow central hyaline band from middle of space 1b to costa, composed of upper spot in space 1b, spots in spaces 2, 3, and cell, and two small spots over cell spot. Incon- spicuous linear spots in spaces 4 and 5, the latter more or less in line with the subequal spots in spaces 6 to 9. On underside of hindwing central band connected to outer spot in space 7, widen- ing towards vein 1b; discal band from vein 7 to vein 1b, with irregular sides, widening towards vein 1b and here almost touching the central band. Male genitalia (figs. 5—8). — Costa bulging so that total height of valve is twice height of cucullus (without spine). Cucullus with a strong apical spine and a few inconspicuous short spin- es. Idenufication. — The species falls within what can be called the teleus group of species, characterized by the uniformly brown upper side with unchequered brownish fringes and spot in space 3 forming part of the central band. By the absence of a costal fold and the con- joined cell spot and outer spot in space 7 on the underside of the hindwing, the species keys to U. cindra Evans with Evans (1952). It differs, however, from this species in the presence of spots in spaces 4 and 5 and a well-developed spot in space 9 of the forewing, and in the bulg- ing costa of the valve. In the latter character it is similar to U. teleus Hübner, which, however, is 1983 more strongly spined in the apical part of the cucullus and has, moreover, the cell spot on the underside of the hindwing central between the inner and outer spots in space 7. In figs. 6—8 the valves of some related species are given for comparison. Material examined. — Holotype, d, Surinam (RMNH). 13. Astraptes cretatus Hayward, 1939. In the male the white tornal area on the un- | derside of the forewing extends up to the cell, in the female it enters the cell almost up to the ra- dius. Costa of forewing on underside hardly with any white. By these characters the speci- | mens are intermediate between spp. cretatus and ssp. adoba Evans, 1952. 14. Bungalotis sipa spec. nov. External characters (figs. 53, 54). — Male, length of forewing 22.6 mm. Ground colour up- | per side orange-brown, dark brown in spaces 7 and 8 of hindwing. Forewing with narrow but conspicuous dark markings in spaces 1b, 2, 3, and 4 to 8, the latter in a curved line, and a dark bar across the cell just before the origin of vein 3; spot in space 2 midway between the origin of vein 3 and the spot in space 3, far from the cell | spot. Hindwing with an irregular series of dark spots in spaces 1b to 6 and a dark spot at end cell. Underside similar, with broad dark suffu- } sion along termen of forewing and along costa and termen of hindwing, and light ochreous È along dorsum of forewing up to middle of space # 1b. Spots in spaces 1b and 2 on underside of } forewing with whitish scales in centre, similarly | on underside of hindwing discal spots in spaces | le to 7 (in space 1c a double spot), basal spot in | space Ic, and spot at end cell; subbasal spot in space 7 without whitish scales. Male genitalia (figs. 9—11). — Uncus rela- tively long and narrow, 1.3 times as long as greatest width, more than 4 times as long as at narrowest point. Cucullus bifurcate apically, lower arm slightly serrate and pointed, upper arm much larger, irregularly serrate, dorsal edge | of cucullus finely serrate. | Identification. — By the position of the spots | on the ae side of the hindwing the specimen | keys to B. quadratum Sepp (cf. Evans, 1952: | 140), but dhe lener is a larger insect (male fore- wing length 26 mm), with a lighter, yellowish | tinge and less conspicuous, broader spots. The | narrow uncus and general shape of the valve are | suggestive of B. borax lactos Evans, which is | DE Jon: Hesperudae of Surinam 253 Fig. 26. Eutychide subpunctata intermedia subspec. nov, Paramaribo (Surinam), inside of left valva. Figs. 27— 29, Phlebodes meesi spec. nov, Lely Gebergte (Surinam); 27, dorsal view of tegumen and uncus; 28, inside of left valva; 29, lateral view of tegumen, uncus and gnathos. Figs. 30—31, Penicula criska extensa subspec. nov, Linker Coppename (Surinam); 30, dorsal view of tegumen and uncus; 31, inside of left valva. All figures drawn at same magnification. only known from Peru, but in that species the cucullus is not as strongly bifurcate and the ex- ternal characters are different, the colour being darker and the cell spot of the forewing being Pa to the inner edge of the spot in space 2 (I do not understand why Evans, 1952, de- scribed lactos as a subspecies of B. borax Evans, as the genitalia are as different from those of B. borax as from those of any other Bungalotis species; it seems better to consider lactos a sepa- rate species). Material examined. — Holotype, d, S. Suri- nam, Sipaliwini, air strip, 14.x.1968, at light, leg. E. H. Jonkers (RMNH). 15. Bungalotis quadratum Sepp, 1848. According to Evans (1952: 140) two subspe- cies can be recognized, viz., ssp. quadratum (Honduras, Colombia, Guiana, Surinam), and spp. barba Evans (Fr. Guiana, Pará, Santarem, Peru). The differences in the male sex are as fol- lows. In ssp. quadratum the ground colour is yellow and the costa of the hindwing (“upf” in Evans must be a misprint for “uph”) broadly dark brown on the upper side. Ssp. barba is dark tawny, with dark brown area on upper side of hindwing usually limited to a small area un- der outer half of vein 7. One male from Parama- ribo (RMNH) caught 5.1.1962, has the yellow colour of ssp. quadratum and the extension ot the dark area on the upper side of the hindwing along the costa as in ssp. barba. The other male from Paramaribo (NZC) caught 23.x.1981, has the colour of ssp. barba and the extension ot the 254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Figs. 32—39. Polites vibicoides spec. nov., Zanderij (Surinam) (even numbers), and Polites vibex catilina, Dom- burg (Surinam) (odd numbers); 32—33, dorsal view of tegumen and uncus; 34—35, lateral view of tegumen, uncus and gnathos; 36—37, inside of left valva; 38—39, apex of antennal club, black = with black scales, stippled = with orange-yellow scales, rest is nudum. All figures drawn at same magnification. dark area of ssp. quadratum. This makes one wonder how far the subspecific distinction is justified. 16. Cogıa hassan Butler, 1870. The specimens from Zanderij are all similar. They agree with ssp. hassan as defined by Evans (1953: 20), and also with the original descrip- tion and figure of Cogia freudiae Williams & Bell, 1931, which Evans, in my opinion, justly synonymized with Cogia hassan Butler. The shape of the valve, with the apex of the cucullus ending in a long spike, is exactly like the draw- ing by Williams & Bell (1931, fig. 3) and unlike the very schematic drawing by Evans (1953). The single male from Temomairem, however, is different. The termen of the forewing is more convex and the lines on the underside of the hindwing are as described for ssp. evansı Bell, 1937, which according to Evans (1953) was fig- DE JONG: Hesperudae of Surinam 255 ured by Draudt (1922, pl. 170d) under the name of “hassan” (the colours of Draudt’s figure must have been invented by the printer, but the posi- tion of the lines on the underside of the hind- wing is clear). The genitalia are similar to those of the specimens from Zanderij. As Zanderij is in the north of the country and Temomairem in the extreme south, a subspecific difference does not seem to be too surprising. Ssp. hassan, however, is the subspecies of north- ern Brazil and ssp. evansi occurs in Bolivia and southern Brazil. The capture of a specimen with characters of ssp. evansi far within the bounda- ries of the distribution area of ssp. hassan sug- gests that the two taxa may be specifically dis- tinct. On the other hand, they could also be merely seasonal forms. The material is still too scanty to make a firm statement. 17. Clito jonkersi spec. nov. External characters (fig. 55). — Male. Length of forewing 15.1 mm. Costal fold present. Nudum 22 segments. Forewing upper side dark brown with hyaline spots in spaces 1b (acco- lade-shaped), 2 (large, V-shaped), 3, 4, 5, and 6— 8 (in line, the latter longer, extending further to termen), and in cell (U-shaped with rounded base). Hindwing upper side dark brown with hyaline spots in spaces 1c (anvil-shaped, just ex- tending into space 1b), 2 (very small, at base), 6 (small, at base), 7 and in cell (filling more than half of cell); inner edge of spots in 1c, 7 and cell in line; faint whitish pointed submarginal spots in spaces 1c—-6. Underside as upper side; lighter shading on forewing in spaces 1b—3 directly distad of the hyaline spots; also faint lighter shading along termen of hindwing in spaces 4— Male genitalia (figs. 12, 13). — Gnathos rela- tively long, almost reaching apex of uncus. Cu- cullus stout, slightly higher than long, the broad apex strongly spined. Costal process short, stout, finely spined. Identification. — Externally the new species comes closest to C. littera anda Evans, 1953. It differs in the white band of the hindwing being narrower and divided by dark veins, and the cell spot of the forewing not being rounded basally, but cut off obliquely. In C. littera littera Ma- bille, 1877, the white band of the hindwing is faint and the inner edge of the cell spot of the forewing forms a straight line perpendicular to the costa. The shape of the valve and length of the gnathos are different from those of all de- scribed Clito species. Material examined. — Holotype, d, Dom- burg, 8.11.1975, leg. E. H. Jonkers (RMNH). Discussion. — According to Evans (1953) the genus Chto is characterized among other char- acters by a nudum of 16 segments. I don’t know how Evans arrived at this figure. In the BM I checked the material arranged by Evans, and found the following numbers of nudum seg- ments: C. littera Mabille, 3, 22; C. bibulus Riley, & 18 E22 EC Hehto) Fabricius iO sompa Evans, d 21; C. zelotes Hewitson, & 21; C. zenda Evans, d and 9 22; C. tuva Evans, d and 9 22. Apparently all species are rarely caught, judged from the very scanty material available. 18. Anthoptus epictetus Fabricius, 1793. According to Evans (1955: 62) the female should be entirely brown. However, in a series of 448 specimens he had only two females. In the series of 15 specimens from Surinam in RMNH there is a single female that is similar to the males except for a yellow stripe on the up- per side of the hindwing from base to termen covering vein 1b. 19. Vinius tryhana Kaye, 1913. The three males in RMNH have identical genitalia corresponding with those of V. tryha- na. One male has the tawny colour on the upper side more extensive, so that the cell of the hindwing is tawny, and the cell of the forewing is tawny with a narrow black central streak. By the latter character the specimen is similar to V. exilis Plötz, 1883, but the genitalia point defi- nitely to V. trybana. 20. Apaustus menes Stoll, 1782. According to Evans’ (1955: 83) key, Apaustus has six nudum segments. In the description of the two species at p. 91, however, A. gracilis Felder is stated to have 4—6 and A. menes 8 nudum segments. The three specimens in RMNH which undoubtedly belong to A. men- es, have 6 nudum segments. 21. Cymaenes geijskesi spec. nov. External characters (fig. 56). — Male. Length of forewing 12.4 mm. Upper side brown; fore- wing with faint ochreous spots in spaces 2, 3 and 6, and traces of spots in spaces 1b and 7, and in cell; hindwing practically unicoloured. Underside as upper side, a little paler, traces of 256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AEL. median spots in spaces 2—5 on the hindwing. Fringes sullied. Nudum with 13 segments of which 9 on apiculus. Male genitalia (figs. 14— 17). — Indentation of uncus not more than one third of total length of uncus + tegumen, twice as long as wide. Cu- cullus greatly expanded to a semicircle with a slightly serrate edge, turned over in horizontal position. Identification. — Rather similar to C. tripunctus Herrich-Schaffer by the absence of markings in spaces 4 and 5 on the underside of the forewing, by the very faint cell spot on the forewing, the faint spots on the underside, and the broadened cucullus. It differs in being smaller (C. tripunctus, 13—15.3 mm), underside more yellowish than greyish brown, and abun- dantly in the genitalia, where the semicircular cucullus is unique for the genus and the indenta- ton of the uncus is much narrower and shal- lower than in C. tripunctus, in which moreover the uncus bulges strongly laterally. In C. tripunctus, and according to Evans (1955) in all other Cymaenes species as well, the nudum consists of 11 segments, of which 8 are on the apiculus. In C. gezjskesi there are two more seg- ments. As the number of nudum segments may vary even within a species, examination of much more material is needed to make sure that this is a constant difference between the new species and its congeners. Material examined. — Holotype, 4, Surinam, Paramaribo-Zanderij, savanne, 27—30.vili. 1964, leg. D. C. Geijskes (RMNH). 22. Vehilius major spec. nov. External characters (figs. 57, 58). — Male. Length of forewing 13.4 mm. Exactly like V. vetulus Mabille, only larger (V. vetulus 10— 12.2 mm) and upper cell spot on upper side of forewing more ns Male genitalia (figs. 18, 19). — Similar to those of V. vetulus, but outline of valve trape- zoid rather than triangular as in V. wetulus. Material examined. — Holotype, d, Surinam, Patamakka Rivier, 4.1x.1969, leg. E. H. Jonkers (RMNH). 23. Mnasilus allubita Butler and Nastra guianae Lindsey, 1925. M. allubita is very difficult to separate from N. guianae. The only external difference I could find is the colour of the palps and cheeks, white in N. guianae, cream-coloured or pale yellow in 1877, 11, 1983 M. allubita. According to Evans (1955) the only difference between Nastra and Mnasilus is the occurrence of a recumbent hair tuft on the up- per side of the forewing in males of Mnasilus. This is correct, but little obvious. It seems exag- gerated to base a generic distinction on this character only. Together with related genera these two are badly in need of revision. 24. Mnasitheus similis spec. nov. External characters (figs. 20, 59, 60). — Male. Length of forewing 12.5 mm. Upper side en- urely dark brown, no spots; slight greenish me- tallic sheen on head, colar and thorax. Under- side dark brown, faintly lighter in tornal half of forewing, in spaces la—c of hindwing, and with very faint discal spots in spaces 2—6 of hind- wing. Forewing with tripartite stigma: a short, obliquely set patch at the base of space 2 and di- rectly under it, a small patch in upper half of space 1b and a patch in lower half of space 1b similar to that in space 2. Male genitalia (figs. 21, 22). — Uncus deeply indented. Valve with cucullus strongly devel- oped in dorso-proximal part, extending far be- yond the ill-developed costa; apex of cucullus upturned and pointed. Identification. — With Evans (1955) the new species keys more or less to M. simplicissima Herrich-Schäffer, which however has an entire uncus and a differently shaped cucullus, without the strong dorso-proximal expansion. The shape of the cucullus is reminiscent of that of M. forma Evans, which does not have the deeply indented uncus of the present species. The shape of the tripartite stigma differs from that of any other Mnasitheus species, the patches in space 1b usually being parallel to vein 1, or (in M. continua Evans) perpendicular to vein 1, and the patch in space 2 either parallel to vein 2 or to the cubitus. Material examined. — Holotype, d, Surinam, leg. E. H. Jonkers (RMNH). 25. Papias phainis Godman, 1900. The single female was identified as belonging to this species rather than to P. phaeomelas Geyer because of the palps being yellower and the hindwing having well visible spots on the underside. Generally, identification of females of this genus as well as of many related genera is hampered by the fact that the female genitalia have not been studied comparatively (and usual- ly are not known at all). DE JONG: Hesperudae of Surinam 257 Figs. 40—44. Cravera rara gen. et spec. nov, Zanderij (Surinam); 40, wing venation of male, spots and brands (hatched) of upperside indicated; 41, inside of left valva; 42, dorsal view of tegumen and uncus; 43, lateral view of tegumen, uncus (u) and gnathos (g); 44, aedeagus. Figs. 41—43 drawn at twice the magnification of fig. 44. 26. Cobalopsis dorpa spec. nov. External characters (figs. 61, 62). — Male. Length of forewing 16.2 mm. Upper side fore- wing dark brown; basal third with inconspicu- ous dark yellow hairs and scaling; semi-hyaline spots in spaces 2, 3 and 6—8, the latter in line (the spot in space 8 is inconspicuous); an opaque spot over vein 1 just beyond middle; fringes pale brown, dark brown at the end ot veins. Upper side hindwing dark brown; incon- spicuous dark yellow hairs and scaling in cell and spaces 1c—5; fringes as on forewing. Un- derside forewing as upper side, narrowly violet along apical half of termen and in spaces 4—5 258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Figs. 45— 48. Surina unica gen. et spec. nov., Surinam; 45, wing venation of male, spots and brands (hatched) of upperside indicated; 46, lateral view of tegumen, uncus and gnathos; 47, dorsal view of tegumen and uncus; 48, inside of left valva. Parts of genitalia drawn at same magnification. midway between the apical spots and termen; area between apical spots and violet scaling along termen darker than rest of wing; fine dark brown terminal line. Underside of hindwing dark brown, veins pale brown; violet scaling along termen, separated from fringes by fine dark brown terminal line; pale median spots with a violet-greyish tinge and whitish outlined in spaces 1b—7, and similar spots in cell and at base of space 7 (vaguely continued into cell and space 8); spot in space 6 detached from both the spot in space 5 and the cell spot; median spot in space 7 large, completely overlapping the spot in space 6; cell spot midway between the basal and median spots in space 7 and detached from either. Female. As male, but spots slightly larger, and on upper side of forewing with faint yel- lowish spots in spaces 4—5, almost midway the apical spots and termen. Male genitalia (fig. 23). — Uncus and gnathos bipartite as in other species of the genus. Aedea- gus dorso-apically with two longitudinal rows of strong spines. Valve peculiarly shaped, more or less like a shoe in lateral view; cucullus occu- pies more than half of entire valve, dorsal and ventral edge parallel, slightly serrate along ven- tral edge, apex obtuse. Costa weakly developed; no costal process. Identification. — The new species agrees with the description of C. prado Evans, but compari- son with the type and further material of the lat- ter species in the BM revealed that in C. prado the underside of the hindwing is dark brown DE JONG: Hesperudae of Surinam 259 55 56 Figs. 49, 50. Elbella bicuspis spec. nov, holotype, upper- and underside. Figs. 51, 52. Urbanus ambiguus spec. nov, holotype, upper- and underside. Figs. 53, 54. Bungalotis sipa spec. nov., holotype, upper- and underside. Fig. 55. Clito jonkersi spec. nov, holotype, upperside. Fig. 56. Cymaenes geijskesi spec. nov, holotype, upper- side. 260 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 61 62 63 Figs. 57, 58. Vehilius major spec. nov., holotype, upper- and underside. Figs. 59, 60. Mnasitheus similis, spec. nov., holotype, upper- and underside. Figs. 61, 62. Cobalopsis dorpa spec. nov, holotype, upper- and underside. Fig. 63. Cobalopsis tanna spec. nov, holotype, upperside. DE JONG: Hesperudae of Surinam 261 with faint lighter brown spots. The markings of the new species are more similar to those of C. catocala Herrich-Schäffer, which, however, has the spot in space 6 on the underside of the hindwing attached to the spot in space 5 and de- tached from the spot in space 7. In the much smaller C. dagon Evans, the spots are arranged as in C. dorpa except that in the former species the spots in spaces 6 and 7 are conjoined to the cell spot. The shape of the valve of C. dorpa is not found in any other Cobalopsis species. Material examined. — Holotype, d, Surinam, Paramaribo, 3.viii.1963, leg. E. H. Jonkers. Par- atype, ®, Surinam, Paramaribo, Ma Retraite, 1.x.1960, leg. E. H. Jonkers. Both types in RMNH. 27. Cobalopsis tanna spec. nov. External characters (fig. 63). — Male. Length of forewing 13.5 mm. Nudum occupying 12 segments of which 9 on apiculus. Mid tibiae sparsely spined. No secondary sex characters. Upper side of forewing brown, possibly with yellowish basal clothing but specimen too worn to make sure; hyaline white spots in spaces 2, 3, 6, and in lower half of cell over origin of vein 3; spot in space 2 slightly closer to cell spot than to spot in space 3; a yellowish opaque spot in space 1b, and a trace of a minute dot in space 7. Upper side hindwing brown with yellowish clothing over greater part of the wing; no spots. Underside of both wings brown, markings as on upper side. Male genitalia (figs. 24, 25). — Uncus and gnathos bifid, uncus arms more widely apart than gnathos arms. Valve seemingly trapezoid in lateral view because apex turns inward. Identification. — The new species agrees well with the description of Cobalopsis as given by Evans (1955), but it does not fit any of the de- scribed species of the genus. As the specimen was collected in a Malaise trap, its condition is rather poor, but it is good enough to make sure that there are no spots on the hindwing. The on- ly known Cobalopsis species with an unmarked hindwing is C. dedecora Plötz, which has the underside of the hindwing reddish-grey and in the forewing only spots in spaces 3, 6 and 7. The presence of a lower spot only in the cell is unique in Cobalopsis. Upper cell spots are found in C. autumna Plötz and C. nero Her- rich-Schäffer, while the female of the first-men- tioned species has a lower cell spot as well; these two species have spots on the underside of the hindwing, and their genitalia are quite dif- ferent, the cucullus being long and narrow. Material examined. — Holotype, d, Surinam, Domburg, 18—23.x11.1963, Malaise trap, leg. D. C. Geijskes (RMNH). 28. Morys subgrisea paradoxa subspec. nov. External characters. — Male. Length of fore- wing 12.5 mm. Upper and underside uniform dark brown. On upper and underside of fore- wing very faint spots in spaces 2 and 3, and al- most invisible in space 6. Faintest traces of me- dian spots on the underside of the hindwing. Stigma on forewing and genitalia as in M. sub- grisea subgrisea Mabille. In size it is closer to the only other described subspecies, M. subgri- sea prada Evans (cf. Evans, 1955: 168). It can be distinguished by the almost complete lack of spots and by the lack of grey scaling on the un- derside of the hindwing, so that the name sub- grisea is not very appropriate, hence the pro- posed subspecies name. Material examined. — Holotype, d, Surinam, Zanderij, savanne, 13—16.vin.1964, leg. D. C. Geijskes (RMNH). 29. Vettius yalta Evans, 1955. The two females directly key to V. artona Hewitson or V. yalta Evans with Evans (1955). They differ from the female of V. artona in be- ing smaller (length of forewing 14—16.4 mm, as against 17.8 mm), in having the spot in space 2 of the forewing more or less under the cell spots (in V. artona mid cell spots and spot in space 3), and in having a paler underside of the hindwing. There are also differences in the genitalia. The allocation of the two specimens to V. yalta must be considered preliminary. 30. Eutychide subpunctata intermedia sub- spec. nov. Discussion. — The three specimens agree with the original description of E. sempa Evans, 1955, except for the cucullus being serrate (fig. 26). Examination of the type and the only other available male of sempa in the BM, however, re- vealed that also in E. sempa the cucullus is ser- rate, be it less obvious than in E. subpunctata Hayward. The spined dorsal projection ot the cucullus, according to Evans’ figures present in E. sempa and absent in E. subpunctata, was pos- sıbly broken in the left valve figured by Evans, as it is present in the right valve of the only specimen of E. subpunctata studied by Evans, 262 TIJpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, more or less concealed by the strong serration of the cucullus. Further differences between £. sempa and E. subpunctata are found in the un- cus arms, which are about parallel in E. sempa and divergent in E. subpunctata, and in the col- our of the wings, E. sempa being darker but with paler area along dorsum on the underside of the forewing, and of the cilia which are paler in E. subpunctata. So far, E. subpunctata and E. sempa are well- defined separate entities. However, the three specimens from Surinam are not only topo- graphically intermediate (E. sempa is only known from Guiana, E. subpunctata from Ar- gentine and French Guiana), but also morpho- logically. The cucullus is more rounded than in both E. sempa and E. subpunctata, the serration is intermediate. The uncus arms are more or less parallel. The pale dorsal area on the underside of the forewing is inconspicuous. The cilia are brownish grey. On the basis of the taxonomically as well as geographically intermediate position of the Su- rinam specimens, it seems most appropriate to unite E. sempa and E. subpunctata with the Surinam representatives into a single, geograph- ically variable species. The Surinam specimens are sufficiently different (see above) to warrant subspecific distinction; the subspecies is named Eutychide subpunctata intermedia here. Thus E. subpunctata consists of the following subspecies: E. subpunctata subpunctata Hay- ward (Argentine, French Guiana), E. subpunc- tata intermedia subspec. nov. (Surinam), and E. subpunctata sempa Evans (comb. nov.) (Guia- na). Material examined. — E. subpunctata sub- punctata: 1 3, French Guiana (BM). E. sub- punctata intermedia: holotype, d, Surinam, Paramaribo, Ma Retraite, 6—8.1.1964, leg. D. C. Geijskes; 2 d, paratypes, same data but 14— 16.1.1964 and 4—8.111.1964, respectively (all types in RMNH). E. subpunctata sempa: 2 3, 1 ©, Guiana, Takutu River (incl. holotype, ¢) (BM). 31. Talides sinois Hübner, 1819. The differences between this species and 7. alternata Bell are slight. Moreover, the differ- entiating characters given by Evans (1955: 226-227) are liable to variation. In the four males from Surinam in the RMNH belonging to T. sinois and/or T. alternata, the shape of the cucullus as well as the length of the tegumen spike is different for each male. There is also 1983 variation in the distance between spots 2 and 3 | of the forewing. It makes one wonder if 7. sinois and 7. alternata are really separate spe- cies. The material listed has been assigned to the two species according to the closest match with Evans’ descriptions, but especially for the males assigned to 7. sinois, this match is not perfect. The single female has been listed provisionally under 7. simois, no differentiating characters having been published for the female sex. 32. Carystus elvira Plötz, 1882. Evans (1955) who had no material of this spe- cies at his disposal remarked: “May be an aber- ration or a faulty figure”, a rather bold CO not seen any specimens. It is true that the figure in Draudt (1923, pl. 189b) does not seem to have entirely right colours, but Evans could not know this. Moreover, the colour differences between Draudt’s figure (probably based on the original figure by Plötz) and the specimen in the RMNH may be a sexual difference, the figure representing a male and the RMNH specimen being female. The female in the RMNH can be described as follows. Head with a conspicuous white spot directly behind the antennae. Length of fore- wing, 21.4 mm. Basal third of costa of forewing, patagia and tegulae with orange hairs. Upper side of forewing brown with sharply defined hyaline spots in spaces 1b (against vein 1), 2, 3, 6 and 7 (last two small, especially in space 7), and two spots in cell slightly basad of spot in space 2. Upper side hindwing brown, light band from base to termen on underside weakly shin- ing through. Underside of forewing as upper side, but orange along costa up to end of vein 12, from there a dull yellowish subapical band, overlapping the spots in spaces 6 and 7 (making them obscure) and ending at termen in space 4/5; pale yellowish suffusion from the spot in space 1b towards the termen. Underside of hindwing with, a warm brown-yellow colour, paler dull yellow from base through cell and space 4/5 to termen, this pale band flanked by dark brown colouring which extends more or less along the veins; space 1b greyish brown. 33. Carystoides basoches Latreille, 1824. The males agree with C. basoches. The fe- males in RMNH differ in size (forewing, 20.1 and 21.5 mm) and spotting: in the smaller speci- men the hyaline central spot on the upper side of the hindwing is relatively small and contin- DE Jong: Hesperiidae of Surinam 263 ued across space 3, in the larger specimen it is a little larger and confined to spaces 4—5. Thus, both females have some characters of C. basoches and some of C. noseda Hewitson. As the males are undoubtedly C. basoches, I have placed the females here as well. 34. Perichares deceptus Butler & Druce, 1872. The cell spots of the forewing are separate in the single male from Surinam. This is a common feature in ssp. drina Evans, but also occurs in ssp. luscinia Plötz. The lilacine areas on the un- derside of the hindwing concord with the situa- tion in ssp. /uscinia (which is only known from S. Brazil). In view of its locality, one would ex- pect the Surinam specimen to be closest to ssp. fulvimargo Butler, which occurs in Colombia, Venezuela, Ecuador and Peru, but the yellow colour at the tornus of the hindwing, character- istic of ssp. fulvimargo, does not occur in the Surinam specimen. More and better preserved material is needed to decide on the systematic position of the population in Surinam. 35. Phlebodes meesi spec. nov. Only a male available. External characters. — Length of forewing, 15.3 mm. Upper side very dark brown; inconspicuous dark ochreous hairs at base of forewing; hyaline spots only in spaces 2 and 3, spot in space 2 larger and less regular than square spot in space 3; no other spots ex- cepting a very faint spot in space 1b indicated by some pale ochreous scales close to vein 1 just beyond its middle; hindwing unmarked. Under- side forewing as upper side, purplish in apical area. Underside hindwing dark brown with purplish gloss, but space 1b and greater part of space 1c plain brown; veins a little paler, no spots. Forewing with sagittate brand over origin of vein 2, and a long narrow brand under basal third of vein 2. Genitalia (figs. 27—29). — Uncus bifid, arms about three times as long as wide, slightly diver- ging. Tegumen with central process, broad at base, much narrower in distal half, bluntly ended, not reaching base of indentation of un- cus. Gnathos consisting of two separate, sharply pointed sclerites. Cucullus with long, narrow, upturned apex; costa hardly developed, at distal end with short, inwards curving flap over basal part of cucullus. Material examined. — Holotype, d, Surinam, Lely Gebergte, 600—700 m, 30.x.1979, leg. G. F. Mees. In RMNH. Discussion. — P. meest differs from all other Phlebodes species (excepting some specimens of the South Brazilian P. smithi) in the absence of apical spots of the forewing and of any spots on upper and underside of the hindwing. In the genitalia the new species seems to come close to P. vira Butler, 1870, P. virgo Evans, 1955, and P. torax Evans, 1955, although in the genitalic drawings by Evans (1955) no trace can be seen of the central process of the tegumen. External- ly these species differ from P. meesi in the un- derside of the hindwing being ochreous-brown with conspicuous yellow veins. 36. Cynea cyrus Plòtz, 1883. The two specimens are more like ssp. hippo Evans, described from Trinidad, than ssp. rhino Evans, described from French Guiana, Upper Amazons and Pará. Length of forewing 21.2 mm. No spot in space 1b on the upper side of the forewing; apical spots in spaces 6—8 (al- most invisible in one specimen). Underside hindwing, spots in spaces 2 and 3 very faint. 37. Penicula criska extensa subspec. nov. Description (figs. 30, 31). — Male. Externally and in the genitalia similar to P. criska Evans, but differs in the shape of the valva, where the cucullus is expanded disto-ventrally, so that the distal edge is almost perpendicular to the ventral edge, and apex not extending above costa. In P. criska criska the ventral edge of the cucullus gently curves to the apex which extends above the costa. Material examined. — Holotype, d, Linker Coppename, Zuidkreek 11.vi11.1943, at light, leg. D. C. Geijskes. In RMNH. 38. Polites vibex Geyer, 1832. The extension of the tawny spots is variable, so that it is not well possible to make a sharp distinction between ssp. praeceps Scudder and ssp. catilina Plötz. This may be due to the fact that Surinam is in the area where both subspe- cies meet, from Ecuador through Venezuela to the Guianas. 39. Polites vibicoides spec. nov. External characters (figs. 64, 65). — A small replica of P. vibex praeceps/catilina. Male. Length of forewing, 10.3—10.5 mm. Markings exactly as in P. vibex, but tawny colour slightly darker and warmer. Extension of tawny colour on upper side variable; median band of fore- wing may be narrower than dark terminal area. 264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 68 69 Figs. 64, 65. Polites vibicoides spec. nov, holotype, upper- and underside. Figs. 66, 67. Cravera rara gen. et spec. nov, paratype, upper- and underside. Figs. 68, 69. Surina unica gen. et spec. nov, holotype, upper- and underside. Female as large as male; on underside of hind- wing band of yellow median spots from vein 2 to vein 8, flanked by dark brown spots in spaces 2, 3, 6 and 7, quite different from the dull grey- ish-ochreous colours of P. vibex catilina fe- males. Male genitalia (figs. 32—37). — Similar to P. vibex. Uncus and tegumen more rounded in dorsal view. Cucullus and costa of valve not suddenly narrowing as in P. vibex. Costa broadened and hollowed apically so as to re- ceive the apex of the cucullus; apical spines of costa directed caudad or more or less turned up as in P. vibex. The complex aedeagus is also similar to the structure found in P. vibex. Identification. — Polites is a mainly North American genus consisting of seven species (Evans, 1955). The only species up to now DE JONG: Hesperudae of Surinam 265 known to occur in South America is P. vibex (distributed from the northeastern U.S.A. to Argentina). The new species cannot be mistaken for P. vibex because it is much smaller (P. vibex from Surinam, length of forewing 13.8—14.8 mm). For the rest, however, they are so similar that they could be considered subspecies it they were not sympatric. Material examined. — Holotype, d, Surinam, Zanderij, savanne, 27—30.viii.1964, leg. D. C. Geijskes. Paratypes, 4 d, 2 ®, same data but 16—20.vin.1964 (1 d), 20—24.viii.1964 (1 dg, 1 9), 24 26.viii.1964 (1 3, 1 9), 47.1x.1964 (1 d). All types in RMNH. Remark. — According to Evans (1955) the Polites species have 11—12 nudum segments. This may hold true for North American species, but in the Surinam specimens of P. vibex and P. vibicoides the number of nudum segments is 13 | (figs. 38, 39). 40. Panoquina panoquinoides minima sub- spec. nov. External characters. — Length of forewing 10.5—11.7 mm. Upper side forewing brown, overlaid with yellow scales and hairs along costa and in basal third. Pale yellow opaque spot in space 1b, pale yellow hyaline spots in spaces 2 and 3, inconspicuous apical spots in spaces 6 and 7 (one or both may be absent); fringes pale ochreous. Upper side hindwing brown overlaid with yellow hairs and scales; fringes pale och- reous. Underside forewing as upper side, darker brown in basal third. Underside hindwing grey- ish brown, veins pale yellow, no spots or at most traces of inconspicuous, streaklike, pale yellow spots in spaces 2 and 3. Male genitalia. — As in P. panoquinoides Skinner. Identification. — Because of its small size it was surprising to find that the new subspecies belonged to a Panoquina species. With 12—14.3 mm for the already described subspecies, P. panoquinoides is the smallest member of the ge- nus, the other species varying from 16 to 25 mm. Apart from its size the new subspecies is characterized by the reduction (in most speci- mens complete absence) of spots on the hind- wing. Material examined. — Holotype, d, Parwa- bos, Kwattaweg naar zee. 8—11.1.1964, in Ma- laise trap, leg. D. C. Geijskes. Paratypes: 1 d, Paramaribo; 11 d, 2 ©, same data as holotype, but 1-A.ii.1964 (1 d), 8—11.11.1964 (2 d), 11— 14.1.1964 (3 8), 14—18.ii.1964 (1 3), 18— 24.41.1964 (1 3), 29.ii 2.11.1964 (1 6, 1 2), 3— 6.111.1964 (2 d, 1 9); 3 bd, Matapica in Malaise trap. All types in RMNH. 41. Cravera gen. nov. External characters. — Palpi second segment flattened, third short, hardly protruding above clothing of second segment. Antennae reaching to origin of vein 10. Antennal club swollen, three times as wide as antennal shaft, length 5/18 ot total length of antenna, constricted be- fore apiculus which is almost twice as long as width of club. Nudum covering 15 segments of which 9 on apiculus. Forewing cell almost as long as dorsum; vein 2 closer to vein 3 than to base; vein 4 midway between veins 3 and 5; vein 5 decurved at origin towards vein 4; termen al- most straight from vein 6 to vein 1, very slightly incurved in space 1b; white alice spots in spaces 1b, 2, 3, 6—7, 8 (may be absent) and cell; spots in 6 amd 7 directed to upper half of ter- men. Hindwing cell about half as long as wing measured along vein 4; vein 7 originates much closer to base than vein 2; termen almost straight (in female slightly curved) from vein 7 to vein 1b; vein 1b produced, longer than vein 1a; white hyaline spots in spaces 2, 3, 5 and 6. Hind tibiae with upper pair of spurs absent. Ab- domen as long as vein la of hindwing. Male with short inconspicuous stigmas over and un- der basal 1/4 of vein 2 and over basal half of vein 1 of forewing (fig. 40). Female with dense long hairs at tip of abdomen. Male genitalia. — Uncus and gnathos bipar- tite; apex of aedeagus with complicate processes and horns. Discussion. — See after description of the ty pe-species. Type-species. — Cravera rara spec. nov. Cravera rara spec. nov. External characters (figs. 66, 67). — Male. Length of forewing 23.9 mm. Upper side black. Forewing cell spot consisting of two spots which just touch and are placed almost parallel to termen. Hindwing with very small spot in space 4. Thorax, abdomen, base of forewing to origin of vein 2, and basal half of hindwing with bluish-grey hairs. Underside brown. Forewing on underside with white suffusion along costa over cell spot, and extensive white suffusion in space 1b under the spot in space 2 distally from and in continuation with the spot in space 1b. Hindwing on underside in addition to the hya- line spots white patches in space 1c (large, be- 266 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, tween spot in space 2 and termen) and space 7 (small, in line with other spots), and a white dot just before end cell. Fringes dark brown, white on forewing at space 1b and on hindwing at spaces 1b, 1c and, very narrowly, 2. Palpi and underside abdomen greyish, underside thorax pale bluish-grey. Antennae upper side at base of club and basal half of shaft chequered, rest black; underside white on basal half of club and upper half of shaft, rest black. Female. Length of forewing 25.5 mm. As male. Cell spot forewing undivided. Hindwing underside with same spots as male, but upper side no spot in space 4. Male genitalia (figs. 41—44). — Tegumen short and broad, with a central indentation in proximal edge, so that proximal part appears to be bilobed. Uncus dorsally with a gutterlike longitudinal depression ending in an apical in- dentation, the two apical processes very much flattened laterally and expanded dorso-ventral- ly. Gnathos consisting of two long, strongly in- curving arms, as long as uncus. Valve trapezoi- dal, cucullus squarish; dorso-distal corner of cucullus slightly prolonged and twisted; dorso- proximal corner overlapped by the weakly de- veloped costa. Aedeagus apically with long curved horns. Identification. — Externally the present spe- cies seems close to Xeniades laureatus Draudt, 1924, known only from a single male caught at Songo (Bolivia). If ever the type of the latter species will turn up, it might prove to belong to the same genus. Differences are found in the presence of a suffused white band externally of the hyaline spots on the underside of the hind- wing, white underside of body and palpi and dark green upperside of body and wing bases in X. laureatus. The differences could even be sub- specific, but without having studied the type, one cannot be sure. Material examined. — Holotype, 6, Surinam, Zanderij, Savannenbos, 28.1.1962, leg. E. H. Jonkers. Paratype, ©, same data. Both types in RMNH. Discussion. — The new genus is undoubtedly closely related to Xeniades, in which genus the apex of the aedeagus can also be provided with long processes. It differs, however, in the ab- sence of the upper pair of spurs on the hind tib- iae, the shorter hindwing cell, the black instead of brown colour of the upper side, the white instead of yellow spots on the upper side of the hindwing, and the direction of the white band on the underside of the hindwing towards the 1983 outer instead of the basal part of vein 8. Evans (1955) without having seen the type of X. laureatus nor any additional material, trans- ferred this species to the genus Vacerra, which differs from Cravera in the possession of the upper pair of spurs on the hind tibiae, in the spots in spaces 6 and 7 of the forewing being di- rected to the lower half of the termen, and in the possession in the male of a long stigma over bas- al half of vein 1 of the forewing and at most an additional short stigma under basal quarter of vein 2. 42. Surina gen. nov. External characters. — Male. Palpi flattened. Antennae longer than half length of forewing, reaching to about origin of vein 9; antennal club constricted before the angled apiculus; nudum 14 segments of which 8 on apiculus. Forewing and hindwing with hyaline spots. Forewing with linear stigmas on both sides of vein 2 and over vein 1. Cell of forewing as long as dorsum. Cell of hindwing just over half wing; termen straight, slightly excavate in space Ic; vein la shorter than vein 1b. Abdomen shorter. than dorsum of hindwing. Mid tibiae spined. Hind tibia with the usual two pairs of spurs. Discussion. — See after description of type- species. Type-species. — Surina unica spec. nov. Surina unica spec. nov. External characters (figs. 45, 68, 69). — Male. Length of forewing 18.9 mm. Forewing upper side brown; ochreous superscaling in basal part, especially along costa; pale yellowish hyaline spots in spaces 1b (against vein 1), 2, 3, 6—8 and two small spots in cell; spot in space 6 slightly out of line with spots in spaces 7 and 8, towards termen; fringes brown, paler towards tornus. Hindwing upper side brown, basal hairs more ochreous tinged; conspicuous, whitish, hyaline spots in spaces 2 and 3, and a vague spot at end cell; fringes pale brown. Forewing underside Brom, paler brown along costa and termen down to vein 2, and from costa to spot 3, leav- ing a narrow darker brown area from outside spot ın space 3 to spot ın space 6; spotting as on upper side, but spot in space 1b in a suffused white patch. Underside hindwing pale brown; spots as on upper side, spot end cell more con- spicuous; indications of small spots in spaces 4-7; ground colour slightly darker around spots. Forewing upper side with a long narrow stigma over vein 1 from just under the spot in DE JONG: Hesperiidae of Surinam 267 space 1b towards base, and narrow stigmas on both sides of vein 2 between the spot in space 2 and the origin of vein 2. Male genitalia (figs. 46—48). — Uncus slight- ly indented; arms divergent, flattened laterally. Gnathos also bifid, just reaching beyond uncus. Cucullus expanded, dorso-proximally with a rounded projection over the weakly developed costa. Aedeagus ventrally broad and flattened; a big, spiky cornutus. Material examined. — Holotype, d, Surinam, leg. Tengberg (RMNH). Discussion. — The flattened palpi, con- stricted antennal club and straight termen of hindwing readily distinguish the species as a member of the Calpodes group (Evans, 1955: 398). It fits, however, none of the genera which are recognized in this group. In Vacerra, e.g., the cell of the forewing is shorter, the abdomen as long as the dorsum of the hindwing, the stig- mas are different, and the apical spots of the forewing are exactly in line. In Niconiades the cell of the forewing is also shorter and the stig- mas different, while generally vein la of the hindwing is longer than vein 1b. Xeniades has more nudum segments, and a conspicuous white band on the underside of the hindwing. Neoxe- niades has also more nudum segments, and the wing bases blue or green. Also the genitalia do not give a clue as to which genus the new spe- cies could belong, so that I decided to erect a new genus for it. A revision of the whole Cal- podes group would be needed to establish the relationships between the genera. 43. Neoxeniades cincia Hewitson, 1871. The male and female agree entirely with the description of this species in Evans (1955), but they have a single cell spot in the forewing. Ex- amination of the three specimens in the BM (all from Pará) showed that also in this material there is a single cell spot, and it is not clear why Evans mentioned “a double hyaline white spot inicella: 44. Neoxeniades braesia Hewitson, 1867. According to Evans (1955) the cell spot of the forewing is divided in ssp. braesia to which sub- species the specimen from Surinam geograph- . ically should belong. This specimen, however, has the cell spot undivided. For the rest it agrees completely with the description. LITERATURE Bell, E. L., 1931. Descriptions of new Hesperiidae from Trinidad, B.W.I., and South America (Lepi- doptera—Rhopalocera). — Journ. New York ent. Soc. 39: 523—530. Brown, F. M., 1941. Some notes on four primary ref- erence works for Lepidoptera. — Ann. ent. Soc. Amer. 34: 127—138. Brown, F. M., & B. Heineman, 1972. Jamaica and its Butterflies. — E.W. Classey, London. Cramer, P., 1775—1780. De Uitlandsche Kapellen voorkomende in de drie Waereld-deelen Asia, Af- rica en America. Vol. 1—4 (in part) (continued by C.J. Stoll). — S. J. Baalde, Amsterdam. For publi- cation dates I have followed Brown (1941). Draudt, M., 1922. Grypocera. In: A. Seitz (ed.), Die Grossschmetterlinge der Erde. Vol. 5: 857—904. — Alfred Kernen Verslag, Stuttgart. , 1923. Grypocera. In: A. Seitz (ed.), Die Gross- schmetterlinge der Erde, Vol. 5: 905—1000. — Alfred Kernen Verlag, Stuttgart. Evans, W. H., 1951. A Catalogue of the American Hesperiidae in the British Museum. Vol. 1, Pyr- rhopyginae. — British Museum (Natural Histo- ry), London. , 1952. A Catalogue of the American Hesperiidae in the British Museum. Vol. 2, Pyrginae, Part 1. — British Museum (Natural History), London. ——, 1953. A Catalogue of the American Hesperiidae in the British Museum. Vol.3, Pyrginae, Part 2. — British Museum (Natural History), London. , 1955. A Catalogue of the American Hesperiidae in the British Museum. Vol. 4, Hesperiinae. — British Museum (Natural History), London. Fabricius, J. C., 1781. Species Insectorum. — C. E. Bohn, Hamburg and Kiel. ——, 1793. Entomologia Systematica, III (1). — C. G. Proft, Hafniae. Geijskes, D. C., 1951. General entomological research in Surinam up to 1950. — Entomology in the Netherlands and their Overseas Territories. IX Int. Ent. Congress, Amsterdam: 46—54. Hewitson, W. C., 1866. Descriptions of new Hesperi- dae. — Trans. ent. Soc. Lond. (3) 2:479—501. Holthuis, L. B., 1958. G. D. Collin’s Fauna Surina- mensis. — De West-Indische Gids 38: 71—85. Hübner, J., 1809. Sammlung Europäischer Schmett- erlinge. — Selbstverlag, Augsburg: 142. , 1818. Zutrige zur Sammlung exotischer Schmettlinge [sic!]. Erstes Hundert. — Selbstver- lag, Augsburg. , 1819. Verzeichniss bekannter Schmettlinge [sic!]. — Selbstverlag, Augsburg: 17—176. ——, 1823. Zuträge zur Sammlung exotischer Schmettlinge [sic!]. Zweites Hundert. — Selbst- verlag, Augsburg. , 1831. Zuträge zur Sammlung exotischer Schmetterlinge. Drittes Hundert. — C. Geyer, Augsburg. Jong, R. de, 1983. Rediscovery of the type of Papilio phineus Cramer and ıts bearing on the genera Phe- miades Hübner and Propertius Evans (Hesperii- dae). — Journ. Lep. Soc. 36: 279— 289 , in press. Notes on “J. C. Sepp, De Surinaamsche Vlinders”. 268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 11, 1983 Kirby, W. F., 1871. A synonymic catalogue of Diur- nal Lepidoptera. — John van Voorst, London. Mabille, P. & E. Boullet, 1912. Essai dé revision de la famille des Hespérides. — Ann. Sci. Nat. Paris 16: 1—159. Möschler, H. B., 1876. Beiträge zur Schmetterlings- Fauna von Surinam. I. — Verh. k.k. zool.-bot. Ges. Wien 1876: 293—352. ——, 1882. Beiträge zur Schmetterlings-Fauna von Surinam. V. Supplement. — Verh. k.k. zool.-bot. Ges. Wien 1882: 3—62. Plötz, C., 1879. Die Hesperiinen-Gattung Pyrrho- pyga und ihre Arten. — Stettin. ent. Ztg. 40: 520—538. ——, 1882a. Einige Hesperiinen-Gattungen und deren Arten. — Berl. ent. Z. 26: 71—82. , 1882b. Die Hesperiinen-Gattung Hesperia Aut. und ihre Arten. — Stettin. ent. Ztg. 43: 314— 344, 436—456. ——, 1883. Die Hesperiinen-Gattung Hesperia Aut. und ihre Arten. Fortsetzung. — Stettin. ent. Ztg. 44: 26— 64. , 1884. Die Hesperiinen-Gattung Apaustus Hub. und ihre Arten. — Stettin. ent. Ztg. 45: 151—166. , 1886. Nachtrag und Berichtigungen zu den Hesperiinen. — Stettin. ent. Ztg. 47: 83—117. Sepp, J. C., 1829—1852. De Surinaamsche Vlinders. — Sepp, Amsterdam. For publication dates, see — De Jong, in press. Stoll, C., 1780— 1791. De Uitlandsche Kapellen voor komende in de drie Waereld-deelen Asia, Africa — en America. Vol. 4 (in part), Supplement (contin- uation of P. Cramer, 1775— 1780). — S. J. Baalde, Amsterdam. For publication dates I have followed Brown (1941). Williams, R. C. & E. L. Bell, 1931. Hesperiidae of the Forbes Expedition to Dutch and British Guyana. — Trans. Amer. ent. Soc. 57: 249—287. Mi DEEL 126 AFLEVERING 12 1983 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING MUS. COMP. ZOOL LIBRARY 0 1984 HARVARD UNIVERSITY INHOUD M. A. LieFrINcK. — Notes on the nomenclature and synonymy of Old World Me- lectine and Anthophorine bees (Hymenoptera, Anthophoridae), pp. 269—284, figs. 1—43. Tijdschrift voor Entomologie, deel 126, afl. 12 Gepubliceerd 15-XH-1983 NOTES ON THE NOMENCLATURE AND SYNONYMY OF OLD WORLD MELECTINE AND ANTHOPHORINE BEES (HYMENOPTERA, ANTHOPHORIDAE) by M. A. LIEFTINCK “Kalliste”, Rhenen, The Netherlands ABSTRACT Tetralonioidella Strand, 1914, is a forgotten name that has fallen into oblivion ever since it was defined and published. The name was given to an anthophorid bee of the Melectinae, which was described in some detail from Taiwan. As a validly proposed name it is resur- rected, its monobasic type-species, T. hoozana Strand, 1914, being recognized as a distinct species, congeneric with, but differing specifically from, all described taxa formerly placed in Protomelissa Friese, 1914 (syn. nov.) or Callomelecta Cockerell, 1926. All ten presently known species are (re)defined, arranged in a key and, where necessary, illustrated by the author. Included are 7. nepalensis spec. nov. (Nepal) and fukienensis spec. nov. (SE China), the latter being the first of its genus recorded from the Chinese continent. A redescription and figures are given of the little known holotype of 7. habropodae (Cockerell, 1929), from Thailand, while the only known specimen of T. iridescens (Friese, 1914), from Taiwan, is probably lost but considered conspecific with 7. hoozana Strand from the same island. A second chapter deals mainly with six undoubtedly validly proposed specific names given by Newman (1835) to trivial variations of Melecta albifrons (Forster, 1771), collected in England. Two of these, alecto and megaera, were first described as new by Newman but quite accidentally proposed also by Lieftinck (1974), to denote two new species found in the Near East. These new homonyms are here replaced by M. diligens nom. nov. and mun- dula nom. nov., respectively. — Lastly, in the anthophorine host bee genus Habropoda F. Smith, the unique type of H. krishna Bingham, 1909, proved to be conspecific with H. apa- telia Lieftinck, 1974, the last-mentioned name thus being placed in the synonymy of H. krishna Bingham. Tetralonioidella STRAND, 1914, A PARASITIC ORIENTAL BEE GENUS RE-INSTATED Since publication of my revision of the melec- tine genus Protomelissa Friese, 1914 (see Lief- tinck, 1972: 267—282, figs. 1—16, pl. 1 figs. 1, 2, with map), some important nomenclatural changes on the genus and species level have again become necessary. As pointed out in that last paper, Friese’s diagnosis of Protomelissa was published in June, 1914. Since the two Tai- wanese melectines Anthophora sauteri Friese (original description published 15 May, 1911), and Melecta formosana Cockerell (the same dated March, 1911), turned out later to be con- specific, formosana became the type-species of Protomelissa. However, at that time it was still unknown that few months earlier (April —May, 1914), E. Strand had already published notes and a description of a very puzzling bee, like- 269 wise found in Taiwan (Formosa) which he “conditionally” named Tetralonioidella, with the monobasic type-species 7. hoozana Strand. As we will see, the unique type of this bee is congeneric with both Protomelissa Friese and the much later described genus Callomelecta Cockerell, 1926 (type-species C. pendleburyi Cockerell, from the Malay Peninsula). The original account of Tetralonioidella hoo- zana Strand was published in German and, for a better understanding of the situation, is here copied verbatim under that species. By the ab- sence of a female, the unique male was mistaken for some non-parasitic, pollen-collecting mem- ber of the Eucerinae presumably related to Tetralonia. Unfortunately, this resulted in the choice of that whimsical generic (or subgeneric) name Tetralonioidella which is, of course, a ver- itable misnomer. It is not strictly a nomen obli- tum since the whole description was simply 270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 overlooked and forgotten ever after its intro- duction, no mention having been made of it in any catalogue or faunal list of the Apoidea. The reasons for this neglect are easily understood because (1) Strand’s story of the affinities of Tetralonioidella is confused, the whole account being full of thoughts couched in decepuve terms leaving no room for conclusions; (2) one of the most outstanding characters of this bee, viz., the presence of a pair of robust mesoscutel- lar processes, was left unnoticed; fand (3) Friese’s almost simultaneously published defi- nition of the genus Protomelissa was not known to Strand. Considering the above facts, it will be clear that Tetralonioidella, though proposed condi- tionally, is the oldest available name amongst those of the included co-ordinate synonyms, i.e. Protomelissa and Callomelecta. Even if the pro- visions laid down in art. 33 (sect. b i and ii) of the law of priority in the Code, are taken into consideration, its validity and re-introduction cannot be called in question. The synonymy, then, is briefly as follows: Tetralonioidella Strand Tetralonioidella Strand, April—May, 1914: 139—141 (type-species: “Tetralonia (?) hoozana” Strand, 1914). Protomelissa Friese, 1 June, 1914: 322, 323 (type-spe- cies: Anthophora sauteri Friese, May, 1911 = Me- lecta formosana Cockerell, Mar., 1911). — Sand- house, 1943: 592 (type-species invalidly proposed: Protomelissa iridescens Friese, June, 1914 = Tetra- lonioidella hoozana Strand, April—May, 1914; iridescens syn. nov. — Lieftinck, 1972: 260, 261 (generic key), 267—270 (diagnostic char.) (type- species: Melecta formosana Cockerell, Mar. 1911). Callomelecta Cockerell, 1926: 621 (type-species: Cal- lomelecta pendleburyi Cockerell, 1926). — Lief- tinck, 1944: 58—62 (gen. & spec. redefined); Lief- tinck, 1972: 269, 270 (synon. notes). The fullest account of the present genus is the one given by Lieftinck (1944, sub Callomelec- ta). It was based on a study of both sexes of three Malaysian species which on that occasion were elaborately described and keyed. These and other species were arranged also in the key published by me at a much later date (1972, sub Protomelissa). Except for specific differences in sculpture, body-colour, pubescent pattern and sexual structures, the above definitions of the male are fully applicable to the type-species 7. boozana, of which sufficient details can be found in the next descriptive key and in Strand’s original narrative copied thereafter. KEY TO THE MALES OF TETRALONIOIDELLA (N.B. — The d of T. tricolor, and the 2 of 5 other species are unknown) 1. Antenna long and slender, surpassing tegu- la; scape short, claviform, little curved, less than three times as long as diameter at apex; anterior face with conspicuous compact | patch of longish, partly raised, silky and finely branched, pale yellow hairs pointing apicad. Segment 2 hairy, extremely short and annular, retracted and occasionally hid- den from view (fig. 20); 3 also short but nearly thrice as long as 2, shiny and cup- shaped, only slightly shorter than its width at apex; 3—13 hairless or almost so, straight and cylindrical; 4—6 slender, 4 up to three times longer than its greatest diameter, 5—6 subequal but shorter than 4; remaining seg- ments successively shorter toward end, more or less squarish, each with lateral cari- na, posterior faces rather flattened and in- creasingly more distinctly crenulated (fig. 13—14, 20). Thorax bulky, clothed densely with long fine plumose hairs. Abdominal tergites not banded, with short varicol- oured pubescence. Inner rami of mid and hind tarsal claws distinctly shorter than outer, both acuminate and claw-like, not squarishtoraxe- shaped 2 — Antenna shorter, not or barely surpassing tegula; scape long and slender, at least three times longer than broad, variably curved, often hairy but lacking conspicuous com- pact brush of finely branched silky hairs an- teriorly; segment 2 often short and annular though not retracted and always plainly vis- ible (figs. 10—12); 4—5 less markedly elon- gated, following segments relatively shorter than in next three species, not noticeably crenulated; last segment only little flattened below, not downcurved. Mandibles uniden- tate. Inner rami of mid and hind tarsal claws much shorter, squarish or axe-shaped (fig. 5). kts dead idk ote 3 2. Integument of head, all thoracic sclerites and propodeum, deep black, only the face reddish black; ground colour of abdomen shining bright orange-rufous above and un- derneath, only the (partly retracted) tergites 4—6 on either side in front of graduli somewhat obscured, as is also the extreme LIEFTINCK: Melectine and Anthophorine bees 271 base of 7. Tergites not banded, almost bare, clothed sparsely with extremely minute ap- pressed pale hairs; graduli at sides of 1, and 2 except upon middle, with narrow but slightly longer silvery yellow appressed hair-lines, also seen on about half of ex- posed portion of 4; hairs longer and sparser on disk of next tergites and on apical ster- nites, integument of the latter somewhat obscured upon middle at base. The follow- ing body parts are also light or dark ferrugi- nous: glossal galea, mandible-bases, la- brum, clypeus and antennae anteriorly, te- gulae, and legs including coxae and trochanters. Antennal scape and flagellar segments posteriorly, brownish. Labrum relatively long: subequal to its width near base, anterior border broadly and deeply emarginate with well-rounded lobes (fig. 19). Mandibles simply sickle-shaped, lack- ing interior subapical tooth (figs. 17, 18). Tergite 7 shallowly excised, broadly V- shaped (fig. 21). Antennal segments (fig. 20) shaped much as in himalayana and formo- sana, the pale silky hair-patch covering front of scape conspicuous, flagellar seg- ments less markedly crenulated but last joint distinctly swollen basally, then strong- ly hollowed out, downcurved and tapering to a point. Thorax posteriorly glossy red- dish black; scutellum behind with median carina thickened at apex; spines long and slender, finger-like, directed straight back, feebly downbent, apices reddened, distinct- ly pointed, shorter than surrounding pubes- cence; metanotum and propodeum entirely smooth and polished. Inner rami of mid and hind tarsal claws but little shorter than out- er. Thoracic pubescence uniformly pale- coloured, long and fluffy, hairs rather shorter and less closely set than in next two speeteslihanlamde.ps. 24am ener: habropodae Ground colour of abdomen less shiny, at least with basal portion of tergites 1—2 dark brown or black, clothed with short ap- pressed tomentum differing in character: hairs either sparsely distributed, shortly branched, thin and darkest on exposed bas- al surface, or longer, more crowded togeth- er and distinctly plumose upon broad, paler-coloured apical margins of tergites; hence only basal part of 1—2 fully exposed and shiny, those of 3—7 withdrawn, leav- ing only dullish distal portions covered with closely set, dark golden-yellow plu- mose hairs. Mandibles with single interior subapical prominence. (For further details, see Lieftinck, 1972). Himalaya and Tai- want: himalayana and formosana . Pubescent colour-pattern of thorax and ab- domen strongly contrasting, pile on thorax long, dense and feathery, fox-red to orange- rufous; and on abdominal tergites very short, scanty and mainly dark on glossy deep black ground, not concealing surface, tergites not or incompletely pale-banded. Three small to medium-sized closely similar species, all with labrum concave dorsally, strongly deeply punctate, the anterior bor- degdistinctlyauptunne disses sage 6 Pubescent colour-pattern of thorax and ab- domen more uniform: pile on thorax dense though somewhat shorter and less bright, ochraceous-buff to ochraceous-orange; very short, less vivid and often forming complete, more or less definite hair- bands on dullish dark brown abdominal tergitesy u. sert cie hr Etsen 4 . Size rather large: body length ca. 13.5 mm; elongate, thorax and abdomen subequally broad, the latter gradually tapering, dull, lacking definite hair bands. Integument of abdominal tergites and sternites distinctly bicoloured: basal (i.e. narrowest) portion of 1—6 dark brown, the distal (postgradular and broader) parts of same semitransparent, distinctly paler, yellowish brown; all ter- gites with extremely short appressed hairs entirely concealing a finely superficially punctate surface, colour throughout ochra- ceous-buff to ochraceous-orange, hairs at bases of tergites 2—4 shortest and but little darker than on remaining parts. Labrum much shorter than broad, widest basally, apical border shallowly emarginate (fig. 4). Antenna relatively short, hardly reaching tegula; scape little curved, straight in dorsal view, 2 small but distinct, slightly broader than long, 3 and 4 elongate, distal flagellar segments only slightly crenulated (figs. 1— 3). Metanotum finely wrinkled, base of pro- podeal triangle shiny though coarsely ru- gose for a distance equaling length of meta- notum, triangle for the rest (save laterally) smooth and polished. Hind tarsal claw, fig. 5. Apex of tergite 7, fig. 6. Hidden sternites and genitalia, figs. 7—9. Dorsal thoracic pubescence relatively short but forming tufts of much longer hair surrounding, and practically concealing, the long curved scu- 272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 cellanspimes maman hoozana Generally smaller, body length 8.5—12.5 mm. Integument of abdominal tergites not noticeably bicoloured, i.e. uniform dark brownish black or black; tergal pubescence either short, decumbent and uneven, or forming more or less definite transverse pale subapical bands. Two very similar, probably nearly related species whose gen- eral appearance, texture and nature of pu- bescent pattern are almost identical ...... 5 . Labrum short, broadest and more parallel- sided near base, and also shorter than in both hoozana (fig. 4) and fukienensis, mid- length to breadth ratio = 60.4 : 100, the emargination smaller and less deep than in hoozana, resembling fukienensis most closely (fig. 27). Antennal segments 3—4 subequally long, both relatively shorter than in fukienensis (fig. 12). Plate-shaped inner rami of mid and hind tarsal claws squarish, apices straight cut off. Apex of tergite 7 deeply broadly emarginate, the small prominent lobes rounded off (fig. 31). Mid-apical plate of sternite 7 parallel-sided (fig. 32). Dorsal thoracic pubescence xan- thine-orange, hairs longer than in fukienen- sis; appressed tergal abdominal hair-bands 1—5(6) but little paler. Antennae and legs throughout dark reddish brown, tarsi be- coming ferruginous distally. Body length 11.5—12.5 mm. For detailed descriptions of both sexes, see Lieftinck (1944: 71—75). Wes tala anette: eget re dt insidiosa Labrum longer than in both insidiosa and hoozana (ratio = 75 : 100), greatest width slightly before midway length, sides more convex. Metanotum and propodeal triangle not very shiny, microscopically tessellate, extreme base of triangle distinctly wrinkled but lacking the strong ridges on either side of the median line seen in insidiosa; for the rest, propodeum, like insidiosa, more shiny and strongly punctate. Basal antennal seg- ments, fig. 11. Plate-shaped inner rami of mid and hind tarsal claws axe-shaped, apices subacute: obliquely cut off. Apex of tergite 7 as in insidiosa, but emargination less wide and deeper, subrectangulate, the lobes more broadly rounded (fig. 28). Mid- apical plate of sternite 7 distinctly con- stricted basally, rather spatulate (fig. 29). Dorsal thoracic pubescence and tergal ab- dominal hair-bands as in insidiosa but all hairs shorter and paler, those covering ter- gites finer and more closely set. Antennae black above, dark reddish brown below; legs also darker than in insidiosa, the apical tarsal segments more reddish brown. Body length 11.5 mm, fore wing 8.5 mm. SE China. 200): TRES fukienensis . Labrum (fig. 22) not much broader than long, widest basally, slightly narrowing to- ward end, with outwardly convex sides, an- terior border strongly upturned, more or less pinched in the median line, the apical emargination small and shallow in full dor- sal view; surface shiny, strongly deeply punctate, punctures circular, smaller than interspaces. Antennal 4 sligthly longer than 3 or 5 (fig. 10, ventral view) or subequal (dorsal view), and also differing from the next ones by having a dense coating of ex- tremely minute, raised, silvery hairs; erect hairs on scape also longer than in next spe- cies. Scutellar spines finger-like, strongly downcurved, apex subacute or rounded, neither compressed nor bifid, surrounded by long fox-red pubescence. Metanotum finely wrinkled; propodeal triangle glossy black, throughout smooth and polished. Dorsal thoracic pubescence as in pendlebu- ryi. Abdominal tergites 1 and most of 2 clothed sparsely with depressed plumose, pale orange-yellow hairs, those on 2 form- ing ill-defined postgradular bands not quite concealing surface; 3—4 similarly banded, subinterrupted by black. Apex of tergite 7 hairless, shaped as in fig. 23. Hidden ster- nites 7—8 and genital capsule, figs. 24 and 25— 26, respectively. Size relatively small, total length 9.5 mm, fore wing 7.7 mm. Ne- pal or ee eee nepalensis Labrum distinctly broader than long, even more markedly so than in hoozana (fig. 4), anterior border either rather widely and deeply emarginate (vulpecula), or almost straight in dorsal view (pendlebury:). An- tennal 3 and 4 subequal in length (figs. 15— 16), raised hairs on scape and next segments all shorter and more sparsely distributed. Abdominal tergites deep black, 2—6 scar- cely or not at all banded, the much shorter tomentum thin and more even .......... 7 7. Anterior border of labrum hardly upturned and almost straight in full dorsal view. Api- cal emargination of abdominal tergite 7 deeply U-shaped, subequal in form to the tubercles (Lieftinck, 1944, fig. 6), the punc- tation similar to nepalensis. Scutellar spines LiEFTINCK: Melectine and Anthophorine bees 273 fi iil INN ( Hh K ANY 6 N R G 4 SSNS G 5 NEGO A Ts YW. A III AN A; in ETN NN LD) 1) | \\ NS 01) | à Fig. 1—9. T. hoozana Strand, 8 structures, holotype Taiwan; fig. 1—3, basal segments of right antenna: fig. 1, lateral view of scape, long sparse raised hairs omitted; fig. 2—3, segm. 2—8, oblique ventral (2) and dorsal view (3); fig. 4, dorsal view of labrum; fig. 5, right hind tarsal claw, oblique inner view; fig. 6, apex of tergite 7, dorsal view, long bristly basal hairs mostly omitted; fig. 7—8, sternites 7 and 8, external view; fig. 9. genital capsule, ventral view of right half, scale line 1 mm. 274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 robust, much shorter than in nepalensis, less downcurved, apex laterally compressed, distinctly bifid forming a pair of little knobs. Metanotum and propodeum dullish, finely tessellate, smooth, but much less shi- ny than in nepalensis, base of triangle coar- sely punctate on either side, these sclerites separated from each other by a deep sulcus. Dorsal thoracic pubescence slightly paler than in vulpecula. Malay Peninsula .…. PARO EI MOORE EE pendleburyi — Anterior border of labrum abruptly up- turned, the latter shaped much as in nepa- lensis but longer, emargination distinctly wider and deeper than in that species. Scu- tellar spines still shorter than in pendlebu- ryi, directed obliquely caudad and but slightly downbent, apex somewhat com- pressed and tapering, slanting and bluntly pointed. Metanotum and propodeum much as in pendleburyi, but basal area of triangle coarsely longitudinally punctate along full breadth of same. Dorsal thoracic pubes- cence bright xanthine orange. Apex of ab- dominal tergite 7 with shallow impunctate median sulcus, otherwise black-haired, emargination shallower and wider than in both nepalensis (fig. 23) and pendleburyi (ie 27), SOMA RR RA vulpecula Tetralonioidella hoozana Strand (figs. 1-9) Tetralonioidella (“Tetralonia ? hoozana n.sp.”) hoo- zana Strand, 1914 (April/May): 139—141 (d For- mosa, Hoozan, ix.1910). Protomelissa iridescens Friese, 1914 (1 June): 324 (d Formosa, Takao, Sept. 1908, H. Sauter). — Sand- house, 1943: 592 (type-species of Protomelissa, not seen). — Lieftinck, 1972: 272 (key), 277, 278 (orig. descr. & notes; type not seen). Syn. nov. Type material. — 1 d (terminalia dissected out, glued on card and pinned on insect’s pin), labelled: “Tetralonia (?) hoozana m. & Strand det.” (Strand’s writing), “Hoozan Formosa H. Sauter Og (prin Ed ate watten) WIP IS” (typewritten on red), “Tetralonioidella hoozana Strand HOLOTYPE (= Protomelissa iridescens Friese), rev. MA Lieftinck 1979”. Holotype d in the former Deutsch. Entom. Institut (Eberswalde). This is the first described and largest member of the genus. It fits the existing amended diag- noses of both Callomelecta and Protomelissa in all important characters and now becomes the type-species of the monobasic Tetralonioidella. Male. — The uniform tint of the body pubes- cence is longest, tufted, and deepest orangish on the thoracic dorsum surrounding the practically invisible mesoscutellar processes, these long feathery hair-tufts being tipped with pale yel- low. The neuration of the fore wing is much as in the drawing given for T. pendleburyi (see Lieftinck, 1972, fig. 11), i.e. the two proximal sides of the 2nd submarginal cell are subequal in | length while the costal and anal sides of the lat- ter are also about equally long though both are slightly less than half as long as the proximal sides. The original description of T. hoozana is very full, as far as it goes, and is best copied along with the author’s comments at full length, as follows. “Gen. Tetralonia Spin. Tetralonia (?) hoozana Strd. n.sp. Ein d von Hoozan IX.1910. Dies Tier ist gewissermaszen ein Mittelding zwis- chen Tetralonia und Anthophora. Eine Anthophora kann es aber nicht sein, weil die Ozellen in gerader oder fast gerader Reihe angeordnet sind (bei Antho- phora bilden sie bekanntermaszen ein ausgespro- chenes Dreieck)!). Fur ein Tetralonia-Mannchen sind aber die Antennen eigentlich viel zu kurz; schon da- durch läszt die Art sich leicht von der sonst offenbar sehr ahnlichen Tetr. himalayensis Rad. unterscheiden. — Von Ancyla Lep. abweichend dadurch, dasz die hinteren Metatarsen des d kaum so lang wie die fol- genden Tarsenglieder zusammen sowie nicht ge- krümmt sind, ferner ist der Clypeus vorstehend und die Körpergrösze ist bedeutender als bei den bisher bekannten Ancyla; soweit ohne Praparation erkenn- bar, sind die Mundteile ziemlich kurz. — Das zweite Geiszelglied ist kaum so lang wie das dritte und auch nicht dünner, das Tier somit, sowie durch die unter sich weit entfernten und subparallelen Augen von Meliturga Latr. leicht zu unterscheiden. Es ist ganz wahrscheinlich, dasz fur dies Tier eine neue Gattung aufgestellt werden musz (die eventuell den Namen Tetralonioidella m. bekommen möge); um dies mit Sicherheit zu entscheiden, wäre aber die Kenntnis auch des © eigentlich nötig. Aber auch wenn dies eine typische Tetralonia sein sollte, dürfte die neue Benennung als Untergattungsname verwend- bar sein. Schwarz; Mandibeln leicht gebräunt ın der Mitte, 1) Anm. Die von Friese in: Verh. zool.bot.Ges. Wien 1911, p. 27, beschriebene Anthophora Sauteri n.sp. ist eher zu Tetralonia zu stellen, weil die Ocellen, wenigstens beim ©, kein Dreieck bilden. In meiner Besprechung der Art im I. Teil dieser Arbeit (in: Supplem. Entomol. II, p. 51) habe ich leider unter- lassen, dies Moment hervorzuheben. LiEFTINCK: Melectine and Anthophorine bees 275 Fig. 10—16. Tetralonioidella species, partial view of right and left d antennae; fig. 10, 7. nepalensis spec. nov., Nepal, right segm. 1—8 showing plainly visible 2nd segment, long sparse hairs at scape omitted; fig. 11, 7. fu- kienensis spec. nov., SE China, dorsal view of right segm. 1—7, same details; fig. 12, T. insidiosa (Lieft.), W. Java, right segm. 2—8, same scale; fig. 13—14, T. himalayana (Ckll.), Bhimtal, antero-ventral view of left segm. 2—13 and 1—7, lateral view, showing retracted 2nd segment (fig. 14 more enlarged, long silky hair-tuft at scape omitted); fig. 15. 7. vulpecula (Lieft.), S. Sumatra, right antennal segm. 2—6, showing cup-shaped 2nd segment; fig. 16, 7. pendleburyi (Ckll.), Larut Hills, Perak, the same of segm. 2—7 showing slightly shorter 2nd segment. 276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 Labrum an der Basis beiderseits gerötet, Fühlergeiszel unten schwach gebräunt, Tegulae bräunlichgelb, der hintere Hälfte der Abdominalsegmente gerötet, die Tarsen leicht gebräunt. Flügel gleichmäszig schwach angeraucht und etwas gelblich schimmernd, Geäder und Mal braunschwarz. Der ganze Kopf hell graugelblich behaart und zwar auch auf dem Labrum lang abstehend, auf dem Cly- peus auszerdem mit anliegender Behaarung. Thorax sehr dicht und lang ziemlich abstehend behaart und zwar auf dem Rücken orangegelblich, an den Seiten und unten heller behaart; die Skulptur daher nicht er- kennbar. Rückensegmente des Abdomen mit feiner, kurzer, anliegender, nicht dichter, goldgelbener Be- haarung, die auf der helleren Hinterhälfte der Seg- mente kaum heller als auf der basalen ist, als Gesamt- eindruck heben die helleren Binden sich daher wenig von der Grundfarbe ab. Die Bauchsegmente verhalten sich wie die Rückensegmente, jedoch ist ihre Behaa- rung in der Endhalfte leicht abstehend und die Seg- mente II und III zeigen auszerdem eine schmale helle Basalhaarbinde. Die Beine kurz und sparlich messing- gelblich behaart. Kopf schmäler als Thorax, aber breiter als lang, mit groszen, vorstehenden, subparallelen (nach unten ganz schwach konvergierenden), innen nicht ausge- randeten Augen und stark vorstehenden, vorn mitten jedoch abgeflachtem Clypeus, dessen Kontur, in An- sicht von der Scheitel, fast trapezförmig erscheint, im Profil scheint die Vorderfläche des Clypeus mit dem Labrum fast einen rechten Winkel zu bilden. Letz- teres beiderseits an der Basis höckerig, am Vorder- rande (im Profil gesehen) leicht vorstehend. Die An- tennen überragen die Mitte des Mesonotum, die Geis- zel zylindrisch, nur das erste und die Basis des zweiten Gliedes etwas dünner als die übrigen; das zweite Geiszelglied ist etwa dreimal so lang wie das erste. Abdomen erscheint in Draufsicht an der Basis breit quergeschnitten, am Ende zugespitzt, das letzte Tergit an der Spitze mitten leicht eingeschnitten. — Die dritte Kubitalzelle ist oben (vorn) so breit wie hinten, die zweite Kubitalquerader ist kurz unterhalb der Mitte stark saumwärts konvex gebogen, die zweite Kubitalzelle ist auf der Radialader nur halb so lang wie auf der Kubitalader. Körperlänge 13.5, Flügellänge 19 mm”. Protomelissa iridescens Friese, — a dubious synonym? For a discussion of this puzzling species, see the above citations, one of which includes a transcription of the original diagnosis. Expect- ing the type not to turn up any more in some European museum or private collection, it must be regarded as lost or destroyed. The locality labels and dates of capture of the types of T. hoozana and iridescens are quite different, so that the chances are very slight that Strand and Friese, at a time when both authors were studying bees from “Sauter’s Formosa Ausbeu- te”, did come across the same specimen and based their descriptions successively on this one individual. On the other hand, with the recent discovery of 7. hoozana, one would be inclined to think sridescens and hoozana are identical species, because the probability of more than two taxa of this rare genus occurring together in such a small island as Taiwan, seems unlikely. Yet the obvious discrepancies in structure pres- ently observed (table 1, especially points 3 and 4) in the types (if correctly stated for irides- cens!), should not be neglected. Despite the in- congruities I have, with all reserve, decided to synonymize these two Taiwanese species. Tetralonioidella formosana (Cockerell) comb. nov. Melecta formosana Cockerell, 1911: 227, 228 (2 Ko- sempo, Formosa). Anthophora sauteri Friese, 1911: 127, 128 (8 Tainan, Formosa). Protomelissa sauteri; Friese, 1914: 323, 324 (62 Tai- nan & Takao, Formosa). Protomelissa formosana; Lieftinck, 1972: 273 (key), 274—277, figs. (6 2 Kosempo, Tainan & Takao; orig. & further descr., synon. & full refs.). Table 1. Differences in structure in the types of T. iridescens and T. hoozana. T. iridescens T. hoozana 1. Abdomen dull and with slight metallic sheen (“Abdomen mit Erzglanz”, and “fast matt, schwach erzfarben schillernd”); 2. Labrum “viereckig, vorn schwach ausgerandet” 3. Antenne rot. . . 2tes Geisselglied (segm. 3) viel kürzer als 3” 4. Apex of tergite 7 small and truncated (“klein und abgestutzt”) 5. Body length 13 mm, width 4.5 mm 1. Abdomen somewhat shiny but lacking any indication of metallic gloss 2. Labrum shaped as in fig. 4 3. Antenna yellow-brown in front, dark brown behind, segm. 3 and 4 subequal in length (figs. 2, 3) 4. Apex of tergite 7 distinctly excised (fig. 6) 5. Body length 13.5 mm, width 4.5 mm LIEFTINCK: Melectine and Anthophorine bees Tetralonioidella himalayana (Bingham) comb. nov. (figs. 13, 14) Melecta himalayana Bingham, 1897: 516, fig. 172, in- sect, with left wing (2 “Kumaon, 5000 ft., and probably throughout the Himalayas at and above that altitude”. Protomelissa himalayana; Liefunck, 1972: 273 (key), 274, figs. 3, 7, 8 (addit. notes & d struct., 49 N & NE India). Additional material. — Series of 12 6, NW India, U.P., Bhimtal near Nainital, 1500 m, 13, 14, 15 and 18.x.1978, M. A. Lieftinck, all at flowers of Labiate shrub Plectranthus coetsa Ham. ex D. Don. (local name “bhanira”). The above Bhimtal males are freshly emerged specimens in perfect condition. They were tak- en within a week’s time, either in the early morning hours around 8.30 a.m., or on a return visit at 17 p.m. (4 on Oct. 14, 5 on Oct. 15, most of the remainder solitarily). All were attracted by flowers of the same plant, which grew in threes or fours at some distance from each other along a shaded forest path at the slope of a rav- ine. The bees flew in company with both sexes of 5 species of Habropoda, viz. H. apostasia m., deiopea (Cameron), hooker: Cockerell, pelmata m., and radoszkowsku (Dalla Torre), the latter being by far the most common of these; fe- males, besides taking nectar, assembled pollen at their hind tibiae and basitarsi, apparently also from Plectranthus. Their high-pitched notes could be heard already at about three yard’s dis- tance. I had asked Mr. Fred Smetacek, the own- er of a small undisturbed patch of forest in the hills above Bhimtal, to keep an eye on the flow- ering season of Plectranthus, in 1979 and 1980. He kindly informed me in his letters that, un- fortunately, no single inflorescence ot plants could be found by him in any part of the same area during these next two years. As pointed out by me earlier (1972: 273— 277, sub Protomelissa), himalayana comes so close to formosana in stature, pubescent colour- pattern and other bisexual characters, that I have failed to discover more features than those already known for their separation. Nothing definite is known of their host relations, but both were associated in the field with Habropo- da species, not with those of Elaphropoda as were other members of Tetralonioidella. Tetralonioidella tricolor (Lieftinck) comb. nov. Protomelissa tricolor Lieftinck, 1972: 273 (key), 278— 277 281. fig. 4 (labrum; 2 Assam, Shillong; notes & host relations). The © holotype in the Berlin museum is still the only specimen so far known. By the absence of a d, nothing definite can yet be said about the nearest relatives of tricolor, but as stated in the original description, we may expect it to come closest to himalayana and formosana. The presence of a conspicuous dense brush-like patch of long hairs in front of the antennal scape of the 6 (loc.cit.: 273), is a unisexual character, so that my subsequent remark (loc.cit.: 278) on its absence in the © as a point of distinction be- tween species-groups, is misleading. Tetralonioidella habropodae (Cockerell) comb. nov. (figs. 17—21) Callomelecta habropodae Cockerell, 1929: 133. — 3 “Siam: Doi Sutep, on summit, Feb. 9, 1928 (Cock- erell)”. — Lieftinck, 1944: 62, 75, footnote (origi- nal material not seen). Protomelissa habropodae; Lieftinck, 1972: 270 (key), 281 (orig. descr. copied, with notes; original material not seen). Type material. — 1 d (holotype), labelled: “Doi Setep Siam feb. 9 Alice Hacker” (?) in un- known writing; and “Callomelecta habropodae Ckll. TYPE”, in T.D.A. Cockerell’s hand; and, “acc. 35740” (AMNH label). Holotype in Amer. Mus. Nat. Hist., New York. The more interesting characters of this re- markable little species are summarized in the preceding key. T. habropodae is known only from the unique holotype d. In addition to the key notes and the original description, copied in full on a previous occasion (Lieftinck, loc.cit. 1972: 281), most of the hitherto unnoticed pe- culiarities of structure are here also illustrated (figs. 17—21). Apart from its smallish size and the predomi- nantly orange-brown colour of the legs and ab- domen, the species is easily distinguished from all allies by a combination of the following char- acters: dense brush-like hair-tuft in front of the antennal scape; characteristic shape of the mid and hind tarsal claws; the unarmed mandibles; and the superficially punctate shiny surface of the first two abdominal tergites. Disk ot labrum (fig. 19) slightly concave, rather shiny, rugosely punctate, colour yellow-brown, its anterior border narrowly ferruginous, long hairs semi- erect, pale yellow, the border fringed with a 278 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 \ iT \ I th | 4/23 A 3 N N È Mh i fi a 26 \ WW Fig. 17—21. T. habropodae (Ckll.), Thailand, & holotype Doi Sutep; fig. 17—18, left mandible, inside and exte- ro-lateral view; fig. 19, dorsal view of labrum; fig. 20, right antennal segm. 1—9, showing long silky hair-tuft at scape and partly retracted 2nd segment, inner (ventral) view; fig. 21, apex of tergite 7, dorsal view, most longish appressed hairs omitted. — Fig. 22—26. T. nepalensis spec. nov., 6 holotype Nepal; fig. 22, dorsal view of la- brum; fig. 23, apex of tergite 7, partly lacking bristly hairs; fig. 24, sternites 7 and 8, external view; fig. 25—26, genital capsule, dorsal and ventral view, scale line 1 mm. LIEFTINCK: Melectine and Anthophorine bees 279 \ } Wit = & Fig. 27—30. T. fukienensis spec. nov, & holotype SE China; fig. 27, dorsal view of labrum; fig. 28, apex of ter- gite 7, long depressed silky hairs mostly omitted; fig. 29, sternites 7 and 8, external view; fig. 30, genital capsule, dorsal and vertral view, scale line 1 mm. 280 row of stiff, parallel, pale golden bristles which are somewhat downcurved. Mandibles smooth, almost impunctate, pale brown, the apex obscured, almost black. Anterior rim of clypeus strong, ferruginous. The neuration of the strongly brownish amber-coloured fore wings is practically identical with that of 7. pendlebu- ryi (Lieftinck, 1944, fig. 1; and id., 1972, fig. 11). Abdominal tergite 7 (fig. 21) almost flat, surface densely rugoso-punctate, interspaces somewhat shiny, hairs pale, depressed. Body length 9.5 mm approx, fore wing 6.8 mm. T. habropodae is undoubtediy most closely related to himalayana and formosana. It is of in- terest to note that in Thailand the species was found flying in company with its supposed host bee, Habropoda sutepensis Cockerell (for the latter, see Lieftinck, 1974: 198, 199, figs.). As we now know, sutepensis is a near ally of H. apostasia Lieft., which in N. India occurred to- gether with Tetralonioidella himalayana. Tetralonioidella nepalensis spec. nov. (figs. 10, 22—26) Type material. — 2 d (terminalia of holotype dissected out, in pinned plastic capsule), E Hi- malayan Range, Nepal, Kathmandu, Godavari, 5000 ft., 6.viii. 1967, Canad. Nepal Exped. (ho- lotype); and same loc., 6000 ft., 24 July 1967, same coll. (paratype). Holotype 4 in Biosyst. Res. Inst., Ottawa (Ontario); paratype d in Mus. Leiden. Male. — Characters as given in the key and as shown in the figures. This is the third of a small group of slender species characterized by a bicoloured pubescent body pattern: a long and dense fox-red vestiture covering the summit of head and all of the tho- racic dorsum and sides, contrasting with an al- most black unbanded abdomen. Differs from the Malaysian pendleburyi and vulpecula in the paler colour of the very short appressed hairs on parts of the abdominal tergites. In vulpecula these short hairs are almost black and evenly distributed, in the other two lighter and most marked at the postgradular lines, especially so in the present new species. In the type the yellowish brown postgradular hairs on tergites 1—4 are broadest and longest at the sides of 1, more or less broadly interrupt- ed by dark brown in the median line on tergites 2—4. The paratype differs from the type in that the pale hairs on 1 cover most of the surface and are evenly distributed, whereas on 2—4 they are TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 somewhat more crowded together, forming in- conspicuous but uninterrupted bands. These complete bands are wholly absent also in pendlebury1. Tetralonioidella vulpecula (Lieftinck) (figs. 15, 3743) Callomelecta vulpecula Lieftinck, 75—77, pl. & figs. (6 2 Sumatra). Protomelissa vulpecula; Lieftinck, 1972: 272 (key), 282, pl. 1 fig. 2(d © Sumatra, refs.). 1944: 68—71 & No additional material. Tetralonioidella pendleburyi (Cockerell) comb. nov. (fig. 16) Callomelecta pendleburyi Cockerell, 1926: 621, 622 (2 Malaya, Selangor). — Lieftinck, 1944: 62—68, | figs. 1—9 & 75—77 (incl. keys Qd and descr. al- lotype d, Perak and Pahang, Malaya). Protomelissa pendleburyi; Lieftinck, 1972: (key), 282, figs. 9—16 (& 2 Malaya, full refs.). No fresh material. For structural details of the only known d of this species, see Lieftinck, 1944 & 1972. On p. 62 line 17 in my 1944 pa- per, “® allotype” should be altered in “& allo- type”. Tetralonioidella fukienensis spec. nov. (figs. 11, 27—30) Type material. — 1 d (terminalia dissected out, in pinned plastic capsule), SE China, NW Fukien, Chungan distr., Bohea Hills, 600 m, 25.1x.1939, T. C. Maa. Holotype d in Mus. Lei- den. Male (slightly discoloured, but otherwise in fair condition). A species sufficiently character- ized in the key. Superficially most closely re- sembling 7. insidiosa, from West Java, but easily distinguished therefrom by comparing the structural illustrations here given for both. Dif- ferences not mentioned in the key are that fu- kienensis, besides being slightly larger 1 in size, Is decidedly a more robustly built species than ın- sidiosa, especially the mid and hind femora and tibiae of fukienensis being markedly more swol- len (expanded) about midway their length, than in the slenderer insidiosa. Tetralonioidella insidiosa (Lieftinck) comb. nov. (figs. 12, 31—36) Callomelecta insidiosa Lieftinck, 1944: 71—75 & 75— 77, pl. & figs. (6 2 Java). LIEFTINCK: Melectine and Anthophorine bees 281 Fig. 31—36. T. insidiosa (Lieft.), & and ® structures, W. Java; fig. 31, apex of d tergite 7, ventral view; fig. 32—33, d sternites 8 and 7, external view; fig. 34—35, & genital capsule, partial dorsal and ventral view; fig. 36, 2 pygidial plate, dorsal view (after Lieftinck, 1944). Protomelissa insidiosa; Lieftinck, 1972: 272 (key), 282, pl. 1 fig. 1 (d © Java, further notes). No more recently collected material. NEW NAMES AND SYNONYMS IN THE GENERA ELECTA ATREILLE 1802, AND HABROPODAF. SMITH, 1854 After publication of my “Prodrome” to a mo- nograph of Palaearctic Melecta species (Lief- tinck, 1980), Mr. D. B. Baker, of Ewell, indi- rectly brought to my notice that two specific names introduced in the Prodrome are preoccu- pied by those given to different taxa in the same genus by E. Newman, in 1835. As the last-men- tioned names were followed by Latin diagnoses and descriptions in English, they are undoubt- edly validly proposed and should take prece- 282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 dence over those accidentally also chosen by myself. By an unfortunate oversight, Newman’s introduction and diagnoses escaped my atten- tion, while his names (extracted from Dalla Torre’s catalogue) were inadvertently consid- ered nomina nuda and therefore left out of con- sideration in my Prodrome. As a matter of fact, on pp. 513, 514 of his Entomological Notes (Newman, 1835), the author characterized six British “species” of Melecta, which he all named. Two of them were described as M. alec- to and M. megaera, both from localities in southern England and admittedly regarded by Newman himself as mere variants of the com- mon Melecta punctata (Fabricius, 1775), a spe- cies now properly called M. albifrons (Forster, 1771)!). The four remaining specimens of Me- lecta, all of them collected in England, were de- scribed in the same way to the former, receiving the names M. atropos, clotho, lachesis, and tisi- phone. Although the validity of this nomencla- ture, like that of the other two, can not be called in question and should be borne in mind by fu- ture workers on the genus, they are of no con- cern in the present context. 1) In view of the many so-called “Spielarten” occur- ring in the entomological literature of the past, I am tempted to quote certain passages in Newman's cheerful introduction to the genus Melecta preced- ing the definition of his novelties, — contempla- tions which may give an idea of his and some con- temporary authors’ appraisal of specific names and the valuation of nomenclatural problems in gener- al: — “It is pleasant to create a smile; and I anti- cipate that many smiles will be accorded me when I coolly assure your readers, that I am going to make six distinct species out of Melecta punctata, and that I cannot, for the life of me, tell to which of these the name punctata properly belongs, and therefore have given it to neither. It is pleasant to see one’s new species given, without comment, as synonyms; and when this happens, and happen it certainly will to my Melectae, I hope I shall take it as good-humouredly as Mr. Waterhouse did, when he beheld his fourteen new Notiophili consigned to utter oblivion”. “In these cases there is this com- fort, that if the new-made species are really species, they will in the course of time be re-admitted: al- low a year for each really new species parted from an old one; then the Notiophili will be re-admitted by the year 1847, and the Melectae — for I reckon them already struck out — by the year 1841”. Indeed, as time went on, re-adjustments in the nomenclature became almost prevailing topics of action, — though not always exactly in the way Edward Newman had predicted! Summarizing the above, it will be clear that the two specific names M. alecto Lieftinck and megaera Lieftinck, are preoccupied and should be substituted by new names, as follows: M. diligens nom. nov. pro M. alecto Lief- tinck, 1980: 164, 186 (keys) & 273 (nom. pre- occ.). — Holotype d, Iraq, in Brit. Mus. (Nat. Hist.), London. M. mundula nom. nov. pro M. megaera Lief- tinck, op. cit.: 175, 185 (keys) & 243 (nom. pre- occ.). — Holotype d, Cyprus, in Mus. f. Na- turk., Berlin DDR. In the same work (1980), the following cor- rections should be made: p. 140 line 12, the wording “specific of varietal” should read “spe- cific or varietal”; p. 290, line 11 from bottom, “figs. 231—232” should read “fig. 231”; p. 291, in the explanation of figs. 231-236, the word “antenna” should, of course, be replaced by “maxillary palpus”, shown in fig. 232. I shall gratefully accept any information about further errors and omissions occurring in this publication. In the genus Habropoda F. Smith, the follow- ing case of synonymy could be established: Habropoda apatelia Lieftinck, 1974: 171, 178 (keys), 215, 216, figs. d 2 Darjeeling, Syn. nov. This taxon proves to be conspecific with the long-lost H. krishna Bingham, 1909 (Rec. Ind. Mus. 2: 366—367, & Darjeeling 7000 ft.). Con- firmed after direct comparison of morphology and internal structures of the well-preserved holotype of krishna (still in coll. Zool. Survey of India, Calcutta) with topotypical holotype of apatelia in the Brit. Mus. (Nat. Hist.) and para- types in the Leiden Museum. ACKNOWLEDGEMENTS My best thanks are due to a number of ento- mologists who, through the years, kindly com- plied with my requests to recover certain melec- tine types and other valuable anthophorid speci- mens in various museum collections abroad. They kindly arranged for long-term loans to be made, and for waiting patiently for the ultimate return of specimens under their care: Messrs. H. Fankhänel (Leipzig, DDR), and G. Morge and J. Oehlke (both D.E.I., Eberswalde), for the loan of the long-concealed type of Tetralo- nioidella ? hoozana Strand. I am also grateful to J. G. Rozen Jr. (Amer. Mus. Nat. Hist., New LIEFTINCK: Melectine and Anthophorine bees 283 Di Fig. 37—43. T. vulpecula (Lieft.), d and 2 structures, S. Sumatra; fig. 37, apex of d tergite 7, ventral view; fig. 38—39, d sternites 7 and 8, external view (sg slightly more downpressed); fig. 40, right lateral view of distal portion of genital capsule; fig. 41—42, apices of & genital capsule, dorsal and ventral view; fig. 43, © pygidial plate, dorsal view (after Lieftinck, 1944). York) for the type loan of Callomelecta habro- podae Ckll.; to J. R. Barron, J. E. R. Martin and W. R. M. Mason (Biol. Res. Sta., Ottawa, Ont.), for specimens collected by members of the Ca- nadian Nepal Expedition 1967, which included the type of Tetralonioidella nepalensis spec. nov.; and to T. C. Maa (Taichung, Taiwan), for his great interest in the cuckoo bees of the pre- sent genus and their hosts, as well as for numer- ous Anthophoridae assembled by him in Fukien during the first years of World War II, on which occasions the type of T. fukienensis spec. nov. was discovered. Lastly, I am indebted to L. B. Holthuis (Leiden), who found out the exact dates of issue of some articles, which appeared almost simultaneously in different European journals. 284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, AFL. 12, 1983 SELECTED REFERENCES Cockerell, T. D. A., 1911. Descriptions and records of bees. 34. — Ann. Mag. Nat. Hist. (8) 7: 225— PDK ——, 1926. Descriptions and records of bees. 113. — Ann. Mag. Nat. Hist. (9) 18: 621, 622 (part). ——, 1929. Descriptions and records of bees. 117. — Ann. Mag. Nat. Hist. (10) 4: 132, 133 (part.). Friese, H., 1911. Neue Bienenarten von Formosa und aus China (Kanton). — Verh. zool.-bot. Ges. Wien 59: 127, 128 (part.). —, 1914. Neue Bienenarten der orientalischen Re- gion. — Deutsch. Ent. Zeitschr.: 322—324 (part.). Lieftinck, M. A., 1944. Some Malaysian bees of the family Anthophoridae (Hym., Apoidea). — Treu- bia (Buitenzorg), hors sér.: 57—138, 79 figs, pl. 42. ——, 1972. Further studies on Old World melectine bees, with stray notes on their distribution and host relationships (Hym., Anthophoridae). — Tijdschr. Ent. 115: 253—322, 2 pls., 57 figs., 1 tab. & 2 maps. (With full bibliography). | ——, 1974. Review of Central and East Asiatic Ha- bropoda F. Smith, with Habrophorula, a new ge- nus from China (Hym., Anthophoridae). — Tijdschr. Ent. 117: 157—224, figs. 1— 100. , 1980. Prodrome to a monograph of the Pal aearctic species of the genus Melecta Latreille 1802 (Hym., Anthophoridae). — Tijdschr. Ent. 123: 129— 349, text-figs. 1—359, pls. 1—8. Newman, E., 1835. Entomological Notes. In: The Entomological Magazine, London 2 (5): 512—514 (part: Melecta). Sandhouse, G. A., 1943. The type species of the gen- era and subgenera of bees. — Proc. U.S. Nat. Mus. 92: 519— 619. Strand, E., 1914. H. Sauter’s Formosa Ausbeute. Api- dae. III. — Arch. f. Naturgesch. 80 A (1): 139— 141 (part.). 9478 034. "e Pri 3 | : è 2 ia a m TE AE 1) ” | i a 14" ; = here AIS (À de a es be pelt AR pal Por ui wi | AE Ke he rf en « n £ Y là L a = DHL Van AT B aah - ‘ #1 ae = 4 ai Pie > ed 7 fi L Sen = d SR, Sr Tad ui i “ I = I d hi | Ni: . JA a, = sal = 4 ni = à 0 È 4 Î È j di n ; ka + È - lai | EE È a è, = N a - DA A Lite > ar i u , x i Lx us LEA gs I ; 5 { > \ ft N x 5 kr TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING REGISTER VAN DEEL 126 * Een sterretje duidt aan een naam nieuw voor de wetenschap * An asterisk denotes a name new to science Of the contribution by R. de Jong — Annotated list of the Hesperiidae of Surinam — of the names of species and subspecies only those of the taxa described as new are included. AMPHIBIA Rana esculenta 75 AVES Apus apus 75 Chlidonias niger 75 Delichon urbica 75 Fulica atra 83 Hirundo rustica 75 Sterna hirundo 75 COLEOPTERA apterus, Lethrus 205, 226, 227 biostius, Typhaeus 205 crassipes, Donacia 83 daphnis, Phanaeis 226 egeriei, Geotrupes 205 farctum, Bolboceras 205 ferrugineus, Bradycinetulus 205 fossor, Typhaeus 206 gaigei, Mycotrupes 205 hiostius, Typhaeus 206, 211, [226, 227 hispanus, Copris 226 hornii, Geotrupes 205 lateridens, Typhaeus 206 momus, Typhaeus 205, 206 mutator, Geotrupes 205, 226 nigroaeneus, Kheper 226 nymphaeae, Pyrrhalta 83 pomorum, Anthonomus 111 retusus, Mycotrupes 205 Sisyphus 226 spiniger, Geotrupes 205, 266 i [sqq stercorarius, Geotrupes 205, [226 sqq stercorosus, Geotrupes 205 typhoeoides, Typhaeus 206 typhoeus, Typhaeus 203 sqq vernalis, Geotrupes 205, 226 youngi, Peltotrupes 205 DICTYOPTERA americana, Periplaneta 227 DIPTERA Agromyza 12 Amaurosoma 10 aquilegiae, Phytomyza 8 bellidina, Phytomyza 8 brunnipes, Notophila 70, 87 Cerodontha 12 confluens, Hydromyza 59 sqq flavipennis, Pycnoglossa 7 griseola, Hydrellia 70 heraclei, Philiphylla 14 Liriomiza 11 livens, Hydromyza 59 sqq minuscula, Phytomyza 8 Pegomya 10 Phytomyza 11, 12 rosae, Psila 14 sylvestris, Cricotopus 72 trifasciatus, Cricotopus 72 Trilobomyza 11, 12 GASTROPODA Lymnaea stagnalis 83 HYDROZOA Cordylophora caspia 89 HYMENOPTERA acalle, Sympiesis 129 Acrolyta 112, 132 Adelura 4 Adesha 175, 176 * A deshoides 175 albifrons, Melecta 282 albitarsis, Euderus 129 albolineata, Adesha 177 alecto, Melecta 282 alternans, Itoplectis 111, 129, [132 285 americana, Orthostigma 13 amplidens, Adelurola 2, 5 angustula, Pentapleura 14 anxium, Agrypon 118, 130, 132 apafelia, Habropoda 282 Apechthis 129 *apheles, Gnaptodon 26, 28, 29 apii, Dapsilarthra 3, 13 Apophua 112, 114 apostata, Diadegma 117, 130, [132 arenicola, Tranosema 117, 130, [132 asiatica, Adelurola 2, 6 *asulcatus, Adeshoides 176 ater, Apanteles 112, 118, 130, [132 atropos, Melecta 282 bachmaieri, Gnaptodon 25, 31 balteata, Dapsilarthra 2, 8 barthii, Dapsilarthra 14 barthii, Phaenocarpa 14 bini, Gnaptodon 39 boozana, Tetralonioidella 269, [272, 274 Bracon 126, 132 brevicornis, Scambus 110, 127, [130, 132 breviradialis, Gnaptodon 28, 30 “brevis, Gnaptodon 28, 30 Callomelecta 270 Campoplex 130, 132 canaliculatus, Habronyx 118, [130, 132 carbonarius, Microgaster 120 carinatus, Plesiobracon 186 *Carinibracon 185, 186, 187 “carpathica, Dapsilarthra 3, 12 cavus, Dibrachys 127, 131, 132 chrysos, Habrocytus 127, 132 cicatricosa, Apophua 114, 129, [132 claripennis, Goniozus 129 286 clotho, Melecta 282 complicator, Lissonota 116, [130, 132 compuctor, Apechthis 112, 132 corvina, Dolichogenidia 120, (131, 132 cruentatus, Charmon 126, 131, [132 dalhousiensis, Dapsilarthra 2, 10 danielssoni, Carinibracon 187 Dapsilarthra 6, 10 decoris, Gnaptodon 25, 29, 31 deserticola, Victoroviella 179 Diadegma 127 “diligens, Melecta 282 dictynna, Dapsilarthra 2, 7 difformis, Compoplex 117, 132 dilecta, Dolichogenidia 120, [130, 132 dimidiator, Agathis 122, 130, [132 Diraphus 27 *erasmi, Gnaptodon 29, 32 errabunda, Lissonota 116 Exochus 118, 129, 132 fenestralis, Diadegma 117, 130, [132 flaviventris, Alysia 11 florimela, Adelurola 2, 5 florus, Colpoclypeus 111, 127, [131, 132 formosana, Tetralonioidella [269, 271, 274 fractigena, Glypta 116 fukiensis, Tetralonioidella 272, [280 fuscula, Dapsilarthra 11 gahani, Dapsilarthra 2, 7 Gelis 112, 132 *georginae, Gnaptodon 29, 33 glaber, Gnaptodon 29 globulipes, Triclistus 117, 118, [129, 132 Glypta 112 Gnamptodon 6 gracilicornis, Alysia 11 Grammospila 6 grandidieri, Malagopsis 177, 181 Habrocytus 131 habropodae, Tetralonioidella [271,275 horstoki, Deltaphyma 182 ictericus, Meteorus 117, 123, [131, 132 indagatrix, Dapsilarthra 2,9 insidiosa, Tetralonioidella 272, [274, 280 L interrupta, Diadegma 116 iridescens, Protomelissa 270 isabella, Dapsilarthra 3, 11 Kenema 186, 188 klemensiewiczii, Graptodes 25, BI krishna, Habropoda 282 lachesis, Melecta 282 laevigata, Dolichogenidia 120, [130, 132 laevipectus, Alysia 13 lanceolator, Lasiophorus 180 Lasiophorus 179 levisulca, Dapsilarthra 2, 8 linearis, Macrocentrus 126, 131, [132 lineatus, Stictopisthus 117, 132 liogaster, Zanzopris 183 Lissogaster 122, 130, 132 longicaudus, Dolichogenidia [120, 130, 132 maculator, Itoplectis 111, 130, [132 megaera, Melecta 282 Malagopsis 181 Mesochorus 130 Mesocrina 6, 8 minutus, Oncophanes 123, 127, [131, 132 mongolica, Gnaptogaster 26 multiarticulata, Phaenocarpa 5 *mundula, Melecta 282 nana, Lysibia 112, 132 Neocarpa 4 nepalensis, Tetralionioidella [272, 280 nepalicus, Gnaptodon 29, 34 nepticulae, Gnaptodon 29 *nieukerkeni, Gnaptodon 29, 34 nigrina, Glypta 114, 129, 132 nowskowskii, Dapsilarthra 7 obscurator, Bracon 126, 131, [132 ochrogaster, Phaenocarpa 11 “orientalis, Gnaptodon 40 pallipes, Macrocentrus 127, 131, [132 pallipes, Triclistus 117, 118, [129, 132 Paraorthostigma 6 pectinicornis, Pnigalio 127, 131, [132 pedata, Glypta 116 pendleburyi, Tetralonioidella [274, 280 pentapleuroides, Dapsilarthra 5 *pilosus, Gnaptodon 28, 34 Plaxopsis 184 Plesiobracon 185, 186 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, 1983 praerogator, Diadegma 116, [130, 132 Protomelissa 269 Pseudomesocrina 6, 8 pulchrigaster, Gnaptodon 29 pumilio, Gnaptodon 29, 35 punctata, Melecta 282 pygmaeus, Diraphus 25, 35 quadridentata, Ascogaster 123, [130, 132 quadridentatus, Apechthis 112, [132 quickei, Kenema 188 resinator, Apechthis 112 rufatus, Apechthis 112, 132 *ruficeps, Gnaptodon 29, 37 rufidens, Ascogaster 123, 130, [132 rufipes, Agathis 122, 130, 132 rufiventris, Dapsilarthra 3, 11 rufiventris, D., f. fuscula 3, 11 sauteri, Anthophora 269, 270 segmentator, Phytodietus 112, (130, 132 semotus, Habrocytus 127, 132 sertifer, Neodiprion 111 servillei, Bracon 183 seyrigi, Malagopsis 182 sicaria, Dolichogenidia 120, [130, 132 silvarum, Mesochorus 117, 132 *similis, Gnaptodon 39, 40 sjoestedti, Plaxopsis 185 striata, Teleutaea 112, 113, 129, [132 subtilis, Dapsilarthra 2, 8 subtilis, Heterolexis 1 sulcifera, Dapsilarthra 14 sylvia, Dapsilarthra 3, 12 tenuicornis, Dapsilarthra 14 Tetralonia 269, 274 Tetralonioidella 269, 270 testacea, Dapsilarthra 8 thienemanni, Porima 1, 3 thoracicus, Macrocenthrus 126, [131,132 tirolensis, Dapsilarthra 3, 12 tisiphone, Melecta 282 tricolor, Tetralonioidella 270, [275 unifossa, Gnaptodon 39 varicoxa, Glypta 114, 116, 129, [132 venatrix, Mesocrina 9 Victoroviella 178 *vlugi, Gnaptodon 28, 38 vulpecula, Tetralonioidella 274, [280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, 1983 westwoodi, Dicladocerus 129 xanthostigma, Apanteles 120, [130, 132 xanthostoma, Sympiesis 127, 1182 *Zanzopsis 183 LEPIDOPTERA Ablepsis 241 Achlyodes 243 Aecas 244 aeneofasciella, Stigmella 37 agrimoniae, Ectoedemia 31 Aguna 251 Aides 248 *ambiguus, Urbanus 239, 252 Amenis 237 amygdali, Stigmella 30 Anastrus 243 angustiorana, Batodes 123 Anisochoria 243 Anthoptus 244, 255 Antigonus 243 Apaustus 244, 255 Artines 244 Aspitha 237 Astraptes 239, 240, 252 Athoptus 253 Atrytone 247 Augiades 237 aurella, Stigmella 31 Autochton 240 bergmanniana, Croesia 122 betulicola nanivora, Stigmella (82537 *bicuspis, Elbella 236, 250 brumata, Operophtera 120 Bungalotis 240, 252, 253 Cabirus 238 Calliades 240 Callimormus 244 Calpodes 248 Cantha 244 caradjai, Ectoedemia 37 carpinella, Stigmella 37 Carrhenes 243 Carystoides 246, 262 Carystus 246, 262 Cecropterus 240 Celaenorrhinus 241 Cephise 241 cerasana, Pandemis 109, 114, [116, 117, 123, 126, 132 confusella, Stigmella 37 Charidia 242 Chioides 238 Chiomara 243 Choranthus 247 Chrysoplectrum 239 Clito 243, 255 Cobalopsis 245, 257, 261 Cobalus 245, 246, 247 Cogia 241, 254 Conga 247 Conognathus 241 continuella, Stigmella 37 Corticea 244 corylana, Pandemis 123 *Cravera 248, 265 criska, Penicula, 247, 263 “subsp. extensa 247, 263 cryptella, Trifurcula 31 Cycloglypha 243 Cyclosemia 242 Cymaenes 244, 255 Cynea 247, 263 *dorpa, Cobalopsis 245, 257 dorycniella, Trifurcula 38 Drephalys 237 Dubiella 246 Dyscophellus 240 Damas 246 Decinea 247 eberhardi, Stigmella 35 Ebrietas 242 Ebusus 246 Ectomis 241 Elbella 236, 250 Enosis 246 Entheus 238, 250 Epargyreus 238 Ephyriades 243 Eracon 241 erythrogenella, Ectoedemia 37 Euphyes 247 Euroto 244, 245 Eutocus 244 Eutychide 246, 261 forterana, Lozotaenia 120 fragariella, Stigmella 31 geminella, Stigmella 31 *geijskesi, Cymaenes 244, 255 Gindanes 242 Gorgythion 242 groschkei, “Ectoedemia” 33 Helias 243 Heliopetes 243 heparana, Pandemis 109, 112, [114, 120, 123, 126, 132 Hesperia 243, 245, 247 hexapetalae, Ectoedemia 30 holmiana, Croesia 132 Hyalothyrus 237, 238 hybnerella, Stigmella 33 Hylophila 247 Jemadia 237 287 Jemado 250 Joanna 247 *Jonkersi, Clito 243, 255 Justinia 246 lapponica, Stigmella 37 lecheana, Pandemis 112, 114, [116, 120, 132 Lento 244 Lerema 245 loranthella, Niepeltia 30 lucella, Teleia 122 luteella, Stigmella 37 Lycas 247 magdalenae, Stigmella 37 mahalebella, Ectoedemia 30, 37 "major, Vehilius 246, 256 malella, Stigmella 33 malinella, Yponomeuta 120 *meesi, Phlebodes 245, 263 Megistias 244, 245 Mellana 247 Metionopsis 244 Metron 247, 248 Milanion 242 mitterbachiana, Ancylis 112 Mnaseas 244 Mnasilus 245, 256 Mnasitheus 245, 256 Mnestheus 244 Moeris 245 Moeros 246 Molo 244 Monca 244 Morvina 242 Morys 245, 261 Mylon 242, 243 Myrinia 242 Myscelus 237 Mysoria 237 naevana, Rhopobota 110, 118, [132 Naevolus 246 Nascus 241 Nastra 244 Neoxeniades 248, 267 Niconiades 248 Nisoniades 241, 242 Nosphistia 236 nubiferana, Hedya 109, 110, HU IEI Nyctelius 248 Nyctus 246 nymphaeata, Nymphaea 83 occultella, Stigmella 32 ocellana, Spilonota 109, 110, [112, 116, 118, 122, 123, PRE den Onophas 246 288 orana, Adoxophyes 109, 111, (114, 116, 117, 120, 123, 126, [127, 129, 132 Orneatus 241 Orphe 246 Orses 247 Ouleus 242 Oxynthes 248 Paches 242 Pachyneuria 242 Pamphila 239, 244, 247 Panoquina 248, 265 panoquinoides, Panoquina 248, [265 “subsp. minima 248, 265 Papias 245, 256 Papilio 239, 240, 242, 243, 246, [248, 250 Parphorus 245 Paramimus 242 Paracarystus 245 Passova 237 Pellicia 242 Penicula 247, 263 Perichares 247, 263 Parihares 247 Phanus 237 Phareas 238 Phlebodes 245, 247, 263 Phocides 237, 250 plagicolella, Stigmella 32 Plumbago 242 podana, Archips 109, 112, 116, [118, 120, 126, 127, 132 Polites 247, 263 Polyctor 241 Polythrix 238, 239, 251 pomella, Stigmella 37 pomonella, Laspeyresia 123 Pompeius 247 porphyrana, Eudemis 112 Porphyrogenes 241 Polygonus 238 poterii, Stigmella 31 Propertius 248 Proteides 238, 246 prunetorum, Stigmella 39 Pyrdalus 241 Pyrgus 243 Pyrrhopyge 236, 237, 248, 250 Pyrrhopygopsis 248 Pythonides 242 Quadrus 242 Quinta 247 “rara, Cravera 248, 265 rhamnella, Stigmella 37 rhamnophila, Stigmella 34 rhombana, Acleris 117, 118 rosana, Archips 109, 112, 117, [118, 122, 132 ruficapitella, Stigmella 33, 37 Salatis 240 Saliana 248 salicis, Stigmella 37 Sarmientoia 240 Saturnus 247 Silius 244 “similis, Mnasitheus 245, 256 *sipa, Bungalotis 240, 252 Sodalia 244 Sostrata 242 Spathilepia 241 spectrana, Archips 118, 132 Spioniades 241 splendidissimella, Stigmella 31, [37 sorbi, Stigmella 37 subgrisea, Morys 245, 261 “subsp. paradoxa 245, 261 subpunctata, Entychide 246, [261 “subsp. intermedia 246, 261 *Surina 248, 266 Synapte 244 Talides 246, 262 *tanna, Cobalopsis 245, 261 Tarsoctenus 237 Telegonus 239, 241 Telemiades 241 Telles 246 Tellona 246 tetraquetrana, Epinotia 122 Thargella 244 Thespieus 248 Thoon 245, 246 Thorybes 240 Thracides 246, 248 Thymele 240 uliae, Stigmella 37 Timochreon 243 Turesis 245 Tityrella, Stigmella 33 Typhedanus 238 uddmanniana, Notocelia 120 Udranomia 237 ulmivora, Sugmella 30, 37 *unica, Surina 248, 266 Urbanus 239, 252 Vacerra 248 Vehilius 245, 256 Venas 244 Vertica 246 Vettius 245, 261 *vibicoides, Polites 247, 263 Vinius 244, 255 Viola 242 TiJDSCHRIFT voor ENTOMOLOGIE, DEEL 126, 1983 Virga 244 viridana, Tortrix 112 Wallengrenia 247 weirana, Strophedra 118 Xeniades 248 Xenophanes 243 xylosteana, Archips 112, 118, [132 Zariaspes 244 Zera 242 ODONATA abbreviatus, Progomphus 137, [138, sqq batesi, Zonophora 148, 150, 156 calippus, Zonophora 148, 149, [150, 154 spp. spectabilis 148, 149, 150, [155 spp. klugi 148, 149, 150, 155 campanulata, Zonophora 145, [148, 150, 151 “spp. annulata 145, 148, 150, [152 ssp. machadoi 145, 148, 150, [152 diversa, Zonophora 145, 148, [150, 153 nobilis, Zonophora 149, 150, [159 *occidentalis, Progomphus 137, [140 phyllochromus, Progomphus [137, 138 polygonus, Progomphus 137, [138 regalis, Zonophora 149, 150, [158 solitaria, Zonophora 149, 157 ssp. obscura 149, 150, 157 supratriangularis, Zonophora [145, 149, 150, 158 surinamensis, Zonophora 145, [150, 156 wucherpfennigi, Zonophora [145, 148, 150, 156 Zonophora 146 ORTHOPTERA bormansi, Ephippiger 91, 97, [98, 102, 104, 106 cruciger, Ephippiger 91, 97, 98, [102, 107 cunii, Ephippiger 91, 97, 99, [102, 104, 107 ephippiger, Ephippiger 91, 93, [97, 98, 102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 126, 1983 ephippiger vicheti, Eph. 97, 98, [99, 104, 106, 107 ephippiger vitium, Eph. 97, 98, [104, 106, 107 provincialis, Ephippiger 91, 97, [99, 102, 104, 106 terrestris, Ephippiger 91, 93, 97, [98, 99, 104, 106 terrestris caprai, Eph. 106 PLANTAE Agrimonia agrimonoides 31 Agrimonia eupatoria 37 Betula 37 Betula nana 32 Betula verrucosa 37 Calluna vulgaris 206 Caltha palustris 61 Carpinus 118 Cladonia 206 Coronilla varia 31 Crataegus monogyna 33 Doryenium germanicum 38 Fagus 118 Fagus sylvatica 33 Filipendula vulgaris 31 Fragaria vesca 31 Glomerella cingulata 88 Loranthus europaeus 30 Lotus corniculatus 31 Malus domestica 37 Malus sylvestris 33 Mentha aquatica 71 Nardus stricta 206 Nymphaea alba 61, 69 Nymphaea candida 61, 69 Nuphar advena 61, 89 Nuphar intermedia 61 Nuphar lutea 59 sqq Nuphar pumila 61, 89 Nupharia rivularis 59 Nymphoides peltata 69 Ostrya carpinifolia 37 Potentilla 32 Potenulla erecta 31 Prunus spinosa 32 Prunus cocomilia 30 Prunus mahaleb 30 Pteridium aquilinum 7 Quercus 33 Quercus pubescens 35 Quercus robur 37 Rhamnus catharticus 37 Rhamnus saxatilis 34 Ribes nigrum 120 Rosa pimpinellifolia 31 Rubus 31, 37 Rubus fruticosus 37 Rumex hydrolapathum 70 Salix caprea 37 Sanguisorba officinalis 31 Scirpus lacustris 70 Sorbus aucuparia 37 Tilia 37 Typha angustifolia 70 Ulmus 30 Ulmus glabra 37 Ulmus procera 37 Vitex agnus-castus 33 289 ids NON een ui tee peat zend wisi pi À a 0 sane 8 u aud, oa | anne) i. spar FLE HELENE mi este gn sh vat why gil ea): mdk vs bike ET Mi DATE NIE anke enen I; | r nn cd IMRE: OE plan uao i l ua pt hore Oy Sw shale Pee MAT ANR BEAN PL AAN) He a 4 vy H Went ge a ‘ ba altri debt Grades pi | ate iu me Milo i Lr a a ‘tame ts RAN i oy NEON, RR Le ERNST MAYR LIBRARY Bi ende FM ee ST LATE gas o sele 4 VA elem A TA kr ¥ LNA ARIA sees ak, Po hoa), L PELLE OO Lon ANCORA PETITE ORN DECT CELIA I SH CENT SRI NOT CA AL as En PROS rte, rdt En apne Aje viv AO TANT EN aasde AERO ande Mets error le Io BEN Het WA ARTE PAT Abra. i YU HOTTE Pate She f doi DETTO WARD PHASE EE schilde) nn ? 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REKEN RATEN tn GROEN EEL ET Wb LTE LATE gu MUR vu +0 DENT DR) He s'u HIER Win. i “if i weg AEN N BR deg SPW gr TETE x it N vn Wiel Ong =u ore CHA Kij È KUNG, Nana, SA les hie COURANTE bbb dj TE ty OASIS dirar 4,1 Wbp ard re the